LIST OF CONTENTS.

1912, Part I. (pp. 1-240).

EXHIBITIONS AND NOTICES.

Page

Tux Secretary. Report on the Additions to the Society's Menagerie during the months

of June, July, August, and September, 1911 .............-.......- Otago Bee Ne t

Mr. James Dunsar-Brunron. Exhibition of skins and skull of a Bushbuck from N.E.

BERL Seats ells aparece tote casi tte elo tee yy late ale tare ale sc fapn PaiMRemeette can TEN Lt ai a a 4

Mr. W. B. Corron. Exhibition of heads and horns of various species of Gazelles from

the Eastern Sudan........- rn Naat ave Neto deles eeheyctals sieve etadeteterteealel: tele stem civiegels etetetmereg Hioie |

Mr. D. Suru-Smitu, F.Z.S. Exhibition of a spirit-specimen of a nestling Australian

Regent-Bird (Sericudus mefinus). (Text-fig. 1.) .........0-2. Dislsters (cape ane ere eee a oe

Mr. R. I. Pococs, F.R.S., F.L.S8., F.Z.S. Exhibition of photographs of, and remarks

upon, a new race of Red-fronted Gazelle (Gazella rufifrons hasteri). (Text-figs, 2, 3.).. 5

Mr. J. Lewis Bonuors, M.A., F.L.S., F.Z.S. Exhibition of, and remarks upon, living

specimens of Rats (Mus rattus) which showed the “‘ waltzing” character .........- 6

Yur Secretary. Report on the Additions to the Society’s Menagerie during the month

of October 1911 .......0... oka doses BOOM e aa tod Shao eine hain wdinie Mihaela eek 142

Mr. RB. I. Pocock, F.B.S., F.L.8., F.Z.S. Exhibition of, awd remarks upon, living speci-

mens of the Common Elephant-Shrew (Macroscelides proboscideus) and the Rock ;

Blephant-Shrew (Hlephantulus rupestris). (Text-figs. 16, 17.). .- 2.0 ..eceese ce eeee 142

Luz Secrerary. Report on the Additions to the.Society’s Menagerie during the months

of November and December 1911 and January 1912 ..... De hls stet oe ceeeee eee 231

Mr. E. G. B. Muave-Watvo, V.P.Z.S. Exhibition of a pale fulvous variety of the

Common Polecat (Putorius putorits) .. 0°. ..... ae ence ren crss ce ieee eee 234

Mr. Rosert D. Carson, C.M.Z.S. An account of retarded development of the foetus in

a Red Kangaroo (Macropus rufus). . co... es ts Sete Pee Sen EER Oo 9: 234

Contents continued on page 3 of Wrapper.

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

ZOOLOGICAL SOCIETY

OF LONDON,

1912, pp. 1-504,

wiTH 60 Puates and 63 TExt-FIGURES.

’

PRINTED FOR THE SOCLETY,

AND SOLD AT THEIR HOUSE IN REGENT’S PARK.

LONDON:

MESSRS. LONGMANS, GREEN, AND CO,

PATERNOSTHR ROW.

of a

> 22 Ss

(orally

OF THE

COUNCIL AND OFFICERS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

COUNCIL.

His Grack THE DUKE OF BeprorD, K.G., F.R.S., President.

Tur Earp or Auramont, F.S.A. | E. G. B. Meape-Watpo, Esq.,

Str Joun Rose BRADFORD, Vice-President. -

K.C.M.G., M.D., D.Sc.,.| Prof: Epwarp\_A. . MINcHIN,.

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

Ricoarp H. Burne, KHsq., | P. CHALMERS MircHewy, Ksq.,

(

M.A. | MRA. DSc.) scons nieae

Lt.-Col. Sir R. Havetock | F.RS., Secretary.

Cuartes, G.C.V.O., M.D. _W. R. Octivin-Grant, Esq.

Aurrep H. Cocks, Esq., M.A. | AvBpEerr Pam, Hsq.

F. G. Dawtrey Drewrrr, Esq., | Aprian D. W. Poxttock, Esq.

Mi At Via): | OLDFIELD T'Homas, Hsq., F.R.S.

CHartes Drummonn, Esq, AntHony H. WHuINGFIELD,

Treasurer. Ksq.

Srr Epwarp Duranp, Bt., C.B. | A. Smrra Woopwarpb, Ksq.,

F. Du Cans Gopmax, Esq., ; LL.D.,F.R.S., Vice-President.

IDV Calli aL Mas), | Henry Woopwarp, Esq., LL.D.,

Sir Epmunp G. Loprmr, Bt., | F_R.S.. Vice-President.

Vice- President.

PRINCIPAL OFFICERS. |

P. Cuaumers Mrrcuenn, M-A.,D.Se., Hon.LL.D., F.R.S.,

Secretary.

Frank E. Bepparp, M.A., F.R.S8., Prosector.

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

Resident Superintendent of the Gardens.

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

Kpwarp G. BouLtencer, Curator of Reptiles.

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

F. H. Waternouse, Librarian.

JouNn Barrow, Accountant.

W. H. Cour, Chief Clerk.

LIST OF CONTENTS.

1912, pp. 1-504.

EXHIBITIONS AND NOTICES.

Page

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

Menagerie during the months of June, July, August,

and September, 1911

Mr. James DunsBAR-Brunton. Exhibition of skins and skull

of Bushbuck from N.E. Rhodesia

Mr. W. B. Corron. Exhibition of heads and horns of

various species of Gazelles from the Eastern Sudan ... 4

Mr. D. Seru-Surru, F.Z.S. Exhibition of a spirit-specimen

of a nestling Australian Regent-Bird (Sericulus melinus).

IES a Tia) i ee, EL rk ao 4

Mr. R. I. Pocoox, F.R.S., F.LS., F.Z.S. . Exhibition of

photographs of a new race of Red-fronted Gazelle

(Gazella rufifrons hasleri). (Text-figs. 2 & 3.)

Roa

eeececese

Mr. J. Lewis Bonnors, M.A., F.L.S., F.Z.S. Exhibition of

living specimens of Rats (J/us rattus) which showed

the “ waltzing” character

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

Menagerie during the month of October 1911

Mr. R. I. Pococn, F.R.S., F.L.S., F.Z.S. Exhibition of

living specimens of the Common Elephant - Shrew

(Macroscelides proboscideus) and the Rock Elephant-

Shrew (Llephantulus rupestris). (Text-figs. 16 & 17.).

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

Menagerie during the months of November and

December 1911 and January 1912

Mr. E. G. B. Mzape-Watpo, V.P.Z.S. Exhibition of a pale

fulvous variety of the Common Polecat (Putorius

ULOVIUS) BN cane FRR eel Mea” SS

Mr. Rosrrr D. Carson, C.M.Z.8. An account of retarded

development of the foetus in a Red Kangaroo (Macro-

UST UF US) a, eens ti SNA EET. a

Dr. A. T. Masrerman, M.A., F.Z.S. Demonstration of

recent investigations on Age-determination in the

Scales of Salmonoids

Mr. Ouprietp THomas, F.R.S., F.Z.S. The Races of the

Buropean Wald’ Sywinmes se an sale eee ee

Mr. A. Rapciyrre Duemore, F.R.G.S. Lantern exhibition

of photographic studies of wild animals in British East

Africa and Newfoundland

Messrs. KE. GERRARD & Sons. Exhibition of the head of a cow

Sable Antelope, the skulls of an Isabelline Bear, a

Leopard, and a Lion, all of which showed some ab-

normality

DO ICI SO ICICI II Tian iO ICnOiO ni iii oICnOICHO CR MPECIry oie OI OO OOO Od

Mr. OxprieLp Tuomas, F.R.S., F.Z.S. Two new Genera and

a new Species of Viverrine Carnivora. (Text-figs.

GIS GS )iecctstartasetnskneeuncaneenens4).til: 56 Rae ae eee

Mr. Guy Dotuman. A new Snub-nosed Monkey

142

231

234

390

498

PAPERS.

. Distant Orientation in Amphibia. By Bruce F. Cum-

mMInGs. (Text-figs. 4 & 5.)..........ccecsceteseereneeeeeeees

. Some Remarks on the Habits of British Frogs and Toads,

with reference to Mr. Cummings’ recent communi-

cation on Distant Orientation in Amphibia. By

Cu Aeonmencnn, LR BPE ihe. isos. sessed ecctvee.

. Game Sanctuaries and Game Protection in India. By

E. P. Srepeine, F.L.S., ¥.Z.8., F.R.G.S.

Pere

. On the Moulting of an Arctic Fox (Vulpes lagopus) in

the Society's Gardens. By R. I. Pocock, F.RS.,

F.LS., F.Z.S., Superintendent of the Gardens. (Text-

GMS ea ta ESN athe s as nsio vice teachable warete cd eeretamte's Sate tara ean ee ©

. On the Moulting of the King Penguin (Aptenodytes

pennanti) in the Society’s Gardens. By Davin Setu-

Smirn, F.Z.S., M.B.O.U., Curator of Birds. (Pl. 1.) .

. On the Presence of two Ovaries in certain British Birds,

more especially the Falconide. By ‘Tl. E. Gunn,

ead is a A (A 2) e e ey ee e

. On some Collembola from India, Burma, and Ceylon;

with a Catalogue of the Oriental Species of the Order.

By A. D. Imus, D.Se., B.A., Forest Zoologist to the

Government of India; late Professor of Biology,

Muir College, and Fellow of the University of

Allahabad. (Pls. VI.—XII. and Text-figs. 14 & 15.)..

. Ontogenetical Transformations of the Bill in the Heron

(Ardea cinerea). By Prof. P. P. Susuxrin, C.M.Z.S.,

hanmovy teussiae) (Pl. MUM ress ilies ccaesvencoseeses

. The Duke of Bedford’s Zoological Exploration of Eastern

Asia.—XV. On Mammals from the Provinces of Sze-

chwan and Yunnan, Western China, By OLDFIELD

SrOMAG) DSuiSee ete. «eres, txec cas eciyincs qotatieesecsuas’

Page

125

127

10.

tae

12:

13,

14.

15.

16.

17.

18.

The Freshwater Crayfishes of Australia. By GEOFFREY

Smirn, M.A., Fellow of New College, Oxford. (Pls.

XIV.-XXVII. and Text-fig. 18.)

weet erst ees eet ert oesene

Structure of the Alimentary Canal of the Stick-Insect,

Bacillus rossii Fabr.; with a Note on the Partheno-

genesis of this Species. By Atrrep EH. Cameron, M.A.,

B.Sc., Fullerton Scholar of the University of Aberdeen

and Research Student in the University of Manchester.

(EIS CNG PIA ONO oe hase peteee wacwine gnosis eee

Diagnoses of new Species of Terrestrial and Fluviatile

Shells from British and German Hast Africa, with the

Description of a new Genus (Hussoia) from Husso

Nyiro River, B.E, Africa. By H. B. Preston, F.Z.8..

(Pls. XXXI. & XXXII)

POOP ee Pee peepee penpererepeserenesese

Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.—III. On a new Genus of Tape-

worms (Ofiditenia) from the Bustard (Hupodotis kort).

By Frank EH. Bepparp, M.A., F.R.S., F.Z.S., Prosector

TO Ne ores” (bettie NO 30))) son sannodoonanonocosoes

On the Milk-Dentition of the Ratel. By R. Lyppxxmr.

(Text-figs. 31 & 32.)

Ceo meme ee meee ns eet pen per ese pero eseeeneroee

On a Further Collection of Mammals from Egypt and

Sinai. By J, Lewis Bonnors, M.A., F.L.S., F.Z.8. ...

Report on the Deaths which occurred in the Zoological

Gardens during 1911. By H. G. Puimer, F.R.S.,

F.Z.8,, Pathologist to the Society

CC i a eC CC eC eC nC ery

Mendelian Experiments on Fowls. By J.T. Cunnine-

SEAM RIVA IM ZiS, Mean atc ct mermcan tie. cc mete aece eee

Studies on Pearl-Oysters and Pearls.—I. The Structure

of the Shell and Pearls of the Ceylon Pearl-Oyster

(Margaritifera vulgaris Schumacher): with an Examin-

ation of the Cestode Theory of Pearl-Production. By

H. Lysrer Jameson, M.A., D.Sc., Ph.D., F.Z.5. (Pls.

XXXIIT.-XLVII. and Text-figs, 33-41.) ...........0005

Page

144

172

183

194

221

235

241

260

vil

Page

19. Mimicry amongst the Blattide; with a Revision of the

Genus Prosoplecta Sauss., and the Description of a new

Genus. By Rospert SuHetForp, M.A., F.Z.S. (PI.

SU Wem, Vext-fgs. 42-46.) oo. cccccecc esse cesccweses 358

20. On the Pairing of Pseudoscorpiones. By H. WaAt..is

Wee ( bext-fias, 47-50.) <2. ccc heccscdenceesves 376

21. A Contribution towards the Knowledge of the Spiders

and other Arachnids of Switzerland. By the Rev. O.

PiIcKARD-CAMBRIDGE, M.A., F.R.S., C.M.Z.S. (Text-

La ema) eer pg he eet eg Oth acca ied ale aye Sls s ses 393

. On the Blood-parasites found in Animals in the

Zoological Gardens during the four years 1908-1911.

By H. G, Pumwer, F.RS., F.Z.8., Pres.R.M.S.,

Pathologist to the Society. (Pls. XLIX.—LY.)......... 406

23. On the Structure of the Internal Ear and the Relations

of the Basicranial Nerves in Dicynodon, and on the

Homology of the Mammalian Auditory Ossicles. By

he Broom, M2D., D:se, CMZ.S8. (Pl LVE and

Pee tite, clel Aies Ne ey tem care Pataca eee sa wr yaice ce eta es 419

24. Zoological Results of the Third ‘Tanganyika Expedition

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

Report on some larval and young stages of Prawns

from Lake Tanganyika. By Prof, G. O. Sars, C.M.Z.8.

ae vlistae Wig Tele Wea yates narra cree n Qenet rail bene ny aticoie cake 426

25. The Classification, Morphology, and Evolution — of

the Echinoidea MHolectypoida. By Herserr L.

Hawkins, M.Se., F.G.S.; Lecturer in Geology, Uni-

versity College, Reading. (Text-figs. 54-60.) ......... 44()

mignabetical List of Comtributors: 22100........0-scenseenes ees, 1X

New Generic Terms ...... sista SRR ace Rew sield eu acsiein akeatip XV1

LAER OMSCLEMUMIC NEES? « \.4cces emesis aa cae oboe bcos cele ae ates aes XV1L

Index of Illustrations ..... NP nineties OE re eS xxix

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ALPHABETICAL LIST

CON T-REbBwU.PO.bs,

With References to the several Articles contributed by each.

(1912, pp. 1-504.)

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

to the Society.

Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea. III.—On a New Genus of Tape-

worms (Otiditenia) from the Bustard (Hupodotis kort).

(Text-figs. 19-30.)

CeCe reece een Hela ene Mea anesseCeeVEKG er dew aCe enews

Bonuorr, J. Lewis, M.A., F.LS., F.Z.8,

Exhibition of living specimens of Rats (Mus ratius)

which showed the “ waltzing” character

were rewesesessoeseere

On a Further Collection of Mammals from Egypt and

Sinai

ENE ey por ce War Tay ee BOR Re Aer LS YC CIEL Fm yr wo ie BC UC CA Ee LBL I a ol

194

BowuLencErR, GrorGE A., F.R.S., F.Z.S.

Some Remarks on the Habits of British Frogs and

Toads, with reference to Mr. Cummings’ recent com-

munication on Distant Orientation in Amphibia

Broom, Rosert, MEADS Disc. CoMEAS.

On the Structure of the Internal Ear and the Relations

of the Basicranial Nerves in Dicynodon, and on the

Homology of the Mammalian Auditory Ossicles. (EE

LVI. and Text-fig. 53.)

TPP emcee em em meee e reese reer esse erreeeeenane

Brunton, JAMES Dunspar-. See DunBar-Brunvon, J.

CAMBRIDGE, The Rey. O. Pickarp-. See PrcKARD-Cam-

BRIDGE, O.

CAMERON, ALFRED H., M.A., B.Sc., Fullerton Scholar of the

University of Aberdeen and Research Student in

the University of Manchester.

Structure of the Alimentary Canal of the Stick-Insect,

Bacillus rossii Fabr.; with a Note on the Partheno-

genesis of this Species. (Pls. XXVITI.-XXX.) ....:.00: Nas

Carson, Rospert D., C.M.Z.S.

An account of retarded development of the feetus in a

Peewee ene Comer mer eerereessees

Red Kangaroo (Macropus rufus)

Corron, W. B.

Exhibition of heads and horns of various species of

Gazelles from the Bastern Sudan

Ahern pre eeeeee peer eeee ae

Page

Ag)

419

an Page

Cummines, Bruce F.

Distant Orientation in Amphibia. (Text-figs. 4 & 5.) 8

Cunnincuam, Josep T., M.A., F.Z.8,

Mendelian Experiments on Fowls .........-:101seeeeeeees 241

Dotuman, Guy, B.A.

A new Snub-nosed Monkey ........-..0cceeeereeeereeneeeees 503

Ducmorg, A. Rapciyrre, F.R.G.S.

Lantern exhibition of photographic studies of wild

animals in British East Africa and Newfoundland ...... 498

DunBAR-BRUNYTON, JAMES.

Exhibition of skins and skull of Bushbuck from

N.E. Rhodesia

GerrRARD, Messrs. E., & Sons.

Exhibition of the head of a cow Sable Antelope, the

skulls of an Isabelline Bear,a Leopard, and a Lion, all of

which showed some abnormality .......- ees yee dics ines 498

Gunn, Tuomas E., F.L.S.

On the Presence of two Ovaries in certain British

Birds, more especially the Falconide. (Pls. I1.-V.)...... 63

Hawkins, Herpert L., M.Sc., F.G.8., Lecturer in Geology,

University College, Reading.

The Classification, Morphology, and Evolution of the

Echinoidea Holectypoida. (Text-figs. 54-60.) ......c00 440

X11

Iuus, Avueustus D., D.Sc., B.A., Forest Zoologist to the

Government of India; late Professor of Biology,

Muir College, and Fellow of the University of

Allahabad.

On some Collembola from India, Burma, and Ceylon ;

with a Catalogue of the Oriental Species of the Order,

(Pls. VI.-XTI. and Text-figs 14 & 15.)

Jameson, H. Lysrer, M.A., D.Se., Phas EeZes.

Studies on Pearl-Oysters and Pearls.—I. The Structure

of the Shell and Pearls of the Ceylon Pearl-Oyster

(Margaritifera vulgaris Schumacher): with an Examin-

ation of the Cestode Theory of Pearl-Production. (Pls.

XXXIII.-XLVII. and Text-figs. 33-41.)

Pee erence ne recrercce

Kew, H. Wautis, F.Z.S.

On the Pairing of Pseudoscorpiones. (Text-figs. 47—

50.)

Te PRB OD GBD DO OF OS OUT C OOOO NOOO COA OGCOOCOoU mo aoa annOdgonG AD

LyDEKKER, RicHarp.

On the Milk-Dentition of the Ratel. (Text-figs. 31 &

B25) eee

nigiak te bcarhc ich eke wheieieleteleseiejeislele e's lejelaialerele see lac, atelis;sfetalersccteteheretete eter clot etaesten tenets

MasterMAN, Arruur T., M.A., D.Sc., F.Z.S.

Demonstration of recent investigations on Age-

determination in the Scales of Salmonoids

Coe ee meee etre scene

Meabze-WaA.po, E. G. B., V.P.Z.S.

Exhibition of a pale fulvous variety of the Common

Polecat: (Patorius putorius)

PROED eee ree err ere se eetreeseceereeree

Page

260

221

390

Page

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

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

Report on the Additions to the Society’s Menagerie

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

1911

Report on the Additions to the Society’s Menagerie

during the month of October 1911

Report on the Additions to the Society’s Menagerie

during the months of November and December 1911

BOC AMON LOU) 5... 3.2.32 cc ements Mas coee sate enon st 231

PicKARD-CAMBRIDGE, The Rey. Octavius, M.A., F.R.S.,

C.M.Z.S.

A Contribution towards the Knowledge of the Spiders

and other Arachnids of Switzerland. (Text-figs. 51 & 52.) 393

PurMer, Henry G., F.R.S., F.L.8., F.Z.S., Pres. R.M.S.,

Pathologist to the Society.

Report on the Deaths which occurred in the Zoological

Gardens dueing U9 iti. a, ate Cees oh. ahhh on. Sidhe cask 235

On the Blood-parasites found in Animals in the

Zoological Gardens during the four years 1908-1911.

ee XEN <)) 2 uss o> Se Ee a oes «a vdks 406

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

of the Gardens and Curator of Mammals.

Exhibition of photographs of a new race of Red-fronted

Gazelle (Gazella rufifrons hasleri). (Text-figs. 2& 3.) ... 5

On the Moulting of an Arctic Fox (Vulpes lagopus) in

the Society’s Gardens. (Text-figs, 6-13.) ...........ccanes DD

X1V

f Page

Pocock, Rreeraxp, I., F.R.S., &e. (Continwed.)

Exhibition of living specimens of the Common Ele-

phant-Shrew (Macroscelides proboscideus) and the Rock

Hlephant-Shrew (Hlephantulus rupestris). (Text-figs.

UG ree MUTE tas Or cere been eects. tality acae caine des. 5's 1 ae eee 142

Preston, HucutB., F.Z.8.

Diagnoses of new Species of Terrestrial and Fluviatile

Shells from British and German East Africa, with the

Description of a new Genus (Hussoia) from the Husso

Nyiro River, B. EH. Africa. (Pls. XXXI.& XXXII.)... 183

Sars, Prof. Grorce O., C.M.Z.8.

Zoological Results of the Third Tanganyika Expe-

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

1905.—Report on some larval and young stages of

Prawns from Lake Tanganyika. (Pls. LVII.-LX.) ...... 426

Seru-Smira, Davip, F.Z.S., M.B.O.U., Curator of Birds

and Inspector of Works.

Exhibition of a spirit-specimen of a nestling Australian

Regent-Bird (Sericulus melinus). (Text-fig. 1.) ............ 4

On the Moulting of the King Penguin (Aptenodytes

pennants) in the Society's Gardens. (PI. 1.) ...0..c.. 60

SHELFORD, Rospert, M.A., F.Z.S.

Mimicry amongst the Blattide; with a Revision of

the Genus Prosoplecta Sauss., and the Description of a

new Genus. (Pl. XLVIII. and Text-figs. 42-46.) ...... 358

SairH, D. SerH-. See Seru-Smirn, D.

Smira, Georrrny, M.A., Fellow of New College, Oxford.

The Freshwater Crayfishes of Australia. (Pls. XIV.—

POG V lle enalMoxt snes). le ee ee ene cae 144

Ey,

STEBBING, Epwarp P., F.L.S., F.Z.S., F.R.G.S.

Game Sanctuaries and Game Protection in India ......

SusHKIN, Prof. Perer P., C.M.Z.8., Kharkov, Russia.

Ontogenetical Transformations of the Bill in the Heron

(Ardea cinerea). (Pl. XITI.)...... ook Bete See See ar eae

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

The Duke of Bedford’s Zoological Exploration of

Eastern Asia.—XV. On Mammals from the Provinces

of Sze-chwan and Yunnan, Western China..................

The Races of the European Wild Swine ..................

Two new Genera and a new Species of Viverrine

eranivoie.,\, (hext-les, 61—Odalamgr em amete sss -0h.e.. faces

Wanpo, E. G. B. Meaps-. See Mrape-WAxpo, E. G. B.

Page

—

iS)

i |

NEW GENERIC TERMS

PROPOSED IN THE PRESENT VOLUME (pp. 1-504).

Page © Page

Chrotogale (Mammalia)......... 499 | Idiomerus (Collembola) ...... 113

Conulopsis (Echinoderma) ... 453 | Megapygus (Echinoderma; ... 449

Dicranocentroides (Collembola) 102 | Melyroidea (Orthoptera) ...... 374

Diplogale (Mammalia) ......... 499 | Otiditeenia (Vermes) ............ 220

Eussoia (Mollusca) ............ 192 | Paracheraps (Crustacea) ...... 161

Heteromuricus (Collembola)... 92 | Pseudocyphoderus (Collembola) 116

SCIENTIFIC

Accipiter

nisus, 67, 70-74, 79.

Achorutes

armatus, 81, 85, 119.

crassus, 119.

hirtellus, 119.

lipaspis, 119.

Achroblatta

luteola, 361.

Acomys

cahirinus, 229.

russatus, 224, 229,

230.

¥ we yptiacus,

230.

— 224,

sellysit, * 224,

Acrobothrium, 274, 279.

Adelopneustes

lamberti, 458.

/Elureedus

melanocephalus, 408.

/élurops

v-insignitus, 402.

Aétobatis

narinari, 274, 283,

284, 289, 291, 358.

Africarion

kemp?, 184, 193.

microgranulata, 184,

microstriata, 184,

193.

Agapornis

taranta, 238.

Agelena |

labyrinthica, 396.

Alza |

keniana, 189, 193.

Algiroides

nigropunctatus, 412.

Allacta, 367. |

Alligator

mississippiensis, 415, |

sinensis, 233. |

Proc. Zoou. Soc.—1912,

| Anomotvnia, 219, 220,

| Anoplotznia, 208, 209,

| Anthus

INDEX

Amadina

Jasciata, 416.

Amaurobius

Senestralis, 396.

ferox, 396.

4-guttatus, 396.

Amblyornis

subalaris, 411, 416.

Amblypygus, 453, 454,

470, 491, 493.

| Amphibolurus

barbatus, 410.

Amydrus

moriv, 233.

Anamesia, 359.

Anaplecta, 367.

decipiens, 365.

vartipennis, 366.

Anas

sparsa, 1.

Anchitherium, 221.

| Ancylus

kempt, 190, 193.

Andigena

bailloni, 409.

| Anisolemnia

distaura, 376.

Anodonta, 301, 316.

isacantha, 210.

pen icillata, 210.

210, 221.

Anorthopygus, 443-445, |

447, 458-467, 473- |

475, 478-480, 486,

490, 493.

orbicularis, 449, 456,

467, 491.

Anoura, 85. |

Anourosorex |

squamipes, 134.

maculatus, 5. |

No. XXXII.

NAMES.

Anthus

rosaceus, 3.

Anura, 85.

Aphelocoma

ultramarina, 408.

Apodemus

agrarius, 135,

chevrieri, 135, 136.

— fergussoni, 135,

136

draca, 137.

JSergussoni, 135, 136.

orestes, 137.

peninsule, 137.

speciosus, 136, 137.

— chevrieri, 136.

— latronum, 137.

— orestes, 136, 137.

— peninsule, 136.

— peninsularis, 137.

Aptenodytes

pennanti, GO.

Aptera

Jusca, 360.

Ara

macao, 415.

Arzoncus

altissimus, 398.

Arctonyx

obscurus, 134.

Ardea

cinerea, 125, 126.

| Areolaria, 361.

bipunctata, 369.

Asagena

phalerata, 397.

Asio

accipitrinus, 3, 73.

| Aspergillus

niger, 239.

Assimania

aurifera, 191, 193.

Astacoides, 148.

bicarinatus, 163.

XVill

Astacoides

plebeius, 157.

serratus, 157.

spinifer, 157.

Astaconephrops

albertisti, 167, 168.

Astacopsis, 150, 151,

1153},

australiensis, 159, 160.

bicarinatus, 163.

Ffranklinii, 145, 146,

148, 152, 154, 156,

160, 161, 170, 171.

— tasmanicus, 146,

149.

kershawi, 146, 149,

152, 160, 161, 170.

nobilis, 157, 159.

paramattensis,

159.

serratus, 145, 146, 147,

149, 152, 157, 159,

160, 161, 170, 171.

— yarraensis, 152.

spinifer, 157, 159.

sydneyensis, 157, 159.

tasmanicus, 156.

Astacus, 144, 488.

armatus, 157.

australiensis, 157.

bicarinatus, 163.

Asterostoma, 448.

Ateles

geoffroyt, 407.

paniscus, 3.

Atemnus, 388.

Atheris

chlorechis, 414.

Attulus

histrio, 402.

Attus

pubescens, 402.

rupicola, 402.

Aulacocheilus, 372.

Aulonia

albimana, 401.

Avitellina, 208.

centripunctata,

216.

157

}

207,

Bacillus

gallicus, 172.

granuatus. 72.

rossti, 172-182.

Balzeniceps, 126.

Balanoglossus, 281.

Balearica

regulorum, 415.

Balistes, 261, 286, 287,

288.

Bathyphantes

concolor, 397.

variegatus, 397.

Bauria, 423.

Bertiella

cercopitheci, 195.

Bitis

arietans, 413.

Blarinella

quadraticauda, 134.

Blatta

germaniea, 181.

Blattella

germanica, 304.

Blauneria

exsilium, 189, 193.

Boa

constrictor, 413.

madagascariensis, 414.

Bolyphantes

alticeps, 397.

Bos

gaurus, 30.

Bubo

capensis, 416.

maculosa, 417.

poensis, 417.

Bufo

calamita, 19.

marinus, 410, 413.

regularis, 414.

vulgaris, 9, 19.

Buteo

vulgaris, 7.

Caccabis

chukar, 41.

Calabaria

reinhardtt, 230.

Calandrelia

acutirostris, 3.

Callilepis

nocturna, 39D.

Calliope

camtschatkensis, 2.

Calliptilus

solitarius, 232.

Calliste

cayana, 409.

cyanoptera, 408, 416.

Jestiva, 416.

Hlaviventris, 409.

guttata, 408.

melanonoia, 416,

thoracica, 415.

tricolor, 416.

Caloblatta, 361.

Calopelia

puella, 416.

Cambarus, 144.

INDEX OF SCIENTIFIC NAMES.

Cam pylothorax, 108.

Cancer

serratus, 157.

Canis

zerda, 225.

Capricornis

milne-edwardsi, 141.

Caratomus, 448, 453,

492, 497.

Cardinalis

pheniceus, 409.

Caridella

cunningtont, 430, 439.

Caridina

nilotica, 427, 430, 431.

— gracilipes, 426.

wyckti, 426.

Carpodacus

meaxicanus, 415.

Carpophaga

concinna, 415.

Caryomys, 139, 140.

Cassiculus

melanicterus, 416.

Cassida, 362.

Cassidodes

ligata, 371.

Cebus

albifrons, 407.

Jatuellus, 3.

hypoleucus, 233.

Centromerus

affinis, 397.

subalpinus, 397.

Cephalobothrium, 285.

abruptum, 284.

aétobatidis, 284, 358.

variabile, 284.

Cephalophus

coronatus, 232.

Ceratameria

(Schottella) maxiina,

119.

Ceratinella

brevis, 398.

Ceratinoptera, 367.

Cercidia

prominens, 399.

Cercopithecus

sabeus, 410, 415.

tantalus budgett?, 142.

Ceryvulus

muntjac, 3d.

Cervus

axis, d2.

duvaucelt, 35.

eldi, 41.

kashineerianus, 232

porcinus, 32.

unicolor, 26.

xanthopygius, 142.

Cheraps, 145, 148, 151,

152, 153, 160, 162,

169.

arwanus, 168.

bicarinatus, 163.

intermedius, 150, 165,

168, 171.

lorentzi, 168.

preissii, 263.

quadricarinatus, 158,

167, 168, 171.

— aruanus, 167.

— lorentzi, 1@7.

quinguecarinatus, 165,

171.

tenuimanus, 166, 170,

Chameleon

senegalensis, 418.

Chapmannia

lapica, 195.

Cheiracanthium

ttalicum, 396.

Cheiracanthus

uncinatus, 266.

Chelifer, 388, 390.

cimicoides, 377.

cyrneus, 377, 381, 382,

384, 486, 387.

latreillii, 377, 880, 382,

385, 386.

multidentatus,

Chernes, 377, 382, 387,

388, 390.

Chironomus, 176.

Choanotznia, 217, 219.

galbuia, 202.

infundibulune,

220.

levigata, 220,

Chodsigoa

berezowskii, 133.

hypsihia, 133, 134.

larvarum, 183.

Cholcepus

hoffmanni, 1.

Chorisoneura, 367.

teniata, 360.

Chrotogale, gen. neyv.,

387,

202,

owstont, 500, 501, 502.

Chrysemys

ptota, 415, 415.

Chrysomitris

cucullatus, 416.

tibetana, 2.

Chrysomma

sinense, 409.

Chrysotis

vantholora, 233.

INDEX OF SCIENTIFIC NAMES,

Chthonius, 381, 388.

tetrachelatus, 403.

Cicinnurus

regius, 408, 416.

Cieurina

cinerea, 396.

Cidaris, 441, 46].

Cinixys

belliana, 413,

Circaétus

gallicus, 73,

Circus

eruginosus, 72, '73, 76,

79.

cineraceus,

79.

cyaneus, 71, 72, 73, 76,

79.

LOT G:

macrurys, 73.

Gistudo

carolina, 411.

Cittoecinela

macrura, 409,

Clemmys

guttata, 414.

leprosa, 413.

Clione, 266.

Clubiona

hilaris, 396.

Clypeaster, 445,

457, 462, 479,

492, 493, 494.

Clypeus, 464.

Coccothraustes

melanura, 415,

Ceculus

echinipes, 403.

Coelophora

formosa, 373, 376.

Ceelotes

atropus, 396.

claustrarius, 396.

pickardii, 396.

terrestris, 396.

Ceenholecty pus, 447, 451,

455, 464, 466, 467,

472, 475, 479, 480,

486, 487, 493.

Jjullient, 467.

macropygus, 430.

Coereba

cyanea,

417.

Colinus

pectoralis, 3.

Gvlobus

matschiei, 142.

Coluber

corats, 412.

guttatus, 413.

leopardinus, 41 1.

409, 416,

Xix

Coluber

longissimus, 413.

nelanaleucus, 413.

ohsaletus, 413.

vulpinus, 2.

Columba

gymnophthalma, 409.

Conoclypeus, 445, 453,

456, 460, 462, 468,

469, 492, 493, 494.

Conylopsis, gen. nov.,

453, 469, 491, 492,

493.

roemert, 453, 495.

Gonulus, 447, 454, 457-

460, 464-480, 487,

490-4938, 497.

alhogalerus, 451, 455,

462, 48].

subconious, 482.

suhratundus, 456, 461,

462, 486.

Copsychus

sauaris, 409, 411,

416.

Coptodiscus, 447, 467.

472, 486, 493.

neme, 450.

Coracias

indica, 417.

Corallus

cooki, 414,

Covronella

getula, 412.

Corvus, 70.

Srugilegus, 73.

Corydia, 361.

petiverana, 362.

Corynephoria

Jjacabsoni, 121.

Cosmozasteria

lateralis, 359.

zonata, 359,

Crateropus

platycercus, 3.

Cremastocephalus

eelebensis, 120.

indicus, 81, 104, 105,

106, 120, 124.

montanus, 81, 105, 106,

120, 124.

Crocidura

attenuata, 184,

— (Crocidura) religi-

osa, 225,

Cratalus

atrox, 412.

confluentus, 414,

horridus, 413,

Crypheeea

silpicola, 396,

xx INDEX OF SCIENTIFIC NAMES.

Curzeus

aterrimus, 409,

Cyanocitta

stellert azteca, 232.

Cyanocorax

luxuosus, 409.

Cyanospiza

leclancheri,

415.

Cyathocephalus, 274, 280.

Cyathophyllus, 274.

Cyclemys

trifasciata, 411.

Cyclosa

conica, 398.

Cyclura

acanthura, 410, 418.

Cygnus

bewicki, 72, '78, 79.

Cylindrophorus, 199.

Cynezlurus

jubatus, 2.

Cyuips

kollari, 174.

Cynixis

belliana, 415.

erosa, 416.

homeana, 415.

Cynognathus, 4238, 424, |

425,

Cyornis

rubeculoides, 2.

Cyphoderodes, 116.

ceylonicus, 121.

Cyphoderus

albinus, 115, 116.

javanus, 121.

simulans, 81, 115, 116, |

121, 125.

Cyrtomorphus, 372.

Dacnis

cayana, 415,

Damonia

reevest, 413.

Davainea, 216.

comitatus, 218.

frontina, 218,

lateralis, 220.

rhynchota, 218.

Degeeria, 99.

Delphinognathus, 428.

Dendraspis

viridis, 233, 414.

Dendreeca

coronata, 142,

Dendryphantes

encarpatus, 402.

Deropeltis

dichroa, 360.

142, 409,

| Dicranocentroides,

Deropeltis

erythrocephala, 360.

paulinoi, 360.

Desorella, 448, 454, 490.

| Desoria, 443.

;

485,

Diademopsis, 484,

Dizea

dorsata, 399.

Dichoceros

bicornis, 238.

gen.

noy., 82, 102, 121.

Ffasciculatus, 81, 102,

120, 128,124.

| Dictyna

flavescens, 396.

uncinata, 396.

viridissima, 396.

| Dicynodon, 419, 424.

latifrons, 420.

Didelphys

nudicaudata, 407.

| Dilepis, 218.

| Diplocephalus

castaneipes, 398.

eristatus, 398.

eborodunensis, 398.

Rochii, 398.

Diplogale, gen.

499.

hosei, 499.

Diploptera

dytiscoides, 361, 366.

Dipeena

braccata, 3897.

Dipsas

plicatus, 317.

Dipus

gerbillus, 226.

Discoholectypus,

487, 491, 493.

meslei, 451.

noy.,

470,

| Discoidea, 443-447, 454,

458-460, 462, 464,

466, 468-471, 474—

77, 479-481, 487,

490, 492-494,

cylindrica, 455-457,

461, 496, 497.

subucula, 451, 452, 472,

478, 497.

Distomum

margaritifactor, 274.

Dolomedes

jimbriatus, 400.

Drapetisca

socialis, 397.

Drassodes

lapidosus, 395.

pubescens, 395.

Drassus

hispanus, 395.

troglodytes, 395.

Drymobius

boddaerti, 418.

Dryotriorchis

spectabilis, 409.

Dymecodon, 130, 131.

Echinanthus, 445.

Hehinites, 452, 477, 494.

Echinobrissus, 489.

Hehinocardium, 441.

Echinoconus, 4438, 445,

453, 495.

abbreviatus, 492.

conicus, 496.

orbignyanus, 492.

| Echinocorys, 478.

| Hchinocyamus, 477, 478,

493, 494.

Echinolampas, 454, 474,

493.

Echinoneus, 448, 444,

452, 459, 462, 467,

490, 497.

Echinus, 474, 476.

Kelectus

pectoralis, 416.

Ectobius, 365, 366.

Elaphodus

cephalophus, 141.

Elephantulus

rupestris, 142, 143,

144.

Emberiza

elegans, 2.

Fucata, 3.

leucocephala, 3.

Emys

lutaria, 411.

orbicularis, 4138.

Engeus, 145, 148, 15],

152, 153, 160, 162,

165.

Eniochobothrium

gracile, 284.

Ennea

microstriata, 183, 193.

Enoploenatha

thoracica, 397.

Entomobrya

anomala, 97.

crassa, 81, 96, 128.

florensis, 120.

halt, 8, 95, 120) 1123:

— lutea, 81, 96.

longicornis, 120.

| Entomyza

cyanotis, 408.

Eos

fuscata, 2.

Eothenomys, 159.

Epeira

adiana, 399.

agalena, 399.

alpica, 399.

earbonaria, 399.

ceropegia, 399.

circe, 399.

cucurbitina, 399.

diademata, 399.

dioidea, 399.

marmorea, 399.

montana, 399.

nordmannii, 399.

pyramidata, 399.

quadrata, 309.

redti, 399.

sclopetaria, 399.

sturmiti, 399.

triguttata, 399.

Epiblemum

scenicum, 402.

Epimys

confucianus, 135.

humiliatus, 135.

norvegicus, 135, 411.

Episinus

truncatus, 396.

Equus

kiang, 2.

Erigone

atra, 398.

dentipalpis, 398.

Jugorum, 398.

remota, 398.

tirolensis, 398.

Erythrura

pealet, 2, 142.

prasina, 499, 416.

psittacea, 416.

Eryx

johni, 414.

Estrelda

melpoda, 408, +11, 415.

phenicotis, 408.

Eumorphus, 362.

Euophrys

ae 402.

Frontalis, 402.

petrensis, 402.

pictilis, 402.

Eupatagus, 475.

Euphonia .

laniirostris, 409.

violacea, 408.

Euplectes

franciscana, 409.

oryx, 416.

Eupleres, 499, 502.

INDEX OF SCIENTIFIC NAMES,

Eupodotis

kori, 194, 220, 221.

Eupsychortyx

nigrigularis, 232.

Eurycotis

floridana, 859, 360.

Euryopis

flavomaculata, 597.

Eussoia, gen. nov., 183,

192.

inopina, 192, 193.

| Eustegasta

buprestoides, 361, 362.

Huzoéa, 427.

Euzosteria

mitchelli, 359.

Falco, 70.

esalon, 73.

cenchris, 73.

peregrinus, 72, 77, 79.

sudbuteo, 72, 73, 77, 79.

tinnunculus, 71, 72, 73,

COD

vespertinus, 73.

| Felis

leo, 498.

nebulosa, 3, 224.

onca, 224.

pardalis, 3, 224.

pardus, 224, 498.

sylvestris, 232.

temmincki, 407.

tristis, 224.

Forficula, 178.

Francolinus

gariepensis, 408, 411.

levaillanti, 409, 411,

417.

Fuligula

baeri, 416.

marila, 417.

Gadopsis

marmoratus, 156:

Galago

crassicaudata, 233.

Galeopithecus, 220.

Galeoscoptes

carolinensis, 409.

| Galerites, 443, 445, 491.

albogalerus, 496.

roemeri, 453, 492.

Galeropygus, 443, 445,

452, 454, 457, 464,

465, 469, 4738, 474,

489.

agariciformis, 455.

dumortier?, 488.

Galictis, 222, 223, 224.

XXx1

' Gallinula

| chloropus, 217.

| Gallus

bankiva, 241, 251, 252,

258.

| Garrulax

albogularis, 409.

leucolophus, 408, 415,

| Garrulus

lanceolatus, 407,

Gazella

dorcas, 4.

isabella, 4.

rujifrons, 4, 5, 6.

— hasleri, 5, 6.

subgutturosa, 3.

tilonura, 4.

Genetta

poénsis, 142.

tigrina, 233.

Geocichla

citrina, 408.

Geopelia

striata, 410.

Gerbillus

gerbillus, 226.

pyramidum, 225,

sellysit, 226.

Gerrhosaurus

nigro-lineatus, 1.

validus, 1.

Ginglymostoma, 282,

285

concolor, 287, 288,

289,

Globator, 4438.

Glugea, 272.

Gnaphosa

badia, 395.

molesta, 395.

petrobia, 395.

tigrina, 399.

Gonatodes

vittatus, 3.

Gongylidium

apicatum, 398.

Goniocidaris, 472.

Gracula

religiosa, 415.

Grus

Jjaponensis, 417,

— monachus, 2.

Gubernatrix

cristata, 408.

Gymnophallus, 327,

Gymnorhina

leuconota, 410, 416.

tibicen, 408.

Hamoceystidium, 415,

Hemoproteus

danilewskyi, 240, 414, |

415, 416, 419.

Haplodiseus, 211.

Harpactes

drassoides, 395.

Hedymeles

ludovicianus;

416.

Heliophanus

@neus, 402.

metallicus, 402.

Helix

zanguebarica, 186.

Hemigale

hosei, 498, 499.

Hemigalus, 499, 503.

hardwickei, 498, 580,

o01, 502.

Hemipedina, 482,

bonei, 484, 485.

Wemithyrsocera

sp., 364, 367.

Heuiitragus

hylocrius, 26:

Hemixus

flavala, 409:

Herpestes

albicauda, 225:

Hestia, 368.

Heteraster, 469.

Heterodon

stmus, 411, 413.

Heteromuricus, gen, nov.,

Sil BEA UAE

cercifer, 81, 92, 120,

TPA, eB}.

Heteromurus

teneicornis, 120.

tetracantha, 120.

(Lempletonin) sp.,

120.

409;

Hexaimitus, 411.

Hilaira

montigena, 397,

Hipposideros

armiger, 128,

Hippotragus

niger, 498.

Holaster, 480,

Holectypus, 448, 445,

447, 454-463, 465,

468, 470, 472, 475

476, 480, 481, 486,

487, 498.

depressus, 450, 455,

463, 464, 466, 471,

478-475.

hemisphericus, 463,

464, 466, 475.

oblongus, 456,

INDEX OF SCIENTIFIC NAMES.

Holectypus

sarthacensis, 4°77.

Holocompsa, 862.

Homalenotus

quadridentatus, 402.

Homopteroidea

nigra, 362.

Homorus

iredalei, 189, 193.

Houbara

macqueent, 416.

Hysena

crocuta, 232:

Hyboclypus, 443, 444,

446.

Hydraspis

hilarii, 414,

Hyla

arborea, 411.

goughi, 2.

rubra, 2.

venulosa, 2, 410, 411.

Hymenolepis

ambiguus, 195.

tetracis, 195.

villosa, 198.

Hypercompsa

Jieberi, 362.

Hyphantornis

spilonotus, 416.

Hypnorna, 361.

amena, 362.

Hyptiotes

paradorus, B98.

Icterus

baltimore, 409.

gularis, 400.

Jamuicat, 408, 415.

Idiogenes

otidis, 195.

Idiomerus, gen. noyv., 82,

IIS}, PAT

pallidus, 81, 114, 121,

124.

Iguana

tuberculata, 414.

Isotoma

crassicornis, 119.

nigropunctata, 81, 90,

119, 122.

palustris, 9B.

quadrioculata, 91.

sita, 81, 89, 122.

Isotomurus

palustris, 81, 98, 120,

22,

— tricuspis, 120.

Ithagenes

cruentus, 2.

Kaliella

consobrina,

193.

depauperata, 186, 193.

iredalei, 187, 193.

higeziensis, 187, 193.

Karyolysus, 412.

Kistecephalus

chelydroides, 422.

Kystocephalus

translucens, 284, 291.

186, 187,

Lacerta

ocellata, 414.

peloponnesiaca, 414,

Lachesis

lanceolatus, 412.

Lagonosticta

senegala, 409.

Lagopus

scoticus, 72, 77, 79.

Lam protornis

enews, 415.

Lanieria, 447, 487,

498.

lanier?, 450.

Lanius

collurio, 401.

Larus

minutus, 72,78; 79.

Leis

dunlopi, 376.

Leistes

gwianensis, 411.

Lemur

coronatus, 407.

mungos, 407.

Leontocebus

edipus, 407.

rosalia, 407:

Lepidocyrtus

braveri, 120.

gavanicus, 120,

javanus, 120.

maxtinus, 94.

robustus, 81, 84, 120.

variabilis, 120.

Leptophis

liocercus, 418.

Leptyphantes

expuncta, 397.

Sragilis, 397.

leprosus, 397.

mengit, 397:

minutus, 397.

pulcher, 397.

tenebricola, 397.

tenuis, 397.

Lepus

egyptius, 231.

sinaiticus, 280.

INDEX OF SCIENTIFIC NAMES.

Leucochiloides

chanlerensis, 188, 1958.

gaziensis, 188, 193.

tredalei, 188, 193.

soror, 188, 198.

Leucodore, 266, 315.

Limnea

kempi, 190, 193.

Limnocaridina

parvula, 452, 485,

440.

spinipes, 427, 459.

Linyphia

emphana, 397.

JSrutetorum, 397.

hortensis, 397.

marginata, 397.

montana, 397.

phrygiana, 597.

pusilla, 397.

triangularis, 397.

Liobunum

limbatum, 402.

rotundum, 402.

Lithyphantes

corollatus, 397.

paykullianus, 397.

Locusta, 178.

Lophocarenum

nemorale, 398.

stramineum, 398.

Lophophaps

leucogaster, 408.

Lophorhina

minor, &,

Lophortyx

douglasi, 3.

Lorieculus

galgqulus, 416.

Lorius

domicella, 417.

Lycaon

pictus sharicus, 142.

— — somalicus,

232,

Lycosa

accentuata, 401.

agrestis, 401,

albata, 401.

amentata, 401.

bifasciata, 401.

blanda, 401.

cursoria, 401.

JSerruginea, 401.

giebeli, 401.

ludovica, 401.

lugubris, 401.

miata, 401.

monticola, 401,

nigra, 401.

paludicola, 401.

| Lycosa

palustris, 401.

pedestris, 401.

pulverulenta, 401.

riparia, 401.

tarsalis, 401.

torrentum, 401.

wagleri, 401.

Lynchia

maura, 416.

Macaea, 128.

Machlolophus

xanthogenys, 2, 417.

Macropus

bennetti, 232.

rufus, 234, 235.

Macropygus, 447, 452,

463, 464, 466, 475,

485, 493.

macrocyphus, 468.

morrisit, 479.

truncatus, 449, 488.

Macroscelides

proboscideus, 142, 148,

144.

Macrotoma

vulgaris, 91,

Manucodia

atra, 409.

Margaritana, 500, 312,

322, 327.

Margaritifera, 303, 304,

316, 323.

margaritifera, 310,

811, 318, 347, 351.

— cumingti, 260, 274,

275, 299, 348.

— mazatlanica, 339.

martensii, 317.

maxima, 261, 295, 305,

306, 307, 318, 347.

vulgaris, 260, 261, 262,

268, 273, 274, 277,

278, 291, 292, 293,

295, 296, 298, 302,

306-309, 311, 314,

327, 328, 329, 331,

332, 337-341, 345-

349, 352-355.

Megabunus

rhinoceros, 402.

Megalema

asiatica, 409.

Megapyga

eximia, 376.

Megapygus, gen. nov.,

447, 449, 452, 457,

464, 466, 469, 479,

480, 486, 490, 493.

XX1l

Megapygus

umbrella,

495.

Melanerpes

uropygialis, 3.

Meleagrina

margaritifera, 295.

Mellivora, 221-224.

Melospiza

georgiana, 232.

pusilla, 409.

Melyroidea, gen. nov.,

O74.

magnifica, 375, 376.

449, 485,

mimetica, 874, 379,

376.

Meriones

crassus, 224, 226, 227,

228.

— pallidus, 226, 227.

— sellysii, 226, 227.

melanurus, 227.

sellysti, 224.

shawi, 226, 227.

— melanurus, 227.

Merula

albocincta, 409.

castanea, 2.

tamaulipensis, 3.

Merychippus, 221.

Mesira, 100.

Meta

menardi, 398.

meriane, 398.

segmentata, 398,

Micaria

breviuscula, 395.

hospes, 399.

pulicaria, 395.

scenica, 395.

Micrommata

virescens, 400.

Micromys

pygmeus, 137.

Microneta

rurestris, 398.

viaria, 398.

Microtus

agrestis, 159.

eva, 140, 141.

irene, 138, 139.

melanogaster, 159, 140.

millicens, 138.

sikimensis, 159.

(Caryomys) alcinous,

140.

(—) eva, 140, 141.

(Eothenomys) smelano-

gaster, 140.

(—) — eleusis, 139.

(—) olitor, 139.

’

XX1V

Micryphantes

gulosa, 397.

Mimeusemia

ceylonica, 863.

Mimocichla

rubripes, 408.

Misumena

vatia, 399.

Modiola, 301.

Molge

cristata, 8, 10.

palmata, 8.

Molothrus

pecoris, 408.

Monieza, 207.

Monopylidium, 218, 219,

220.

cayennense, 217.

infundibulum,

LNG.

macracanthum, 217.

marchali, 217.

rostellatum, 217.

secundum, 217.

Moschops, 423.

Moschus

sifanicus, 141.

Motacilla

feldeggi, 2.

Mungos

albicauda, 225.

Mus

alexandrinus, 7.

chevriert, 135.

gentilis, 228.

musculus, 228.

— gentilis, 228.

ouang-thome, 135.

plumbeus, 135.

rattus, 6, 7, 228.

— alexandrinus,

228.

— tectorwin, 6, 7, 228.

rufifrons, 6.

tectorum, 7, 228.

Muttua

margaritifere, 327.

Myiophoneus

temmincki, 407.

Myiozetetes

similis, 3.

My liobatis

maculata, 284.

Myrmecophaga

tridactyla, 407.

Mysis, 426.

Mytilus, 266,

275-278,

300, 301,

316, 324,

348.

216,

270, 272,

293, 294,

306, 308,

336, 347,

Mytilus

edulis, 267, 307, 327, |

302.

Myzantha

garrula, 408.

Nemorhedus

cinereus, 141.

Naia

tripudians, 411, 413.

Nasillus

gracilis, 129,

Natalina

permembranacea, 188,

193

Neanura

corallina, 81, 85, 87,

88, 119, 1122:

fortis, 119.

mtermedia, 81, 87, 88,

1P9) 1122:

pudibunda,

cela}, IIS), Tey

Nemastoma

dentipalpe, 403.

quadripunctatum, 403.

Nematogmus

sanguinclentus, 398.

Nesoceleus

Sernandine, 232.

Nettium

torquatum, 142.

Neurotrichus, 131.

Nucifraga

caryocatactes, 415.

Nucleolites, 489, 493.

Nucleopygus, 443, 490.

Nyctalus

labiatus, 129.

plancyt, 129,

Obisium, 388.

qugorum, 403.

Odontophorus

capueira, 410,

Cidemia

nigra, 409.

Oides

biplagiata, 300, 376. |

Oligolophus

morio, 402.

— alpinus, 402.

— glacialis, 402.

— palliata, 402.

Onychiurus

(Lipura) fimetarius,119. }

Onychogale

frenata, 407.

Oochoristica, 217,

81, 86, 87, |

INDEX OF SCIENTIFIC NAMES.

Ophiosaurus

apus, 414.

Orchesella, 113.

Oreoneta

fortunata, 397.

Oreospiza

chlorura, 409.

Orthopsis, 471.

Ostrea, 301.

Otaria

pusilla, 2.

Otiditenia, gen. nov.,

196, 220.

eupodotidis, 194-221.

Otoeyon

megalotis, 232.

Oudemansia

cerulea, 119.

Oudenodon

kolbei, 422, 425.

Oviclypeus, 453.

Pachnepteryx, 361.

Pachyclypeus, 443, 445.

Pachygnatha

degeerti, 398.

Pachyrhynchus, 368.

Palemontes

varians, 438.

Palinurus, 145.

Paracheraps, gen. nov.,

145, 148, 151, 16n,

165, 168.

bicarinatus, 145, 150,

152, 153, 162, 163,

169, 170, 171.

Paradisea

raggiana, 408.

Paranephrops, 144, 167.

Parastacus, 144.

| Paratropes, 361

Pareiasaurus, 421.

Paronella

bornert, 81, 106, 114,

120, 124.

dahlii, 106.

gracilis, 81, 109, 120,

124.

insignis, 81, 112, 120,

124, 125.

phanolepis, 81,110, 124.

travancorica, 81, 108,

124,

Parotia

lawesi, 408, 417.

Passer

arcuatus, 417.

Patella, 476.

| Pecten, 301.

Pelecanus

roseus, 1.

INDEX OF SCIENTIFIC NAMES,

Pellenes

lapponicus, 402.

Pelmatosilpha, 359.

Periplaneta, 178.

Sortipes, 359.

orientalis, 181.

Perisphaeria, 363.

Phalangium

brevicorne, 402.

opilio, 402.

parietinum, 402.

Phasianus

Sormosanus, 3.

Phileus

chrysops, 402.

Philodromus

alpestris, 400.

collinus, 400,

emarginatus, 400.

laricium, 400.

lividus, 400.

margaritatus, 400,

Philodryas

schotti, 233.

Pholeus

phalangioides, 396.

Phonipara

canora, 415.

Phonygama

chalybeata, 408.

Phoraspis, 361.

Phyllodromia, 178.

Phyllonethis

lineata, 397.

Pileus, 443, 444, 447,

463-466, 469, 474,

476, 485, 486, 4938.

pileus, 449.

Pinna, 302, 308.

Pionopsittacus

pileatus, 408.

Pirata

latitans, 400,

Pisaura

mirabilis, 400.

Pitangus

derbianus, 3.

Pithecus, 128.

Pithodia, 494.

Pituophis

sayi, 410, 413.

Placuna, 273, 274.

placenta, 260, 299, 345,

348, 351.

Planispira, 185.

Planorbis

kigeziensis, 190,

193.

sperabilis, 190, 193.

Plasmodium

kocni, 414, 415.

| Plasmodiuin

precox, 240, 414, 415,

418.

Platybunus

pinetorum, 402.

triangularis, 402.

Platycercus

icterotis, 408.

Plectoptera, 366.

Plesictis, 225.

Plesiechinus, 448, 451. |

452.

Plotus

anhinga, 126.

Podicipes

cristatus, 72, 77, 79.

Podoenemis

expansa, 413.

Podura

armata, 8d.

palustris, 93.

Pceocephalus

fuscicollis, 416.

Polyzosteria

cuprea, 358.

limbata, 358.

nove-zealandiea, 3d9.

Porphyrio

madagascariensis, 416.

Porzana

pusilla, 2.

Pourtalesia, 496,

Pratincola

caprata, 2.

Presbytis, 503.

Prioptera

sinuata, 871, 376.

Pristes

cuspidatus, 284.

Procavia

sp., 231.

burtoni, 231.

ruficeps, 231.

Prosoplecta, 358, 363,

364, 367.

bipunctata, 363, 368,

369, 371, 376.

coccinella, 363, 3868,

369, 376.

celophoroides, 369, 373,

376.

gutticollis, 369, 372,

710.

ligata, 368, 369, 371,

72.

megaspila, 370.

mimas, 369, 373,

376.

nigra, 369, 372, 376.

nigroplagiata, 369, 370,

371, 376.

XXV

Prosoplecta

nigrovariegala, 366.

quadriplagiata, 369,

376.

rufa, 369, 371, 376.

semperi, 368, 369, 371,

376.

trifaria, 368, 370, 372,

376.

Prosthesima

apricorum, 399.

latretllii, 395.

nigrita, 39D.

petiverii, 399.

prefica, 395.

talpina, 399.

Protanura

krepelini, 119.

Protocyamus, 452, 478,

494.

Psammomys

obesus, 228.

Pseudachorutes

anomalus, 81, 88, 119,

122.

Pseudaspis

cana, 413.

Pseudocyphoderus, gen.

nov., 89, 116, 121.

annandalei, 81, 116,

121, 125.

Pseudoglomeris, 363.

Pseudoleistes

guirahuro, 238,

Pseudophyllodromia

parilis, 362.

Pseudosira, 99.

indra, 81, 100, 101,

120, 123.

Psittinus

incertus, 233.

Ptenothrix

gracilicornis, 121.

— gibbosa, 121.

Ptenura, 97.

Pteroglossus

torquatus, 408.

Pteroplatea

micrura, 284.

Ptilotis

Jusca, 416.

Ptyodactylus

lobatus, 414.

Putorius

putorius, 234,

Pycnonotus

Jocosus, 415,

| Pygaster, 443-449, 454,

457-466, 469-480,

484, 488-490, 493,

495.

XXV1

Pygaster

relictus, 452.

reynest, 483.

semisulcatus, 448, 451,

454, 455, 461, 468,

475, 482, 485, 488,

— conoideus, 456.

umbrella, 452,

(Macropygus)

noides, 455.

(—) truncatus, 454.

(Megapygus) macro-

cyphus, 458.

laga-

(—) wmbrella, 454,

466, 468.

Pygastrides, 452, 462.

Pygurus

blumenbachi, 474.

Pyrina, 443, 444, 457,

458, 462, 464, 466,

469, 470, 480, 490,

493,

desmoulinst, 489,

Pyrrhulopsis

taviunensis, 9,

Python

molurus, 412.

reticulatus, 413.

sebe, 411, 414.

spilotes, 4138,

Rallus

aquaticus,

Ramphastos

dicolorus, 409.

Rana

catesbiana, 411.

esculenta, 19, 410, 411.

temporaria, 19.

Rangifer

tarandus, 232.

Rhinoceros

wnicornis, 30.

Rhinolophus

acrotis brachygnathus,

225.

cornutus pumilus, 128.

ferrum-equinum, 128.

Rhinopithecus

avunculus, 503.

bieti, 503, 504.

roxellane, 504.

Rhinoptera

javanica, 283, 284, 286,

288, 289.

quadriloba, 279,

Rhodites

rose, 174.

Rhynchobothrius, 280.

72, 78, 79.

Rhynchonax

andersont, 1380, 131.

Rhytidoceros

undulatus, 142.

Robertus

lividus, 397.

Ruticilla

rufiventris, 142.

Salmo

fario, 406.

Saltator

senilis, 409,

Sarcophilus

satanicus, 232.

Scardafella

sguamosa, 415.

Sceloporus

clarkii, 413.

Sciurotamias

davidianus consobrinus,

154.

Sciurus

castaneoventris

hotet, 134.

Scolopax

rusticula, 71, 72,

ao:

Scops

bakkamena, 2.

leucotis, 416.

Scopus, 126.

Scotopelia

bouvieri, 416,

Scutella, 476.

Segestria

senoculata, 395.

Segmentina

eussoensis, 191, 193.

kempt, 191, 193.

Seira

brahma, 81, 99, 120

123,

Frigida, 81, 97, 123.

sumatrana, 120.

(Sira) annulicornis,

120.

Semnopithecus

ptleatus, 1.

Sericulus

melinus, 4.

Serinus

canicollis, 409.

Setigera

phanolepis,

120, 124.

tarsata, \20.

bon-

81, 110,

travancorica, 81, 108, |

IAD), eae

INDEX OF SCIENTIFIC NAMES.

Sialia

sialis, 409.

Simia sp., 128.

rhesus, 128.

sylvana, 128.

Sinella

curviseta, 101.

hofti, 102.

montana, 81, 101, 128,

myrmecophila, 102.

Singa

albovittata, 899.

hamata, 399.

Sistrurus

miliarius, 413.

Sitala

iredalet, 187, 193.

Sitta

castaneiventris, 2.

Sitticus

longipes, 402.

Sminthurides

appendiculatus, 81,117,

121, 125.

Sminthurus, 117.

Sorex

bedfordie, 132.

cylindricauda, 132.

religiosa, 228.

wardi, 182.

— fumeolus, 132.

Soriculus

irene, 132, 138.

macrurus, 132, 138.

Spalax

egyptiacus, 230.

typhus, 230.

Spatangus, 479, 493.

Spermophila

castanetventris, 409.

minuta, 408.

Sphezrium

kigeziensis, 192, 193.

Spheniscus

humboldti, 61, 62.

Sphenodon, 421.

Spirocheta

recurrentis, 410,

Staurotypus

treporcatus, 414, 415.

Steatoda

bipunctata, 397.

Steatomys

pratensis, 410, 411.

Sternothzrus

niger, 414.

Stilesia, 208.

hepatica, 207.

sjostedti, 216.

INDEX OF SCIENTIFIC NAMES.

Stoparola

melanops, 415.

Strepera

_ fuliginosa, 408.

Strix

Hamimea, 416.

Styloctetor

broccha, 398.

(Hilaira) carli, 398.

Succinea

kempi, 189, 198.

princet, 189, 193.

Sula

capensis, 1,

Suricata

suricatta, 407.

Sus

aper, 391, 392.

attila, 391, 392, 393.

celtica, 391, 392.

europeéus, 391, 392.

scrofa, 390, 591, 392.

— beticus, 391, 392,

393.

castilianus,

392.

— scrofa, 391, 392.

setosus, 391, 392.

Synageles

venator, 402.

291,

Tadorna

tadornoides, 416.

Tenia

marchali, 2\7.

saginata, 195, 196,

Teeniura

melanospilos, 286.

Tanagra

episcopus, 409, 417,

palmarum, 416.

Tapinocephalus, 425,

Tapinocy ba,

affinis, 898.

pallens, 398.

Tapinopa

longidens, 397.

Tarbophis

fallax, 412.

Tarentola

annularis, 414.

mauritanica, 412,

Tarentula

accentuata, 401,

aculeata, 401.

andrenivora, 400.

barbipes, 400.

lessertii, 400, 404,

405.

miniata, 400.

Tarentula

nemoralis, 400.

pulverulenta, 400.

trabalis, 400.

Tayassu

tajacu, 407.

Tegenaria

campestris, 396.

derhamii, 390.

domestica, 396.

pusilla, 396.

tridentina, 396.

‘Termes

redemanni, 117.

Testudo

angulata, 411.

Tetragnatha

extensa, 398.

Tetragonocephala, 284,

Tetrao

tetria, 72, 77, 79.

Tetrarhynchus, 269.

sp., 288, 292.

balistidis, 286.

evinaceus, 287, 350.

minus, 279.

pinne, 286.

unionifactor, 268, 278, |

279-283,

347, 361.

Teutana

grossa, 397.

Textor

adlector, 415,

Textrix

denticulata, 306.

Thanatus

arenarius, 400.

Sormicinus, 400.

Thapsiella

millestriata,

193:

opposita, 186, 193.

Tharrhaleus

perdoni, 2,

285-290,

186,

Theganopteryx, 865, 366, |

367.

Thelotornis

kirtlandi, 238,

Theridion

bigibbum, 396, 403.

blackwallii, 397.

nigro-punctatum, 397.

nigro-variegatum, 397.

pallens, 404.

riparium, 390.

sisyphium, 397.

Thomisus

onustus, 099,

Thylacinus

cynocephalus, 232.

XXVil

Thyrsocera, 362.

Thysanoteenia,

218.

Tiarabothrium

Javanicum, 284.

Tigellinus

saxicola, 397.

Tinnunculus

alaudarius, 416.

Tomocerus, 115.

vulgaris, 81, 91.

(Macrotoma)monlanus,

120.

Toxostoma

bendirei, 3.

Trachyeystis

tredalei, 187, 193.

Tragulus

javanicus, 407.

Trematopygus, 466.

Trichoglossus

nigrigularis, 416.

Trichomonas, 411.

Trichorypha, 106.

Triconchus

mobi, 397.

Tringa

minuta, 2.

Trirachodon, 425,

Trochosa

leopardus, 400,

terricola, 400.

Tropidonotus

fasciatus, 415.

Trygon, 261.

kuhli, 284.

warnak, 284,

walga, 283, 284.

Trypanosoma

avium, +10.

Zewisi, 410.

rotatorium, 410.

214,

Tupinambis

tequexin, 412.

Turacus

corythaix, 416.

erythrolophus, 3.

macrorhynchus,

416.

persa, 416.

Turdus

migratorius, 408,

mustelinus, 408.

Tylocephalum, 262, 275,

287.

dierama, 284,

hkuhli, 284.

ludificans,

408,

279-285,

282-293, 296, 347,

848, 352, 358.

margaritifere, 274.

XXVU1

Tylocephalum

minus, 279,

285, 286,

292, 298,

352.

pingue, 279.

warnak, 284, 358.

(Letragonocephalum)

abtobatidis, 283.

(—) trygonis, 283.

Tympanistria

bicolor, 415.

282, 283,

288, 290,

337, 347,

Urobrachya

albonotata, 408.

Urocissa

occipitalis, 410.

Uropsilus, 127.

soricipes, 129,

USI.

Urotrichus, 130, 131.

Ursus

isabellinus, 498.

130,

Varanus

bengatensis, 412.

niloticus, 414.

varius, 412.

Vidua

paradisea, 417.

Vipera

ammodytes, 414.

russelli, 414.

Vitrina

compacta, 184, 193.

Vivipara

rubicunda kisumiensis,

191, 198.

Vulpes

hengalensis, 252.

lagopus, 55, 238.

zerda, 225.

Withius, 388.

Xenylla

brevicauda, 895.

grisea, 809.

humicola, 85.

maritima, 85.

mitida, 85.

obscura, 81, 84, 119,

Xiphidiopictus

percussus, 232.

Xysticus

bifasciatus, 399.

cristatus, 399.

erraticus, 399.

INDEX OF SCIENTIFIC NAMES.

Xysticus

gallicus, 399.

glacialis, 399.

kochit, 399.

lanio, 399.

lateralis, 399.

luctator, 399.

pini, 399.

Zamenis

constrictor, 413.

dahli, 414.

jlagelliformis, 413.

gemonensis, 414.

hippocrepis, 412.

mucosa, 413.

Zilla

montana, 398.

stroemiti, 398.

«z-notata, 398.

Zingis

kempt, 185, 193.

papyracea, 185, 193.

planispira, 185, 195.

Zodarion

gallicum, 396.

Zora

maculata, 396.

Zosterops

virens, 233.

INDEX

PEE UST RAPIONS:

Aceipiter nisus, Pls. II., IV., p. 63.

Africarion kempi, Pl, XXX1. p. 183.

—— microgranulata, Pl. XXXI. p. 183.

— microstriata, Pl, XXXT.sp. 183.

Alea keniana, Pl. XXXI. p. 183.

Anaplecta decipiens, Fig. 44, p. 365.

Ancylus kempi. Pl, XXXT. p. 183.

Anisolemnia distaura, Pl. XUVIILI.

p. 358.

Anorthopygus, Figs. 55, 56, 58, 59,

pp. 459, 464, 473, 479.

Aptenodytes pennanti, Pl. I. p. 60.

Ardea cinerea, Pl. XIII. p. 125.

Assimania aurifera, Pl. XXXT. p. 183.

Chelifer latreillii, Rigs. 47, 50, pp. 380,

386.

Chorisoneura teniata, Fig. 43, p. 365.

Chrotogale owstoni, Figs.61-63, pp. 500-—

502.

Circus eruginosus, Pl. ITT. p. 63.

—— ctneraceus, P|. III, p. 63.

—— cyaneus, Pls. III.-V., p. 63.

Clypeaster, Fig. 59, p. 479.

Celophora formosa, Pl. XLVITI. p. 358,

Coenholectypus, Fig. 59, p. 479,

Conoclypeus, Fig. 57, p. 469.

| Conulus, Figs. 55-59, pp. 459, 464,

Asticopsis franklinii, Pls. XIV., XV., |

XXV., XXVI., p. 144.

serratus, Pls.

XXVL., p. 144.

Bacillus rossii, Pls. XXVIII-XXX.,

p. L72.

Blawieria exsilium, Pl. XXX. p. 183.

Blood - parasites. Pls. XLIX.-LYV.,

p. 106.

Caricella

. 126.

Cephilobothrium, Fig. 84, p. 285.

aétobatides, Pl. XLVII. p. 260.

Cherips intermedius, Pls. XXIV.,

XAXVIL., p. 144.

—— quinquecarinatus, Pls. XXIIL.,

XXIV., XXVIL., p. 144.

fenuimanus, Pls, XXII., XXVIL.,

. 144.

Chelier cyrneus, Figs. 48-50, pp. 882,

384 386.

cunningtoni, Pl. LVIII.

469, 478, 479.

albogaterus, Wig. 54, p. 455.

| Cremastocephalus indicus, Pl. IX. p. 80.

—— kershawi, Pls. X1X., XX., p. 144. |

XVI-XVIIL., |

montana, Pl, 1X. p. 80.

Cygnus bewicki, Pl. TIT. p. 63.

Cynognathus, Fig, 53, p. 424,

| Diagram of the adoral surface in some

Holectypoida, Fig. 54, p. 455.

of the perignathic girdles in some

Holectypoida, Fig. 55, p. 459.

showing some characteristic plates

of the ambulacra in some Holecty-

poida. Figs. 56, 57, pp. 469, 473.

showing the shape and position of

the periproct in some Holectypoida,

Fig. 56, p. 464.

Dicranocentroides

VITI.-X., p. 80.

Dicynodon, Fig. 58, p. 424.

Discoidea, Figs. 55-59, pp. 459, 464,

469, 473, 479.

Jusciculatus, Pls,

| ——— eylindrica, Fig. 54, p. 455.

Elephantulus rupestris, Fig. 17, p. 143.

Ennea microstriata, Pl. XXX. p. 185,

XXX

Entomobrya crassa, Pl. VII. p. 80.

kal, Pls. VII., VIII, p. 80.

Hussoia tnopina, Pl. XXXII. p. 1838.

Falco peregrinus, Pl. IIT. p. 63.

subbuteo, Pl. III. p. 63.

—- tinnunculus, Pl. IL. p. 63. |

Galeropygus, Figs. 57, 58, pp. 469, |

473.

agariciformis, Fig. 54, p. 455.

Galictis, Fig. 31, p. 222.

Gazella rufifrons haslert, Figs. 2, 3,

pp. 5, 6.

Hemoproteus danilewskyt, Pl. LIT.

. 406.

Hemithyrsocera sp., Fig. 42, p. 364.

Heteromuricus ceroifer, Pls. VIII., 1X.,

p- 80.

Holectypus depressus, Figs, 54, 56, 58,

pp. 455, 464, 473.

hemispherieus, Figs,

pp. 464, 473.

Homorus iredalei, Pl, XXX1I, p, 183.

56, 58,

Idiomerus pallidus, Pl. IX. p, 80;

Fig. 15, p. 114.

Isotoma nigropunctata, P\. VIL. p. 80,

siva, Pls. VI., VIL., p. 80.

Isotomurus palustris, Pls. VI,, VIL, |

p: 80. |

Kaliella consobrina, Pl. XXXI. p, 183,

depauperata, Pl. XXXI. p. 183.

tredalet, Pl. XXXI. p. 183.

— kigeziensis, Pl, XXX]. p. 183.

Lagopus scoticus, Pl. 111. p. 63,

Larus minutus, Pl. ILL. p. 63.

Leis duntopi, Pl. XLVILII. p. 388.

Leucochiloides chanlerensis, Pl. XXX.

p. 183.

gaziensis, Pl. XXXI. p. 183,

wredalei, Pl. XX XI. p. 183,

soror, Pl. XXXTI. p. 183.

LTimnea kempi, P\. XXXII. p. 183,

LTimnocaridina parvula, Pls. LiX., LX.,

p. 426,

spinipes, Pl. LVIL. p. 426.

Macropygus, Fig. 56, p. 464.

Macroscelides proboscideus,

p. 143.

Map of Australia, showing distribution

of Orayfishes. Fig. 18, p. 149.

of Oriental Region, showing

localities at which Collembola were |

obtained. Fig. 14, p. 82. |

Margaritifera margaritifera, Fig. 38. |

p. all.

Fic. 16, |

| Plans showing horse-trough, ete.

_ Podicipes cristatus, Pls. IIT., 1V.,

INDEX OF ILLUSTRATIONS.

Margaritifera maxima, Figs. 36, 37,

pp. 805, 306.

—— vulgaris, Pls. XXXIII-XXXvV.,

XXXVIT.-XUL., p. 260; Fig. 35,

p. 302.

Megapyga eximia, Pl. XLVITL. p. 358.

Megapygus, Figs. 56, 57, pp. 464, 469.

Mellivora, Figs. 31, 32, pp. 222, 223.

Melyroidea magnifica, Pl. XUNIIT.

p- 358.

—— mimetica, Pl. XLVIII. p. 358;

Fig. 46, p. 874.

Mytilus edulis, Pl. XXXTV. p, 260.

Natalina permembranacea, Pl. XXXI.

p- 188.

Neanura corallina, Pl. VII. p. 80.

intermedia, Pls. VI,, VIL., p- 80.

pudibunda, Pl. V1. p. 80.

Oides biplagiata, Pl. XLVIII. p. 358.

Otiditenia eupodotidis, F igs. 19-380,

pp. 194, 197-202, 204, 206, 208, 212,

215.

Oudenodon holbei, Pl, LVI. p. 419.

| Paracheraps _bicarinatus, Pls. XXT.,

XXVIL, p. 144.

Paronella borneri, Pls. X,, XI1., p. 80.

—— gracilis, P|. XI. p. 80.

—— insignis, Pi. XI. p. 80.

——-phanolepis, Pls. X,, XI, p. 80,

= travancorica, Pls. 1X., X., p. §0.

Pearls. Pls. XXXV.-XLVL., p. 160;

- Figs, 38, 39-41, pp. 277, 318, 819,

340.

Phylogenetie table of the Holectyyoida

and their allies. Fig. 60, p. 498.

Pileus, Fig, 57, p. 469. |

Planorbis kigeziensis, Pl, XXXII.

p- 183.

sperabilis, Pl. XXXII. p. 183.

used

for homing experiments with Am-

phibia, Figs. 4, 5, pp. 12, 15.

Plasmodium precox, Pl, LILI, p, 406,

ip. 65,

Prioptera sinuata, Pl, XLVIIL. p| 358.

Prosoplecta bipunciata, P1, AN. Il.

p- 308.

coccinella, Pl. XLVIIL, p. 38.

celophoroides, P\. XLVITL. }, 358,

—— guiticollis, Pl. XLVIIL. p. 8,

mimas, Pl. XLVITT. p. 858,

nigra, Pl. XLVILIL. p. 358.

—— nigroplagiata, P\, XLVIII, }, 358.

—— nigrovariegata, Fig, 45, p, a

quadriplagiata, Pl. XLV IT

p. 358.

INDEX OF ILLUSTRATIONS.

Prosoplecta rufa, Pl. XLVILII. p. 358.

senperi, Pl. XLVILI. p. 358.

trifaria, Pl, XLVIIL. p. 358,

Pseudachorutes anomalus, Pl. VI. p. 80.

Pseudosira indra, Pls. VII., VIIL.,

p. 80.

Pygaster, Figs. 55-59, pp. 459, 464,

469, 473, 479.

-—— semisulcatus, Fig. 54, p. 455.

(Macropygus) laganoides, Fig. 54,

455.

(Megapygus), Fig. 59, p. 479,

Pyrina, Fig. 57, p. 469.

Rallus aquaticus, Pl, ILI, p. 63,

Scolopax rusticula, P\. III. p. 68.

Segmentina eussoensis, Pl. XXXII,

p. 183.

kempi, Pl. XXXII. p. 183.

Seira brahma, Pl. VIUI. p, 80.

frigida, Pl. VIII, p, 80.

Sericulus melinus, Fig. 1, p. 4.

Sinella montana, Pls. VILL., [X., p, 80,

Sitala iredalei, Pl. XXXI. p. 183.

Spatangus, Fig. 59, p. 479.

Spherium kigeziensis, Pl. XXXI. p, 183.

XXX1

Suceinea kempi, Pl, XXXIT. p. 183.

princei, Pl. XXXII. p. 183.

Tarentula lessertii, Fig. 52, p. 405.

Tetrao tetrix, Pl. III. p. 63.

Thapsiella millestriata, P}.

p. 183.

opposita, Pl. XX XIT. p. 185.

Theridion bigibbum, Fig. 51, p. 403.

Trachycystis iredalei, Pl, XXXII,

p. 183.

Tylocephalum, Big. 34, p. 285.

ludificans, Pls. XXXIII., XLVI,

XLVII., p. 260.

minus, Pl. XX XIII. p. 260,

uarnak, Pl, XLVIL. p, 260,

XXXII.

Vitrina compacta, P|. XXXII. p. 185.

Vivipara rubicunda kisumiensis, Pl,

XXXII. p. 183.

Vulpes lagopus, Figs, 6-13, pp. 56-59.

Xenylla obscura, Pl. VI. p, 80.

Zingis kempi, Pl. XXXIT. p. 183.

papyracea, Pl, XXXII. p. 185.

-—— planispira, Pl, XXXII. p, 183,

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

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PROCEEDINGS

OF TIE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF TILE

ZOOLOGICAL SOCIETY OF LONDON.

EXHIBITIONS AND NOTICES.

October 24, 1911.

Sir Jonun Rose Braprorp, K.C.M.G., M.D., F,R.S.,

Vice-President, in the Chair,

Tue Secrerary read the following report on the Additions that

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

June, July, August, and September, 1911 :—

JUNE.

The registered additions to the Society’s Menagerie during

the month of June were 602 in number. Of these 164 were

acquired by presentation, 70 by purchase, 265 were received on

deposit, 32 in exchange, and 7] were born in the Gardens.

The number of departures during the same period, by death

and removals, was 328.

Amongst the additions special attention may be directed to :—

A collection of Mammals, Birds, and Reptiles from Africa

(chiefly South), presented to H.M, King George by various donors,

and deposited in the Gardens by His Majesty on June 2nd,

including 2 Eastern White Pelicans (Pelecanus roseus), 3 Cape

Gannets (Sula capensis), 1 Scarce Duck (Anas sparsa), | Robust

Lizard (Gerrhosaurus validus), and | Black-lined Lizard (Gerrho-

saurus nigro-lineatus), all new to the Collection.

2 Capped Langurs (Semmnopithecus pileatus) from the Himalayas,

new to the Collection, purchased on June 6th.

1 Hoffmann’s Two-toed Sloth (Chalepus hoffmanni), from

Brazil, purchased on June 17th.

Proc, Zoon, Soc, —1912, No. I. |

2 THE SECRETARY ON ADDITIONS TO THE MENAGERIE.

A collection of rare Indian Birds presented by E. J. Brook, Esq.,

F.Z.S., on June 6th and 19th, including 2 Grey-headed Ouzels

(Merula castanea), 2 Pied Bush-Chats (Pratincola caprata),

1 Ruby-throat (Calliope camtschatkensis), 1 HKlegant Bunting

(Hmberiza elegans), 1 Jerdon’s Accentor (Tharrhaleus jerdoni),

2 Yellow-cheeked Tits (J/achlolophus xanthogenys), 2 Chestnut-

bellied Nuthatches (Sitta castaneiventris), 1 Black-headed Wagtail

(Motacilla feldeggi), 1 Blue-throated Flycatcher (Cyornis rube-

culoides), 2 Sikkim Siskins (Chrysomitris tibetana), 1 Collared

Scops Owl (Scops bakkamena), and a pair of Blood Pheasants

(Lthagenes cruentus), all new to the Collection.

2 Peale’s Parrot-Finches (Hrythrura pealei), from the Fiji

Islands, new to the Collection, deposited on June 14th.

2 Taviuni Parrakeets (Pyrrhalopsis taviunensis), from Taviuni,

Fiji Islands, new to the Collection, presented by Dr. P. H. Bahr,

FE.Z.S., on June 6th.

1 Pigmy Crake (Porzana pusilla), and 3 Little Stints (7ringa

minuta), from India, new to the Collection, deposited on June 6th.

2 Hooded Cranes (Grus monachus), from Japan, purchased on

June 6th.

JULY.

The registered additions to the Society’s Menagerie during the

month of July were 444 in number. Of these 174 were acquired

by presentation, 62 by purchase, 56 were received on deposit,

12 in exchange, and 140 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 278.

Amongst the additions special attention may be directed to :—

1 African Cheetah (Cynelurus jubatus), from Berbera, Somali-

land, purchased on July 22nd.

3 Cape Sea-Lions (Otaria pusilla), from South Africa, received

from Dr. Louis Péringuey, F.Z.8., for H.M. THe Kine’s African

Collection, on July 29th.

1 Tibetan Wild Ass or Kiang (Zquus kiang), 3, born in

the Menagerie on July 13th.

7 White-rumped Lories (Hos fuscata), from New Guinea, new

to the Collection, | purchased and 6 deposited on July 13th.

1 Wolf-Snake (Coluber vulpinus), from North America, new to

the Collection, received in exchange on July 8th.

A. Collection of rare Reptiles and Batrachians from Trinidad,

presented by Dr. Lewis Henry Gough on July 11th, including

18 Warty Tree-Frogs (Tyla venulosa), 1 Daudin’s Tree-Frog

(Hyla rubra), and 16 Tree-Frogs (Hyla goughi), all new to the

Collection.

AvGusT.

The registered additions to the Society’s Menagerie during the

month of August. were 229 in number. Of these 85 were

acquired by presentation, 21 by purchase, 8 were received on

deposit, 62 in exchange, and 53 were born in the Gardens,

THE SECRETARY ON ADDITIONS TO THE MENAGERIE, 3

The number of departures during the same period, by death

and removals, was 244.

Amongst the additions special attention may be directed to :—

1 Red-faced Spider-Monkey (Ateles paniscus) and 1 Brown

Capuchin (Cebus fatuellus), from British Guiana, presented by

Messrs. John Reed and W. A. Boyd on August 3rd.

1 Clouded Tiger (Felis nebulosa), from British North Borneo,

presented by Almeric Paget, Esq., F.Z.S., on August 29th,

1 Ocelot (Felis pardalis), from Para, presented by F, J. Weldon

Taylor, Esq., on August 16th.

1 Lesser Superb Bird-of-Paradise (Lopharhina minor), from

S.E. New Guinea, new to the Collection, presented by E. J. Brook,

Ksq., F.Z.S., on August 12th.

1 Red-crested Touracou (Z'uracus erythrolophus), from Angola,

deposited on August 31st.

A Collection of American Birds, including 2 Bendire Thrashers

(Toxostoma bendirer), 4 Gila Woodpeckers (Jelanerpes uropygialis),

and 4 Douglas’ Quails (Lophortyx douglasi), all new to the

Collection, received in exchange on August 3rd.

2 Broad-tailed Babblers (Crateropus platycercus), from Gambia,

new to the Collection, presented by Dr. E. Hopkinson, D.S,O.,

F.Z.S., on August 7th.

1 Striped Gonatode (Gonatodes vittatus), from Trinidad, new to

the Collection, presented by Dr. L. H. Gough on August 31st,

SEPTEMBER.

The registered additions to the Society’s Menagerie during the

month of September were 361 in number, Of these 137 were

acquired by presentation, 39 by purchase, 39 were received on

deposit, 125 in exchange, and 21 were born in the Gardens,

The number of departures during the same period, by death

and removals, was 254.

Amongst the additions special attention may be directed to :—

1 Persian Gazelle (Gazella subgutturosa), from Arabia, presented

by Rear-Admiral Sir E. J. W. Slade, K.C.I.E,, M.V.O., R.N.,

F.Z.S., on September 7th.

2 Tamaulipas Tawny Thrushes (Merula tamaulipensis), 2 Derby

Tyrants (Pitangus derbianus), 1 Giraud Tyrant (Jyiozetetes

similis), 4 Black-breasted Colins (Colinus pectoralis), fram Mexico ;

4 American Barn-Owls (Asio accipitrinus), from North America,

1 Hodgson’s Pipit (Anthus rosaceus), 1 Indian Tree-Pipit (Anthus

maculatus), 2 Grey-headed Buntings (Hmberiza fucata), 1 Pine-

Bunting (Zmberiza leucocephala), and | Hume's Short-toed Lark

(Calandrella acutirostris), from India, all new to the Collection,

and received in exchange on September 27th.

2 Formosan Pheasants (Phasianus formosanus), from Formosa,

new to the Collection, presented by W. H. St. Quintin, Esq., F.Z.S.,

on September 12th.

4 MR. D. SETH-SMITH ON A NESTLING REGENT-BIRD.

Mr. James DunsBar-Brunton sent for exhibition two skins and

a mounted skull, with horns, of Bushbuck shot by him in North-

Kast Rhodesia.

. W. B. Corron, of the Indian Civil Service, exhibited a

number of heads and horns of various species of Gazelles which

he had obtained in the Eastern Sudan, and gave a brief account

of their habits and distribution. The specimens included

Gazella isabella, from the hills behind Suakin and the route

from Sinkat to Kassala, and eleven specimens, of which he under-

stood nine to be Gazella tilonura and two to be Gazella rufifrons,

from the Atbara, Settit, and Rahad.

Mr. Cotton mentioned that some sportsmen believed isabella to

be identical with dorcas, and advanced a decided opinion that

tilonura was merely a local form of rufifrons. This opinion was

fortified by consideration of the extreme variation of type in the

specimens exhibited, of which all but one had come from the

same locality, namely the Atbara and Settit.

Mr. D. Serx-Smiru, F.Z.S., Curator of Birds, exhibited a

spirit-specimen of a nestling Australian Regent-Bird (Sericulus

melinus) (text-fig. 1), which had been hatched in the aviary of

Mr. Reginald Phillipps, of 26 Cromwell Grove, West Kensington,

during the past summer, The male parent of this bird was also

bred in the same aviary in 1906, and is the only specimen of this

fine Bower-Bird ever bred and reared to maturity in captivity.

Text-fig, |

Sericulus melinus, ten days old.

Two young birds were hatched this year (1911), but succumbed

to the effect of a thunderstorm in July, when about ten days old.

The nestling is chiefly remarkable for the length and thickness

of fluffy down on the feather tracts, especially upon the head.

MR. R. I, POCOCK ON A NEW RED-FRONTED GAZELLE, a)

The exhibitor remarked that thanks to Mr. Phillipps’

generosity, the parents of this young bird now formed part of

the Society’s collection, and were amongst the rarest of the

exhibits, since at the present time they were probably the only

living examples of this fine species in captivity.

Mr. R. I. Pocock, F.R.S., F.L.S., F.Z.S., Superintendent of

the Gardens, exhibited two photographs (text-figs. 2 & 3) of a

male specimen of Red-fronted Gazelle (Gazella rufifrons) which

came from Kano in Northern Nigeria, and was presented to the

Society in 1908 by Col. Julian Hasler. This specimen apparently

represented an undescribed race which he proposed to name

and to diagnose as follows :—

GAZELLA RUFIFRONS HASLERI.*

Abstract P. Z. S. No. 100, p. 47 (Oct. 31st).

Back ochre-fawn in colour with a well-defined paler area

between the richer tint of the dorsal area and the infero-lateral

Text-fig. 2.

Side view of Red-fronted Gazelle, Gazella rufifrons hasleri.

black stripe. This stripe, as in the typical form from Senegambia,

with a narrow border of fawn below. Basal inch of the tail the

same tint as the back, the rest black. Only a faintly defined dark

* The complete account of this new subspecies appears here, but since the name

and a preliminary diagnosis were published in the‘ Abstract,’ it is distinguished by

being underlined,—Eprror,

6 MR. J. LEWIS BONHOTE ON

mark bordering the white of the back of the thighs. Forehead

a darker and richer colour than the neck and cheeks, a few white

hairs between the horns. The whole of the upper side of the

nose from the preorbital glands to the nostrils snow-white like

the chin. The glands themselves and the area round the eye

creamy white or very pale fawn. No black on the legs or feet ;

the glandular tafts below the knees present as in all the examples

of G. rufifrons that had been exhibited in the Gardens.

Text-fig. 3.

Head of Red-fronted Gazelle, Gazella rufifrons hasleri.

This Gazelle differed from all previously described specimens of

G. rufifrons in the whiteness of the upper side of the nose.

Mr. J. Lewis Bonnors, M.A., F.L.S., F.Z.8., exhibited living

specimens of Rats (us rattus) which he had bred in the course

of his experiments and which showed the “ waltzing” character

well known in a variety of the domestic mouse, but which had

not hitherto been recorded in rats.

The original stock from which these rats had been bred was

caught wild at Giza near Cairo (see P. Z.8. 1910, p. 664), and the

experiments were being conducted to study heredity in two

varieties found in a wild state, namely, Mus rattus teclorwm with

pure white underparts, and Mus rattus alewandrinus, im which

the hairs of the underparts were entirely slate-eoloured or

had slate-coloured bases. ‘The experiments were also being

conducted to test the inheritance of a fawn-coloured individual

which had appeared asa “sport” in the first generation from

pure wild parents (P. Z.8. 1910, p. 638). With regard to the

heredity of the two normal varieties the white-bellied form (JZ. r.

tectorum) was found to be apparently a simple Mendelian

dominant to the dark-bellied form (J/. rv. alewandrinus), the

WALTZING RATS. 7

dark-bellied ones always breeding true and the heterozygous

light-bellied ones giving a proportion of pure alexandrinus.

This entirely bore out the suggestion put forward by Mr. Bonhote

in a former paper (loc. cit.) that the main varieties (or sub-groups

as he had called them) of Mus rattus were true Mendelian

mutations. It would thus be understood how forms (which were

at present considered as closely allied species) and which differed

only in small but constant characters, were able to exist side by

side under practically identical conditions without those characters

becoming blended or one of them being lost, as would be bound to

occur if natural selection were the only active force in evolution.

With regard to the heredity of the fawn—the original specimen,

a male, which showed the characteristic white underparts of J/.

teclorum (its parents), was mated to an alewandrinus 9 , and the

resulting F, generation, some thirty in number, were all typical

white-bellied tectorwm.

Five pairs of these were mated and gave in the F, generation :—

17 Tectorum, 5 Alexandrinus, 7 Fawn 'T., 1 Fawn A.

the expectation

being 18 A 6 PS 6 4 2 a!

In addition, there were three individuals that died before it

could be determined whether they were alexandrinus or tectorum,

but they were not Fawn, as these could be distinguished at birth.

The fawn colour, which was probably due to the absence of black,

was, therefore, recessive to the normal grey colouring but might

occur in either of the normal forms. Thus a rat having an

absence of black and presence of the Alexandrine (slaty underparts)

character was whitish fawn in colour, since the number of hairs

which should contain black was much greater and in the absence

of the black factor these hairs were colourless.

It might be noticed that in young Fawn Rats the eye was

ruby coloured, as in the case of the Cinnamon Canary, becoming

quite dark as maturity was reached.

The “waltzing” rats, of which altogether four had been

produced, all appeared in the F, generation, but were not all from

the same parents. Of the matings for the F, generation only

one (daughter x father) produced sound young. Apart from the

“‘ waltzing ” character, three individuals were born blind, and in

two of these cases the eyes were undeveloped and the optic nerve

absent, and in addition many of the young that were reared were

so weakly that they had to be killed, and others died before being

weaned.

From certain causes, therefore, which are not very clear, partly

due to inbreeding, partly probably to environment, and partly pos-

sibly to their breeding at too early an age (6 months), a degenerate

and defective generation was produced, and one of the results

was to bring about in certain individuals a character (‘ waltz-

ing”), which in a nearly allied species was shown * to have a

Mendelian inheritance.

* Darbishire, ‘Biometrika,’ i. pp. 101, 165, 282 (1902), ibid. iii. p. 1 (1903) ;

G. von Guaita, Ber. Naturg. Ges. Freiburg, x., x1. (1898) (1900).

8 MR. B. F. CUMMINGS ON

PAPERS.

1. Distant Orientation in Amphibia.

By Bruce F. Commines *.

[ Received September 1, 1911: Read October 24, 1911.)

(Text-figures 4 & 5.)

INTRODUCTION.

The observations and experiments detailed below were made in

North Devonshire during the course of the last two years. The

Amphibia chiefly used were two species of Newts. The British

Newts have a peculiar geographical distribution in these Islands,

and, living in the north of Devon, I was fortunate in being able

to obtain large quantities of two species, Molge cristata Laur., and

Molge palmata Schneid. It has long been the custom of field

naturalists, year after year with all the precision of Gilbert

White of Selborne, to note the date of the first return of Frogs

and Efts to their breeding-ponds as indicative of the return of

spring. It occurred to me to enquire how the Amphibia find the

water again after having left it in the autumn of the preceding

year. Nothing appears to be known of this subject, although the

problem of these amphibian migrations is a simple one compared

with that of the migrations of birds, which has attracted, and still

is attracting, so much attention from the students of animal life.

No one has shown how the Amphibia find the water, and it

is not definitely known whether the Amphibia return to the same

piece of water each spring, nor whether every animal or only

a lucky small percentage finds water as the breeding season comes

round, nor precisely how extensive their perambulations on land

may be during the autumn months before hibernation sets in.

Many larval Newts, and a few adult Newts of Molge palmata

remain in the water hibernating at the bottom of the pond. But

these are exceptional. The majority of our Amphibia do leave

the water, and large numbers annually find their way back to it

in the following spring. The Newt is essentially a nocturnal

animal, and a visit to a newt-pond with a lantern after dark,

whether in spring or autumn, is sufficient to show that night is

the time when the migrations are carried on. The answer te

the question how these amphibian migrations are performed

must involve points of importance to amphibian psychology and,

when elucidated, the subject will form an interesting chapter

in their naturai history. The literature is scant and vague.

G. J. Romanes t thought that Frogs had a distinct idea of locality.

He based this idea on the cases reported to him by some of his

correspondents, where Frogs, removed a distance of 200 or 300

* Communicated by T. A. Cowarp, F.Z.S.

+ G. J. Roimanes, ‘ Animal Intelligence,’ p. 254.

AMPHIBIAN MIGRATION, 9

yards from their habitual haunts, returned to them again and

again. He also expressed the opinion that Frogs were able

to perceive moisture from a great distance, and he quotes Warden

who in “ An Account of the United States,” vol. ii. p. 9, says

that a pond containing some Frogs having dried up, the animals

“made straight for” the nearest water, though it was eight

kilometres away.

CoMPENSATORY HBAD-MOVEMENTS.

In all experiments with the Amphibia it is necessary to beware of

error through compensatory head-movements, which, if neglected,

give « false complexion to the results. No mention of them is

made by Romanes’ correspondents and no details of the experi-

ments are given. It isastonishing to find how persistently Newts

will return and immediately re-enter their pond, after they have

been removed from it to some considerable distance. Even when

placed heading directly away from the water, they turn round and

walk towards the water, as if endowed with a quite uncanny

knowledge of the geographical features of the neighbourhood of

their breeding-pond. The explanation partly lies in the head-

movements contrary to the movement of rotation, set up by the

motion of the experimenter’s hand as he turns the Newt round

to face away from the water. The Newt mechanically responds

by turning round, with the head towards the water. These move-

ments are very pronounced in the Newt on account of its linear

shape. A Newt, on a rotating disc which is being moved back-

wards and forwards through an angle of 180°, responds correctly

if the motion is slow, 7.e., it turns its head regularly in the

opposite direction to the backward and forward movements of the

dise. If the motion of the dise be fast, it overtakes the slow

head-movements of the Newt, and the Newt hecomes confused and

escapes from the predicament by lowering its chin so as to place

it in contact with the moving disc. No response is then given.

Continuous rotation in one direction produces no very marked

external signs of giddiness. But in the Common Toad (Bufo

vulgaris) these signs are marked. While the disc is moving the

head is motionless. As soon as it stops, the Toad begins to crawl

round in circles moving in the direction the dise had been moving.

One of these animals, after rotation of one minute, in this way

crawled through four circles! If a Newt be rotated through 90°

in a trough with perpendicular walls two inches high, I have seen

it turn in response to the rotation and climb over the side.

GEOTAXIS.

This subject must also be taken into consideration, It must

be borne in mind that the ground around most ponds is sloping.

Most fresh water lies at the bottom of hollows or at the foot of

slopes. Newts are positively geotactic. They preferred to walk

10. MR. B. F. CUMMINGS ON

downhill when I tested them in the spring, but in the autumn

this preference changed and they went up. In work with

an inclined plane, Molge cristata was chiefly used. The Newts

were kept in water in large earthenware pans under similar

conditions so far as possible. The observations were made in

the early morning or at night, always in subdued light. The

plane was of wood and measured 125 centimetres by 82 centi-

metres 5 millimetres. A line was drawn transversely across the

plane, and each Newt on removal from the pan was allowed

to crawl on a level surface adjoining until manceuvred with the

finger into the right direction, when it was swiftly lifted on to the

incline and left in a horizontal position. In this way I cut out

errors arising from compensatory head-movements. An interval

of ten minutes was allowed each animal before being tried again.

This prevented errors arising from simple habit formation, which

in the Newt is rapid.

Table I. shows the results of experiments performed in the

spring when the Newts, used singly, walked downhill in 80 per

cent. of the tests. Similar results were obtained by using Newts

in series; twenty Newts were used. The number of positive re-

sponses was in this way reduced to 67 per cent., which perhaps

is nearer the truth. Table II. was obtained in the autumn

when the Newts were endeavouring to leave the water for winter

quarters. The Newts were then negative and walked uphill

72 per cent. of the times. In this table the Newts were used in

series, the first row being worked by twenty Palmates and the

rest by twenty Crested.

TART E

neice Incline. Eoeion at | Down. Up. | Neutral.) Total.

1 16° Horizontal. 12 (0) (0) 12

1 16° | Facing Up. 1 0 0 1

iL 122 Horizontal. 11 1 0 12

2 10° a 10 1 1 12

2 10° | Facing Up. 2 il 0 3

3 10° Horizontal. 12 2 1 15

3 10° o 6 1 0 7

4 5° : 10 1 3 14

4 5° | Facing Up. 3 0) 0 3

4, Ba Horizontal. i 3 3 13

Motalsieees. 74 10 8 92

AMPHIBIAN MIGRATION. 11

TABLE IT,

Incline.| Position. Down. Up. | Neutral. Total. |

8° Horizontal. 4 19 1 24

g° _ 8 33 0 41

8° FS 13 26 4 43

Motalsiisec- 25 78 5 108

The real cause of the Newts normally walking downhill may

lie in the weakness of the limbs. Yet, from a few observations

made on the strong-limbed Toad placed horizontally on an incline,

IT am induced to think that it also usually goes down, if given

the choice. But, as in the case of the Newt, the cause of this’

behaviour of the Toad when placed horizontally on an incline

receives a simple mechanical explanation. The kick given by the

hind leg, which is higher up the incline than the other, would

naturally tend at the moment of the hop to turn the animal down-

wards. The Toad always hops downhill, but it crawls and rarely

hops wp a gradient. On an incline of 12° one of my Toads would

begin hopping downhill. If it was then teased on one side so as

to send it up, the Toad would endeavour to continue hopping

uphill, but when once facing up, the gait very quickly changed

to a crawl. It may be observed that when endeavouring to hop

up, the hind limbs cannot be brought up under the body from a

state of extension unless they scrape the ground. After they have

once scraped the ground in this way, the Toad ceases to attempt

to hop, but crawls. It would appear, therefore, antecedently

probable that in the spring, at the time of their greatest activity,

the general tendency of the Toad also would be downhill towards

water, rather than up.

One more observation is worth recording. Some Newts, when

placed on a level surface or on only a slight incline, would never

start off straight in any direction. They always turned, and some

of their turns were very elaborate. They would meander about

the centre of the board around and up and down, before setting

out in any particular direction. When once started, however,

they went ahead. I regard this elaborate turning merely as

a complex withdrawing response. Similar behaviour has been

noted in the Crayfish *.

All those Newts which winter on the slopes around the pond

could, by their geotactic responses, easily find their way back to

the water in spring. They would simply walk downhill.

* J. Carlton Bell, “Reaetions of the Crayfish,’ Harvard Psychological Studies,

Vol. 11. 1906, p. 615.

12 MR, B. F. CUMMINGS ON

Homine Experiments, 1910.

The first step was to find a pond suitable for experimental

purposes, which was not easy in this neighbourhood where ponds

are not common. Eventually I found a pond, which though

not by any means the. best possible, yet fulfilled sufticiently

the required conditions. It was conveniently near home, it

contained a good number of Palmate Newts, and was small

enough to admit of a thorough dredging. Text-fig. 4 represents

the general nature of the surroundings. The sides of the pond

were bricked up, and there was a thick sediment of mud at

the bottom. The three sides A, B, C, rose perpendicular from

the water for a foot, while at the fourth and remaining side the

water was level with the edge. The depth was 18 inches on an

average, but it varied of course at different states of the weather

and at different periods of the spring season. The area of the

surface-water exposed was 4 sq. metres 60 sq. decimetres.

Yext-fig. 4.

ROAD TALL [HEDGE

cc.

GENTLE INCLINE QB -- GATE

C SILOS (Ul? ===

STEEP GRADIENT DOWN

Bl 1% Pa oft bie

GRASS FIELD

—_— === —- _

Plan showing pond and its surroundings used for homing experiments

Nos. 1-10.

The pool occupied a very sheltered position and was protected

from all winds except those from an easterly direction. For

dredging the pool I used a large prawning net on a long pole.

At first I could not be sure that the pool was efficiently dredged,

as it was impossible to see whether all Newts had been satis-

factorily removed. Subsequently I was in several ways able

to show that the pool could be thoroughly cleared, when necessary,

of all Newts, if the dredging was carefully done.

AMPHIBIAN MIGRATION, ies

In the experiments the method was to collect a number

of Palmate Newts, mark them and set them loose near the pool at

different distances from it. Hach separate batch of Newts was

given its own special mark, so that the animals when recovered

could be immediately assigned to their respective batches. After

several unsuccessful attempts at ringing with fine aluminium wire,

I came to the conclusion that the simplest and best way of

marking them was by amputating the toes of the feet in a variety

of combinations.

The toe was snipped off close to the base with a pair of fine

scissors. In the case of some controls, operated on and kept in

an observation-pan, these animals could be easily distinguished

from unmarked ones even after the lapse of two months, and

this period of time for the purpose of the experiments was ample,

so that I had nothing to fear from regeneration. ‘Two days

after the operation, the Newts were set loose at nightfall on

the ground near the pool. They were always placed heading

towards the pool, and care was taken not to influence the immediate

course of the Newts through the reactions of compensatory head-

movements.

Experiment 1. In this experiment twenty-one Newts were

collected from a large pond 10 miles away from the experimental

pond. This collection was divided into three lots of seven each,

and each batch bore its distinguishing mark, On April 26th:—

Batch A was placed 15 yards from the edge of the pool on the

incline X.

Batch B was placed ,, _ ,, a x a on the

incline Y.

Batch C was placed 10 yards up the steep slope Z.

Result. Next morning none of the marked Newts had suc-

ceeded in finding the water, On April 29th one of Bateh C was

recovered, and one of Batch B on the following day, On May 13th

one of Batch A was taken. Thus only three were recovered.

Experiment 2. A collection of twelve Newts was made from

the pond 10 miles away, marked and treated as usual. On

May 8th six were placed 5 yards away at Y, and six 10 yards

away at Z. These two lots were marked distinguishably one from

another and from all other lots, as in all other experiments with

one exception.

Result. Next day none had found the water. No signs

of them on May 14th. On May 29th one, the only successful

animal, was obtained. It was one of the Batch at Z,

Kxperiment 3. Seven more marked and imported Newts were

placed at X, 5 yards from the pool, on May 26th, One was

recovered on May 29th,

Recorded below are the experiments with Newts collected from

the experimental pool itself and set free in its precincts after the

usual treatment.

14 MR. B. F. CUMMINGS ON

Experiment 4. Fourteen Newts were collected and on April 30th

seven were placed 10 yards away at Z, and seven 5 yards away at Y.

Result. No Newts had returned by the next day, but seven were

recovered on May 4th and four more on May 14th, making 11 out

of the 14. In this experiment I did not take the precaution of

making the two lots distinguishable when marking them, so that

I could not refer the animals recovered to their respective

batches.

Experiment 5. Six natives were collected and set loose on

May 12th, 10 yards away at Z.

Result. None had returned on the ]3th, three were recovered

on the 14th. On May 29th the fourth returned, and on June Ist

the fifth was taken.

Experiment 6. On May 30th six natives were placed at X,

5 yards from the pool.

Result. On June Ist five were recovered,

After this date the approach of summer weather made further

work for the year prejudicial to reliable results.

Homine Experiments, 1911.

I took the precaution of beginning earlier in the spring this

year, and the following few experiments which I was able to carry

out shew that a greater proportion of imported Newts was

recovered.

Experiment 7. On March 26th six natives were set loose, 12

yards away from the pool along the incline at X.

Result. Next day none had returned. Two were caught on

March 29th, one on March 30th, one more on April 5th, and the

fifth, and last to be recovered, on April 9th.

Haperiment 8. On April 2nd six natives were set loose along

the same incline but 24 yards from the pond.

Result. The next day the only successful animal was recovered.

Resuming with imported Newts, the following experiments

were made :—

Experiment 9, On April 2nd six imported Newts were placed

6 yards from the pond, along the incline at X, six more 12 yards

away along the same incline.

Result. The next day none had returned. On April 7th one of

the 6-yards’ lot was obtained and one more on the 12th. Three

of the 12-yards’ batch were obtained, one on April 5th, one on

the 7th, and the third on the 12th.

Experiment 10. On April 16th 24 imported Newts were set

loose in the neighbourhood of the pond in four Batches :—

A. 6 were placed 5 yards along the incline X.

B. 6 ” ” 12 ” ”? oP) ” X.

C. 6 ” ” 24 ” ”? ” ” xe

IDE a » 40 yards up the steep gradient Z.

AMPHIBIAN MIGRATION. 15

Result. On April 18th one of Batch B was caught and two of

Batch A. On April 22nd one more of Batch A was obtained.

Keperiment 11. In this experiment four full-grown Palmate

Newts were taken from a horse-trough about one foot deep near

the experimental pond. ‘The trough was only 2 inches above

ground level. Text-fig. 5 shows the nature of the ground around

this trough and the point (P) at which these four marked Newts

were set loose, together with four young Frogs, also caught in the

trough, the eight specimens being the whole of the amphibian live-

stock which the trough contained. The Frogs were young—not

more than 10 or 11 months old and hatched the previous spring,

judging from their size. These eight specimens were set loose on

April 16th.

Text-fig. 5.

GRASS FIELD

GRASSY HEDGE BANK

TROUGH eg

SS, VOFF Ginches|; ss: 7r.o

‘es

Seare LANE coe

GRASS FIELD SLOPE DOWN

Plan showing the horse-trough and its surroundings used for homing experiment

No. 11.

The spot where they were set loose was deliberately chosen as

the most unfavourable one from which the Newts might be

expected to be able to return. They were set heading down the

slope away from the trough,

Result. The trough remained empty until May 16th, when one

of the four Newts was recovered,

16 MR. B. F. CUMMINGS ON

Houmipiry ExPErimMeEnNts.

In order directly to test Newts for a supposed faculty for

perception of humidity in the atmosphere, I planned apparatus to

work with electric fans driving two air currents which were to be

kept at an even temperature and velocity. One current was to

be thoroughly saturated by passage over a series of porous plates

containing water, and the other to be kept dry. The Newt

to be tested was to walk down an inclined runway, and to be

exposed on one side to the wet current and on the other to the dry.

At the bottom of the runway it was free to move in the direction

of either current. Fortunately for me, before carrying out this

elaborate apparatus in detail, I took the precaution of making a

few of what I considered would prove only preliminary trials

with a simpler arrangement in which the temperature factor was

not eliminated. I quickly formed the opinion either that Newts

were unable to detect moisture at a distance and slight variations

in temperature, or that my method of experimenting was un-

satisfactory.

Although the existence of this faculty—which Romanes

favoured-—seems improbable in the face of the facts recorded in the

two previous sections, an ingenious experimenter might perhaps

obtain more encouraging results. I think, though, such a per-

ceptive power in Newts would prove to be of too slight a nature

to be easily demonstrated by a laboratory experiment, and

especially by one such as mine, where a turn right or left at the

end of a runway, after a comparatively short exposure to the

conditions, was to decide each test. =

DISCUSSION,

The evidence presented, fragmentary as it is, lends support to

the hypothesis that Newts and presumably other Amphibia

possess a homing faculty. Table III, is interesting, particularly

that part dealing with the imported Newts, as one would have

been led to expect more of these to return if only by chance ;

for they were set loose at very short distances from the pond (with

one or two exceptions), But then the small size of the pond must

be remembered, particularly in comparing with the account in

Romanes’ ‘Animal Intelligence’ where no mention of the size of

the pond concerned is given. It will be seen that more imported

Newts got back in 1911 than in 1910, This may be explained

by the fact that experiments in 1911 were begun earlier than

in 1910, when the Newts might have already done some

breeding before being canght. Their desire to find water would

then be proportionately less, This qualification does not apply

to the natives, as the same natives were used several times over

and their breeding was therefore prevented up to May in both

years, Furthermore, in the experiments of 1911, the Newts,

both imported and native, were set loose, with one exception,

along the incline X. This incline was undoubtedly the most

AMPHIBIAN MIGRATION, 17

TaBe ITT,

(a) Imported Newts.

Exp. | Number set tye Distance in yards} Number

1910, loose. Position. | from the pool. recovered.

tL eg 7 xX 15 2

Bh 7 ¥ 15 0)

eee 7 Z 10 1

Fee 6 NG 5 0)

4 ee 6 Z 10 1

a 7 X 5 | 1

1911. |

Ls eee 6 xX 6 2

ee 6 xX 12 3

TO"... 6 xX 6 3

LOR 6 x 12 | 1

Lon. 6 xX 24 (0)

10. 6 Z 40 0

76 14

(b) Native Newts,

} |

| Exp. | Number set aa Distance in yards} Number

1910. loose. Position, from the pool. recovered.

4. 7 Z 10

4. q Y 5 BL

ws. 6 Z 10 5

Gre 6 x 5 5

1911

(he 6 x 6 5

Sine 6 x 24 1

38 27

favourable position for the Newts from which to find the pool.

It was a gentle incline down, and they would naturally walk down

it. On the other hand, if they turned tq the right, they were

confronted with the steep gradient Z which in all probability

they would not negociate. The hedge on the left was the only

thing which may have lured them away from the right path.

In 1910 the number of natives in the pool was twenty-two. In

1911 this supply had fallen to twelve, and two of these still

bore marks, sufficiently clear for me to be able to identify them

as belonging to batches of native Newts set free the prevjous

year. ‘hese two at all events must have wintered near the

pool and returned to it in the spring. The decrease in the

supply of natives in the spring of 1911 may be explained when

I say that seven of the original twenty-two were accidentally

killed before they could be returned to the pond at the close of

Proc. Zoou, Soc.—1912, No. II. 2

18 ON AMPHIBIAN MIGRATION.

the 1910 work. It is probable from this that the number in any

one pond remains fairly constant. It will be noticed that of the

native animals set loose in Experiment 11 only one got back.

These were set loose, however, as will be seen in text-fig. 5, in the

worst possible position I could find from which the Newts—even

natives—might be expected to return. ‘The slope led away down

the field from the pond and the field was separated from the

trough by a drop of several feet and a hard road three yards across.

A decisive result brought out by these homing experiments is

that Newts certainly have no very definite faculty or instinct for

detecting water even from a short distance (see Table JIT.) ; and

in this connection attention may be drawn to the length of time

usually taken by the native Newts in finding water from short

distances. They perhaps rely on a combination of small clues,

for, in addition to the downhill tendency and a small homing

faculty, a supposed sensitiveness to atmospheric moisture at a

distance and to slight variations in temperature, I can suggest as

auxiliary factors (but can adduce no evidence for or against) :—a

sense of smell for detecting stagnant water and a sense of hearing

for discovering other Amphibia already in the water, or, in those

Amphibia which live in streams, for discovering running water.

Either all or a few of these may provide the animals with

clues, small though some of them would be. In this paper a com-

bination of the downhill tendency and a homing faculty is

considered as the chief factor.

I was very anxious to make more experiments with natives,

but the stock became exhausted and this brought my work to a

premature close.

CONCLUSION.

This paper does not pretend to present a conclusive case on

the subject of amphibian migration. But the results of the

experiments are suggestive, and the opinions I have formed

from them are :—

1. That Newts are helped to find water in which to breed

by their marked tendency to walk downhill. Those animals

hibernating on the slopes around their pond would simply have

to walk downhill. A good many young Newts I have found

hibernating in this way, close to the water of their native pond.

It may be a wise precaution, because young Newts of the year

would not have any knowledge of locality.

2. That there is a small homing faculty in Newts, which

probably consists in a topographical knowledge of the locality

where the breeding pond is situated.

3. That they are able to ‘““home” from only short distances

and that, therefore, both their topographical knowledge and their

terrestrial wanderings in autumn and winter must be rather

circumscribed.

4, That, speaking generally, the same Newts return to the same

pond each year.

5. That those Newts which wander away from the precincts of

ON TILE HABITS OF BRITISH FROGS AND TOADS. 19

the pool, become “lost” and only regain the water by chance or,

for a season, may even fail altogether to reach water and to

propagate their kind. Field naturalists are well acquainted with

the Newt’s capacities for travelling.

6. That further experiments on a more extensive scale ought to

settle this matter; suitable ponds, however, are difficult to find,

particularly in this part of the country.

I hope those living in more favoured localities will be induced

to carry these preliminaries to more decisive results.

In conclusion, I should like to thank Dr. R. M, Yerkes, of

Harvard University, for the encouragement he has given me in

this undertaking, and my sister for her seryices in obtaining

Specimens and in dredging.

2. Some Remarks on the Habits of British Frogs and Toads,

with reference to Mr. Cummings’s recent communication

on Distant Orientation in Amphibia. By Gree A

Bouencsr, F.R.S., F.Z.8,

[Received November 6, 1911; Read November 21, 1911.]

In the course of the discussion which followed Mr. Cummings’s

communication at the Meeting on October 24, I made some

remarks on the habits of Batrachians, which the Secretary has

asked me to embody in a short paper for the information of those

who might feel inclined to carry on further observations in this

country on the interesting subject with which Mr, Cammings

has dealt.

There is great diversity among Tailless Batrachians in the

possession, lack, or degree of instinctive foresight in selecting

places for the deposition of their eggs. Leaving out the

thoroughly aquatic Edible Frog (Rana esculenta), which has only

doubtful claims to be regarded as British, two of our species,

the Common Frog (Rana temporaria) and the Natterjack Toad

(Bufo calamita), belong to the category of the least gifted in this

respect, whilst the third, the Common Toad (Bufo vulgaris),

behaves differently, and is besides possessed of an instinct for

orientation which appears to be highly developed. The fact that,

after the perfect condition is attained, Tailless Batrachians,

unlike the Tailed, do not regenerate lost parts, would render the

Common Toad further suitable for experiments on migrations,

after the amputation of a finger or toe, as a distinctive mark.

The Natterjack is, with us, a very local Species, but the Common

Toad and the Common Frog are ubiquitous and, as young or

outside the breeding-season, are often found far from water, to

which they have, of course, to resort for oviposition, How the

three species behave in connection with this function I will tell

briefly from my own observations.

The Common Frog exercises little discrimination in the choice

20 MR. G. A. BOULENGER ON THE

ne)

often spawning in temporary winter pools which are sure to dry

up before the time at which the larve have completed their

metamorphosis, or on the edge of a pond where, the level of the

water falling after dry weather, the egg-clumps remain stranded.

No doubt the frogs are attracted to the shallow, slanting borders

in preference to the deeper parts with abrupt banks by the

comparative warmth of the water under the sun-rays of late

winter or early spring, with the unfortunate result which may be

witnessed almost year after year in some places, when, following

a spell of dry weather, the borders of ponds are covered with the

stranded jelly-like masses of frog-spawn drying away in the sun

before the liberation of the larve. On my country rambles in

the end of March or beginning of April, I have often rescued

the progeny of many frogs by removing such doomed egg-masses

from these fatal beaches to deeper water close by, which the

mother would have chosen had she been gifted with the instinct

possessed by the Common Toad.

The migrations of the Common Frog towards the water must

take place, to a great extent, in the autumn, as many, perhaps

the majority of individuals, hibernate buried deep in the mud at

the bottom of ponds, sluggish but not dormant as in some higher

animals. Some years ago, just before Christmas, after a period

of severe frost, which had lasted more than a week, I was walking

on the ice of a small and shallow pond in Belgium, when my

attention was drawn to a bright red creature, first taken for a

gold-fish, moving under the ice, which very nearly extended to

the bottom of the pond ; this was a Common Frog, and on looking

closer, | discovered hundreds of others, less conspicuous owing to

their yellowish, brown, or olivecolour. Some were very handsome

specimens, which I wished to secure. A pick-axe was fetched

from a neighbouring farm and a hole made in the ice, through

which I could introduce my arm and reach the bottom ; but the

frogs were so active that they swam away and not one could be

caught. I decided to return to the pond after the thaw, which

I did, but no more frogs were to be seen; they had evidently

retired to their hibernating-quarters in the mud and in holes

under water. The reason why they came out when the pond was

frozen nearly to the bottom, must be attributed to the reduced

oxygen in the water, which made the frogs, breathing as they do

under such conditions chiefly by the skin, feel uncomfortable and

desire to escape.

The Common Frog is the earliest breeder among European

Batrachians, spawning taking place in the South of England

from between the end of January and the end of March, the date

depending almost entirely on the temperature and _ therefore

varying considerably from year to year. Should the winter be a

mild one, the breeding may be over by the middle of February ;

recurrence of cold weather after the frogs have begun spawning

may cause them to return to their winter-quarters, with the

of water in which to deposit its eggs, provided it be not brackish,

HABITS OF BRITISH FROGS AND TOADS, 21

result that the breeding-season is broken up into two or three

periods within a couple of months. When the first appearance

of mild weather is much retarded, the spawning takes place for

all individuals within a very few days, with an ensemble which is

not customary for the species in our uncertain climate.

The Narrersack Toap does not undertake lengthy migrations

for the purpose of depositing its eggs. It usually lives in

colonies in sandy localities, such as dunes on the sea-coast, sand-

quarries, heath-land, etc., which must be in close proximity to

at least some sort of shallow pool, even of an intermittent kind.

This toad shows not the slightest discrimination in the choice

of water, and will not move any distance in search of better

conditions if only it can find close by a little water in side ditches

ov even in cart-ruts of a road, whilst an excellent pond may be

available a hundred yards off. ‘The development of the eggs and

larve, itis true, is much more rapid than that of earlier fire eeders

like the Common Frog and the Common Toad, but nevertheless,

in some places, a great many broods are destroyed through want of

foresight on the part of the mother. Should, however, the pools or

ponds in which the Natterjack is accustomed to breed, and around

which it has settled, disappear for good, by the agency of man

or otherwise, the whole colony will move off after a time to a

more suitable locality. This I have observed in sand-quarries 1n

Belgium and France, some of which are provided with stagnant

water whilst others are not; those which have no water or from

which the water has disappeared for a year or two, are without

Natterjacks at all times of the year, thus showing that this

gregarious Batrachian only settles down in such places as will

afford a site for spawning within a few yards, whilst half a mile’s

journey is nothing to aCommon Toad. It may also be mentioned

that the Natterjack, in common with the Palmated Newt, has no

objection to brackish water and is therefore often found in great

abundance in close proximity to the sea. The greatest number I

have ever seen was on the little tle de Bast, opposite Roscoff, on

which, for the reason that the pools are brackish, neither frogs

nor the common toad exist.

The Natterjack has no such restricted breeding-season as our

other Tailless Batrachians. The females do not go to the water

until ready to spawn, and the time at which the ova descend into

the oviduects varies according to individuals, from between the

middle of April to the beginning of July, sometimes even later.

The Common Toap, on the other hand, is remarkable for the

fixity in the time at which it seeks the water for the purpose of

reproduction and for the shortness of the period within which all

the females get rid of their eggs. Exceptions of later breeders,

which have been observed, are so rare they may well be said to

confirm the rule. The same may be said of isolated pairs

occasionally found in places to-which toads are not in the habit

of resorting to: breed.

The breeding-season in the South of England, the North of

22 ON THE HABITS OF BRITISH FROGS AND TOADS.

France, and Belgium, falls within the last days of March and

the first days of April, and, with the exception of severe frosty

weather, which rarely occurs at that season, irrespective of the

temperature. Then certain ponds or deep flooded quarries will

be found alive with hundreds or thousands of toads which have

congregated from the neighbourhood, often from a radius of half a

mile or more. Thither all the toads have travelled with remark-

able directness, passing other ponds or ditches of which they

might avail themselves were it not for the instinct which leads

them to seleet a place offering all guarantees for the successful

rearing of their progeny. If a male meets a female en route, he

vigorously clasps her under the axils and accomplishes the rest of

the journey on her back. Much of the travelling takes place at

night, but individuals are also met with in the daytime, hopping

along towards the meeting-place. If a high road should run not

far from the pond which is the rendezvous, numbers of crushed

corpses of toads, run over by motor-cars or other vehicles, will be

found for a distance of perhaps two or three hundred yards, the

crushed toads being more and more numerous as the road nears

the pond.

If pairing toads are taken from the place selected by them for

spawning, and removed to a neighbouring garden with a small

artificial pond in which, we should think, they might comfortably

conclude their breeding-operations, they will often leave and

start off in the direction whence they were brought. It does not

matter to them whether this be up or down hill.

Some years ago I made an experiment on the instinct of

orientation in this toad. I took a number of pairing individuals

out of a pond frequented by the species, which was only a short

distance from another in which frogs spawn but to which toads

never resort. I turned them loose on a monticule midway between

the two ponds, from which neither could be seen, and watched

their movements. All, after a little hesitation or after a few

hops in the opposite direction, took the right orientation and

made their way straight towards the pond whence they had

been taken. I experimented on single individuals, on pairs, and

on groups of individuals, with the same result. In this case, it

was evident that the toads were not influenced by hygroscopic

sensations, since there was water in both directions. Whether

the sounds uttered by their fellows in the pond were a guidance

to them, seems to me doubtful, considering the very feeble

voice of the Common Toad, the males of which, as is well known,

are devoid of vocal sacs; if so, it would denote a very acute

sense of hearing in toads. J am, however, conviuced, from other

observations, that even at a greater distance, from which no

such sounds could be heard, the-toads would have taken the

right direction.

I strongly recommend the Common Toad as the most suitable

Batrachian on which to institute series of experiments on

distant orientation.

ON GAME PROTECTION IN INDIA. 23

3 Game Sanctuaries and Game Protection in India.

By BE. P. Sressine, F.L.S., F.Z.8., E.BR.G.S.

[Received June 6, 1911: Read October 24, 1911.]

I. Inrropuctory REMARKS.

Could some of the great sportsmen who lived and revelled in

the land half a century ago revisit the scenes of their former

exploits, how, it may be wondered, would their favourite shooting-

grounds strike them at the present day ¢

We have all read our Forsyth’s ‘ Highlands of Central India,’

Sterndale’s ‘Seonee or Camp Life in the Satpuras,’ Saunderson’s

‘Thirteen Years’ Sport in India, Simson’s ‘Sport in Eastern

Bengal,’ Pollok’s ‘Sport in British Burma,’ Pollok and Thom’s

‘Wild Sports of Burma and Assam,’ Pollok’s ‘ Sporting Days in

Southern India,’ Kinloch’s ‘ Large Game Shooting in Thibet, the

Himalayas and Central India,’ Col. Fife-Cookson’s ‘ Tiger Shooting

in the Dun and Alwar,’ Baker's ‘ Wild Beasts and their Ways,’

Eardley-Wilmot’s ‘ Forest Life and Sport in India,’ ete. What

lesson does a perusal of these fascinating volumes teach us—

inevitably tell us? That the Game of India is on the decrease

and on a very rapid decrease, and that the good old days of yore

are gone, never to return.

That the modern rifle has to some extent been responsible

for the present state of affairs is beyond cavil—its accuracy

and also the cheapness with which the more roughly made

forms can be purchased. The native shikari has now to some

extent replaced the old blunderbuss of his father’s days by a

breech-loader, and when possessed of such kills an infinitely larger

head of game in the yearas a consequence. The weapon itself costs

Rupees 45 only, but it is doubtless the price of rartridges which

mercifully prevents the breech-loader from coming into as general

use amongst this class of men as would otherwise be the case.

But the startling decrease which the head of game existing in

India has undergone during the last two or three decades

cannot be attributed only to the improved accuracy of the

weapons with which the modern-day sportsman is armed. The

opening out of the country and the consequent restriction of

the animals is largely responsible. For instance, Bengal and

Assam, e. g. the Western Duars, no longer contain sufficiently

extensive jungles to harbour rhinoceros and buffalo. The great

increase’ in the number of sportsmen who visit the jungles

annually on sport intent, an increase brought about chiefly by

the greatly improved communications owing to railway and road

development, has also been a great factor in the case ; and, finally,

the infinitely greater number of competent native shikaris in

existence; 1 write “competent” in the sense merely to express

their power to hill game, The vast majority of these men

are poachers pure and simple, as were their fathers and fathers

24 MR. E. P. STEBBING ON

fathers before them. Formerly, however, owing to their anti-

quated low-power weapons, the damage they were capable of doing

was of a negligible quantity: nowadays it is far otherwise, and

the methods to be put in force to deal with them form one of the

most difficult problems those responsible for the upkeep of the

game in the forests, and country generally, have to solve.

The plea ever placed in the forefront by such men is that the

guns are required to protect their crops, and this plausible excuse

has been accepted by Local Government after Local Government ;

and we can quite see the difficulties that have faced the latter, and

still do so, in a settlement of the question. It cannot, however,

be said to have been ever satisfactorily or fairly faced, and this

inaction on the part of the central authority has checkmated the

efforts of many a Collector and Forest Officer in his attempts

to keep down the number of (poaching) guns in a district. A

sympathetic Government has always been too eager to listen to

the tales of destruction to crops, and the District Officer, without

local knowledge, prefers to err on the side of liberality, and so

readily grants licenses to applicants.

We all know the way these license gun-holders go to work. A

machan (platform) is built on a krown deer-run on the edge of the

forest and just without its boundary, if not inside, with the con-

nivance of the Forest Guard. The shikari occuptes his post in the

late afternoon, and by sunrise next day several bucks and does

may be lying round the machan ; the skins, horns, should there be

any of the latter, and the flesh are taken off to the bazaar, where

a ready sale is found for them throughout the country. The meat

is sold locally, the skins and horns being bought by middlemen

for export. It is a common thing to see on the platform at way-

side stations near forest-areas piles of skins and horns booked,

and openly booked, in defiance of all rules and regulations, to some

large centre.

I would not be understood to say that it is the native shikari

alone who acts in this way. It is an open secret that the native

soldier of shikar-loving propensities, as also his British brother,

will act in an exactly similar manner on occasion. Once, however,

this matter 1s properly faced, the latter class of offenders can

easily be coped with. For the non-military native offender a

license to protect his crops should be given only after careful

personal enquiry on the ground by the District Officer. Also the

sale of venison in the open market should be made a criminal

offence.

The whole crux of the position is, of course, the necessity for

regulating the number of animals shot, so as to prevent deterio-

ration or extermination of the game. On the whole, the European

is more destructive than the native to the animals which are

greatest in need of protection. Rules and the proper control

and management of shooting-grounds can control the European.

The native is, however, not so easily dealt with. In order

therefore to arrest the slaughter which takes place ostensibly to

GAME PROTECTION IN INDIA. 95

protect crops, some special measures are necessary. Wherever

it can be proved that game is no longer destructive, the licenses

should be cancelled and the weapons called in. In other cases

where destruction is still being done the guns must be retained.

Since, however, these weapons are given merely for the protection

of the crops, they should be restricted to that purpose and be

rendered unfit for any other. This can be easily done by cutting

down the gun-barrel to 18 inches or 2 feet:

A further point of importance in connection with the pro-

tection of game is to be found in the sale of ammunition. It is

a question for consideration whether this should not come into

the hands of Government and be directly regulated by it. This

question scarcely comes tnder the head of Game Sanctuaries,

but it is of very considerable importance in the interests of

Game Preservation.

The apathy whieh the proper protection of the Game of the

country has met with in the past at the hands of Government is

almost incredible. In many parts of the counti'y there are forests

in blocks of very considetable area. Rules under the Forest Act

have been in force in these forests for years, a sixth of British

India being under the Act and Rules. It would have been

sufficient merely to have enforeed these rules in the spirit as also

in the letter; and adequate protection would have been afforded

to species which are now, owing to this apathy and neglect,

within a measurable space of deterioration, if not of total

extinction.

Lord Curgon, so long ago as December 1901, when replying to

the Burma Game Preservation Association at Rangoon, said he

was in ‘close sympathy ” with the aims of that body.

That he recognised the importance of the subject is shown by

his opening remarks, in which he said:—‘‘ Among the many

memorials which the enterprising inhabitants of Lower Burma

have showered tpon me, and to all of which it has not been

possible for me to give a verbal reply, I have selected yours as

one of those to whith I should.” He then observed that the

great importance of the question of Game Preservation in India

is one that, in his judgment, appealed not only to the sportsman

but also to the naturalist and the friend of animal life.

Further on he stated that facts pointed entirely in the direction

of the “ progressive diminution of wild life in India,” and gave

examples in support of this belief. He then mentioned some of

the ‘artificial and preventable” causes of this diminution, and

also of those which are “ natural and inevitable.” He admitted

that hitherto the attempts made by Government to deal with

the question by legislation or by rules or notifications based on

statutes had been somewhat “ fitful and lacking in method,” and,

after mentioning some of these attempts, he said :—‘‘'The general

effect of these restrictions has been in the right direction, but I

doubt if they have been sufficiently co-ordinated, or if they have

gone far enough, and one of my last acts in Simla; before I had

26 MR. E. P. STEBBING ON

received or read your memorial, was to invite a re-examination

of the subject, with the view of deciding whether we might not

proceed somewhat further than we have already done.”

His idea was ‘to frame some kind of legislation of a permissive

and elastic nature, the provisions of which should be applied to

the various provinces of India, in so far only as they were adapted

to local conditions.” Unfortunately, however, he did not deal

with one of the most difficult sides of the question, viz., the co-

operation of Native States ; he merely observed that this question

is ‘‘ somewhat complicated,” and expressed his belief that ‘ The

Government would meet with the willing co-operation of the

Chiefs.”

There does not appear to be at the present moment, notwith-

standing the Viceroy’s action, so far back as 1901, any Act or

Regulation issued by the Government of India, defining the

principles governing the protection of game, and the means by

which such protection should be effected.

Of late, however, matters for the protection of game have to

some extent considerably improved, and Local Governments

throughout the country have revised their Game Rules, and in

some cases have ordered the formation of Game Sanctuaries in

addition to limiting the number of head of game to be shot in a

district or block of forest to a definite number per year. Further, in

certain provinces sportsmen are only allowed to kill individually

a certain head of each different species of animal, thus elimi-

nating the worst feature of the old-time sportsmen—the butcher,

whose boast was not the size of the trophies he obtained so much

as the nwmber of animals he had killed. For the departures

thus made throughout the country I think a due meed of credit

should be accerded to the Nilgiri Game Association. Inaugurated

about 1885, this Association has now for years not only

protected the game of the Plateau which the sportsmen and

the Todas between them were surely exterminating, but has

enabled an increase to be maintained and recorded. The last

annual reports of the Association point to a satisfactory increase

in the head of Saddlebacks (Hemitragus hylocrius) and the

Sambhar (Cervus wiicolor). For some years past the number of

such to be shot by each sportsman has been regulated under the

authority of the Association, directly supported by Government.

The departure thus initiated in the distant Southern Plateau

was followed in the far North when the game of Kashmir was

threatened with extinction owing to the annually large recurring

influx of sportsmen who visited the Fair Vale. Game Protection

in Kashmir now forms a separate Department of the State, as, I

think, it should in British India, and one which has fully achieved

under its able head the objects anticipated from its inauguration.

The enlightened ruler of Chamba State also took up the question,

and prohibited all shooting except on passes issued on his own

authority.

Whilst such laudable commencements were thus made to

GAME PROTECTION IN INDIA, 27

preserve the game of areas which, owing to their peculiarly

favourable climatic conditions for the European sportsman, were

threatened with extinction, the Local Governments in India were

very apathetic in the matter. Game Rules were in existence

for the Forest Reserves of the country, but they related chiefly to

a close season, the latter in some cases only applicable to the

females, and the same was the case for the open country, where the

rules usually related to birds only. These regulations were, how-

ever, openly broken, and the penalties in existence were practically

rarely put into foree, except by some exceptionally energetic

officer ; and even then an appeal was usually upheld and the

orders passed reversed.

At length, however, the apathy that hung over this question

gave place to some show of interest, which was followed by

activity on the part of the Government of India, on. whom it had

slowly dawned that there was perhaps some reason for the outcry,

increasing in intensity each year, that the game of the country

was doomed and that but a few years separated it from ex-

tinction. Local Administrations were addressed on the subject: of

the Rules and Regulations in force in their Presidencies and

Provinces under the Forest and other Acts, and as to the steps

necessary to be taken to prevent the extinction of the several

heads of game, excluding carnivora, This led to many separate

enquiries being undertaken throughout the country, to a prolific

correspondence in the Press, of which desultory rumblings are

still heard, and to many improvements being initiated in the

Shooting and Game Rules throughout India. Jam aware that

I am laying myself open to serious attack in thus stating the

case, but it is maintained that any and every rule that is made

with the idea of protecting the game of a country is a step in the

right direction, and therefore advantageous both to the sportsman

and the game itself, however hard it may seem to fall on a

particular body of individuals or a particular individual, This

point will be referred to at some length later on, but I would

like to appeal here to the public spirit and fairness of all true

sportsmen to aid to the best of their abilities the present

endeavours of the Government of India in the direction of

arresting the destruction of Wild Animals. I feel sure that the

Government will approach this matter in a much firmer spirit if

they feel that they have behind them the good wishes of a solid

mass of sportsmen throughout the country.

What is required is to fix the close seasons definitely, and the

Government of India have now, as we shall see, proposed to

legislate to give power to fix a close season for different kinds

of game.

In a series of articles published in the ‘ Indian Field’ between

the dates 6th June, 1907, and August 1907, a writer thus

approached this question :—

‘“We presume that the Imperial Government, as proprietor of

all the game in the country, is the most important party concerned

28 MR. E. P. STEBBING ON

in its preservation, and that in effecting this object it has every

right not only to safeguard this valuable asset, but also to obtain

a reasonable profit therefrom. Less than half a century ago

this principle was acted upon when forest lands were taken

possession of, and now the villager has to pay for every bundle

of grass and bamboo that he obtains from Government forests

for building his humble abode.” This latter, by the way, is not

a true representation of the actual facts of the case, for many

forests are burdened with “rights,” under which large amounts

of forest products are annually given away free to villagers by

the Government.

“The agricultural community is certainly the next important

body whose interests will be affected. In European countries, in

England for example, the population have a firm conviction that

the right of killmg game belongs to them, contrary to the

practice of centuries. In India, no such difficulties surround the

game-protection question. The occupiers of the soil have no

conviction whatever that they possess the right of property in

the game animals and birds that abound in their fields and

jungles. They are coneerned only about the protection of their

crops from the raids of these animals. If this right of protection

is recognised, and the Government which assumes ownership of

all game acknowledges its responsibility in this respect, it can be

confidently stated that no apprehension need be felt regarding

the attitude of the Indian rural population towards the proposed

law. Legislation for the protection of game in this country

will not create any feeling comparable to that which still exists

in the minds of the people regarding the action taken by

Government when forest lands were resumed fifty years ago.”

The analogy, it may be pointed out, scarcely holds, since the forests

of the country are protected solely in the interests of the people

themselves and posterity, whilst the protection of game is under-

taken in the interests of the sportsman and in that of science.

“The third class which will be affected by this law is sportsmen,

European and Indian. In the former category should be placed,

first, the official who is backed by Government authority and his

favoured friends who are given all the facilities which his position

allows; next, the solitary hunter with his small camp and all

the Game Regulations of the Province to keep him in the right

path. The subdivisions in this class are numerous, and include

the unemployed colonel putting in his time, the subaltern out

on ten days’ casual leave, the humble clerk who can occasionally

get away from his office for a week, and the British soldier let

lose in parties for fixed periods. .

“Next comes the Indian shikari, who is also numerously sub-

divided. There is the rich land-owner, possessed of unlimited

sporting appliances and his host of followers; the professional

game-killer who lives by his licensed gun; the small landholder

who does not know how to use a firearm, but whose dignity is

enhanced by the possession of weapons,—freely lent to his

GAME PROTECTION IN INDIA. 29

underlings for supplying his kitchen with fresh meat; and,

finally, the half-starved villager of the meaner castes, who is

given a gun for use at night in the fields, but which weapon

is regular ly and illegally used for the destruction of game animals

of every prohibited class. In this inventory, the Gurkha soldier

does not find a place, for he belongs to a class which he amply

fills by himself with his small but very important personality.

He deserves separate notice. From the banks of the Sarda on

the frontier of Nepal, to the banks of the Indus, the battalions of

these gallant little men are scattered in cantonments all along

the outer spurs of the Himalayan range. In seven or eight of

these locations there are at least fourteen thousand of these

disciplined warriors, who, in the absence of opportunities for

spilling human blood legitimately, are given a free hand for

slaughtering wild animals, along five hundred miles of the best

hunting erounds i in Upper India.”

I propose, then, to consider first the question of the formation

of Game Sanctuaries, defining a Game Sanctuary and the dif-

ferent ways of forming them, and detailing what has been done

under this head in the several Presidencies and Provinces of the

country and what in my opinion it would seem still remains to

be done.

I shall then give an abstract of portions of the proposed new

Indian Game Act, making some suggestions for specifying more

distinctly than the Act does at present the various classes of

game, and suggesting that the game animals and birds, as also

useful insectivorous birds, shall be severally mentioned in the

Act by name. I shall then indicate some specific suggestions

for the formation of close seasons and for the closing to shooting

of species which have been decimated by drought, anthrax, etc.,

and for the definite regulation from year to year of the number

of head of particular species toa be shot in given areas; concluding,

finally, with some remarks on the subject of the proportion of the

head of game of a district to be shot respectively by the district

officials and the outside sportsman.

II. Tae Game SANCTUARY.

The idea of the Game Sanetuary was a natural outcome of

the indiscriminate slaughter to which wild animals have at all

times and in all countries been subjected by man, So long as it

was man imperfectly armed against the animal with his natural

sagacity or fierceness to protect him, conditions were equal, or

in favour of the animal, and there was no reason for intervention.

From the day, however, of the introduction of the breech-loader

and the repeater and a whole host of perfectly built weapons of

every kind, enabling man to kill with comparative ease and

certainty, the odds were against the animal and the question of

affording some degree of protection to the game of a country

became of paramount importance; and, curiously enough, the

30 MR. E. P. STEBBING ON

question became most vital in the more uncivilized, uninhabited,

and wilder portions of the globe. Such shooting ‘grounds were

open to one and all, just as for centuries the shooting in India had

been open, with the result that the modern rifle soon threatened

the extinction of all game. That modern conditions have rendered

this quite feasible the two well-known and oft-quoted instances

afforded by the practically extinct American bison and the

extinct quagga of South Africa sufficiently illustrate.

In India we have come within measurable distance of exter-

minating the rhinoceros (Rhinoceros unicornis), which, together

with the elephant and the gaur or Indian bison (os gaurus),

would without protection probably soon disappear from the

jungles which have known them for so long.

It may be mentioned here, in passing, that Indian ruminants,

such as the gaur, sambhar, etc., and the elephant, have always

had to contend against a sickness resembling anthrax, and also

against drought, which at times have decimated their numbers

over large tracts of country.

With a view to affording a certain protection to animals

of this kind and of giving a rest to species which have been

heavily thinned in a district by indiscriminate shooting in

the past or by anthrax, drought, etc., the idea of the Game

Sanctuary was introduced into India (and in other parts of the

world) and has been accepted in many parts of the country. The

sanctuary consists of a block of country, either of forest or grass-

land, ete., depending upon the nature of the animal to which

sanctuary is required to be given; the area has rough boundaries

such as roads, fire lines, nullahs, etc., assigned to it, and no

shooting of any kind is allowed in it if it is a sanctuary pure

and simple; or the shooting of carnivora may be permitted, or of

these latter and of everything else save certain specified animals.

Sanctuaries may be formed in two ways :—

I. The area is automatically closed and re-opened for certain

definate periods of years.

Il. The area is closed until the head of game has become

satisfactory, and the shooting on the area is then regulated, no

further closing taking place, save for exceptional circumstances.

I. The Sanctuary is automatically closed and re-opened for a

definite period of years.—The Sanctuary is notified for a period of

years: this period would naturally be variable, but it is of import-

ance, I think, that it should not be placed at too great a length, or the

animals in the sanctuary, so long immune from danger, ona on

the re-opening of the area be so unused to the sporteman that

they would be shot down in avery short space of time. Probably

the period during which a block of forest is closed to all shooting

should never exceed, at the most, three years. That good sportsman

Sir John Hewett considers that a period of five years for a sanctuary

is too long, He thinks that the ground of the sanctuary should

be changed every two or three years, probably the former, and

GAME PROTECTION IN INDIA. wil

that the animals would soon learn where the sanctuary was. He

also agrees that before opening a sanctuary to sportsmen the

area should be beaten through so as to distribute and disperse the

game, and not have them collected together Noah’s-ark-fashion on

« large scale for the first permit-holder who enters to shoot down

with ease.

Whilst, however, this system of opening and closing areas to

shooting is best adapted to some localities and to certain classes

of game, it is quite inadequate for the satisfactory protection of

others. In many parts of India I would favour the second

suggestion as being by far the most satisfactory in the long run

and in some cases essential.

Il. The area is closed until the head of game has become

satisfactory, and the shooting on the urea is then definitely

regulated, no further periods of closure being enforced save for

exceptional circumstances,

The length of time a Sanctuary should be in existence is of

very considerable importance, and to a certain extent is intimately

dependent upon a knowledge of the habits of the animals for

which the sanctuary is formed. The period of closure to be

effective must depend :—

(1) On the condition of the head of game of the area when the

sanctuary is first formed.

(2) On the nature of the animal, e. g., the rhinoceros, with a period

of gestation of two years and a period of fifteen years before it

reaches maturity, would require practically permanent closure of

its haunts to produce any appreciable result, as has, in effect, been

carried out in Goalpara in Assam,

The procedure followed should usually be determined by the

condition of the head of game on an area. There would be no

question of fixing a definite period for the sanctuary in the first

instance. When the requisite effect on the game had resulted

from its formation, careful and efficient rules and management

should be sufficient to keep up the head of game, and it would

not be necessary to continue the rigid exclusion of sportsmen.

It would be sufficient to limit the number of head of each species

to be shot each year, as is done in many parts of the Central

Provinces. When the limit had been reached the shooting of

that species in that locality would cease for the year,

Once a sufficient head of game has been established in a

locality, it is questionable whether regulated shooting each year

would not have a better effect than the alternative proposal of

closure for a term followed by a period of unchecked shooting.

It would certainly minimise the chance of the animals becoming

too tame.

The size of a Sanctuary must, of course, entirely depend on local

conditions and on the nature of the animals to be protected.

Such animals as the rhinoceros or gaur, which are of an extremely

shy disposition and are given to roaming considerable distances,

would require an area of considerable dimensions, whereas chital

oy MR. E. P. STEBBING ON

(Cervus axis) and hog-deer (Cervus porcinus) would require a

comparatively small one.

Pheasants, again, would not require large areas, and the same

applies to the hill sheep and goats—a nullah or certain nullahs

being prescribed as closed to shooting, as, in fact, is done in

Kashmir. :

Game Sanctuaries may be of several kinds :—

1. Entirely closed to all shooting.

2. Closed to beating only.

3. Closed to the shooting of certain species of game.

4. Closed to shooting of all game, save noxious ones, such as

carnivora, pig, etc.

The question of enforcing the sanctuary law against shooting

is one of some difficulty. In Reserved Forests it is comparatively

easy, Since all shooting without special passes in such areas is for-

bidden and the granting of these would be stopped for sanctuaries.

Outside, however, the matter is by no means so simple, and

the people of the country, particularly the shooting element,

will require a careful education if they are to understand and

respect the sanctuary, should it be formed in Governmeut Waste

Land. It will be necessary to fully explain the uses of sanctuaries,

and the reason for closing the areas as soon as attempts have been

made to form them.

At present anyone may enter on land, which is not reserved

forest, and shoot. ‘To alter this would at once curtail what is a

prescriptive right, and this is the main obstacle to the introduction

of a Game Law. Rich and poor alike enjoy this privilege, and

although the occupier may in time come to learn that shooting

rents can add to his income, or reserve his waste land for his own

shooting and close it to the general public, as is done in many

cases in the Dun below the Mussoorie Hills, it will be difficult to

introduce restrictions on areas in which shooting is practically a

right in all but name,

It is, we fear, hardly to be expected that the question of the

formation of sanctuaries and their closing will be received without

apposition throughout the country even amongst the Europeans,

but I am of opinion that the matter is one of such great

importance that the outcry of the few interested people opposed

frem personal motives to their formation on Government Land,

both Reserve Forest and Waste Land, should not be allowed to

blind the public generally to their immense value. It is con-

ceivable that the Zemindar and large landed private proprietors

would in course of time follow an example so set when its value

made itself apparent to them.

It has been said that some Provinces, as also Native States,

have already taken up and dealt with this matter, and it will be

interesting here to see, province by province, exactly what progress

has been made towards carrying out the policy of game protection

hy this means.

GAME PROTECTION IN INDIA. ap

IVI. Tar Existing GAME SANCTUARIES IN THE Country.

A. Bririsa Inpra.

1 take British India first, and deal with Forest Areas only,

which are practically the only areas in which Sanctuaries for

game have been formed.

1. The North-West Frontier Province.

The only Forest Division in the North-West Frontier Province

is that of Hazara, and the Deputy Conservator of Forests in

charge says that no Gaine Sanctuaries have been formed in that

Division.

2. Punjab.

A number of Game Sanctuaries have been formed in the

Punjab, most of the Divisions containing one or more, in which

the shooting of certain game is prohibited. Throughout the

Punjab a close season is in force in all Reserved and Protected

Forests for all game from March 15th to September 15th every

year.

; Rawalpindi. The Lalachitt, a Reserve, has an area of 93,000

acres, Half of this is closed alternately as a Game Sanctuary for

a period of three years. The Barakao plantation of 200 acres is

entirely closed to all shooting throughout the year.

Jhelam. In the Jhelam Division there are six sanctuaries

entirely closed to shooting. These comprise the Reserves known

as Rakb Ara Jadid (12,172 acres), Rakb Samarkand (south, 9196

acres), Rakb Phadial (10,722 acres), Rakb illa (24,314 acres),

takb Nili (south, 17,114 acres), and Rakb Bazza (3742 acres).

Chenab. About 15,000 acres of the Pabbi Reserve and 525

acres of the Thatla Fagirullah Reserve are closed to all shooting.

Kangra. This is a Division with much scattered forest, and

a large number of small areas have been made into Game Sanc-

tuaries. A number of these have been closed for forestry reasons

rather than directly for the protection of game. However, the

results are the same, as they become de facto Game Sanctuaries.

The game to be protected varies from the hill sheep and goats,

oorial, serow, gural, etc., to pheasants, kalig, chukor partridge,

kakar (Cervulus muntjac), etc. The following is a list of these

sanctuary blocks, with their areas in acres :—

Dharamtal (634), Khanni (698), Tatal (1228), Chatril (1228),

Mehdhar (367), Balnhi (312), Swarka (47), Bhali and Dolba

(1266), Sildah (266), Ballah (371), Talava Gurial (788), Kalak

(256), Mastgarh (240), Tilli (1013), Darang (558), Habrol (480),

Gumar (400), Khardwar (123), Lohara (2580), Punjab (3610),

Dharni (2822), Karnpur (3321), Bindraban (2946), Bohan (250),

Phakhloh (55), Chagrin and Bhernal (285), Tather (60), Chowkid-

harin (61), Sariali (61), Tiri (52), Nobian (179), Khalig (56),

Jamli (46), Khamahar (378), Baldoa (565), Lohjang (1082), Sam-

lian (537), Tiamal (135), Bakarar (455), Behan (127), Naum (257),

Proc. Zoou, Soc.—1912, No. ITI. 3

34 MR. E. P. STEBBING ON

Santhla (403), Saleti (83), Aloh I and IT (62), Kandbari (5066),

Soupdhar and Pundban (10,831), Boh (7834), Narwana (151), and

Chanderban (4484).

Simla. The Division is practically all situated in the hills,

the forests being mainly in Native States. The Deputy Com-

missioner, as Political Officer, issues passes for shooting, which

cover everything outside certain specified preserves.

The patch of sal-forest known as Kalesar at the foot of the

Simla Hills on the west bank of the Jumna River, area 11,570

acres, is closed to all shooting, as is also the Jagadhari or

Kalanour plantation, an area of 197 acres. So also no shooting

is permitted in the forests of the Simla Catchment Area. The

forests of the rest of the division are open. Kalesar contains

sambhar and chital, hog-deer and kakar; the Jagadhari plantation

pheasants etc.; and the Simla Catchment Area gural, serow,

pheasants, and chukor.

Lahore. The areas closed to shooting in the Lahore Division are

mostly plantations. In the well-known Changa Manga planta-

tion of Sissu and Mulberry, situated at about 40 miles from Lahore,

Blocks III and XIV (area 4424 acres) are closed to shooting, as

is also Shahdara Plantation close to Lahore (1828 acres) and the

Shadhanwali Plantation area (3059 acres).

It will be seen from the above that in the Punjab considerable

progress has been made in the endeavour to protect the game of

the Province.

The game to be protected is nilgai in Changa Manga and

game birds ete. elsewhere.

3. United Provinces.

Curiously enough, there are as yet no Game Sanctuaries in the

United Provinces, and this in spite of the fact that many of the

forests are probably amongst the most intensely shot over in the

country. It is understood that this question is now receiving

the consideration of the Lieutenant Governor and his Conser-

vators of Forests, and it is probable that before long the game

of the United Provinces will be given that amount of protection

which is essential if its deterioration and disappearance is to be

prevented. Were it possible to close blocks of considerable size

in shooting grounds like those, e. g., of the Dun (Siwalik Division),

there might possibly be some hope of saving the wild animal fauna

from extinction; that it is perilously near this, all who are

familiar with the conditions and the absurd so-called shooting

‘‘rights” these forests are burdened with are well aware.

With Sir John Hewett, the Lieutenant Governor, I would

advocate the closing of blocks of forest in every shooting division

in the Province for periods of two (I advocate preferably three)

years, the areas to be carefully selected and listed and closed

and opened in rotation. If an example as to the necessity of this

step being taken is required, it is only necessary to compare, say,

the Patli Dun with the ruined Dehra Dun forests as an example

of what unrestricted shooting, without protection, must lead to.

GAME PROTECTION IN INDIA. BY)

4. The Central Provinces.

As a whole, the Central Provinces may be considered to be the

most advanced so far as game protection is concerned. The

shooting regulations will be referred to at a later stage. They

were revised by the Chief Commissioner in his Notification

No. 1263, dated 28th October, 1907. For my present purpose it

is sufficient to notice that they provide that areas or blocks of

forests may be closed to shooting absolutely for purposes of forest

management or as sanctuaries for the protection of game, other

than carnivora, for the destruction of which special permits may

be issued. The list of closed forests or blocks is prepared each

year in October by the Conservators and is published in the

Central Provinces Gazette, and copies are hung up in the Deputy

Commissioner’s and Forest Ofticer’s offices.

Tt will thus be obvious that the Game Sanctuaries in the

Central Provinces are formed automatically by the closing alter-

nately of different forests or blocks of forest yearly. As a

matter of fact, however, most of the present sanctuaries, though

in many instances reduced in size, have been game sanctuaries

since 1902, though a few others have been added later. It would

be better if these areas were closed for periods of not more or

less than three years. Of course, in the case of areas closed

for purposes of forest management it is possible that they are

closed for a considerable period of years, but nothing is said

on this score in the rules nor as to the length of time blocks

are closed for purely sanctuary purposes.

In addition to the automatic closure and opening of blocks

there are other most valuable restrictions for the preservation

of game, and I believe that I am correct in stating that this

procedure is now applied to most of the blocks, instead of

automatically closing and opening them. In any particular

block or series of blocks only a certain head of any particular

species may be shot. As soon as this number has been reached,

that species is closed to shooting for the year. This rule might

well be introduced elsewhere in the country. It is really better

than the procedure of the Nilgiri Game Association of allowing

each sportsman to shoot one or two head of each species, since

the latter safeguard is in force also in the Central Provinces.

The permit of each sportsman is endorsed with the number of

head he may shoot, e.g. one bison, one sambhar, two chital, four

other deer, and carnivora ad. lib., provided the maximum number

of head of the species allowable to be shot in the year has not been

already reached. This latter information is supplied him either

by the divisional officer or by the Range officers in the areas for

which his permit is made out. Were not this latter provision in

force, one sportsman might shoot the whole number of, say, Bara-

singha (Cervus duvauceli) permissible for the year and thus close

this particular animal to succeeding guns for the rest of the

season—a somewhat unfair and onerous restriction.

In the Central Provinces excluding Berar in 1909 there were

24 Game Sanctuaries with an area of 1445 square miles in the

36 MR. E. P. STEBBING ON

Northern Circle closed to shooting, but permits could be obtained

to kill earnivora, and 293 sq. miles in the Southern Circle, or a

total of 1738 sq. miles in the Province.

Other areas are closed conditionally, subject to the issue of a

permit or to sportsmen being exempt from obtaining one.

As already stated, the blocks vary from year to year, and the

list is made out annually in October and published.

The game protected are bison, buffalo, sambhar, barasingha, and

chital, and, in fact, all horned game.

Berar.

The revision of the shooting rules in the Central Provinces in

August 1905 was followed by a similar revision in those of the

forests of the Berar Circle in the following October, the rule

regarding the formation of Sanctuaries being made applicable

here. The number of individuals of each species to be shot in a

year is laid down and notified. An area of 885 sq. miles was

closed to all shooting (save carnivora on permit) during 1909.

Shooting in the other forests was subject to permits being taken

out in the ordinary manner.

The game to be protected are chiefly sambhar and chital.

Bengal.

Bengal has recently taken up the question of Sanctuaries and

under Notification No. 1992 For., dated 6th Apvil, 1907, the

following have been formed :—

Darjiling Division.

Nenchal Forest. The boundaries of this Reserve are :—

North—Pashok Road from Jore bungalow to the 3rd mile.

East — From the 3rd mile to Lalkhunti where the road crosses

the saddle (by the Rangbi path).

South—From Lalkhunti to the old military road.

West -— The old military road to Jore bungalow.

Puri Division.

The Bharatpur, Jaimangal, and Majna forests are Game

Sanctuaries.

In Angul and Puri the shooting of bison is prohibited.

The shooting of the rhinoceros, buffalo, and female bison is

entirely prohibited throughout Bengal.

{ would suggest that in certain parts of the Province further

Sanctuaries are urgently needed. For instance, the game of the

hill forests of Darjiling, Kurseong, and Tista Divisions, especially

the two former which are surrounded by planters, requires

protection if it is not to be completely exterminated. The

Senchal Sanctuary can scarcely be considered sufficient in a

country as rugged and difficult as the Himalayas. The plains

and forests of both Kurseong and Tista Divisions were once noted

for the game they held, but a very different state of affairs now

exists and the accounts of old sportsmen are like legendary fables.

GAME PROTECTION IN INDIA. 37

On the other side of the Province to the west, the opening out of

Chota Nagpur by railway extension has thrown the forests open

to a far larger number of rifles than was ever to be seen in that

part of India a decade ago or less.

Singbhum, Palamow, Hazaribagh. The game in all these forests

now requires pr otection if it is not to be wiped out by the rifles

of such large centres as Calcutta etc., now brought within easy

rail distance.

Rules such as are in force in the Central Provinces would be

equally valuable and applicable to these parts of Bengal.

Eastern Bengal and Assam.

This province may be put almost on a par with the Central

Provinces so far as the provision made for Sanctuaries for game

is concerned, The work has not been carried out in all the

divisions, but the ones most shot over are well provided in this

respect, if we except Chittagong.

The provision of Sanctuaries, however, was not made any too

soon, if, indeed, it can be caneidened to have been introduced soon

enough. ‘The apathy of the authorities has led to the deteriora-

tion of the stock of game almost to the verge of extinction. The

forests are in large blocks, and it would have been suthcient in

the past to enforce the rules under the Forest Act. Neglect to

do so has reduced the numbers of such animals as rhinoceros,

buffalo, and bison to such small figures that deterioration, if not

extinction, now threatens these species.

The following Sanctuaries exist in the divisions detailed below :—

Khasi and Jaintia Hills. The Riat Khwan Sanctuary of 969

acres in this division is closed to all shooting, with the idea of

preserving game in general and not of one or more particular

species only. The Laitkar | Sanctuary of 432 acres is closed to all

shooting, with the idea of protecting insectivorous birds in addi-

tion to game in general. The Riat Laban of 995 acres is a

sanctuary similar to the Laitkar.

Nowgong and Sibsagar. An immense Sanctuary known as the

Kaziranga Sanctuary of 57,273°6 acres is closed to all shooting

except permit-holders,

Nowgong. Lowkhawa Sanctuary of 25,760 acres is closed to all

shooting, save that ailosred to permit: holders.

Kamr up. The North Kamrup Sanctuary of 57,600 acres is also

closed to shooting other than that allowed to authorised permit-

holders.

Goalpara. Three large Sanctuaries have been created at the

instance of Mr. W. F. Perrée, Divisional Forest Officer in this

Division, probably one of the most visited from outside of any

in Assam. The sanctuaries were in the first instance, I believe,

designed to prevent the rhinoceros from being exterminated, but

the buffalo and gaur are also carefully protected. Although

shooting is said to be permitted to permit-holders I believe that,

asa matter of fact, these sanctuaries have been entirely closed to

all shooting,

38 MR. E. P. STEBBING ON

The Sanctuaries comprise :— _

(i.) Portions of the Raipur and Kachugaon Reserves (area

69,120 acres).

(ii.) A portion of the Raipur Reserve (area 30,040 acres) and

a portion of the Bizni Reserve of 36,840 acres.

Jalpaiguri and Baksa Duars. J believe it is intended to form

Sanctuaries in these two Divisions, newly acquired by Assam

under the Partition. No Sanctuaries have been formed as yet,

although they are, we understand, under consideration.

These forests, however, are nominally Sanctuaries already for

certain animals, since the shooting of rhinoceros, buffalo, and bison

is entirely prohibited. The forest blocks are, however, too small

to furnish sufficient space for animals of such wandering dis-

positions. In former times the animals had access to the adjacent

Bhutan Hills in the hot weather, to which they used to repair

to escape the fly season. This outlet has now been cut off, owing

to the settlements of Nepalis on the Outer Hills.

Chittagong. This is another of the ceded divisions. It might

be suggested that a Sanctuary or two should be formed in some of

the Reserve Forests in the Collectorate to protect the bison and

sambhar, both from the over-keen European sportsmen and the

neighbouring villagers, who are apt to help themselves and shoot

anywhere. The neighbouring Hill Tracts also afford excellent

areas for game protection.

Bombay.

In Bombay the question of Game Sanctuaries has received

considerable attention, special areas having been set aside in

which only pig, carnivora, and other destructive animals may be

shot. To do so, however, entails a special endorsement being

first made by the Divisional Forest Officer on the holder’s shooting-

permit, save in the case of Gazetted Officers whose jurisdiction

extends to the Sanctuary in question. Such special permission

cannot be given for a longer period than one month in each case.

In the Southern Circle there are four Sanctuaries, situated in

the Belgaum, Kanara Northern, Kanara Southern, and Kanara

Western Divisions, the areas being 26 sq. miles, 330 sq. miles,

400 sq. miles, and 216 sq. miles respectively.

These Sanctuaries have been specially made to afford protection

to sambhar, chital, and bison.

In the Sind Circle a number of Game Sanctuaries have been

formed in the Jerruck, Hyderabad, Naushahro, and Sukkur

Divisions.

The chief game for which the closure has been prescribed is

the “phara.” The period for which the Sanctuary is formed is

one of five years, at the end of which the Sanctuary will be thrown

open and other forests closed.

The Sanctuaries are as follows :—

Jerruck Division.

The Sanctuaries in this Division are eleven in number :—Sonda

GAME PROTECTION IN INDIA. 39

(area 2612 acres), Aligunj (1199), Shah Lanko (1743), Garko

(4092), Chacb (1661), Marho Kotri (1668), Allahbux (2045),

Bohadipur (2763), Sukhapur (3878), Hazari (2483), and Shatoh

(277).

Hydrabad Division.

Tn this Division there are fourteen Sanctuaries :—Abad Thebat

(640), Rajri Khasai (457), Budhopur Vaechero (748), Railo Boda

(1322), Mahomed Khan (181), Kathri (756), Miani (5312), Elehi

Hingoroni (288), Nurketi (4254), Kot Dhingano (1893), Kacho

Lakha (4705), Mehrobpur (3012), Mari (17,651), Sai (5901).

Naushahro Division.

Fourteen Sanctuaries as follows:—Kocho Khairoders No. 1

(4118), Khairoders No. 2 Phulel and Dulotpur (3570), Lalia

(2324), Dalipota (867), Kundoh (5965), Khairodero No. 1

(1650), Keti Lalia (1159), Karampur (1095), Gajidero (4336),

Budhodero (1555), Nari (1522), Kasim Sah (2107), Akil (397),

Sharifpur (212).

Sukkur Division.

All the forests of the Sukkur Division have been closed to

shooting for five years, as big game (phara) has become scarce.

These forests number 64 in all, covering an area of 273,497 acres.

Madras.

Reference has already been made to the excellent Game Asso-

ciation of the Nilgiris, Under the rules of this society, and the

spirit in which they are worked out and observed, protection

may be said to have reached a high state of perfection in the

Ootacamund Hills.

The large resident European population in the hills has, of

course, rendered the introduction and strict observance of the

rules a much easier business than is likely to obtain in areas

which are only visited by occasional sportsmen, who have not

even the interest of the local resident officials, who are in a way

permanent in the locality, in the upkeep of the head of game,

and therefore clamour for free shooting, equal rights, ete.

To some extent the lead of the Nilgiri Game Association has

been followed by the Forest Department, but at present apparently

only in a small degree.

In the Southern Circle the only Sanctuaries existing in 1906

were situated in the Nilgiri District as follows :—

Nilgiri Game District. The whole Nilgiri Game District was

closed to the shooting of pea-hens altogether and jungle-hens for

one year. This was done by the Game Association.

Todanad Division. The Marlimund Plantation (17 acres) and

the Kanabettri Forest (500 acres of the Segur Reserve) were

closed to all shooting.

The Governor’s Shola (33), Sheffield Plantation (25), Tiger

Shola (20), Kurnutherguli Shola (78), and the Sholas on the east

40 MR. E. P. STEBBING ON

and south sides of Hembar Hill (36), were closed to small-game

shooting only.

South-Hast Wynaad. The Benne (10,344), Madumali (46,639),

and the Kombarakkollai (1771) Reserves were closed to bison-

shooting for one year from Ist June, 1906. Owing to the heavy

mortality, from anthrax, bison had experienced a few years

previously and to the increased shooting, this step was a much-

needed one; the year could well be extended to three or more.

Kundahs. The Kundah Reserve of 40,379 acres was closed to

beating for big game except tigers and panthers, and to the

shooting of small game.

Paranginad. A number of Game Sanctuaries have been formed

in this Division. In the Longwood Reserve the portion North

and Kast of a line from Kengarai Sign Post to the top of the hill

and Longwood No. 1 (30 acres); Nedukaduhalla (885). Sundatti

Reserve and addition (123 and 35), Kunshola Nos. I and Il

(158), Madanad (722) and addition (708), Kodanad valley (294)

and additions I and II (245), Nedugula (88) and addition (113),

Avarahalla (121), Gudakahalla No. Il north of Kilkotagiri

bridle-path (70), Seven mile tope reserve (2), Kannerihodai (16),

Nlavandmund Shola and additions I and II (118), Sullicodu

Nos. Land II (308), Attukadu (71), Warbreccan (17), Sinnattu

and addition (358), Uppatti Shola and addition (67), Doddakavu

(62), Curzon Valley Block IIT west of Kilkotagiri-Curzon bridle-

path (895). All these are reserves and are closed to beating for

big game except tiger and panther; and are also closed to small-

game shooting. In addition Ralha (382) and Sims Park (83) are

closed to small-game shooting only.

The total area thus closed as Game Sanctuaries in the Nilgiri

District in 1906 totalled 107,083 acres.

In the Central Circle only two districts were reported to have

constituted Game Sanctuaries.

North Arcot District. In the North Arcot Division various

additions to the shooting licenses had been made which will be

considered under the shooting-permits. The number of head of

particular game to be shot is limited in the case of certain species,

as is the case in the Central Provinces. Special sanction has to

be obtained for beating; and sitting up over water and salt licks

is strictly prohibited.

The following reserves are closed to shooting, without the

special permission in writing of the Collector and District Forest

Officer :—

Chamala Reserve of 19,142 acres; Javadi Reserve of 403,566

acres; and the Musalimdugu Reserve of 5542 acres.

The game for which these Sanctuaries are especially created are

sambhar, deer, black buck in the case of the Chamala Reserve,

bison and deer in the Javadi Reserve, and deer and hare in the

Musalimdugu Reserve.

In addition, 71 blocks of forest are closed to shooting during

the fire season from Ist March to 30th June.

GAME PROTEC'TION IN INDIA. 4]

Tanjore District. The Point Calimere Reserve of 4204 acres is

closed to the shooting of black buck, chital, and hare. Pigs and

partridges may be Shick without Sagan and three blag buck

for each license given. All oe deer, does, and young animals

may not be shot between the Ist Febru: wy and 31st July.

Burma.

Very little has been done as yet in Burma towards the forma-

tion of Game Sanctuaries. Fortun: ately Burma was but recently

disarmed and Government has not given licenses in anything

hike the numbers granted elsewhere. In many parts game is

believed to have increased, but the European sportsman has had

little mercy on some species, and the fine herds of brow-antlered

deer (Cervus eldi) in Lower Burma were almost exterminated

before measures of protection were introduced. Deterioration

must of necessity follow, even if the species be not entirely

exterminated.

No Game Sanctuaries exist in the Pegu, Tenasserim, and

Northern Circles.

In the Southern Circle, Upper Burma, the Maymyo Park Reserve

was notified as a Sanctuary for game in an interesting notification

which appeared on page 421 of the Burma Gazette for the 7th April,

1906. This Sanctuary is in the Mandalay Division, its extent

being approximately 7? miles. The notification being the first

of its kind in Burma, it will be of interest to reproduce it here.

“Tt is hereby notified under Rule 2 (2) of the rules under the

Forest Act 1902 that, with the previous approval of the Local

Government, the Maymyo Park Reserve is closed to the issue of

licenses for shooting or fishing.”

The Conservator in issuing a notice on this subject stated :

“The object of this is to provide a sanctuary for pea-fowl and

other game (chiefly barking deer and partridges) and to thereby

enhance the pleasure of the numerous sections of the public who

utilize the Reserve for riding and take an interest in the animals

therein. It is hoped that the public generally will co-operate

with the Officers of the Forest Department in seeing that the

intention of the above notification is carried out.”

The terms of the notification will not be enforced as regards

snipe-shooting in the swamps, which may as heretofore be carried

on without license at the proper season, nor will it be enforced as

regards the shooting of dangerous carnivora. For the purpose of

notification, the following are the rough boundaries of the Sanc-

tuarles :—

North. The Lashio Road from the Circular Road to Yegyano.

Kast. Bames’ ride from Yegyano to Kadettaw cut and the

Circular ride from Nyaungni Pagoda to the Mandalay Road near

Pwedaung quarry.

West. The Mandalay Road to the Circular road and the latter

to its Junction with the Lashio Road.

42 MR. E. P. STEBBING ON

B. NATIVE STATES.

1. Nashmir.

Allusion has already been made to the fact that the progressive

Native State of Jammu and Kashmir has a Game Preservation

Department as a separate part of the administration of the State,

with an officer and special staff in charge of the work. ‘This, I

think, is the ideal at which British India should, and I believe

will, eventually inevitably work up to.

The Department issues each year a printed copy of the Game

Laws notification, which are applicable to everyone, save those

persons especially excepted by order of His Highness the Maha-

raja in Council.

The Rules do not apply to the State Game Reserves, for which

no shooting-permits are issued.

The existing State Game Reserves, about 300 sq. miles in

extent, are :—

(1) Chashma Shahi, and (2) Dachgam, and the ridge of hills

between these Rakhs, (3) Khonmoo, and the grass-farm between

Sangri and Chak Khonmoo, (4) Khru, (5) Tral-eum-Kerrim, and

Punjhair as far as the Bhoogmor Road, (6) Achabal, (7) Kool-

gam, (8) Kandi and Khoras in the Uri Nullah, the lower boun-

dary of which is the new road to the Hajipir and Hokarsar Jhil.

N.B.—The basin above Pandrittan is a Sanctuary for chukor.

In General Raja Sir Amar Singh’s Jagir and in the Iaqua of

the Raja of Poonch no one is allowed to shoot without the

permission of the respective Rajas.

The following Nullahs will be considered as the private

shooting-grounds of their respective Rajas, viz., Kapalu, Khar-

mung, Shigar, Rondu, Kiris, also that portion of the Mantho

and Satpur Nullahs demarcated in 1905, for the Rajas of Tolti

and Skardu, respectively. These nullahs will be closed for

shooting, except with the permission of the Rajas concerned.

The following Nullahs are closed until further orders as Game

Sanctuaries, and no shooting is permitted therein, nor is any

grazing allowed :—

I. The Cretchen basin in the Lidder Valley from Bhatkote

to Kitchroo (Srinagar Khass). Area 20 sq. miles.

II. The Kiar Nullah in the Duchan District of Kishtwar.

Area 120 sq. miles.

III. The Gueo Nai in the Wardwan (Kishtwar). This is the

nullah which joins the left bank of the Wardwan River one

march above Maru Wardwan. Area 48 sq. miles.

TV. and V. Nimo and Masho in Ladakh. Area 40 and 36 sq.

miles.

VI. Melangan in the Kajnag (Mozufferabad). Area 28 sq.

miles.

GAME PROTECTION IN INDIA. 43

VII. The Siranda branch of the Erin Nullah (Baramoula).

Area 32 sq. miles.

VIIT. Alchori and Hashopa and intermediate ground in the

Shigar Valley ( Baltistan). Area 50 sq. miles.

IX. Doosoo in the Nowboog district (Srinagar Khass). Area

60 sq. miles.

X. Shelterin Astor. Area 30 sq. miles.

XI. Soomjam and the Sapphire Mines Nullah in Kishtwar.

Area. 35 sq. miles.

XII. Gabdoori and Kunirawul in the Shamshibiri. Area

23 sq. miles.

As will be seen when we come to a consideration of the

shooting rules in force, driving game is only permitted in certain

localities at certain periods, the number of head to be shot on

a license is restricted per species and neither females nor young

may be shot.

2. Chamba.

Chamba is another of the favoured shooting grounds of the

Kuropean from the climatic point of view, and under the late

Raja the game was being rapidly destroyed. Under its present

enlightened and most hospitabie ruler conditions have altered,

and shooting is now prohibited save on permits issued by the

Raja himself.

Sportsmen have found little difficulty in obtaining such, the

ground and number of head to be shot being indicated on the

permit. The method of protection followed is on the lines of

the Kashmir Shooting Regulations.

3. Hyderabad (Deccan).

Whilst there are no special Game Sanctuaries in the Hydera-

bad State Forests, the recently amended Game Regulations and

the periodical opening and closure of certain tracts to shooting

go to prove that His Highness the Nizam is fully aware of the

necessity of affording protection to the wild game of his territory

if it is not to be exterminated by over-shooting.

The Game Preserves of the State are of four classes :—

1. His Highness the Nizam’s own Preserve.-—This Preserve is

situated in the Warangal Division in the neighbourhood of the

Godavery Valley Railway, and has an approximate area of 828 sq.

miles. Protection in this Preserve is provided to all game, and

the shooting in this tract is reserved entirely for His Highness

the Nizam or His Excellency the Minister. No permits to shoot

can be obtained.

Il. His Highness ihe Nizam’s State Preserves.—These are

situated in the vicinity of Yellander and Singarein. All game is

protected, and permission to shoot can only be obtained from the

Nizam or his Minister. This area is kept for the State guests,

44 MR. E. P. STEBBING ON

and a permit would never be granted under ordinary circum-

stances to the casual sportsman.

Til. The Open Ferests—The Open Forests are divided into

three Divisions. These are opened each year from Ist March

to 31st July and for ten days at Christmas in rotation, 7. ¢. each

division is open for one year and closed for two. The divisions

are each divided up into separate blocks. Sportsmen wishing to

shoot in any special] block register their names, together with a

fee of R.25 before the lst December. The names of persons

applying for the blocks, in the event of there being more than

one, are publicly balloted for by lot on a date fixed somewhere

about Ist January and the permits are issued accordingly. Persons

drawing a block pay an additional fee of R. 25 for each member

of the party intending to shoot. Those failing to draw a block

are refunded their deposited fee.

Anyone not occupying his block within a month of March Ist

may be made to forfeit his right to do so, in which ease the block

is given to the next name drawn

Priority of claim to an open block is always allowed to a

person resident within H.H. the Nizam’s territories.

The number of tigers allowed to be shot in a block is limited

to a total average of. two per each rifle in the party.

Deer are not, and never have been, protected in any way, and

as nearly every person in the State is armed with a firearm of

some sort, from the latest thing in Rigby cordite rifles to a

horse-pistol, practically the only game existing outside the State

Preserves is confined to carnivora and bears.

IV. The Jaghirs.—Jaghirdars have the right to give or refuse

shooting-permits to applicants to shoot in their land, as they

think fit. For others permission to shoot in a Jaghir is a private

matter between the sportsman and the Jaghirdar.

4. Pannah State (Bundelkhand).

In the Pannah State there are two State Sanctuaries of about

50 sq. miles apiece, which have existed from time immemorial.

No one is permitted to shoot in these, save the Maharaja and the

Political Agent, so that they are not Game Sanctuaries in the

true sense of the word. The license granted for shooting in other

parts of the State prohibits the killing of tiger, chital, stag,

sambhar (when hornless or in velvet), and doe sambhar through-

out the year.

In a letter answering some enquiries put to him on the question

of Game Protection in the Central India States, Mr. R. M.

Williamson, I.F.8., who has a considerable experience of these

States, wrote :—‘‘ Generally there is no protection of any sort of

game outside the special State Preserves, except tiger, the killing

of which is reserved for the ruler. This state of affairs is general

in the Native States of Central India, and it is impossible to effect

any improvement in this respect till additional forests ave reserved

GAME PROTECTION IN INDIA. 45

and brought under efficient protection. My experience both

here (Rewah) and in Bundeikhand is that big game (save carni-

vora) is much scarcer than in British Indian Forests, 7. ¢., that

the proportion of carnivora to other kinds of big game is far

higher in these States than in British India.”

5. Bhopawar Agency.

No regular Game Sanctuaries have yet been formed in the

; P Bis

Bhopawar Agency, but the Game Rules are undergoing revision,

and with this revision the formation of Sanctuaries is intended.

A close season is in foree, but it does not extend to solitary bison

stag or male antelope, florican or quail.

6. Rewah State.

Shooting of all kinds is prohibited in the Reserved Forests

without special permission, which it is very difficult to secure.

The area of the reserves totals, however, only 600 sq. miles.

Elsewhere there is no game protection whatsoever for game, save

tigers, the shooting of which the Maharaja reserves for himself

and his guests.

7. Cochin.

The Conservator of Forests of the Cochin State informs me

that no Game Sanctuaries have as yet been established in the

Cochin State.

&. Mysore.

In Mysore seven areas have been set aside as Game Sanc-

tuaries. No special rules have been framed for these areas, but

the protection of bison, sambhar, and other deer is chiefly aimed

at. It is apparent, however, that shooting is allowed in the areas,

but only with the special permission of H.H. the Maharaja.

It seems a pity not to prohibit all shooting so long as an area

is a sanctuary and to rotate the sanctuaries on a three or five

yearly system.

These sanctuaries are situated as follows :—

1. Mysore District.—This District contains four sanctuaries,

Kakenkote, Begur, Berambadi, and Chamarajanagar, of 55, 42,

102, and 98 sq. miles in area respectively.

2. Hassan District. —One Sanctuary, by name Hirikalgudda, of

27 sq. miles in area.

3. Kadur District.—One Sanctuary, Tegurgudda, of 9 sq. miles

in extent.

4. Shimoga District.—One Sanctuary, Sakrebyle, of 15 sq. miles

in area.

9. Travancore.

There are no Game Sanctuaries in Travancore, and none are

considered necessary. The Conservator of Forests writes: “There

are no Game Sanctuaries in the Travancore Forests, nor are they

consilered necessary, because the area of evergreen forests where

46 MR. E. P. STEBBING ON

the game can take refuge is so extensive that there is no danger

of its extermination. In the Reserved Forests extending over

one-third of the State, hunting and shooting are only permitted

after a license has been obtained.”

TV. Tue Proposep InpIAN GAME ACT.

I shall now briefly glance at the provisions of the New Indian

Game Protection Act.

The following is an abstract of the Proposed Indian Game

Protection Act :—.

1. (1) To be called the Game Protection Act.

(2) Includes all India, Baluchistan, Santhal Parganas, and

Spiti.

(3) To come into force when published in the Gazette of

India.

2. “Game” and “large animal” to have meanings defined by

Local Governments. In the absence of such definitions they

shall mean :—

(a) Game: The following in their wild state :—

(1.) All pigeons and sandgrouse.

(I1.) Peafowl, jungle-fowl, pheasants, partridges, quail,

spurfowl, florican, and their congeners.

(IIT.) Geese, ducks, and their congeners.

(IV.) Woodcock and snipe.

V.) Hares.

on Asses, all kinds of rhinoceros, buffalo, bison, oxen ;

all kinds of sheep, goats, antelopes, and their

congeners; all kinds of gazelles and deer.

(VII.) Any other animals the Local Government may

notify.

(b) (1.) “ Large animal” shall mean any animal included in

subclause (VL.) above.

(IL) “Specified kind” means any kind of game, large

animal, or fish, distinguished by species,

immaturity, or sex.

3. This Act does not affect :—

(a) The pursuit, capture, or killing of :—

(1.) Game by non-commissioned officers or soldiers, on

whose behalf regulations have been made.

(11.) Any animal for which a reward may be claimed

from Government.

(1I1.) Any large animal in self-defence.

(1V.) Any large animal by a cultivator or his servants,

whose crops it is injuring.

(b) Anything done under license for possessing arms and

ammunition to protect crops, or for destroying dangerous

animals, under the Indian Arms Act (XI. of 1878).

“1

GAME PROTECTION IN INDIA, 47

ProurIBIToRY PROvISIONS.

No firearms shall be used or carried for killing game, without

license under this Act, or under the Indian Arms Act.

The Local Government may, by notification, prohibit, within

any local area, for any period, the capture, or killing, or

attempt to capture or kill :—

(a) All game, or any specitied kind.

(b) All game, or any specified kind, by any special means or

manner, either absolutely or except in accordance with

a license under this Act.

(1) The Local Government may prohibit the capture or

killing (or attempt to capture or kill) of any fish, during

any part of the year, in any specified stream, or in the

headwaters of any specified river, or in any specified

part thereof.

(2) Any notification under this section shall be conclusive proof

of the matters stated therein.

A Local Government may notify prohibition of the possession

or sale of all game or fish, within Municipal or Cantonment

limits, or in the importation of any kind of plumage of any

specitied kind of game.

Provisions As To LICENSES.

(1) Licenses shall be granted under this Act, by authorities

empowered under it, save as provided under sub-

section (2).

(2) A Local Government can authorise a Commissioner or, in

Madras, the Collector, under orders of the Governor-

General in Council, to grant licenses under this Act, for

the use or carrying of firearms for killing game,

throughout the whole of British India.

(3) Under rules framed by a Local Government, the Collector,

or any authority thereby empowered, may cancel any

license granted under this Act :

(a) When the holder has been convicted under this Act.

(b) When any term of the license has been infringed.

(1) In addition to conditions prescribed by a Local Government,

every license shall be deemed to be subject to the conditions

set forth in any notification in force for the time being

under section 5.

(2) Every license shall specify :—

(a) The area in which it is in force.

(b) The instruments to be used, and in the case of a weapon

to which the Indian Arms Act (IX. of 1878) applies,

the person or persons by whom alone it may be used

or carried.

48 MR. E. P. STEBBING ON

game, the periods, and the methods of capture or

killing, to which alone it is to apply.

(3) When firearms are authorised, the license may limit the

number of any specified kind of large animal which may

be killed, and the fee payable for each such animal

killed.

(4) Every such hunter, beater, or assistant, aiding the licensee,

shall be protected by such license.

10. Every license under this Act is non-transferable, and may

be granted for any period not exceeding a year, and may be

renewed on payment of a fee of twenty rupees for each

issue, and for renewal, provided :—

(1) That the Governor-General may notify a higher rate of fee

in the Gazette of India, and that security shall be

deposited on each issue or renewal, or from any specified

class of persons.

(2) That a fee of only five rupees be payable by an owner of

agricultural land; the license to be in force exclusively

within the village or villages in which such land is

situated, and within a local area, if it includes more than

one village, not exceeding fifteen hundred acres.

(3) That a Local Government may exempt any specified class

of person, in any specified area, from payment of fees on

licenses not authorising the use of firearms.

11. Every licensee is bound :—

(a) to produce his license for inspection, when demanded, by

any Magistrate, Police officer not below the rank of

Inspector, or in a forest by any Forest officer not below

the rank of Extra Assistant Conservator ;

(b) to deliver up his license on expiry of the period for which

it was granted, or when leaving India, whichever is

sooner, to the Collector or any authority authorised to

grant a license, or tothe principal Police officer of a

district.

12. Every licensee shall :—

(a) keepan account of any specified kind of large animal killed,

giving sex, date, and place ;

(6) produce such account on demand made by any Magistrate,

Police officer not below the rank of Inspector, or in a

forest by any Forest officer not below the rank of Extre

Assistant Conservator ;

period for which the license was granted, or on leaving

India, whichever is sooner, to the principal Police officer

of the District or Presidency town in which he then 1s.

GAME PROTECTION IN INDIA. 49

13. (1) A licensee under this Act is exempt from the provisions

of the Indian Arms Act as regards taking out a license

for the same weapon, and the same area, under that

Act.

(2) Every license granted under Forest regulations for the

capture or killing of game, shall have the same effect as

a like license granted under this Act.

RULEs.

22, (1) A Local Government may make rules for the purposes

and objects of this Act.

(2) Such rules may :—

(a) declare the authority by which the licenses may be

granted or cancelled ;

(6) prescribe terms and conditions under which such

licenses are granted ;

otherwise than by a Magistrate or Court.

V. Some CriricismMs on THE PRoposEp ACT.

It will be of interest to consider in some slight detail several

of the provisions of the proposed Act.

The Provincial Rules at present in force under which all Game

Protection work is carried out are comprised in the Arms Act,

Forest Act, and Fisheries Act.

The proposed new Act extends to all India, with the exception

of Burma and, of course, the Native States. I have shown

that some of these latter are already doing excellent work in Game

Protection, and others will doubtless follow a firm lead set them

by the Imperial Government.

It is not intended here to offer any seriously destructive

criticisms on the above provisions of the suggested Game Act.

There are one or two points to which, however, I think

attention might with advantage be drawn. In the second clause

the meanings attached to “ game,” “ large animal,” and “ bird ”

are distinctly and regrettably vague. I am unable to see any

reason for this. If the drafters of the Bill were to apply to any

zoologist in the country who has a practical working and sporting

knowledge of the game life of India they could be furnished with

detailed lists of animals both large and small: and by “animals ”

Proc. Zoou, Soc.—1912, No. IV. 4

50 MR. E. P. STEBBING ON

IT here mean “mammals” classed, say, into some such groups

as, €.9.;

(a) Carnivora. Hach species in the country to be quoted.

(b) Herbivora. do. do. do. do.

The various deer, antelope, goats, and sheep are all perfectly

well known, and the preparation of lists detailing each animal

by name is an absolutely easy matter.

extinction of which from a sporting point of view is far from

desirable.

(d) A General Group which may be made to include the

rest of the Mammalia. This would allow protection to be ex-

tended, should it be deemed necessary from the point of view of

the preservation of the species in the case of rare species now

perhaps being exterminated for the value of their fur or for other

reasons, to animals not at present included in the sportman’s

category of Game.

Turning to the Birds. There is no distinction made between

migratory birds and non-migratory birds, and no mention made

at all of Jnsectivorous Birds, and yet the distinction is one of

enormous value in a great agricultural country like India, where

the benefit the cultivators must derive annually from insectivorous

birds is quite incalculable.

I suggest that the Birds be sharply defined into groups and

the names of all the game-birds and of all the chief insecti-

vorous birds be definitely given in the Act. This likewise is a

matter of the greatest simplicity, since there would be no difticulty

in drawing up such lists.

With a proper classification by name of all our animals and

birds, sections and subsections 2 (a) (I.) to (VI.) would be revised

and (VII.) and (6) (I.) and (II.) could be eliminated if considered

necessary.

Turning now to section (3). Allowing that it is necessary to

make separate rules for the Army, I think that the Act should

specifically lay down that permits may not be given for parties of

more than say 4-6 men from a cantonment to go out together to

shoot inanyarea. At present it is well known that at times parties

of from 15 to 20 or more men go out into a block of forest and

drive the game systematically into a cul-de-sac and then slaughter

the animals in numbers. The Gurkha is particularly addicted to

this form of “sport” during the rainy season, when in the parts of

the country where they are cantoned it is generally impossible

for the European to go near the Terai forest owing to its great

unhealthiness. Parties of military men should be small and the

number of head they may shoot should be distinctly laid down on

the permit, and penalties be enforced if this number is exceeded.

I think the Game Act might embody some such definite ruling

for the whole country.

3. (II.) Lam of opinion that Game rewards in general should

be abolished and that no provision on the subject should be

included in the Game Act.

It would be quite within the power of the Local Government to

GAME PROTECTION IN INDIA. 5]

issue rewards for the destruction of a particular species which is

on the increase and becoming a danger either to public life or

property or to the sporting interests of a particular area of

country.

Also, save in exceptional cases, e.g. rogue elephants and man-

eaters, I would abolish the giving of a reward for every tiger,

leopard, wild dog or wolf slain.

Where any of these animals were becoming a pest or scourge to

the community or endangering the head of game of other species

in any locality, the Local Government should notify or empower

its officers to notify a reward or scale of rewards to remain in

force until the danger is past and the balance of power between

man and animal or animal and animal is once again normal. The

rewards on the prescribed animals should then be taken off.

Every shooting season nowadays sees an army of eager sports-

men competing for blocks and shooting-permits, and surely the

giving of the old-time reward for a tiger is quite unnecessary.

I would leave the grant of rewards or offer of rewards to the

discretion of the District officer or Forest officer. They would

when necessary prescribe such and such an animal to be a man-

eater or cattle-lifter of notoriety and would fix a reward upon the

animal, procuring, if considered necessary, the sanction of the

Commissioner or Conservator to their doing so. Why Government

should nowadays pay a reward of from Rupees 20 to Rupees 50

for a tiger which may be a pure game-eater and rarely if ever

touch a cow (and there are numbers of such) is beyond my com-

prehension. Sportsmen will not slack off if the rewards are

withdrawn. Many a district official would be only too delighted

if they would. Once a man-eater or a noted cattle-lifter is pro-

claimed, then make it worth the sportmen’s while to collect to

tackle him by giving straight off a large reward commencing at

R. 200 and going rapidly up to R.500. It would be a far more

satisfactory way of working the reward system both from the

point of view of the cultivator, the man who lives on the soil, and

that of the sportsman, and, I think, would probably be less costly

to Government.

Or rewards might be offered only for tigers in a district or parts

of a district where a noted man-eater or cattle-lifter has made his

home. For every tiger killed in this area a suitable reward might

be given, say R. 50, with the larger reward to be paid to the

sportsman who bagged the particular man-eater or cattle-lifter

prescribed. This would probably be the best method, since it

would tempt sportsmen to have a try for the man-eater, knowing

that they would receive a certain reward for each tiger killed, even

if they should not be lucky enough to kill the prescribed beast.

VI. Some SUGGESTIONS FOR THE PRESERVATION OF THE GAME OF

THE CouUNTRY IN THE INTERESTS OF THE GAME ITSELF AND

OF THE SPORTSMAN.

The maintenance of a proper head of a particular species on the

areas it affects in the country, 7.¢., in its own particular habitat,

52 MR. E. P. STEBBING ON

is the first point for consideration, as we elsewhere mentioned,

in the interests of the preservation of the species from

a. Deterioration,

b. Extinction.

The first point to be considered to ensure this object being

attained is the question of close seasons.

It is not of advantage, as was done in many Presidencies and

Provinces, to divide the year into two periods, calling one the

close season and the other the open and apply them to all animals

alike. Such a policy in a country of such varying conditions and

climate as India is obviously not only inadequate but may do as

much harm as good, for the breeding-seasons of many animals

and birds vary totally from those of others, whilst some species

breed at one time in one place and either later or earlier in other

parts of the country. ‘Take, for instance, the three chief deer

in the Central Provinces (sambhar, barasingha, and chital). Of

these the sambhar sheds its horns the earliest in the season (late

April), the barasingha next, and the chital last, the latter a

month or two, or more, later than the sambhav.

The close season for one obviously commences earlier than does

that for the others. At the same time I am not prepared to

say that the close season for sambhar in the Central Provinces

would be the same for the United Provinces Terai or for the

Assam Forests.

The drawing up of the close seasons for both animals and birds

is a work for the Local Government. With the knowledge we

now have of their habits, the preparation of such memoranda

presents no difficulties whatsoever. For instance, in the Central

Provinces Shooting Rules excellent Appendices are given in which

the varying dates for the close seasons for the game-birds of the

Province are laid down.

In drawing up the close seasons, therefore, I would prepare

tables showing separately the close season for each animal and

each game-bird of the Province. I also suggest that a separate

Appendix be attached to the Shooting Rules giving the names

of the insectivorous birds of the Province and enlisting the

sympathies of all sportsmen in their protection. A list of the

small predatory mammals and predatory birds such as hawks could

also with advantage be attached to the Shooting Rules, with a

recommendation to sportsmen to shoot such pests when met with.

There appears to be little danger at present of their numbers

decreasing to a dangerously low figure, and their depredations

touch the cultivator in one way, though not perhaps such an

obvious one, almost as closely as those of the large carnivora.

Having disposed of the question of the proper drawing up and

enforcement of the correct close season for each particular species

of animal and bird, we now come to a consideration of a question

which is of almost equal importance.

In past years but scant attention has been paid to the severe

attacks of a disease having kindred affinities to anthrax which

GAME PROTECTION IN INDIA, 53

appears at intervals and takes a heavy toll of the head of game

(such as bison (gaur), buffalo, sambhar, ete.) on the area it affects.

Further, in years of severe dr ought the mortality amongst the

wild animals of the country affected is often very heavy, and

in the past this factor has received no consideration from the

authorities in the interests of the wild game.

I suggest that in the wake of such calamities a very careful

and detailed inspection of the area or areas affected should be

undertaken, with the object of ascertaining which species have

suffered and to what extent. Until such survey has been carried

out, no shooting-permits should be issued for the area or areas.

After the survey the species which have suffered severely should

be notified, as also the area affected, and this area should be entirely

closed to the shooting of these particular species for such period

as will ensure their multiplication to the number of head it is

required to maintain on the area. In closing such area the

notification should distinctly state the reason for the closure. No

true sportsmen would be found to cavil at such a procedure.

The period of closure would require careful consideration. I

remember reading a year or two ago that the Nilgiri Plateau was

closed to the shooting of sambhar for one year. There would seem

to be little use for such a procedure, as no growth of horn of any

value to a sportsman can be put on by a stagina year. Of course,

such a provision might be of use in the case of areas which are

repopulated from neighbouring forests outside the limits of the

area for which the provisions are in force, and this may have been

the object on the Nilgiri Plateau. Still it should be borne in mind,

I think, that it is of little use closing areas or closing the shooting

of particular species unless the time that species will take to pro-

duce the desired result is carefully taken into consideration. And

nowadays such knowledge is available, since we have a first-rate

working knowledge, thanks to many enthusiastic and hard-working

sportsmen, of our game animals and birds.

In the case of an area affected by anthrax, for example. It

would be necessary to close the area to the shooting of the deci-

mated species for such a period as would allow a two-year old

sambhar stag to produce a decent head, 7. e. from 4-5 years. For

bison or gaur the period would be longer, and so on. Sportsmen

may decry this suggestion and say that it is toostrict. But why ?

The true sportsman, I take it, only wishes to shoot good heads.

Supposing, for instance, one or two good sambhar heads only are

left in an area. Only a man or two can get them. A sportsman

may follow who has come expressly to the area to obtain a good

sambhar. The anthrax attack was unknown to him. The result

is that he wastes both his time and money, and has a very good

complaint against the District Officials, who he very aeuiie con-

siders should have notified that the area contained no good heads

or, as we suggest, should have closed it to shooting of this species.

Some three or four years ago anthrax swept through a part of

the Central Provinces and portions of Madras, resulting in a

heavy mortality amongst the wild game, especially bison and

54 ON GAME PROTECTION IN INDIA.

sambhar. I know of one or two recent cases where men coming

from a long distance have taken blocks of forest in these areas, not

knowing of the anthrax attack. Take the one animal, sambhar.

One or two decent heads were bagged, but after that the most

indefatigable and laborious search, both in plain and hill forest,

resulted only in the disclosure of the fact that though young

2-3-year old stags were numerous, large ones were almost absent.

It may be said that this in itself acts as an automatic closure.

But it does not, as there is a considerable number of sportsmen

who will shoot the smaller head, as an examination of trophies

in the bungalows of every station in India will only too clearly

disclose. The entire closure of the species to shooting after such

visitations is the only fair action to take, both in the interests of

the animal and of the sportsman.

And now to turn for a moment, in conclusion, to the question of

the sportsman—the outside sportsman, not the District official—

and the rules under which he ean enjoy sport in a district.

The rules under which the District official enjoys sport in his

district are, I think, quite fair in most if not all Provinces.

I think, however, that the outside sportsman has often a

justifiable complaint, though more often than not he goes the

wrong way about in making it, and so puts himself out of court.

The whole matter really turns, and must always turn, on the

number of individuals of a particular species it is permissible to

shoot on a given area. This number can only be fixed by the

District Officers on the spot. There can be no cavil against this,

as they are, or ought to be, the best judges on the question.

In fact, as matters in game protection at present stand,

and in the absence of a separate Game Protection establishment,

there can be no appeal from their decision.

Probably the best and most elastic method for the outside

sportsman is to give him a block or blocks, according as to what is

available on receipt of his application, and to enter on his permit

the number of individuals of any one species he may shoot and

the number of different species. This number would, of course,

vary according to the length of time for which the permit was

issued, but would never exceed a fixed maximum for each species.

So far so good.

But it will doubtless soon be found necessary to definitely limit

the number of head of a species to be shot in any one area in a

year, as is done, in fact, in the Central Provinces. I¢ is in this

limitation that complaints arise and causes for friction come in.

For instance, supposing twelve sambhar may be shot in any

particular block. A military man, whose leave season will not

open before the 15th April, applies for and is allotted a block. He

arrives to find the maximum annual number of the animal it is

permitted to kill already reached and is debarred from shooting

that particular species. It is quite conceivable that he might find

more than one species in the same condition. In fact, the total

number of head of a particular species might be easily shot off by

the local District ofticials in the first couple of months of the open

ON THE M@ULTING OF THE ARCTIC FOX. 55

season, the animal becoming then de facto closed to shooting for

the rest of the open season.

This is where the shoe pinches the heel of the military man

very hard, as also, of course, that of his civil brother when

shooting on areas outside of his jurisdiction. ‘To remedy a state

of affairs which is undoubtedly a real grievance, I suggest that

the number of individuals to be shot in a particular block or

area in any one year should be allotted in a fixed proportion

throughout each month of the close season for that animal, say

2 or 3 or 4 per month, according to the total number notified as

shootable during that season, any balance remaining from any one

or more months being, of course, carried forward and distributed

throughout the remaining months of that shooting-season.

This would give the hot-weather military sportsman, who in

pursuit of his favourite pastime is ready to put up with many and

decided discomforts, an equal chance with his civil brother, who

is not so tied during the cold weather. The suggestion is made

simply with the idea of giving a fair chance to all.

But I would suggest a further step. I would allot a certain

proportion of the head of a particular species to be shot in an area

to the local District officials, the balance going to the outside

sportsman. The District officials could be left to make their own

arrangements as to when their proportion of head was to be shot,

but I think that in the case of the outside sportsmen the

number to be shot should be allotted throughout the shooting

months, so as to give an equal chance to all the block-holders.

No reflection is intended on the District officials by any of these

suggestions. They are made only in the interests of that

particular quality all Englishmen pride themselves in possessing

Fair Play.

4. On the Moulting of an Arctic Fox (Vulpes lagopus) in the

Society’s Gardens. By R. I. Pocock, F.R.S., F.L.S.,

F.Z.S., Superintendent of the Gardens.

[Received October 24, 1911: Read November 7, 1911.]

(Text-figures 6-13.)

The following account of the seasonal change of colour and the

phases in the moult of an Arctic Fox (Vulpes lagopus), which had

been living in the Society’s Gardens since 1904, is based on

observations made during the summer of 1906.

The white winter coat was retained unchanged until the middle

of May. In the latter half of that month it began to come off on

the outside of the fore and hind legs, on the back of the ears, on

the muzzle, and on the sacral region in front of the root of the tail.

By the end of May the sacral patch had extended as a narrow strip

down the thigh, joining the moulted area on the outer side of

the lower portion of the leg. The hair was thinning on the back

of the head, and as the white coat parted with the movements of

56 MR. R. I. POCOCK ON THE

Text-fig. 6,

Arctic Foxes showing the persistence of the thick coat in both “ white” and “ blue

varieties until the early summer, May 5th.

Text-fig. 7.

Arctic Fox showing the beginning of the moult. June 5th.

MOULTING OF THE ARCTIC FOX. 57

Text-fig

Arctic Fox showing the shedding of the white coat over the hind quarters,

June 15th,

Text-fig. 9,

Arctic Foxes, the white specimen showing the further shedding of the coat over

the hind quarters and along the back. June 18th,

58 MR. R. I. POCOCK ON THE

Text-fig. 10.

Ayctic Fox showing the shedding of the white coat along the back. June 18th.

Text-fig. 11.

Arctic Fox showing the white coat adhering to the tail, to the throat, and to the

sides of the body. June 21st.

MOULTING OF THE ARCTIC FOX, 59

Text-fig. 12.

Arctic Fox with moult nearly complete, the white coat still adhering to the tail,

but on the body reduced to a small patch behind the shoulder. July Sth.

Text-fig. 13.

Arctic Fox with the dark coat almost entirely replaced by the new white coat.

October 16th,

60 MR. D. SETH-SMITH ON THE

the animal, the dark summer coat could be seen beneath. This

condition lasted until about the middle of June. The change

thereafter proceeded more rapidly. The coat was moulted all

along the middle line of the back from the root of the tail to the

nape of the neck. It then came off on the sides of the neck, the

shoulders, and the belly, leaving a white mat on the throat, the

sides of the body, and tail. By the end of the first week of July

the patches on the throat and on the sides were much reduced,

but the tail was still white. By the middle of July the animal

was in full summer coat. The summer coat lasted without clearly

marked change for about two months only, but, owing to the

gradual nature of its transition to the winter coat, it was not

possible to state exactly when the change began. Like the

winter coat, the summer coat was moulted. It did not, however,

come off in tufts, but was gradually replaced uniformly all over

the body, the first indication of the replacement being an apparent

increase in the thickness and paling in the colour of the summer

coat. To ascertain that the summer coat was really being shed,

it was found necessary to catch the animal, when by pulling the

coat it was found that the dark hairs were coming away.

The changes in the colour and coat of this animal were watched

for several: years after 1906, when the photographs were taken,

and practically no variation in the details of the process or in the

time of their occurrence was observed.

In a blue variety of the Arctic Fox which lived for several

years with the one above described, the coat was shed in the same

way and at the same time as in the other, but there was no

change of colour except such as accompanied the replacement of

the faded winter coat by the darker-tinted new summer coat.

For neither of these specimens was the locality known; but

several specimens of the species from Hudson’s Bay that the

Society possessed in 1910-1911 turned white in the winter.

I am indebted to Mr. P. W. Farmborough, F.Z.S., for permission

to reproduce the unique series of photographs illustrating the

facts recorded in this account, with the exception of text-fig. 13.

5. On the Moulting of the King Penguin (Aptenodytes

pennanti) in the Society’s Gardens. By Davin Srru-

Smiru, F.Z.8., M.B.O.U., Curator of Birds.

[Received October 21, 1911: Read November 7, 1911. |

(Plate I.*)

Under the above title Mr. W. E. de Winton published a paper

in 1898 7 in which he described the moult of a specimen of A pteno-

dytes pennanti, which the Society at that time possessed, the only

previous paper on the moult of Penguins being one on the change

* For explanation of the Plate see p. 62.

+ P. Z. S. 1898, p. 900.

a - ai re aa 1

4 j oe

i =

' : pan se x i f sb ox

ae f ¥

F4 .

al * a

ae ee is eee eet ee

MOULTING OF THE KING PENGUIN. 61

from the immature to the adult plumage of Humboldt’s Penguin

(Spheniscus humboldti) by the late Mr. A. D. Bartlett *.

[ propose to supplement these papers with some observations

I have made on the moult of a King Penguin recently acquired

by the Society.

The specimen observed by Mr. de Winton did not moult until

it had been in the Gardens some sixteen months, when it began

to look “ seedy,” the feathers losing their lustre and the bird

sitting moping with half-closed eyes. This state of things went

on for fully a month before any feathers were shed. When it

began to moult the feathers of the back and wings became

‘“as brown as withered leaves, so that the bird looked as if it

were covered with mud.” The tail-feathers were the first to be

shed, and it was observed that the bird assisted the removal of

the feathers by pushing them off with its beak. ‘This moult took

place in August-September and occupied about a month.

The example from which I have taken my notes was presented

to the Society by Senor Clemente Onelli, and arrived at the

Gardens on February 11th, 1911, since which date it has gone

through two complete moults, the first in March, and the second

in August-September. The moulting of this species seems to be

very irregular in captivity. ‘The bird observed by Mr. de Winton

arrived in the Gardens in June 1897, but did not begin to

moult until August of the following year. It lived until

January 1903, but did not repeat the process of moulting, its

plumage gradually fading to a sandy-brown colour, so that when

two new birds were received in 1902 they were hardly to be

recognised as belonging to the same species. Our present

specimen has, as before mentioned, completed the process twice

in six months.

This irregularity is doubtless due to the changed climatic

conditions under which the birds live here. In all probability

they normally moult once during the year, but on this point we

have no evidence. On both occasions upon which our present

specimen has moulted here, the process was completed in just

four weeks. In neither case did the plumage appear faded to any

appreciable extent before the moult, the feathers being merely

somewhat dirty and stained with fish. Before moulting the bird

took an extra quantity of fish, devouring as many as fifteen herrings

a day, but just before the moult commenced its appetite failed

and it was with difficulty that the keeper persuaded it to take

even one fish a day. The bird began to look very puffed out all

over and its eyes became dull.

The tail-feathers were the first to be shed, and the feathers

began to come off from the lower regions generally and the

process to be gradually continued upwards. The undersides of

the wings, however, were clean moulted at an early period.

It was observed that the feathers were not shed easily, but had

* P.Z. 8. 1879, p. 6

62 ON THE MOULTING OF THE KING PENGUIN.

to be helped by being pushed off by the bird’s bill, or scratched off

with the feet. The new plumage was apparently fully grown before

the old feathers were shed, this giving rise to the very puffed out

appearance of the bird just before the actual feather-shedding

commenced. The throwing off of the old plumage proceeded

gradually in an upward direction, and when it had been going on

for about a fortnight the bird presented the appearance of wearing

a cape (PI. I. fig. 1), the lower regions being clean-moulted while

the old feathers still adhered to the upper parts. The bird con-

tinued to rub off the feathers where it could reach them with

its bill, the continual moving of the head and neck causing

those on the throat to fall off and leave a triangular patch of

the new plumage visible.

Bartlett noticed that in the case of Spheniscus the feathers of the

flipper-like wing flaked off in masses, and I took particular notice

of the King Penguin to see if the same phenomenon occurred,

and with the aid of the keeper, Pace, was able to secure a clump

of shed feathers which had come off en masse from the wing (fig. 2).

The feathers on the forehead were pushed up into projecting

ridges, which can be seen in the photograph (fig. 1), by the new

plumage and probably flaked off in a mass, though we were un-

able to find them. Most of the feathers of the head and neck were,

however, scratched off with the bird’s feet, the photograph (fig. 3)

- showing the process of removing these feathers. Figures 3 and 4

show the bird nearly clean-moulted, patches of the old plumage

remaining only on the back of the head, the chest, and the back,

parts that could not easily be reached with the bill. Those on

the head were finally scratched off with the feet, but how the

others were removed, or whether they actually fell off finally

without assistance, is uncertain.

The appearance of the shed feathers (fig. 5) is remarkable, the

main shaft being extraordinarily broad and flattened out like a

scale. The stem is bent away from the shaft almost at right

angles, and to the end of the shaft is affixed the sheath of the

new feather which, in growing, has pushed off the old. This

accounts for the fact of the feathers failing to fall off easily, but

having to be helped off by the aid of the bird’s bill or feet. The

two feathers at the top of the figure are from the tail, the bottom

one from the back, and the others from the abdomen.

EXPLANATION OF PLATE I.

Moult of the King Penguin (Aptenodytes pennanti), from photographs

taken by the author.

Fig. 1. Two weeks after commencement of moult.

Fig. 2. Moulted feathers from the wings.

Fig. 3. Three weeks after commencement of moult. Scratching off the old

plumage from the head.

Fig. 4. Same period as fig. 3.

Fig. 5. Moulted feathers of the King Penguin.

1D. 4H. {), MEN. I. 10.

PAIRED OVARIES OF ACCIPITER NISUS (2a-14:a)

AND FALCO TINNUNCULUS (1b.- 8b)

Sa SARS ES

SSNS.

SS =

PAIRED

1912, Pl. I.

CIRCUS, FALCO, ETC.

OVARIES OF

IP Bo 8 WIA, 2, WY,

OWNRUWES) O15 U. ANCCIPMNBIR INSUS., 2. \CMRCUIS) CWAINIE US.

So PODICIFIES CRISTAMUS.

PBS, ISB, 2h. Ww,

OVARIES OF CIRCUS CYANEUS

ON THE OVARIES IN CERTAIN BRITISH BIRDS. 63

6. On the Presence of two Ovaries in certain British Birds,

more especially the Faleconide. By 'T. HE. Gunn, F'.L.8.*

[Received October 23, 1911 ; Read November 7, 1911. ]

(Plates II.—-V.7)

In all vertebrated animals the reproductive organs or gonads

are paired symmetrical glands lying to the inner side of the

Wolttian body and in front of the kidney in the dorsal aspect of

the body-cavity.

In the early stages of development these paired glands are

found in every vertebrate from Fishes upwards. Morphologically

indifferent at first, they become differentiated later on into the

essential male (testis) or female (ovary) glands.

In exceptional cases a further change takes place leading to the

suppression of one of the female gonads with its appendage, e. ¢.,

certain Elasmobranch fishes, and—it is said—the whole class

Aves. The suppression is supposed to be connected with the

large size of the eggs developed by the individual, e. g., Scylliwm

in fishes and Uria in birds.

In the embryo chick (of the fowl) the ridge of germinal

epithelium is sufficiently differentiated by the fifth day to determine

whether the primordial germ-cells of which it is largely composed

are destined to furnish the individual with the permanent ova

characteristic of the hen, or the spermatozoa of the cock bird : in

other words, the ovaries are distinct from the testes at this

date.

With regard to the male organs, their development pursues a

normal course for the remainder of embryonic life, and the male

chick steps out of the shell with two symmetrically placed and

permanent functional testes.

With the female organs a further negative change takes place.

The development of the ovary on the right side comes to a

standstill and the organ atrophies, leaving little or no trace of its

former presence. The oviduct dwindles with the ovary and is

finally lost almost entirely; vestiges may be found more especially

at the cloacal end of the tube, but it is never complete from end

to end as a functional duct.

Exactly when this degeneration begins, and at what period of

embryonic life the chick will be found with only the single left

ovary, are apparently uncertain ; but it is an accepted fact that

the newly hatched female chick has but one ovary—that on the

left. side—and this one gland has to serve for the purposes of

reproduction during the whole of her ovum-bearing existence.

Morphologists and ornithologists alike are agreed that adult

female birds have but one functional ovary. Very few authors

* Communicated by F. Mentrerry Oativiz, F.Z.S.

+ For explanation of the Plates see p. 79.

64 MR. 'T. E. GUNN ON THE

make any reference even to remains of the right ovary being

found on dissection, much less to a perfect functional organ,

though they may allude to the persistence of portions of the

Miillerian duct on that side.

Balfour * writes :—“ In birds the left ovary alone is found in the

adult, and is attached by the mesovarium to the dorsal wall

of the abdominal cavity on the left side of the vertebral

column.”

Milnes Marshall? :—“ In the embryo fowl there are two ovaries,

but in the course of development the right ovary disappears

and in the adult hen the left ovary is alone present. Of the

two oviducts the right one is rudimentary ; the left one, which

alone is functional, forms in the adult hen a wide convoluted

tube.”

And again ¢ :—“ In the female, or hen bird (of the fowl) .....

the Miillerian duct of the right side, like the ovary,

disappears, though traces of it may persist in the adult—the

left Mullerian duct becomes the oviduct.”

Wiedersheim § :—“ In Sauropsida, as in other vertebrates, the

form of the gonads is influenced by that of the body ; thus

in Chelonians they are broad, while in the snake and snake-

like lizards they are more elongated and, as well as in other

lizards, are asymmetrical, the organ of one side lying more or

less in front of that of the other.

“More room is thus obtained for the development of the

ovaries, and in cases where the eggs are very large the organs

of one side tend to disappear as in certain elasmobranchs.

In birds, for instance, the left ovary only is completely

developed and functional... .

“Tn birds the right oviduct as well as tha right ovary

becomes more or less completely degenerated.”

Lillie || :—‘*‘The organs of reproduction of the hen are the ovary

and oviduct of the left side of the body. Although the

right ovary and oviduct are formed in the embryo at the

same time as those on the left side, they degenerate more

or less completely in the course of development so that only

functionless rudiments remain.

‘This would appear to be correlated with the large size of

ege and the delicate nature of the shell, as there is not

room for the eggs side by side in the lower part of the

body-cavity.”

Alex. Macalister 4] :—‘‘ There are two fasciculate ovaria, of which

%* ©Blements of Embryology, p. 11. Foster and Balfour, 1883.

+ ‘Vertebrate Embryology,’ pp. 228-229. Milnes Marshall, 1893.

{ Ibid. p. 320. fy.

§ ‘Comparative Anatomy of Vertebrates,’ p. 474. Wiedersheim (adapted by Prof.

W.N. Parker, 1907). ; se

\| ‘ Development of the Chick,’ p. 21. Lillie, 1908.

€ ‘An Introduction to the Systematic Zoology and Morphology of Vertebrate

Animals,’ (organs of reproduction in Birds) pp. 159-160. Alex. Macalister, 1878.

OVARIES IN CERTAIN BRITISH BIRDS, 65

only one (the left) is developed and functional. The

right oviduct remains a hydatid; sometimes is developed

anomalously, while the left atrophies; rarely the two remain.

(Pigeons and Parrots).”

Bowdler Sharpe *:—“The right ovary of birds is always

atrophied, and it is only in rare cases that rudiments of it

are found (namely in the diurnal Raptores). The right

oviduct is not so completely atrophied as is the ovary of the

same side.”

In the recently published ‘Grouse in Health and Disease’

Dr. EK. A. Wilson writes (of that bird) t:—“There is but one

ovary, and it lies always on the left side of the backbone

of the bird. This development of the ovary, only on one

side, is the reason for advising the examination to be made,

as detailed above, on the left side always.”

This last quotation is interesting in connection with the subject

of paired ovaries, since over 2000 grouse passed through the hands

of the Grouse Disease Committee and were carefully examined ;

of these nearly a quarter (476) were females, and not a single

example of a persistent right ovary or its remains was found ¢.

It is to be noted, however, that if the sexing were carried out, as

Dr. Wilson recommended, by examination of the left side, persistent

right ovaries might have been overlooked.

Newton $:—‘ In the female a pair of ovaries are developed, but

with rare exceptions only that on the left side becomes

functional. In young birds both oviducts are almost equally

developed, but the right one soon becomes reduced to an

insignificant ligamentous strand along the ventral side or

part of the kidney.

“This one-sided suppression of the organs may possibly he

referable to the inconvenience that might be caused were

each oviduct to contain an egg ready to be deposited.”

From the above quotations it will be seen that the majority of

the writers do not admit the presence of a right ovary at all ir

the adult female. Sharpe speaks of rudiments || of the righ.

ovary being retained in the diurnal Raptores, Newton very

guardedly refers to ‘rare exceptions” where both gvaries are

present, but he does not mention in what species of bird these

exceptions were found. It is indeed rather daubtful if these

instances came under his personal observation; had they done so

he would probably have published the names of the species and

* Cassell’s Natural History, vol. iii. p. 251. KR. Bowdler Sharpe, 1883.

+ ‘Grouse in Health and Disease,’ vol. i. p. 60.

t Out of 17 female grouse that have passed through my hands in the last foyr

years, three have shown well-marked paired ovaries.—T. E. G.

§ ‘Dictionary of Birds,’ (“reproductive organs”) p, 782. Alfred Newtan, 1894,

|| Italics are mine (T. E. G.).

Proc, Zoou. Soc.—1912, No. V. 5

66 MR. T. E. GUNN ON THE

details of the parts as he found them. Rev. F. C. R. Jourdain *

suggests that Newton's exceptions might have been drawn from

a German source 7.

Professor Taschenberg, the writer of the article referred to by

Jourdain, says :—

“Tt happens not very rarely that remains ¢ of the right ovary

are retained, especially in the sparrow-hawk and _ buzzards,

much more irregularly in other accipitres, and still more

rarely in the owls.”

Oeteris paribus, a bird possessing two functional ovaries 1s clearly

more fitted for the reproduction of its kind than the bird with

only one. If disease or injury destroys the one ovary and spares

the other, the bird can still fulfil the chief duty of its life,

the bearing of ova. If the one ovary becomes exhausted (¢. g.

destruction of first nests) the other would act as a reserve to

draw on in eases of necessity. If the fittest female is to survive

in the long run, one would suppose that the bird that possessed

the two complete genital tracts, the foundations of which were

laid in the very early embryo, would surpass its fellow which had

squandered fifty per cent. of its reproductive capital while still in

the shell. That that is not the case we know.

Speaking in general terms, the rule holds good that adult

females have but one ovary and one oviduct, those on the left

side.

Why has the second ovary been abolished ¢

Tt has been suggested §, with some show of probability, that the

suppression is in connection with the passage of the egg through

the oviduct; that the danger involved by two eggs engaging

simultaneously in the two oviducts would be very great, and

would lead to fracture of the delicate shell, or rupture of the walls

of the containing duct; to peritonitis, intestinal obstruction or

some such erlamity, and in any event death.

Tf such an accident—a fully-developed egg in the lower part of

either oviduct at the same time—were to happen, no doubt the

results would be untoward. A Guillemot with two full-sized

eggs in its abdomen would presumably burst.

But is it necessary to suppose that such a sequence would follow

the presence of two separate genital tracts ?

IT think not. The number of cases collected in this paper goes

far to prove that death and disaster are neither the necessary

nor even the common result of paired ovaries. It is contradicted

by the frequency with which the Falconide are found with paired

ovaries which are obviously functional. That a single ovary

permits the bird to perpetuate its kind is true enough. The

+ ‘Natureeschichte der Vogel Mitteleuropas,’ vol. i. p. 60, 1905. [ Dr. Gadow, in

Bronn’s ‘ Thier-Reich, Vogel,’ p. 842, quotes from Stannius, and adds mstances from

his own observation of the persistence of the right ovary, particularly in diurnal

Lirds of prey —Epiror P.Z.8. }

¥ Italics are mine (T. E. G.).

§ Newton, ‘ Dictionary of Birds,’ p. 783.

OVARIES IN CERTAIN BRITISH BIRDS, 67

excretion of urea can be carried on by one kidney if the other be

destroyed by disease, or removed in its entirety by the surgeon;

but the animal with one kidney only is admittedly less well off

than the animal with the normal pair. The life would be called

a bad one at any vertebrate life insurance office, and the premium

charged would be correspondingly high. ‘Two ovaries may not be

a necessity, but they must be better than one. The suppression

of the second ovary appears to be a retrograde step for which it is

hard to find any adequate reason, and is almost without a parallel

in vertebrate embryology.

Tn dissecting an immature female Sparrow-hawk (Accipiter nisus)

(plumage of the first year) shot on the 9th of January, 1892, [

found both ovaries equally developed, containing small eggs of

uniform size.

In the following April a second example came under the notice

of Mr. F. Menteith Ogilvie, who had happened to be with me

when I skinned and examined the former specimen. He made

the following note :—“ April 11th, 1892, I dissected A. nisus 2

Both ovaries well developed, eggs numerous and of various sizes_—

I think the forward condition of the ova shows the bird would

have bred this season, though it was certainly only a last year's

bird.”

From 1892 up to the present date I have paid special attention

to the sexual organs of all birds passing through my hands, in

order to investigate the condition of the female generative system,

the frequency with which the right ovary was found to persist,

and the species in which such persistence occurred. In every case

where paired ovaries were found I took notesand madeas accurate

sketches as I could with the specimen on the table before me.

In 1895 and again in 1903 I recorded a number of these

instances in two papers read at meetings of the Norwich Science

xossip Club*, Since then further examples have been added,

bringing the total number of specimens with paired ovaries in my

series to.45, These are grouped in bulk in Table I. (see p. 72),

and are separately treated in detail in the Appendix.

Neither Table I. nor the Appendix includes several instances in

which I have found the two ovaries in certain nestlings,

On July 7th, 1909, six nestling Sparrow-hawks with their

parents (second year’s plumage) were sent me from Snffolk

(see Appendix 10"). In sexing the nestlings I found that five

were females and the sixth, a much smaller bird, a male. All

the five female nestlings had paired ovaries, the glands averaging

half an inch in length. ‘They were equally developed and easy to

recognise. ‘The excessive proportion of females over males in the

Sparrow-hawk in this instance is noteworthy—I have observed

the same fact on a former occasion t, in which, out of six nestlings,

four were females and two were males.

* Report of Proceedings, May 1895 ; May 1903.

+ ‘ Zoologist,’ 1885, p. 51.

68 MR. T. E. GUNN ON THE

Exceptions to the rule that breeding females possess but a

single left ovary must be very considerable. Since especially

looking for paired ovaries I have found them comparatively

common—the natural inference is that for many years I had

overlooked them.

The number of examples of paired ovaries, apparently functional

and ready for ovulation in this series, negatives the idea of the

danger to life necessarily involved by the possession of a paired

generative system. If death were the common result of a right

and left ovary (with the corresponding ducts) it would seem likely

that specimens would have come to hand before now demonstrating

the pathology of the fatality and its mode of occurrence. I have

never handled such a bird in the flesh, nor seen any reference in

print to dissections illustrating death from this cause.

Is it not a reasonable conception that the nervous system

would govern the activity of the ovaries—and of course the

oviducts—during the period of turgescence associated with the

breeding season, utilizing the left ovary only, or the right ovary

only for the egg-supply and the other for a reserve—or in cases of

necessity utilizing both ovaries

(a) In most of the later examples of paired ovaries in a state

of activity in my series—birds that were obtained in June and

early July and had then laid their full complement of eggs for

the season—all the evidence goes to show that though two ovaries

were present only one has been utilized for the production of eggs

(cf. 1°: 8%); the other ovary developing eggs up to a point—the

eggs then ceasing to grow in size and finally shrinking with those

in the working ovary to the small undeveloped ova common to

birds in the non-breeding season.

In some of the earlier examples—birds obtained between March

and early May (cf. 5°; 1%; 2*; 2")—both ovaries are well and

equally developed with eggs of approximately equal size in either

ovary—so that at this time it is impossible to guess which ovary

is to furnish the season’s eggs, or whether the supply is to be

drawn from both.

This seems to point to the nervous system determining which

ovary shall be finally selected for the year’s output.

Both ovaries are developed up to a certain point; and then

one comes to a standstill. while the requisite number of eggs

in the other continue to increase in size until they are ripe for

the oviduct. After they are shed both ovaries rapidly undergo

the normal process of involution and become comparatively

insignificant.

According to this theory the regulating nervous stimulus could

be switched off one ovary and on to the other as best suited the

requirements of the organism.

(b) If both ovaries are to share in the production of one clutch

of eggs, then one can imagine each oviduct in turn being inhibited,

while an egg is engaged in the duct of the opposite side, much as

OVARIES IN CERTAIN BRITISH BIRDS. 69

a signalman refuses to accept a new train until his section is

cleared of the old one.

serve two ovaries. There is not much evidence for or against this

supposition.

In some cases the appearance of the paired ovaries rather

suggests that both had taken part in discharging ripe ova, while

only one oviduct shows much development.

On the other hand, the normal involution of the oviduct is so

extraordinarily rapid, that in some specimens with well-developed

ovaries, which are known to have passed the last eggs of their

clutch quite recently (3°), the oviducts on both sides are merely

represented by ill-defined strands quite difficult to differentiate

macroscopically from the surrounding tissues, and but little more

marked than the oviduct (or ducts) would be in the autumn and

winter months.

The upper portion of the oviduct is provided with a mesentery

which is sufticiently long to allow a very considerable amount of

free play. There seems no physical objection to the open end of

the tube (ostiwm abdominale) swinging across the mid-line of the

spine *, and grasping the ripe ovum of the opposite ovary with

nearly the same facility as the ovum of its own side.

Extra-tubal gestation, by which I mean the shedding of a ripe

ovum into the peritoneal cavity, is, so far as we know, an

exceedingly rare accident among birds. Dr. Wilson? mentions

such a case.

Two functional ovaries and one duct collecting eggs from both

sides would be exactly the conditions which would predispose to

such an occurrence.

Returning to the paired ovaries, it is manifest that these cases

are not exceedingly rare among many quite widely separated

groups of British birds. In one family, the Falconide, such

instances may be termed positively common. One cannot help

realising that if 45 examples come under the notice of one man

in the course of nineteen years, double ovaries must be something

more than of occasional occurrence.

The obvious conclusion seems to be that they are not found

because they are not looked for.

In the writings of English ornithologists I can find but one

specific record (and that a very recent one) of paired ovaries.

Dr. GC. B. Ticehurst ¢ deseribes three cases in which he found

the right ovary persisting—all three examples were Sparrow-

hawks.

* Except in the Falconide the examples of paired ovaries generally show an

asymmetrical state of affairs, with the right ovary below the lett, and at the same

time carried over somewhat laterally from right to left. In such cases as these the

left oviduct would have no further to go for the eggsfrom the right ovary than ior

those from the left.

+ ‘Grouse in Health and Disease,’ p. 183.

+ ‘British Birds,” November 1910, p. 188.

70 MR. T. E. GUNN ON THE

Some very excellent notes on the subject by Signora C. Picchi

appear in the July (1911) number of ‘ British Birds,’ pp. 45-49.

This lady gives 27 examples of a persistent right ovary, the

large majority of them (25 out of 27) are found among the

Falconide, especially the genera Falco and Circus.

For the purposes of this paper I have reduced Signora C. Picchi’s

notes to a tabulated form (‘Table I., p. 73) in order to render it

readily comparable with the table (Table I.) constructed from my

own series.

Ornithologists expect to find either two testes or one ovary. The

intestines and peritoneal folds are raised and gently turned over

from the (bird’s) left to right, which of course exposes the left half

of the dorsal wall of the body-cavity. In the case of the female

the left ovary is satisfactorily identified, and there the dissection

usually stops.

The conditions obtaining on the right side of the body-cavity

are not investigated at all,—a second ovary may or may not be

present, but by this method of examination it is effectually hidden

by the pile of intestines pushed over from the left side.

The percentage of double ovaries found in this paper in certain

genera in the family (Ci ircus 9 out of 12, Accipiter 14 out of 20)

is so large that one is forced to the conclusion that individuals

of the same genera dying in captivity would also furnish a

considerable percentage of paired ovaries.

In the 45 instances of double ovaries referred to (Table I.,

group 1) nearly three quarters (33 out of 45) are of one family,

the Falconide.

In Signora C. Picchi’s notes (Table II., group 1) 27 examples

are given, and of these 25 are drawn from the same source.

The remaining twelve examples in my series (Table I., group 2)

represent seven species that, with the exception of the two Grouse,

are widely separated. The seven species in fact belong to six

different orders.

The remaining two in Signora C. Picchi’s notes (Table I.,

group 2) also occur in two very diverse species, one of them being

an Owl (Striges) and the other a Rook (Corvus).

Oddly enough, I have no note of paired ovaries among the

Strigide ; but that may be due to so few of these birds passing

through my hands in recent years, owing to the wise protection

which is now almost univer sally sor ded owls in this country.

Table I. (T. E. G.) and Table II. (C. Picchi) show clearly

enough that the right ovary does persist in not a few cases; that

it is sometimes developed equally with the left and sometimes very

considerably exceeds that gland.

1° (Circus cineraceus), in which the right ovary has several large

ripe ova whereas those of the left ave small.

8° (dAccipiter risus). The right ovary is but fragmentary, and

shows only three ova: one as large as an ordinary sized marble,

another about one third that size, and the third about as large as

hempseed, ‘Those of the left were quite small. The right

ovary had evidently been the source from whieh the season’s eges

OVARIES IN CERTAIN BRITISH BIRDS. 71

had been derived, and had become nearly exhausted : the left ovary,

on the other hand, showed little more activity than the quiescent

gland of the autumn months.

7 (Falco tinnunculus). The left ovary contained a cluster of

eggs of even size with one slightly larger—the left oviduct well

developed. The right ovary contained but six very small eggs,

and the oviduct was represented by only a narrow ligamentous

strand,

14" (Accipiter nisus). Both ovaries were present, the right half

the size of the left, and contained small eggs only.

Of the oviducts the right was double the width of the left and

had the appearance of having recently passed eggs.

12° (Accipiter nisus). Both ovaries and their corresponding

ducts present, the left oviduct being the wider.

This female had probably quite recently passed her full clutch

of six eggs, which were taken from the nest by the keeper at the

same time that he killed the parent bird. ‘The remaining eggs in

both ovaries were all small. The left oviduct had quickly become

reduced in width after passing the last of the six eggs, which event

must have taken place not more than two days before.

From the greater width of the left oviduct in 7, of the right in

14°, of the left in 12°, we may reasonably infer that they had

severally been the latest oviducts in active service.

This indicates that either the right or the left oviduct may be

the functional one—and that one or other may work as the

maturity of the ova in each respective ovary requires their

service.

In conclusion I would again draw attention to Tables I.

and IT.

In Table I., to take two extreme instances, out of sia female

Hen Harriers (Circus cyaneus) examined five had a persistent right

ovary : out of fifty female Woodcock (Scolopax rusticula) examined

only one retained the gland on the right side.

If the figures in the two tables are combined we get the following

results :—

Grand total, 72 examples of paired ovaries out of 212 females

examined, made up of two sub-divisions A and B.

Poe, MOE kefopato Gy GRA Aree aoe See Re POn ES Pe aoe 56 out of 98.

B. Species from any family other than the

i the Verornt (0 (2 = Sos eee ee ee a , Bees Vivien, LAOud, OF LAD)

This shows in rather a startling manner the frequency with

which paired ovaries are found in the Falconide (or rather in

the genera which have come under examination) as compared

with any other family of British birds.

There is a second point of some interest in regard to the

anatomical position of the ovaries. In the Falconide these are

symmetrically piaced and are arranged, almost without exception,

parallel to each other, and on about the same level, lying on

the kidney of the same side and separated from each other by the

vertebral coluinn,

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greater part

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ovary more or less overlapping the other.

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to its final disappeatance:

importance as furnishing more space for the development of the

uncertain.

ovaries.

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niversity of

original drawings

74 MR. T. E. GUNN ON THE

APPENDIX.

SparRow-Hawk (Accipiter nisus). ai

(PI. II. figs. 29-14"; Pl. IV. fig. 1.)

12, Date, January 9th, 1892. Locality, Suffolk. Age 8 to 9

months. Both ovaries present, each containing small eggs.

2, May 21st, 1894. Suffolk. In second year’s plumage.

Paired ovaries of equal size, being filled with uniform eggs.

3°, November 8th, 1900. Norfolk. In second year’s plumage,

showing both ovaries: eggs small, and uniform in size.

4°, December 20th, 1906. Derbyshire. Two years of age.

Right and left ovaries nearly equal in size. Right a trifle the

larger, both filled with cream-coloured eges.

5¢, February Ist, 1907. Suffolk. Entering its second year.

Both ovaries present, the right being larger, and the ova uniform

in size.

62, October 3rd, 1908. Suffolk. In its first year’s plumage.

Two small thin ovaries present. Right larger than left. Eggs

very small.

7*, October 7th, 1908. Norfolk. First year’s plumage. ‘Two

ovaries present, right trifle the larger. Eggs small.

8", May 7th, 1909. Suffolk. In its second year’s plumage.

Two ovaries with eggs, the right having three eggs, one as large

as an ordinary marble, the second about one third its size, and

the third about the size of a hempseed. The left ovary contained

two as large as hempseed, the remainder very small,

92, * June 9th, 1909. Suffolk. Two years old. Two ovaries

present, the right being larger and containing one large egg, the

remainder very small and presenting three different colours, pale

yellow, cream, and black.

102. July 7th, 1909. Suffolk. In second year’s plumage.

Two ovaries present, the right as large again as the left, both

containing small cream-coloured eggs. ‘The full clutch of six

nestlings was sent with the parent birds. Of these five were

females and one a male; all five females had well-defined paired

ovaries.

11. May 26th, 1911. Suffolk. Two ovaries, apparently

exhausted, and too decomposed to preserve.

122,* May 26th, 1911. Norfolk. Not less than three years old.

Both ovaries nearly equally developed. The left oviduct much

wider than the right. This bird had laid its full complement of

six eggs; quite fresh when taken by the keeper.

* Preserved as a museum specimen.

+ In the fractions at the head of each species the numerator indicates the number

of examples of paired ovaries found, and the denominator the total number ¢xamined,

¢. €. 99 1s interpreted 20 females of 4. nisus examined and 14 found with paired

ovaries.

OVARIES IN CERTAIN BRITISH BIRDS. 75

132,¢ May 27th, 1911. Norfolk. Age two years. Two

ovaries present and equally developed. I examined this bird the

day after death ; it had been exposed to the sun, so was somewhat

decomposed, but I was able to preserve the ovaries. Its six fresh

eggs were also taken by the keeper.

144,¢ June 8th, 1911. Suffolk. Second year. Both ovaries

present. The left containing small and larger eggs. The right

with small eggs only. The two oviducts well developed, that

of the right being twice the width of the left. The ostium

abdominale of the left duct is well shown in this specimen.

12°

Kustrew (Palco tinnunculus).

(Pl. II. figs. 1-8.)

1>, January 15th, 1901. Norfolk. Adult. Two ovaries

present, but not equal in size, the right ovary being about half

as large as the left, but the oviduct was more prominent in the

right.

2b,* June 7th, 1909. Suffolk. Adult plumage. Two ovaries

present, the right small, the left quite four times as large as the

right. Eggs of two sizes, the larger of a pale cream, a few of the

smaller that were scattered were of a rich yellow colour.

3>,* June 7th, 1909. Suffolk. Adult. Two ovaries, the

right a small dark gelatinous mass with small eggs ; the left about

six times as large and contained eggs of various sizes of a yellow

colour, a dark gelatinous mass being in the centre.

4b, May 24th, 1910. Norfolk. Adult plumage. Two ovaries

about equal in size, the eggs in the right ovary somewhat the

larger.

5b, May 25th, 1910. Suffolk. Second year’s plumage. Two

ovaries present. The right in a fragmentary state but containing

a few eggs of two sizes. The left ovary was about six times larger,

with eggs of various sizes, none large,

6».§ January 6th, 1911. Norfolk. Second year. Both ovaries

present. The left as large again as the right. Eggs small and of

uniform size. Both oviducts thin ; the left very attenuated.

7>,* May 10th, 1911. Suffolk. Two years old. Left ovary

full of eggs of uniform size, with the exception of a single larger

one. Left oviduct exceedingly well developed, right ovary small

and fragmentary; six small eggs only; right oviduct a fine

ligamentous strand.

8>.* May 26th,1911. Suffolk. Age two years. Both ovaries

present. The left larger and full of eggs, the right ovary smaller

with fewer eggs, two of which were, however, larger than those in

* Preserved as a museum specimen.

+ Stereoscopic photograph of the ovaries in situ.

{ Sections of the right and left ovaries; photographs of the same,

§ Stereoscopic photograph of the ovaries in situ.

76 MR. 1. E. GUNN ON THE

the left. Both oviducts represented by thin strands only, the

left being more evident.

Hen Harrier (Circus cyaneus).

(Pl. ILI. figs. 1°-5°; Pl. IV. fig. 2; Pl. V. figs. 1-4.)

le. November 4th, 1893. Norfolk. Second year. Both

ovaries present; eggs of a rich cream colour, small and of uniform

size.

2°, April 9th, 1900. Suffolk. Probably three years old.

This bird possessed two ovaries which contained eggs of two

S1Zes.

3°, January 10th, 1908. Norfolk. In second year’s plumage.

Both ovaries present, the left one as large again as the right.

Both filled with small eggs of uniform size.

4c, January /th, 1907. Suffolk. In its second year. Two

ovaries present ; the left slightly the larger. Full of small

eggs.

“Be, *+ March 30th, 1911. Norfolk. Age two years. Both

ovaries equal, containing cream-coloured eggs of two sizes; the

majority smal], the right ovary having more larger-sized eggs than

the left. The two ovaries were packed closer together than any

I had hitherto seen,

Glu

MarsH Harrier (Circus eruginosus),

(Pl. III. fig. 14.)

1¢, April 17th, 1907. Derbyshire. Two years old. Both

ovaries present, the left a trifle the larger; both full of small eggs

of wniform size.

Montacu’s Harrier (Circus cineraceus).

(Pl. III. figs. le-3e.)

1°. May 15th, 1903. Suffolk. Adult, three to four years old.

Both ovaries present, the right containing much larger eggs

of varying sizes, the largest just half an inch in diameter, and

most of them of a bright orange colour. The left ovary contained

small eggs of a yellowish cream colour.

2°. July 3rd, 1909. Suffolk. Two yearsold, The two ovaries

present. Eggs very small, uniform in size, and of two shades of

colour—pale and deep yellow.

3°. June 17th, 1911. Suffolk. Adult, probably three years.

Two ovaries equal; containing a mass of eggs of various sizes and

* Sections of the right and left ovaries; photographs of the same.

+ Stereoscopic photograph of the ovaries in sitz.

t Sections of the right and left ovaries: photographs of the same.

OVARIES IN CERTAIN BRITISH BIRDS. 77

of pale yellow and flesh colour; the left oviduct a fine strand only,

that of the right very fine and thin.

Hospy (Falco subbuteo).

(Pl. ITT. fig. 14.)

1f, June 6th, 1907. Suffolk. Two years old. Two ovaries

present; the left contained five large eggs, and the right some

very small ones.

PrrecRIne Fancon (Falco peregrinus).

(PI. IIT. fig. 18.)

1¢. November 22nd, 1907. Suffolk. Three years old. Two

ovaries present, the right small and thin, with small eggs; the

left was quite four times larger and had some eggs of two sizes.

Great Cresrep Grese (Podicipes cristatus). .

(Pl. III. figs. 15, 24; Pl. IV. fig. 3.)

1h. * November 26th, 1910. Suffolk. Two ovaries, the right

below the left and overlapped by it; both filled with small eggs.

‘The upper (left) ovary the larger.

oh + February 28th, 1911. Norfolk. Probably three years old.

Two ovaries present, the left slightly larger than the right; eggs

of various sizes and of a yellow colour. The right ovary below the

left and slightly overlapped by it.

. 3

Rep Grouse (Lagopus scoticus). 7

(Pl. III. figs. 13-35.)

li. October 26th, 1907. Argyllshire. Two ovaries, situated

side by side, the right double the length of the left. Both filled

with small eggs of uniform size.

9i. October 26th, 1907. Argyllshire. Two ovaries; same

remarks as above.

3), November 29th, 1910. Arygllshire. ‘I'wo ovaries of equal

size and rounded form. ‘The left ovary is superior and 1s

overlapped by the right ovary, which lies nearly directly below

it.

Brack Grouse (7etrao tetrix), =

@BL OD fe: 1)

1k, August 30th, 1907. Staffordshire. Probably three years old,

* Stereoscopic photograph of the ovaries in situ.

+ Preserved as a museum specimen,

78 MR, T, E. GUNN ON THE

This bird had partly assumed the plumage of the male with its

black breast and forked tail, the rest of the plumage being a

mixture of black cock and grey hen.

Two ovaries present and equally developed, the left slightly

overlapping the upper part of the right ovary ; both contained eggs

of uniform size.

Bewicn’s Swan (Cygnus bewickr). =

(PST ates)

1, January 5th, 1900. Scotland. Adult. It possessed two

ovaries ; one placed in front and nearly covering the other. The

front and larger one was undoubtedly the right ovary and contained

eggs of various sizes; the left was sinnalltors containing eggs of

uniform size.

Water Ratu (Lallus aquaticus).

(BITE figs)

1™.* + December 27th, 1907. Suffolk. Both ovaries present,

the left slightly the larger, and above the right, which it overlaps

at the upper part. Each ovary is filled with small eggs of uniform

size.

2m, December 26th, 1910. Norfolk. Two ovaries equally

developed, and as in previous instance, one is situated immediately

above the other, eggs being uniform in both.

=1/ bo

Wooncock (Scolopax rusticula).

(RUST hea.)

50°

1", December 9th, 1909. Argyllshire. Two ovaries present,

and situated one above the other, the left overlapping the upper

part of the right. This latter ovary is slightly the larger and

seemed partially twisted at its extremity; both full of small

uniform eggs,

Lirrte Guuu (Larus minutus).

(PISeies ae.)

1°. February 5th, 1902. Suffolk. Bird in immature plumage,

i. e. first year. The right and left ovaries are present, and meet

in an apex above with a small blood-vessel overlying the junction.

Both ovaries filled with small eggs.

2°, January 6th, 1906. Suffolk. Adult. Both ovaries present,

and arranged side by side; the right is twice the length and size

of the left, and both are full of small cream-coloured eges

of uniform size.

* Stereoscopic photograph of the ovaries én site.

+ Preserved as a museum specimen,

OVARIES IN CERTAIN BRITISH BIRDS. 79

EXPLANATION OF THE PLATES.

For further details see Appendix (p. 74).

Puate II,

Paired Ovaries.

Figs. 2"-14". Sparrow-Hawk (Accipiter nisus).

1’-8". Kestrel (Falco tinnunculus).

(v., vight ovary. J., left ovary.)

Pruate III.

Paired Ovaries,

Vies. 1°-5°. Hen Harrier (Circus eyaneus).

1", Marsh Harrier (C. @ruginosus).

1-3". Montagu’s Harrier (C. cineraceus).

1. Hobby (Falco subbuteo).

1%, Peregrine Falcon (Falco peregrinus).

1",2". Great Crested Grebe (Podicipes cristatus).

1-3), Red Grouse (Lagopus scoticus).

1". Black Grouse (Tetrao tetrix).

1'. Bewick’s Swan (Cygnus bewicki).

1,2". Water Rail (Rallus aquaticus).

1°. Woodcock (Scolopax rusticula),

1°,2°. Little Gull (Larus minutus).

(r., right ovary. J., left ovary).

PuaTeE IV.

Paired Ovaries.

Fig. 1. Sparrow-Hawk (Accipiter nisus) X 1}. Both ovaries present, the right

Fig.

nearly exhausted and containing small eggs only. Both oviducts well

developed ; the right three or four times the size of the left and apparently

the last in use; the left oviduct is well defined, the ovarian end of the

duct is turned directly downwards and to the left, and the Ostiwn abdo-

minale is clearly seen. (Appendix 14.)

2. Hen Harrier (Cireus cyaneus) X1}. Symmetrical paired ovaries equally

developed and containing eggs of approximately the same size.

(Appendix 5°.)

3. Great Crested Grebe (Podicipes cristatus) X13. Asymmetrical paired

ovaries. The right ovary lies below the left, and is mostly in the lett half

of the body-cavity. (Appendix 2",)

PLATE V.

Microscopie sections of the ovaries of the Hen Harrier (Circus cyaneus).

(Appendix 6°). :

1. Left ovary X4} showing general ovarian structure.

2. A selected ovum from fig. 1 showing germinal spot, protoplasm, germinal

epithelium, and vascular connective tissue capsule (X69).

8. Right ovary «5,

4, A selected ovum from fig, 3 50,

80 PROF. A. D. IMMS ON

On some Collembola from India, Burma, and Ceylon ;

with a Catalogue of the Oriental Species of the Order.

By A. D. Imus, D.Sc., B.A., Forest Zoologist to the

Government of India ; late Professor of Biolony , Muir

College, and Fellow of the University of Mibhae es

[Received May 29, 1911: Read November 7, 1911.]

(Plates VI.-XITI. and Text-figures 14 & 15.)

CONTENTS,

Page

Nemintroductonysemeanks peeeeeeeeeeeemeeneteateernettcetars 80

II. Description of the Species ........ ea eT oe

Ill. A Catalogue of the Oriental Collembelae Be Hieticmasane 118

IV. A Summary of General Conclusions ................:0008 121

VY. Explanation of the Plates ............... FOREN On SoRCE 122

1. Invropuctory REMARKS,

During the last decade the Collembola have attracted a con-

siderable amount of attention both from morphologists and

systematists. Collections of these primitive insects have been

studied from various regions of the world. Hitherto, however,

T am not aware that any species of the Order have been known

and recorded from the Indian Empire. The present paper is

intended as a small contribution towards a knowledge of the

Collembola of that extensive region.

Iam indebted to Dr. N. Annandale for handing over to me

for examination the specimens contained in the collections of the

Indian Museum, and to Mr. E. E. Green for forwarding me two

species from Ceylon. In addition to the species received from the

above two sources, I have myself collected a number of examples

in several parts of India, from Allahabad in the “ plains” up to

an altitude exceeding 12,000 feet in the Himalayas. The area

thus worked over lies within two zoo-geographical regions. The

collecting, however, has only been done during casual intervals,

and gener. ‘ally when I was occupied in searching for other Romaine

of animallife. I wish to acknowledge the facilities for consulting

entomological literature that were “afforded me by the libraries

attached to the Indian Museum, Caleutta, and the Agricultural

Research Institute, Pusa.

Altogether 571 specimens of Collembola have been examined

and from among these, 4 genera and 27 species are described as

new, and 3 species were already known.

* Communicated by A, KE, Surprny, M.A., F.R.S., F.Z.S.

PZ.o. 19l2-PUNV1,

E.Wilson, Cambridge.

A.D.Imms ad nat.del.

ORIENTAL COLLEMBOLA.

P45. 1912. P). Vir.

(TaroKsl

occ

A.D.Imms ad nat.del.

E.Wilson, Cambridge.

ORIENTAL COLLEMBOLA.

PZ lol2 Fl Vik

uae

hy)

WEP

—s

A.D.Imms ad nat.del. E.Wilson,Cambridge.

ORIENTAL COLLEMBOLA.

ie Aes ele kale

E Wilson,Cambridge.

A.D.Imms ad nat.del.

ORIENTAL COLLEMBOLA.

PAu. ISA Pixs

wor a

Wyia

Uh,

Lena

yy iY

ii is YUGVe

it Y

Mi} t

Vy av

ql Nithy i

aN yyy \ =

——__——.

i) " ‘i

WOVE

yates

Arve (i,

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Oak tay SEN

bay

4 t

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VW AW) Af

nit Y, MW

to SM agi

ANA YOR TTR

NUE i

AN Ky Nt

A.D. Imms ad nat.del.

E.Wilson, Cambridge.

ORIENTAL COLLEMBOLA.

PB Z.S. 1912. P1.XI.

Se Ss 3

a SCE

ee SS

SSS

E.Wilson, Cambridge.

ORIENTAL COLLEMBOLA.

A.D.Imms ad nat.del.

PAS? iz (Pl XI,

E.Wilson, Cambridge.

A.D.Imms ad nat.del.

ORIENTAL COLLEMBOLA.

™

f i :

: a

‘ Bis

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ORIENTAL COLLEMBOLA, 81

A.—Palearctice Species.

Entomobrya crassa, sp. 1.

Isotoma siva, sp. n.

Tomocerus vulgaris Tullb,

Sinella montana, sp. n.

Seira frigida, sp. n.

B.— Oriental Species.

NXenylla obscura, sp. n.

Achorutes armatus Nicolet.

Pseudachorutes anomalus, sp. n.

Neanura corallina, sp, n.

V. intermedia, sp. n,

iV. pudibunda, sp. n.

Jsotoma nigropunctata, Sp. Nn.

Heteromuricus cercifer, gen, et sp. n.

Isotomurus palustris Miill.

Lepidocyrtus robustus, sp. n.

Entomobrya kali, sp. n,

HL. kali var. lutea, nov.

Seira brahma, sp. n.

Pseudosira indra, sp. v0.

Dicranocentroides fasciculatus, gen. et sp. n.

Cremastocephalus montanus, sp. n.

C. indicus, sp. n.

Paronella borneri, sp. n.

P. travancorica, sp. n.

P. gracilis, sp. n.

P. phanolepis, sp. n.

P. insignis, sp. n.

Idiomerus pallidus, gen. et sp. n.

Cyphoderus simulans, sp. n.

Pseudocyphoderus annandalei, gen. et sp. n.

Sminthurides appendiculatus, sp. n.

The fact that so large a proportion of the species are new is

scarcely remarkable, owing to the great diversities of climate and

soil found in the various areas from which the specimens were

obtained ; vide text-fig. 14, p. 82.

The limits of the northern boundary of the Oriental Z00-ge0-

graphical region are difficult to define owing to the fact that

members of the Oriental fauna penetrate, in places, for long dis-

tances up into the hot confined valleys of the Himalayas; while,

on the other hand, the Palearctic fauna descends the southerr

slopes of that range to within the limits of forest growth. As

Blanford * remarks, above the limit of forests the fauna is purely

Palearctic, all the Oriental types having disappeared, he forest

* “The Distribution of Vertebrate Animals in India, Ceylon, and Burma.”

Phil. Trans, Roy. Soe. vol. 194, 1901, p. 347.

Proc, Zoou, Soc.— 1912, No. VI. 6

82 PROF. A. D. IMMS ON

limits in the E. Himalayas may extend up to an elevation of

14,000 feet, while in the W. Himalayas it attains in places an

altitude no higher than 9000 feet.

Text-fig. 14.

Se Ms ae 3

NaraGhal. Skunseon ee,

Sel Betal (Wovofe ae

Ganger R Z fo

Allahabad oe SS a

cats Dh

f Na

Rhambade aa S <

) \

a oe: ie \ve Moulmein

c \

Weare mel

\ / )

f {

So q \

Map showing the localities from which the Collembola treated in this paper were

obtained. Mountains are represented by the dotted lines. The thick line

represents the boundary between the Oriental and Palearctic regions according

to Blanford (Phil. Trans. Roy. Soc. B, vol. 194, 1901).

The Palearctic Collembola considered in the present paper were

all obtained from around Badrinath, and the country north of that

locality, in Br itish Garhwal. They were met with at elevations

varying from 10,200 feet to over 12,000 feet, and at a minimum

elevation of 500 feet above the Imits of forests. None of the

species obtained from this area were found in the Oriental region,

either among the lower slopes of the Himalayas or from “ tropical

India.” These Palearctic forms are all referable to well-known

genera.

Among the Oriental Collembola it has been found necessary to

create doame <a new genera, i.e.: Heteromuricus, Dicranocentrordes,

Idiomerus, and Pseudocyphoderus. The remaining species belong

to genera whose range extends into at least one other zoo-

geographical region,

Among the new forms described the most remarkable is Hetero-

MUPICUs cercifer, gen.et sp.n. It is unique among Collembola in

ORIENTAL COLLEMBOLA, 83

possessing a median cercus to the fifth segment of the abdomen.

A new sub-family—the Heteromuricinw—has been created for its

reception. A second new genus, Pseudocyphoderus, with a single

Species, occurs in nests of Termites near Lake Chilka.

In no instance has a new genus been created unless I had at

my disposal at least five specimens, and many of the new species

have only been founded after an examination of a series of twenty

or more examples. At least nine apparently new species, and

two new genera, have been left undescribed owing to there being,

in these particular instances, only one or at the most two speci-

mens of each at my disposal. A few years’ experience with the

Collembola is sufficient to teach that, apart from colour and size,

such structural features as the number of teeth to the claws of the

feet, and the relative lengths of the antennal joints, frequently

cannot be relied upon for specific purposes, unless several spe-

cimens are at hand for comparison.

Unless definitely stated otherwise, the descriptions have been

drawn up from specimens preserved in alcohol. In this connection,

it is important to take into account the fact that a variable amount

of contraction of parts frequently takes place. The relative

lengths of the trunk segments, and the joints of the antenne, are

difficult to represent with complete numerical correctness, owing

to the contraction (or telescoping) that takes place in many in-

stances from this cause. However, in almost every instance the

numerical ratios given in this paper have only been arrived at

after having made a series of measurements on several specimens.

These measurements were obtained by the aid of a Leitz drawing

apparatus and a Zeiss micrometer. In all cases the measurement

of the relative lengths of the trunk segments has been taken

along the mid-dorsal line of the body.

As regards the terminology employed, I have in every instance

used the word body to denote the whole insect, excluding, of course,

the appendages, and the word trunk to mean the thorax and

abdomen. As regards the configuration of the furcula, I have

described that organ in the extended attitude, projecting back-

wards. The terms dorsal and ventral and anterior and posterior

are used with reference to the organ in that position. The

use of these terms in this connection has been made solely for

purposes of description, without any reference to their being of

morphological value.

The measurements of the length of the different species refer

to the body only, and do not include either the antenne or

furcula.

Owing to the fact that it has not been possible in India to

consult certain Papers of the earlier zoological literature, a few

of the references to the original descriptions ' of genera and species

have been quoted on the authority of Tullberg’s “ Sveriges

Podurider”; these are denoted thus 7.

The majority of the specimens are to be found in the collection °

of the Indian Museum, Calcutta, and the reference numbers. to

84 PROF, A, D, IMMS ON

that collection are quoted in each case. Wherever material has

allowed, a series of co-types has also been deposited in the

Natural History Department of the British Museum, South

Kensington. Such species are indicated thus *.

[l. Drscriprion OF THE SPECIES,

Sub-order ARTHROPLEON A Born.

Fam. Popuritp« Lbk.

Sub-fam. Hyroaasrrurina Born.

Genus Xunyuua Tullb.

+Xenylla 'Tullberg, “ Om Skand. Podurider af underfam,

Lipurine,” Akad. Afh, Upsala, 1869, p. Nl.

*XENYLLA OBSCURA, Sp.n. (PI. VI. figs. 5-9.)

Denies furcule cum mucronibus tibia longitudine cquales.

Spine anales parve, arcuate, papillis crassis affine. Pili clavati

in tibiis nulli. Long. 75 mm.

Head.—Vhe eyes as in other species of the genus.

Legs.—The claws of the feet all similar, unarmed. Near the

apex of each tibia is a slender tapering seta (fig. 9); denent hairs

absent.

Furcula.—The dens and mucro together equal to the tibia in

length—a little longer than the manubrium. The muerones not

fused with the dentes, slender, and very slightly curved at their

apices (figs. 5 & 7).

Hamula.—A little shorter than the dens; the ram tri-dentate,

the innermost tooth the largest (fig. 8).

Anal Spines—Small and stout, curved. The papille from

which they arise small and short (fig. 6).

Coloration.—Leaden purple with a number of small irregular

ellowish markings on the dorsal aspect of the head and trunk.

The legs, fureula, and ventral aspect of the body pale, almost

white. The eyes on a black patch on each side of the head.

Length ‘75-1 mm. ; average length *75 mm,

Highteen specimens from Simla, altitude circa 7000 ft., where

they were found floating in large numbers on the surface of pools

of a stream (V’. Annandale, May 11th, 1908).

No. ~ Indian Museum Coll.

This species is a little smaller than most species of the genns.

It resembles Yenylla humicola (O. Faby.) Tullb. in having the

mucro free and not fused up with the dens, though the separation

in Y. humicola is rather more marked when viewed from the

dorsal aspect than in the present species. From 1’, /awmicola it is

further distinguished—(q) by the absence of tenent hairs from

ORIENTAL COLLEMBOLA. RD5

the extremity of the tibia; (b) in the combined length of the dens

and muecro not exceeding that of the tibia; and (c) in the anal

spines being larger and more strongly curved. YX. grisea Axels,

and X. maritima Tullb. differ from XY. obscura in having the

mucro and dens fused together, and in the anal papille being

larger. X. brevicauda Vullb. and X. nitida Tullb. are both

easily separable from the present species on account of their

having a much smaller fureula,

Sub-fam. Acuorurina Born.

Genus AcuornutTes Templ.

Achorutes Templeton, “Thys. Hib.,” Trans. Ent. Soc. Lond.

vol. 1. 1835, p. 96 (ad partem),

Achorutes Schott, “ Zur Syst. und Verbreit. Pal. Collem.,”

Kongl. Svensk. Vet.-Akad. Handl. 1893, vol. xxv. p- 80.

Acnorures Anmatos Nicolet.

Syn. 1842. Podwra armata Nicolet, Rech. pour servir a hist.

des Podur. p. 57, pl. v. fig. 6.

+ 1847. Achorutes armatus Nicolet, Essai s. classif. de l’ordre

des Thys. p. 378.

One hundred and forty specimens, Peradeniya, Ceylon (Z. #,

Green, 1905).

[ have compared these specimens with European examples of

Achorutes armatus and find that they agree in all details of

structure. In a letter dated Noy. 2nd, 1907, Mr. E. E. Green

remarks that “this little species appears occasionally in vast

numbers on the silt left in ditches after rains. It could then be

collected by the bucketful. It has a peculiarly unpleasant

odour of its own, which it has communicated to the spirit in which

it has been preserved.”

It has been previously recorded from the Oriental region by

Oudemans £ from Sumatra. :

Sub-fam. NEANURINA Born,

Genus NEANURA Macgillivray.

+ dnowra Gervais, Une quinz. d’espéces des Ins. Apt., 1842,

p. 45.

t Anura Nicolet, Essai s. classif. de Vordre des Thys., 1847,

p. 386.

Neanura Macgillivray, “A Cat. Thys. N. America,” Can.

Ent. xxiii. 1891, p. 267.

* NEANURA CORALLINA, sp. n. (PI. VII. figs. 23-26.)

Latera corporis parvis tuberibus instructa. Unguiculus inermis.

} Apterygota des Indischen Arch., in Max Weber's ‘ Zool. Ergebnisse einer Reise

in Niederl.-Ostind.’ Hft. i. p. 89.

86 PROF. A. D. IMMS ON

Organa postantennalia desunt. Ocelli in utroque latere capitis 2.

Color coralium. Long. 2-2°3 mm.

Head.—Broader than long, somewhat triangular in form

(fig. 23). The eyes two in number on either side, situated at the

base of a rounded dorsal tubercle which is surmounted by a

central seta (fig. 25). Post-antennal organs wanting.

Antennce.—Approximately equal in length to the head; the

joints related to one another in length as 8:11:7:9. The

articulation between the third and fourth joints imperfectly

developed, and only visible on the ventral aspect. A small

trilobed sense-organ at the apex of the terminal joint (fig. 24).

Trunk.—The segments mutually related in length as 6:8:

9:8:8:8:8:6:4. The three thoracic segments each provided

with a pair of small, dorso-lateral hemispherical protuberances,

surmounted by set. The first three abdominal segments have

each a similar pair of protuberances, together with a larger pro-

tuberance placed externally to the latter (fig. 23). The lateral

margins of the fourth abdominal segment produced into a series

of three such protuberances; the fifth and sixth abdomimal

segments each with a pair of similar bodies, only larger and

dorso-lateral in position. The cuticle investing the body and

appendages ornamented with minute closely-set tubercles (fig. 25).

Legs.—Short and stout, sub-equal in size; the hind pair of

tibie as long as the femora, the tibie of the first and second

pairs of legs shorter than the femora (fig. 26). The claws alike,

large and stout, slightly curved and minutely tuberculated ;

teeth wanting.

Coloration —Bright coral-red when alive (Green), but the

pigment is completely soluble in alcohol, giving the latter a

pinkish-orange colour.

Length 2-2-3 mm.

Fifty-two examples from Peradeniya, Ceylon, 1500 ft., where

it is common under stones, logs of wood, and dead leaves (£. L.

Green).

No. 7 Indian Museum Coll.

The relative lengths of the joints of the antenne and legs

exhibit considerable diversity in alcohol specimens owing to the

variable amount of contraction undergone.

NEANURA PUDIBUNDA, sp.n. (Pl. VI. figs. 10-12.)

Organum postantennale ellipticum, inchoatum. Ocelli in utroque

latere capitis 3. Unguiculus superior uno dente armatus, inferior

in choatus. Color coccineus. Long. 1°5-2:25 mm.

Head.—Somewhat broader than long, triangular. Hyes

spherical, three on each side. The post-antennal organs rudi-

mentary and appear to be represented by a patch of modified

cuticle close to the outer side of each group of eyes (fig. 10).

Antenne.—In length very nearly equal to the head; the

articulation between the third and fourth joints imperfectly

ORIENTAL COLLEMBOLA. 87

developed, and only noticeable on the ventral aspect. The second

joint the longest; the fourth joint with a small trilobed apical

sense-organ (fig. 10).

Lrunk.—The segments related to one another in length as

4:7:7:6:6:6:6:7:6,and provided with lateral tufts of

long sete. The postero-lateral margins of the fifth abdominal

segment produced into a setigerous protuberance on either side ;

the sixth abdominal segment with a pair of such protuberances

on each side (fig. 12).

Legs.—Short and subequal. The superior claw provided

with a single tooth on the inner margin near the base. The

inferior claw probably represented in a vestigial condition by a

small basal process (fig. 11).

Coloration.—Scarlet when alive (Annandale); specimens in

alcohol are white.

Length 1-5—2:25 mm.

Eight specimens taken on bats’-dung in total darkness in the

Khayon Caves, near Moulmein, Lower Burma (WV. Annandale,

March 7th, 1908).

No. — Indian Museum Coll.

Dr. Annandale informs me that the Khayon Caves are of no

great extent, though their inner parts are quite dark. There are

no features about Veanura pudibunda that point to its being a

true cave form, and most probably it is only a recent migrant

thither.

* NEANURA INTERMEDIA, sp. n. (PI. VI. figs. 13, 14; Pl. VII.

fig. 20.)

Tubera corporis desunt. Unguiculus inermis. Organa post-

antennalia nulla. Ocelli in utroque latere capitis 2. Color coralium.

Long. 2 mm.

This species differs from Veanura corallina, sp. n. in the absence

of dorso-lateral protuberances from the body, and in possessing

longer sete to the antenne. From Neanura pudibunda, sp. n. it

differs in the number of the eyes and in the absence of the tooth

from the inner margin of each of the claws of the feet (fig. 14).

It resembles Veanwra corallina very closely in the important

structural features of the claws of the feet, the number of the

eyes, and the absence of post-antennal organs. It resembles

Neanura pudibunda in possessing two pairs of terminal setigerous

protuberances to the abdomen, in the absence of the dorso-lateral

protuberances from the abdomen, in the length and general

disposition of the sete on the antenne (fig. 13), and in the

general form of the body. It is thus intermediate in its characters

between the two preceding species of the genus, though it differs

from both in the above mentioned features.

Colour.—Coral-red ; in alcohol specimens quite white.

Length 2 mm.

Eleven specimens from near Bhowali, Naini Tal District, in

88 PROF. A. D. IMMS ON

the Himalayan foot hills of Kumaon, circa 5000 ft. It is very

local, and occurs under damp loose bark and in decaying stems of

Euphorbia (A. D. Imms, July 1909).

No. = Indian Museum Coll.

A Key to the three Species of the Genus Veanura.

A. Ocelli two on each side of the head. No post-antennal organs.

as Body without dorso-lateral protuberances ..................... I. intermedia.

. Dorso-lateral protuberances present ..................0......... IV. corallina.

1B}, one three on each side of the head. Post- omental organs

DNESCIIG Sct Saraki Nae coarse nyc mit Ohne re ae ee N. pudibunda.

Oudemans* has described a single species of this genus (iV.

fortis) from the Oriental region, where it occurs in Java, Sumatra,

and Saleyer.

Genus PseupAcnorutEs Tullb.

Pseudachorutes Tullberg, “ Fort. 6fver Sv. Podurider,” Ofvers.

Kongl. Vet.-Akad. Forhandl. xxvii. 1871, p. 155.

Pseudachorutes Borner, “‘ Das System der Collembolen,” Mitt.

Naturhist. Mus. Hamburg, xxiii. 1906, p. 164 (= 2 Gnatho-

cephalus Macg.).

PSEUDACHORUTES ANOMALUS, sp. n. (PI. VI. figs. 1-4.)

Corpus tuberculatum. Tumores utriusque organi post-antennalis

17. Onguiculus inermis. Pili clavati in tibiis nulli. Anten-

narum articulus quartus duobus preecedentibus longior. Long.

1-1°25 mm.

Head.—TVhe eyes eight in number on each side. Post-

antennal organs oval in form, each with seventeen tubercles

(fig. 2).

Antenne.—The joints related in length to one another

as 0:6:4:11; the third and fourth joints partially fused

together : a small tri-lobed apical sense-organ, and a second sense-

organ situated ashort distance below the apex of the antenna.

Ibnrasied with a few short slender hairs; the cuticle tuberculated.

Trunk.— Almost entirely glabrous, only a few odd scattered

hairs being present. The cuticle uniformly tuberculated (fig. 2).

Leqs. —Short and stout; the cuticle not tuberculated. A

few scattered sete on the femora and _ basal joints, and a double

circlet of sete near the distal extremity of each tibia. The claws

similar on each pair of legs, large and stout, more than one half

the length of the tibia, unarmed (fig. 3). Morag hairs absent.

Parente: —Short and stout (fig. 4), not quite reaching up to

the apex of the abdomen ; the cuticle uniformly tuberculated. The

manubrium and dentes about equal in length, and each approxi-

mately two and a half times the length of the mucro. The mucro

* Oudemans in Weber’s ‘ Zool. Ergeb. einer Reise in Niederland.-Ostind.’ Hft. i.

p. 91.

ORIENTAL COLLEMBOLA, 89

(fig. 1) large and blade-like, its surface partially sculptured with

small tubercles similar to those found elsewhere; at its apex is a

rounded curved tooth.

Coloration.—In alcohol specimens dull brick-red above and

pale dirty cream-colour beneath ; the antenne somewhat darker

than the rest of the body and with a purplish suffusion. The

legs and furcula whitish.

Length \-1:25 mm.

Two specimens taken on the surface of water at Kurseong,

EK. Himalayas, 5000 feet (V. Annandale, July 4th, 1908).

~ 4395 ;

No. ty Indian Museum Coll.

)

This species differs from Tullberg’s original diagnosis of

the genus in having the cuticle of the whole of the body and

appendages, with the exception of the legs, tuberculated.

Family Enromopryip# D.T.

Sub-fam. [sovomin® Schiff.

Genus Isoroma Boutlet.

Isotoma Bourlet, Mém. sur les Podures, 1839, p. 23 (ad

partem).

IsoroMA siva, sp.n. (Pl. VI. figs. 16-18; Pl. VII. fig. 19.)

Setosa. Segmentum quartum abdominis triplo longius quam

tertium. Antenne capite duplo longiores ; articulus quartus tertio

Jere duplo longior. Ocelli 12: 6 in utroque latere capitis. Organum

postantennale nullum. Dentes mucronum tres, wrus post alterwm

inserti. Long. 1:25-1:5 mm.

Head.—Slightly longer than broad, as long as the thorax

(fig. 19). The eyes six in number on each side (fig. 18); post-

antennal organs wanting.

Antennew.—In average length measuring “5 mm.; the joints

related to one another proportionately in length as 5:8:8: 14.

Trunk.—The segments velated to one another in length as

9:8:5:7:7:20:5:1 (fig. 19). Invested with plumose

hairs.

Legs.—Sub-equal, clothed with plumose hairs; the claws to

each of the pairs similar. The superior claw elongate and greatly

acuminate, with two extremely minute teeth near the apex, and

a third tooth near the base. The inferior claw unarmed. A

single very long tenent hair in relation with each foot (fig. 16).

Furcula.—Approximately equal im length to the antenne ;

clothed with plumose hairs. The dentes one half longer than the

manubrium. The mucrones small, tridentate; provided with a

prominent, upwardly directed terminal tooth, and immediately

in front of it isa second tooth pointing obliquely forwards, and

in close relation with the latter is a backwardly directed spiniform

tooth (fig. 17).

90 PROF. A. D. IMMS ON

Coloration.—When alive dull reddish with a purplish tinge.

In alcohol specimens dull brick-red, with the head paler and

inclining to yellowish. Theantennz and legs dark purplish; the

furcula white. The eyes on a black patch on each side of the

head, the two eye-patches united together by a transverse black

band situated just behind the points of origin of the antenne

(ise etO)).

Length 1:25-1:5 mm.

Five specimens taken under stones along the edge of a

mountain stream at Badrinath, Garhwal Himalaya, 10,300 ft.

(A. D. Zmms, May 27th, 1910).

No. ane Indian Museum Coll.

TsOTOMA NIGROPUNCTATA, sp. n. (PI. VII. figs. 27-29.)

Setosa. Segmentum quartum abdominis fere triplo longius quam

tertium. Ocelli4:2 in utroque latere capitis. Dentes furcule

manubrio longiores ; mucrones tridenticulati. Long. 15-2 mm.

Head.—Vhe eyes two in number on each side, placed one behind

the other. The post-antennal organs very smali, annular (fig. 27).

Situated on the dorsal aspect of the head are large curved setz,

ciliated along one side at their apices.

Antenne.—Slightly longer than half the total length of the

head and trunk; the joints related to one another in length as

3 8 (G8 Gai

Trunk.—Vhe segments related respectively in length as 6:5:

3:4:5:13:3:1. <A prominent “collar” of sete along the

anterior border of the mesothorax, similar to those occurring on

the head. A few scattered setze over the general surface of the

body, and a tuft of plumose hairs at the apex of the abdomen.

Legs.—The claws of the feet similar on each pair of legs

(fig. 28). The superior claw slender, strongly curved and acumi-

nate; armed with one large tooth towards the base, and a

minute tooth immediately in front of the latter. The inferior

claw linear and acuminate, unarmed. No tenené hairs; in the

position occupied by them is a slender tapering seta.

Furcula.—Slender, the dentes related in length to the manu-

brium as 5:4. The mucrones (fig. 29) tridentate, armed with a

slender curved terminal tooth, in front of the latter is a shorter

and stouter vertical tooth, and at the base of the mucro isa slender

backwardly directed spiniform tooth. Arising from the dens, at

a distance from the apex equal to three times the length of the

mucro, are several long compound (plumose) hairs. These extend

backwards, parallel with the long axis of the furcula, reaching

nearly to the apex of the mucro (fig. 29).

Coloration. —Straw-coloured with a slight brownish tinge ;

the legs and spring whitish. When viewed under an 3 in. objective

deposits of fine brown granules are seen beneath the cuticle, and

to these the brownish tinge owes its origin. The deposits are

for the most part arranged segmentally in transverse bands. The

ORIENTAL COLLEMBOLA, 9]

eyes densely pigmented, appearing as two black dots on each side

of the head.

Length 1-5-2 mm.

Three examples, taken under stones at the edge of a spring in

the Kurseong District, E. Himalayas, 4700 feet (V. Annandale,

March 25th, 1910).

8603 ;

No. a Indian Museum Coll.

al » . . . . . . .

This species is readily distinguishable from Jsotoma quadri-

oculata Tullb. by the fact that the dentes are much longer than

the manubrium, and that the mucrones are tridentate.

Sub-fam. ToMocErRIN« Schiff.

Genus Tomocerus Nicolet.

Tomocerus Nicolet, Rech. p. serv. & hist. des Podur., 1841,

p..67.

Tomocervs vuLearis Tullb.

Syn. 1871. Macrotoma vulgaris Tullberg, “ Fort. ofver Sv.

Podurider,” Ofvers. Kong]. Vet.-Akad. Férhandl. xxvii. p. 149.

1893. Tomocerus vulgaris Schott, “Zar Syst. und Verbreit.

Paleare. Coll.,” Kong]. Svenska Vet.-Akad. Handl. xxv. p. 41.

A form closely resembling the type species and differing only

in the following points :—

a. Smaller in size.

6. Ten instead of 12-16 spines to the dentes.

c. The basal tooth of the mucro larger and more pointed

than is represented in Tullberg’s figure of 7’. vulgaris *.

Length 3 mm.

Two specimens, taken under stones near the edge of a mountain

stream at Badrinath, Garhwal Himalaya, 10,300 feet (A.D. Jmms,

May 27th, 1910).

No. ee Indian Museum Coll.

In both examples the antennz possessed only three joints,

which were related to one another in length as 7:12:70. The

antenne themselves measured 2°2 mm. long.

On account of the small size of the specimens and their

possessing only three joints to the antenne, instead of the normal

number of four, I believe that they are immature individuals

of the above species. They are probably to be regarded as a

Himalayan variety of the same, but this point cannot be definitely

determined until adult specimens have been discovered.

Sub-fam. Hereromuricin#, sub-fam. nov.

This sub-family is characterised by the presence of a median

cercus to the fifth abdominal segment.

* Sveriges Podurider, pl. iv. fig. 9.

92 PROF. A. D. IMMS ON

Genus HETEROMURICUS, gen. nov.

Mesonotum non prominens. Segmentum abdominale quartum

quam tertium paullo lougius. Antenne quinque articulos habent.

Ocelli 16: 8 in utroque latere capitis. Organa postantennalia

carent. Seqmentum abdominale quintum medio cerco instructunr.

Cutis squamosa.

The presence of a single median cercus to the fifth abdominal

segment separates this genus from other known genera of

Collembola. In possessing five-jointed antenne, and in the body

being scaled, it shows perhaps closer relations with Heteromurus

Wankel than with any other genus.

* HETEROMURICUS CERCIFER, sp. n. (Pl. VIII. figs. 49-51;

Pl. LX. figs. 52-54.)

Antennarum articulus quartus longissimus, quam tertius duplo

longior. Cercus segmento abdominuli tertio longitudine cequalis.

Unguiculus superior duobus minutis dentibus armatus ; unguiculus

injerior lanceolatus, acuminatus, inermis. Mucrones dentibus

duobus atque seta spiniforme una instructt. Long. 2 mm.

Head.—Inclined at an angle of 45° with the long axis of the

body ; invested with scales and scattered sete. The eyes eight in

number on each side ; post-antennal organs wanting.

Antenne.—Kqual in length to the furcula; five-jointed (fig. 52),

the joints respectively related in length as 1: 10:12:25: 14.

The basal joint small and annular, provided with a whorl of

short spine-like sete; the second and third joints scaled; the

fourth and fifth joints clothed with closely-set whorls of short,

curved hairs.

Trunk.—Densely clothed with scales (figs. 51 and 52); the

scales at the hinder extremity of the body, surrounding the base

of the cercus, larger than those found elsewhere. The segments

mutually related in length as 6:6:5:6:8:12:4:1. Arising -

from the dorsal aspect of the fifth abdominal segment is a

prominent median cercws (figs. 50 and 52) nearly equal in length

to the third abdominal segment. The cereus densely clothed with

scales, and provided ventrally with long slender sete, possibly

sensory. Along the anterior border of the mesothorax is a

“collar” or “frill” of stout sete, and a tuft of similar sete at

the extremity of the abdomen,

Legs.—Sub-equal ; the two basal joints clothed with setz, the

remaining joints sealed down to the claws; interspersed among

the scales are numerous hairs and set (fig. 49). The superior

claws of the feet with two small teeth along the inner margin

(in five specimens one or other of the teeth were absent). ‘lhe

inferior claws large, lanceolate and acuminate, unarmed; those of

the third pair of legs a little longer than the corresponding claws

of the preceding pairs,

Furcula.Slender, reaching forwards to the ventral tube ;

densely clothed ventrally with scales. ‘he dentes related in

ORTENTAL COLLEMBOLA. 93

length to the manubrium as 4:3. The mucrones small, armed

with a curved terminal tooth, a single dorsal tooth and a _ basal

spiniform tooth (fig. 54).

Coloration. —Ground-colour of the body and fureula varies

from whitish to dull ochre-yellow; the legs, antenn, and cereus

bluish-violet. The eyes on a black patch on each side of the

head. The ground-colour of the body varies according to whether

the specimens have been denuded of their scales or not.

Length varying from 1:5—2°5 mm. (excluding cercus)

length 2 mm.

Thirteen specimens, taken under dead leaves at Calcutta

(Jndian Museum Collector, Jan. 14th—20th, 1908, and Feb. 18th,

1910).

2 4AAS °

No. = Indian Museum Coll.

> average

Sub-fam. EnromoBry1n& Schiff.

Genus Isoromurus Born.

Isotomurus Borner, ‘* Neue altw. Collem., nebst Bemerk. z.

Syst. der Isotom. und Entomob.,” Sitz. Gesell. naturf. Freunde zu

Berlin, 1903, p. 129.

*JIsoromurus PALusTRIS Miill. (Pl. VI. fig. 15; Pl. VII.

gs. 21, 22.)

rSyn. 1776. Podura palustris Miiller, Zool. Dan. Prodr..

Havnie, p. 184.

1873. /sotoma palustris Lubbock, Monogr. Coll. and = Thys,

p- 169. r

HHead.—The eyes eight in number on each side (fig. 22).

Antenne.—A little longer than the thorax, the joints related

proportionately in length as 4: 8:9: 9 (in one example they

were related as 3:6:7:8).

Trunk.—Clothed with plumose hairs. The third abdominal

segment a little longer than the fourth.

Furcula.—As long as, or a little longer than the antenne:

reaching to the ventral tube. The dentes approximately twice

the length of the manuhriwm.

Coloration.—Ochre-yellow, either with or without a few small

irregular scattered black markings on the dorsal aspect, which

coalesce in some specimens to form blotches. The antenne and

furcula paler; the antennz in two examples tinged with purple.

Length 2 mm.

Ten specimens, taken on the surface of water at Calcutta t

(Undian Museum Collector, Sept. 1st, 1908, and Sept. 22nd, 1909).

No. = Indian Museum Coll.

The specimens agree in all essential details of structure with

European forms of the species.

t I. palustris has been previously recorded from the Orientai region by Bérner,

from Java.

94 PROF. A. D. IMMS ON

Genus Lepipocyrrus Bourlet.

Lepidocyrtus Bourlet, Mém. s. les Podurelles, 1839, p. 15.

Lepidocyrtus Borner, ‘“ Das System der Coll.,” Mitt. Naturhist.

Mus. Hamburg, 1906, xxii. pp. 164 and 174. [Including

Pseudosinella Schiff. arn Acanthurella Born. |

LEPIDOCYRTUS ROBUSTUS, sp. n.

Segmentum abdominale quartum, thoracem, et segmentum

abdominale primum longitudine equans. Unguiculus superior

dentibus duobus parvis armatus ; unguiculus inferior lanceolatus,

inermis. Long. 3°6 mn.

Antenne.—Three times longer than the head, the joints related

to one another in length as 2:3:3: 4.

Trunk.—The segments related respectively as 16:6:4:5:4:

26:2:1. The fourth abdominal segment six times the length of

the preceding segment.

Legs.—The denna similar on each of the pairs of legs; the

superior claw armed with two small teeth situated respectively

from the base and apex of the claw, at distances equal to one

third the length of the latter. The inferior claw lanceolate,

unarmed. A single tenent hair in relation with each foot.

Furcula.—Vhe dentes very nearly twice the length of the

manubrium; the mucrones tridentate, similar to the typical

form found in the genus.

Coloration.—The ground colour yellowish, the appendages

paler. The third joint of the antennz with an apical suffusion

of violet-black, the fourth joint almost white with a sight basal

suffusion of violet. The eyes ona black patch on each side of

the head; a purplish lateral suffusion on each side of the head

behind the eye-patch. An extensive suffusion of the same colour

on either side of the mesothorax, and a broad conspicuous band

of similar colour along the distal portion of the fourth abdominal

seoment., The femora of the hind pair of legs almost entirely

violet.

Length 3°6 mm. (including the head).

One example, taken under dry leaves and stones on the edge

of a jungle stream, Maddathoray, W. base of W. Ghats, Travancore,

S. India (WV. Annandale, November 18th, 1908).

No. ae Indian Museum Coll.

This species 1s closely allied to LZ. maximus Schott *, from the

Kamerun. It is separable, however, on account oo the great

size of the fourth abdominal segment; in the inferior claws of

the feet being lanceolate, with its lower margin curved instead of

being straight ; and in the stouter tenent hair.

* “Tnsektenfauna von Kamerun: Collembola,’ Bihang till K. Sy. Vet.-Akad.

Handl., Bd. 19, Afd. iv. no. 2, p. 11, pl. mi

Je)

ORIENTAL COLLEMBOLA,

Genus Enromoprya Rondani.

+ Entomobrya Rondani, Dipterol. Ital. Prodr. vol. iv.

Degeeria Nicolet, Rech. p. s. a Vhist. d. Podur., 1842, p. 70.

Entomobrya Borner, ‘ Das Syst. Coll.,” Mitt. Naturhist. Mus.

Hamburg, 1906, xxii. p. 164, [Including Homidia Borner and

Sinella Brook. |

*ENTOMOBRYA kama, spam. (Pl. VIL. fig, 33;° Pl. VIEL,

figs. 34-36.)

Antenne truncum longitudine fere aequantes. Segmentum

abdominale quartum plus dimidia trunci parte occupans. Mucrones

denticulis duobus atque seta spiniforme una instructi. Flava;

anteriore parte capitis, marginibus segmenti secundi thoracis,

segmentis secundis tertiisque abdominis, et fascia transversa posteriore

quarti, nigris. Long. 175-2 mm.

Head.—Slightly longer than broad ; clothed with short, scattered

plumose hairs and a dorsal tuft of very long, stout setee, ciliated

along one side at their apices. The eyes eight in number on

each side.

Antennee.— Usually very nearly equal in length to the body

excluding the head, but in some specimens they exceed the length

of the body. Four jointed, densely clothed with hairs, dispersed

among which are slender sete. The joints related to one another

in length as 5:6:6:11; the ring-like basal joint, typically

present in the genus Hntomobrya, is absent.

Trunk.—Fusiform, clothed with short, curved plumose hairs,

scattered among which are slender sete. The thorax and first

two abdominal ‘segments provided dorsally with very long, stout

sete, ciliated along one side at their apices (fig. 36); a tuft of

similar setz at the apex of the abdomen. The trunk segments

related proportionately in length as 11:5:3:6:6:45:5:1

(fig. 34).

Legs.—Clothed with hairs among which are slender sete.

The superior claws moderately slender and bidentate along the

inner margin (fig. 33); the teeth situated respectively from the

apex of the claw at a distance equal to one quarter and one half

the total length of the latter. The inferior claws a little more

than half the “length of the superior claws, acuminate and sharply

pointed, the margins without any teeth. A single tenent hair

in relation to each foot.

Furcula.—As long as the trunk excluding the mesothorax ;

densely clothed with hairs, dispersed among which are slender

sete. The dentes a little longer than the manuhriwm, very slender,

and each is provided with a double row of small peg-like spines

along the proximal half of its inner aspect. The smwerones tri-

dentate, with a curved terminal tooth, a stout erect conical

middle tooth, and an oblique acicular posterior tooth (fig. 35).

Coloration.—Light ochre-yellow marked with patches of violet-

black. The eyes on a large irregular black area on each side; a

96 PROF, A. D. IMMS ON

small violet-black patch between the bases of the antennz and

frequently prolonged into a narrow streak on each side to unite

with the eye-patch. A triangular area of the same colour on the

middle of the hind border of the mesothorax, and a slight suffusion

on each side near the outer margin of that segment. The meta-

thorax and the first abdominal segment entirely yellow; the

second abdominal segment violet-black, with the exception of a

narrow irregular yellow area along its anterior margin ; the third

abdominal segment entirely deep violet-black; the posterior

half of the fourth abdominal segment densely suffused with

violet-black; the fifth and sixth abdominal segments yellow.

The first and second joints of the antenne yellow, the second

joint in most specimens with a slight violet suffusion at its apex ;

the third and fourth joints purplish.

Length 175-2 mm.

Fifty-one specimens taken under dead leaves in Calcutta, where

it appears to be very plentiful (/ndian Museum Collector, January

16th, 17th, 18th, and 20th, 1908, and February 18th, 1910).

Nos. a and — Indian Museum Coll.

In all the individuals examined the colour pattern was found

to be very constant, practically no variation being observed. In

specimens that had been kept in alcohol for two years, the

ground colour is much paler and cream-coloured. The long, stout

sete (fig. 36) fall off very readily in alcohol specimens, and very

many specimens have lost them altogether. In many cases the

antenne are very much shrivelled in saleohol, and in a large

proportion of the specimeus the tenent hair is either broken or

lost from one or more of the legs.

ENTOMOBRYA KALI LUTEA, Var. NOV.

This differs from the typical form in having the fourth abdominal

segment entirely yellow.

One specimen taken among low herbs and grass at Simla circa

7900 ft. (V. Annandale, May 12th, 1908),

No. 88t4

6 Indian Museum Coll.

EnroMOBRYA CRASSA, Sp.n. (PI. VII. figs. 30, 31.)

Segmentum tertium abdominis quartum longitudine Sere equans.

Mucrones denticulis duobus atque seta spiniforme una instructe.

Color fava-viridis. Long. 1:5 mm.

Head.—The eyes eight in number on each side (fig. 31); the

post-antennal organs wanting.

Antenne.—The joints mutually related in length in the pro-

portion of 4: 6:6: 9.

Trunk.—Provided with pilose hairs of various lengths. The

segments related to one another in length as 8:7:4:5:8: 10:

33 2.

Legs.—Clothed with pilose hairs among which are a few stouter

ORIENTAL COLLEMBOLA, 97

sete. The superior claw of each foot slender and acuminate, with

a small tooth situated at the middle of the inner margin. In

some examples there is a second, and much smaller tooth, placed

half way between the former tooth and the apex of the claw.

The inferior claw of each foot slender and tapering, a little more

than half the length of the superior claw, unarmed, A single

very slender tenent hair to each toot.

Furcula.— 7 —9 mm. long; the manubrium one half the length

of the dens. The muero =}; mm. long (fig. 30), tridentate, with a

slender, curved terminal tooth, a vertical and somewhat stouter

tooth anterior to the latter, and a minute backwardly directed

spiniform tooth.

Coloration.—When alive, dull dark green to the naked eye,

In alcohol specimens, pale greenish yellow suffused with dark

indigo-blue. The eyes on a black patch on each side of the head,

the two patches joined together by a transverse band, which

passes across the head just behind the bases of the antenne,

On the middle of the dorsal side of the head is a prominent

black sagittate marking thus |, with its apex directed backwards,

The antenne and legs darker than the body, somewhat purplish

in colour. The furcula yellowish white,

Length varying from 1:5-1-8 mm.

Six examples, taken in ants’ nests under stones about half a mile

below the base of the Satopanth Glacier, Garhwal Himalaya,

12,500 ft. (A. D. Imms, May 25th, 1910).

8609

No. Sia Indian Museum Coll,

This species differs from typical members of the genus #nto=

mobrya, and resembles the genus Orchesella, in the very short

fourth abdominal segment, It agrees with the genus Hntomobrya

in the characters of the antenne and furcula, in the eyes, and

in the absence of post-antennal organs, Hntomabrya anomala

Carpenter * similarly possesses a relatively short fourth abdominal

segment. The latter species, however, may ultimately be

separated into a new genus on account of its possessing six- jointed

antenne.

Genus Serra Lubbock,

Seira Lubbock, * Notes on the Thysanura,” pt. iv., Trans. Linn.

Soc. 1870, vol. xxvii, p. 279, pl. 45 (= Ptenuwra, Templ., Born.).

Sira Tullberg, “ Sveriges Podurider,” Kongl, Svensk, Vetensk,-

Akad. Handl, 1872, vol, x, p. 41, pl. vi.

SEIRA FRIGIDA, sp.n. (PI. VIII. figs. 41, 42.)

Unguiculus superior tridenticulatus ; denticuli perparvi, ita colla-

cati ut unus post alierwm insertus sit. Unguiculus inferior

lanceolatus, inermis. Mucrones breves, bidentati. Articulo quarto

* “On two new Irish species of Collembola,” Sci. Proc. Roy. Dublin Soe, yol, xi,

(n. s.) 1906, p. 40, pl. 11.

Proc. Zoou. Soc.—1912, No, VII. 7

98 PROF, A. D. IMMS ON

antennarum primum et secundum longitudine cequante. Tibice

pilis clavatis singulis instructe. Long. 2°5 mm.

Head.— As long as the combined length of the meso- and meta-

thorax. The eyes as usual in the genus.

Antenne.—Kqual in length to the furcula (in some examples

slightly shorter than that organ). ‘The joints related to one

another in length as 5: 8:9: 13.

Trunk.—Invested with scales, among which are prominent

curved sete, ciliated at their apices along one side, and disposed

in the following manner : a prominent * frill” or ‘ collar” along

the anterior border of the mesothorax, and a second group of such

sete near the posterior margin of that segment. A few scattered

setee on the metathorax, and a tuft of similar but shorter sete

at the extremity of the abdomen. The segments related to one

another in length as 8:6:5:5:5: 23:5: 3.

Legs.—The claws of the feet similar on all pairs of legs

(fig. 41). The superior claws moderately slender, sightly curved

at their extremity, and armed with three minute teeth along the

inner margin. The inferior claws lanceolate, entire, slightly

curved at their apices. A single tenent hair in relation to each

foot.

Furcula.—In length measuring *8 mm.; the manubriwm related

to the denies in length as 13: 19—or approximately as 2: 3.

The dentes without ventral scales; slightly curved upwards at

their apices (fig. 42). The mucrones only imperfectly separated

off from the dentes, armed with a prominently curved terminal

tooth and a basal spiniform tooth. The latter directed back-

wards in an oblique fashion, almost reaching to the apex of the

mucro (fig. 42).

Coloration.—Dirty yellowish white, marked with irregular

patches of blue-black disposed in the following manner :—A

prominent patch on either side of the head enveloping each

eye-group, and united by a transverse band, which crosses

the head immediately behind the bases of the antenne. <A

slender Y-shaped marking on the middle of the dorsal aspect

of the head. A pair of irregular lateral patches on the meta-

thorax and on the first three abdominal segments; the third

abdominal segment with a median unpaired patch near its

posterior border. The fourth abdominal segment marked with

several irregular lateral and median areas partially confluent with

one another, and varying in different specimens; a short trans-

verse band near the posterior end of the segment. The fifth

abdominal segment with a pair of prominent lateral patches near

its posterior margin. ‘The sixth abdominal segment with a pair

of small lateral spots.

The legs, antenne, and furcula yellowish white, similar to the

ground colour of the body. The antennz and legs conspicuously

marked with blotches of blue-black; in one very dark example

these markings on the antenne were confluent, the latter

appearing entirely blue-black.

ORIENTAL COLLEMBODLA. 99

Length varies from 2°1—2°8 mm.

Four specimens, taken in ants’ nests under stones on a moun-

tain side a short distance below tie base of the Satopanth Glacier,

Garhwal Himalaya, circa 12,300 ft. (A.D. Zmms, May 27th, 1910).

8608

No.

In one example the lateral blue-black markings on the meta-

thorax and the first abdominal segment, together with the median

posterior patch on the third abdominal segment, were entirely

absent.

Indian Museum Coll.

SEIRA BRAHMA, sp.n. (PI. VIII. figs, 43, 44,)

Unguiculus superior bidenticulatus ; unguiculus inferior lan-

ceolatus, mermis. Articulus quartus antennarum longissimus,

secundus et tertius mter se longitudine sub-cequales. Mucrones

breves tridentati. Tibi pilis clavatis singulis instructe. Long,

1:5 mm.

Head. —The eyes eight in number on each side, the anterior

four in each group the largest. No post-antennal argans

present.

Antenne.— A little longer than half the length of the body,

the joints related in length to one another as 3:8: 8or9: 14,

Legs.—The claws of the feet similar on each of the pairs of

legs (fig. 43). The distal extremity of each tibia provided with

a single extremely slender tenent hair. The superior claws armed

with two small teeth, one of which is situated from the base at

a distance equal to one third the total length of the claw. The

other tooth is situated at a similar distance from the apex of

the claw. The inferior claws lanceolate and unarmed.

Furcula.—Reaching to the ventral tube; slender, The dentes

related in length to the manubrium as 6:5; tapering to their

extremities. The mucrones tridentate, armed with a curved

upwardly directed terminal tooth, a median tooth slightly inclined

in a forward direction, and a backwardly directed basal spiniform

tooth (fig. 44),

Coloration.—Ground colour pale yellowish dusted over with

indigo-blue, the insect appearing slate-grey under a hand-lens,

The antennz indigo-blue, the legs and furenla whitish. The

intersegmental areas of the body yellowish, The eyes gn a black

patch on each side of the head.

Length 1:5 mm.

Five examples, taken crawling up the surface of whitewashed

walls ina bungalow at Allahabad (A. D. Jmms, September 20th,

1907).

No 8601

naa Indian Museum Coll,

Genus PsrEupbostira Schibtt,

. Pseudosira Schott, “ Insektenfauna von Kamerun: Collembola,”

Bihang till K. Svensk. Vet.-Akad. Hand], 1893, Bd. 19, Afd. iv,

p. 10, taf. 11. figs. 1-11.

100 PROF. A. D. IMMS ON

Pseudosira Birner, “ Das Syst. Coll.,” Mitt. Naturhist. Mus.

Hamburg, 1906, xxii. p. 164 (including Aesira Stscherbakow

| = Lepidocyrtinus Borner ]}).

* PsEUDOSIRA INDRA, sp.n. (Pl. VII. fig. 32; Pl. VIII. figs.

37-40.)

Unguiculus superior tridenticulatus; denticuli perparvi, ita

collocati ut unus post alterwm insertus sit. Unguiculus inferior

lanceolatus, inermis. Mucrones breves, falciformes. Tibice pilrs

clavatis singulis instruct. Articulo quarto antennarum longissimo,

primum et secundum longitudine equante. Long. 15 mm.

Head.—A little longer than the mesothorax; clothed with

scales, among which on the dorsal aspect are long stout sete,

ciliated along one side at the apex. The eyes eight in number on

each side; no post-antennal organs (fig. 37).

Antenne.—As long as, or, in some examples, a little longer than

the furcula. The joints related respectively in length as

4:7:70r8:11. The first two joints clothed with scales, the

third and fourth joints clothed with small hairs.

Trunk.— Clothed with scales of somewhat variable shape, but

for the most part oval or linear-oval. The scales are finely and

faintly striated, with a relatively long, and very slender pedicel

(fig. 40). Sete (fig. 39), similar to those found on the head,

form a kind of “frill” or “collar” along the anterior edge of

the mesothorax ; a few are also scattered over the general surface

of the body, and there is a terminal tuft at the extremity of

the abdomen. The trunk segments mutually related in length as

Me ROueA weg Or Ola).

Legs.—Al\ the feet similar ; the distal extremity of each tibia

provided with a single tenent hair. The superior claws slender

and acuminate, armed with three small teeth along their inner

margin. The inferior claws linear and acuminate, acicular,

unarmed (fig. 38).

Hamula.—Vhe corpus with a median stout anterior seta placed

in front of the rami. Hach ramus quadridentate.

Furcula.—Slender, reaching to the ventral tube. The manw-

briunmy somewhat shorter than the dentes; scaled. The dentes

clothed ventrally with scales. The macrones hook-shaped

(fig. 32).

Ooloration.—Greyish white when denuded of the scales ; when

the greater number of the scales are present the ground colour

appears markedly brown. Theantenne tinged with violet, a slight

violet suffusion on the mesothorax, and some small lateral patches

of the same colour on either side of the abdomen. The furcula

white. The eyes on a black patch on each side of the head.

Length vavies in different examples from 1°25—2 mm.

Five specimens on the surface of the pool in the “com-

pound” of the Indian Museum, Caleutta (4. D. Imes, December

31st, 1909). The specimens were apparently immature.

ORIENTAL COLLEMBOLA. 101

No. 8602

Thirty specimens taken under dead leaves and at the bases of

the leaves of a palm tree, in the “compound” of the Indian

Museum, Calcutta (Jndian Museum Collector, July 28th, 1909,

and eet 18th and 21st, 1910).

No. hae Indian Museum Coll.

The antennze of this species are very variable; in three speci-

mens the second and third antennal joints were equal in length ;

one example possessed only three joints to the antenne; and

another specimen had three joints to the right antenna and

four to the left. These anomalies are most likely to be explained

as being due to the results of regeneration after an Injury.

The specimens were in each case, so far as could be ascertained,

quite mature.

Pseudosira indra does not agree fully with the diagnosis of the

genus given by Schott. The chief points of difference are:

(a) the presence of teeth to the superior claws of the feet ;

(6) the relative length of the joints of the antenne ; and (c) the

great length of the fourth abdominal segment. As Se ‘hott erected

the genus on a single specimen only, some of his generic

characters will, I believe, prove to be of nothing more than

specific value,

Borner * separates Psewdosira from Setra principally by the fact

that the dentes are scaled ventrally in the former and not so in

the latter genus. This character along with the hook-like mucro

renders the genus Pseudosira easy of recognition.

Indian Museum Coll.

Genus SINELLA Brook.

Sinella Brook, “On a new genus of Collembola allied to

Degeeria,” Journ. Linn. Soc., Zool. xvi. 1882, p. 541.

SINELLA MONTANA, sp.n. (PI. VIII. fig. 48; Pl. IX. figs. 56, 57.)

Ocelli nulli. Unguiculus superior denticulis quattuor armatus.

Mucrones furcule dente uno atque seta spiniforme wno instructi.

Omnino alba. Long. 2 mm.

Head.—Vhe eyes and post-antennal organs absent.

Antennw.—Measuring *9 mm. long; the joints related respec-

tively in length as 8:13:13: 26; the terminal joint tapering

somewhat distally. Clothed with pilose hairs; sets are present

among the hairs on the first three joints, and on the proximal

portion of the fourth joint.

Trunk.—Clothed with short pilose (compound) hairs, among

which, on the head and mesothorax, are numerous large erect

sete similar to those of Sinella curviseta Brook (fig. 57a). The

extremity of the abdomen provided with longer compound hairs

(fig. 57 te The segments | related to one another in length as

perce Ohs Bix Dh bee SAEAE : 5S

* Das System der Collembolen, pp. 164 and 174.

102 PROF, A. D, IMMS ON

Legs.—Superior claws of the feet elongate and acuminate, with

two large slender proximal teeth; in front of the latter are two

minute teeth, the distal one extremely small and situated just

behind the apex of the claw. The inferior claws flattened, bifid

at their apex ; resembling those of Sinella hofti Schiff. (fig. 48).

Tenent hairs absent, their place being occupied by a slender

tapering seta. The legs are clothed with plumose compound hairs

similar to those found on the trunk, and among them, on the

inferior surface of the tibiz, are spine-like sete (fig. 48).

Furcula.—Equal in length to the antenne ; the manubriwm

related in length to the dentes as 2:3; clothed on its dorsal

aspect with long plumose hairs. The mucrones resembling those

of S. héfti; provided with a single stout and prominently curved

terminal tooth, and a basal backwardly directed spiniform tooth

(fig. 56).

Coloration.—Entirely white.

Length in adult examples 2 mm.; in young specimens

1-15 mm.

Two adult specimens and four young specimens; taken in an

ants’ nest under stones on a mountain-side near Badrinath, Garhwal

Himalaya, circa 10,300 ft. (A. D. mms, May 27th, 1910).

No. Indian Museum Coll.

This species is closely allied to Sinella héfti Schaff., but differs

in the claws of the feet. It is readily separable from S. myrmeco-

phila Reuter, which similarly occurs in ants’ nests, by the

characters of the mucro and feet.

The young examples differ from adult specimens chiefly in

having the first antennal joint proportionately shorter.

Genus DicRANOCENTROIDES, gen. nov.

Mesonotum non prominens. Segmentun. abdominale quartum

longius dimidia parte trunci. Antenne dimidia corporis parte

longiores, quadriarticulate. Ocelli 16 : 8 in utroque iatere capitis.

‘urcula latitedine apici fere equalis. Dentes spinosi ; mucrones

lati denticulis armati. Cutis squamosa.

This genus agrees with Dicranocentrus Schott in the dentes

being armed with simple spines, and in the length of the antenne.

With Campylothorax Schott it agrees in the great length of the

fourth abdominal segment, in the form of the furcula, which

scarcely tapers in width up to the apex, and in the form of the

mucrones. It is separable from the latter genus on account of

the thorax not being flexed upon itself, and the relative shortness

of the antenne.

* DICRANOCENTROIDES FASCICULATUS, Sp.n. (PI. VIII. figs. 45-47 ;

PX tes sO Ona DD am bln ONG sie. Sm)

Antenne articule ultimo omniwm longissimo ; ceteris iter se

ORIENTAL COLLEMBOLA. 103

longitudine cequalibus. Unguiculus superior duobus dentibus

armatus ; wnguiculus inferior lanceolatus. Mucrones lati denti-

culis quinque. Long. 2°5-3°5 mm.

Head—Kqual in length to the mesothorax. The eyes eight

in number on each side ; post-antennal oryans absent.

Antenne.—Slightly longer than half the total length of the

body (including the head). The first three joints subequal in

length, the terminal joint a little longer than the preceding

ones. The first two joints clothed with long and conspicuous,

erect, almost black sete; the joints in consequence appearing

greatly swollen to the naked eye, and like “ bottle brushes ”

when viewed under the low power of the microscope (fig. 68).

Trunk.—Clothed with both hairs and scales with many

transitional structures between the two (fig. 47). The segments

related to one another in length as 12:5:4:5:5:44:4: 2;

the fourth abdominal segment longer than half the total length

of the trunk.

Legs.—Long, the third pair longest of all and extending to

the apex of the abdomen. A single long tenent hair in relation

with each foot. The superior claw of the first and second pairs

of legs long, and gradually tapering to a point (fig. 45), armed

with two teeth along its inner margin; one tooth situated at a

distance from the base of the claw equal to one third of the

total length of the claw, the second tooth placed at the same

distance from the apex of the claw. The ixfertor claw lanceolate

and acuminate, unarmed, but in occasional specimens its ‘inner

margin shows minute rudimentary serrations. The superior claw

of the third pair of legs slightly broader than that of the pre-

ceding pair; the inferior claw with a minute tooth at its

base on the inner margin.

Ventral Tube.—Long and cylindrical, the vesicles bilobed

(fig. 68).

Furcula.—Reaching to the ventral tube ; clothed ventrally with

scales. The dentes scarcely narrowing to their apices, a little

longer than the manwbriwm, armed with a longitudinal row of

short stout lanceolate spines along the middle of the inner

lateral margin of each (fig. 46). Towards the apex of the dens

the spines become replaced by stout setee. The mucrones with

two large terminal teeth, two smailer dorsal, sub-apical teeth,

and a lateral tooth (fig. 55).

Coloration.—Seen with the naked eye when alive, it appears

black with a conspicuous yellow band across the abdomen. It

varies from very deep purple-brown to black, with an extremely

variable arrangement of the colour-pattern, In the majority

of individuals, the base of the metathorax, and the first and

second abdominal segments are pale yellow suffused with purplish

brown. The first two joints of the antenne are similar to the

ground colour of the body, the third and fourth joints vary

from yellow to dark violet. At the base of each joint, in five

out of the six specimens, there is a narrow transverse band of pale

104 PROF, A. D. IMMS ON

yellow. The legs and furcula vary from pale dirty yellow,

with purplish or violet suffusions, to deep purple.

Length 2°5-3°5 mm.

Six specimens, taken under damp dead leaves, chiefly of Quercus,

in forest at Bhowali, Himalayan foot-hills of Kumaon, cirea

5700 ft. (A. D. Imms. ‘October 23rd, 1909).

No. ae Indian Museum Coll.

Genus CREMASTOCEPHALUS Schott.

Oremastocephalus Schott, ‘‘ North American Apterygogenea,”

Proc. Cal. Acad. Sci. 2nd ser. vol. vi. 1896, p. 175.

Oremastocephalus Schiiffer, ‘ Collembola des Bismarck-Archi-

pels,” Arch. f. Naturgesch., 1898, p.-406.

CREMASTOCEPHALUS INDICUS, sp.n. (PI. 1X. figs. 58, 59.)

Ocelli 16:8 in wtroque latere capitis. Unguiculus superior

duobus parvis dentibus armatus; unguiculus inferior imermis.

Pretarsi pilis clavatis singulis imstructi. Mucrones furcule

trilobati. Long. 1°5 mm.

Head.—The eyes eight in number on each side ; the post-antennal

organs absent.

Antenne.—Long and slender, equal to the combined length

of the trunk and furcula. The joints related proportionately

in length as 5:7:4:7. The two basal joints armed with

long slender sete.

Th ‘unk.—Densely covered with fine hairs and slender sete.

The segments mutually related in length in the proportion of

Tike MakineD tay OOM E 3t2)

Legs.—The superior claws moderately stout, armed with two

minute teeth (fig. 58): in 15 per cent. of the specimens one or other

of these teeth was absent. The iferior claws broad, obliquely

truncated distally, unarmed. <A single, very stout, tenent hair to

each foot arising from the pretarsus.

Furcula.—Reaching to the ventral tube. The manubriwm

related in length to the dentes in the proportion of 8:11. The

mucrones (fig. 59) quadrangular, with the distal border trilobed ;

in some specimens the lobes appeared to be worn down and

absent. A single sinall scale-like appendage at the apex of each

dens on the dorsal side.

Coloration.—Ground colour varying from cream colour to

yellow. The eyes on a conspicuous black patch on either side of

ae head. The lateral margins of the thorax and first abdominal

segment edged with indigo-blue ; a few scattered patches of the

same colour over the rest of the abdomen. The antennz

suffused distally with violet ; the legs and furcula white.

The coloration, however, is extremely variable, and a detailed

description of the various forms that occur would occupy con-

siderable space. In several instances almost all traces of the

indigo-blue markings were absent; this reduction of the colour-

ORIENTAL COLLEMBOLA, 105

pattern is more evident in the Allahabad specimens. On the

other hand, in several of the Bengal specimens the markings are

much enlarged and intensified.

Length varying from 1 mm. to 1°75 mm.; average length 1-5 mm.

‘Twenty-five specimens, taken at night crawling up the surface

of a whitewashed outer wall of a bungalow, illuminated by electric

light, Allahabad (A. D. Jmms, September 22nd, 1907).

mous

Twenty-six specimens, taken under dead leaves, Calcutta ; for

the most part poorly preserved (Zidian Museum Collector, Jan. Ist,

16th, and 18th, 1908).

Indian Museum Coll.

CREMASTOCEPHALUS MONTANUS, sp.n. (PI. IX. fig. 60.)

Ocelli 16:8 in utroque latere capitis. Unguiculus superior

duobus parvis dentibus armatus; unguiculus inferior mermis.

Pretarsi pilis clavatis singulis instructi. Mucrones furcule tribus

dentibus armati. Long. 2-2°5 mm.

Head.—The eyes eight in number on each side ; post-antennal

organs absent.

Antenne.—The first two joints related proportionately in

length as 5:7; the remaining joints missing in the specimens

examined.

Trunk.—The segments related in length in the proportion of

5:3:3:4:1:15:2:1. Clothed with fine hairs and slender

sete.

Legs.—The superior claws moderately stout, armed with two

small teeth ; one tooth situated from the base of the claw at a

distance equal to one third the length of the claw; the other

placed at a similar distance from the apex. The inferior claws

broad, resembling those of C. indicus (fig. 58), only slightly more

acuminate; unarmed. A single ¢enent hair to each foot very

stout, and arising from the pretarsus.

Furcula.—The mucrones inclined at an angle of 30° with the

dentes, tridentate (fig. 60). At the apex of each dens is a scale-

like appendage, equal in length to the mucro.

Coloration.—Straw-coloured inclining to yellow. The lateral

margins of the thorax and the first segment of the abdomen

edged with dark violet. A few dorso-lateral markings of the

same colour over the rest of the abdomen, and a proximal and

distal suffusion to each of the tibiae. The two basal antennal

joints inclining to pale yellow ; the furcula white.

Length 2-2'5 mm.

Three examples, taken among damp soil under stones and

leaves at Kurseong, E. Himalayas, 5000 ft. (2. H. Gravely,

March 25th, 1910).

+ O08 - . ;

No. “ Indian Museum Coll.

Cremastocephalus montanus is closely related to the preceding

species (C. indicus), but can be readily separated by the form of

106 PROF. A. D. IMMS ON

the muero, In C. montanus the mucro is relatively short, promi-

nently tridentate, and the seale-like appendage is equal in length

to that organ. Tn CO. indicus the mucro is longer, is not toothed

but merely lobed, and the scale-like appendage is considerably

shorter.

Genus PARONELLA Schott (sens. Cat.).

Paronella Schott, Insektenfauna von Kamerun: Collembola,”

Bihang till K. Sv. Vet.-Akad. Handl., Bd. 19, Afd. iv. p. 14,

taf. iv.

Paronella Schiiffer, ‘ Die Collembola des Bismarek- Archipels,”

Arch. £. Naturgesch. 1898, p. 408. (Including Trichorypha

Schott, loc. cit. p. 16, taf. v.)

Schiffer described Paronella dahlii from the Bismarck Arch.,

which is intermediate in its characters between Paronella

and Trichorypha. I have, therefore, followed him by including

Schott’s two genera in the single genus Paronella. Schott states

that the ocelli are four in number on each side in Paronella, but,

nevertheless, figures eight ina group! ‘This latter number obtains

in Paronella dahlii.

* PARONELLA BORNERI, sp. n. (PI. X. figs. 70-74; Pl. XI.

figs. 75, 76.)

Segmentum abdominale quartwm dimidiam trunci partem

occupans. Antenne corpore longiores. Ocelli 16:8 in utroque

latere capitis. Unguiculus superior denticulis tribus (vel dwobus)

instructus ; unguiculus inferior inernis. Mucrones lati. Long.

37d mm.

Head.—Longer than broad, approximately equal in length to

the thorax; inclined at an angle of 45° with the longitudinal

axis of the body. A group of strongly chitinised sete between

the eyes and directed forwards towards the bases of the antenne.

The eyes eight in number on each side (fig. 74); post-antennal

organs absent.

Antenne.— Very long, the length apparently varying according

to age, and sometimes exceeding that of the body and furcula

taken together. The first two joints sub-equal in length,

the first joint provided with a number of very long slender setz

on its inner and ventral aspects. The third joint a little more

than one half the length of the second. The fourth joint long

and slender; variable, but usually equal to the combined length

of the first two joints; slightly but irregularly annulated, and

densely clothed with setose pile. (Vide fig. 75.)

Trunk.—Hlongate fusiform in shape, straight (fig. 75). The

segments related proportionately in nee as (9302.45 12) 3 2eae

or in other examples as 8:4:3: 2 fF 8 ve :2:1; the fourth

abdominal segment occupying from 4 to 24 of the total length of

the trunk. An abundant covering of eles hairs, and sete

(fig. 73).

ORIENTAL COLLEMBOLA. 107

The scales lanceolate, the hairs finely plumose (compound).

Groups of strongly chitinised curved sete are present along the

anterior border of the mesothorax, forming a “ frill” or “collar,”

and at the extremity of the abdomen.

Legs.—Long and slender; the tibize divided by means of

movable joint into a longer proximal and a shorter distal

segment. The femora of the first pair provided with several

extremely elongate slender (sensory ?) sete along their inner

aspect (fig. 76). The superior claws straight, as long as the width

of the distal joint of the tibia at the base (fig. 71); armed with

two teeth along the inner margin—one tooth situated at a

distance from the base of the claw equal to approximately one

third the total length of the latter, the second tooth situated at a

similar distance from the apex of the claw. In many examples

there is a minute tooth placed between the distal. tooth and the

apex of the claw. The inferior claws straight and acuminate.

In relation with each foot is a single stout tenent hair, broadly

expanded at its apex. Pseudonychia long.

fentral Tube-—Moderately long, cylindrical. The vesicles

were retracted in all the specimens examined,

Hamula.—Situated on the anterior third of the fourth abdo-

minal segment. The corpus somewhat mammilated, armed with

a stout, median backwardly directed spine. The rami short and

hoe each provided with four small teeth (fig. 72).

Fureulu.— Long and slender, as long as the trunk-region.

The dentes parallel-sided or only very slightly tapering towards

their apices, clothed with numerous long hairs. The dentes

related in length to the manubriwm as OT | 22. The mucrones

stout and broad, wedge-shaped in sectional area; armed with two

large apical teeth, a lateral inside tooth, and a row of three dorsal

teeth (fig. 70). The distal extremity of the mucro armed with a

very stout rod-like seta on its inner side towards the ventral

aspect. At the base of each mucro on the dorsal aspect of

the dens is a scale-like organ* (fig. 70).

Coloration.—The ground colour varying from dirty cream

colour to yellow, with indigo or violet-black markings disposed in

the following manner :—A lateral area on either side of the head

embracing the eye-group ; a few small patches at the bases of the

antenne ; and irregular lateral markings on each of the thoracic

and abdominal segments which, however, are scarcely visible

dorsally. On the dorsal aspect of the fourth abdominal segment

are a few bilaterally symmetrical markings, and a lateral patch

on either side of the fifth segment. The femora marked with

a distal band of violet; a small proximal band and a more

extensive distal band of the same colour on the first joint of the

tibia. A pale violet suffusion on the second (or distal) joint of

the latter (fig. 76).

The general colour pattern, however, is very variable, and for

* Termed by Schafler “ Schuppenformiger Anhang.”’

108 PROF. A. D. IMMS ON

this reason it has only been possible to describe it in a general

fashion. The markings on the legs are an exception, being

remarkably constant.

Length varying from 2—4°5 mm. ; average length 3°5 mm.

Twenty-seven specimens from Nara Ghat, in the Terai, Nepal

Undian Musewm Collector, February 25th and 26th, 1908); and

two immature specimens from Butal, also in the Terai, Nepal,

taken by the same collector (February 12th, 1908).

Nos. = and - Indian Museum Coll.

This species shares the characters of the genera Paronella and

Campylothorax. It resembles the latter genus, and differs from

typical members of Paronella in the great size of the fourth

abdominal segment. It is readily separated from Campylothorax

by the fact that the metathorax is straight and not curved upon

itself.

PARONELLA TRAVANOORICA, sp. n. (PI. IX. figs. 62-66; Pl. X.

fig. 67.)

Segmentum abdominale quartum 2 partem trunci occupans.

Antenne corpore breviores. Ocelli 16 : 8 in utroque latere capitis.

Unguiculus superior denticulo wno armatus ; wnguiculus inferior

acuminatus, mermis. Mucrones lati, rectangulares. Long. 3°5-4:5

mn.

Head.—Clothed with scales. The eyes eight in number on each

side (fig. 64) ; post-antennal organs absent.

Antenne.—A little shorter than the body. The joints related

in length as 7:8: 6:15 (fig. 67). The basal joint clothed with

sete and acuminate scales; the distal three-fourths of the

terminal joint shghtly and irregularly annulated.

Trunk.—Clothed with scales and scattered sete. The scales

(fig. 66) linear or linear-oval in shape. A group of strongly

chitinised sets forming a kind of “collar” or “frill” along the

anterior border of the mesothorax, and a tuft of similar sete at

the extremity of the abdomen. The segments mutually related in

length as 10:5:3:5:5:22:4:1; the fourth abdominal

segment occupying two-fifths the total length of the body (fig. 67).

Legs.—Sub-equal. A single tenent hair at the distal extremity

of each tibia. The superior claws of the feet (figs. 62 and 63)

nearly straight, broad at the base; a single minute tooth on the

inner margin near the base of the claw. In two specimens, on

the first pair of legs, there was present a minute tooth situated in

front of the first tooth, and separated from it by a distance equal

to one third the total length of the claw. The inferior claw

lanceolate and acuminate, unarmed. Pseudonychia large.

Ventral Tube.—Long, with highly protrusible bilobed vesicles ;

the anterior lobe of each four times the length of the posterior

lobe (fig. 67).

Hamula—The corpus with a stout median anterior seta

ORIENTAL COLLEMBOLA. 109

situated anterior to the rami. ‘The rami armed with four small

teeth.

Fureula.—Reaching to the ventral tube; clothed with long

sete (fig. 67). The mucrones quadrangular, armed with four

terminal teeth and a small lateral tooth on each side (fig. 65).

Coloration.—Purple-brown, somewhat paler in the mid-dorsal

region. ‘The head and first joint of the antenne darker than the

rest of the body. ‘The first and second antennal joints with

a distal band of cream-colour, the third and fourth joints entirely

cream-coloured with a slight purplish suffusion. The basal joints

of the legs, together with the femora, purplish brown ; the femora

with their apices cream-coloured. The tibize cream-coloured with

a proximal and distal band of purple. The ventral tube suffused

with purple. The fureula pallid with light purple suffusions.

Length varying from 3°5-4°5 mm.

Four specimens, taken among dry leaves and stones on the edge

of a jungle-stream at Maddathoray, W. base of W. Ghats,

Travancore, S. India (V. Annandale, November 18th, 1908).

No. Indian Museum Coll.

* PARONELLA GRACILIS, sp. n. (Pl. XI. figs. 77, 78.)

Segmentum abdominale quartum segmentis preecedentibus tribus

duplo longius. Antennw corpore longiore. Ocelli 16:8 im

utroque latere capitis. Unguiculus superior denticulis tribus

(vel duobus) instructus ; unguiculus inferior terms. Mucrones

lati. Long. 5 mm.

Head.—Considerably longer than broad. The eyes eight in

number on each side ; post-antennal organs absent.

Antenne.—Longer than the body, in some cases as long as the

combined length of the body and furcula. In full-grown examples

they vary from 6°5-7°5 mm. in length. The relative lengths of

the joints varying from the proportion of 15:15:10: 27 to

17: 16:10:30. Densely clothed with hairs ; on the basal joint

lanceolate scales are present among the hairs.

Trunk.—Clothed with small lanceolate scales, among which are

numerous hairs; a “fringe” of sete along the anterior border of

the mesothorax. The segments related proportionately in length

as 9:6:4:5:2:22:2:1. The fourth abdominal segment

double the length of the three preceding segments.

Legs.—Long and slender, clothed with slender, elongate sete.

The tibie divided by a joint into proximal and distal portions,

related respectively in length in the proportion of 2:1. A single

long stout tenent hair in relation with each foot. The superior

claws moderately slender, straight (fig. 77); armed with two

teeth, one placed at a distance from the base equal to one-third

the total length of the claw, the other placed at a similar distance

from the apex. Between the distal tooth and the apex of the

claw is a minute tooth which, however, is not always present.

110 PROF. A. D. IMMS ON

The inferior claws markedly acuminate, unarmed. Pseudonychia

large, projecting laterally.

Ventral Tube—1'75 mm. long, slender, cylindrical. The

vesicles each subdivided into a long anterior lobe and a shorter

posterior loke.

Furcula.—Reaching to the ventral tube; average length

2:25 mm. The dentes slightly tapering towards their extremities ;

related in length to the manubrium in the proportion of 7 : 5.

The mucrones (fig. 78) complex, wedge-shaped when viewed

in section; armed with a prominent terminal tooth, and a small

ventral tooth applied to the base of the latter; two lateral teeth

on the inner side of the mucro. The dorsal edge of the mucro

provided with four teeth. The apex of the dens provided with

a scale-like organ on the dorsal side, and a stout rod-like seta on

its inner aspect.

Coloration.—The ground colour varying from cream to pale

yellow, darkening according to the number of scales present.

‘he eyes on a black patch on each side of the head. The antenne

a little darker than the ground colour of the body, inclining in

some examples to pale brown; the basal joint longitudinally

streaked with violet-black. Body-markings varying from violet-

black to almost black, giving the insect a mottled appearance to

thenaked eye. The sides of the head and the lateral margins of the

thorax and first abdominal segment violet-black. A few lateral

markings of the same colour on the remaining abdominal

segments. The only dorsal marking is a narrow irregular trans-

verse streak crossing the head behind the bases of the antenne.

The femora marked with a distal band of violet; the proximal

tibial joint with both proximal and distal bands of the same colour ;

the distal tibial joint with a violet suffusion across the middle.

Tn very pale examples the body-markings are entirely absent,

only the legs retaining the usual coloration. In very dark

specimens the markings along the sides of the trunk are con-

fluent, and are united by transverse bands crossing the two

thoracic segments, and each of the first three segments of the

abdomen.

Length varying from 5—5'5 mm.

Twenty-two examples, taken among damp dead leaves in forest

of rhododendron and oak at Bhowali, Himalayan foot-hills of

Kumaon, 5700 ft. (A. D. Imms, October 18th-23rd, 1909).

No. hes Indian Museum Ooll.

* PARONELLA PHANOLEPIS, sp, n, (Pl. X. fig. 69; Pl. XI.

fig. 79.)

Unguiculus superior duobus parvis dentibus armatus ; wnguiculus

inferior lanceolatus, imermis. Mucrones lati, quattuor apicalibus

et tribus dorsalibus denticulis armati. Denticuli dorsales ita

collocati ut wnrus post alterum insertus sit. Articulus quartus

ORIENTAL COLLEMBOLA, 111

antennarum secundo et tertio longitudine equus, vel paullo longior.

Sete corporis longe, in fasciis instructe. Long. 3°5 mm.

Head.—Invested with scales and provided with a prominent

dorsal tuft of large sub-erect sete. The eyes eight in number on

each side ; no post-antennal organs.

Antenne.— Varying in length from 4 to 4:5 mm.; the joints

very variable in length. The first two joints sub-equal; the

fourth joint at least as long as the combined length of the second

and third joints. The exact numerical proportions in the length

of the antennal joints of four typical specimens were 50 : 50 : 32: 89;

good 2/37 295s 03) 2RAE Ge B8ie* and 5): 51's 35 86) ‘The

two proximal joints clothed with scales and hairs, the distal joints

entirely clothed with hairs of various lengths.

Trunk.—The segments related proportionately in length as

14:9:6:11:6:37:5:2. Clothed with small lanceolate scales

densely packed together; in the mid-dorsal line the scales

are considerably larger and oblong-ovate in shape. A prominent

investiture of large and very conspicuous sub-erect set@ with

curved extremities disposed in the following manner :—A “collar ”

along the anterior margin of the mesothorax, a few scattered

setz of similar type on the dorsal aspect of the segment and a

group near the posterior border. Similar groups are situated

near the posterior margins of the metathorax and the first two

abdominal segments. ‘he third abdominal segment with a few

scattered sete only. The fourth abdominal segment with a

conspicuous tuft of longer and more slender sets about the

middle of its dorsal aspect, and a fringe of similar set, directed

backwards, along its posterior and postero-lateral margins. The

fifth and sixth abdominal segments densely clothed with setz and

partially concealed by them.

Legs.—The femora and basal joints scaled. The tibie distinctly

separated into proximal and distal joints; the former related in

length to the latter as 19 : 7 on the third pair of legs, and as 17 : 7

on the first pair of legs; clothed with hairs and sete of various

lengths. The superior claws of the feet lanceolate and acuminate,

armed with twosmall teeth (fig.69); one tooth situated at adistance

from the base of the claw equal to one third the total length of

the latter, the second tooth situated at a similar distance from

the apex of theclaw. The inferior claws lanceolate and acuminate,

unarmed. Pseudonychia large, at least one half the length of the

inferior claw. <A single stout tenent hair to each foot.

Furewa.—In length varying from 2°5 to 2-75 mm.; the ratio

of the length of the manubrium to that of the dens varying from

11: 12to 3:4; in the majority of examples, however, the ratio is

as 4:5. The muero large and somewhat plate-like; armed with

an outer and inner apical tooth, each provided with a slender

lateral tooth prolonged down the mucro in the form of a ridge.

The inner apical tooth is continuous at its base with the dorsal

plate-lke portion of the mucro, The latter bears two large

Wily PROF, A. D. IMMS ON

backwardly directed teeth, situated just behind the inner apical

tooth, and a small acicular tooth near to the base of the mucro.

At the apex of the dens, on its inner aspect, is a small scale-like

organ (fig. 79).

Coloration.—Colour when alive leaden purple (in some speci-

mens almost black) with a slight metallic sheen. The antenne

and legs purple-black. ‘The furcula greyish white. In alcohol

specimens the colour is purple-black when but few of the scales

have been lost. In the majority of cases the specimens become

denuded of a large proportion of the scales, and the body-colour

then appears mottled and streaked with shades of brown and

purple-black.

Length varying from 3-3°75 mm.

Thirteen examples, taken on the inside walls of an old bungalow,

probably residing in decaying beams and rafters; Bhowali,

Himalayan foot-hills of Kumaon, circa 5700 ft. (A. D. Imms,

June 12th—29th, 1910).

ie, ee

Perfect examples of this species are easily recognised by the

conspicuous tufts of sub-erect sets ; in alcohol specimens the

latter fall off with extreme readiness and are usually absent.

In five of the specimens the two distal antennal joints are

lemon-yellow, and very conspicuous in consequence. Whether

this is a sexual or varietal difference I am unable to say.

Indian Museum Coil.

* PARONELLA INSIGNIS, sp.n. (Pl. XI. fig. 80; Pl. XII. figs. 81

& 82.)

Unguiculus superior uno dente armatus ; unguiculus inferior

aculeatus, inermis. Mucrones lati, duobus denticulis apicalibus

et tribus denticulis dorsalibus armati. Denticuli dorsales ita collocati

ut unus post alterwm sit insertus. Long. 37 mm.

Head.—A little longer than the mesothorax; covered with

scales, among which are a few scattered sete. The eyes cight in

number on either side ; post-antennal organs absent.

Antenne.—Very long and slender, exceeding 5 mm. in length.

The joints mutually related in length in the proportion of

LieW 34) 23%

Trunk.—longate, linear, clothed with scales. The scales

small and broadly lanceolate ; a tuft of sete at the extremity of

the abdomen. The segments related in length as 10:8:5:6:

5g DS 3) 32) (ame, iO)

Legs.—All the legs similar; the tibie distinctly divided into

proximal and distal portions by a definite joint. The proximal

portion a little more than twice the length of the distal portion

(i.e.,a8 15:7). The superior claws (fig. 81) of the feet slender

and tolerably long, armed with a single tooth situated at a

distance from the apex of the claw equal to one third the length

of the claw. In some examples there is also a rudimentary

tooth placed at a similar distance from the base of the claw.

ORTENTAL COLLEMBOLA, 115

The inferior claws nearly as long as the superior claws, unarmed ;

their distal portion abruptly narrowed, becoming acicular. A

single long stout tenent hair in relation to each foot.

Ventral Tube.—Long and cylindrical, emitting a pair of elongate

vesicles, each of the latter subdivided into a longer and shorter

lobe (fig. 80).

Furcula.—As long as the abdomen ; the manubrium related in

length to the dens as 10:13. The mucro (fig, 82) broad and

flattened, inclined at an obtuse angle with the long axis of the

dens ; armed with an outer and inner terminal tooth and a row

of three sharply pointed teeth along its dorsal edge, situated one

behind the other. The apex of the dens provided with stout

elongate sete slightly longer than the mucro (fig, 82),

Coloration.—The gvound colour varying from light yellow to

brown-yellow with markings of brownish purple, The antenne

purplish ; the posterior half of the head, together with the

lateral and anterior margins of the mesothorax, and the first

three abdominal segments, suffused with brownish purple. The

fourth abdominal segment suffused with the same colour along

its sides, and along its anterior and posterior margins, A broad

transverse band of similar colour crosses the middle of the seg-

ment, but is interrupted in the mid-dorsal line. The eyes ona

black patch on each side of the head. The fureyla and ventral

tube yellowish white. The Jegs with the tibial joints almost

entirely dark purple except for a whitish suffusion at the apex of

their distal joint. The femora suffused with purple, especially

towards their apices.

Length. Average length measures 3:7 mm.

Five examples, taken from among dry leaves and stones along

the edge of a jungle-stream at Maddathoray, W. base of W. Ghats,

Travancore, 8. India (VV. Annandale, November 18th, 1908).

No. a Indian Museum Coll.

In none of the five specimens were the antenne perfect, except

in the case of the example figured on Pl. VI. The number

of joints (3) is exceptional among Collembola, and it is not

unlikely that the antenna has been injured and has not re-

generated the full number of joints (4), Such examples are

known to be not infrequent in Jomocerus, Orchesella, and other

genera.

Genus IDIOMERUS, gen. nov,

Mesonotum prominens, conicus. Ocelli 16: 8 in utraque latere

capitis ; organa postantennalia carent, Antenne quinque-

articulate ; articulis basalis perparvus, ultimus longissimus.

Cutis squamosa.

This genus is easily recognisable by the remarkable form of the

mesonotum, which projects upwards in the form of a cone above

the level of the rest of the body (text-fig, 15).

Proc. Zoou, Soc.—1912, No. VIII. 8

114 PROF. A. D. IMMS ON

* TDIOMERUS PALLIDUS, sp. n. (PI. IX. fig. 61.)

Segmentum abdominale quartum fere dimidiam partem truncr

occupans. Unguiculus superior duobus parvis dentibus armatus ;

unguiculus inferior lanceolatus, inermis. Mucrones lati. Long.

35-5 mm.

Head.—Longer than broad, equal to the combined length of

the metathorax and first three abdominal segments ; clothed with

scales. The eyes eight in number on each side ; the post-antennal

organs absent.

Antenne.—Five-jointed, a little longer than the body. The

basal joint small, and concealed (text-fig. 15); the terminal joint

the largest, variable in length. The relative lengths of the joints

areas 1] : 24:21:17 : 32.

Trunk.—Clothed with scales; a “collar” of long setz along the

anterior margin of the mesothorax, and a tuft at the extremity of

the abdomen. The segments related to one another in length in

the proportion of 16:6:4:5:4:36:4:1. The fourth abdo-

minal segment occupying a little less than half the length of the

trunk, equal to the total length of the preceding trunk-segments.

Text-fig. ho.

Outline figure of Idiomerus pallidus, showing the characteristic shape of

the mesonotum,

Legs.—The legs sub-equal ; a single tenent hair at the apex of

each tibia. The sawperior claws moderately slender, acuminate }

armed with two minute teeth, inserted one behind the other on

the inner margin. The inferior claws lanceolate and acuminate,

unarmed. The psewdonychia large. The tibie biarticulate,

the proximal and distal portions related to one another in length

in the proportion of 15: 7.

Ventral 7ube.—Long and cylindrical.

Hamula.—Closely resembling that of Paronella borneri (Pl. X.

fig. 72); the corpus carrying a stout median seta between and in

front of the rami. The rami each armed with four small teeth.

Furcula.—Reaching to the ventral tube, clothed ventrally with

lanceolate scales. The dentes only slightly tapering distally; the

ORIENTAL COLLEMBOLA. 115

manubrium related in length to the dentes in the proportion of

26:29. The mucrones broad (tig. 61), armed with three large

rounded terminal teeth, a lateral tooth on the outer side, and an

erect dorsal tooth with a spiniform tooth at its base.

Yoloration.—The ground colour varying from a dirty cream-

colour to light yellow, with the furcula and ventral tube paler.

A few small purple-black markings on the head between the eyes,

and around the bases of the antenne. <A longitudinal. stripe

of the same colour along the greater part of the length of the

last pair of femora. In two specimens the other femora were

similarly streaked. The eyes ona black patch on either side of

the head.

In specimens retaining their full complement of scales the

ground colour appears darker and inclined to a brownish tinge.

Length varying from 3°5—-5 mm.

Five examples, taken among dry leaves and stones along the

edge of a jungle-stream at Maddathoray, W. base of W. Ghats,

Travancore, 8. India (VY. Annandale, November 18th, 1908).

No. a Indian Museum Coll.

In one example out of the five the upper claw of the feet was

devoid of teeth, and in a second specimen there was only ¢

single tooth present. The mucrones, however, supply much more

constant specific characters. =m

Genus Cypnoperus Nicolet,

Cyphodeirus Nicolet, Rech. pour serv. & Vhist, des Podur.,

Neuchatel, 1841, p. 63 (ad partem). 4

Cyphoderus Tullberg, “ Fort. 6fver Sv. Podur,”, Ofvers, Kongl.

Vet.-Akad. Forhandl. xxvii. 1871, p. 150,

CYPHODERUS SIMULANS, sp. n. (Pl. XII. figs. 90, 91.)

Omnino albus. Ocellinulli. Unguiculus superior uno magno et

duobus parvis dentibus instructus. Long. 1 mm,

Head.—Vhe eyes and post-antennal organs absent.

. Antenne.—Four-jointed, the joints related respectively in

length as 2: 7:3: 10; the terminal joint slender and tapering

distally. The length of the antenna is approximately half the

total length of the thorax and abdomen.

Trunk.—Closely resembling C. albinus Nic.

Legs.—All similar. The superior claw long, provided with

a very large basal tooth and two small distal teeth ; the inferior

claw bidentate at the apex, the ventral tooth being considerably °

shorter than the dorsal; a single very slightly curved ‘tenent

hair (fig. 90).

Furcula.—In length slightly exceeding the antenne, The

mvucrones (fig. 91) very long and slender, and when viewed

dorsally, quite straight ; the terminal and dorsal teeth similar to

those of C.albinus. The manubrium and dentes clothed ventrally

116 PROF. A, D. IMMS ON

with scales; the dentes provided with a double row of six large

scales, which are somewhat spear-shaped with a prominent mid-

rib; to the inner side of each mucro is a large terminal scale

Gigs i):

Coloration.—When alive, white (Annandale); in alcohol, pale

dirty cream-colour.

Length 1 mm.

Four specimens on bats’ dung in total darkness in the Khayon

Caves near Moulmein, Burma (WV. Annandale, March 7th, 1908).

No. == Indian Museum Coll.

The absence of eyes and pigmentation are characteristics of a

true cave-dwelling animal. Dr. Annandale informs me, however,

that the Khayon Caves are of no great extent, though their inner

parts are quite dark. :

Cyphoderus simulans is readily distinguished from C. albinus

Nic.f in possessing a somewhat longer superior claw to the feet,

with two small additional teeth. The latter, however, are difficult

to detect on account of their delicacy and transparency.

Genus PsEUDOCYPHODERUS, gen. nov.

Mesonotum non prominens. Segmentum abdominis quartum

quintuplo longius quam tertium. Antenne dimidiam partem

corporis cequantes, infra caput insert. Mucrones minuti. Manu-

brium biarticulatum. Ocelli et organa postantennalia carent.

Squamis et setis instructus.

This genus may be readily separated from Cyphoderus Nic.,

and Cyphoderodes Silvestrit, by the fact that the mouth and

frontal region of the head are completely ventral in position, and

consequently, the points of insertion of the antenne are situated

on the ventral aspect of the head.

* PSEUDOCYPHODERUS ANNANDALEI, sp.n. (Pl. XII. figs. 87-89.)

Antenne articulis tribus ultimis inter se longitudine sub-equal-

ibus. Unguiculus superior quadridentatus ; unguiculus inferior

bifidus. Dentes dimidiam partem manubri equantes ; quingue

squamis externis, tribus internis, et una squama terminali instructi.

Mucrones quadridentati, dentes ita collocati ut wnus post alterum

insertus sit. Long. 1 mm.

Head.—Considerably broader than long, the facial region,

together with the mouth, almost entirely ventral in position.

The eyes and post-antennal organs absent.

Antenne.— Arising from the ventral surface of the head, four-

jointed, the joints related to one another in length as 6: 14: 14:13.

Trunk.—Densely covered with small scales, among which are

some short scattered hairs of variable length. The head and

trunk segments related respectively in length as 15:12:7:4:4:

+ Vide Borner, “Zur Kenntnis der Apterygoten-Fauna von Bremen und der

Nachbardistrikte,’ Abh. Nat. Ver. Bremen, Bd. xvii. p. 71. fig. 28.

; a : > I? s

+ In * Termitenleben auf Ceylon.” von K, Escherich, Jena 1911, p. 244.

ORIENTAL COLLEMBOLA. wy

5:26:2:2. The fourth abdominal segment at least five times

as long as the preceding segment.

Legs.—Provided with short straight set. The superior claw

of the third pair of legs stout and thick, except at the apex where

it becomes swollen and membranous (fig. 87); one large acicular

tooth situated at the middle of the inner margin, a minute tooth

immediately in front of the former, and a small tooth close to

the apex of the claw on either side. The inferior claw wide and

plate-like, bifid. The claws of the first and second pairs of legs

smaller and somewhat shorter. A single tenent hair in relation

to each foot.

Furcula.—Moderate in size, stout, *4 mm. long, scaled ven-

trally. When closed up beneath the abdomen the mucrones

reach to the anterior border of the fourth abdominal segment.

The manubrium divided imperfectly into proximal and distal

portions (fig. 88). The dentes just about half the length of the

manubrium, short and stout; each provided along its outer

aspect with a dorsal row of five very large scales, a row of two

similar scales along the inner side, and a small terminal scale

just beneath the mucro (figs. 88 and 89). The proximal inner

scale equal in length to the proximal outer scale; the outer and

inner apical scales longest of all, the inner slightly longer than

the outer. The mwuero very small and armed with four teeth

(fig. 89); a very small terminal tooth, and three dorsal teeth

situated one behind the other; the posterior and middle teeth of

the row curved and pointed, the anterior (or proximal) tooth

blunt and rounded,

Coloration.—In alcohol, white.

Length 1-1:15 mm.

Twelve specimens taken from a nest of Termites at Rhamba,

south end of Lake Chilka, N.E. Madras (VV. Annandale, March

3rd, 1910). The Termite has been subsequently identified by

Prof. Silvestri as Termes redemanni Wasm.

N 444

Oo. — Indian Museum Coll.

Sub-order SYMPHYPLEONA Born.

Fam. SMintHvuRID# Lbk.

Sub-fam. SMINTHURIDIN2 Born.

Genus SMINTHURIDES Born.

+ Sminthurus (Smynthurus) Latreille, Hist. Nat. 1804, T. viii.

p. 79 (ad partem).

Sminthurides Borner, “ Zur Kennt. der Apteryg.-Fauna von

Bremen,” Abh. Nat. Ver. Bremen, 1901, Bd. xvii. p. 91.

* SMINTHURIDES APPENDICULATUS, sp.n. (PI. XII. figs. 83-86.)

Articulus ultimus antennarwm longissimus, vie annulatus.

Pili clavati in tibiis nulli. Ungwiculi superiores inermes ; ungur-

culi inferiores in setam longam prolongati. Unguiculi infertores

118 PROF. A. D. IMMS ON

pedum posteriorum etiam tribus appendicibus filiformibus instructi.

Mucrones furcule lati, laminati. Long. 5-75 mm.

Antenne.—Slender, the joints related to one another in length

as 8: 11:22:32; the terminal joint only with slight indications

of annulation (fig. 83).

Trunk.—The abdomen provided dorsally with a few short,

curved, scattered hairs.

Legs.—The first and second pairs similar (fig. 85); the superior

claws very long, at least two and a half times as long as the

breadth of the tibia, slightly curved at their apices, unarmed.

The inferior claw approximately one fifth longer than the superior

claw, setiform and whip-like, usually with a minute tooth on its

ventral aspect towards the base. The superior claws of the third

pair of legs shorter and smaller than those of the preceding

pairs; the inferior claws whip-like, armed at about the middle of

their length with a group of three filiform dorsal appendages, and

a small tooth situated close to the latter on the ventral surface

(fig. 84). No tenent hairs to the feet. The tibie of the third

pair of legs provided with three apical sense organs (#) on the

inner side (fig. 84).

Ventral Pube.— Very short, without elongate vesicles.

Purceula.—The dentes, without. their mucrones, equal in length

to the abdomen; slightly eurved ventralwards. The mucrones

very large (fig. 86), at least as wide as the maximum width of

the dens, lamellate.

Coloration.—Ground colour leaden, the legs and spring paler.

The eyes on a black patch on each sideof the head, The antenne

dark leaden coloured with a purplish suffusion. A pale yellow

dorsal area on the head, bearing a small bluish-purple patch

between the eyes. The trunk indigo-blue dorsally, with small

pale yellow markings.

Length varying from -5—-75 mm.

Twenty-one specimens taken on the surface of water at Calcutta

CUndian Museum Collector, January 9th, 21st, and 22nd, 1908).

: 4392

Nos. BR and

3 :

16 ig Indian Museum Coll.

TIl. A CATALOGUE OF THE ORIENTAL CoLLEMBOLA.

In defining the area comprised within the Oriental region, I

have followed Blanford * in taking for its northern boundary the

limits of forest growth in the Himalayas ; while as regards the

south-eastern boundary, I have followed the later views that

are wellsummarised by Pelseneer T and supported by the results

of the “Siboga” expedition. Pelseneer proposes the name of

& Weber’s Line” for the south-eastern boundary, which is regarded

as passing east of Timor and through the Banda and Molucca Seas.

* “The Distribution of Vertebrate Animals in India, Ceylon, and Burma.”

Phil. Trans. Roy. Soc. vol. 194, 1901, p.347. : pe

+ “Ta Lignede Weber, Limite Zoologique de Asie et de l’Australie.” Bull. Belg.

Acad. 1904.

ORIENTAL COLLEMBOLA. 119

With the small amount of knowledge of Oriental Collembola

at -one’s disposal it is impossible to make any generalisations.

Two features, however, stand out as notable:—(1) The paucity of

the members of the Sub-order Symphypleona, which is represented

by only three genera, and as many species, out of a total of fifty-

three species of Oriental Collembola. (2) The relative pre-

ponderance among species of the genus Paronella.

OO Ee eS)

14.

Y5.

16.

Sub-order ARTHROPLEONA Born.

Fam. Popurip# Lbk.

Sub-fam. HypoGastrurIn«% Born.

. Xenylla obscura, sp.n. India (W. Himalayas).

Sub-fam. Acnorutin Born.

. Protanura krepelint Born, Java. (Borner; Mitt. Natur-

hist. Mus. Hamburg, 1906, xxiii. p. 169.)

. Oudemansia cerulea Schott. Thousand Islands. (Schott ;

Ent. Tidskr. 1893, p. 172.)

. Achorutes lipaspis Born. Java. (Borner; Mitt. Naturhist.

Mus. Hamburg, 1906, xxii. p. 170.)

. A. hirtellus Born. Java. (Borner; loc. cit. p. 170.)

. A. armatus Nic. Sumatra. (Oudemans; Zool. Ergeb.

einer Reise in Niederl.-Ostind., Hft. i. p. 89.) Ceylon.

. A. crassus Oud. Sumatra. (Oudemans; loc. cit. p. 90.)

. Ceratameria (Schéttella) maxima Schott. Java. (Borner ;

Mitt. Naturhist, Mus. Hamburg, 1906, xxiil. p. 167.)

Sub-fam. NEANURIN«A Born.

. Pseudachorutes anomalus, sp.n. India. (BK. Himalayas.)

. Neanura corallina, sp. n. Ceylon.

. WV. pudibunda, sp.n. Lower Burma.

. WV. intermedia, sp. n. India (W. Himalayas).

. WV. fortis Oud. Java, Sumatra, and Saleyer. (Oudemans ;

Zool. Ergeb. einer Reise in Niederl.-Ostind., Hft. 1. p. 91.)

Sub-fam, OnycuruRiIn& Born.

Onychiurus (Lipura) fimetarius Burm. Sumatra, (Oude-

mans ; loc. cit. p. 90.)

Fam. ENtToMOBRYID& D. T.

Sub-fam. Isoromina Schiff.

Isotoma crassicornis Schott. Sumatra. (Schétt; Ent.

Tidskr. 1893,.p. 172.)

I. nigropunctata,sp.n. India (E. Himalayas).

120

Wha

18.

PROF, A. D. IMMS ON

Sub-fam. TomoceriIn& Schaff.

Tomocerus (Macrotoma) montanus Oud. Sumatra. (Oude--

mans; Zool. Ergeb. einer Reise in Niederl.-Ostind., Hft. 1.

10 Ske)

Sub-fam. Herrromuricin#&, sub-fam. nov.

Heteromuricus cercifer, gen. et sp. n. India (Bengal).

Sub-fam. EnroMosryiIn& Schiff.

. Isotomurus (Isotoma) palustris Mill., India (Bengal). Sub.-

sp. tricuspis Born. Java. (Borner; Mitt. Naturhist. Mus.

Hamburg, 1906, xxiii. p. 173.)

. Lepidocyrtus robustus, sp. n. India (Travancore).

. L. brauert Born. Seychelles. (Borner; Mitt. Naturhist.

Mus. Hamburg, 1906, xxiii. p. 176.)

. L.javanus Born. Java. (Borner ; loc. cit. p. 176.)

. L. variabilis Oud. Sumatra and Java. (Oudemans; Zool.

Ergeb. einer Reise in Nieder].-Ostind., Hft. i. p. 84.)

. L. javanicus Oud, Java. (Oudemans ; loc. cit. p. 85.)

. Entomobrya kali, sp. n. India (Bengal). Var lutea nov.

India (W. Himalayas).

. E. florensis Oud. Floves. (Oudemans; loc. cit. p. 86.)

. LE. longicornis Oud. Sumatra and Java, (Oudemans ; loc.

CEL Ds Sin)

. Sera (Sira) annulicornis Oud. Java, (Oudemans ; loc. cit.

oh (Sidhg)

. S. sumatrana Oud. Sumatra. (Oudemans; Joc. cit. p. 88.)

. S. brahma, sp.n. India (United Provinces).

. Pseudosira indra, sp.n. India (Bengal).

. Dicranocentroides fasciculatus, gen. et sp.n. India (W.Hima-

layas).

. Heteromurus tenwicornis Born, Java. (Borner; Mitt.

Naturhist. Mus. Hamburg, 1906, xxiii. p. 177.)

. H. tetracantha Born. Java. (Bérner; loc. cit. p. 177.)

. H. (Templetonia) sp.? Java. (Oudemans; Zool. Ergeb.

einer Reise in Niederl.-Ostind., Hft. i. p. 89.)

. Cremastocephalus celebensis Schiif. Celebes. (Schaffer ;

Archiv f. Naturges., 1898, p. 407.)

. C. montanus, sp. n. India (E. Himalayas).

. C. indicus, sp. n. India (United Provinces and Bengal).

. Paronella tarsata Born. Java. (Borner; Mitt. Naturhist.

Mus. Hamburg, 1906, xxiii. p. 177.)

. P. setigera Born. Java. (Biérner ; loc. cit. p. 178.)

, E. borneri, Sp. a. Niepail:

. P. travancorica, sp. n. India (Travancore).

. P. gracilis, sp. n. India (W. Himalayas).

. P. phanolepis, sp.n. India (W. Himalayas).

5. P. insignis, sp.n. India (Travancore).

ORIENTAL COLLEMBOLA, P21

46. [diomerus pallidus, gen. et sp.n. India (Travancore),

47. Cyphoderus simulans, sp.n. Burma.

48. C. javanus Born. Java. (Borner ; loc. cit. p. 180.)

49. Pseudocyphoderus annandalei, gen. et sp. n. India (N.E.

Madras).

50. Cyphoderodes ceylonicus Silv. Ceylon. (Silvestri ; in Ter-

mit. auf Ceylon von E. Escherich, 1911, p. 245.)

Sub-order SYMPHYPLEONA Born.

Fam. SMINTHURID2@ Lbk.

Sub-fam. SMINTHURIDIN&Z Born.

51. Sminthurides appendiculatus, sp. n. India (Bengal).

Sub-fam. CorynepHorin« Abs.

52. Corynephoria jacobsoni Abs. Java. (Absolon; Entom.

Ztg. Wien, 26, 1907, p. 338.) .

Sub-fam. Dicyrtromin# Born.

03. Ptenothrix gracilicornis Schiiff., subsp. gibbosa Born. Java.

(Borner; Mitt. Naturhist. Mus. Hamburg, 1906, xxiii.

p. 185.)

IV. A Summary or GENERAL ConcLusSIONS.

1. Four genera and twenty-seven species of Collembola are de-

scribed as new, and three species belonging to as many genera

were already known, Out of a total of thirty-one species, five

are Palearctic, and were obtained above the limits of forest-

growth in the Himalayas. The remaining species are Oriental.

2. The Palearctic species are all referable to well-known genera,

and were not met with in the Oriental region.

3. Among the Oriental species it has been found necessary

to erect four new genera, 1. e.:—IJdiomerus, Dicranocentroides,

Heteromuricus, and Pseudocyphoderus. The remaining species

all pertain to genera whose range extends into at least one

other zoo-geographical region.

4. Among the new forms discovered, the most remarkable is

Heteromuricus cercifer, gen. et sp.n. It is unique among Collem-

bola in possessing a median cercus to the fifth abdominal segment.

A new sub-family—the Heteromuricine—is proposed for its

reception. Psewdocyphoderus gen. nov., with a single species, is

described from the neighbourhood of Lake Chilka, where it

occurs in Termites’ nests.

5. The total number of Collembola known from the Oriental

region amounts to fifty-three species comprised within twenty-

seven genera, Of these only three genera, each with a single

species, are members of the sub-order Symphypleona.

122 PROF. A. D. IMMS ON

V. EXPLANATION OF THE PLATES.

The figures were made with a Leitz drawing apparatus under various magnifi-

cations. Wherever arrows are represented they indicate the direction of the anterior

end of the body.

Prate VI.

Pseudachorutes anomalus, sp. un.

Fig. 1. The right mucro viewed from the lateral aspect.

. The eyes and post-antennal organ of the right side.

. The right foot of the first pair of legs.

. An outline figure of the furcula seen from the dorsal side.

Bm owrDr

Xenylla obscura, sp. nD.

. An outline figure of the dorsal aspect of the furcula.

. A dorso-lateral view of the anal spines and their papilli.

. The mucro and apical portion of the dens of the left side viewed from the

inner aspect.

. The hamula.

. The right leg of the first pair.

£6 CO TO Ot

Neanura pudibunda, sp. n.

Fig. 10. The antenna, eyes, and post-antennal organ of the left side.

11. The left foot of the third pair of legs.

12. The insect viewed from the dorsal side.

Neanura intermedia, sp. n.

Fig. 12. The left antenna seen from the dorsal aspect.

14, The left foot of the first pair of legs.

Tsotomurus palustris Mull.

Fig. 15. The mucro, together with the apex of the dens ; right side.

Isotoma siva, sp. 0.

Fig. 16. The left foot of the first pair of legs.

17. The mucro and apex of the dens of the left side; viewed from the outer

side.

18. The eyes of the right side.

Puate VII.

Tsotoma siva (continued).

Fig.19. The insect seen from above.

Neanura intermedia (continued).

Fig. 20. The eyes of the left side.

Tsotomurus patustris (continued).

Fig.21. The right foot of the first pair of legs seen from the inner side.

22. The eyes and post-antennal organ of the right side.

Neanura corallina, sp. n.

Fig. 23. The insect viewed from above.

24. The left antenna seen from the ventral aspect.

25. The eyes of the left side.

26. The third leg, left side.

LIsotoma nigropunctata, sp. 0.

Fig. 27. The eyes and post-antennal organ of the right side.

28. The right foot of the third pair of legs seen from the imner aspect,

29. The apex of the dens of the right side together with the mucro.

Fig. 30.

31.

Fig. 48.

Fig. 49.

50.

61,

Fig. 62.

63.

54,

Fig. 55.

ORIENTAL COLLEMBOLA. 123

Entomobrya crassa, sp. n.

The mucro and distal portion of the dens of the right side.

The eyes of the right side.

Pseudosira indra, sp. n.

. The left mucro together with the apical portion of the dens, viewed from the

inside,

Entomobrya kali, sp. n.

. The right foot and apex of tibia of the third pair of legs; inside view.

Prats VIII.

Entomobrya kali (continued).

. The insect viewed from above. (The specimen figured has lost the longer

sete from the body.)

The left mucro and apical portion of the dens; inside view.

Typical sete from the mesothorax.

Pseudosira indra (continued).

. The eyes of the right side.

. The right foot of the third pair of legs.

. A typical seta from the anterior margin of the mesothorax.

. A typical body-scale.

Seira frigida, sp. n.

. The left foot of the third pair of legs.

The mucro and apex of the dens of the left side; inside view.

Seira brahma, sp. n.

The left foot of the third pair of legs.

. The left mucro and apical portion of the dens; viewed from the inside.

Dicranocentroides fasciculatus, gen. et sp. n.

The right foot of the first pair of legs.

. A portion of the inner aspect of the dens showing the characteristic

spines.

- Hairs from various regions of the body. (a) From the apical joint of

the antenna. (6) From the tuft on the second antennal joint. (c) An

acuminate hair from the legs.

Sinella montana, sp. n.

The right foot and apex of the tibia; third pair of legs. Sp., Spine-like

setae.

Heteromuricus cercifer, gen. et sp. n.

The right foot of the third pair of legs.

The cereus viewed laterally.

One of the larger body-scales.

Puate IX.

Heteromuricus cercifer (continued).

The insect viewed from the right side. (The head is inclined at a greater

angle with the trunk than is represented.)

The eyes of the right side.

The left mucro viewed from the outer aspect.

Dicranocentroidés fasciculatus (continued).

The left mucro viewed from the outer side. d., dens; 7. ¢., lateral tooth,

55 a, Body-scales,

Fig. 69.

Fig. 70.

ile

72.

73.

74.,

Fig. 75.

76.

Fig, 77.

78.

Fig. 79.

Fig, 80.

ON ORIENTAL COLLEMBOLA,

Sinella montana (continued).

The right mucro seen from the outside.

. (a) A seta from the mesothorax. (6) A seta from the extremity of the

abdomen.

Cremastocephalus indicus, sp. n.

. The right foot of the third pair of legs.

. The apex of the dens with the mucro of the left side; inner aspect.

Cremastocephalus montanis, sp.n.

. The apex of the dens with the mucro of the left side; inner aspect.

Idiomerus pallidus, gen. et sp. n.

. The right mucro viewed from the outer side.

Paronella travancorica, sp. vi.

. The left foot of the second pair of legs; inside view. ps., pseudonychium ;

t.h., tenent hair.

. The left foot of the second pair of legs viewed from above. ps., pseudo-

nychium ; ¢.h., tenent hair.

. The eyes of the right side.

. The left mucro seen from its outer aspect.

. Body-scales.

PLATE X.

Paronella travancorica (continued),

The insect seen from the right side.

Dicranocentroides fasciculatus (continued).

The insect seen from the right side. (The legs have been drawn too short,

the hind pair should reach almost to the extremity of the abdomen.)

Paronella phanolepis, sp. n.

The left foot of the first pair of legs.

Paronella borneri, sp. n.

The mucro and apex of the dens viewed from the outer side; right side.

d., dens; m., mucro; s.o., scale-like organ.

The foot and distal extremity of the tibia of the first pair of legs.

The hamula. The apex of the organ is directed towards the hinder end of

the animal. c., corpus; 7.. ramus; s., seta.

Hairs and scales. (a) From the anterior border of the mesothorax. (8)

From the fore leg. (¢c) From the general surface of the body.

The eyes of the right side.

Poate XI,

Paronella borneri (continued).

The insect viewed from above.

Entire leg of first pair.

Paronella gracilis, sp. n.

The left foot of the first pair of legs seen from the inner side,

The left mucro seen from the inner side.

Paronella phanolepis (continued).

The right mucro seen from the outer side.

Paronella insignis, sp. n.

The insect viewed from the left side.

Tee eis), WS) a, WT, XCHULL

West, Newman lith.

DEVELOPMENT OF THE BILL IN ARDEA CINEREA.

ON THE BILL OF THE HERON. 125

Puate XII.

Paronella insignis (continued).

Fig. 81. The right foot of the third pair of legs, seen from the outer aspect.

82. The right mucro seen from the outer aspect.

Sminthurides appendiculatus, sp. n.

Fig. 83. The left antenna seen from the inner side.

84. The left hind foot, inner side.

85. The left foot of the first pair of legs seen from the inner side.

86. The right mucro seen from its outer aspect. d.¢./., dorsal inner lamella;

d.o.1., dorsal outer lamella; v.2., ventral lamella.

Pseudocyphoderus annandalei, gen. et sp. n.

Fig. 87. Foot of third pair of legs.

88. The fureula viewed from the dorsal side. #.s., terminal scale.

89. The left dens and mucro from the outer side. ¢. s., terminal scale.

Cyphoderus simulans, sp. n.

Fig. 90. The left foot of the second pair of legs.

91. The left mucro seen from its outer aspect.

&. Ontogenetical Transformations of the Bill in the Heron

(Ardea cinerea.) By Prof. P. P. Susugin, C.M.Z.S.,

Kharkoy, Russia.

{Received June 3, 1911: Read November 7, 1911. |

(Plate XITT.*)

The birds of the subfamily Ardeine are known to have a conical,

pointed, spear-shaped bill with a simple rhamphotheca. The

curious fact that these features are acquired only in a late

post-embryonic stage seems to have attracted little attention. I

have been able to trace a gradual development of these features

in a series of embryos and young birds of Ardea cinerea,

collected and generously presented to me by one of my friends,

Mr. J. G. Sobolev, of Moscow. 3

In an embryo, in which the feather-papille on the back have

just assumed a conical shape (length of the gape about 12 mm.),

the bill is straight and rather slender, the tip of the upper jaw

is conspicuously swollen (not only owing to the presence of the

so-called egg-tooth), and the culmen is concave; a furrow runs

from the nostril to the base of the swollen tip (Pl. XIII, fig. 1).

In an embryo about one day before hatching (length of gape

22mm.) the form of the bill has already changed ; it is thicker,

its swollen tip is not so prominent and is slightly hooked, and the

culmen is not so coneave. The horny sheath of the bill is mani-

festing itself; the covering of the tips of both jaws is thicker and

more solid, and the covering of the tip of the upper jaw is marked

off by a raised area; a furrow anterior to the. nostril, and a space

* For explanation of the Plate see p, 126,

126 ON THE BILL OF THE HERON.

of softer and finely wrinkled skin between the eye and the nostril

divide the covering of the upper jaw into a dorsal and a paired

lateral plate. The covering of the mandibular rami is marked by

the presence of some parallel oblique ridges; these are perhaps

still more conspicuous just at the time of hatching, but quickly

disappear afterwards (fig. 2).

One day after hatching (length of gape 25mm.) the bill is

longer and conspicuously thicker, and the swelling of the tip has

almost disappeared. The covering of the tip of the upper jaw is

still marked off by its texture and relief, and the tip-covering of

the mandible is marked off by two irregular crack-like furrows

as well as by its solid texture. The division between the upper and

lateral plates of the upper mandible is plainly visible (fig. 3).

About three days after hatching (length of gape 32 mm.) the bill

has nearly assumed its conical shape, but is still relatively short

and thick and the tip is still conspicuously hooked. The horny

covering of the mandible-tip is still marked off by irregular

furrows; in the upper jaw it is no more distinct; the lateral

furrows before the nostril, dividing the upper and paired lateral

horny plates, are still clearly visible (fig. 4).

These changes seem to deserve attention on more than one

ground. The simple rhamphotheca proves to be only a late stage

of the compound one, as very clear vestiges of separate pieces are

visible in younger stages. In the upper jaw we find a tip piece,

a dorsal piece, and a paired lateral piece ; in the mandible there

are a tip piece and a paired lateral piece. The form of the Ardeine

bill proves to be a derivative one. The presence of the tip-hook

in the nearest relatives of the Herons, Scopus and Baleniceps, 1s

hence a primitive feature. In younger stages the shape of the

bill, combined with its compound rhamphotheca, recalls very

nearly that of a Cormorant, especially of a young one about the

time of hatching, and also of the nestling of a Frigate-bird, and

of a Pelican about the time of hatching. This points once more

to the affinities of the Ardez and Steganopodes. ‘The change in

shape of the bill in the Ardeine is obviously due to their habits

of spearing their prey instead of sjatching as most of the Stega-

nopodes do, The peculiar heron-lke shape of the bill in the

Darter (Plotus anhinga) is connected with the same habits, and

the likeness of the bill’s shape in the Herons and Plotus is to be

considered as a case of parallel evolution. The transitory ridges

on the mandibular rami recall remotely—by their direction also

—the structures so much developed in the Anseres, and also,

perhaps, the indentations of the bill in the fossil Odontopteryzx.

EXPLANATION OF PLATE XIII.

Fig. 1. Bill and head of an embryo of Ardea cinerea, length of gape 12 mm.

1X2.

2. The same, one day before hatching. 1X1°75.

3. Young bird, one day after hatching. 1X1‘.

4, The same, about three days after hatching. 1X1‘.

ON MAMMALS FROM WESTERN CHINA. NOL

9. The Duke of Bedford’s Zoological Exploration of Hastern

Asia.—XV. On Mammals from the Provinces of Sze-

chwan and Yunnan, Western China. By OLp¥FiELp

Tuomas, F.R.S., F.Z.8.*

[Received September 13, 1911: Read October 24, 1911.]

After the departure of his assistants, Dr. J. A. C. Smith and

Mr. Kingdon Ward, down the Yang-tze, with the remarkable

collection of mammals which formed the subject of paper

No. XIII. of the present series, Mr. Maleolm Anderson worked

on by himself froin September 1910 to the end of March 1911,

when he brought back to Shanghai the series on which the

present paper is based.

During this period he explored a number of localities in

Western Sze-chwan and N. Yunnan, and has been rewarded

by finding a considerable and quite unexpected number of species

additional to those previously sent. He has been especially

fortunate in discovering novelties among the Shrews and Voles,

while of the peculiar little Insectivores allied to Uropsilus he has

sent home the material on which I have found occasion to base

two new genera.

Altogether the set consists of about 160 specimens belonging to

33 species, and no less than eleven prove to need description as new.

To our great loss and regret Mr. Anderson now proposes to

give up the arduous life of the field collector, and this is

therefore the last paper to be written on his collections for the

Duke of Bedford’s Exploration of Eastern Asia. On this account

it is suitable to give a few words of appreciation for the manner

in which he has risen to the opportunity afforded by our

President’s continued generosity.

From the commencement of the work in 1904 to the sending

home of this last valuable collection in 1911 Mr. Anderson has

worked for the Exploration with superb enthusiasm and success,

and in the extent to which his collections have revolutionized our

knowledge of an extended part of the earth’s surface he has

made a record which, so far as I know, has never been equalled,

the nearest approach to it being Mr. OC, H. B. Grant’s collections

for Mr. Rudd’s Exploration of §8, Africa T.

In turn Mr. Anderson has visited and collected in Japan and

its islands, both the small southern ones and the large northern

Hokkaido and Saghalien, Korea, N. China and Mongolia, Shen-si,

Shan-si, Kansu, Sze-chwan and N, Yunnan, his collecting-localities

being dotted through all the previously most unexplored parts of

this little-known area.

Over 2700 mammals have been collected, besides many birds,

and, as is natural, a large number of novelties have been found

among them.

* Published by permission of the Trustees of the British Museum,

+ Cf. P.Z.S. 1908, p. 555.

128 MR. OLDFTELD THOMAS ON

On this occasion it may be useful to give a list of the papers

which have been written on the mammals collected by Mr.

Anderson :—

J. Japan. ~P.Z. 8. 1905, vol. 1. p. 331. 60ispp.

IT. Korea and Quelpart. 1906, p. 858. 9 spp.

III. Mindanao. 1907, p. 140. 6s

IV. Saghalien and Hokkaido. 1907, p. 404, 22 spp.

V. Korea (II.). 1907, p. 462. 13 spp.

VI. Shantung. 1908, p. 5. 6 spp.

Vil. Tsu-shima. 1908, p. 47. 11 spp.

(VIII. Fishes of Korea. 1908, p. 59

TX. Mongolian Plateau. 1908, p. 104. 9 spp.

X. Chih-li and Shan-si. 1908, p. 635. 20s

XJ. Shan-si and Shen-si. 1908, p. 963. 33 spp.

XII. 8. Shen-si. 1910, p. 635. 3 spp. nn.

XIII. Kansu and Sze-chwan. 1911, p.158. 48 spp.

XIV. S. Shen-si. 1911, p. 687. 31 spp.

XY. Sze-chwan and Yunnan. 1912, p.127. 32 spp.

The localities at which the present collection was made being

somewhat scattered, and but little known, it may be helpful to

give a list of them :—

Chin-fu-san, a mountain near the city of Nan-chwan, south of

Chung- king, not far north of the Sze-chwan— eae chow

bor den.

Hwa-yin-san, a limestone peak 50 miles N.K. of Chung-king.

Caves near Pen-hsien, about 35 miles N.W. by N. of Chen-tu.

Wei-choe, about 60 miles N.W. of Chen-tu, in the narrow

valley of the Si-ho (or Sung-pan-ho).

Wen-chwan, 12 miles lower down the Si-ho.

Yuen-ching-hsien, 8.W. of Ya-chow. ‘Part of the humid

range in which Mount Omi lies.”

Ning-yuen-fu, 8. of Fu-lin, on the Tung-ho.

Chao-tung-fu, N. Yunnan, about 103° 40’ E., 27° 30’ N.

1, Smt * sp. (S. rhesus group).

2. 2678. Mts. 30 miles 8.W. of Kia-ting-fu, Sze-chwan.

2. RHINOLOPHUS FERRUM-EQUINUM Schr.

3. 2592 (in al.), Caves near Pen-hsien, 35 miles N.W. of

Chen-tu. 3000.

3. RHINOLOPHUS CoRNUTUS PUMILUS K, And,

2. 2587, 2588, 2589, 2590 (in al.), 2591 (in al.) Caves near

Pen-hsien, 35 miles N.W, of Chen-tu, 3000’.

4, HiIpPosIDEROS ARMIGER Hodgs,

o . 2585,2586, Caves near Pen-hsien, Central Sze-chwan, 3000’.

* Simia Iinneus, Type S. sylvana, the Barbary Ape; Macaca and Pithecus

auctorum; see Thomas, P. Z.S. 1911, p. 125.

MAMMALS FROM WESTERN CHINA. 129

5. NycoraLus pLANcyi Gerbe.

3. 2677. Ya-chow-fu, Sze-chwan. 2500".

This Bat is just distinguishable—by its rather smaller size—

from the Nepalese WV. labiatus Hodgs., with which it shares the

reduced length of the outer incisors.

6. Urorsinus soricirrs M.-Edw.

3. 2605, 2637. Wei-choe, on Si-ho River, W. Sze-chwan.

9000'—12,500'.

Much to my surprise, these specimens show that, in spite of

the nearness of the localities and the almost complete identity

of external and cranial characters, the examples from Mt. Omi

referred in my previous paper to U. soricipes, with a note on

the discrepancy in the dental formula, belong to a distinct

species altogether—that discrepancy being the distinguishing

characteristic.

For these two specimens, like that figured by Milne-Edwards

and another one from Pére David’s series kindly lent me. by

Prof. 'Trouessart, have no trace either of the minute p* or 1,, both

present in the Omi species, nor any space in which they could

have stood. We must therefore accept the formula given by

Milne-Edwards as correct, and assign these examples to his

species.

This being the case, the Omi animal, with a constantly different

formula, will need not only description as a new species, but the

formation of a special genus for its reception.

And, furthermore, a single example of the group from Chin-fu-

san proves to have yet another formula, the small upper premolar

being present, and indeed better developed than in the Omi form,

the additional lower incisor is absent, while, alone in the group,

a p, 1s present.

The three animals may be briefly diagnosed as follows :—

URopsILus.

Teeth aX 2 = 34, neither p’, p,, nor i, present, the adult

formula being therefore

Lia a) I LO. & ee

. ae ei ae eee ba Bee

U. soricipes. Size fairly large. Colour near “ sepia ”; underside

of tail lighter than upper. (Winter specimens only seen.)

Nasinuus Thos.

Teeth = x 2 = 38; p’ and p, present; no i, (detailed formula

below).

Type. . gracilis.

Proc. Zoou. Soc.—1912, No, IX. !)

130 MR. OLDFIELD THOMAS ON

RuyncHonax Thos.

Teeth + x 2=38; p* andi, present; no p,. Complete dental

formula as follows :—

(alee 2 SE a Cale Oh asia

(ert a NK aac) (ne | DNS Bel Lee a8

ee Pr Sos es dh Co

i eames one | Slee tg «%3

Urea i Ce a

Type :—RHYNCHONAX ANDERSONI.

Uropsilus soricipes Thos. P. ZS. 1911, p. 163, nee M.-Edw.

R. andersoni Thos. Abstr. P. Z.8. 1911, p. 49 (Oct. 31).

Size comparatively large. Colour dark, approaching “ clove-

brown,” the lighter specimens near ‘“ bistre.” Under surface dark

slaty. Tail scaly, almost naked, scarcely lighter below than

above (summer specimens only seen). Teeth described /. ¢., but

the dental formula there given proves to be incorrect and should

be as above.

Dimensions of the type, measured in the flesh :—

Head and body 70mm. ; tail 67; hind foot 15-5.

Skull: greatest length 21:7; basal length 17:5; zygomatic

breadth 11-6; interorbital length 5°5 ; breadth of brain-case 11:3 ;

upper tooth-series 9°7.

Hab. Omi-san, Omi-hsien, 8. Sze-chwan, 9500",

Type. Adult male. B.M. No. 11.2.1.25. Original number 2504,

Collected 15 August, 1910. Nine specimens examined.

In naming this final species after Mr. Malcolm Anderson, I

would recall not only the wonderful success he has made of the

whole expedition, but also the extent to which he has increased

our material, and with it our knowledge, of this particular group

of Mammals. Practically the whole of our magnificent series of

Urotrichus was obtained by him, including the specimens on

which three new subspecies were founded, all our Uropsilus,

Rhynchonax, and Nasillus were sent by him, while even our

examples of Dymecodon were obtained by a Japanese whom he

had taught to collect.

7. NASILLUS GRACILIS.

Rhos, Abstr As02 19 lileap. AON (Oct..3!).

@, 2566. Mt. Chin-fu-san, near Nan-chwan, S.E. Sze-chwan.

4000’. 20 September, 1910. B.M. No. 11.9.1.13. Type.

A comparatively small species, with peculiar dental formula.

Size decidedly less than in the other species, the hind foot

1-2 mm. shorter, and the skull smaller in all dimensions.

General colour above near “sepia,” very much as in Uvropsilus

soricipes, Rhynchonax andersoni being darker, Under surface

MAMMALS FROM WESTERN CHINA, eat

slaty. Hands and feet pale brown. ‘Tail uniformly brown above

and below.

Skull shorter and decidedly narrower than in either of the other

species ; the brain-case less expanded laterally.

Dentition :—

Mier eee te eat ee 1 Hoe arene? 10 ¢

iS ee et a gas oe

Upper p* (the tooth which is absent in Uropsilus soricipes

and minute in Rhynchonax andersoni) comparatively large, as large

as the small anterior premolar in the horizontal area. Below

there is no trace of a minute incisor (i,) behind the large anterior

tooth, but, on the other hand, alone in the group, a p, is present

behind the tooth wrongly taken in my previous paper for p,, but

really p,; it is slightly smaller than the corresponding tooth in

the upper jaw and is therefore the smallest tooth present in the

animal.

Dimensions of the type, measured in the flesh :-—

Head and body 66 mm.; tail 55; hind foot 13:5; ear 9.

Skull: greatest length 20°5; basai length 16°5; zygomatic

breadth 10; interorbital breadth 5; breadth of brain-case 10 ;

upper tooth-series 9:1.

Hab. and Type as above.

This delicate little species proves to be of much interest, as its

peculiar dentition has caused the discovery of the error in the

dental formula of Rhynchonaw (under the name of Urepsilus)

previously published, and indicates that that animal has in the

lower jaw the very unusual premolar formula 1. 2.0.4, a formula

which in Uropsilus oceurs in the upper jaw as well,

This being the case, it is natural to examine again the

dentitions of the nearly allied Uvrotrichus, Dymecodon, and

Neurotrichus, and after some consideration [ venture to put

forward the following as their formule, modifying those I had

previously published :—

Urotrichus :—

eee ee | Jay:

ST aia cade ait palin Bi:

Dymecodon :—

pee 2 58 ih gy De py hs

Neurotrichus :—

Lees 1 OrGe O 4 Yeas 9

a = ———— eS

ee ee eer ore M. > 3-39 * 36.

It will be seen that, arguing from the analogy of the minute

teeth of Rhynchonax, which are missing in Uropsilus, | suggest

that it is the third lower incisor which is missing in Urotrichus,

32 MR. OLDFIELD THOMAS ON

and that in all, when one premolar is absent, it is the third and

not the second, whether above or below.

Of course these formule are still merely provisional, and liable

to modification when good examples of the respective milk-

dentitions are available for examination.

8. SorEX CYLINDRICAUDA M.-Edw.

@. 2672. Near Yuen-ching-hsien, Sze-chwan. 5200’.

9. SorEX WARDI FUMECLUS.

Thos. Abstr. P. Z.S. 1911, p. 49 (Oct. 31).

S. 2606, 2616, 2627, 2657. 2. 2638, 2656. Wei-choe, on

Si-ho River, W. Sze-chwan. 6000'—-11,000'.

Larger and darker-coloured than true wardi, the brain-case

broader. Fur about 6 mm. in length (winter). General colour

nearly the same brown as in S. bedfordie or a little greyer,

decidedly darker than in S. wardi. Dark dorsal line well marked

in all the specimens, rather more diffuse than in the other striped

species, but this may be due to the greater length of the winter

coat. Under surface markedly greyer than upper (‘‘ smoke-

grey”), while in S. bedfordie there is little difference between

the two.

Skull, like that of S. wardi, at once distinguishable from that

of S. bedfordie by its lower and less vaulted brain-case. From

that of the type of S. wardi it is separated by the rather larger size

and by the distinctly greater breadth of the low flat brain-case.

Dimensions of the type, measured in the flesh :—

Head and body 60 mm.; tail 60; hind foot 13; ear 8.

Skull : condylo-basal length 18:1 ; condylo-incisive length 18:5 ;

breadth across brain-case 8-4; upper tooth-series 8°1 ; height of

brain-case from basion 4°6.

Hab. as above.

Type. Adult male. B.M. No. 11.9.8.17. Original number

2627. Collected 24 November, 1910.

This little striped Shrew has a considerable resemblance to

S. bedfordie, but its flattened skull indicates that it is really

more allied to S. wardi, of which it may be considered a darker-

coloured representative.

10. SoRICULUS IRENE.

Thos. Abstr. P. Z.S. 1911, p. 49 (Oct. 31).

3. 2674. @. 2673. Yuen-ching-hsien, S.W. ‘Sze-chwan.

5200’.

General characters of S. macrurus, but brain-case much lower.

Size about as in S. macrurus. Fur soft and fine; hairs of back

(winter) about 6-5 mm. in length; about 5°0 mm. in what I

believe to be summer specimens of the same species. General

colour above uniform slaty grey (grey no. 4); under surface paler,

MAMMALS FROM WESTERN CHINA, 133

near smoke-grey. Hands and feet whitish, a httle darkening on

the metatarsals. Tail long and slender, uniformly grey-brown

above, whitish below. In the summer specimens the underside

of the tail is scarcely lighter than the upper.

Skull quite like that of S. macrurus, except that the brain-case

is conspicuously lower, the height nearly a millimetre less, the

length and width being about the same. Teeth rather more

heavily pigmented.

Dimensions of the type, measured in the flesh :—

Head and body 60 mm.; tail 90; hind foot 16; ear 10:5.

Skull : condylo-incisive length 17-8; condylo-basal length 17-2;

greatest breadth 9; height of brain-case from basion 4°9; length

of upper tooth-series 7-4.

Hab. of type as above ; specimens from Mt. Omi believed to be

the same.

Type. Adult female. B.M. No. 11.9.8.22. Original number 2673.

‘ollected 13 January, 1911.

The specimens from Mt. Omi collected in August, 1910, are

darker-coloured and their tails are scarcely lighter below, but

they agree with the Yuen-ching examples in all other characters,

and it seems probable that their differences are merely seasonal.

They were formerly identified by me with S. maerurus of

Darjiling, their distinctive character, the much lower brain-case,

not becoming evident until the arrival of the present additional

material,

11. CHopsicoa HypstBiA de Wint.

@. 2604, 2642. Wei-choe, on Si-ho River, W. Sze-chwan.

5900'—7000'.

These Shrews, collected in mid-winter, bear the same relation in

colour to de Winton’s type that the winter specimens of Sori-

culus irene do to those which I consider to be summer skins of

the same animal, their grey being similarly paler and their tails

more whitish. The type locality is by no means distant, nor is

that of Ch. berezowskii, which, as previously stated, I believe to be

of the same species. :

The skulls of these specimens, however, being perfect, indicate

that the Chodsiyoa previously obtained by Mr. Anderson in

Chih-li, and assigned to Ch. hypsibia, must be separated speci-

fically. It may be called

CHODSIGOA LARVARUM.

Thos. Abstr. P. Z. 8. 1911, p. 49 (Oct. 31).

General characters of Ch. hypsibia, brain-case narrower and less

flattened.

Fur close and fine; hairs of back about 3°8 mm. in length.

General colour above ‘ mouse-grey,” rather lighter, apparently

“ smoke-grey ” below. Hands and feet whitish, Toil brown

above, dull whitish brown below.

134 MR. OLDFIELD THOMAS ON

Skull, although showing the peculiar flattening and concavity

of the upper profile-line characteristic of the genus, not so

extreme as that of Ch. hypsibia, the top of the brain-case fairly

vaulted, and the sides not splayed out. Teeth quite as in

Ch. hypsibia.

Dimensions of the type, measured in the flesh :—

Head and body 68 mm.; teil 50; hind foot 14; ear 8°5.

Skull: condylo-incisive length 19:8 ; condylo-basal length 18°8 ;

greatest breadth across brain-case 8°8; height of brain-case from

basion 5; upper tooth-series 8°6.

Hab. Traperial tombs, 65 miles E. ef Peking, Chih-li. 1000".

Type. Adult female. B.M. No. 8.8.7.21. Original number 1559.

Collected 25 September, 1907, by M. P. Anderson. Two

specimens.

12. BLARINELLA QUADRATICAUDA M.-Edw.

9.2567. Near Nan-chwan, S.E. Sze-chwan. 4000’.

13, CrocriDURA ATTENUATA M.-Edw.

@. 2565, 2572, 2573. Chin-fu-san, near Nan-chwan, 8.E.

Sze-chwan. 4000’.

3g. 2575, 2581. @. 2576. Mts. 50 miles N.E. of Chung-king.

3000!.

14. AnouRosoREX squamipEs M.-Edw.

@. 2570, 2574. Chin-fu-san, nedr Nan-chwan, 8.E. Sze-

echwan. 4000’.

Q. 2577, 2582. Mts. 50 miles N.E. of Chung-king. 3000’.

3d. 2675. Near Yuen-ching-hsien, W. Sze-chwan.

3. 2693, 2697, 2698, 2708. 2. 2707. 21 miles N.E. of

Chao-tung-fu, Yunnan. 5800’.

15. Arcronyx opscurus M.-Edw.

. 2651, 2662, 2663. Wei-choe, Si-ho R., W. Sze-chwan.

16. ScruRUS CASTANEOVENTRIS BONHOTEI Robinson & Wrought.

| 9. 2676. Yuen-ching-hsien, S.W. of Ya-chow, Sze-chwan.

3000’.

@. 2679. Ning-yuen-fu, 8. of Fu-lin, S.W. Sze-chwan.

This subspecies has only been recently distinguished on the

basis of Sze-chwan specimens jpuesene ed to the Museum by

Mr. F. W. Styan.

“ A forest-loving species; rare. ”__ ME. BEA,

17. SCIUROTAMIAS DAVIDIANUS CONSOBRINUS M.-Edw.

2. 2669. Wen-chwan-hsien, Si-ho R., Sze-chwan. 6000’.

MAMMALS FROM WESTERN CHINA, 135

18, Epimys numiiiatrus M.-Edw,

2. 2701. 21 miles E. of Chao-tung-fu, N. Yunnan. 5800’.

This is an interesting animal as being a wild species closely

allied to the ubiquitous pest, /. norvegicus, from which it chiefly

differs by its smaller size. The flesh-measurements of Mr, An-

derson’s specimens are as follows :—Head and body 160 mm. ;

tail 130; hind foot 29; ear 17.

Milne-Edwards’s J/us plumbeus appears to be the young of the

same form, and it is also probable that his J/. owang-thome is not

distinguishable from it.

19. Eetys conructanus M.-Edw.

3. 2568, 2569 (young). Chin-fu-san, near Nan-chwan, S.E.

Sze-chwan. 4000’ (September).

3. 2583. 2. 2584 (young). Hwa-yin-san, 50 miles N.E. of

Chung-king. 4000" (October).

dg. 2594, 2600, 2602, 2603, 2641, 2643, 2658. 92. 2599,

2659. Wei-choe, on Si-ho R., W. Sze-chwan. —5900'—7000'

(November—December).

3. 2682. 21 miles E. of Chao-tung-fu, Yunnan (March).

The September and October specimens are distinctly spinous,

the others quite soft-furred.

20, APODEMUS CHEVRIERI M.-Edw.

3. 2595, 2596, 2608. Wei-choe, Si-ho R., W. Sze-chwan.

5900'—7000'.

db. 2680, 2681, 2682, 2683, 2688, 2689, 2691. 2. 2686,

2687, 2690. 30 miles 8.E. of Ning-yuen-fu, 8. Sze-chwan. 4500’.

Be enlo. ai2e, ©, 2112, 2hloe 2pli, ates, 2024, 2727.

Chao-tung-fu, N. Yunnan. 6400’.

A. agrarius group. ‘Tail and ears comparatively short.

M? without antero-external secondary cusp. M®* with only two

internal projections.

On finding both this species and the next in considerable

numbers in the Sze-chwan collection, I have thought it advisable

to verify the determination of Milne-Edwards’s Mus chevrieri,

which, thanks to the kindness of Prof. Trouessart, I have now

been able to do.

To my great regret I find that my suspicions about the

determination were justified, and that, instead of being the

common and widely spread species allied to A. speciosus, as has

been supposed, chevrieri is the rarer shorter-tailed form only

recently sent us by Messrs. Anderson and Fergusson, the mouse I

described as A. fergussoni being the same species, though perhaps

subspecifically separable.

Fortunately, like as the two species are externally, the determi-

nation of specimens is easy enough by the dental characters

given above, characters which show clearly in Milne- Edwards’s

type, although the teeth are worn down.

136 MR. OLDFIELD THOMAS ON

In all papers previous to this therefore* the name chevriert

must be entirely withdrawn from the member of the speciosus

group to which it has been wrongly attached.

With regard to fergussoni, more material from Kan-su is

needed before its characters can be elucidated ; but there is some

evidence that the northern specimens of chevrieri, e. g. those from

Wei-choe, have less fulvous suffusion on the hinder back than

the southern ones, the type of fergussoni being completely without

it. For the present, therefore, we may provisionally call the

Kan-su form Apodemus chevrieri fergussoni, with the Wei-choe

specimens as connecting intermediates.

21. APODEMUS SPECIOSUS PENINSUL& Thos.

3g. 2578, 2580. 9. 2579. Hwa-yin-san, 50 miles N.E. of

Chung-king. 3000’.

a. 2598, 92647, 2649) 2652, 26538012661.) 2. 2593" 2507,

2601, 2644, 2645, 2646, 2648, 2650, 2655, 2660. Wei-choe,

Si-ho R., W. Sze-chwan. 5900'-7500'.

A. speciosus group. ‘Tail and ears comparatively long. Molars

more complicated; m? with antero-external secondary cusp;

m* with three internal projections.

22, APODEMUS SPECIOSUS ORESTES.

Nos, Aas, 12/4, So WOU, tos AD (Certs sill),

A. s. chevrieri M.-Edw., Thos. P.Z.S. 1911, p. 172 (in part.).

3. 2610, 2622, 2624, 2636, 2654. 9. 2623, 2625, 2626,

2635. Wei-choe, R. Si-ho. 7000'-12,000'.

As explained above, the common long-tailed Field-mouse of Sze-

chwan proves to have no claim to the name chevriert, hitherto

supposed to be applicable to it, and we have now to consider

what it should be called.

On laying out again the fine series received from Mr. Anderson,

we find, to start with, that the specimens from Mt. Omi are -~

uniformly dark-coloured and clearly form a race distinct from the

pale 4. s. peninsularis.

This may be called

A. 8. ORESTES.

Size about as in A. s. peninsule or rather larger. Tail longer

than head and body. Lars of medium size. Colour very dark,

the back (im summer specimens) slightly more rufous than

Ridgway’s “sepia.” Under surface grey no. 7.

Skull slenderly built, of medium size.

Dimensions, taken in the flesh :— Extremes

Type. of Omi series.

Hlendanid jue cytee- nase eae 93mm. 85 — 98mm.

etl) ee LR ee. ae ae Ion, 102 -125 ,,

itlanid hoot! vase. wek 2 eae 24 ,, 22°5— 25 ,,

UE As Fea RAR CMO Fora te R 16 15:5-— 18

7 9

* From Barrett-Hamilton, P. Z.S. 1900, p. 418 onwards.

MAMMALS FROM WESTERN CHINA. 37

Skull: greatest length 27°5 mm. ; condylo-incisive length 24°8 ;

interorbital breadth 4:7; palatilar length 11°38; palatal fora-

mina 5:7; upper molar series 4-2.

Typical locality. Mt. Oi, W. Sze-chwan. 6000'-10,000'.

Type. Adult male. B.M. No. 11.2.1.170. Original number

2535. Collected 18 August, 1910.

This subspecies differs from peninsule by its darker colour and

from the W. Fokien draco B.-Ham. by its larger size, longer tail,

and less rufous tone.

The Wei-choe series of the present collection contains examples

agreeing with peninsule and others with orestes, and on exami-

nation it proves that all of the latter are from altitudes above

7000', while all of the former are below that height. On the

other hand, on Mt. Omi those from 6000' are of quite the same

colour as those from 9000’, so that it cannot be said that the

darker colour is a direct result of the altitude.

23. APODEMUS SPECIOSUS LATRONUM.

Thos. Abstr. P.Z.8. 1911, p. 49 (Oct. 31).

go. 2726, 2728. Chao-tung-fu, Yunnan. 6400’.

(Series from Ta-tsien-lu previously enumerated.)

A large form, with large ears and comparatively short tail.

Size decidedly larger than in A. s. orestes. General colour above

near “ broccoli-brown ”; under surface grey no. 7, the hairs slaty

basally, whitish terminally. Ears very large, their visible surfaces

black. Hands and feet white. Tail ranging from a little shorter

to a little longer than the head and body, blackish above, white

below.

Skull conspicuously larger than in the other W. Chinese forms,

as large as in true Japanese speciosus.

Dimensions :— Extremes of

Type. Ta-tsien-lu series.

Wead and body :.5..--+=5- 107 mm. 95-108 mm.

I Tl Seattle cera TOUTS LOO] 3

EMO MOOh Stott cies oss acess 253 24- 26 ,,

| rig leemtetas eek sit enaeeg teeing ay 18- 20 ,,

Skull: greatest length 29°6 mm. ; condylo-incisive length 26°6 ;

interorbital breadth 4:3; palatilar length 12°8 ; palatal foramina

6:1; upper molar series 4°6.

Typical locality. Ta-tsien-lu, W. Sze-chwan. 9000’,

Type. Adult male. B.M. No, 11.2.1.156. Original number

2377. Collected 30 June, 1910.

This distinct large-eared form would seem to range over the

mountainous country to the west and south of the area inhabited

by A. s. orestes.

24, Micromys pyemaus M.-Edw.

3. 2684, 2685. 30 miles 8.E. of Ning-yuen-fu, $.W. Sze-

chwan. 4500’,

138 MR. OLDFIELD THOMAS ON

25. MIcROTUS MILLICENS.

hos: Abstr PoZe Ss. 191 tip. 491(@cta 2).

3. 2613, 2614, 2615, 2621, 2639. 9. 2632, 2640. Wei-

choe, on Si-ho R., W. Sze-chwan. 12,000'-12,500’.

A greyish species with a delicate flattened skull, long tail, and

extra angle on m°.

Size medium, about as in J/. irene. Fur long, soft, and loose ;

hairs of back (in winter pelage) about 12 mim. in length. General

colour above rather darker than ‘ hair-brown,” passing into

‘“*smoke-grey ” below. Hars scarcely projecting above the fur,

well-haired, greyish brown. Head and feet glossy greyish white,

varying to greyish brown ; soles with 6 pads. Tail comparatively

long, well-haired, grey-brown above, whitish below. Mammz

2—2=8.

Skull rather hghtly built, with a long flattened oval brain-case

without marked ridges or angles. Upper profile-line flat or

even slightly concave at a point above m*. Rostrum of medium

length, narrow. Interorbital region smooth, not ridged, flat-

tened above. Angles of brain-case scarcely perceptible. Posterior

palate normal, the lateral pits of medium depth.

Incisors rather thrown forward. Molars narrow. M!' with the

usual five spaces, all separated, a scarcely perceptible trace of a

supplementary postero-internal projection. M? with a large

supplementary postero-internal salient angle, making three angles

on the inner side. M?* narrow, with the usual transverse first

space, the second and third small, united to each other, and a

posterior U ; three inner and three outer salient angles (the whole

somewhat as in Blasius’s figure 221 *, except that the anterior

transverse lobe is narrower, and the second and third spaces are

united). M, with a large posterior transverse lobe, four lateral

closed triangles in front of it, and a fifth united to the anterior

trefoil, there being in all five inner and four outer salient angles

(near fig. 193 of Blasius, if the first lateral closed triangle were

open to the anterior trefoil). M, with the anterior two of the

usual five spaces united to each other; three inner and three

outer angles.

In short, the most salient features of the dentition are the

development of the extra postero-internal angle on m” and the

junction of the second and third spaces of m* and first and

second of m,,.

Dimensions of the type, measured in the flesh :-—

Head and body 90 mm.; tail 53; hind foot 18-5; ear 14.

Skull : condylo-basal length 24°3 ; condylo-incisive length 24°6 ;

zygomatic breadth 14:2; nasals 6:7; interorbital breadth 4:2 ;

brain-case, length 12:5, breadth 11°5, height including bulle 9,

height of supraorbital edge above root of m* 6-9; palatilar length

12:5; palatal foramina 4:8 ; upper molar series (crowns) 5:6.

Hab. as above.

* J. H. Blasius, Naturg. der Sdugethiere Deutschlands, 1857.

MAMMALS FROM WESTERN CHINA, 139

Type. Adult male. B.M. No. 11.9.8.105. Original number

2615. Collected 20 November, 1910.

This striking species, from the considerable altitude of 12,000’,

is readily recognizable by its smooth flattened skull and long

tail, the latter character separating it at once from J/. irene, to

which alone of the Chinese species hitherto described it has any

resemblance.

In its possession of an extra angle on m’* it has a curious like-

ness to many of the species of Hothenomys and Caryomys, this

character in true Microfus being only hitherto known in

M. agrestis L. and sikimensis Hodgs.

26. Mricrorus (EoTHENOMYS) MELANOGASTER ELEUSIS.

Thos. Abstr. P. Z.S. 1911, p. 50 (Oct. 31).

3. 2694, 2695, 2696, 2700, 27038, 2704, 2709, 2710, 2711.

@. 2699, 2702, 2705, 2706. 21 miles east of Chao-tung-fu,

N. Yunnan. 5800’.

[?¢d. 2571. Chin-fu-san, near Nan-schwan, 8.E. Sze-chwan.

4000’. |

Longer-tailed than true melanogaster, with more complicated

m’.

General characters quite as in true melanogaster, though the

colour may average slightly more greyish brown. Head and feet

dark. Tail decidedly longer than in melanogaster, the shortest

measured 43 mm., the majority about 46 or 48, and the longest

55; on the other hand, in a number of melanogaster the majority

are 34-36, a single isolated example 43.

Skull and teeth asin melanogaster, except that m’ is in nearly all

cases more complicated, the posterior U-shaped lobe being rather

a C, so that there are four inner salient angles instead of three.

Dimensions of the type, measured in the flesh :—

Head and body 98 mm. ; tail 55; hind foot 17; ear 11-5.

Skull: condylo-incisive length 24°7 ; basilar length 22; zygo-

matic breadth 15; nasals 7°4; interorbital breadth 4:2; breadth

of brain-case 12°3; height of crown above alveolus of m* 7-7;

palatilar length 11:8; palatal foramina 5:1; upper molar series

(crowns) 5:6,

Hab. East of Chao-tung-fu, N. Yunnan. 5800’,

Type. Adult male. B.M. No. 11.9.8.111. Original number

2696. Collected 13 March, 1911.

The Chin-fu-san specimen is a little indeterminate in character,

the teeth of the two sides not being quite alike.

27. Microrus (KoTHENoMys) OLITOR.

Thos. Abstr, P.Z. S.,1911, p. 50 (Oct. 31).

Hamdten 2.2714, 2716, 2719, 2720) 2721, 2725... Chac-

tung-fu, Yunnan. 6700’.

A short-tailed sepia-brown Lothenomys without supplementary

projection on in’,

140 MR. OLDFIELD THOMAS ON

Size rather smaller than in J/. (Z.) melanogaster. General

colour a Jess warm brown than in that animal, the upper surface

nearest to ‘‘sepia”; under surface dark slaty grey (rather more

smoky than “grey no. 5”). Hands and feet brown, soles with

6 pads. Tail dark brown above, little lighter below.

Skull rather smaller than that of J/. melanogaster, but similar

to it in general form ; the bulle not quite so large.

Teeth agreeing with those of melanogaster in the junction to

each other of nearly all the opposite spaces of the teeth, and

therefore equally differing from the members, now four in number,

of the subgenus Caryomys. But they are readily distinguished

by the fact that m* has not got the characteristic supplementary

postero-internal salient projection occurring in both m' and m*

of melanogaster, and here present of full size in m*; as a result,

there are three internal angles on both these teeth. M* more

complicated than in melanogaster, with four salient angles on each

side; it consists of three separated dentine spaces, the first with

one inner and two outer angles, the middle one with one external

and one internal angle, and the posterior with two internal and

one antero-external angle. Lower teeth essentially as in melano-

gaster, the spaces all opposite and united.

Dimensions of the type, measured in the flesh :—

Head and body 82 mm.; tail 34; hind foot 16; ear 9.

Skull: condylo-incisive length 24; zygomatic breadth 15-4;

nasals 6°8; interorbital breadth 4; height of crown from alveolus

of m* 7-5; breadth of brain-case 11; palatilar length 11-5 ; palatal

foramina 4°7; upper molar series (crowns) 5:5.

Hab. as above.

Type. Adult female. B.M. No. 11.9.8.122. Original number

2714. Collected 19 March, 1911.

The dentition of this Vole is interesting, as it conforms strictly

to the Hothenomys as opposed to the Caryomys type, while pre-

senting such differences from that of J/. (1.) melanogaster as

readily to distinguish the two species.

“Trapped in the open fields of the little cultivated plaim sur-

rounding the city.”.—M. P. A.

28. Mrcrorus (CARYoMYS) ALCINOUS.

Thos. Abstr. P. Z.S. 1911, p. 50 (Oct. 31).

3. 2609, 2611, 2612, 2617, 2618, 2620, 2628, 2629, 2630, 2631,

2633, 26384. 2. 2619. Wei-choe, Si-ho R., W. Sze-chwan.

8000'-12,000'.

Essential characters as in J/. (C.) eva, but colour much darker.

Proportions, skull, and teeth all about as in eva. Fur long

and soft; hairs of back about 9-10 mm. in length (winter).

General colour above uniform dark bistre-brown, conspicuously

different from the pale brown J/. eva; under surface dark

smoky grey, decidedly darker than Ridgway’s ‘“smoke-grey.”

Ears about as long as the fur, their backs with some whitish

MAMMALS FROM WESTERN CHINA. 141

hairs on them, making an inconspicuous grey patch. Hands and

feet dark brown. ‘Tail as long as in JZ, eva, blackish above, little

lighter below.

Skull apparently very much as in MZ. eva, though the bull are

perhaps a little smaller.

Dimensions of the type, measured in the flesh :—

Head and body 90 mm. ; tail 56; hind foot 17; ear 11:5.

Skull: greatest length 24; condylo-incisive length 23:8 ; zygo-

matic breadth 14; nasals 7; interorbital breadth 4:3; breadth of

brain-case 11°3; height of crown from alveolus of m* 7:1 ;

palatilar length 11:2; palatal foramina 5; upper molar series

(crowns) 5:7.

Hab. as above.

Type. Adult male. B.M. No. 11.9.8.1386. Original number

2631. Collected 24 November, 1910.

The close agreement of this species with JM. (C.) eva in its

proportions, notably in the unusually long tail and in its skull

and dentition, is remarkable, as in its colour it is so entirely

different that one would have expected to find distinguishing

characters in other respects.

29. GAPRICORNIS MILNE-EDWARDST David.

©. 2667. Wen-chwan-hsien, Si-ho R., W. Sze-chwan.

“Tnhabits the forests and cliffs at altitudes above 6000’. 1t

is not so common as Vemorhedus cinereus.

‘* Chinese name ‘ Ngai-lii’? = Cliff Donkey, probably so called

because of the long ears.”— M. P. A.

30. NA&MORHEDUS CINEREUS M.-Edw.

3. 2665, 2666. 9. 2664, 2671 (imm.). Wen-chwan-hsien,

Si-ho R., W. Sze-chwan. 6000'—7000’.

31. ELAPHODUS CEPHALOPHUS M.-Edw.

3. 2668. Wen-chwan-hsien, Si-ho R., Sze-chwan. 8000.

In the dark winter pelage, that figured by Milne-Edwards being

in the rufous summer coat.

“Very shy and retiring, but not really rare.”—M. P. A.

32. Moscuus siFanicus Buchn.

3. 2670 (imm.). Wen-chwan-hsien, Si-ho R., W. 8ze-chwan,

142 MR. R. I. POCOCK ON ELEPHANT-SHREWS.

EXHIBITIONS AND NOTICES.

November 21, 1911.

S. F. Harmer, Esq., M.A., Se.D., F.R.S., Vice-President,

in the Chair.

Tue Secretary read the following report on the additions

made to the Society’s Menagerie during the month of October

1911 :-—

The number of registered additions to the Society’s Menagerie

during the month of October last was 287. Of these 141 were

acquired by presentation, 42 by purchase, 34 were received on

deposit, 28 in exchange, and 42 were born in the Gardens.

The number of departures during the same period, by deaths

and removals, was 391.

Amongst the additions special attention may be directed to :—

1 Budgett’s Cercopitheque (Cercopithecus tantalus budgetti), 2

Matschie’s Guerezas (Colobus matschiei), and 1 Shari River

Hunting-Dog (Lycaon pictus sharicus), new to the Collection,

from the Northern Congo, deposited on October 12th.

1 Duke of Bedford’s Wapiti (Cervus xanthopygius) 3, from

Manchuria, deposited on October 9th.

1 Waterhouse’s Genet (Genetta poénsis), from 8. Nigeria, new

to the Collection, presented by W. A. Lambourne, Esq., on

October Ist.

1 Indian Redstart (Ruticilla rufiventris), new to the Collection,

received in exchange on October 25th.

3 Myrtle Warblers (Dendraca coronata), new to the Col-

lection, from North America, presented by HE. Hollman, Esq., on

October 25th.

4 Peale’s Parrot-Finches (Zrythrura pealei), from the Fiji

Tslands, presented by Dr. Philip H. Bahr, F.Z.S., on October

19th.

1 Rainbow-Bunting (Cyanospiza leclancheri), from. Western

Mexico, new to the Collection, received in exchange on October

6th.

1 Malayan Wrinkled Hornbill (/hytidoceros wndulatus),

received in exchange on October 6th.

2 Ring-necked Teal (Netiiwn torquatum), from South

America, new to the Collection, received in exchange on

October 20th.

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

Gardens, exhibited a living specimen of the Common Elephant-

Shrew (Macroscelides proboscideus) (text-fig. 16), which had been

captured at Tuin in Bushmanland and presented to the Society

by Capt. H. A. P. Littledale, who sent at the same time a

second example of the same species, an example of the Rock

Elephant-Shrew (Hlephantulus rupestris) (text-fig. 17), also from

MR. R. I, POCOCK ON ELEPHANT-SHREWS. 143

Bushmanland, and a number of small rodents, most of which

belonged to species new to the Society’s list. After drawing

attention to certain interesting morphological points connected

with the group of Insectivora to which Elephant-Shrews belong,

and remarking that the Society had never previously exhibited

Text-fig. 16.

Common Elephant-Shrew (Macroseelides proboscideus).

Text-fig. 17.

Rock Elephant-Shrew (ZJephantulus rupestris).

specimens of the family Macroscelididee, Ma. Pocock pointed out

that in their general appearance and behaviour these animals

much more closely resembled Mouse-like rodents than Shrews,

on account of the length and texture of the fur, the prominence

and size of their staring eyes and of the pinna of the ear, the

144 MR. GEOFFREY SMITH ON THE

only feature that recalled the Shrew being the long, cylindrical,

twitching snout. They were also quite like rodents in their

quadrupedal gait, the ground being traversed by the ordinary

running action or by lightning-like leaps from point to point.

They did not raise their fore-quarters from the ground more

frequently than is the custom with typical rodents, and were

never seen to hop on the hind legs alone, like Jerboas and

Kangaroos, as they have been declared to do in some natural

histories.

In appearance the two examples of MJacroscelides proboscidews

differed considerably from the one representative of Hlephantulus

rupestris. In the former the eyes were smaller and the ears

more widely separated and more concealed in the hair of the

sides of the head. In Elephantulus rupestris there was a

conspicuous light ring round the large eye, the ears were

more erect, and separated by a much narrower space on the top

of the head. It was noticeable, too, that, whereas the Rock

Elephant-Shrew lay hidden beneath a heap of hay during the

daytime, the two Common Hlephant-Shrews preferred to huddle

together in the open part of the cage, evincing a dislike to push

beneath the hay and refusing to remain under it when it was

placed over them. The differences between the living animals,

indeed, quite bore out the view, based upon the structure of the

skull, that the two species belonged to different genera.

PAPERS.

10. The Freshwater Crayfishes of Australia.

By Grorrrey Sura, M.A., Fellow of New College, Oxford”.

[Received October 20, 1911: Read November 21, 1911. ]

(Plates XIV.-XXVII.7 and Text-figure 18.)

1. Introduction.

The study of Freshwater Crayfishes has been distinguished

by the labours of Huxley; the detailed work of Ortman and

Faxon has made us acquainted with the North-American species

of Astacus and Cambarus, and Faxon has reduced the South-

American genus Parastacus and the New Zealand Paranephrops to

order, but what Huxley wrote in 1879 concerning the Australian

Orayfishes, ‘‘ that their nomenclature requires thorough revision,”

is almost as true to-day as thirty years ago.

The following memoir does not pretend to be an exhaustive

monograph of the anatomy or of the systematic classification of

the Australian Crayfishes, but by publishing the series of accurate

* Communicated by the Secretary.

+ For explanation of the Plates see pp. 170-171.

PeeZeon lO laneee |

ASTACOPSIS FRANKLINII. dy,

ALV.

Be 4 ole el ey,

ASTACOPSIS FRANKLINII. a

PaeG.o. LOA, el

ASIACOPSIS SERRATUS: 2

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ASTACOPSIS SERRATUS. o:

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ASTACOPSIS SERRATUS. J

PeZe on 1902, le) OMe:

ASTACOPSIS KERSHAWI. oe

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ASTACOPSIS KERSHAWI. 6.

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PARACHA-ROPS BICARINATUS, J.

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CHASROPS TENUIMANUS. J.

Pea so. 192, Pio

ty,

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7 4

CHAZROPS QUINQUECARINATUS. Jf a1

ie SS MICH VEEN I Olas ae SOIVNIGVOINdVNO SdOMWHO T

Ow Ne ClGLe S90: ‘el

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West, Newman lith.

ASTACOPSIS FRANKLINIL.

bisa Spegil: Zen anege, O80

17 ff

23 Bs,

7-44 ASTAGOPSIS FRANKLINII. 12-14. A. SERRATUS.

45-24. PARACHARAPS BICARINATUS.

West,Newman lith.

34,

as West, Newman lith.

25- 29. CHAARAPS QUINQUECARINATUS. 80.C. TENUIMANUS.

' 381-33 C. QUADRICARINATUS. 34. C. INTERMEDIUS.

es *

Pas. 10a. Peel, -

FRESHWATER CRAYFISHES OF AUSTRALIA. L435

and beautiful drawings made by Mr. W. G. Browning under my

direction, illustrating the forms and chief varieties of all the

known species, by arranging these species in systematic genera

according to their structural affinities, by recording, as far as is

known, their geographical distribution, and by indicating the

light which they throw upon the geographical problems of

Australia, it is hoped that a secure foundation may be laid

on which local naturalists can build. In this memoir, only

the Freshwater forms are dealt with, comprising the genera

Astacopsis, Vheraps, and Paracheraps, the treatment of the land-

Crayfishes of Australia included in the genus Hngeus being

postponed to a future paper, These land-Crayfishes, which are

a highly specialised group, probably derived from the freshwater

form Paracheraps bicarinatus, are confined, in their distribution

to the Victorian Highlands, Gippsland, and Tasmania.

The material on which this} memoir is fouirded, was in part

co.lected by myself in Tasmania and Australia during my visit

to those countries in the years 1907-8, but by far the largest

number of specimens are derived from the very fine collection

belonging to the Melbourne Museum, which Professor Baldwin

Spencer entrusted to me for description, This collection, which

has been gradually accumulated during twenty years by the

efforts of a number of Victorian naturalists, especially Mr.

Kershaw and Mr, Fulton, is particularly representative of

Victorian forms, though specimens from other States are not

altogether absent. 1 have been able to supplement this

Pallection by specimens from New South Wales, through the

kindness of Professor Haswell, from Western Aceiralin. throug]

7 Woodward of Perth, and from New Guinea, througk

. Gestro of Genoa.

sab appears probable that as the population of Australia

increases, the Freshwater Crayfishes might assume as much

importance, as an article of food, as the lobster in America or

England. There is no true lobster in the Australian seas ; its

place is taken by a form allied to our Rock Lobster (Palinurus),

which is altogether inferior as an article of diet. In the first

place, it possesses no large claws and the flesh is altogether coarser

than in the lobster. The Australian Freshwater Crayfishes, on

the other hand, are, on the whole, superior in flavour and texture

to our marine lobsters. Several of the forms, notably Astacopsis

serratus of the Murray River and A. franklinii of the North-Tas-

manian streams, attain a very large size indeed (Pls. XIV., XVI.),

the body reaching a foot in length and the whole animal weighing

as much as eight or nine pounds, thus rivalling a fine lobster.

These animals not very frequently reach the Melbourne market,

and the Tasmanian Crayfish, which is the finest of all, is, I

believe, only occasionally caught and eaten by settlers in the

bush or a few prospectors. The labour of obtaining these large

Crayfishes, for they are not very abundant, has hitherto prevented

their becoming an article of commerce. It would probably be

Proc, Zoou. Soc.—1912, No. X. 10

‘6 MR, GEOFFREY SMITH ON THE

practicable to farm these animals artificially in ponds supplied with

running water, such as are used at trout-hatcheries. In the

case of the marine lobster this is impossible, since the young

hatch out as larvee which at first live a pelagie existence ; but this

difficulty is absent from the culture of the Freshwater Crayfish,

since the young leave the egg in a condition similar to the adult

and pass a certain amount: BE time after hatching attached to

the swimmerets under the tail of the parent. Much work would

have to be done upon the habits, food, rate of growth, ete., before

the undertaking could be placed upon a practical footing, butt the

enquiry would he well worth undertaking. It may be suggested

that the best form for experimenting with would be cen large

Tasmanian Crayfish (Astacopsis Jranklini), which occurs in the

neighbourhood of Launceston, and in this town there is already

a Government trout- hatchery which has been very successful.

One further suggestion may be made: the Freshwater Crayfish

industry im Germany, Russia, France, and England has from time

to time been seriously damaged and in places extinguished by

the occurrence of a plague, probably of bacterial nature ; it would

therefore be well to investigate the parasites of the form to be

cultivated and the best means of protecting it against bacterial

diseases.

There is a curious fact connected with the occurrence and

nature of the large Crayfishes of Australia and Tasmania. ‘lhe

genus Astacopsis is represented by three distinct species, each of

whieh oceurs under two forms, a large and a small, which appear

to oceupy different areas of distribution.

Thus in Tasmania there isthe large 4. franklini (Pl. XTV.), which

is found only in the streams on the north and north-west. coast,

near their entrance to the sea, and in these situations it appears

impossible to obtain small specimens. The small Tasmanian

Crayfish (Pl. XV.) (var: A. tasmanicus), which cannot be sepa-

rated from the large form by any important characters, save

that of size, occurs in the mountain streams of the ceutiliusomn and

central part of the island, where it breeds, but never grows to

more than about five inches in length. Parallel facts occur in

A. serratus (Pl. XVI.), which occurs as a large form in the

Murray and Paramatta Rivers but is represented by smaller, less

spiny forms further inland (Pls. XVIL., XVIIL.). A. hershaawi

las a large form in the Moe River of Gippsland (Pl. XTX.) and

a small form in the Narracan (Pl. XX.) and smaller streams.

The most obvious explanation of this phenomenon is that the

large form, e. g. the large Tasmanian Crayfish, really does breed

in the north coast streams and that its young stages in growth

are passed there and could be found by assiduous search. The

only other alternative is that the young of the large form are

always destroyed near the mouths of the rivers by Blackacn and

other predaceous forms which prey upon them, and that their

numbers are recruited by a few individuals of the small mountain

forms which find their way froin the inland streams to the mouths

PRESIIWATER CRAYFISHES OF AUSTRALIA, 147

of the rivers. ‘The species, in this case, would be always

recruited from the small inland forms. Whether this suggestion

holds good or not, the problem of why small individuals have not

hitherto been found in the streams where the large Tasmanian

and Murray River Crayfishes occur requires settling one way or

the other.

Since this question at present remains open and J have found

it impossible to discover morphological points of any value between

the large and sinall forms of these species, | have not treated

them as distinct species. This leads to the inclusion under one

specific name of the very widely distributed form A. serratus,

which occurs in the Murray River and its tributaries, in the

Victorian rivers such as the Yarra, Plenty, and Bunyip Rivers,

again in the Paramatta River at Sy dney and in the Blue Moun-

tains, This is a truly immense range for a species such as this

to occupy, which, so far as we know, never forsakes the water for

any length of time and has been presumably slowly distributed

by the “alow alteration and communication of different river-

systems. Many of these widely separated communities of Cray-

fishes cannot have interbred for centuries, almost for geological

periods, and yet they have retained the common specific characters

with remarkable constancy. It is true that an immense range of

variation in size and in the degree to which the spines are

developed is met with in these local groups, but these characters

show graduations, and the development of the spines is so variable,

being aien asymmetrical on the two sides of one specimen, that

they “do not offer constant specific characters for separating the

various groups. It appears to me certain that it is impossible to

separate the large Paramatta Crayfish from the large Murray

River form, and ‘yet these two large rivers belong to two different

systems, their nearest point of approach being in the Blue Moun-

tains on different sides of the watershed. With the material at

my disposal [am unable to split up these various local specimens

of A. serratus into a number of subspecies, although I do not

wish to prejudice the work of some future investigator who, with

more material at command, may feel himself competent to do so.

Il. The Relationships of the Australian Crayfishes

and their Geographical Distribution.

The Crayfishes of Australia, Tasmania, and New Guinea belong,

in common with those of New Zealand, South America, and Mada-

gascar, to the family Parastacide, which differs from the Crayfish

family of the Northern Hemisphere, the Astacide, ina number of

important characters which were first pointed out, clearly by

Huxley. The diagnostic features of the Parastacide given by

Huxley (P. Z. 8. 1878, p. 775) are as follows :

“ The podobranchize are devoid of more ‘ieee a rudiment of a

lamina, though the stem may be alate. The podobranchia of the

first maxillipede has the form of an epipodite; but in almost

10*

148 MR. GEOFFREY SMITH ON THE

all cases it bears a certain number of well-developed branchial

filaments.

“The first abdominal somite possesses no appendage in either

sex ; and the appendages of the four following somites are large.

The telson is never completely divided by a transverse suture.

“‘More or fewer of the branchial filaments are terminated by

short hooked spines; and the coxopoditic sete, as well as those

which beset the stems of the podobranchie, have hooked apices.”

The Astacide possess the converse of these diagnostic

characters.

The Astacidee and Parastacide, the one family occurring in

the Northern Hemisphere, the other in the Southern, are there-

fore separated by important characters, and it is very prebable

that they have been independently evolved from marine lobster:

like ancestors which already differed in these characters before

they took to a freshwater life.

The occurrence of Parastacide in Australia. New Zealand, and

South America, with an aberrant genus (Astacoides) in Mada-

gascar—that is to say, in countries which are now separated

by wide stretches of ocean—is a striking fact in geographical

distribution, but it does not stand alone, the distribution of

many freshwater fish, crustacea, molluscs, etc., having a similar

character in the Southern Hemisphere.

These facts, taken in conjunction with geological evidence, have

led many naturalists to assume a much greater extension of the

Antarctic Continent in past times which is supposed to have been

connected with South America, Australia, and New Zealand, and

possibly at a very remote period with Madagascar, thus permitting

the migration of land and freshwater animals to and from those

countries. In the case of the Parastacide the only alternative

theory is that the South-American, Australian, and New Zealand

genera have been independently derived from some common

marine ancestor.

Our concern here, however, is not so much with the origin of

the Parastacidz in the remote past, but with the inter-relation-

ships of the Australasian genera and their probable evolution and

migrations.

We can distinguish four genera of Australasian Crayfishes—

Astacopsis, Paracheraps, Cheraps, and Kngeus (the last-named

genus we will consider as a single entity, though it may be found

convenient hereafter to split it up into several subgenera).

The members of the genus AsTacopsis are characterised by the

development of spines or tubercles upon the body and limbs, and

by certain features in the gills and appendages which are fully

set forth in the diagnosis of the genus on p. 154. They inhabit

swift-moving streams and rivers; they are not found in ponds and

water-holes, and they are not known to forsake the water for any

period of time. Their distribution is as follows :—-In Tasmania

there occurs 4. franklinii (Pls. XIV., XV.), the largest Crayfish

in the world. It is confined to the rivers and streams upon the

FRESHWATER CRAYFISHES OF AUSTRALIA. 149

north and west coasts. In the highland streams of the south

and centre a small form occurs (var. tasmanicus) which may or

may not be reckoned as a separate species (see p. 156). A. ser-

ratus (Pls. XVI.-X VIII.) occurs as a large form in the Murray

River and in the Paramatta River at Sydney. Asa small or

medium-sized form it occupies the Victorian Highlands, and as a

small form it also occurs in the Blue Mountain streams. 4. ker-

shawi (Pls. XTX., XX.), which is, in many respects, intermediate

between the above two species, lives in a few Gippsland rivers, thus

occupying also an intermediate geographical station. The genus

Astacopis therefore runs from the mouth of the Murray River

southwards to Gippsland and Tasmania and up the east coast to

Sydney. Its centre of distribution would appear to be the Murray

River and its tributaries, and it is confined to the south-eastern

corner of Australia and to Tasmania, where the climate is tem-

perate and clear swift-running streams are abundant.

Text-fig. 18.

Sketch Map

AUSTRALIA

Map of Australia show ng areas of distribution of Crayfishes.

The genus Cu#rars (see p. 165 and Pls. XXIIXXIV.),

although departing in many important features from A stacopsis,

yet shows a trace of agreement with this genus, especially in the

150 MR. GEOFFREY SMITH ON THE

fact that the last posterior arthrobranch is not rudimentary. Its

members inhabit running water, and they are confined in dis-

tribution to the west and north coasts and to New Guinea, being

entirely absent from the southern and eastern districts oceupied by

Astaccpsis. In the south the arid coast-line, fringing the Great

Australian Bight, constitutes a wide and insurpassable barrier

between the genera Astacopsis and Cheraps. On the east coast

Cheraps is absent and Astacopsis does not appear to occur much

north of Sydney.

The species C. intermedius (P]. XXIV. fig. 2) from Western

Australia is of great interest, because it forms a_ perfect

transition to the genus Paracheraps. (C. intermedius retains the

diagnostic features of a Cheraps, but it presents a remarkable

approach in general facies and in a number of points to Para-

cheraps bicarinatus.

The genus ParacH ®raPs, consisting of the single species P. bicar-

inatus (Pl. XX1.), is closely allied in all its features to Cheraps,

and the alliance is made. more obvious by the existence of

the before-mentioned C. intermedius, which may reasonably be

regarded as the ancestral form from which P. bicarinatus has

been derived. /. bicarinatus is the most widely distributed

Crayfish on the Australian continent, being the only form which

penetrates into the interior of the continent. It is, in fact,

universally distributed all over the continent, occurring with

Cheraps in the west and north and with Astacopsis in the south

and east. It does not occur in Tasmania or in New Guinea. The

relationship of this genus and species is obviously with the

western Cheraps, and it is also equally certain that it is a secon-

dary derivation from Cheraps, which has been able to spread

across the deserts eastwards and invade Victoria in the south

by becoming adapted to live under semi-desert conditions. The

alternative supposition, viz. that Paracheraps is the ancestral

form and has given rise to Cheraps, is clearly negatived by the

impossibility of thus accounting for the fact that, although it

ranges as a continuous species all over the continent, it should have

given rise to several species of Cheraps only in the west. It would

be difficult also on this supposition to account for the fact that

Cheraps differs less from Astacopsis than Paracheraps does. In

fact, if we take the now almost universally distributed Para-

cheraps as representing the ancestral form of Australian Cray-

fish, the entire isolation, both structurally and geographically, of

Astacopsis and Cheraps becomes unintelligible.

It is worthy of notice, also, that Paracheraps apparently does

not occur either in New Guinea or Tasmania, and, although stress

cannot be laid on this fact, it is possibly another indication of

the modern origin of Paracheraps, after the separation of New

Guinea and Tasmania from the mainland of Australia.

The genus Exexvs, comprising the land-burrowing Crayfishes,

which have gone a step beyond Paracheraps in their independence

of water, are confined to Victoria, Gippsland, and Tasmania. From

FRESHWATER CRAYFISHES OF AUSTRALIA. 151

their habits and anatomical structure they have probably been

derived directly from Paracheraps, some of the species graduating

very perfectly into the latter genus. The burrowing land-Cray-

fishes are therefore the most modern product of all. Although

it is not my intention to give any description of the genus Engeus

in this paper, certain of their characters may be mentioned

which point to their high degree of specialization. Their whole

appearance and form are very different from the other Crayfishes,

the thorax being extremely deep dorso-ventrally, a character

which is correlated with the very great size of the gastric mill.

The abdomen is much reduced in size. Certain of the species

show a peculiar reduction in the gill-formula, the last pleuro-

branch being entirely absent, while in some species the exopodite

of the maxillipede is completely absent. The body is entirely

free from spines or ridges.

The above survey and an examination of the diagnoses of the

several genera afford a remarkably clear picture of the evolu-

tionary relationships of the genera described, which we may

represent in the form of a table, thus :-—

Chera ps. Engeus, Paracheraps. Astacopsis.

3 i

ad 2 3

a A

Y A

Se 5 ve

ee: ae

Say we

Pale i b wat

ater | i

Common ancestor.

We may attempt to reconstruct also the means by which these

genera have reached their present geographical distribution. The

outstanding fact is clearly the complete isolation in structure and

distribution of the western Cheraps from the eastern Astacopsis.

This sharp distinction between a South-eastern and a North-

western Australian fauna does not apply only to Crayfishes, but

is practically universal, and in looking for a cause the attention

of naturalists has been called to the existence in Central Australia

of Cretaceous deposits, which are held to indicate the existence of

a sea in Cretaceous times dividing Australia into a western and

an eastern island. We may therefore start with the hypothesis

that the separation and divergent evolution of Astacopsis and

Cheraps dates from the separation of Eastern and Western

Australia by the Cretaceous sea. It would appear that, with the

152 MR. GEOFFREY SMITH ON THE

filling in of this sea and the junction of the eastern and western

islands, no feasible passage was formed by means of which

the eastern Astacopsis could invade the territory of the western

Cheraps or vice versa. The nature of this early barrier is un-

certain, but when once the desert condition was established, the

central deserts have consituted a barrier against a direct eastern

or western migration : on the south-west coast the desert and sem1-

tropical region of the Australian Bight prevents a northern passage

of Astacopsis into the territory of Cheraps, while to the north-

east the semi-tropical coastal streams do not appear to have been

colonized either by Cheraps moving southwards or Astacopsis

migrating northwards. -Astacopsis, m fact, remains a typically

temperate genus, Cheeraps a semi-tr opical or tropical one, and the

barrier of temperature seems to determine the distribution of

these two forms with complete rigidity.

An offshoot from Cheraps, however, viz. Paracheraps bicarinatus,

on the filling in of the Cretaceous sea, has gradually extended its

range from the west eastwards, invading the central districts of

Australia and penetrating thence southwards and eastwards so

as to become almost coterminous in distribution. with A stacopsis.

In the very centre of the Astacopsis distribution, viz. in Victoria,

P. bicarinatus is particularly abundant, and here it has given rise

to the burrowing land-Crayfishes, Hngewus, whose headquarters are

in this region. If it is true that P. bicarinatws does not occur in

New Guinea or Tasmania, the view that this form is of compara-

tively recent origin is amply confirmed, the date of its origin and

migration being fixed as subsequent to the separation of New

Guinea and Tasmania from the mainiand in Tertiary times. We

are, however, here confronted with a difficulty, since, although

P. bicarinatus apparently does not occur in Tasmania, one or two

species of Hngeus, which we must hold to be derived from

P. bicarinatus, are common in Tasmania. The habits, however,

of Engeus are peculiar, and it is quite possible that individuals of

this genus have been carried across Bass’s Straits from Gippsland

in floating tree trunks, at the roots of which the ‘“ land-crabs ”

frequently make their burrows.

Without unduly pressing this hypothesis, it nevertheless offers

an explanation of the curious fact of the occurrence of Engcus in

Tasmania and the absence of its parent form, P. bicarinatus from

that island.

The occurrence of 4. kershawit in Gippsland, related by many

of its features, e. g. the absence of spines on the abdomen and the

truncated shape of the rostrum, to dA. franklinw in Tasmania,

cannot be passed over without reference. It is apparently con-

fined to Gippsland, but it is surrounded and in close contact

with the much more widely distributed A. serratus. Its

relationship to the Tasmanian form indicates a close connection

at some not very remote time between Gippsland and Tasmania,

a connection which may be detected im the similar geological and

r FRESHWATER CRAYFISHES OF AUSTRALIA, Los

physioeryg ice! characters ‘of these two countries. It seems

probable that when the two countries were in actual continuity

they together formed a zoological district with many features in

common, distinct from the surrounding countries of Victoria and

New South Wales.

By théeforegoing lines of argument we are able to establish on

a secure basis the general course of evolution and the routes of

dispersal of the Parastacine Crayfishes of Australia. We are

able to establish with certainty that the widely distributed Para-

cheraps bicarinatus is a comparatively recent derivation from

the Western-Australian Chewraps, and that the land-Crayfishes,

Engeus, ave a still more modern derivative from Paracheraps.

We are left, therefore, with Cheraps and Astacopsis as the two

primitive representatives of Australian Crayfishes, which, both

by their complete isolation from one another and by their wide

distribution, betray a great antiquity. The question as to which

of these two primitive genera is the most primitive and represents

to the greatest extent the original ancestor of the group is a very

obscure question. We may, however, make some suggestions for

the solution of this problem.

Since Crayfishes in general are emphatically not tropical forms,

and since the Australian Crayfishes are oniy represented in

Northern Australia by a single species, C. quadricarinatus, it is

certain that this form is only a northern straggler, and that New

Guinea and North Australia are not the centre of distribution of

the group. Granted that the centre of distribution is somewhere

in the south of the continent, have we any light to guide us in

choosing between Oheraps or Astacopsis as occupying most nearly

the original area of distribution? The genus Astacopsis, on the

whole, now lives under conditions more generally characteristic

of Freshwater Crayfishes than Cheraps. It is characteristic of

cooler regions and is particularly abundant in mountain streams of

great rapidity and clearness; while the western Charaps inhabits

rivers, more sluggish and clouded in nature, which rather fitfully

irrigate a parched country. It seems that Cheraps is already in

the grip of those circumstances attendant on a lack of water

which have finally resulted in the production of such specialized

forms as Paracheraps and Engeus. This is no more than a

suggestion, but if it is true we are led to the conclusion that the

more southern Astacopsis, inhabiting the temperate and well-

watered mountainous regions of South and South-eastern Australia,

retain to the greatest extent the original characteristics and

distribution of the ancestral form. If this is so, and if the

Bassian Subregion is really the centre of distribution of the

Australian Parastacide, we may perhaps include these animals in

the array of alpine plants and animals characteristic of this

region, which were probably once distributed across the Antarctic

Continent and reached their present distribution in South

America, South Australia, and New Zealand by this means.

ye! MR. GHOFFREY SMITH ON THE

ITI. Systematic Account of the Genera and Species of

Australian Parastacide.

Genus Asracopsis Huxley (P.Z.8. 1878, p. 764).

Arthrobranch of last leg but one not rudimentary or even much

reduced. Ala of podobranchs small and inconspicuous. Short

and stout hooks only present at the end of a few branchial fila-

ments fringing the stem of podobranchs (Pl. XX VI. fig. 10),

being absent in all other qills.

Hooked set on coxopodites and on gill-stems not very sharply

recurved (Pl. XX VI. fig. 11).

Mandibles with three prominent teeth and smooth lower

midce (EIS XeXGVis fie .16)).

First maxilla with endopodite consisting of base and distinct

flagellum united on to it (Pl. XXV. fig. 5; Pl. XXVI

fig. 9).

Second ,maxillipede with penultimate segment bearing an

upper lobe, which projects forwards as far as terminal segment

(EIPEGaVE fet):

Third maxillipede with serial row of bristles on outer face of

third and fourth segments, and a sparse fringe of bristles on

inner face. Exopodite does not equal third segment in length

GEE XG erie: 1):

Great chela with the carpus rather short and stout; much

tubereulated, with a row of tubercles on its outer border as well

as on the inner (Pls. X1LV.—XX.).

Succeeding legs with terminal and subterminal joints well

provided with pencils of bristles. Opening of vas deferens

situated on a short simple papilla.

The lateral keels of rostrum are spiny or tuberculated. Lateral

earina on carapace is also spiny or tuberculated. Carapace and

branchiostegites tuberculated or spiny. The sternal keel is

depressed and blunt (PR 2GQV Tefigal4)

The first abdominal segment carries lateral spines; the suc-

ceeding segments are spiny, tuberculated, or setose.

The membranous portion of telson and uropods is short

compared to the upper calcified portion, and the median spine on

the endopodites of the uropods is, in consequence, situated

distally.

ASTACOPSIS FRANKLINI. (Pls. XIV., XXV.,& XXVL., figs. 7-11.)

The Large Tasmanian Crayfish.

(Gray, Eyre’s Journals of Expeditions of Discovery into Central

Australia, i. p. 409, 1845.)

The rostrum is rather broad and tumid, and ends in a short

median spine; the lateral keels are rounded, and each keel

carries four or five blunt spines or tubercles.

FRESUWATER CRAYFISHES OF AUSTRALIA. Mays

On the carapace immediately lateral to the posterior outer

border of the rostral carina, a tubercle, often double, is present,

and posteriorly in a line with this tubercle is another rounded

tubercle, and behind this again a rounded boss. The disposition

of these tubercles is subject to some variation. Laterally

and anteriorly, the cephalothorax has a few blunt spines and

tubercles, and there are numerous blunt tubercles on the

branchiostegites.

Ventrally the interantennal spine is rather long and narrow,

and not produced laterally at the base. The chitinous margin

of upper lip has large Jateral arches and a small median arch

(PI. XX VI. fig. 7).

The inner border of the antennal scale widens rather suddenly

at the base of the terminal spine (Pl. XX VI. fig. 8).

The sternal keel is of the usual Astacopsis type, but is sharper

than in succeeding species.

The first abdominal segment bears laterally a prominent

forwardly directed spine.

The second abdominal segment has a varying number (4-7)

of small sharp spines on each lateral border.

The succeeding abdominal segments are smooth, but laterally

each segment is produced into a small sharp spine, often with

another: smaller spine just above it.

The exopodite of the uropod has a row of teeth between

the hard and membranous portions, which graduates: internally

into a row of bristles. The endopodite has a small spine close to

the distal border in the middle line.

The telson has two lateral serrations, but is otherwise smooth.

The great chela has the pincer studded with tubercles and a few

brushes of sete. ‘Two tubercles in the pincer, one on the upper

or outer and one on the inner pincer, are greatly enlarged,

especially in the larger specimens. The inner edge of the carpo-

podite has three or, at most, four serrations. The surface of the

carpopodite is not greatly pitted.

The two anterior walking legs have only a few spines, but

numerous brushes of bristles. The two posterior legs have more

numerous spines, which are not very long or conspicuous.

The colour is uniform dark green,

The largest specimen obtained by me weighed just under

eight pounds, and measured 16 inches from rostrum to telson.

Small specimens less than 8 inches are very rarely obtained, and

have not been seen by me.

Localities.—Several specimens were taken by me from the

Brid and Muddy Creek, Bridport, Tasmania—the latter stream

being a very small muddy rivulet. They are also reported from

numerous streams and rivers along the north coast, and from the

Gordon River on the west coast, but they are unknown from the

south, east, or central districts of Tasmania,

156 MR. GEOFFREY SMITH ON THE

The Small Tasmanian Crayfish, Var. vasMANIcUS. (PI. XY.)

(Erichson, Archiv fiir Naturgeschichte, 12 Jahrg. p. 94.)

This species (¢), which occurs on Mount Wellington and

the southern and central ranges of Tasmania, reproduces on a

small scale almost all the characters of the large northern

and western 4. franklinii, so that it may perhaps be considered

as only a small variety of the large form. The following

characters, which may, however, be merely differences due to

growth, may be referred to. There is a great reduction of the

tubercles upon the carapace and branchiostegites. The upper

margin of the carpopodite of the great chela has about six

serrations ; there is no pronounced enlams ement of two tubercles

in the pincer. ‘The inner border of the meropodite has about six

spines. The surface of the carpopodite is greatly pitted. The

spines on the walking-legs are reduced to very minute points,

The colour is green.

The length of fully adult specimens is never more than

5 inches.

Localitves.—Streams on Mount Wellington, Tasmania (1907).

A small specimen with the tubercles more prominent than

usual, measuring 100 mm., from Traveller River, Lake St. Clair,

Tasmania (1893). Two small specimens from Zeehan, West

Coast of Tasmania.

Remarks on the above two species.—There are, in reality, no

distinctive characters, beyond those due to size, by which the

above two forms can be separated. The fact, however, that the

small and large forms occupy different stations in different parts

of the island appears to preclude the possibility of the small

specimens being merely not fully-grown individuals of the large

form. A most curious fact is that I was unable to obtain

the young stages in growth of the large A. franklinii in the

rivers and streams, where the large individuals were fairly

common. The localities from which the large 4 .franklinii have been

obtained are invariably in the mouths of the streams near the

sea, and in these localities it is impossible, apparently, to obtain

small specimens. The small A. tasmanieus, on the other hand, is

always found inland in mountain-streams, often near their sources.

It is therefore within the bounds of possibility that the large

A. franklinii represents those individuals of the small A. tasmanicus

which have succeeded in reaching the mouths of the rivers, but

that the young which they produce at these situations do not

arrive at maturity, owing to the presence of predatory fish,

e.g. Gadopsis marmoratus. The species would then be entirely

Teer ‘uited from the young of the small form, which breeds up in

the mountains. Exactly parallel facts are met with in the other

species of Astacopsis now to be described. A careful examination

of this question should be undertaken by a local naturalist.

FRESHWATER CRAYFISHES OF AUSTRALIA, LHF

ASTACOPSIS SERRATUS. (Pls. XVI.-XVIIL. & sacs ae 12-14.)

The Murray River Cray fish.

(Shaw, Zoology of New Holland, vol. i. p. 407, pl. iii., 1843.)

Nomenclature.

Names given to Murray River form :

Cancer serratus Shaw, loc. cit.

Gray, Eyre’s Journals of Expeditions of Discovery into

Central Australia, vol. i. p. 409, 1845.

Astacus armatus Von Martens, Ann, Mag. Nat. Hist. ser. 3,

vol. xvii. p. 359, 1866.

Astacoides spinifer Heller, Novara Reise, Crustaceen, p. 102,

Taf, ix., 1865:

Astacoides serratus McCoy, Prodromus of the Zoology of

Victoria, vol. 1. pl. 15, 1885.

Names given to Paramatta River and other forms :

Astacopsis spinifer Spence - Bate, ‘Challenger’ Reports,

vol. xxiv. pl. xxvill., 1888. (Large Paramatta form.)

Astacopsis nobilis Dana, United States Exploring Expedition

—Crustacea, Part I. p. 526, pl. xxxiii., 1852. (Locality

given as New South Wales. A not very clear figure

of a 5-inch specimen.)

Astacus australiensis Milne-Kdwards, Hist. Nat. Crust. vol. ii.

p. 332, pl. xxiv. (Small Paratnatts type.)

Astacopsis paramattensis Spence-Bate, loc. cit. p. 202, pl. xxiii.

(Small Paramatta specimen.)

A. sydneyensis Spence-Bate, loc. cit. p. 204, pl. xxill. (Very

smal] Paramatta specimen. )

Astacoides plebeius Hess, Archiv fiir Naturg. xxxi. p. 164,

Taf. 7, 1865. (From Sydney. Small Paramatta (?)

form.)

Rostrum elongated ; spines on rostral keel sharp and long:

posterior border of keel tumid.

A sharp spine is present on carapace just posterior and

lateral to the end of rostral keel, and another sharp spine is present

posterior to this and confluent with a rounded boss.

There are a few blunt tubercles and two or three sharp spines

on the antero-lateral part of the carapace, and the branchiostegites

are not only tuberculated, but carry a superior row of very large

spines.

The interantennal spine is rather short and broad, with pro-

duced lateral angles at the base (Pl. XX VI. fig. 12).

The chitinous margin of the upper lip has small lateral arches

and a large concave median arch (fig. 12).

The inner border of the antennal scale does not suddenly

expand at the base of the terminal spine (fig. 13).

The sternal keel is depressed and blunt (fig. 14).

The first abdominal segment has two very large spines on each

dorso-lateral margin.

158 MR. GEOFFREY SMITH ON THE

The second abdominal segment has two (sometimes three)

smaller lateral and two large dorso-lateral spines on each side.

The next three abdominal segments have two large dorso-

lateral spines, and one (sometimes two) small lateral spine on

each side.

The sixth abdominal segment carries two small lateral spines

or tubercles on each side.

The telson has two (sometimes more, a variable number)

median spines.

The number and arrangement of the above spines is sometimes

variable and even asymmetr ical.

The great chela has two enlar ged tubercles in the pincers, both

on the ‘biting- edge of carpopodite. The lower external border of

the carpopodite has a row of sharp and large spines continued right

down to the posterior border. The upper internal border has

about five marked serrations. The carpopodite is flattened,

and does not carry any tubercles on its other surfaces. The meso-

podite has three large spines on its inner border and two on its

outer.

The first two walking-legs are nearly free from spines, but the

last two carry long and prominent spines, especially on the last

joint but one.

Colour (as given by McCoy).—The anterior legs, the middle

of the back, and the apices of the spines and tubercles rich

creamy white or ivory-coloured ; the ground-colour of the other

legs, the sides of the carapace, and the abdomen pale Prussian-

blue of varying intensity in different individuals, or sometimes

mottled with dull olive-green. Semicorneous flexible edges

of tail-fin brownish.

Largest specimen obtained measured 10 inches (about 230

mm.); the smallest 7 inches.

Locality of type specimens.— Murray River.

Local Varieties.

From VICTORIA.

(«) Bunyip River... One specimen, January 1880, measuring

140mm. It resembles the Murray River form, except that the

spines on carapace and branchiostegites tend to be replaced

hy blunter tubercles, also the dorsal spines on abdominal segments.

On second abdominal segment there are four lateral spines on

left side, three on the right, thus illustrating the variability and

fr equent asymmetry of these spines.

(6) Yarra River. Several specimens, 1905. A large specimen

(Pl. XVII.), measuring 150 min. , has a single spine on each side

of first abdominal seoment, three lateral spines on second, and

the dorsal spines on abdominal segments | clearly marked but

reduced, The smaller specimens, about 100 mm. in Jength, have,

in some cases, a single spine on first abdominal segment, in

others two; there is a great reduction of dorsal spines on the

FRESHWATER CRAYFISHES OF AUSTRALIA. 159

abdominal segments. ‘The Yarra River Crayfish las been

deseribed and figured by MeCoy (‘ Prodromus of Zoology of

Victoria,’ vol. u. pl. 160) as Astacopsis serratus var, yarraensis,

and he refers to its brilliant blue colour, but he otherwise regards

it as merely a variety of A. serratus from the Murray Bevery

specimens (1897), one measuring 135, the other 100 mm. Both

very slinilar to above. There are four spines on second abdo-

minal segment in one specimen, three in the other. The dorsal

abdominal spines are well marked, but reduced.

(d) Plenty River. One specimen (1896), measuring 160 mm.,

has two spines on right side of first abdominal segment, one on

the left. ‘There are four spines on right side of second abdominal

segment, three on the left. The dorsal abdominal spines and

those on carapace are rather reduced,

From New Sourn WALEs.

(e) Paramatta River, Sydney. The Large Paramatta River

( ray fish has been figured by Spence-Bate (‘ Challenger’ Reports,

Vol, sxive ps 194, pl. xxviii.) under Heller’s name of A stacopsis

spinifer. He regards it, in common with McCoy and others, as

identical with ihe Murray River form. I have not seen the

large Paramatta Crayfish, but from the excellent figure given by

Spence- Bate 1 cannot detect any difference between it ‘and the

Murray River form.

(f) Paramatta River. The Small Paramatta Crayfish

described and figured by Spence-Bate (doc. cit.) as Astacopsis

paramattensis. It is about 100 mm. long, and is distinguished

from the large form (e) by complete reduction of spines on cara-

pace and abdomen to similarly situated small tubercles. It is,

however, very probable that this specimen, coming as it does from

the same river as the large form, only represents a stage

in growth of the large Paramatta Crayfish. Dana’s A. nobilis

and Milne-Edwards’s 4. australiensis both probably belong here.

A very small specimen (about 50 mm.), described and figured

by Spence-Bate (loc. cit.) as A. sydneyensis, is also probably only a

very young Paramatta River form.

(g) The Blue-Mountain Crayfish (P\. XVIII.), found in small

streams in the Blue Mountains. The largest specimen obtained

by me measured 110 mm. It resembles in every particular

Spence-Bate’s small Paramatta Crayfish (/). The spines on the

carapace, branchiostegites, and abdomen are reduced to small

blunt tubercles. This is particularly clear if we compare a

specimen of this Crayfish with a similarly sized Victorian

specimen, é. g., from the Yarra River. In the latter the dorso-

lateral and lateral abdominal spines are far better developed

than in the Blue-Mountain or small Paramatta forms. he

practical identity of the Blue-Mountain Crayfish with the small

Paramatta form deseribed by Spence-Bate seems to me to

make it undesirable to invent another name for it,

160 MR. GEOFFREY SMITH ON THE

Note,—Nobili, in his paper in the ‘ Annali del Museo Civico di

Genova,’ vol. xl. p. 244, 1901, describes a specimen of Astacopsis

identical with Milne-Edwards’s 4. australiensis, as coming from

Sorong, New Guinea. The occurrence of an Astacopsis in New

Guinea is so utterly at variance with all the known facts of

the distribution of Australian Crayfishes that I am unable to

accept the locality of this single specimen as correct, especially as

all the subsequent expeditions to New Guinea have failed to find

any Astacopsis there, although Cheraps in abundance have been

obtained. Dr. Calman has suggested to me that Sorong was

probably wrongly read for Sydney, or else that the label for this

specimen had been somehow transposed, Until further evidence

is forthcoming as to the occurrence of an Astucopsis in New

Guinea, the locality given for this single specimen, which is

identical with the small Paramatta form of -4. serratus, must be

received with the greatest scepticism.

Remarks on the above species.—F¥or including all the above forms

under one species, A. serratus, ranging from the Murray River

to Sydney, I shall be blamed by many systematists, but the

problem is one of peculiar difficulty. The Freshwater Cray-

fishes, like so many of the large Decapods, begin breeding long

before they have attained their limit of size; we are therefore

often puzzled to know whether a particular set of specimens

represents a separate species, er only not fully-grown individuals

of a species which progressively alters as it grows older. It

seems undoubted that the large Murray River and the Para-

matta River Crayfishes are identical. It is true that the

geographical separation of the Murray and Paramatta is not so

vreat as it looks, as the Lachlan River, a tributary of the Murray,

vises in the Blue Mountains on the other side of the watershed

to that on which the Paramatta rises. The Yarra, Bunyip,

and other rivers of the Victorian Highlands are similarly

divided from the Murray tributaries, and here, although the

differenees are very slight, it may be possible to separate a true

variety of Crayfish inhabiting these rivers from the Murray

River form. The small Blue-Mountain Crayfish bears much the

same relation to the big Paramatta form as the small Tas-

manian Crayfish to the big one.

Unlike the genera Cheraps and Hngeus, they are not known to

leave the water and migrate across the land, so that the various

waces of this species must have been isolated from one another

for very long periods.

ASTACOPSIS KERSHAWL, sp, n. (Pls. XIX., XX.)

The Large Gippsland Crqytish. (PE exexXe)

The rostrum is broader than in A. serratus, with blinter

tuberculated spines on its keels, thus approaching 4. franklinit.

The spimes on the carapace are replaced by blunt, rounded

“tubercles and ridges. Similarly, the sharp spines present on the

FRESHWATER CRAYFISHES OF AUSTRALIA, 161

branchiostegites of A. serratus are replaced by blunt, often tumid

tubercles.

The interantennal spine, the margins of the upper lip, and the

antennal scales approach the condition found in A. franklinii,

and the sternal keel is sharper than in 4. serratus,

The first abdominal segment has only one lateral spjne (jn twa

out of the three specimens this spine was absent on ane side).

The second abdominal segment has three lateral spines on each

side and one dorso-lateral; the dorsal spine present in 4, serratus

is replaced by a very large tumid tubercle which is joined hy a

ridge to its fellow of the opposite side,

The succeeding three segments haye qne tq thpee lateral spines

on each side and a dorso-lateral, the dorsal spine of A. serratus

being replaced by a tumid tubercle, The sixth abdqminal segment,

telson, and uropods are provided with several small spines. The

chela is less elongated and more stoytly built than jn 4, serratus,

and very often carries spines or tubercles on the dactylopodite

and on the surfaces of the carpopodite which are nqn-tuberculous

in A. serratus. .

The largest specimen was 10 inches long.

Locality.—Moe River, Gippsland. Three specjmens egllected

by Mr. Kershaw,

Local Variety,—The Small Gippsland Crayfish. (Pl. XX.)

Several rather small specimens from the Narracgn River and a

number of other small Gippsland streams (largest specimen

5 inches in length) agree with A. kershawi in the broad truncated

form of the rostrum, sharpness of sternal keel, and replacement

of spines by blunted tubercles, thus differing in diagnostic

characters from the smaller specjmens of A, serratus from the

Victorian Highlands. Exeept in point of gize, it is impossible tq

separate these specimens from the large A, kershawi, of which

they are probably only the not fully grown individuals.

Remarls an the abave species,—There can be no dqubt that

this Gippsland Crayfish js specifically distincf from A. serratus,

It is of considerable interest to observe that if approaches in

many respects the Tasmanian Astacopsis with which it is

geographically related, being separated by the comparatively

modern Bass’s Straits,

Genus PARACHARAPS, gen, nov,

Arthrobranch of last leg but one rudimentary, consisting of a

stout fleshy pedungle tipped with a few minute filaments which

end in long curved hooks (Pl. XX VI, figs. 16, 18). Ala on

anterior podobranchs js broad and conspicuous, and carries small

filaments. The majority of gijll-filaments attached to the stems of

all the gills are furnished with long recurved hooks at their ends,

The other filaments are frequently pointed at the end,

The hooked sete on coxopodites and on gill-stems are very

sharply recurved (fig. 17),

Proc, Zoou. Soc.—1912, No, XI. 1}

162 MR. GEOFFREY SMITH ON THE

The antennal scale is broad, with the inner margin widening

suddenly after the terminal spine (fig. 20).

Mandibles with two prominent teeth and a serrated lower

edge (fig. 24).

First maxilla. with endopodite consisting of a basal segment

and a minute terminal segment (figs. 15, 23),

Second maxillipede with penultimate segment bearing an upper

lobe which does not project forwards nearly so far as the tip of

the terminal segment (fig. 22).

Third maxillipede without a serial row of bristles, but with the

face of the third and fourth segments provided with a double

fringe of slender crowded hairs. The exopodite exceeds the third

sesment of endopodite in length (fig. 21),

‘Great chela with the carpus long and stout, especially in the

male; on the underside a longitudinal ridge is present, often

with slight excavations on either side of it, The surface is

smooth ; “there is no row of tubercles on the outer border, but

there is a conspicuous fringe of downy hairs on the inner border.

Succeeding legs with terminal and subterminal joints smoother,

with very few bristles.

Opening of vas deferens situated on a long projecting and

complicated papilla.

The keel of the rostrum is flat and smooth; the lateral carina

on carapace also continuous and smooth; there is no median

carina.

The distance between the tip of the rostrum and the cervical

suture is shorter than between the cervical suture and posterior

border of carapace.

The carapace is smooth; the abdominal segments are smooth

and non-setose.

The sternal keel is fairly prominent and sharp.

The interantennal spine is rounded, and does not end in a

sharp point (fig. 19). :

The membranous portion of telson ‘and uropods is long

compared to the upper calcified portion, and the median spine on

the endopodites of the uropods is in consequence situated mesially.

Remarks on the above genus.—In creating the above genus for

the reception of the single species, P. bicarinatus, I have been

influenced by the following facts. Although P. bicarinatus does

not differ in any very striking characters from the species which

[ retain in Erichson’s genus Cheraps, yet if it were included in

this genus it would occupy an altogether isolated position, both

as regards structure, geographical distribution, and the fact that

a it has probably been derived the large and varied genus

Engeus. For convenience’ sake, therefore, and for the purpose of

bringing out its peculiar importance in the evolutionary history

of the Australian Crayfishes, I have after much hesitation

decided to separate it off from the related forms retained in the

genus Cheraps. Since P. bicarinatus is one of the oldest-known

species from Australia, T had originally intended keeping the

FRESHWATER CRAYFISHES OF AUSTRALIA. 163

name Cheraps for it, and placing the Western-Australian species

in a new genus, Paracheraps. But, as M. Roux, of the Basle

Museum, pointed out to me, the term Cheraps was first used by

Erichson (Arch. f. Naturg. xii. 1846, p. 101) for the species

C. preissii, which, whatever it may be, does not seem to be

identical with P. bicarinatus. From Erichson’s diagnosis of the

genus Cheraps and of the single species C. preissii, which he

includes in it, [ find it quite impossible to discover what species

of Australian Crayfish he was dealing with. No figuresare given,

and the only characters mentioned which are of the slightest

diagnostic value are the facts that the animal came from Western

Australia, that the tail-fan was in part membranaceous, and that

the antennal scale was ‘‘ egg-shaped and pointed.” No mention

whatever is made of any keels on the carapace, a point noticed by

Haswell in his ‘ Catalogue of the Stalk- and Sessile-eyed Crustacea

of the Australian Museum,’ who gives as a character of C. preissii

“the absence of keels on carapace (?).”

The balance of evidence seems favourable to the idea that

Krichson’s really worthless description of C. preissii does refer to

a species of Cheraps and not to Paracheraps bicarinatus, so

that the term Cheraps must be kept for the Western-Australian

species.

PARACHAZRAPS BICARINATUS Gray. (Pls. XXII. & XXVI.

figs. 15-24.)

The Yabber.

(Eyre’s Journals of Expeditions of Discovery inta Central

Australia, vol. i. 1845.)

Nomenclature :—

Astacus bicarinatus Gray, loc. cit. p. 410, pl. iii. fig. 2.

Cheraps bicarinatus Von Martens, Monatsbericht Akad.

Wiss. Berlin, 1868, p. 617.

Astacopsis bicarinatus Haswell, Australian Museum Cata-

logues, v. Crustacea.

Astacoides bicarinatus McCoy, Prodromus of the Zoology of

Victoria, vol. i. pl. 29, 1885. «

The rostrum is without pronounced keels; it terminates in a

spine, just below which two small lateral spmes indicate the

beginning of the much-reduced keel.

The lateral carina on the carapace is a coniimuous blunt ridge.

There are no other tubercles or spines on the carapace or body

and there are no hairs except at the lateral borders of the

carapace and abdominal segments and on the limbs. * The surface

of the carapace is, however, usually pitted superficially.

The telson is broadly ovate, and there are two small spines

laterally at the junction of the hard and membranous portions.

There are no median spines.

In the great chela the dactylopodite is not larger than the

164 MR, GEOFFREY SMITH ON THE

carpus; there is a row of about 8-10 tubercles on the inner

border of the carpus and just above this row there is a thick

earpet of downy hair. An inconspicuous tuberculation and a

certain amount of downy hair is present between the pincers.

‘The meropodite has three tubercles and a certain amount of down,

and the inner border of the ischiopodite has the usual double row

of tubercles. ‘

‘he succeeding legs are remarkably smooth and free from

hairs, though the last two joints of the last two legs are fairly

hairy.

The sternal keel is fairly sharp, but not very prominent, and

the basal joints of the legs in the neighbourhood of the keel are

hairy.

The interantennal spine is rounded and does not end in a

sharp point (Pl. X XVI. fig. 19).

The largest specimen is 6 inches in length.

Colour “(MeCoy). —The whole body and abdomen is dull

pale olive, varying in some speeimens to greenish horn-colour ;

membranous part of tail-flaps pale brown; anterior part of legs

bright blue ; basal and outer portion of bead mottled with scale-

shaped spots of dull ochreous yellow ; skin of joints bright red.

Locality. —The type specimens are from the pond in the

Melbourne University Grounds. Similar forms from various

localities in Victoria and also from the mud-flats on Murray

River. ‘The species is also widely distributed all over Central,

Northern, and Western Australia and in Queensland. It is, so far

as is known, absent from Tasmania and New Guinea.

Local Varieties :—

(a) Queensland. One specimen measuring 43 inches found

(Oct. 1891) in a field on a hillside at Coou. The rostrum is

rather blunt and truncated; the dactylopodite is longer than

carpus; and there are very well-marked pits on the great chela

and on the carapace.

(6) Central Australia, Specimens collected during the Horn

Expedition sre described by Spencer and Hall (‘ Report of

the Horn Expedition to Central Australia,’ Part II. Zoology :

Crustacea, p. 244) as being similar to ordinary Victorian species.

Shght differences are found in size of chela (a very variable

character according to growth and sex), absence of lateral spines on

rostrum, and the rostral keel is rather well marked. The authors

do not, however, propose making a new species for this form.

Remarks on the above species.—If we take into account the

habits of this species, which lives in the banks of ponds, quarry-

holes, and other stagnant waters, and is occasionally found walking

about in fields, there is no difficulty in understanding its wide

distribution all over the continent of Australia. It was found

in abundance by the Horn Expedition in Central Australia, where

it is said to be eaten regularly by the wandering tribes of blacks,

who know it as the Yabber. It is not easy to see how it can

survive in these arid and desert regions, lable to the most

FRESHWATER CRAYFISHES OF AUSTRALIA, 165

prolonged droughts, where all except the deepest water-holes

dry up, but presumably its burrowing habits save it. It seems

probable, from an examination of the anatomical features of the

land-Crayfishes of Victoria and Tasmania belonging to the genus

Engeus, that this latter genus has been derived from Cheraps

by an intensification of the burrowing habit and of the structures

associated with it.

Genus CHrars.

(Erichson, Arch. f. Naturg. xii. 1846, p. 94.)

Arthrobranch of last leg but one somewhat reduced, but

without fleshy peduncle and consisting of fairly numerous small

filaments springing from a very short bane. These filaments are

pointed at the end (Pl. XXVII. fig. 25). The ala on the podo-

branchs is broad as in Paracheraps. Many of the gill-filaments

attached to stem of podobranchs carry long curved hooks as in

Paracheraps ; the filaments on the other gills are either round or

pointed, but do not carry hooks (fig. 26).

The hook-setee on coxopodites are sharply recurved.

The antennal seale is not broad as in Paracheraps, but broadens

gradually after terminal spine (fig. 28).

The lateral keels of the rostrum are usually very sharp and

conspicuous, but they may be absent (C. intermedius).

The lateral carina on carapace is well marked, and may be verv

sharp and prominent, and there may also be a well-marked

median carina.

The distance between the tip of the rostrum and the cervical

suture is distinctly longer than that between the cervical suture

and the posterior border of carapace.

The sternal keel is sharp and very prominent (fig. 31).

The interantennal spine ends in a sharp point (fig. 27).

The great chela has the carpus either very long and slender,

and unprovided with a carpet of downy hairs, or else provided

with hairs and stoutly built.

In all other respects the generic characters, e. g. of the mouth-

parts, telson, and uropods, etc., agree exactly with Par acheraps.

Remarks on the above genus. ~The diagnostic characters by

which Cheraps may be distinguished from Parachen ‘aps are

(1) the nature of the posterior arthrobranch, (2) the antennal

scale, (3) the distance between cervical suture to rostrum and

cervical suture to posterior border of carapace, (4) the interantennal

spine, and (5) the sternal keel.

CHERAPS QUINQUECARINATUS. (Pls. XXIII. & XXVII. figs.

25-29.)

The Gilgit.

(Gray, Eyre’s Central Australia, vol. i. p. 410, pl. iii.).

The rostrum is moderately excavated, with ravher preminent

166 MR. GEOFFREY SMITH ON THE

keels ; it ends in a sharp spine, with two lateral serrations on

each side.

The lateral keel on the carapace is prominent and arched

outwards ; anteriorly it does not terminate in a prominent spine.

There is a well-marked median keel.

There is no prominence on the carapace behind the postorbital

spines, nor are there any tubercles upon the carapace, branchio-

stegites, or abdomen ; nor are there any bunches of hairs.

The first maxilla has the terminal segment or flagellum of the

endopodite entirely repressed (Pl. XX VII. fig. 29).

The third maxillipede has its inner borders abundantly clothed

with hair.

On the telson there are two lateral spines, but no medio-

lateral ones.

The great cheliped is long, but not slender ; the carpus is very

long, but stout; the dactylopodite is only about one-third as long

as the carpus. The pincer is slightly tuberculated internally and

there is a carpet of fine hairs upon the inner margin of the carpus.

There is a row of small tubercles upon the upper ridge of the

ischiopodite.

Greatest length 43 inches.

Locality.w— Western Australia, in streams tributary to the

Swan River.

CHHERAPS TENUIMANUS, sp.n. (Pls. XXIT. & XXVII. fig. 30.)-

The rostrum is deeply excavated in the middle, with sharply

upstanding lateral keels. The rostrum ends in a sharp spine, and

there are three lateral serrations on each side.

The lateral keel on the carapace is prominent and arched

outwards, ending anteriorly ina prominent spine. The median

keel is also much pronounced.

There is a distinct prominence on the carapace running back

from the postorbital spine, on which a few small tubercles are

situated. There are also a few small tubercles on the branchio-

stegites, and the body is covered with numerous groups of short

inconspicuous hairs.

The first maxilla has the endopodite consisting of a broad basal

‘segment, with the flagellum represented by a small conical segment,

the base of which is much smaller than the top of the segment

with which it articulates (Pl. XX VII. fig. 30).

The third maxillipede is only sparsely provided with hairs.

On the telson, besides the two lateral spines at the junction of

the calcified and membranous portions, there are two median

spines.

The great cheliped is fairly long and very slender. The carpus

is long and slender; the dactylopodite is more than half as long

as the hand. ‘There are no enlarged tubercles on the pincer and

there is no carpet of fine hairs on the inner margin of the carpus.

The row of tubercles on the upper ridge of the ischiopodite is

reduced to one or two small tubercles.

FRESHWATER CRAYFISHES OF AUSTRALIA. 167

The largest specimen measures 62 inches. Specimens in British

Museum about 11 inches.

Locality.— Margaret River, Western Australia.

CHERAPS QUADRICARINATUS. (Pls. XXIV. fig. 1; XXVIT.

figs. 31-33.)

(Von Martens, Monatsbericht Akad. Wiss. Berlin, 1868, p. 617.)

Nomenclature : —

Cheraps quadricarinatus Vou Martens.

Astaconephrops albertisii Nobili, Annali del Museo Civico di

Genova, x]. 1899 (1901), p. O44,

The keels of the rostrum are well marked and continued far

backwards on to the carapace; the rostrum ends in a sharp spine,

and there are three sharp lateral serrations on each side of the

rostrum.

The lateral keels on the carapace are well marked and sharp ;

they are arched outwards and end anteriorly in a sharp spine.

There is no median keel.

The carapace and branchiostegites are smooth, except for the

presence of three sharp spines laterally on each branchiostegite

just posterior to the cervical suture.

The first maxilla has the endopodite consisting of a base and

a fairly distinct flagellum jointed on to it.

The third maxillipede has the third and fourth segments

sparsely fringed on the inner and outer face with bristles

(Pl. XXVIT. fig. 32

The great cheliped is very slender and small, with slender

carpus; the carpus is smooth except for a fine serration on inner

border, and there is no carpet of downy hairs upon it. A pad of

fine hairs is, however, present on the inner face of the meropodite

between the spines. The row of tubercles upon the upper ridge

of the ischiopodite is represented by a single spine.

The succeeding two chelate legs are provided with a few hairs

on the terminal segment. The two posterior non-chelate legs

have a fringe of hairs on the last two segments.

The telson is without median spines.

Length about 4 inches.

Locality.— From Cape York, N. Australia (Von Martens), and

Katau, New Guinea (Nobili). Also from Aru Islands (var.

aruanus) and Mainikion, Baie Etna and Sabang, New Guinea

(var. lorentzi) (Roux).

Remarks.—Vhrough the kindness of Dr. Gestro, of Genoa,

I have been able to examine the single New Guinea specimen

described by the late Dr. Nobili. In the characters of the gills,

coxopoditie sete, mouth-parts, antennal scale, interantennal spine,

sternal keel, and general structural features it falls clearly within

the diagnosis of our genus Cheraps. Dr. Nobili, in regarding it

as allied to the New Zealand Paranephrops, was apparently led

astray by not having ever seen any of the Western Australian

168 MR. GEOFFREY SMITIL ON THE

Paracheraps. My identification of the New Guinea form with

Von Martens’s Cheraps quadricarinatus appears to me certain

from his description of the keels, tail-fan, and chelipeds of his

Cape York specimens.

Since the above was written, Dr. Jean Roux, of Basle, has

described certain specimens belonging to the genus Cheeraps

which are evidently closely allied to Cheraps quadricarinatus.

He has also re-examined the specimen described by Nobili as

Astaconephrops altertisti, and comes to the same conclusion as

myself that this specimen is an undoubted Cheraps, practically

identical with C. quadricarinatus. Dr. Roux remarks, however,

that Nobili’s specimen has three serrations on the rostrum,

whereas C. quadricarinatus only possesses two, and he therefore

proposes to keep Nobili’s species as valid. Since no other

differences are to be observed, it seems to me desirable to examine

many more specimens of both varieties before accepting Nobili’s

species on this single difference.

Dr. Roux also creates two new species, separate from quadri-

carinatus, for his specimens from the Aru Islands and for those

from other localities in New Guinea: Here, again, he admits the

practical identity of his forms with quadricarinatus, but calls

attention to certain differences in the proportions of the cheliped,

especially in the male. He also finds in the males of his

C. aruanus and C. lorentzi that the internal border of the carpus

of the cheliped in the male possesses smooth soft areas, which

have not been observed in C. quadricarinatus or albertisii. Tb

seems possible that these areas may be otly developed periodically,

as Calman suggests, and that they do not represent specific

characters. Without ventuiing to dogimatise on the subject, it

seems premature to accord these varieties of C. guadricarinatus

more than subspecific rank, especially as the variability of the

cheliped in the Crayfish according to age and individuality is

notorious: (See Dr. Jean Roux (15, 16),. Zoologischer Anzeiger,

Bd. xxxvii. Nr. 5, p: 104, Feb. 1911, and Notes from the Leyden

Museum, vol. XxXiii. p. 81,1911.) Calman (17) has examined a

number of specimens from the Mimika River, New Guinea, and

has observed soft areas on the claws of the inale identical with

those observed by Roux, and he also notices variations in the

proportions of the chela, which, howeVer, graduate into one another.

He is emphatically of the same opinion as myself that all these

specimens from New Guinea and the adjacent islands hitherto

described are not sufficiently distinct to warrant their separation

into more than One species.

T am therefofe all the more inclined for the rete tO preserve

the name gtiadricarinatis for all these forms.

CuaitaPs INTERMEDIUS; sp: ns (Pls. XXIV. fig. 2; XXVII.

fig. 34.)

The rostrum ig fat and qtite nnexeavated, without any lateral

FRESHWATER CRAYFISHES OF AUSTRALIA. 169

keels upon it. It ends in aspine and there are two inconspicuous

lateral serrations on each side. It closely resembles the rostrum

of Paracheraps bicarinatus.

The lateral keels on the carapace are blunt and deepened,

ending posteriorly in a rounded boss. There are no other

tubercles, prominences, or groups of hairs upon the body. So far

the external features are practically identical with P. bicarinatus,

but it can be at once distinguished by the generic character of

the length between tip of rostrum and cervical suture being

greater than that between cervical suture and end of carapace.

The first maxilla has a very small pointed terminal segment on

the endopodite, which articulates by a broad base on to the basal

segment (Pl. XX VII. fig. 34),

The third maxillipede is fairly well provided with hairs on its

inner borders.

There are no median spines on telson.

The great cheliped is stout and broad; the hand itself being

broad and rather flattened; the dactylopodite is about equal in

length to the inner border of the carpus. The finger is definitely

tuberculated internally, often with two enlarged tubercles. There

is a small carpet of fine hairs on the inner margin of the carpus.

On the under side the carpus has a pronounced longitudinal

ridge with marked excavations on either side of it.

The upper ridge of the ischiopodite has a row of small round

tubercles upon it.

Length 4 inches. ‘

Locality.—Two specimens from Western Australia, April 30th,

1880. No further locality given.

Remarks on the above species.—This species is of considerable

interest, as it forms a transition from the genus Chwraps to

Paracheraps bicarinatus: In general appearance, in the features

of the claws, carapace, rostrum, etc., it agrees exactly with

Paracheraps; in the fact that the posterior arthrobranch is not

rudimentary, the sternal keel is sharp and very prominent, the

interantennal spine is pointed and not rounded, the antennal

scale is not conspicuously bitoad, and in the length between tip

of rostrum and cervical suture being greater than between latter

and end of carapace, it exhibits the characteristic features of

the genus Cheraps.

List oF LIrerRATURE.

Sitaw:— Zoology of New Holland, Vol. i., 1843.

Gray.—Eyre’s Journals of Expeditions of Discovery into

Central Australia, Vol. i., 1845.

3. Ertcuson.—Archiv fiir Naturgeschichte, Jahrg. 12, 1846.

4. Dana.—United States Exploring Expedition: Crustacea.

Part I., 1852.

Mitye-Epwarps.:— Hist: Nat. Crust., Vol. 11. p. 332,

170 MR. GEOFFREY SMITH ON THE

6. Hess.—Archiv fiir Naturgeschichte, Jahrg. 31, 1865.

7. Heriter.— Novara’ Reise, Crustaceen, 1865.

8. Von Marrens.—Ann. Mag. Nat. Hist. ser. 3, Vol. xvil.,

1866.

9. Von Marrens.—Monatsbericht Akad. Wiss. Berlin, 1868.

0. Huxiry.—Proe. Zool. Soc. 1878, p. 764.

1. McCoy.—Prodromus of the Zoology of Victoria, Vol. 1.,

1885.

12. SpencE-Batse.—‘ Challenger’ Reports, Vol. xxiv., 1888.

13. Hasweiu.—Catalogue of the Australian Museum: Crustacea.

1890.

14. Nopit1.—Annali del Museo Civico di Genova, Vol. xl.,

1899-01.

15. Roux.—Zoologischer Anzeiger, Bd. xxxvil. Nr. 5, p. 104,

Feb. 1911.

16. Rovx.—Notes from the Leyden Museum, Vol. xxxui. p. 81,

OUBIE

17. Canman.—Ann. Mag. Nat. Hist. ser. 8, Vol. viil., Sept.

SOIL.

EXPLANATION OF THE PLATES.

IP, SOW,

Astacopsis franklinii, male. Large form. +} nat. size. North Tasmania.

Jett, 2k

Astacopsis franklinii, female. Small tasmanicus form. Nat. size. South

Tasmania, Mt. Wellington.

IP. DOW IL

Astacopsis serratus, male. Large form. % nat, size. Murray River.

Pity SWIO

Astacopsis serratus, female. Small form. # nat. size. Yarra River.

Pr. XVIII.

Astacopsis serratus, male. Small form. Nat. size. Blue Mountains.

; IP, 2D,

Astacopsis kershauwi, male. Large form. 4 nat. size. Moe River, Gippsland.

Pu. XX.

Astacopsis kershawi, male. Small form. Nat. size. Narracan River, Gippsland.

IP, DO:

Paracheraps bicarinatus, male. % nat. size. Melbourne University pond.

Pru. XXII.

Cheraps tenuimanus, male. $ nat. size. Margaret River, Western Australia.

—

FRESHWATER CRAYFISHES OF AUSTRALIA. 1

Pr. XXIII.

Cheraps quinquecarinatus, male. $ nat. size. Swan River, Western Australia.

—

. 2.

33.

34.

Pu, XXIV.

Cheraps quadricarinatus, male. # nat. size. From Katan, New Guinea.

Thoracic legs not represented. From a specimen lent by Dr. Gestro.

Cheraps intermedius, male. Nat. size. Western Australia, locality not

known,

Pr. XXV.

. Third maxillipede of A. franklinii.

Second mavillipede of ditto.

First maxillipede of ditto.

Second maxilla of ditto.

First maxilla of ditto.

Mandible of ditto.

PL. XXVI.

Head of A. franklinii from ventral view. 'To show interantennal spine and

upper lip.

. Antennal seale of A. franklinii.

“ndopodite of first maxilla of 4A. franklinii.

Termination of branchial filament of A. franklinii.

. Termination of coxopoditie seta of 4. franklinii.

. Head of A. serratus from ventral view. ‘To show interantennal spine and

upper lip.

Antennal scale of A. serratus.

View of sternum of thorax from ventral surface, showing sternal keel and

attachment of thoracic legs in A. serratus.

Endopodite of first maxilla of Paracheraps bicarinatus.

Termination of branchial filament of P. bicarinatus.

Termination of coxopoditic seta of P. bicarinatus.

Terminal portion of the last arthrobranch of P. bicarinatus.

Interantennal spine of P. bicarinatus.

. Antennal scale of P. bicarinatus.

Third maxillipede of P. bicarinatus.

. Second maxillipede of P. bicarinatus.

Kirst maxilla of P. bicarinatus.

Mandible of P. bicarinatus.

Pr. XXVIN,

Last arthrobranch of Cheraps quinquecarinatus.

Termination of branchial filament from pleurobranch of C. quinque-

carinatus.

Interantennal spine of C. quinquecarinatus.

Antennal seale of C. quinguecarinatus.

Endopodite of first maxilla of C. quinquecarinatus.

Endopodite of first maxilla of C. tenvimanus.

View of sternum of thorax from ventral surface, showing sternal keel and

attachment of thoracic legs in C. guadricarinatus.

Third maxlilipede of C. quadricarinatus.

Second maxillipede of C. quadricarinatus.

Endopodite of first maxilla of C. intermedius.

7/2, MR. A, E. CAMERON ON

11. Structure of the Alimentary Canal of the Stick-Insect,

Bacillus rossii Fabr.; with a Note on the Parthenogenesis

of this Species. By ALFRED H. Cameron, M.A., B.Se.,

Fullerton Scholar of the University of Aberdeen and

Research Student in the University of Manchester *.

[Received October 6, 1911: Read November 21, 1911.]

(Plates XX VITI.-XXX.7)

L[ntroductory:

The following description of Bacillus rossti, a species of the

Phasmide, is translated from Girard’s ‘ Traité Elémentaive d’Ento-

mologie’ (p. 100) (Paris, 1879).

‘‘The genus Bacillus is of speeial interest, as it includes the

only species of Phasmid known to occur in Europe; this 1s

Bacillus rossii Fabr., green, greyish green, or brownish red, with

integument either smooth or bearing slight projections, antennz

composed of 12 to 23 segments, with a dorsal ridge, with slender

legs angularly striated, the anterior pair very long, the two

posterior pairs of femora tridentate below. The male is from

45 to 55 mm. long, the female from 58 to 108 mm, ‘The species

is native to South Europe and North Africa, and was discovered

by Rossi in the gardens and heaths of Tuscany ; it is common on

the green hedgerows bordering water-filled ditches ; it is found

also in Dalmatia and in Istria. Near the town of Pola this

motionless Phasmid oecurs on the Montpellier rock-rose; but so

great is its resemblance to the twigs among which it lives that it

is very difficult of recognition; its enemies, besides parasitic

larvee, are the Praying Mantis and the Green Lizard. A smaller

race of Bacillus rossii exists with antenne of 12 segments, thorax

bearing minute prominences, the female 54 to 67 mm. in length,

of which several authors make a distinct species under the name

of B. gallicus Charp., 8. granulatus Brullé, found in Greece at

the time of the French expedition, also in Andalusia (Rambur),

in North Africa, in the South-East and South-West of France,

at Nice, at Cannes, at Hyéres, near Toulon. In the early spring

the insects may be captured by shaking the bushes over an open

umbrella, when there fall into it both adults which have hiber-

nated and larvee in various stages of development; the larvee

being more delicate than the adults preserve badly when collected,

becoming twisted and folded. The species is said to be found in

isolated examples as far north as Orleans.

‘“‘B. rossti is found in Algeria, in places which are covered with

grasses and shaded from the light.

“The genus includes one-fortieth of the species of Phasmidee of

* Communicated by Professor Sypney J. Hickson, F.R.S., F.Z.S.

+ For explanation of the Plates see p. 182.

PZ 4912 Pl xx Vile

a Bs

uti

A.E.C.del. West, Newman lith.

ALIMENTARY CANAL OF BACILLUS ROSSII.

' tia ‘ i

eee ‘ t wil ;

ae i

i 5 wy

{

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: 5

‘i t

ew at

Tos as;

P.Z.S.1912. Pl. XXIX.

e\e> i

Y |

Cn Mp.

ben nn — = —— arn

yey ECigaa ae

| Sllebinn yee Ble [c,

a eeCUL Ey Wy

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——

A.E.C. del. West, Newman lith.

ALIMENTARY CANAL OF BACILLUS ROSSI.

‘ PZ.5. Leta Pl Axx.

ay,

Ai QO

OD Iss

‘ A.E.C. del. West, Newman lith.

Be ALIMENTARY CANAL OF BACILLUS ROSSII.

THE STICK-INSECT. Ng

the Old World, and it has a wide distribution, occurring in Europe,

Africa, East Indies, Ceylon, Java, China, Sandwich Islands,

Australia, Island of Hamoa, and New Zealand.”

In the course of the present year several English collectors

have had in their possession specimens which were observed

to deposit their ova during the months of February, March, and

April, and the hatching of the ova began in the latter part of

June. It is highly probable that this insect, alien to the British

fauna, made its entrance into this country along with fruit-cargoes

and nursery-stock imported from France. ‘There is not much

chance of its becoming established here since, being accustomed

to mild temperatures, it fares rather badly under the changeable

conditions of the English climate. In captivity they require

close attention; but by supplying them regularly with fresh

food-plants and by keeping the temperature of their surroundings

at about average house-temperature they appear to thrive.

Under these conditions they are quite healthy, and the mortality

is very low.

Note on Parthenogenesis,

A very curious phenomenon in connection with the repro-

duction of Bacillus rossii may be worthy of note. Although my

specimens were in the third generation reckoning from the first

generation in captivity, no male had been observed, and perhaps

parthenogenesis had been the normal method of reproduction

previous to this, as my observations were necessarily limited by

the fact that I had no record of the generations preceding the

first one in captivity. Von Baehr (1907) records that in 1903

he received specimens of B. rossii whi¢gh belonged at least to the

seventh parthenogenetic generation, being descendants of a half-

grown female captured in Dalmatia in 1896, But the latter, he

says, may itself have arisen from an ovum parthenogenetically

produced, so that in all probability it was not the first in the

ancestral line to produce ova which had not been fertilised by

the sperm.

These facts seem to lend support to the supposition that the

female of B. rossit can dispense with the GAR altogether and

that virgin, rather than sexual reproduction is the normal

method adopted by this insect of perpetuating its race. We

cannot state with any degree of confidence why the insect has

betaken itself to the parthenogenetic production of eggs, but that

advantages accrue we are tolerably certain. In cases where,

owing to some cause or other, there is a scarcity of males, the

method would be a decided acquisition, since the great majority

of the more prevalent females would stand a very poor chance of

ever meeting a male and of the consequent enjoyment of sexual

intercourse.

I think that it is open to make the hypothesis that both sexes

of B. rossii originally existed in equal numbers. Owing to some

cause with which we are not acquainted, the males began to

174 MR. A. E. CAMERON ON

diminish so that the dominant female, in order to safeguard the

species against utter extirpation, began to reproduce partheno-

genetically. We must understand that the process has been a

very gradual one, proceeding slowly throughout several centuries,

the male finally becoming almost extinct.

Another interesting question intimately connected with the

subject of parthenogenesis may be asked. Why does the female

produce ova from which only females arise? While I offer no

suggestion, I may be permitted to allude to the same phenomenon

which occurs regularly in many of the Cynipide or Gall-fiies.

The male of Cynips kollari, the maker of the marble-gall of the

oak, has never been observed, although many entomologists have

given this species their close attention, and it is now generally

believed that it does not exist, but that parthenogenetic repro-

duction is the only method of preserving the species.

The following is quoted from Adler and Straton’s ‘ Alternating

yenerations’ (Oxford, 1894).

“Tt would appear that in Cynips kollari the sexual generation

is wholly subordinated to the asexual, and in /thodites rose, which

forms the pretty Bedeguar galls on the rose-tree, the process is still

going on, and the males are becoming functionless and extinct.

... Lt is difficult to believe that the agamous (or asexual) can be the

primitive form ; or that the perfectly formed sexual organs could

have been evolved unless the sexual had been the earlier

generation.”

Alimentary Canal.

While at the Royal College of Science, London, in the beginning

of 1911, it was suggested to me that the alimentary canal of

Bacillus rossti would repay careful observation. The material at

hand I subsequently worked up in the Zoological Department

of Manchester University, and I here take the opportunity of

expressing my indebtedness to Professor Hickson as well as to

Mr. Manean for many timely hints.

In the Orthoptera viewed as a whole the alimentary canal

shows a very uniform structure; but in the various families

there are many secondary differences, sometimes even in the same

family, necessitating special description. The gizzard is im-

portant in the Orthoptera, and there are diverse degrees of

complication in its chitinous armour. The intestinal cca vary

ereatly in number, there being eight in Mantidw and Blattide,

six in Acridiide, and one pair ot lateral ceca in Locustide and

Gryllide. The diverse variations of structure in the gizzard,

its different degrees of complexity added especially to the pre-

senee or absence of intestinal ceca, suggested to Bordas (1897) a

te]

method of dividing the Orthoptera into two large groups :

1. The Acolotasia, or Orthoptera without intestinal caxea ;

2. The Colotasia, or Orthoptera with intestinal ceca more or

less numerous,

THE STICK-INSECT. 175

The Malpighian tubes in all Orthoptera except the Forficulide

ave very numerous, and are generally grouped in six bundles

opening at the summit of six tubercles, as in the Locustide and

Blattide. In all the Gryllide, on the contrary, the Malpighian

tubes are arranged in a single bundle opening into the dilated

extremity of an unpaired excretory canal. Very characteristic

is the presence of a “ciliated border ” (un revétement cilié)

projecting from the cells of the mid-intestine and _ lateral

appendages.

The alimentary canal of B. resszi is divided into three parts, as

in all arthropods.

1. Fore-gut, ectodermic, corresponding to the stomodeum, and

comprising the mouth , pharynx, cesophagus, and crop (Pl. XX VIIT,

fig. 1); the gizzard is ‘absent.

a Mid-g ut, endodermic, corresponding to the mesenteron,

beginning “posterior to the crop and ending at the insertion of

the Malpighian tubes. The lateral ceca, ” generally developed

from ie anterior part of the mid-gut, are rudimentary in B. rossii

(fig.

3. tad gut, ectodermic, corresponding to the proctodeum,

beginning at the i insertion of the Malpighian tubes and including

small intestine and rectum (fig. 1).

In many respects the alimentary canal is peculiar, and the main

features which contribute to its uniqueness may be tabulated

as follows :—

1. The rectilineal shape of the gut, there being no con-

volutions of the posterior region (fig. 1).

. The complete absence of the gizzard.

. The rudimentary nature of the mid-gut digestive ceca

(figs. 1 & 3, C.).

4. The presence of numerous peculiar conical tubular.

organs on the posterior position of the mid-gut

Cig. t,o ka.)

In preparing sections for histological examination of the gut

various stains were employed. Grenacher's hematoxylin was

especially g good in the differentiation of chitin, while Heidenhain’s

iron- -hematoxylin proved very effective where epithelium was

concerned. Professor Hickson’s brazilin stain and borax carmine

were also used to advantage. To get the best results the gut

required rather careful tre atment, and in the matter of a fixing

agent I found that Schaudinn’s fluid was all that could be

desired,

It is generally accepted that the mid-gut of insects is endo-

dermic in its origin, but in the ease of B. rossii Heymons (1897)

demonstrated that it was really ectodermic, being developed from

two epithelial outgrowths of the stomodeum and _ proctodeum

which surround the yolk and become united. This manner of

formation, he maintains, may be found to hold good for all

Phasmidee.

oo bo

176 MR, A. E. CAMERON ON

The fore-gut up to and including the crop is of rather simple

structure. As a rule the histological structure of the insect

alimentary canal is always the same: on a layer composed of

muscular and connective tisssue there rests an epithelium which

secretes a strong chitinous lining. This cuticle is raised into

straight longitudinal ridges which bear numerous minute horny

denticulations. The pharynx of B. rossi (fig. 1, Ph.) is very

short and is lodged in the posterior cephalic region, the cesophagus

(fig. 1, CH.) occupying the elongated prothorax and passing

insensibly into the somewhat dilated crop (fig. 1, Cr.) in the

region of the mesothorax. Lastly, the crop joins the mid-gut

(fig. 1, Gm,) in the region of the metathorax, and, indeed, the

internal wall of the crop is telescoped into the cavity of the

mid-gut as a cone-shaped prolongation (Riissel), But in addition

to what may be called the primary direct extension of the crop

(Pl. XXIX. fig. 5, Ov, and Ov,) there is also a secondary

indirect extension (fig. 5, Ov, and Ove,). The chitinogenous

epithelial cells of the wall of the crop extend backward into

the mid-gut for a short distance as far as the point A denoted

in the figure. They then become folded back on themselves up

to the point B, and turning once more form the long dorsal pro-

longation which reaches a comparatively long way into the mid-gut.

At C the cells are again reflected, and finally at D join with the

larger epithelial cells of the mid-gut. The chitinous cuticle lining

the prolongations is continuous with that of the crop, From fig. 5

it will be observed that the extensions are not symmetrical but

are much more pronounced dorsally than ventrally. This un-

symmetrical arrangement is known to occur only in the few

Phasmids in which the alimentary canal has been investigated,

and the exact reason of the greater development of the dorsal

lamina is not known. In the larve of Chironomus the esophageal

~ telescoping is uniform and symmetrical, and I believe that this is

the case with the majority of insects where the telescoping occurs.

Heymons (1897) gives it as his opinion that the elongate

dorsal lamina (Verschlussklappe) functions as a closing-valve,

preventing the back-flow of digested food from the mid- to the

fore-gut. But Sinéty (1901) does not agree with this interpre-

tation, as he is convinced that if the direction of the food-current

were to be reversed this flap would be overcome by the pressure.

The bundles of circular muscle surrounding the anterior part of

the mid-gut would be quite effective in checking the reversed

food-current if any sueh occurred.

The mid-gut of B, rossij is divided into two distinct parts, of

which the anterior is characterised by very prominent transverse

folds (P], XXVIII. fig, 1, Gm, ; fig. 3, Gm,; Pl. X XIX. fig. 5);

while the posterior (fig. 1, Gm,), besides being narrower, is easily

distinguished by the presence on its external walls of numerous

(about 59) conical, tubular organs (fig, 1, Ta,),

In all Orthoptera as before stated, except the Phasmide, the

surface of the mid-intestine is enlarged by diverticula of various

{THE STICK-INSECT. Wize

shapes developed from the anterior end, eight long cylindrical

ceca in Blattide and Mantide, two large rounded ceca in

Locustide and Gryllide, or again six pouches in Acridiidee, which

may be further provided with inferior appendages. In B. rossii

these ceca (fig. 1, fig. 3, fig. 5, C.) exist merely as small lobes

clustered together in small numbers and very rudimentary.

These ceca must not be confused with the gland-like structures

(fig. 1, Ta.) which occur just anterior to the Malpighian tubes.

Indeed it is not definitely known whether the function of these

latter so-called “ glands” is really glandular, secreting a digestive

* juice which is poured into the intestine, or excretory, eliminating

waste products like the Malpighian tubes. At their point of

attachment to the gut these organs are pear-shaped, tapering

abruptly into a small tubular thread, the calibre of which is

about half the size of that of the Malpighian tubes amongst

which they pass. Heymons (1897) has shown that in origin and

morphology these tubular appendages bear a very clue resem-

blance to the Malpighian tubes; but physiologically he regards

the two sets of organs as differ ent, since they are dissimilar

in their behaviour towards injections of coloured particles.

Generally speaking, it was found that the tubular appendages

were the less sensitive, but they may eliminate some substances

quite as actively as the Malpighian tubes, as, for instance, Ehrlich

blue. If thei function be excretory, it is less general than that

of the ordinary organs of excretion.

The internal wall of the mid-gut is a TEED of a layer of

“

large cylindrical epithelial cells (Pl. XXX. fig. 5, E; fig. 6, E

and E ,) provided with a “striated border ” (fig. 5 and fig. 6, Ch.)

and further they are characterised by a continual secretion of

globules of mucilage (fig. 5 and fig. 6, Mg.), which are deposited

in the intestinal cavity. Cuénot (1895), amongst other authors,

describes the presence of young cells in this epithelium undergoing

mitotic division and gradually replacing the older cells, These are

of the nature of small interstitial cells with rather dense, deep-

staining nuclei; but I myself have not observed any mitotic

figures in them. There is still dispute on the question of these

so-called ‘ germinative-centres,” and many authors, notably

Frenzel (1886) and Visart (1894), consider them as glandular

crypts.

There is no protective chitinous cuticle lining the internal

walls of the mid-gut, but its place seems to be falen by a very

slender peritrophic membrane (figs. 5 and 6, Mp.) which is quite

detached from the epithelium. It takes its origin at the point

where the fore-gut epithelium joins that of the mid- gut, and

extends as far as the hind-gut, which it enters for a ‘short

distance. Thus the food passes through the alimentary canal

without the delicate epithelial cells running any risk of abrasion

from jagged particles eaten by the insect in the course of its diet.

This membrane (Trichter) is certainly not a prolongation of the

chitinous cuticle of the crop, as Mecanikow and Schneider (1890)

Proc, Zoou, Soc,.—1912, No, XII, 12

178 MR. A. E. CAMERON ON

believed, but seems to be secreted by the most anterior epithelial

cells of the mid-gut. The question may be asked: How does

S. rossu assimilate its food? ‘This can only be explained by a

knowledge of the laws of osmosis, so that the digestive juices

secreted by the epithelium and the fluid juices which result from

their action must pass through the membrane, the former to act

on the food substances, the latter to be absorbed.

Schaeider (1890) and Adlerz have observed the presence of a

peritrophic membrane in diverse Orthoptera, Phyllodromia, Peri-

planeta, Bacillus, Locusta, Forficula. ‘This has been verified by

Cuénot (1895), who adds some details as regards its origin of

which little is known.

Plateau (1875), in his splendid work ‘ Recherches sur Jes

phénomenes de la digestion chez les msectes,’ has shown that

digestion first begins in the crop, where the alkaline or neutral

secretion of the salivary glands acts on starchy substances,

changing them to glucose. The processes of digestion are con-

tinued in the mid-gut, the epithelium of which also secretes a

fluid with alkaline or slightly acid properties which has the power

of changing albuminoids into peptones and of emulsifying fats.

Petrunkewitsch (1899) held that in certain Orthoptera the

crop was the principal organ for the absorption of digested food,

citing as his proof the presence of fat in the epithelial cells. But

Sinéty (1901) demonstrated clearly that the fat globules here

present are really elaborated by the epithelial cells from materials

extracted from the blood, and thus the epithelium of the crop is

functionally comparable to the fat-body. Again, many authors

adopted the idea that owig to the shortness of the mid-gut

supplementary absorption must be carried out in the crop

(Plateau [1876] and Jousset de Bellesme in the Blattide), or

even in the hind-gut (Plateau [1878] and Frenzel [1886]). But

from all known laws of osmosis it is highly improbable that this

can be the case, for the thick chitinous cuticle lining both crop

and hind-gut is Impermeable to dissolved substances. Again, it

is nothing short of absurd to suggest that absorption can go on

below the point of insertion of the Malpighian tubes which mark

the posterior limit of the mid-gut, and invariably pour their

waste-products into the alimentary canal at this place.

In #. rossi the Malpighian tubes: (Pl. XXVIII. fig. 1 and.

Pl. XXX. fig. 7, Mt.) are very numerous and are disposed re-

gularly in a circle round the anterior end of the hind-gut, into

which they open in groups of three to six. These groups, occurring |

to the number of 20 to 30, are equidistant from each other, and

the tubes composing any one group fuse at the apex of a small

conical tubercle which arises as an evagination from the gut

and is traversed by a minute duct (fig. 7, Cd, and Cd,). The con-.

stituent cells of the Malpighian tubes also possess the “ ciliated

border” characteristic of the epithelial ceils of the mid-gut, but

the “cilia” in both cases do not possess any power of motion

even in an indifferent medium. It cannot be argued that the

fact of the insect being dissected would cause the “cilia” to

cease vibrating immediately, as the cilia on the gill-plates of

THE STICK-INSEC'T. 179

Unio, the fresh-water mussel, continue their vigorous move-

ment long after the removal of the gills from the animal,

In conclusion I may state that the different authors who have

investigated the matter refer to the “ciliated border” under

various names, such as “striated plate ” “ciliated plate,” “ plate

composed of free-filaments or rods,” “ brush-like border”

(Hirchensaum of Frenzel); but all these terms convey to our

minds exactly the same idea.

The hind-gut, including the small intestine (Pl. XXVIII. fig 1,

Is.) and rectum (fig. 1, R.), is, like the fore-gut, lined with chitin.

In the small intestine I distinguish two regions—the anterior one

broad, with small internal longitudinal folds, and the posterior

narrow, the circumference of which is greatly lessened by the

deep infolding of the epithelium, the lumen appearing only as

six narrow bays. Just anterior to the narrowing a circular band

of muscle surrounds the small intestine and by its contraction

shuts off the anterior from the posterior region. A curious

valvular apparatus (PI. XXX. fig. 9, F.) occurs at the anterior

oritice of the rectum in the shape of six muscular pyramidal

projections inserted by one of their surfaces in the internal wall

of the gut, and with the surface adjacent to the rectum slightly

hollowed out. The small intestine can be completely cut off from

the rectum by a muscular band on the external wall, which when

it contracts brings the six valves together so that the passage is

completely closed.

The rectum (fig. 1, R.) bears internally six longitudinal thicken-

ings of large epithelial cells (Pl. XXX. fig. 8, KE.) which have a

glandular function, and between the epithelial cells and the

basement membrane (fig. 8, Mb.) there ramifies a system of

trachee, suggesting a rather perfect aeration of rectal tissues.

Musculature of the Alimentary Canal.

As always in insects, the gut is provided with two sets of

muscles, longitudinal and circular, which extend along its whole

length. But in B. rossii there is a marked deficiency of longi-

tudinal muscles in the region of the fore-gut, and their place

seems to have been in great measure taken by six broad bands of

short oblique fibres which belong morphologically to the circular

system (PI. XXVIII. fig. 2, Mo.). On the mid-gut the bands of

circular muscles are set wide apart, producing the characteristic

furrowed appearance of this region of the intestine.

The longitudinal muscles present some interesting peculiarities.

They extend all along the intestine with the exception of the

anterior part, as slender fibres equidistant from each other, and

numérous branches are given off from the main strands. These

branches may anastomose where they run close together, or they

may taper off into extremely fine threads. The longitudinal

muscles seem to play an important part in connection with the

morphological significance of the pyriform organs of the posterior

region of the mid-gut. he latter are observed to arise either in

close proximity to one, or equidistant from two longitudinal

i2*

180 MR. A. E. CAMERON ON

muscles. In the first case a single branch is given off from the

main fibre and may be traced to the surface of the pear-shaped

dilatation ; in the second case two branch fibres may be supplied,

one from each of the two nearest longitudinal muscles. Under a

nerve stimulus these branches are said to be the mechanism

whereby the mid-gut appendages are animated with a movement

analogous to that observed in the case of the Malpighian tubes

(Sinéty [1901)).

In the anterior region of the mid-gut there is a distinct dorso-

longitudinal furrow (Pl. XXVIII. fig. 1, F, and fig. 3, F,) in

which numerous longitudinal muscles are congregated, and toa

less extent the same phenomenon is observable in the ventro-

longitudinal furrow. The fibres which are situated dorsally

and laterally on the mid-gut are united into three, then into

a single strand which proceeds anteriorly to its attachment in

the pronotum. Those fibres lying ventrally are united into a

somewhat more slender strand.

As they pass backward to the hind-gut the longitudinal

muscles become grouped into six strong bands applied symmetri-

cally to the wall of the gut, and giving off distally cutaneous

branches which proceed outwards.

Summary.

The alimentary canal of B. rossii presents us with much that

is of interest. Its rectilineal shape is what we would naturally

expect from the narrow elongated form of the insect. But the

absence of a gizzard, together with the rudimentary digestive

ceca and the presence of tubular filiform organs at once arrest

attention. The anterior portion of the mid-gut is puckered owing

to the isolation of the circular muscle-bundles. The Malpighian

tubes are curiously grouped in bundles of from three to six, each

bundle opening into the gut by a common duct. The small

intestine is separated from the rectum by six powerful pyramidal

valves. The internal wall of the crop telescopes into the lumen

of the mid-gut, and the telescoped portion is reflected on itself in

a curious manner.

The fact that the males of B. rossii are disappearing points to

parthenogenesis not being the primitive method of reproduction,

but suggests that the species has become adapted to it through

the failure of sexual reproduction,

LITERATURE.

1841, Durour.—Recherches anatomiques et physiologiques sur

les Orthopteres, les Hymenoptéres et les Névropteres.

Mem. Ac. Se. Paris, t. viii. p. 265.

1858. Srropor.—Recherches sur les sécrétions chez les insectes.

Ann. Sc. Nat. (4) t. x. p. 251.

1875. Puarrav.—Recherches sur les phénoménes de la digestion

chez les insectes. Mém. Acad. Roy. Belgique, t. x1.

lre partie.

1876.

1878.

1879.

1882,

1886.

1886,

1889.

1890.

1894.

1895.

1896.

Teo.

1897.

1899.

1900.

1901.

1904,

1907.

THE STICK-INSECT. 181

Puateau.—Note sur les phénoménes de la digestion chez

la Blatte américaine. Bull. Acad. Roy. Belgique, (2)

tai. p.. F206.

ScurvpLer.—Beitriige zur Kenntniss der Malpighischen

Gefiisse der Insekten. Zeits. fiir wiss. Zool., 30 Bad.,

p-. 087.

Girarp.—Traité élémentaire d’Entomologie, vol. 1. Paris.

VIALLANES.—Recherches sur l’histologie des insectes et

sur les phénomeénes histologiques qui accompagnent le

développement post-embryonnaire de ces animaux.

Ann. Sc. Nat.

Frenzev.—Kiniges iiber den Mitteldarm der Insekten

sowie iiber Epithelregeneration. Arch. ftir mikr, Anat.,

26 Bd., p. 229.

Mratn and Denny.—The Structure and Life-History of

the Cockroach. London.

Scuirror.—Beitriige zur Histologie der Insekten. Zool.

Jahrb. (Abth. fiir Anatomie), 3 Bd., p. 611.

Scunemper._Ueber der Darmkanal der Arthropoden.

Zool. Beitriige, Bd. 1.

Visartr,—Contribuzione allo studio del tubo digerents

degli Artropodi, Atti d. Soc. Toscana di Se, Natur.,

vol. xint,, p..20.

Cutnor.—Etudes physiologiques sur les Orthopteres.

Arch. de Biologie, t. xiv.

Borpas.—Considérations générales sur l'appareil digestif

des Phasmide. Bull. Mus. Paris.

Borpas.—L’appareil digestif des Orthopteres. Ann. Se,

Nat. t. v.

Hrymons.— Ueber die Organisation und Entwickelung von

Bacillus rossii Fabr. 8.B. Ak. Berlin.

Curnor.—Sur la détermination du sexe chez les animaux.

Bull. Se. France et Belgique, t. xxxil.

Dominique. — Parthénogénése et thélytokie chez _ les

Phasmides. Bull. Soc. Sc. Nat. Ouest Fr., t. xi.

PerrunKewrtscu.—Zur Physiologie der Verdaming bei

Periplaneta orientalis und Blatta germaniea. Zool. Anz.

1899, p. 137.

Kuri. —Biologisches iiber Bacillus rossii. Ent. Zeitschr.

nos. 16-17.

Srvtry._Sur la parthénogénése des Phasmes. Bull. Soc.

Ent. France, p. 194.

Sintiry.—Les tubes de Malpighi chez les Phasmes, Bull.

Soc. Ent. France, p. 350.

Srviry,—Recherches sur la biologie et Vanatomie des

Phasmes. L’Université de Paris. Cellule XTX.

Goprrman.—Beitriige zur Kenntniss von Bacillus rossit.

Arch, f. Entw.-Mech., Bd. xii.

Hennecuy.—-Les Insectes. Paris.

y. Baur—Zool. Jahrb. (Anat) vol, xxiv.

182

ON THE STICK-INSECT.

EXPLANATION OF THE PLATES.

Pruate XXVIII.

Fig. 1. Alimentary canal of Bacillus rossii. Md., mandibles; Ph., pharynx ;

Fig. 2

M., muscles of the pharynx; €., esophagus; Cr., crop; Gmyj, anterior

region of the mid-gut; Gmg, posterior region of the mid-gut; F., dorso-

longitudinal furrow in which several longitudinal muscles are congregated ;

C., rudimentary ceca of the mid-gut; Ta., tubular appendages of the

mid-gut with thei filiform prolongations; Mt., Malpighian tubes ;

Is., small intestines; R., rectum, with the rectal longitudinal folds Rf.

. Portion of the external wall of the crop. Mc., circular muscle; Mo., band of

oblique muscle fibres, of which there are six, extending longitudinally along

the cesophagus and crop, and displacing in great measure the ordinary

longitudinal muscles ; Rme., circular muscles reflected to show one of the

_bands of oblique fibres.

Vig. 3. Crop and anterior part of the mid-gut. Cr., crop; Gn., ganglion ; Tr., traches ;

Fig. 4

Gm., mid-gut; C., rudimentary ceca of the mid-gut; FI., longitudinal

muscles grouped together in the dorso-longitudinal furrow; M1,, longi-

tudinal muscles lying outside the furrow.

PuatEe XXIX.

. Transverse section of the crop. Mc., circular muscles; MI., longitudinal

muscles; Pm., peritoneal membrane; E., epithelial cells which secrete the

chitinous lining Ch.

Fig. 5. Longitudinal section of the posterior region of the crop and anterior region of

the mid-gut. Mc., circular muscles of the crop; C., rudimentary digestive

ceca; Me., mucilaginous globules secreted by the epithelial cells of the

ceca as also by those of the mid-gut. In the mid-gut these globules

occupy the space between the epithelial layer and the peritrophie mem-

brane. They are not included in the figure. M1., longitudinal muscles;

Ch., chitinous lining of the erop secreted by the chitinogenous cells cc.,

extending with the “valvular” prolongation into the mid-gut; Mp., peri-

trophic membrane; E., epithelium of the mid-gut with Cb., the ciliated

border; Ov,, primary dorsal cesophageal “ valve” with the corresponding

ventral small “valve ” Ovya.

Fig. 6. Transverse section of the anterior region of the mid-gut. E., epithelium;

Cb., ciliated border; Ey, epithelium of an adjacent fold which has also been

cut through; Ml; and Mls, the longitudinal muscles of the dorsal and

ventral longitudinal grooves of the mid-gut; Mc., circular muscles;

Mp., peritrophic membrane; Pm., peritoneal membrane; Mg., globules of

mucilaginons secretion.

PratE XXX.

Fig. 7. Transverse section through the junction of mid- and hind-guts. E., epithe-

lium; .Ml., longitudinal muscles; Pm., peritoneal membrane; Mp.,

peritrophic membrane; Ch., chitious lining of the hind-gut; Mt., Mal-

pighian tubes; Cd) and Cds, common ducts of several Malpighian tubes.

Fig. 8. Transverse section of the rectum. E., band of large epithelial cells, of which

there are six; between each two bands there is a non-epitheliated

interspace I, where the chitmous intima Ch. becomes corrugated and is

~~usely applied to the basement-membrane Mb.; Tr., trachea; M., muscular

layer.

Fig. 9. Section through small intestine at the orifice of the rectum. F., one of the

six projections or valves which shut off the small intestine from the rectum ;

H., epithelium secreting the chitinous intima Ch.; Mce., circular muscle;

M1., longitudinal muscle; Tr., trachea.

IP (Aa) LO PIZeeae

ATL.Searle delet hth. Huth, imp.

NEW AFRICAN TERRESTRIAL AND FLUVIATILE SHELLS.

Pea Ss OZ eos

AH.Searle delet lith.

NEW AFRICAN TERRESTRIAL AND FLUVIATILE SHELLS.

ON NEW 'FERRESTRIAL AND FLUVIATILE SHELLS, 183

12. Diagnoses of new Species of Terrestrial and Fluviatile

Shells from British and German East Africa, with the

Description of a new Genus (Mussora) from the Eusso

Nyiro River, B.E. Africa. By H. B. Preston, ¥.Z.8.

[Received September 27, 1911 : Read November 21, 1911. |

(Plates XXXI. & XXXIL*)

The species described in the present paper were all, with one

exception, collected by Mr. Robin Kemp during his more recent

travels in British East Africa. As they represent only a very small

portion of the large number of species sent home by Mr. Kemp,

it will be readily seen what a vast field there is for investigation

in the mollusean fauna of this almost hitherto unknown concho-

logical province.

ENNEA MicrostriavA, sp. n. (PI. XXXI. fig. 7.)

Shell scarcely rimate, ovate, edentulate, moderately thin, cream-

coloured; whorls 6, the first four small, regularly increasing, the

fifth large in proportion, gibbous, the sixth proportionately

longer though not quite so broad, marked especially on the fifth

whorl with oblique, distant, regular growth-lines, and sculptured

with closely-set, microscopic, serateh-like strie ; suture impressed,

closely and rather finely crenellate below ; umbilical area

represented by a somewhat broad depression, sloping to an

extremely narrow, elongate, and shallow fissure ; columella rather

vertically descending in a gentle curve ; labrum narrowly out-

wardly expanded, very slightly reflexed ; aperture subquadrate.

Alt. 7°5, diam. maj. 5 mm.

Aperture: alt. 2, diam. 2 mm.

Hab. Dar-es-Salaam, German East Africa (Connolly).

NATALINA PERMEMBRANACEA, sp.n, (PI. XXXI. figs. 20, 204,

20 b.)

Shell perforate, suborbicular, with almost planulate spire,

membranaceous, pale brownish horn-colour ; whorls 5, marked

with closely-set, oblique, arcuate, transverse wrinkles ; suture

impressed, very narrowly margined below ; umbilicus moderately

narrow, deep; columella outwardly expanded above, vitreous,

descending in an oblique curve, diffused above into a thin, glassy,

ill-defined callus which reaches the upper margin of the labrum ;

labrum membranaceous, slightly retlexed, receding below, projecting

above; aperture ovate.

Alt. 6°25, diam, maj. 14°25, diam. min, 11°75 mm,

Aperture : alt. 6°25, diam, 7°25 mm,

Hab. Kigezi, extreme 8.W. Uganda, at an altitude of 6000 ft.

(Robin Kemp).

* For explanation of the Plates see p, 193,

184 MR. H. B. PRESTON ON NEW

AFRICARION KEMPI, sp.n. (PI. XXXI. fig. 3.)

Shell subovate, with moderately depressed spire, yellowish

brown; whorls 3, marked with arcuate, wrinkle-like growth-

ridges; suture impressed, narrowly margined below; columella

vertically descending above, somewhat obliquely curved below ;

labrum simple, projecting in front, receding above and below

aperture rather squarely ovate,

Alt. 7, diam. ma}. 14, dtam. min. 10°75 mm.

Aperture: alt. nearly 7, diam. 8 mm.

Hab. Between Entebbe and Mbarara, S.W. Uganda (fobim

Kenp).

AFRICARION MICROSTRIATA, Sp.n. (PI. XXXT. figs. 4, 4a.)

Shell rather small, thin, moderately globose, with small and

depressed spire, pale yellowish horn-colour ; whorls 3, the first

two very small, the last proportionately very large, marked with

transverse, arcuate growth-wrinkles and microscopic, closely-set,

wavy, spiral striz; suture impressed; base of shell rather

inflated ; columella descending in a strong curve; labrum acute,

receding below and sharply above, very prominently projecting

in front; aperture very broadly and somewhat compressedly

sublunate.

Alt. 5°75, diam. ee 11-25, diam. min. 8° mm.

Aperture : alt. 5°75, diam. 6: 25 mm.

Hab. Between Mbarara and Kigezi, extreme 8.W. Uganda

(Robin Kenp).

AFRICARION MICROGRANULATA, sp.n. (PI. XXXT. figs. 5, 5a.)

Shell differing from 4. microstriata in its much finer sculpture,

the spiral strie being very much finer and still more closely set ;

inoreover, they are crossed by fine, oblique, transverse striz, thus

presenting under the microscope a finely granular appearance.

The present species is also rather larger, the last whorl is some-

what more inflated, and the labrum less prominently projecting

in front; the aperture is much higher in proportion to its breadth,

and the columella is even more curved than is the case with

A. microstriata.

Alt. 8, diam. maj. 12, diam. min. 9 mm.

Aperture : alt. 8, diam. 7 mm.

Hab. Wigezi, extreme 8.W. Uganda, at an altitude of 6000 ft.

(Robin Kemp).

VirRINA COMPACTA, sp.n. (Pl. XXXII, fig. 11.)

Shell semiorbicular, not very thin, pale brown; whorls 27,

rapidiy increasing, smooth but for radiate lines of growth, the

last whorl subangulate at the periphery; suture impressed,

slightly crenellate and narrowly margined below; base of shell

rather inflated; columella descending in a very gentle curve,

narrowly outwardly reflexed and diffused into a light callus whieh

veaches the upper margin of the labrum; Jabrum simple, the

margins converging; aperture broadly subovate.

TERRESTRIAL AND FLUVIATILE SHELLS, 185

Alt. 4:5, diam. maj. 9, diam. min, 7 mm.

Aperture : alt. 4, diam. 4°75 mm.

Hab. Between Mbarara and Kigezi, extreme 5.W. Uganda

(Robin Kemp).

ZINGIS KEMPI, sp.n. (Pl. XXXII. fig. 14.)

Shell small, perforate, thin, rather depressedly turbinate,

greenish olive; whorls 33, the earher whorls somewhat pitted,

the later sculptured with “oblique, 1 radiate growth-plice, the last

whorl subangulate at the periphery ; suture impressed ; base of

shell slightly inflated ; umbilicus narrow, deep, slightly overhung

by the outward expansion of the columella ; columella somewhat

br ‘oudly outwardly expanded, descending in a very slight curve ;

labrum acute, simple, receding below; aperture obliquely

sublunate.

Alt. 2:5, diam. maj. 5°25, diam. min, 4°5 mm.

Aperture : alt. 2°25, diam. 2 mm.

Hab. Between Mbarara and Kigezi, extreme 8.W. Uganda

(Robin Kemp).

ZINGIS PAPYRACEA, sp.n. (Pl. XXXIT. fig. 15.).

Shell rimate, very thin, globosely turbinate, pale yellowish

horn-colour; whorls 5, the first four regularly increasing, the

last large, inflated, marked with transverse growth-lines, and

minute, wavy, transverse striz ; suture impressed, very narrowly

margined below ; perforation very narrow, almost covered by the

outward expansion of the columella; columella outwardly

reflexed and vertically descending above, curved below; labrum

thin, very slightly retlexed, especially towards the base; aperture

very broadly and compressedly sublunate.

Alt. 8°25, diam. maj. 13°5, diam. min. 11°25 mm,

Aperture: alt. 6°75, diam. 6°25 mm.

Hab. Kigezi, extreme 8.W. Uganda, at an altitude of 6000 ft.

(Robin Kemp).

ZINGIS PLANISPIRA, sp.n. (Pl. XXXIUI. fig. 16.)

Shell perforate, thin, almost membranaceous, very depressedly

tarbinate, semiorbicular, pale yellowish brown, painted with a

super-sutural line of dull reddish purple, which appears on the

last whorl as a narrow, super-peripheral band; whorls 43, some-

what rapidly increasing, the last large and slightly descending in

front, sculptured throughout with radiate wrinkles; suture

impressed ; umbilicus moderately wide, deep ; columella dese ending

in a Sharp curve, outwardly rather bro adly expanded; labrum

acute, reflexed throughout ; aperture very broadly and depressedly

sublunate.

Alt. 6°75, diam. maj. 13°75, diam. min, 11 mm,

Aperture; alt. 5, diam. 6 min.

HTab, Between Entebbe and Mbarara, Uganda (Pobin Kemp),

In shape resembling the Kastern Helicoid genus Planispira,

186 MR. H. B. PRESTON ON NEW

THAPSIELLA MILLESTRIATA, sp. n. (Pl. XXXII. fig. 12.)

Shell perforate, very depressedly turbinate, thin, pale yellowish

brown; whorls 47, regularly and rather rapidly increasing, marked

throughout with radiate and very minute and closely-set, wavy,

spiral strize; suture impressed, margined below; umbilicus narrow,

deep, partly concealed by the outward expansion of the columella ;

columella outwardly expanded above, very obliquely descending,

a well-defined, outwardly projecting callus reaching from it to the

upper margin of the labrum; labrum thin, acute, receding below ;

aperture very broadly and compressedly sublunate.

Alt. 4, diam. maj. 8, diam. min. 6°75 mm.

Aperture: alt. 3°5, diam. nearly 4-25 mm.

_ Hab. Near Kigezi, extreme S.W. Uganda, at an altitude of

6000 ft. (obi Kemp).

THAPSIELLA OPPOSITA, sp. n. (Pl. XXXII. fig. 13.)

Shell differing from Helix zanguebarica Craven * in being

spirally sculptured throughout and in having no transverse striz

on the spire, whereas 17. zanguebarica has no spirals on the spire

but is strongly transversely striate, the basal strize are not so

apparent as those on H. zanguebarica; in other respects there is

a great resemblance between the two species.

Alt. 1°75, diam. maj. 1:5 mm.

Hab. Mt. Kenangop, Aberdare Range, British Hast Africa

(Robin Kemp).

It may be as well to mention here that through the courtesy

of Mr. KE. A. Smith I have microscopically examined the type

specimen of Mr. Craven’s species in the British Museum, which

is, as Stated above, basally spirally striate, though no mention is

made of this important character in his description.

KALIELLA CONSOBRINA, sp.n. (Pl. XX XI. fig. 11.)

Shell rimate, conical, pale brownish horn-colour; whorls 54,

sculptured with fine, oblique, slightly wavy, closely-set, transverse

strie, the last strongly carinate at the periphery; base of shell

rather inflated, marked with lines of growth and fine, wavy, spiral

strie ; suture impressed, narrowly but strongly callously margined

above; umbilicus reduced to a mere chink; columella outwardly

expanded and very vertically descending above, oblique below ;

labrum simple; aperture angularly sublunate.

Alt. 3°75, diam. maj. nearly 4, diam. min. 3°5 mm.

Aperture: alt. 1:25, diam. 1:25 mm.

Hab. Between the Igembi Hills and Nyeri, British East Africa

(Robin Kemp).

KALIELLA DEPAUPERATA, sp.n. (PI. XXXI. figs. 13, 134.)

Shell differing from A’. consobrina in its smaller size, it having

one whorl less, and in its proportionately broader shape, in being

very bluntly carinate at the periphery, and in its rather lighter

colour; moreover, it lacks the spiral basal strize of that species.

* Proc, Zool. Soc. London, 1880, p. 217, pl. xxii. figs. 4 a, 4, ¢.

TERRESTRIAL AND FLUVIATILE SHELLS. 187

5, diam. min. 2°5 mm.

Alt. 2, diam. maj. 2°7

Igembi Hills and Nyeri, British East Africa

TTab. Between the

(Robin Kemp).

KALIELLA KIGEZIENSIS, sp.n. (Pl. XXXI. fig. 14.)

Shell allied to K. consobrina, but imperforate, narrower in

form, with proportionately higher aperture and rather finer

transverse sculpture; the columella also descends vertically

throughout its whole length.

Alt. 3, diam. maj. 3°25, diam. min. 3 mm,

Aperture: alt. 1°25, diam, 1 mm.

Hab. Between Mbarara and Kigezi, extreme S.W. Uganda ;

also at various points in the neighbourhood of Kigezi itself

(Robin Kemp).

KALIELLA IREDALEI, sp.n. (Pl. XXXI. fig. 12.)

Shell rimate, small, conically turbinate, light reddish brown ;

whorls 6, rather convex, the last not carinate at the periphery,

sculptured with very fine, somewhat oblique, transverse strive ;

base of shell marked with lines of growth and sculptured with

slightly distant, fine, wavy striez; suture well impressed, not

margined ; umbilicus very narrow, partly concealed by the outward

expansion of the columella; columella outwardly expanded,

vertically descending in a gentle curve above, oblique below ;

labrum simple; aperture sublunate.

Alt. 2°5, diam. maj. 2°75, diam. min. 2°5 mm.

Hab. Between the Igembi Hills and Nyeri, British Kast Africa

(Robin Kemp).

The very characteristic sculpture of this pretty little species is

only visible under the microscope.

SITALA IREDALEI, sp. n. (Pl. XXXI. fig. 15.)

Shell rather acuminately turbinate, with convex base, pale

greyish yellow ; whorls 6, somewhat rapidly increasing, the last

inflated, sculptured with microscopic, spiral striz and oblique,

transverse riblets or creases; suture impressed; columella

vertically descending, slightly angled below ; aperture ovate.

Alt. 4°25, diam. maj. 4°25 mm.

Aperture: alt. 2, diam. 1°75 mm.

Hab. Mt. Kenangop, Aberdare Range, British East Africa

(Robin Kemp).

TTRACHYCYSTIS IREDALEI, sp.n. (Pl. XXXII. figs. 8, 8a, 80.)

Shell small, depressedly suborbicular, almost planulate, reddish

brown; whorls 3, the apical whorl large, smooth, the remainder

sculptured throughout with rather fine and closely-set, obliquely

arcuate, transverse costule, between which ocevr very fine,

transverse striz, crossed by extremely fine spirals, both these

last and the transverse strie being only visible with the aid of

the microscope; suture well impressed ; umbilicus very wide and

188 MR, H. B. PRESTON ON NEW

rather deep; columella very oblique; labrum simple; aperture

broadly sublunate.

Alt. °75, diam. maj. 2, diam. min. 1:75 mm.

Hab. Between the lgembi Hills and Nyeri, British Kast Africa

(ftobin Kemp).

LEUCOCHILOIDES CHANLERENSIS, sp.n. (Pl. XX XI. fig. 16.)

Shell small, rimate, cylindrically fusiform, slghtly shining,

reddish brown ; whorls 5, regularly increasing, the last ascending

in front, marked with oblique, transverse growth-lines; suture

well impressed ; umbilicus very narrow; columella curved ; labrum

rather narrowly expanded, whitish, not reflexed; aperture sub-

circular, bearing a single, nodulous denticle just below the point

of insertion of the labrum with the parietal wall.

Alt. nearly 4, diam. maj. 1°75 mm.

Aperture: alt. °75, diam. nearly -75 mm.

Hab. Chanler Falls, Eusso Nyiro, British Kast Africa (Rodin

Kemp).

LEUCOCHILOIDES IREDALET, sp.n. (PI. XXXII. fig. 18.)

Shell differing from ZL. chanlerensis in its blunter form, more

swollen whorls, deeper suture, more open umbilicus, straighter

columella, and narrower and more erect labrum ; moreover, it lacks

the nodulous denticle which in the present species is replaced by

an erect white denticle situate low down on the parietal wall well

within the aperture, and a smaller erect squarish denticle on the

columella also situate well within the opening.

Alt. 3°5, diam. maj. 1°75 mm.

Aperture: alt. °75, diam. nearly ‘75 mm.

Hab. Kusso Nyiro, British East Africa (Robin Hemp).

LErUCOCHILOIDES soroR, sp.n. (PI. XX XI. fig. 17.)

Very closely allied to Z. chanlerensis and possibly only a variety

of that species; it differs, however, in its larger size, more open

umbilicus, though having a proportionately narrower base, and in

having an additional whorl, the aperture is also more ovate than

is the case in that species.

Alt. 6, diam. maj. 2°25 mm.

Aperture: alt. 1:5, diam. 1°25 mm.

Hab. Chanler Falls, Eusso Nyiro, British East Africa (2obin

Kemp).

LEUCOCHILOIDES GAZIENSIS, sp.n. (Pl. XX XI. fig. 19.)

Shell small, ovately fusiform, pale reddish brown; whorls 5,

convex, marked with fine, oblique, transverse striz ; suture deeply

impressed; umbilical area broadly depressed: labrum with

converging upper margin, white, rather broadly expanded ;

aperture ovate.

Alt. nearly 4°5, diam. maj. 2°25, diam. min. 2 mm.

Aperture: alt. 1, diam. -5 mm.

fab. Gaa, British East Africa (Robin Kemp).

TERRESTRIAL AND FLUVIATILE SHELLS, 189

ALA KENIANA, sp.n. (PI. XXXI. fig. 6.)

Shell minute, cylindrically ovate with very obtuse apex, scarcely

rimate, moderately thin, pale reddish chestnut ; whorls 4, convex,

shouldered above, marked with fine, oblique, transverse striz and

slightly malleated; suture well impressed; columella whitish,

internally broad; labrum erectly reflexed, the margins joined by

a thin a callus ; aperture very broadly inversely auriform.

Alt. 2, diam. maj. 2°25 mm.

Hab. Mt. Kenia, at an altitude of from 6000 to 9000 ft. (Robin

Kemp).

Homorvs IREDALEI, sp. n. (Pl. XX XI. fig. 10.)

Shell moderately large, subulately fusiform, with minute apex,

covered with a yellowish periostracum, on the lower whorls closely,

obliquely, transversely banded with dark blackish purple; whorls

10, flattened, the first minute, the second proportionately large,

the pom ainder regularly increasing, smooth but for lines “of

growth ; suture lightly impressed, faintly crenellate below ;

columella short, gently curved, abruptly truncate, diffused above

into a thickish, well-defined callus which reaahies the upper

margin of the labrum ; ; labrum simple; aperture elongately ovate.

Alt: 31°25, diam, maj. 9:25 mm.

Aperture: alt. 8°5, diam. 4°5 mm.

Hab. Between Mbarara and Kigezi, extreme 8.W. Uganda

(Robin Kemp).

SuccinEA KEMPI, sp.n. (Pl. XXXII. figs. 2, 2a.)

Shell ovate, opaque, cream-coloured ; remaining whorls 2, the

first very small, the last comparatively very large, marked only

with lines of growth ; suture impressed, broadly margined below ;

columella very obliquely curved ; labrum simple, acute ; aperture

dilated, ovate.

Alt. 7°25, diam. maj. 5:5, diam. min. 3:25 mm.

Aperture: alt. 6, diam. 4 mm.

Hab. Between Entebbe and Mbarara, 8.W. Uganda (Robin

Kemp).

A remarkable form, easily recognizable by its minute spire and

enormously large last whorl.

SUCCINEA PRINCEI, sp.n. (Pl. XXXII. figs. 3, 3.)

Shell ovately fusiform, very thin, pale yellowish horn-colour;

whorls 22, the last very long, polished, smooth but for lines of

growth ; columella arched ; aper ture dilated, very lar ge.

Alt. 7°25, diam. ee 3°D mm.

Aperture: alt. 5, diam. 2°5 mm.

Hab. Nakuru, British East Africa (Robin Kemp),

BLAUNERIA EXSILIUM, sp. n. (Pl. XXXI. fig. 8.)

Shell subulate, thin, pale yellowish horn-colour; whorls 8,

smooth but for lines of growth, polished, shining, flat; suture

190 Mi. H. B. PRESTON ON NEW

linear, broadly margined below; columelia arched, white,

extending into a thick, white, narrow, well-defined callus, bearing

a single plait above, obliquely truncate below; labrum acute,

whitish ; aperture narrowly inversely auriform.

Alt. 4°5, diam. maj. 1:25 mm.

Aperture: alt. 1-5, diam. -5 mm.

Hab. Gazi, British Hast Africa (Robin Kemp).

ANCYLUS KEMPI, sp. n. (PI. XXXI. figs. 2, 2 a.)

Shell rectangularly ovate, depressed, somewhat sinuous on the

left side, with subcentral apex, olive-brown, marked with concentric

growth-lines, and indistinctly sculptured towards the margin with

faint, radiate striz, which are more visible on the interior surface

of the shell; margin acute; interior of shell polished, shining,

ale brown.

Alt. 1:25, diam. maj. 4°25, diam. min. 3°25 mm.

Hab. Kigezi, extreme 8.W. Uganda, at an altitude of 6000 ft.

(Robin Kemp).

LIMN#A KEMPI, sp.n. (Pl. XXXII. fig. 1.)

Shell small, ovately fusiform, thin, pale reddish brown ; whorls 3,

the last large and somewhat elongate, marked with fine, silky,

transverse growth-strie; suture well impressed; columella

obliquely descending, not twisted or arched above, diffused into a

very thin, well-defined, parietal callus which reaches the upper

margin of the labrum; Jabrum simple, acute, somewhat dilated

at the base; aperture very elongately ovate.

Alt. 7-5, diam. maj. 4 mm.

Aperture: alt. 5, diam. 2°5 min.

Hab. Kisumi, Lake Victoria Nyanza, British East Africa

(Robin Kemp).

PLANORBIS KIGEZIENSIS, sp.n. (Pl. XXXII. figs. 5, 5a, 5 6.)

Shell small, depressed, suborbicular, with concave spire, thin,

pale greenish grey; whorls 4, regularly and rather rapidly

increasing, the last obtusely angled above, somewhat sharply

angled below, sculptured with fine, closely-set, arcuate, transverse

striz; suture well impressed; base of shell not very convex ;

umbilical depression very wide, shallow; columella descending

very obliquely and diffused above into a projecting callus which

reaches the upper margin of the labrum ; labrum simple, acute,

projecting in front, receding above and below, the margins

converging ; aperture subelliptical.

Alt. 1, diam. maj. nearly 3°5, diam. min. 3 mm.

Hab. Kigezi, extreme S.W. Uganda, at an altitude of 6000 ft.

(Robin Kemp).

PLANORBIS SPERABILIS, sp. n. (Pl XXXII. figs. 4, 4a, 4 0.)

Shell very small, depressedly orbicular, thin, greyish-yellow

horn-colour; whorls 3, rather rapidly increasing, marked with

TERRESTRIAL AND FLUVIATILE SHELLS. 191

somewhat strong, radiate growth-lines; base of shell slightly

concave ; labrum receding below, the mar gins joined by a thin

callus ; aperture subcircular.

Alt. nearly ‘5, Sa maj. nearly 2, diam. min, 1°) mm.

Aperture : alt. 25, diam. ‘25 mm.

Hab. Gazi, British’ East Africa (Robin Kemp).

SEGMENTINA EUSSOENSIS, sp.n. (Pl. XXXII. figs. 6, 6a, 66.)

Shell depressedly orbicular, with concave spire and almost

planulate base, pale yellowish wax-colour, polished, somewhat

shining; whorls 43, rapidly i increasing, the last large and rounded,

angled above and bluntly carinate at the per iphery, below which

ae base of the shell is nearly flat; suture rather lightly impressed,

margined below; umbilicus wide, shallow; labrum thin, 1 rapidly

receding below ; aperture depressedly and angularly sublunate.

Alt. 1°5, diam. Ea 5°25, diam. min. 4°75 mm.

Aperture: alt. 1°25, Santi: 1 mm.

Hab. Chanler Falls, Eusso Nyiro, British East Africa (Robin

Kemp).

SEGMENTINA KEMPI, sp.n. (Pl. XXXII. figs. 7, 7a, 7b.)

Shell small, suborbicular, planulate above and below, with

concave spire, Avhite: polished, shining; whorls 4, the first three

small, regularly increasing, the last very large, founded above,

sharply angular below, indistinctly sculptured with microscopic,

silky, transverse strie ; suture impressed, irregular; umbilicus

moderately narrow, and deep; columella obliquely descending,

angled below, extending above into a parietal callus which joins

the upper or sutural margin of the last whorl well within the

shell; labrum acute, receding below ; aperture triangular.

Alt. 1, diam. maj. 3°75, diam. min. 2°25 mm.

Hab. Kigezi, extreme S.W. Uganda, at an altitude of 6000 ft.

(Robin Kemp).

VIVIPARA RUBICUNDA, v. Martens, var. KISUMIENSIS, var. n.

(Pl. XXXIT. fig. 9.)

Shell differing from the typical form in its larger size, darker

colour, and in ‘being proportionately higher and narrower, the

apertur e is also more ovate and the peristome edged with black.

Alt. 28, diam. maj. 15°75, diam. min. 12 mm.

Aperture: alt. 10, diam. 7 mm.

Hab. Kisumi, Lake Victoria Nyanza, British East Africa

(Robin Kemp).

ASSIMANIA AURIFERA, sp.n. (Pl. XXXI. fig. 9.)

Shell turbinate, perforate, thin, semitransparent, dark brownish

horn-colour, minutely and densely freckled with golden yellow;

whorls 53, regularly increasing, shouldered above, convex, very

minutely spirally striate; suture well impressed ; umbilicus

moderately wide, deep; columella very oblique, obtuse-angled,

192 MR. H. B. PRESTON ON NEW

slightly outwardly reflexed; labrum simple, acute; aperture

irregularly roundly ovate.

Alt. 3:75, diam. maj. 2°75 mm.

Aperture: alt. 1-5, diam. 1 mm.

Hab. Gazi, British East Africa (Robin Kemp).

Evssora, gen. n.

Shell solid, conical, resembling Assemania, but without oper-

eulum, though fluviatile in its habits.

Type of genus, 4. inopina.

The genus, as above stated, bears a great resemblance to Assi-

mania, but though I have been able to examine a jarge number of

specimens collected alive, and which still contained the decaying

animal, J have been unable to find any trace of operculum : more-

over, on physiographical grounds alone, I should be loth to refer

it to that genus, which is essentially a coast form, whereas the

po at which the present specimens were collected is at least

375 miles from the nearest point on the African sea-board, and

it is extremely probable, judging by the maps at my disposal,

that the connection by water between the locality where the

present species was collected and the sea is very much more than

that distance ; the Eusso Nyiroafter flowing through Lake Lorian,

is at present believed to flow out of that lake as the Wakes

River, a tributary of the Juba, which it joins not far from its

mouth and which forms the north-eastern boundary-line between

British East Africa and Italian Somaliland.

Pending the examination of the animal, which in the specimens

sent home by Mr. Kemp was not in sufficiently good condition

for dissectional purposes, I propose to place the genus temporarily

in the vicinity of the Assimanieide.

KUSsOIA INOPINA, sp. n. (PI. XXXIT. fig. 10.)

Shell small, rimate, turbinately conic, dark reddish brown ;

whorls 54, regularly increasing, flattish, except the last, which is

somewhat globose, marked only with transverse growth-lines ;

suture impressed ; umbilicus very narrow, almost covered by the

dilation of the columella; columella descending in a rounded

curve, rather erectly dilated, diffused above into a thin, parietal

callus; labrum simple; aperture subovate.

Alt. 3°5, diam. maj. 2°5, diam. min. 2 mm.

Aperture: alt. 1°75, diam. 1°25 mm.

Hab. Banks of the Eusso Nyiro River, Br “us East Africa

(Robin Kemp).

SPHARIUM KIGEZIENSIS, sp. n. (Pl. XX XI. figs. 1, 1 a.)

Shell very small, triangularly ovate, yellowish red ; both valves

closely concentrically striate, the striz being rather coarser

towards the umbonal region; umbones rather large, not prominent ;

dorsal margin sharply arched ; ventral margin gently rounded ;

anterior side rather abruptly descending ; “posterior side sub-

rostrate, somewhat angularly rounded ; cardinal tooth in right

TERRESTRIAL AND FLUVIATILE SHELLS. 193

valve small, oblique, slightly notched anteriorly ; cardinal teeth

in left valve very small, converging; lateral teeth in right valve

very weak, in left valve weak, somewhat curved; scars very

lightly impressed ; interior of shell somewhat granular.

Long. 3, lat. 3°5 mm.

Hab. Kigezi, extreme S.W. Uganda, at an altitude of 6000 ft.

(Robin Kemp).

EXPLANATION OF THE PLATES.

Proate XXXI.

Fig. 1. Spherium kigeziensis, X 5.

Was > is hinge, X 5.

2&2a. Ancylus kempi, X 5.

_ 3. Africarion kempi, nat. size.

4, x microstriata, X 14.

4a. 4 5 sculpture, X 12.

5 a6 microgranulata, X 1.

5a. a S sculpture, X 12.

6. Alea keniana, X 10.

7. Ennea microstriata, X 2%.

8. Blauneria exsilium, X 5.

9. Assimania aurifera, X 6.

10. Homorus iredalei, nat. size.

11. Kaliella consobrina, X 6.

12. ‘A iredalei, * 6.

13. a depauperata, X 6.

Hsia, 55 i sculpture, X 12.

14. 5a higeziensis, X 6.

15. Sitala iredalei, X 4.

16. Leucochiloides chanlerensis, X 7.

17. os soror, X 4.

18. a iredalei, X 7.

; 53 gaziensis, X 6.

20. Natalina permembranacea, X 1s.

PAO aR Er spire, X 1}.

20Ibe" ss % base, X 15.

PLatE XXXII.

Fig. 1. Limnea kempi, X 3.

2 &2a. Suecinea kempi, X 25.

3&3a. 5 princei, X 25.

4, Planorbis sperabilis, X 8.

La. a 3 spire, X 8.

Ab. a - base, X 8

5. 4 kigeziensis, X 6.

Site 5 . spire, X 6

55. = base, X 6.

6. Segmentina eussoensis, X 3.

6a. 5 H spire, X 3.

6 b. Pe a base, X 3.

if i kempi, X 6.

7 ” ” spire, x 6.

“Aloe fs »» base, X 6.

8. Trachycystis iredalei, X 8.

8 x = spire, X 8.

8 + os base, X 8.

9. Vivipara rubicunda, var. kisumiensis, nat. size.

10. Hussoia inopina, X 5.

11. Vitrina compacta, X 2.

12. Thapsiella millestriata, X 2.

13. ES opposita, X 8.

14. Zingis kempi, X 4.

15. 4, papyracea, X 13.

16. » planispira, X 13.

Proc. Zoot. Soc.—1912, No. XIII. 13

194 MR. F. E. BEDDARD ON A

13. Contributions to the Anatomy and Systematic Arrange-

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

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

[Received November 1, 1911: Read November 21, 1911.]

(Text-figures 19-30.)

II]. On a New Genus or Tarpeworms (Ofiditenia) FROM THE

BustarD (Lupodotis kort).

An example of the South-African Bustard (Hupodotis kort)

on April 27th, 1911, voided a number of white curiously-shaped

bodies, the nature of which I did not recognise until they were

submitted to microscopic examination. The shape of these bodies

may be learnt by an inspection of text-figure 19. They are

separate mature proglottids of a tapeworm, of which I obtained

complete specimens later (on May 7th), when the bird died. Since

the Bustard was acquired by the Society on October Ist, 1910, 16

seems likely that the infection did not take place in the Society’s

Gardens, but that the parasites were already present when the

bird arrived.

Text-fig. 19.

Ripe and detached proglottids of Otiditenia ewpodotidis.

A most useful and presumably comprehensive list of the tape-

worms of birds has been quite recently published by Prof. Fuhr-

mann*. From this it would appear that only the following species

have been found in the Otidiformes, and that none at all have been

met with in Hupodotis kori—if it has been examined from this

* “Die Cestoden der Vogel,” Zool. Jahrb. 1908, Suppl.-Bd. x.

NEW AVIAN TAPEWORM. 195

point of view. The species are /diogenes otidis, Chapmannia lapica,

Hymenolepis villosa, H. tetracis, and H. ambiguus. There is no

doubt that the species which forms the subject of the present

communication to the Society is none of these; nor can I reconcile

its characters with those of any other genus of 'Tapeworms of the

group Tetracotylea (of which group it is undoubtedly a member)

which have been described up to the present. I shall, however,

defer the attempt to fix its systematic position until I have

described its anatomical characters.

I have examined feur complete specimens of this worm, besides

some fragments which may or may not have belonged to those

specimens. The largest was about 3 inches in length, and the

greatest diameter of the posterior segments was 4 mm. ‘The

diameter of the body gradually increases from the head up to the

point of greatest diameter, and then remains the same ; there is

not, as in so many tapeworms, a narrow anterior region of the

body, which suddenly widens within the limits of two or three

proglottids. The general shape is thus much like that of certain

species of Bertiella, for example of B. cercopithect*. There is,

however, no further resemblance between the present genus and

Bertiella in external form. The worm is white in colour; but

there is some black pigment in the head behind the suckers, and

the ripe proglottids are of a brick-red hue.

The scolex of this new worm—for which I propose the generic

and specific names of Otiditenia eupodotidis, as indicative of its

habitat—is not particularly small nor, on the other hand, con-

spicuous by its large size. I have not found the least trace of

hooks anywhere on the suckers. The head seems be completely

unarmed as to the suckers, but the rostellum has hooks.

The four suckers are, as is so often the case, extremely mobile,

and can be protruded a long way from the head as cylindrical

processes. In this case they look rather forwards than outwards.

When retracted and in a normal condition the suckers look out-

wards and slightly upwards. When fully protruded the suckers

give to the head of this worm an appearance very like that of the

genus of Tetraphyllidea, Cylindrophorus *.

I have investigated the structure of the rostellum and lateral

suckers by transverse sections through the head. There is nothing

remarkable about the structure of the suckers. The rostellum,

however, demands some consideration. When the worm is ex-

amined with a lens or with a low power only of the microscope,

the appearance presented is that of a median sucker such as occurs

sometimes, though, as it is stated by Kiichenmeister ¢, not always,

in Tenia saginata. The hooks are, in fact, so minute that I was

unable to detect them when studying the worm in that fashion.

Nor can they be detected save with high-powered lenses. The

varying condition of the retraction of the median sucker, or rather

* See Beddard, P. Z.S. 1911, p. 640, text-fig. 151.

+ Bronn’s ‘ Klassen u. Ordnungen des Thierreichs,’ Bd. iv. Taf. 42. fig. 14,

{ ‘Die Parasiten des Menschen,’ Leipsic 1881, p. 140, pl. ii. fig. 3.

13*

196 MR. F. E. BEDDARD ON A

rostellum, renders it apparently absent or more feebly developed

in some individuals than in others. And there is here a resem-

blance to the otherwise not closely allied Tenia saginata.

This median sucker is, indeed, a not much more developed

structure than the rudimentary rostellum of Z’enia saginata. In

this latter species Railliet* speaks of “‘le rostre faisant défaut et

se trouvant remplacé fréquemment par une légere dépression ” ;

while Leuckart 1, in defining the species, states that “‘the head has

a flattened crown with a pit-like hollow in the middle.” On the

other hand, Kiichenmeister, while emphasizing the occasional

absence of any such depression in both text and figure, also

figures { an organ very like a sucker, and certainly more than a

mere depression. ‘The structure, however, is not, according to

Leuckart §, a sucker, but a rudimentary rostellum with a distinct

lenticular muscular mass. Leuckart’s figure of a section through

this ““sucker” is copied in Bronn’s ‘'Thierreichs’ (Bd. iv. p. 1203).

The accompanying text-figure (text-fig. 20) shows a median

longitudinal section through the scolex of Onditenia, illustrating

the: rudimentary rostellum and enabling it to be compared with

the adjacent suckers. The simplicity ‘of the really sucker-like

rostellum will be apparent from this figure, which represents

the rostellum in a contracted state withdrawn into the scolex.

Inasmuch as the sete border the rostellum in a double circle,

they are not visible in such a preparation, only one or two being

seen in each section at the two corners of the rostellum. The

rostellum itself is obviously (to my mind) no more, or very little

more, developed than that of Tcenia saginata, already referred to;

its muscular layer is not thicker than that of the suckers, as will

be seen from my figure. This obviously contrasts greatly with

complicated and more typical rostella ||.

It will be observed, moreover, that the floor of the cavity into

which the rostellum is retractile—which floor constitutes the

rostellum—is hardly, if at all, thicker than the lateral walls of the

cavity; that the whole structure, in fact, does not greatly differ

froma sucker. It may be borne in mind that this is a possible

origin of the rostellum, though the theory that it is the meta-

morphosed remains of a pharynx has been put forward. On the

other hand, the nuclei are more densely packed in the rostellum

than in the suckers, as the figure referred to shows. This

difference is, however, perhaps partly due to the more contracted

condition of the rostellum, and possibly also to its greater con-

tractility. Again, the layer of chitin covering the rostellum is

very much thicker than that which lines the suckers—the latter

being, indeed, inappreciable. ‘There is some black pigment in the

head behind the suckers.

My preparations unfortunately do not allow me to give any

* “Tyraité de Zoologie Médicale et Agricole,’ ed. 2, Paris 1895, p. 232.

+ ‘The Parasites of Man’ (Engl. ed. transl. by Hoyle), Edinburgh 1886, p. 407.

{t Loe. cit. pl. vi. fig. 8.

is Loe. cit. p. 435.

|| Cf, e.g.) Bronn’s ‘ Thierreichs,’ Bd. iv. figs. 56, 57, 58, pp. 1826-1328.

NEW AVIAN TAPEWORM., 197

accurate account or figures of the rostellar hooks. I can only

say that they are curved and hook-like, not straight, and very

numerous, certainly at least 150 in number, and probably more,

and that they are disposed in an alternating fashion, forming a

double crown.

Text-fig. 20,

a : oa

Bea Pad

Longitudinal section through scolex of Otiditenia ewpodotidis.

R. Rudimentary rostellum (the hooks are too small to appear). S. One or

the suckers. NV. Lateral nerve-cord of one side.

The longitudinal section from which the annexed text-figure of

the rostellum was drawn also shows very well the existence of

a neck in this tapeworm. It varies much in length, though it

is never longer than the length of three or four proglottids

immediately following upon it. On the other hand, it is some-

times very short, not more than the length of the first proglottid.

If this had not been observed in the living worm, where the neck

is much more distinct, as such, from the proglottids than in aleohol-

198 MR. F. E. BEDDARD ON A

preserved material, it might easily be supposed that this species

was characterised by the absence of a neck. Such a mistake may

have been made with other species.

Text-fig. 21.

Two examples of Otiditenia eupodotidis differing slightly in progressive

increase of size of proglottids.

The general character of the segmentation and the form of the

individual proglottids is shown in text-figure 21, which represents

two individuals which are slightly different in these characters.

In one of them the segments increase in length more rapidly than

in the other; but in neither are the posterior segments longer

than they are broad. Nor do they even get to be so long, except

in the case of the detached proglottids. As the segments advance

towards maturity the projection of one segment over that which

follows is seen to be more marked, and it forms quite a sharp-edged

frill, as is shown in the drawings referred to, The completely

NEW AVIAN TAPEWORM. 199

mature proglottids are represented in several aspects in text-

figure 19 (p. 194). These proglottids, as there depicted, have

a very peculiar form, and have quite lost the flattened appearance

of the anterior region of the strobila. They are, indeed, almost

spherical in general outline. The projecting frill remains, but its

edges are closely adpressed. A single proglottid presents much

the appearance of a full sack with the lips of the mouth closely

drawn together.

Left-hand figure a ventral view of eight nearly mature proglottids of

Otiditenia eupodotidis.

Right-hand figure a lateral view of the same showing genital pores.

The last external character to be noticed is the position and

character of the genital pores. These are irregularly alternate.

The pores are simple orifices upon the lateral surfaces of the

proglottids, and are not in any way prolonged into a spout-like

outgrowth. They are, however, situated on an area which covers

the whole of the lateral face of the segment, and is slightly raised ;

it is also whiter in colour than the rest of the (rather yellowish)

body. Finally, the segment is longer from before backwards on

the side which bears the genital pore; this inequality of the two

sides of a segment enables the position of a genital pore to be

fixed without actually inspecting the orifice (text-fig. 22).

In transverse sections of the body before the reproductive

organs ave to be seen, except in rudiment, the cortical parenchyma

will be observed to be fully as thick as the medullary parenchyma.

Indeed, anteriorly in the segment before the overlapping frill has

detached itself the cortical parenchyma is actually the thicker.

The longitudinal muscle-fibres seen in this layer are disposed

200 MR. F. E. BEDDARD ON A

singly in the more peripheral regions; more centrally they are

ageregated into bundles, the number of fibres being larger in the

deeper strata. In the largest I have counted 14 or 15 fibres.

Within this layer and bounding the medullary parenchyma is a

iayer of delicate circular fibres. In older proglottids the number

of longitudinal fibres to a bundle may be considerably greater and

the strength of the circular layer has increased. This latter fact

is particularly to be noted in the segments which are nearly ready

for detachment.

The water vascular tubes are the usual four longitudinal trunks,

of which—as is again usual—the dorsal is smaller and has much

thicker walls. The larger ventral tubes are connected by a rather

wide commissural vessel in each segment. The dorsal vessels are

apt to vary a little in their relation to the ventral vessels; but

they are never superposed, ?. e. directly dorsal and ventral to each

other, The dorsal vessel, in fact, lies to the inside of the ventral

Text-fig. 23.

Transverse section through an immature proglottid of Otiditenia eupodotidis.

The larger ventral water vascular tubes lie below and rather to the outside

of the smaller dorsal vessels.

vessel, sometimes absolutely parallel with it and sometimes more

dorsally inclined. The capacity of the large ventral vessel tends

to vary; I have seen it to be almost obliterated for a short space.

In the anterior segments of the body there is much less difference

in calibre between the water vascular vessels than is to be seen

later.

The testes of this tapeworm lie quite posteriorly in the segment,

and thus behind all the other parts of the generative system.

In sagittal sections of young proglottids in which the testes are

mature or nearly so, but in which the uterus is not yet much

NEW AVIAN TAPEWORM. 201

developed, a straight line joining the posterior angles formed by

the outgrowths of the segments dorsally and ventrally is just

in contact with the posterior boundary of the mass of testes.

Anteriorly they are well separated from, though very close to, the

ovaries. ‘The testes, in fact, in the genus described in the present

paper do not straggle over the segment as they are represented to

Text-fig. 24.

Upper figure a horizontal section through four not completely ripe proglottids

of Otiditenia ewpodotidis.

Ov. Ovary. T. Mass of testes. U. Uterus.

Lower figure a portion of a transverse section through a proglottid, showing the

division of the layer of circular muscles (C.m.) to enclose the young uterus.

202 MR, F. E. BEDDARD ON A

do in the allied genus Choanotenia—for example, in the species

Ch. galbula and Ch. infundibulum figured by Cohn *

Text-fig. 25.

Horizontal section through three not fully mature proglottids of

Otiditenia eupodotidis.

d. Dorsal water vascular tube. W. Ventral ditto. 7. Mass of testes.

U. Uterus.

* “Jur Anatomie etc. der Vogelcestoden,’ Nova Acta Ac. Nat. Curios. Ixxix. Be

pl. 31. fig. 40 & pl. 32. fig. 47.

NEW AVIAN TAPEWORM. 203

The testes of Otiditania (text-fig. 24, p. 201) form, in fact, a very

compact mass, which almost suggests a single organ whose com-

pactness and general characters form a very distinguishing mark

of this genus. In the sagittal sections already referred to, the

mass of testes is seen to form a band rather lenticular in outline,

2. é. tapering dorsally and ventrally, which extends without inter-

mission from the dorsal to the ventral muscular body-wall. In

transverse sections of the worm the testicular mass is seen equally

well to form an unbroken dorso-ventral layer occupying the

whole of the central parenchyma between the larger ventral water

vascular vessels, to the outside of which the testes do not extend.

The wall of testes which thus exists—-marking, as has already

been said, the posterior region of the proglottid—is a very thin

one. It is formed of only a single layer or in parts of two layers

of testes.

In their general form the testes are oval. The very small

amount of space occupied by the testes in an antero-posterior

plane is striking ; but it is greater in the less fully than in the

more fully mature proglottids. In the former the antero-posterior

diameter of the group of testes is a little more than one-third of

the length of the segment; in the latter perhaps one-tenth, and

the disproportion becomes even greater in absolutely ripe seg-

ments, where, indeed, the testes finally disappear, being apparently

pressed out of existence by the swelling proglottids.

In such tapeworms as I am acquainted with by first-hand

knowledge the testes are scattered in the medullary parenchyma,

though often closely adpressed ; but there is no particular modi-

fication of the medullary parenchyma in the region occupied by

the testes. The same conditions appear to obtain in many forms

described by others.

In Otiditenia, however, the testes lie in a very distinct cavity,

which has in sagittal sections of the entire proglottid a lenticular

outline already mentioned. This cavity is not everywhere apparent

as actual spaces lying outside of the testes ; it is for the most part

filled up with an interstitial tissue, which separates the individual

testes and in which they lie closely packed together. Although

this interstitial tissue is not very different in appearance from

the medullary parenchyma, it is separated from it by a distinct

boundary-line and is more faintly stained by carmine, and does

not seem to contain any calcareous corpuscles. The distinctness

of the mass of testes, as a whole, from the surrounding regions

of the proglottid will be apparent from text-figure 26 (p. 204).

Here and there the interstitial tissue seems to be replaced by

actual cavities which contain no fluid that takes a stain. They

appear, in fact, to be empty.

Since the testes le quite posteriorly in the segments, it is

obvious that they must be close to the transverse vessels of the

water vascular system, which unite the ventral vessels. ‘The

spaces to which I have referred, I believe, entirely belong to

the transverse water vascular trunks which are large in the

204 MR. F. E. BEDDARD ON A

present species. Though the individual testes appear at first

sight to be actually bathed by the fluid of the excretory tube, a

more careful examination has always shown a layer of interstitial

tissue separating the actual testes from the water vascular spaces.

The testes, however, do project with their covering into the spaces

(see text-fig. 25, p. 202) here and there, and there is thus a very

close anatomical relation between them.

Text-fig. 26.

sit

Sagittal section through three proglottids of Otiditenia eupodotidis.

T. Mass of testes. U. Uterus.

In longitudinal horizontal sections (text-fig. 25) it is obvious

that the testes of Otiditenia form a layer which is, as a rule, only

one thick, though here and there two testes lie one in front of the

other. The distinction between the interstitial tissue separating

NEW AVIAN TAPEWORM. 205

the individual testes and the general medullary parenchyma. is

strikingly obvious in such sections, inasmuch as the whole area

can be seen in a single section extending from one side of the

body to the other. Such sections also show particularly well the

fact that the testes extend rather beyond the large ventral water

vascular vessel on the fore side, where, indeed, they occasionally

nearly touch the outer skin. The extension is not quite so great

on the opposite side, though the mass of testes usually bulges into

the vessel, and, as already mentioned, presents very much the

appearance of lying in it. It is to be noticed also that in these

lateral extensions of the testes the whole mass gets very much

wider from back to front, the testes being three or four deep.

These outgrowths intrench upon the area occupied by the coils of

the vas deferens, and push laterally and flatten out that coil.

The extension of this testicular mass laterally is highly suggestive

of its being a single organ growing as a whole and not merely an

association of separate and quite individualized testes. The inter-

stitial substance which divides the testes presents the appearance

in the direction of its fibrils of having flowed laterally, carrying

with it the enclosed testes. Multiplying the 30 or so testes

which are visible in a horizontal section by the 6 or 7 which are

visible in a sagittal section, we get a total of 200 or so, which may

be slightly increased to include the testes found in the lateral

extensions of the testicular area just referred to.

The ovaries are very obvious in the segments where they occur ;

but they disappear early in the strobila and are represented

thenceforward by mere vestiges. The ovaries are to the fore side

of the segment and closer to that margin of the strobila than the

vitelline gland, to which, however, they are closely contiguous.

They commence on the level of the receptaculum and lie on both

dorsal and ventral sides of that portion of the ovidueal tube,

hence getting a double character—the ventral half (or more

ventral ovary, if we speak of two) is considerably the larger.

They lie posteriorly in the segment, but between the testes and

the uterus. They by no means occupy the whole breadth of the

medullary parenchyma in a sagittal section, as do the testes when

fully developed.

The cloaca genitalis opens on to the exterior directly. There

is no papilla-like process bearing a genital orifice. As already

mentioned, the pores are irregularly alternate in position, and I

have seen as many as six consecutive pores on the same side of

the body. The cavity of this conjoined region of the genital ducts

is rather wide and of some length; the lumen is slit-like to

circular, but the walls never seem to be crumpled so as to give a

star-like appearance to the lumen. The cavity is lined by a very

thick layer of cuticle which stains very deeply in carmine. The

walls of the cloaca genitalis contain at some little distance from

the chitinous lining a sheath of loosely packed large muscle-fibres,

which completely encircle the tube and run in a longitudinal

direction. The fibres form several layers when seen in transverse

206 MR. F. E. BEDDARD ON A

section, and are very conspicuous on account of their deep stain-

ing with carmine. The cloaca genitalis is very nearly as deep as

Text-fig. 27.

Cirrus sac of Otiditenia eupodotidis.

C. Extensions of genital cloaca. C.S. Expanded region of cirrus sac. x2. Narrow

region of sperm-duct. « in genital cloaca points to spines on “ penis.”

NEW AVIAN TAPEWORM. 207

the length of the cirrus sac. When the entire system of genital

ducts is seen in longitudinal section, 7. e. in transverse sections of

the proglottid, it will be observed that the cirrus sac projects into

the cloaca genitalis, and that the latter is therefore prolonged

backwards as a kind of collar surrounding the protrusible part of

the cirrus sac. Dr. Gough has figured in Stilesia hepatica* a

similar backward growth of the cavity of the cloaca genitalis. In

such longitudinal sections of the cloaca genitalis of Otiditenia it

will be observed that the cavity is not so regular as I have seen

it in transverse sections ; there are lateral foldings of the lumen

which are probably paired and thus produce no appearance of

crumpling when the organ is viewed in transverse sections.

The vagina and cirrus sac, of course, open into this cloaca, the

vagina nearer to the external orifice of the cloaca genitalis.

In following out a series of sagittal sections of the proglottids,

which are, of course, transverse to the direction of the cloaca

genitalis, it will be seen that, as a rule, the cirrus sac becomes

separate from the vagina in a position which is at first obliquely

anterior to the vagina. This is the same on both sides of the body ;

but it is not invariably that the cirrus sae thus opens into the

genital cloaca anteriorly of the vagina. I have seen one case (text-

fig. 28, p. 208) in which the vagina lay anteriorly to the cirrus sac,

and in this case an obliquity occurred in the relative positions of

the two tubes, but the obliquity was in the reverse direction to that

shown in the normal arrangement. The obliquity, that is the

more dorsal or ventral inclination of the cirrus sac to the vagina, 1s

already known to vary; thus, in Monieza the position in relation

to each other of the two tubes varies according as to whether they

belong to the right or left side of the body, while in Avitellina

centripunctata Dr. Gough has figured, in his paper already referred

tot, an alternation of this kind in successive segments and on the

same side of the body. There is, however, here no question of a

reversal of an anterior to a posterior position such as I have found

in Otiditenia eupodotidis.

In one case I found another remarkable variation. This was

the complete absence of the proximal part of the vagina; into the

cloaca genitalis opened only a cirrus sac. However, as the series

of sections in which I cbserved this variation was traced further

towards the middle line of the body, the median region of the

vagina, which is of small calibre, came into view. In any case,

the wide region of that tube which, in normal cases, opens into

the cloaca genitalis was completely absent, save, indeed, for a few

traces of its muscular longitudinal sheath.

The vagina issues from the cloaca genitalis in an oblique

position, as seen when the worm is studied in a series of sagittal

sections, save, of course, for the exceptions mentioned above. — It

runs parallel with and later in close contact with the dense coil of

* “A Monograph of the Tapeworms of the Subfamily Avitellininz,” Quart. Journ.

Mier. Sci. vol. lvi. 1911, pl. xii. fig. 16, at.

+ Loc. cit. pl. xiv. fig. 43.

208 MR. F. E. BEDDARD ON A

the vas deferens on the ventral side. At first the vagina is wide,

with a much plicated lining epithelium surrounded by a thick

layer of muscles. It then suddenly becomes much narrower—

these two regions of the vagina being thus analogous to the wider

distal sac-like part of the cirrus and the narrow proximal part

of the same. The narrow region of the vagina joins the wider

region at an angle. Followed back the narrow part of the vagina

dilates with moderate abruptness into the large receptaculum

seminis, which commences at a point about on a level with the

pore side of the large ventral water vascular vessel.

Text-fig. 28.

Sagittal section showing different position of cirrus (C.) and vagina (V.) in

two segments of Otiditenia ewpodotidis.

The cirrus sac among the Tetracotylea occurs in at least two

types of structure. One, the most general, is typified, for example,

by that seen in Stilesia and represented in a figure in Dr. Gough’s

paper upon that genus and Avitellina, and discussed in a footnote

concerning the cloaca genitalis. The second type, which is cer-

tainly not so general, is seen in a genus Anoplotenia, the anatomy

NEW AVIAN TAPEWORM. 209

of which I have lately given an account to this Society *. In the

former type, as will be seen in Dr. Gough’s paper referred to, the

cirrus sac contains a narrow cirrus which is protrusible from

the sac and is of pretty well the same dimensions, through the sac

not being at all wider at its free extremity. In Anoplotenia the

cirrus-apparatus is divided into two regions; there is distally a

protrusible sac of much greater calibre than the slender cirrus

which is continuous with it posteriorly. The protrusible sac

appears to be an ingrowth into the cirrus sac of the cloaca geni-

talis. It is this kind of terminal male apparatus which we find

in the tapeworm which forms the subject of the present com-

munication to the Society. I have examined into its structure

both by transverse and longitudinal sections, and have found that

it presents certain differences from the cirrus sac of Anoplotenia.

It has already been said, in describing the cloaca genitalis, that

the end of the cirrus sac protrudes into that cavity ; how far it

ean be protruded without any further temporary dislocation of

its parts I do not know. It is evident, however, that, as with

Anoplotenia, so in Otiditenia, there is a terminal section of this

apparatus which can be protruded, though I have not met with

specimens in which the protrusion is actually effected. 'Text-

fig. 27 (p. 206) shows a longitudinal section through a cirrus

sac that is seen in a transverse section of the proglottid con-

taining it. It will be seen that the whole organ is of con-

siderable size, though not so large proportionately as it is in

Anoplotenia. It is, however, of much the same shape, being a

slightly elongated sphere, though flask-shaped when fully retracted.

The sperm-duct in this particular specimen approaches the cirrus

sac from one side, and then sharply curves to enter it. The point

of entry is at the extreme posterior end, 7. e. that furthest removed

from the orifice of the cirrus sac into the cloaca genitalis,

The sperm-duct is a good deal coiled within the cirrus sac and

has clear slightly stained walls. It does not, however, become

enlarged anywhere to form a vesicula seminalis. The transition

between the sperm-duct and the cirrus is abrupt. The cirrus is

not, however, of greater calibre, or at most of very slightly greater

calibre, than the sperm-duct with which it is continuous. The

abruptness of the transition lies in the fact that the walls of the

cirrus are thick and very deeply stained and the lumen propor-

tionately reduced. The cirrus is very much shorter than the

section of the sperm-duct which lies within the cirrus sac ; it is

slightly coiled. The cirrus itself again terminates abruptly in a

wider sac. As will be seen in text-figure 27, this latter sac is

flask-shaped, and, in fact, broadly corresponds in outline with

the entire cirrus sac. The wider part of the flask lies within the

wider part of the cirrus sac and the narrower part in the neck

of the latter.

The whole sac is composed of a deeply staining epithelium,

* P.Z.S. 1911, p. 1003.

Proc. Zoou. Soc.—1912, No. XIV. 14

210 MR. F. E. BEDDARD ON A

which is quite continuous above with the lining of the genital

cloaca and below with the epithelium of the cirrus sac. The

epithelium appears to be quite the same throughout this whole

tract. It is clear, in fact, that both the narrow cirrus and

the flask-shaped pouch into which it opens form one structure

possibly invaginated from the genital cloaca, which is quite distinet

morphologically from the sperm-duct. The cirrus sac itself 1s

algo divisible into two regions when it is occupied by the fully

retracted cirrus, as shown in text-figure 27. The almost spherical

distal part leads through a narrower neck to the cloaca genitalis.

The spherical bulb is covered externally by a thin layer of muscle-

fibres. The interior of the sac is filled with a tissue like that seen

in the cirrus sac of other tapeworms, for example, in Anoplotenia,

where I have represented its principal characteristics *.

There are many nuclei belonging apparently to delicate muscular

fibres and forming a padding tissue between the contained coils

of the male efferent system. Running from the wall of the

cirrus sac furthest from the pore are muscular slips which are

attached to the nearest end of the wide sac into which the cirrus

opens and are doubtless retractile in function. The narrower

neck-part of the cirrus sac is tubular and has a much thicker

coating of muscles, which are circular in direction. This layer is

one with the thin muscular coat of the bulb-region of the cirrus

sac. Between this thick muscular coating and the terminal part

of the male efferent apparatus which lies within it is seen a

delicate muscular layer with abundant interspersed nuclei like the

packing tissue of the bulb of the cirrus sac. The fibres, however,

have a definitely longitudinal direction, and would seem to be

part of the retracting apparatus.

It has been mentioned that the cirrus sac projects into the

cloaca genitalis. The surface which thus projects is covered on

that part of it which faces the external pore by close chevawx

de frise of minute conical spines, which are not implanted hooks

such as often occur upon the cirrus in tapeworms, but rather

appear to be outgrowths of the cuticular covering. They suggest

to some extent a comparison with a group of fine sete described

by Fuhrmannt in the two species Anomotenia penicillata and

A. isacantha in an apparently similar situation.

The vas deferens immediately on leaving the cirrus sac forms a

dense coil filling up a good deal of space. The tube is here wider

than the sperm-duct within the cirrus, and the nuclei of the cells

which constitute its walls are very obvious; the cells themselves

are rather clear. I could find no vesicula seminalis along the

course of the sperm-duct.

The wierws of this worm, together with its contents—the ripe

or ripening embryos—shows certain definite peculiarities. At its

first appearance the uterus is sac-like and narrow, lying in front

of the ovaries and testes but still not very near to the anterior

# P. Z.S. 1911, p. 1015, text-fig. 215.

+ Centralbl. f. Bakt. u. Parasit. xlv. 1908, p. 516.

NEW AVIAN TAPEWORM, 211

border of the segment. In this stage, when it is empty of eggs,

it forms a narrow transversely running tube. In transverse

sections of a slightly older uterus, such as that which is repre-

sented in text-figure 24 (p. 201), the position of this part of the

reproductive organs is seen to be remarkable. If such sections be

examined from in front backwards, the uterus is the first part of

the generative system to come into view ; for, although posterior

in position in its segment, the uterus lies in front of the ovaries

and testes. In such sections it appears first of all dorsally as a

narrow and transversely elongated sac, not lying in the medullary

parenchyma, but lying within the internal and circular layer of

muscles which delimit the medullary parenchyma from the cortical

layer—the body-wall of the worm. The uterus, so to speak, splits

this circular layer of muscles into two layers, fibres being recog-

nisable both dorsally and ventrally of it. It has probably pushed

its way into this situation; but located there the appearance

given is most suggestive of a coelomic cavity lying between the

body-wall and a central hypoblastic mass comparable to the

reduced gut of certain simple Planarian worms (e. g. Haplodiscus).

In such sections, moreover, the lumen of the uterus is seen to be

not quite continuous from side to side of the body. Muscular

fibres run across dorso-ventrally, which are of the nature of

strands rather than septa. These are plainly shown in the

accompanying illustration (text-fig. 29). Moreover, the epithelial

lining of the uterus is quite alone in these transverse sections, its

nuclei being very deeply stained.

This epithelial lining is continued over the strands of muscular

tissue which partly divide up the cavity of the uterus. Although

the uterus lies, as already said, below the dorsal parietes, there

are hereand there a few outpocketings of the cavity of the uterus,

spherical in form, which push their way into the thickness of the

dorsal parietes. Each of these contained only one or two groups

of ovarian and vitelline cells, and in the case of one uterus T only

found two of these small blind outgrowths. These transverse

sections also show another fact in the constitution of the growing

uterus, and that is its increased dorso-ventral diameter laterally,

both on the pore side and on the opposite side. These dilated

regions of the uterus were rather more liberally traversed by

transverse strands than the rest of the cavity. The lateral

widening of the uterus is also very plainly to be seen in horizontal

sections, where it is visible even in rather younger proglottids

than those just referred to. These longitudinal horizontal sections

(see text-fig. 25, p. 202) also show very well the extension of the

uterus on both sides of the body up to but not beyond the large

ventral water vascular tubes. The cavity of the uterus is thus apt

to be a little shorter from side to side than the space occupied

by the testes. On the other hand, when viewed in a series of

longitudinal sagittal sections, the uterus will be seen to have a

greater dorso-ventral extension than the space occupied by the

testes. In the three figures (text-figs. 24, 26, 29), which show

14*

I12, MR. F. E. BEDDARD ON A

the gradual increase in importance of the uterus, it will be also

observed that the total area occupied by the uterus and the rest

of the generative organs diminishes in its proportion to the length

of the segment in the older proglottids. It will also be noted

that in the oldest proglottid figured (text-fig. 29) the testes are in

course of disappearance.

Text-fig. 29.

Nearly mature proglottid of Otiditenia ewpodotidis, sagittal section.

U. Uterus.

In the ripe proglottids which are in course of being detached

the condition of the eggs is remarkable in several ways. The

NEW AVIAN TAPEWORM. VALI)

segments, as has been pointed out in considering the external

characters of this worm, are enormously swollen so as to be

almost globular in form, except for the frill-like posterior end of

each proglottid, where it ‘overlaps the one next in order behind it.

It would be easily supposed that this tension of the proglottids is

simply a matter of their being gorged with ripe eggs, such a

dilatation of proglottids seen in other tapeworms being due to

this cause. A microscopical examination, however, of such pro-

glottids shows that this conjecture is wrong. By far the greater

part of the interior of the proglottid contains no uterus or egg at

all. Fully three-quarters of the area of the proglottid is quite

empty of eggs, and the medullary substance only contains a few

fragmentary remains of other parts of the sexual system—as I

suppose, for I have not identified the vestiges in question with any

certainty. Moreover, the dilatation is not due to the imbibition

of fluid during the processes of preservation. It was as obvious

in proglottids shed during the life of the bird as in those extracted

from the gut after death.

I have examined such proglottids in both transverse and longi-

tudinal sections. Both sets of sections prove that the ripe eggs

(or rather embryos) are only present posteriorly in the segment.

In one proglottid, which was completely cut into sections and

mounted upon ten slides, no less than seven of these slides

showed no embryos at all scattered through the parenchyma ;

in another proglottid, however, treated in the same way, embryos

were found to lie in the more anterior region, though here also

the most anterior part of the proglottid was devoid of eggs.

In both of these specimens, however, and in several others which

I have examined by means of sections, the ripe embryos or masses

of embryos by no means occupied the whole of the parenchyma.

There were large tracts of parenchyma quite uninvaded by the

uterus or its remains.

The fact that the masses of eggs may in some cases lie rather

more anteriorly in the segment than they are apt to do in less

fully ripe proglottids is, as I think, in favour of Dr. Benham’s

view that a scattering of eggs takes place owing to a rupture of

the uterus when the proglottids come apart *. The parenchyma

in these ripe proglottids must now be considered, as its condition

is doubtless related to the changes in the uterus that remain for

deseription. In younger proglottids, such as those represented in

text-figure 24, the parenchyma is firm in appearance, moderately

stained by carmine, and with abundant very well-stained nuclei ;

these nuclei seem to have a prevalently dorso-ventral arrangement

in lines. Calcareous corpuscles are abundant, especially towards

the posterior region of the segment, where the generative organs

ave situated. In still older proglottids, which, however, are not

yet detachable from the strobila, such as those represented in

text-figure 29, the medullary parenchyma has taken on a

* Quart. Journ. Mier, Sci. voi. xliii., 1900.

214 MR. F. E. BEDDARD ON A

different appearance. It is much more distinctly fibrous than in

younger proglottids, as is plainly shown in the text-figure just

referred to. The fibres run, on the whole, dorso-ventrally, and

thus appear transverse in sagittal sections. The fibrous appear-

ance of the parenchyma is particularly obvious where the fibres

are frayed out, owing to the breakage of the section. In such

sections the caleareous corpuscles are still quite plain, chiefly

massed posteriorly in each segment. And, finally, in proglottids

of this age the uterus retains the perfect regularity of its shape.

We may now revert to the ripe detached or quite easily detach-

able proglottids. The same fibrous appearance of the medullary

parenchyma is plain; but, owing to the swelling of the whole

proglottid, the fibres no longer run entirely transversely in a

dorso-ventral direction, but tend, in places at least, to be more

circular in the way in which they lie.

This is especially the case in the neighbourhood of the uterus.

In these segments, however, we can no longer speak of a uterus

like that of the earlier proglottids represented in the series of text-

figures already dealt with (text-figs. 24-29). In the completely

mature proglottids (text-fig. 30) under consideration the embryos

are in partly or wholly detached masses of roughly spherical form

consisting of more or fewer individual embryos. These spherical

sacs, though of course vestiges of the uterus, seem to have no

definite walls of their own, but to be bounded only by the fibrous

tissue of the medullary parenchyma. There are comparatively

few of them, and those so small as to contaih only one embryo are

very few. The spherical masses of embryos are encircled by the

fibres of the medullary parenchyma in such a way as to produce

the impression of a definite sheath, which is perhaps particularly

obvious in the case of two such masses lying close together. This

statement applies to the larger as well as the smaller masses of

ova. There is not, however, any very great development of this

sheath, the character of its fibres being quite like that of the

surrounding parenchyma. Otherwise, we might speak of par-

uterine organs ; and, in any case, it is to be pointed out that we

may have here just the commencement of the formation of

paruterine organs, which is carried much further, but along the

same lines, in such a genus as Thysanotenia*. I take it, how-

ever, that there is nothing sufficiently definite to allow of the

assertion that the present genus or species is to be characterised

by the possession of paruterine organs. Before comparing the

history of the development of the uterus in this species with that

of other forms it will be necessary to follow the development of

the contained eggs.

In the uterus, when in such a stage of development as is

represented in text-figure 24, there are comparatively few eges.

In many sections the cavity of the uterus, which is not yet very

spacious, is seen to be completely empty. When ova are present

* See Beddard, “On Two Genera of Tapeworms,” P, Z.S. 1911, p. 994,

NEW AVIAN TAPEWORM. 215

they are small and delicate-looking, and not accompanied by much

interstitial matter. The uterus itself at this stage has not very

marked epithelial walls. The appearance of the ova is, indeed,

that of masses of the ovary transferred bodily to the uterus.

Later on the lining epithelium of the uterus is quite obvious

especially so far as concerns the nuclei of the same. In the

uterus of this and some later stages the individual eggs (or

embryos, as the case may be) are by no means always in close

contact with each other. They are separated by and imbedded in

a great deal of interstitial substance, which, as a rule, stains very

faintly with carmine.

Text-fig. 30.

Mature proglottid of Otiditenia ewpodotidis, longitudinal section.

U. Chambers into which the uterus becomes divided.

It contains, however, a good many nuclei which do stain, and

is thus evidently of a cellular nature. In the uteri of this

216 MR. F, E. BEDDARD ON A

particular series of proglottids (shown in text-figure 25, p. 202) the

accumulation of interstitial cells was chiefly in the uteral expan-

sions close to the water vascular vessels. In these regions of the

uterus the lumen was often entirely occluded by the densely

packed contents. In the uterus of a later stage—that which is

represented in text-figure 26-—the embryos were firmly imbedded in

the interstitial substance throughout the whole organ, so far as I

could see. This substance appears much denser in the uterus of

these older proglottids—so much so that cavities are left here and

there from which an embryo has fallen out in the course of

preparing the sections, which preserve exactly the spheroidal shape

of the embryo. The interstitial matter also seems to take stains

more readily at this stage, but exhibits no nuclei and no structure

that I was able to unravel.

In the fully mature proglottids (see text-figure 30) this inter-

stitial substance is still present, and the embryos are seen in the

same way to be embedded in it, leaving cavities of exactly their

shape when they chance to have fallen out. The interstitial sub-

stance, however, appears here to be rather laxer and is not so

deeply stained. It is very possible that this matter lying between

the eggs is a source of food-supply for the growing embryos. In

Avitellina centripunctata Gough has described the fact that the

eges, after they have arrived in the uterus, ‘‘ become surrounded

by smaller cells derived from the walls of the uterus,” as has been

pointed out by Fuhrmann in the case of Stilesia sjistedti. These

cells are figured by Gough* as lying between and apparently

completely filling the interstices between the eggs. These cells

have conspicuous nuclei and are held by Gough to perform a

nutritive function with regard to the eggs.

This final disposition of the uterus, or perhaps rather of its

contents, in Otiditenia eupodotidis is not exactly like anything

that is known to me at first hand or from the figures and

descriptions of others. It is, however, suggestive of the figures

that have been published of Monopylidium and even of Davainea.

In his definitions of both of these genera (which are placed in

separate families) Ransom } describes the fate of the uterus in

almost exactly similar words: viz. “ Uterus breaks down into

(numerous) egg-capsules, each containing one or more (several)

eggs.” This is a little misleading in that the sharply contoured

and thick-walled capsules of certain species generally assigned to

the genus Davainea are different from the figures given of Mono-

pylidium infundibulum ete. The definition would, moreover,

apply to Otiditenia, in which, as I maintain, the conditions are

really rather different from both.

In several species of Monopylidium the ripe eggs are scattered

and imbedded singly in the medullary parenchyma, so far

* Gough, Quart. Journ. Micr. Sci. vol. lvi. pl. xiv. fig. 49.

+ “The Tenioid Cestodes of North-American Birds,” Bull. U,S. Nat, Mus. no. 69,

1909.

NEW AVIAN TAPEWORM. 217

suggesting the genus Oochoristica *. Of MW. rostellatum Fuhrmann

writes f that each egg is surrounded by “une parenchyme vacuo-

laire qui forme ..... des cellules hexagonales.” This is repre-

sented in a figure which suggests a definite limiting membrane to

each of these “cellules,” and thus separating them from the

general medullary parenchyma. In J/. cayennense, M. secundum,

and M/. macracanthum the same author { finds the same singly

imbedded ripe ova. And this condition appears to characterise

some other species of the genus Monopylidiwm. There is,

however, another species of tapeworm recently described from

Gallinula chloropus under the name of “ Tenia marchali” by

Mola §, which Fuhrmann || refers to the genus Monopylidium, in

which the disposition of the ripe ova is rather different. Mola,

in fact, figures the ova as not scattered singly through the paren-

chyma, but arranged in groups of one to six. Furthermore, he

does not represent the parenchyma surrounding them as vacuolar

(as does Fuhrmann in his species), but as distinctly fibrous. The

state of affairs is therefore obviously much more like that of the

species with which | am concerned in the present paper. Mola,

however, does not show any tendency for the fibrous parenchyma

to enwrap the bundles of eggs such as I have described above in

Otiditenia. There is, furthermore, no suggestion given of an

epithelial lining to the cavities which lodge the eggs in

M. marchali.

Some important histological details are added by Clere 4] with

reference to Monopylidium infundibulum.

In this species we have, as it would appear, an agreement with

M. marchali in the fact that several ova are lodged together in

one lacuniform cavity of the medullary parenchyma. But these

cavities are not isolated ; they form a complicated and irregular

network of lacune**. Later still, however, the individual ova

become more completely separated the one from the other, and

the typical (¢) character of the genus Monopylidium is arrived at.

It may also possibly be the case with IM. marchali. 1 have, how-

ever, no reason to suppose that the metamorphosis of the uterus

in my genus Oliditenia proceeds any further than is indicated in

* Of, e.g., Beddard, P. Z. S. 1911, p. 633, text-fig. 150, e.

+ Rev. Zool. Suisse, t. xvi., 1908.

{ Centralbl. f. Bakt. Parasit. Bd. 45, 1907.

§ Bull. Ac. Belg. 1907, p. 886.

|| Die Cestoden der Vogel, Zool. Jahrb. 1908, Suppl.-Bd. x.

§] Rev. Zool. Suisse, t. xi. 1903, p. 354. Ransom places this species in the genus

Choanotenia on the grounds of his own observations as well as those of Cohn (Nova

Acta Ac. Nat. Cur. Bd. Ixxix., 1901), which seem to show that the uterus is per-

sistent “and possesses an irregularly lobulated cavity incompletely subdivided by

infoldings from the wall.” This is, according to Ransom, a character of Choano-

tenia, and contrasts with the breaking up of the uterus in Monopylidium, which I

have referred to above. I may point out, in support of Clerc, that Cohn’s figure (loc.

eit. pl. 32. fig. 47) may well indicate an incompletely mature uterus, since he

represents in the same proglottid both ovaries and testes. In Otiditenia, at any

rate, the ovaries and testes have disappeared when the uterus is fully mature. Cohn’s

figure may correspond to a stage figured in text-figure 30 of the present memoir,

** Toc. cit. vol. xi. figs. 73, 77, 85.

218 MR. F. E. BEDDARD ON A

text-figure 30. Clere also figures somewhat similar changes

in the uterus of a species of Dilepis* (not named), where the

scattered ova actually penetrate the cortical parenchyma. This

fact I have also observed in Otiditenia.

There are, therefore, clearly differences of importance between

the ultimate fate of the uterus in the two genera—as I regard

them—WMonopylidium and Otiditenia. In the former the eggs

are in most (? in all) species finally scattered through the paren-

chyma unaccompanied by any other cells such as occur with the

ova in Otiditenia, in which genus the eggs are at the very

end grouped into many more or less isolated but not well-marked

capsules. There remains a comparison of the uterus of Otiditenia

with that of Davainea. With regard to certain species of the

latter there is no particular comparison possible. But Clere has

figured the egg-capsules of Davainea frontinat, which do seem to

present certain similarities to the egg-capsules of Otiditemia

eupodotidis. In the former worm the egg-capsules differ from

those of such a species as D. comitatus t in that the individual

eggs do not lie closely together within the capsule, but are more

or less widely separated with a good deal of interstitial matter

between them, which the author regards as a part of the medullary

parenchyma which has undergone alteration, This and the

enclosed eggs is surrounded by a definite layer of cells, which are

not to be looked upon as a part of the persisting walls of the

uterus, since in that case they would not enclose a portion of the

medullary parenchyma. If this interpretation of the facts be

correct, there is clearly no possible correspondence with the

phenomena which I myself describe in the present paper. It

seems to me, however, to be not impossible that the supposed

altered parenchyma-cells which lie amongst the ova are really the

equivalents of the similarly placed cells in my genus Oéeditenia.

Moreover, Ransom § speaks of cells accompanying the scattered

ova of Davainea rhynchota.

I am disposed, however, to think that there is another view to

be taken of the egg-capsules of Davainea frontina, as figured by

Clerc. From the account that he gives of them in the text and

of the investigations of others on the same subject—to which we

may add those of Leuckart ||—it seems clear that the egg-capsules

correspond to the paruterine organs of a genus recently described

by myself 4] under the name of Thysanotenia, as I have already

suggested. In this case we cannot agree with the statement of

Ransom, in defining the subfamily Davaineine, that the “ uterus

breaks down into numerous egg-capsules.” For in Thysanotenia

there is, I think, no doubt that the cases in which the eggs finally

come to lie have nothing whatever to do with the uterus, and are,

* Clerc, loc. cit. pl. xi. figs. 75, 76.

+ Loc. cit. pl. xi. fig. 80.

{ Ransom, Bull. U.S. Nat. Mus. no. 69, p. 17, fig. 8, cap.

§ Loe. cit. p. 14.

|| Quoted in Bronn’s ‘ Thierreichs,’ p. 1445,

4) P. Z. 8. 1911, p. 1000.

NEW AVIAN TAPEWORM. 219

in fact, paruterine organs. This is not the case with the sacs of

eges in Otiditenia.

We may abstract from the foregoing account of the anatomy of

this worm the following réswmé of its structural characters :-—

Length about 3 inches; greatest breadth 4mm. Scolex with

feebly developed rostellum with a marginal double circle of very

numerous and small hooks. Suckers unarmed and rather large.

Neck present and short. Proglottids broader than long, except “at

the extreme end of the body, where they become rather longer

than broad. Completely mature proglottids almost spherical,

except for the frill which overlaps the following proglottid. ‘Two

excretory tubes on each side, of which the ventral are very wide

and the dorsal narrow; the two are at times parallel, but occasion-

ally the dorsal tube is really somewhat dorsal in position. Ventral

vessels joined by a wide cross-trunk at the posterior end of each

proglottid. Generative pores alternate irregularly; the generative

ducts pass between the dorsal and ventral trunks of the excretory

system. ‘The generative pores are not borne at the end of out-

growths of the proglottid. The testes lie at the extreme end of

the proglottid and form a vertical wall of over 200 separate

gonads, which, however, are included in a mass of medullary tissue

rather different from the medullary parenchyma generally. The

sperm-duct forms a large coil, and is also slightly coiled within

the rather large cirrus sac. In the latter the actual cirrus is

short and is connected with a wider (? eversible) tube also con-

tained within the cirrus sac. The cloaca genitalis is deep. The

ovary and vitelline gland lie in front of the testes. The vagina is

furnished with a, lar ge receptaculum seminis. It opens into the

cloaca genitalis behind and at right angles to the cirrus sac. The

uterus lies in front of the ovary, and in mature segments the

whole reproductive system is restricted to the posterior fifth or so

of the proglottid. The uterus is a narrow transverse tube, wider

at the two sides. Later it is broken up into several more or less

spherical compartments. The eggs are imbedded, within the

uterus, in a mass of cellular tissue whose origin is uncertain, and

which later loses its cellular character,

We have now to consider whether the species described in the

present communication can be referred to any known genus or

whether it will be necessary to form a new genus for its reception.

The abstract of the anatomical characters that has just been given

shows that the worm is undoubtedly a member of the large family

Hymenolepidid or perhaps of the Davaineide. This conclusion

need not be proved in detail. Furthermore, it is clear that the

features which distinguish it, viz. the following assemblage of

characters, necessitate its inclusion in one of the genera Mono-

pylidium, Choanotenia, or Anomotenia of the Hymenolepidide,

or to a new genus altogether. These characters are:—Armed

rostellum and unarmed suckers; large number of proglottids ;

alternate generative pores; absence of paruterine organs; testes

behind ovaries; breaking up of uterus in ripe segments; and in

220 ON A NEW AVIAN TAPEWORM.

a number of points which we shall have to consider in fixing the

position of this worm. It is true that the three genera mentioned

are separated by somewhat slender characters, which is emphasized

by the fact that Choanotenia infundibulum has been referred by

Fuhrmann to the genus Monopylidiwm, and that the same author

has left the position of Choanotenia laevigata of Rudolphi in

doubt.

All the three genera, however, seem to differ from that which

forms the subject of the present paper by the strong muscular

rostellum and by the relatively small number of the hooks,

arranged in a single or a double crown. Moreover, the seg-

ments in representations of these genera are figured as con-

siderably longer in proportion to their breadth than they are

in Otiditenia. Finally (so far as concerns the more important

point), the uterus appears to be a persistent and continuous sac in

Choanotenia and Anomotenia, while the eggs in Monopylidium

come to be scattered singly through the parenchyma. As I have

endeavoured to show on a previous page *, the development of the

uterus in Otiditenia is different. Nevertheless, I am disposed to

place Otiditenia more nearly in the neighbourhood of these genera

than in that of any others.

With regard to the Davaineide the present genus does un-

doubtedly show certain points of likeness to the genus Davainea,

the definition of which genus is not, however, a matter of great

ease. Otiditenia certainly lacks hooks upon the suckers, the

existence of which in Davainea forms a part of its generic

definition. But it must be remembered that D. lateralis from the

mammal G'aleopithecus has suckers without hooks. The breadth

of the rostellum and its numerous hooks is a point of similarity to

Davaimea, in which genus the rostellum may be rather rudi-

mentary (cf. Zschokke +), The principal reason, however, against

assigning this worm from Hupodotis kori to the genus Davainea

is the persistence of the uterus to the stage of the perfectly ripe

proglottids, whereas in “the genus Davainea a definite functional

uterus is not developed” (Ransom) £, and there are also paruterine

organs.

The generic § characters of Otiditenia will be, as I believe, the

following :—

Scolex with wide but rather rudimentary rostellum armed with

over & hundred minute hooks arranged in two rows. Suckers

unarmed. Neck present, not long. Proglottids for the most part

wider than long, only as wide or slightly wider than long at the end

of the body. Genital pores irregularly alternating. Detached pro-

glottids very swollen, almost spherical. Water vascular vessels two

pairs, ventral much the larger, nearly or quite lateral to each other ;

transverse trunks also wide. Genital organs not mature very early

* P. 210.

+ Centralbl. f. Bakt. u. Parasit. 1895, p. 634.

{ Bull. U.S. Nat. Mus. no. 69, 1909, p. 14.

§ I do not feel able to differentiate the specific characters.

ON THE MILK-DENTITION OF THE RATEL. 22/1

im strobila. Testes numerous, at least 200 lying posteriorly in a

thin vertical plate extending between water vascular vessels and

even beyond them laterally, imbedded in interstitial tissue different

from medullary parenchyma. No vesicula seminalis, sperm-duct

with large coil; cirrus sac large, containing a looped part of sperm-

duct, short cirrus, and an evaginable sac derived from cloaca geni-

talis. Ovaries and vitelline gland in front of testes. Uterus in

Jront of ovuries, a narrow transverse sac with cavity partly divided

by ingrowing trabecule. Later uterus converted into several spherical

sacs. Kggs thin-walled, surrounded by nutritive (?) cells.

Hab. Hupodotis kori.

The most noteworthy characters of the genus, so far as the

external characters go, appear to be the rather wide and

rudimentary rostelluin, which is, nevertheless, armed with very

numerous though very minute hooks. The internal structure is

remarkable for the restriction of the gonads to the extreme

posterior part of each segment, and to the fact that the very

numerous testes arranged only one deep are imbedded in a very

different kind of medullary parenchyma, which is obvious to the

eye on account of its feeble staining: by the peculiar form of

the cirrus sac and the inclusion within it of an outgrowth of the

genital cloaca, which is probably protrusible like the somewhat

similar “‘ penis” of Anoplotenia: and by the fate of the uterus and

the presence of interstitial cells lying among the eggs.

14. On the Milk-Dentition of the Ratel.

By R. LypDEKKER.

[ Received September 30, 1911: Read November 21, 1911.]

(Text-figures 31 & 32.)

Many instances are known among extinct mammals in which

the milk-dentition presents primitive features entirely lost in the

teeth of the permanent series. In the equine Merychippus, for

example, the cheek-teeth of the milk-series are of the short-

crowned cementless type of Anchitheriwm, whereas those of the

permanent set show the high-crowned and cemented type cha-

racteristic of the more specialised representatives of the family

Kquide *. So far, however, as I am aware, no such atavistic

features have been recorded in the case of any existing mammals ;

and it is therefore of interest to bring to notice what appears to

be a case of this nature.

In the ninth edition of the ‘ Encyclopedia Britannica,’ Sir

William Flower classed the Ratels (M/ellivora) with the Badgers

in the subfamily Meline; this classification was followed in

Blantord’s volume on Mammalia in the ‘ Fauna of British

India’ + and in Flower and Lydekker’s ‘Study of Mammals’ ¢.

* See Lull, Amer. J. Science, vol. xxiii. p. 177, 1907.

+ Page 175, 1888. { Page 576, 1891.

De, MR. Re LYDEKKER ON THE

When re-arranging, some fifteen years ago, the exhibition series

of mammals in the British Museum I came, however, to the

conclusion that this was incorrect; and in the label on the

genus Mellivora 1 stated that although Ratels were generally

classed with Badgers, it appeared, from thei peculiar type of

colouring and the form of the upper molar, that they ought to be

placed next the Tayra and Grison (Galictis) in the subfamily

Mustelins, This revised classification (which renders the

definition of the last-named subfamily more concise) was followed

in the revision of Sir William Flower’s article ‘‘ Carnivora,”

contributed by myself to the eleventh edition of the ‘ Encyclopedia

Britannica,’ and likewise in my account of the Mustelide in

‘Harmsworth’s Natural History’*. It has also been adopted

in Max Weber's ‘Siugetiere, who probably follows Winge in

this respect.

Text-fig. 31.

Left upper dentition of Mellivora and Gialictis.

A. Permanent teeth of Mellivora.

B. 3; . Galictis. :

C. Milk-, and some of the permanent, teeth of Mellivora.

Thus matters stood till a few days ago, when I observed

among a series of specimens sent to the British Museum by Miss

Olive MacLeod from the Lake Chad district the skull of a Ratel

in which the milk-dentition is just being replaced by the

permanent set; the upper carnassial and molar being protruded,

but the milk-carnassial being still retained, as are the milk-molars

in advance of this tooth and the canines.

* Vol. i. p. 513. In revising the 8th ed., 1906, of the ‘Guide to the Mammal-

Galleries in the British Museum (Nat. Hist.),’ I unfortunately allowed Mellivora

to retain its old position (p. 45).

MILK-DENTITION OF THE RATEL. 223

In the permanent upper dentition of Ratels the carnassial

(p. 4), in common with that of nearly all other living Mustelines,

has the inner tubercle placed close up to the front edge of the

tooth, while the molar (m.1) is characterised by the antero-

posterior diameter of the inner half of the crown being greatly in

excess of that of the outer half. In the milk-dentition, on the

other hand, the carnassial (which in most Carnivora is a replica

of the permanent one) has its inner tubercle placed near the

middle of the blade; while, as I gather from another specimen,

the inner half of the molar is much narrower than the outer,

this tooth having, in fact, what may be called the typical

carnivorous triangular form.

Now in both the foregoing respects the aforesaid milk-teeth

correspond in general characters with their permanent repre-

sentatives in Galictis; the upper carnassial of that genus being

peculiar among existing Mustelines on account of having the

inner tubercle placed near the middle of the blade. ‘There are,

of course, differences in regard to the details of these teeth, and

also in respect to the orientation of the molar, when the milk-

series of Mellivora is compared with the permanent set of

Galictis; but the resemblance is such as to leave little doubt as

to genetic affinity between the animals to which they respectively

belong. And it would thus seem that the milk-teeth of the

Ratel and the permanent ones of the Tayra represent a common

primitive type, which has been superseded by a more advanced

modification in the permanent teeth of the Ratel.

Text-fig. 32.

Outer side of left upper milk-dentition of Mellivora,

showing the bifid or bicuspid canine.

I find nothing very noticeable in regard to the hinder lower

milk-teeth of the Ratel; but the upper canine has a distinctly

bifid crown, as in certain Bats, and there are indications of a

similar bifurcation in the corresponding lower tooth. Whether

this is a primitive or a specialised feature, it is, in this case,

difficult to say ; it is almost certainly the former in Bats.

In referring to the teeth of Galictis as primitive in comparison

with those of Mellivora, it should be mentioned that the

comparison must be limited to those genera, as the permanent

carnassial of Tertiary Mustelines, such as Plesictis, is of the

Mellivora-Mustela type. That Mellivora is a more specialised

224 MR. J. LEWIS BONHOTE ON

form than Galictis is evident, not only from the dental features

already mentioned, but from the normal absence of the second

lower molar (m.2) and likewise by the shortness of the tail

and the disappearance of the ear-conchs; both the latter features

being, of course, adaptations to a burrowing life. In connection

with the absence of the second lower molar, it is interesting to

note that in one specimen in the British Museum (No. 9.7.19.1)

this tooth is retained on the left side. It is very small, like the

corresponding tooth of Galictis, and had come into use before the

carnassial was fully protruded, so that it would have been shed

early.

Although Galictis is now unknown north of Mexico, or

thereabouts, it occurs fossil in the Jater Tertiaries of the United

States; and this leads to the idea that Galictis and Mellivora are

divergent members of a common stock which, like the Leopard

(Felis pardus and FP. onca) and Ocelot (7. pardals, F. tristis, and

F. nebulosa) groups, once inhabited a large area in Asia, whence

it reached America by way of Bering Strait, and, having made

its way into South America, died out in the north of the

New World.

15. On a Further Collection of Mammals from Egypt

and Sinai. By J. Lewis Bonnorr, M.A., F.L.S.,

VAIS

[Received October 24, 1911: Read February 6, 1912. ]

The following is an account of a small collection of mammals

which has been sent home during the past two years by Capt.

Flower. The most notable specimens are the Meriones crassus

and Acomys russatus from Sinai, the type locality of these species.

Apart from their extreme rarity in collections, the acquisition

of these animals has enabled me to identify definitely the

Meriones of Lower Egypt with Pomel’s JM. sellysti and also to

describe the form found in the Sudan as a new race. The

specimens of Acomys russatus prove to be quite different in size

and colour from those obtained near Cairo by Mr. Nicoll and

myself, which latter are therefore described under the name

A. r. egyptiacus *.

I must express my indebtedness to Capt. Flower and Mr. Nicoll

for their kindness in allowing me to work out the collection and

more especially for bringing home some of the specimens alive,

and thus enabling me to carry on some observations and experi-

ments on which I shall hope to have something further to

record in the future.

* The complete account of these two new subspecies appears here; but since the

names and preliminary diagnoses were published in the ‘Abstract,’ No. 103, 1912,

they are distinguished by being underlined.—Eprror.

MAMMALS FROM EGYPT AND SINAI,

RHINOLOPHUS ACROTIS BRACHYGNATHUS KX, Anders.

Rhinolophus acrotis brachygnathus K. Anders. Ann. Mag. N. H.

ser. 7, vol. xv. p. 73 (1905); Bonh. P. Z. 8S. 1909, p. 788.

One specimen received from the Delta Barrage, 4th March,

1911.

Crocrpura (CRoC.) RELIGIOSA Geoffr,

Sorex religiosa 1s, Geoftr. Mém, Mus, xv. p, 128, pl, iv. fig, 1

(1827).

Crocidura (Croc.) religiosa Is. Geoffr,; de Wint, in Anders,

Zool. Egypt, Mamm. p. 168 (1902); Bonh, P. Z.S, 1909, p, 790.

Two examples, both males, of this minute species were brought

in from Aburoash near Cairo. Their measurements are as

follows :—

No. “ A.” Head and body 50 mm,; tail 37; hind foot 9; ear 7.

No, “ B.” Head and body 55 mm,; tail 40; hind foot 10; ear 8.

The skulls of both are in very good condition and show them

to be quite adult. The skulls are identical in size, and measure :

Greatest length 16 mm., greatest breadth 7; breadth across

maxille 5; interorbital breadth 4; breadth of snont 2; tip of

incisors to tip of large premolars 3°5,

CANIS ZERDA Zimin,

Canis zerda Zimm, Geogr. Gesch, ii. p, 247 (1780),

Vulpes zerda Zimm. ; de Wint, in Anders, Zool, Egypt, Mamm.,

p. 233 (1902).

3d near Muut, Oasis of Dakhel, 9th May, 1911,

This specimen was procured by Mr. Harding King, who has

kindly presented it to the British Museum,

MUNGOS ALBICAUDA Cuv.

Herpestes albicauda G. Cuv. Régne Anim. ed, 2, i, p. 158

(1829); de Wint. in Anders. Zool. Egypt, Mamm, p. 193 (1902).

Capt. Flower brought back a very young example of this species

from Abu Usher on the Blue Nile, taken on the |st November,

1910.

GERBILLUS PYRAMIDUM Geoftr,

Gerbillus pyramidum Is. Geoftr, Dict. Class, H. N. vil. p. 321

(1825); F. Cuv. Trans. Zool. Soc. ii. p, 141, pl. xxv, figs. 6-9

(1838) ; de Wint. in Anders. Zool. Egypt, Mamm, p, 255 (1902) ;

Bonh. P. Z. 5. 1909, p. 791.

Three specimens of a large Gerbille said to have come from near

Alexandria I have provisionally placed under this species, They

are fully adult, but show a considerable range in size, the female

being considerably smaller than the two males. From a com-

parison of these specimens with the series in the British Museum

it seems probable that there are several forms of this species, but

Proc. Zeoi. Soc.—1912, No. XV. 15

226 MR. J. LEWIS BONHOTE ON

the material at hand is not at present sufficient to admit of any

definite conclusions being arrived at.

The measurements are as follows :—

3. No. 358. J. L. B.: Hd.& b. 128 mm.; tail —; h.ft. 38; ear 17.

é No. 359 J... B.: Hd G& b. 126 mm. ; tail 143; htt. 34 eansie

2. No. 360, J. L. B.: Hd. & b. 116 mm.; tail 143; h.ft.32; ear 17.

Least supra- Length of

Skulls Greatest Greatest orbital Basal Length of molar

Sees est length. breadth. breadth. length. nasal. series.

Sa Borskse Bh, Wien, 19 18 30 15 6

GMO SO aes, 20 8 30°5 15 6

ORS COM Some: 18°5 7 28 13°5 i)

GERBILLUS GERBILLUS Oliv.

Dipus gerbillus Olivier, Bull. des Soc. Phil. Paris, u. p. 121

(1801).

Gerbillus gerbillus Oliv.; de Wint in Anders. Zool. Egypt,

Mamm. p. 252 (1902); Bonh. P. Z. 8. 1909, p. 792.

The collection contains two more of this pretty and common

Gerbille, one from the desert near Giza and the other from the

Khargeh Oasis. In spite of its wide range this species is

remarkably constant both in size and coloration.

MERIONES CRASSUS Sundev.

Meriones crassus Sundeyv. K. Vet.-Ak. Handl. p. 233, pl. ii.

fig. 4, craniuin (1843).

Gerbillus sellysii Pomel, C. R. Acad. Sci. xii. p. 654 (1856).

Meriones shawi Riipp.; de Wint. in Anders. Zool. Egypt,

Mamm. p. 266 (1902) (partim).

T have lately received from Capt. Flower an example of Meriones

erassus from Tor in Sinai. As this species was originally

described from Sinai, this example represents the typical form of

the species. It differs from that found m Lower Egypt in

being slightly larger, with a more pointed snout and a much

longer and stouter tail. The Egyptian specimens recorded by

me in a former paper agree well with a series from Tripoli in

the British Museum, which are practically topotypes of Pomel’s

<pecies, and I have now no hesitation in referring them to this

form. This I was unable to do before, owing to my having no

typical crassus with which to compare them.

The form of Meriones found in the Sudan is much paler than

that found in Lower Egypt, and I propose tor it the name

MERIONES CRASSUS PALLIDUS.

Abstract P. Z. 8. 1912, p. 3 (Feb. 13).

Meriones crassus sellysii Pomel, Bonh. P. ZS. 1909, p. 793

(parti).

. Very similar. to df. crassus sellysii, but much paler, and the

MAMMALS FROM EGYPT AND SINAI. 227

snout is, in life, rather more pointed, though this character is

not obvious in the skull. In size they are a tvifle larger and

with a rather shorter tail.

The skull differs from that of W. c. sellysii in having a rather

longer and narrower appearance. This is chiefly due to the

slightly greater length of the auditory bulle, which also project

further behind from the base of the skull, and the skull itself

is slightly narrower at its posterior margin.

Measurements of the type (in the flesh).—Head and body

127 mm.; tail 110; hind foot 28; ear 17.

Skull of type—Greatest length 39 mm., basal length 34,

greatest breadth 23 ; palatal length 17; diastema 10; length of

bulla 16-5; length of molar series 5°5.

Type. Coll. J. L. B. No. 313. ¢. Atbara, Sudan. Collected

by Capt. S. S. Flower.

The genus Meriones is one which offers great difficulties to the

systematic worker, as the different forms resemble each other

very closely and have at various times been burdened with many

names and but imperfect descriptions.

The forms dealt with at present are, however, fairly distinct ;

the larger size and long and stout tail easily distinguish the

typical crassus. M. c. sellysii (Tripoli and L. Egypt) 1s smaller,

with a less pointed snout, rather paler in colour, and with longer

and more conspicuous black tips to the hairs of the back.

M. c. pallidus (Sudan) resembles M. c. sellysir, except in its much

paler coloration. In life, however, the Sudan form has a more

pointed snout, approaching in this character true crassus. From

a comparison of the measurements below it will be seen that it

is also slightly larger.

Head and body. Tail. Hind foot. Lar.

M. crassus, lexample? .... 1389 mm, 148 33°5 18

M. c. sellysii, av. 4examples 126°5 ,, 120 29 bib

M.c.pallidus,,, 4 5, Heyl 3 Tia! PST 18-2

M. shawimelanurus, lex.2 139 ,, 135 32 il

In order to prevent any misunderstanding in the future, | may

mention that these measurements and other particulars are all

taken from wild specimens and do not in any way refer to a

large series of these animals that I have bred in confinement.

MERIONES SHAWI MELANURUS Rupp.

Meriones shawi Rozet, Voy. rég. d’Alger, p. 243 (sine descr.)

(1833); Cuvier, Legons d’Anat. Comp. iv. 2nd ed. 2nd _ part,

p- 456 (1835); Duvernoy, Mém. Soe. Strasb. iii. p. 22, pls. 1 et 2

1842).

ae melanurus Riipp. Mus. Senck. iii. p. 95, pl. 7. fig. 3

(1845).

a. Q ad. Near Alexandria, Egypt.

This species, which is the one figured by Dr. Anderson, is easily

Lae

228 MR, J. LEWIS BONHOTE ON

distinguished from the foregoing, as it is a larger and stouter

animal, more closely approaching Psamunomys in general build.

In colour it is darker than J. crassus, the dark tips to the hairs

being longer and much more conspicuous, especially on the tail.

The skull is more stoutly built and slightly longer, the extra

length being almost entirely due to the longer molar series.

The teeth themseives, except in size, resemble those of

MW. crassus.

The most obvious skull-difference, however, is to be found in

the auditory bulle, which are much smaller and do not extend

beyond the back of the skull.

The measurements of this specimen have been given under the

preceding species to facilitate comparison.

The skull-measurements are as follows :—Greatest. length

40 mm., basal length 35, greatest breadth 23; palatal length 19 ;

diastema 11; length of bulle 15; length of molar series 6.

PsAMMOMYS OBESUS Cretzschm.

Psammomys obesus Cretzschmar, Rupp. Atlas, p. 58, pl. 22

(1828); de Wint., Anders. Zool. Egypt, Mamm. p. 274 (1902);

Bonh. P. Z.5. 1909, p..793.

Three specimens of the typical form from Maryut, L. Egypt.

MUS RATTUS 'TECTORUM Savi.

Mus tectorum Savi, Nov. Giorn. Pisa, Feb. 1825.

Mus rattws Linn. ; de Wint., Anders. Zool. Egypt, Mamm. p. 274

(1902); Bonh. P. Z.S. 1909, p. 793; id. ibid. 1910, pp. 638, 651.

Mr. Nicoll shot an example of this form some distance up a

tree. I have been trying to obtain evidence as to which of the

two forms of Mus rattus climbs trees or whether both do it.

This is the first definite evidence that tectorwm is the climber,

although there is a certain amount of indirect evidence on that

point (see P. Z.S. 1910, p. 655). The question which requires

settling is, does M. r. alewandrinus usually climb trees? On this

point there is no evidence, direct or indirect. In confinement

both forms climb with equal facility.

MUS MUSCULUS GENTILIS Brants.

Mus gentilis Brants, Muizen, p. 126 (1827).

Mus musculus Linn.; de Wint., Anders. Zool. Egypt, Mamm.

p- 277 (1902); Bonh. P. Z.8. 1909, p. 794.

Two examples of this race of House-mouse, in which the hairs

of the under parts are white to their bases, were brought back by

Capt. Flower from Khartoum in the autumn of 1910.

Mus muscutus, albino var.

Mr. Nicoll received from his correspondent Signor A. J. Balboni

MAMMALS FROM EGYPT AND SINAT, 229

in the Wadi Natron two examples of white mice, with the note

that they were quite common in the Salt Company’s buildings.

These are apparently ordinary albino mice, similar to those which

are domesticated throughout Europe. There is, however, no

evidence to show whether these arose “ spontaneously” or were

imported by some previous occupant of the buildings and then

escaped. I have asked Mr. Nicoll to try to procure further

information on this point and also as to whether normal-coloured

and piebald individuals are also found.

Acomys RussATuS Wagner.

Acomys russatus Wagner, Abh. Akad. Miinchen, iii. p. 195,

pl. 3. fig. 2 (1840); Tristram, Fauna Palestine, p. 11, pl. 3. fig. 1

(1884).

I. have received a pair of these rare mice alive from Sinai, the

typical locality. This pair, which I received on May 3rd, gave

birth on the 10th to three young. These were born, as in the case of

A. cahirinus*, in a very forward state of development, being very

large and covered with hair and spines; they grew very rapidly and

were about half grown and well able to fend for themselves when a

fortnight old. They differ strikingly in coloration from the adults,

being of a uniform pale grey throughout; when three weeks old the

new adult coat of a lighter tint foie ale on the under parts,

and a week later the first:trace of the yellow colour of the upper

parts began to show on the shoulders; this coat gradually spread

backwards towards the tail in a broad stripe along the centre of

the back, the adult dress being finally completed along the flanks

at the age of six weeks, by which time they were full-grown. I

have been unable to note the exact period of gestation, but it is,

I fancy, slightly longer than in the case of 4. eabir’ nus.

This species is of a deep reddish brown on the back, each spine

having a darker apex; the head is greyer and the ‘under parts

are greyish white, there being no sharp line of demarcation

between the colours of the upper and under parts, as is the case in

all other species of this genus, except A. cahirinus. The tail is

short, being only about two-thirds the length of the head and

body, and appears almost black, although it is sparsely covered with

minute whitish hairs.

In my previous paper? I referred a specimen of an Acomys

taken just outside Cairo to dA. cahirinus. Since then I have

received a second example obtained by Mr. Nicoll in the Wadi

Hof near Helouan, and through the kindness of Capt. Flower I

have also received, as noted above, a pair of Acomys russatus alive

from Sinai.

This being the typical locality of A. russatus, the live ones are

topotypes, and they differ in several particulars from those found

* P.Z.S. 1911, p. 5. + P.Z.S. 1909, p. 795.

230 ON MAMMALS FROM EGYPT AND SINAT.

in Keypt; the Egyptian form is therefore undescribed, and I

propose for it the name

ACGOMYS RUSSATUS AGYPTIACUS.

Abstract P. Z.S. 1912, p. 3 (Feb. 13).

Very similar to Acomys russatus Wagner, but smaller and much

brighter and yellower in coloration. Character of the fur spiny

throughout, except on the vent, shoulders, thighs, and side of the

face. General colour above bright orange-brown, each spine with

a minute tip of dark brown, which, however, is not sufficient to

affect the general coloration. Hands, feet, and under parts

grevish white, shading into the colour of the upper parts with no

definite line of demarcation. Skin of ears and tail black, thickly

covered in the case of the ears with greyish hair on both surfaces

and in the case of the tail with short greyish spines. There is a

small but conspicuous white spot below the eye.

The skull, which I described in my previous paper, has the

snout rather shorter and broader than its allies and the bull

considerably larger, while the teeth and length of the molar series

are also of greater dimensions.

Measurements of type (in flesh).—Head and body 91 mm. ; tail,

damaged ; hind foot 17; ear 16.

Skull—Greatest length 27 mm., basal length 22, greatest

breadth 13; palatal length 12°5; diastema 7-5; length of molar

series 5; length of nasals 10.

Tope. Coll. J. L. B. No. 306. ¢ adult. Wadi Hof, near

Helouan. Collected by Mr. M. J. Nicoll, 29th October, 1909.

As compared with the true 4. rwssatus this form would appear

to be rather smaller and much brighter in coloration.

Measurements of an adult ¢ (in the flesh) of the typical race

are: Head and body 115mm.; hind foot 19 ; ear 18; it is therefore

considerably larger than the Egyptian form. Except in size the

skulls of A. r. egyptiacus aud the typical form are very similar.

The skull-measurements of A. russatws, typical race, are as

follows :—Greatest length 30 mm., basal length 25, greatest

breadth 15; palatal length 13°5; diastema 8; length of molar

series 5°5; length of nasals 11-5.

SpALAX aGypT1Acus Nehring.

Spalax typhus Pall., Anders. P.Z. 8. 1892, p. 472.

Spalax egyptiacus Nehring, SB. Gesellsch. naturf. Fr. Berlin,

1897, p. 180; id. ibid. 1900, p. 210; de Wint. in Anders. Zool.

Egypt, Mamm. p. 168 (1902).

Two examples of this scarce and little-known Rodent were

received from Marytit in the Delta.

Lepus stnarricus Hemp. et Ehr.

Lepus sinaiticus Hemp. et Khr. Symb. Phys., Mamm. u. pl. 14.

THE SECRETARY ON ADDITIONS ‘TO THE MENAGERIE. I31

fig. 1 (1830); de Wint. in Anders. Zool. Egypt, Mamm. p. 322

(1902).

3. Sinai, 23rd May, 1911.

This specimen, which was brought back from Sinai, through the

courtesy of Capt. J. Falconer Bey, is in very rough and worn

pelage. It has not yet been recorded from Egypt, and may be

distinguished from L. wgyptius by its paler coloration and shorter

hind foot.

PROCAVIA sp. ?

A. fine example of a Hyrax, which was taken near Ras Gurdi,

between Keneh and the Red Sea, was recently sent home. As

the locality is very near the spot where the examples of P. burtoni

mentioned in my previous paper came from, it is more than likely

that this specimen also belongs to that species. It shows, how-

ever, the yellow spot round the dorsal gland which is absent in

other examples of P. burtoni, although present in 2. ruficeps ; on

the other hand, the head does not show the rufous tinge character-

istic of the last-mentioned form. Unfortunately the skull, which

would have settled the difficulty, was accidentally lost. I incline

to the belief, however, that it is P. burtoni, and if so it must be

borne in mind that when in worn pelage this species may show

the yellow dorsal spot. The teeth, however, would form a definite

distinguishing character.

EXHIBITIONS AND NOTICES.

February 6, 1912.

E. G. B. Meavre-Waxpo, Esq., 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, 1911, and January, 1912 :—

NovEMBER,

The registered additions to the Society’s Menagerie during the

month of November were 247 in number. Of these, 119 were

acquired by presentation, 28 by purchase, 17 were received on

deposit, 77 in exchange, and 6 were born in the Gardens.

W3sy THE SECRETARY ON ADDITIONS 'TO THE MENAGERIE.

The total number of departures during the month, by death

and removals, was 261.

Amongst the additions special attention may be called to the

following :—

1 Spotted Hyena (Hifena crocuta) and 1 Crowned Duiker

(Cephalophus coronatus), from the Gold Coast, presented by

Capt. C. H. Armitage, D.S.O.; on November 6th.

1 Kashmirian Deer (Cervus kashmeerianus), from Kashmir,

presented by H.G. the Duke of Bedford, K.G., President of the

Society, on November 16th.

4 Reindeer (Rangifer tarandus), from Lapland, presented by

the European Construction Co., on November 11th.

1 Tasmanian Wolf (Thylacinus cynocephalus), 5 'Tasmanian

Devils (Sarcophilus satanicts), and 2 Bennett's Wallabies

(Macropus bennetti), fro. Tasmania, purchased on November

21st.

A collection of small rodents containing several species new to

the Collection, and specimens of two species of Elephant-Shrews,

both new to the Collection, from Bechuanaland, presented by

Capt. H. A. P. Littledale, F.Z.S., on November 18th.

A Hunting-Dog (Lycaon pictus somalicus), from Somaliland,

presented by Lieut. H. C. Dobbs, I:A., on November 21st.

A Bengal Fox (Viilpes bengalensis), from Bombay, presented by

Alfred W. Hanckel, Esq:, on November 10th;

A Long-eared Fox (Otocyon megalotis), from South Africa,

forwarded by Dr. Louis Péringuey, C.M.Z:S.,on November 18th.

2 Ruffed Lorikeets (Calliptilus solitarius), from the Fiji Islands,

new to the Collection, received on deposit on November 6th.

1 Aztec Jay (Cydnocitta stelleri azteca), from Mexico, and

2 Swamp-Sparrows (Melospiza georgiana), from North America,

new to the Collection, received in exchange on November 8th.

2 Cuban Banded Woodpeckers (Wesoceleus fernandine), 1 Cuban

Green Woodpecker (Xiphidiopicus percussus), and 4 Black-throated

Crested Quails (Hupsychortyx nigrigularis), the latter from

Central America, all new to the Collection; received in exehange

on November 22nd.

DECEMBER,

The registered additions to the Society’s Métiagerie during

the month of December were 88 in number: Of these, 38 were

acquired by presentation, 2 by purchase, 31 were received on

deposit, 14 in exchange, and 3 were born in the Gardens.

The total number of departures during the month, by death

and removals, was 211.

Amongst the additions special attention may be called to the

following :— .

1 Wild Cat (felis sylvestris); fvom Inverness-shire, purchased

on December 20th:

THE SECRETARY ON ADDITIONS TO THE MENAGERIE, Tae

4 Arctic Foxes (Vulpes lagopus), from Spitzbergen, deposited

on December 11th.

6 Green White-eyes (Zosterops virens), from South Africa,

new to the Collection, received in exchange on December 16th.

1 Concave-casqued Hornbill (Dichoceros bicornis), from India,

received in exchange on December Ist.

4 Chinese Alligators (Alligator sinensis), from China, deposited

on December 18th.

1 Kirtland’s Tree-Snake (TZhelotornis kirtlandi), 1 Green

Tree-Snake (Dendraspis viridis), from Dunkwa, presented by

Dr. H. G. F. Spurrell, F.Z.8., on December 16th.

J ANUARY.

The registered additions to the Society’s Menagerie during the

month of January were 88 in number. Of these, 50 were

acquired by presentation, 3 by purchase, 11 were received on

deposit, 5 in exchange, and 19 were born in the Gardens.

The total number of departures during the month, by death

and removals, was 267,

Amongst the additions special attenion may be called to the

following :—

1 White-throated Capuchin (Cebus hypoleucus), from Panama,

presented by Francis Wright, Esq., on January 31st.

2 Grand Galagos (Galago crassicaudata), and 1 Blotched Genet

(Genetta tigrina), from the ‘Transvaal, presented by Commander

F. J. Moseley, on January 20th.

1 African Red-winged Starling (Amydrus morio), from

S. Africa, new to the Collection, received in exchange on

January 3rd.

1 Yellow-rumped Hangnest (Pseudoleistes guirahwro), from

S. America, new to the Collection, presented by Miss Wi

Bromwich on January 22nd.

1 Yellow-lored Amazon (Chrysotis xantholora), from Paraguay,

purchased on January 4th,

2 Blue-rumped Parrakeets (Psittinus incertus), from Malacca,

received in exchange on January 3rd.

1 Abyssinian Love-bird (Agapornis taranta), new to the

Collection, deposited on January 3rd.

1 Calabar Snake (Calabaria reinhardti), from W. Africa, new

to the Collection, presented by S. H. Carnelley, Esq., on January

16th.

1 Schott’s Snake (Philodryas schotti), from S. America, new

to the Collection, received in exchange on January 18th.

(st)

vse

MR. R. D. CARSON ON A RED KANGAT.OO.

A Colour Phase of the Poiecat.

Mr. E.G. B. Mrapr-Waupo, V.P.Z.8., exhibited a mounted

specimen of a pale fulvous variety of the common wild Polecat

(Putorius putorius), belonging to the Rev. D. Edmondes Owen,

of Llandovery, 8. Wales. The animal resembled in colour a well-

known variety of the Badger, and was interesting because the

Polecats in that particular district were frequently of this

peculiar variety and had been well known locally for years.

Retarded Development in a Red Kangaroo.

Tue SECRETARY communicated the following account of retarded

development of the foetus in a Red Kangaroo, by Mr. Roper D.

Carson, C.M.Z.8. :—

A female Red Kangaroo (Macropus rufus) was received at the

Philadelphia Zoological Garden, June 5, 1908. On December 15,

1908, the movement indicating a young one was observed in her

marsupium ; this young one became ¢ No. 21 on our records.

The father of this young one came August 6, 1908, and died

October 28, 1908, seven weeks before ¢ No. 21 was noticed.

3S 21 ceased going into his mother’s pouch June 6, 1909, and

stopped nursing about November 7, 1909.

On September 25, 1909, movement was again observed in the

female’s pouch and proved to be what became on our records

3 No. 22. i

As the father of the first one, ¢ No. 21, had been dead 11 months

less 3 days-‘and no other male had been with this female nor in

the collection, except the older young one, which was still nursing

and very small and immature when the second young one was

first observed, the question is which animal was the father of the

last born ¢

Tt does not appear possible that it was the older young male, as

that must have been at least two months younger still when

conception took place.

As there is no possibility of this female having been impregnated

by any other male of her own or any other species, the only

explanation appears to be that of retarded development of one

egg or foetus.

In Darwin’s ‘ Descent of Man,’ under “ Manner of Development”

it is stated that ‘ In various Mammals the uterus graduates from

a double organ with two distinct orifices and two passages as in

the marsupials into a single organ . . . . as in the higher apes and

man”; also 27 lines further on are six lines indicating possible

functional development in the double uterus,

Did the adult male impregnate this female twice, one egg

ON DEATHS IN THE GARDENS DURING 1911, 2355)

developing in one half of the uterus while the other lay dormant

or developed more slowly in the other half during the months

necessary to account for the delay in birth ?

Male No, 22 is still living, and is a typical Macropus rufus.

PAPERS.

16. Report on the Deaths which occurred in the Zoological

Gardens during 1911. By H. G. Puimmer, F.R.S.,

T.Z.8., Pathologist to the Society.

[Received February 3, 1912: Read February 6, 1912.]

On January Ist, 1911, the number of animals in the Zoological

Gardens was 3184, and during the year 3493 animals were

admitted, making a total of 6677 for the year.

The number of deaths during the year has been 1647—-that is,

a death-rate of 24°6 per cent.; but if from the above total we

deduct 801 animals which did not live for six months after their

arrival in the Gardens—that is, the time at which we find they

have either got over their journeys, or died from any disease

they brought, with them, or have got fairly used to their new

environment—the percentage of deaths is reduced to 12°6, which

is considerably less than those of the past three years.

The following figures will show the general results of the last

four years :—

1908. 1909. 1910. 1911.

Total number of animals ......... HOO edo 0st SORE OO

Mibsudl MEAG OS: &, doc, cere fared owes de bar z.,. VAR2 ns LOG Ae UGAE

Percentage of total deaths ...... 31 28 28 24°6

Percentage of deaths, excluding

those which occurred in animals

which had not been six months

im the Garagens 4.2) siesc sh ideces i/ 17°8 16:4 12°6

In any case the figures for 1911 show a considerable im-

provement on those of the last three years, and if the relative

numbers be borne in mind, the improvement is much more

marked.

The total deaths are divided as follows: Mammals 359, Birds

849, Reptiles 439,

The following tables show the facts ascertained in outline.

Table I. sets forth the actual causes of death in each of the three

groups specified. Under Reptiles are included Amphibia.

236 MR. H, G. PLIMMER ON DEATHS

TaBLE I.— Analysis of the Causes of Death.

Reference

following.

Diseases, Mammals.| Birds. | Reptiles.| to Notes

er ce ee en ieee

1. Microbic or Parasitic |

Diseases.

Muberculosisqeee ee eee eee eee: 18 106 23 1

MiyCOSis: ay cael er eee eeniceen aa 3 103 ae 2,

JEAYAMTNOUE, — g5c00s cososonancns aodede 48 81 93 3

Septicsemiac: eae eee 3 & a 4

EAIUSGESS pont tee sea ocr Peatnnee 1 1 1

| IRericardutisime eee eee 8 3 1 5

Beritonitis saber sot escenario: 3 Wee

| Stomlatitismeerresce eeu eee 4

| Pleumitiswseee wes koiecsc cen sine: 2

JONERHOTM EIEN) Ganon onoeos aaa anasas ene 1 6

| Webydatids! Saves soccciesinsveneeeee 3 Bat Ae

WIGS Bis ce aeenistate canteens a 3 7 7

Heemogregarines bap SOMA OSE ON 3 a GY

Meallaria 5 ten Sn ontt orc eet cce cae il

Cy sbhitisiece cee: soos 1

| 2. Diseases of ee

Organs.

Broncho-pneumonia ............... Al as ce, weal 8

Congestion of lungs ............... 27 106 23 | 9

iAttelectasishh.. Aa aty.et tee: 2 - ake |

IBSROMGLMNG) ss 5ccod05 o0 v50 onbeeonse nee 4, me cee

| 3. Diseases of the Heart. |

Fatty degeneration ............... 2 4 2 |

4. Diseases of the Liver. ;

Fatty degeneration ............... 3 22 5

ee HAA aaocndenensateducineccen 2 6 |

Cirrhosis .. es 1 bag |

Angioma.. ; nseaes 1 il

5. Diseases of Alimentary 7 Tract.

Gastritis .. is uy ee 1 1

Gastric ulceration ................. | 8 Ba it

Gastro-enteritis arene eee ee 15 6 16 | 10

tention cet et a ents 39 214 20) | 11

Intussusception..................... 2 a itl

Intestinal obstruction ...........,) 1 1 a ||

| 6. Diseases of Urinary ess | |

Nephritis ..... sss sisi 48 33 8

Fibrosis of kidneys” auosescuacoeuee| 3 8 ae |

SOT Es eer) eens eee 2 12

Inflammation of oviduct ...... 2

7. Various.

CERRENNOMNE, noc csoga0000 s080000900 004 on 2 ae 1 13

SARCOMAS nce cuore eae El ose 2 on 14

SenilesDecayscnees-ce eset: 2 abe ose

Anemia without ASS 1 3

CAUSE Fei. scented eeaeey MU ar

Injuries discovered post - 4 6 = 9°

12) a

mortem

Besides those tabulated above,

120 animals were killed by order or by companions,

15] » ded of malnutrition and starvation,

1 animal was not examined,

138 animals were too stale for detailed examination,

these completing the total.

IN THE GARDENS DURING 1911. 237

In Table I, the classification is made into those diseases which

actually caused the death of the animals, but in a large number

of the Mammals, Birds, and Reptiles other lesions were present

which helped towards the fatal issue, and these are classified in

Table II. If this Table be taken together with Table I. a much

more accurate estimate of the amount of disease in the Gardens

will be arrived at,

TABLE I1.—Other Diseases found in the animals tabulated in

Table I.

Rafarehes

Diseases. Mammals. Birds. Reptiles to Notes |

| | following.|

ADA MENCULOSIS ts Cie ec tre ae cscs ees eons 10

f UE GIG: Le dadhre nasi aes neecensesnnesies eote=|

WIPNOMINODNAN dco 42 eioks sheaves want uemeb eaten 5

ESTP WHINE eee cee ee ee soot beeen cee Moses anse i

RSMICALGIAS Ie. gate «vs teaser aces toe earsctasesiseree| 5 f

3 |

Bp) was

| LEE a0 )en Boocdo pop uooeea conaadeeeeoe: serra‘

BICUBICISI Messen scence ae bateee Pan cates ae | Bh uF

| IBETOR EST HRIIE) gat tom are Ub ReREMR AC aRlcr came cia ines ae | ae 4 :

IN ERED Ee SR Re ie ee ner eae eer vere Mes | 28

[TS ewevisey es Sa gta dA ee A 1 30

NMIONTIS MRC ceE ef etnwleccnata doc duast weautelssects 13 ) ls

Hamogregarines .. Rete ached: ; *.

Pu A OS OMICS) yet. cc-cistriackisencstle yiooe sone His iL

SROMLURIUIS HEER saad achis Neaceneuestecentss

JNTICTEISES doce HRC ROBO EDC E DE GEE CELE eee oreo ere

Splenic infarcts.. |

Cystitis ..

Ste Ee Eas See a ae og

ERGIAC EMU Steep certts ceastecicae seine etc meca ele cCeee

Broncho-pneumonia rect bata

Conpestion: of lungs.c-.2\s5.c2-on ces satarsees

(Bidemmoh lumegs (asi. e.s-sctesatoraesp oe etee

Hin By SOMA 2 sees iens esos ean e-eeiee eens se

Hydrothorax

Dilated heart Eco poate tracts

IAG ReTONI aE ee wate.pe cases rectaseree aecaeonlees se

MirGbyalmetegasetasscseert- stk ereeaete c-sisecese 13 bf || 28

Me pabihisweemsre Mette ae scssetakles secs se 1] 236

Gastritis ....... eM SEs Pad 3 soe

Gastric AUC RAR NOTIN sc ceric 8s cigs esc aas 23 2 3

PA LOMUGI Sw ee cecd dinerarc siete air cans ag ptete ae 21 43, 18

Imtussuscep tions ss eter feeeccc nce s<< saree «ss 8 1 1

noe :

eben we po:

co Or Ole

Wr bn kOD-

ise)

Ve)

Nephritis ....... RAM h BEES 20 15

Fibrosis of kidneys .. 1 adach Mica ats 4

Cystic Kime ysieg ae ced Sekacat oc aerc doce los 1 - Age

INGEIEERe concen cee - o ee ats banc 4 4 11

EMATOUTS:, tiasda ned ea ee eee tad akislebe ce tertes 3 id An

Myelitis scent ar cc Sun RO ReC open Caner 2

IRTCKEES cc. che caracee esas cect ssiteetsseee| 24 ae

Watfembolisna Gece. ese. aee coe Bh a 1 16

238 MR. H. G. PLIMMER ON DEATHS

Taste LIJ.—Shows the Distribution of Diseases causing Death

amongst the principal Orders of Mammals.

Diseases. 5 5 = s S =

a | Biel 5S obits Seep alate

& oO a} =) fea} =

ANTOSROCUOS Bean oonco:c0 sen sandecone sop bese Weg 6 2 2 iL

MiyiCosis ees, aiece pecesenaeenemncve died are mien abn 3

TEINOW NOMI). Lac ccacooccaadoneabopcnsasccoouuele : Ld 17 7 7 2 iL

Sepiiesemilareecerer eee settee eerwec sacl: 2 nae

UNV OSOESIS cas hoe cdh apr aire hon eno REE SUB OTe eee Ihe coe Ke - 1 Eee 4d:

TREEIGRINGHIINS: ecco contea sescnesooeoueen coseae 2 4 1 an Bae 1

JEEIHIRCTAT PIS an boRetent naan eceCoeee teonntee so ronlin = cas ae can 3 ‘af es

Bersih seer ee i 1 a

DTS HOMTASTS Peace) aac ene eRe ed eae oe il

LEONG IS) sacatea on epaden pee sodenosnceedees came a Sac Bs 1

(ONS SIS aneree cu aeemnsneanboane nacsaoant caneaal he eae Se 1

Broncho-pneumonia .....................| 17 10 3 5 2

Clos AOM OF WOES | soo n5eenstnoose 200 one | 14 | nS 3 sae

JNPSINGINSIEY ooo bods dhacuanadmaceos end aon pbs Ei: 1 il bis

IBTONCHIGISI EEL ee mee emcees een: 3 1

Wabby Meanti eeacasccvscseene ten sacteeceeeelty Mee iene

IM at fey LIVOTA aoe ceeds na seeonesuecaee seeeees 1 1 1

JEU olininks teeta dane dastans anuashcasnes aeabae fie atc sn 2 :

(Choa NORIS: Ore INNER .ecne nsnbcoscs cooscnunsobel! ‘Gos ai ce Hl

ANTENA, O1 WII sao condom cag nciancbesodescae i” os a a oe

Gastric ulceration ............... ae Micpee 3 4 se 1

GCHOEDROGIMEEONIIS coo ancoonanonso cbeace aeoces 3 7 3 1 1

TBP TUIGISUDIS is acetates Sam men Ren nnCmUe coeecertnn aes) 9 4 4, 1 2

| 1

GHPTESURGS DOOM 445.05 cns00nb00 cop cod e960

Intestinal obstruction..................--. eh 1 a0 ae Nee 3

Nephritis .. Ce RR AE Te 10 7 1 4

Fibrosis of Kidneys Go aacaas cece tn eee 2 1 a Be nee

Stone.. ID Fete Lateran ate: i 1

Carcinoma LAS Hae aR ane 2

Seal CIB CENY Goo one bdaooe cogeobbacasy coonas.aal bate 2

UNTIED Id haat aeesscee sosg me mecabcerta seeds cee 1

JPR EHEES ose6p seston soupges ceande vadeads> sual) B00, 1] 2 ee wy 3

Notes on the foregoing Tables.

I. General.

The following calculations will show better what the figures on

page 235 mean. On the average of the past three years, taking

into consideration the different numbers of animals in each year,

the figures for 1911 should have been, roughly :—

4 Noy ie(Clern Ste eae a agnciace qopeduanooncasocs 2020

Percentage of total deaths ............... 36

Ditto, excluding those animals which

had not been six months in the

Gardens ywakneet Veena pesca tet ett 20

So that the improvement is really much greater than appears

from the mere figure-statement.

IN THE GARDENS DURING 1911. 239

The principal percentage improvement is in the Mammals,

which come out for 1911, as compared with the three previous

years, roughly as 22 is to 31.

The following table shows the deaths amongst the Mammals

from the numerically most important diseases for the four years

1908-1911 :—

1908: 1909. 1910: “1911.

Total number of animals .... 5608 5303 5540 6677

arberculosis e sedi. relies. 59 ye 18 18

Pneumonia: eh sedesee oe AZ 43 31 48

SOpticswilacivs: juss. cte cananss 12 15 7 a

Broncho-pneumonia ......... 44 DI 50 4]

Congestion of lungs ......... 29 38 4] 27

Gastro-enteritis . occu. .scseee 23 11 19 1s:

HNGEOLIGIS: fo vdst oncceremsetor ent (2 46 32 39

RSS NIbIG él van: nd. dopateak 15 4 6 3

The Birds do not show any percentage reduction of mor ‘tality ;

and the great relative mortality of birds in the Gardens is due,

in my opinion, largely to the overcrowding of many of the

aviaries.

The Reptiles show the largest percentage mortality ; this is,

in my opinion, partly due to the fact that so many of them

arrive heavily laden with parasites of all sorts—some having

as many as four different species—and partly to the present

unphysiological method of feeding them.

IT. Special.

1. The percentage of tubercle has fallen to hs per cent. in

the Mammals, 13-3 per cent. in the Birds, and is 5-4 per cent. in

the Reptiles, which is relatively less than last year all round.

In 6 Mammals and 41 birds it was a general tuberculosis,

In a Lion it was of the human type with cavitation of lungs,

and in 7 Birds it was of bovine type.

A remarkable case occurred in a young Duck three weeks old,

in which the lungs, air-sacs, and kidneys were affected, and the

growths were already caseating.

2. As before, all the mould-diseases are grouped here. In the

Mammals—2 Wallabies and a Kangaroo—it was of the same

type as described before, and due to the same organism. The

percentage among the Birds remains unchanged: three cases

were due to Aspergillus niger.

3. The deaths from pneumonia still remain very high, roughly

about the same percentage as last year. My impression, from

the post mortems, is that it is becoming more virulent in the

xardens: there have been more secondary pneumococcal affections,

240 ON DEATHS IN THE GARDENS DURING 1911.

such as enteritis, nephritis, and gastric ulceration. Many of the

Mammals were very rickety and so predisposed. In three of

the Reptiles it was due to worms: in all the other cases pneumo-

coccal.

4, One, ina Japanese Ape, was due to pyorrheea alveolaris :

the other two to deep abscesses.

5. Hight of these cases were due to pneumococcal infection.

6. In an Anoa, in which the liver was crammed with flukes.

7. The three birds had all syngamus.

8. Two of these were influenzal, and five had rickets.

9. Nine of the Mammals were badly rickety: in the Birds

there was also, in most cases, cedema of the lungs.

There has been a notable decrease of this condition in the

Mammals and Birds.

10. The cause of this condition has been traced definitely in

one instance, separately to shot, straw, grass, in two to sand, and

in seven to worms.

11. The number of cases of enteritis is relatively about the

same as last year. In the following instances the cause could be

definitely traced: 15 were due to worms, 3 to flukes, 11 to

decomposed food, 14 to sand, and 6 to foreign bodies.

12. This peowrredh in a Fox anda IResnoacien Devil.

13. The two cases in Mammals were in a Wallaroo and in a

Kangaroo ; in both the primary growth was in the stomach :

the former there were secondary deposits in glands and cael

and in the latter in liver and glands. The reptilian case was a

glandular cancer in the stomach of a Tortoise.

14. The two cases of sarcoma in Birds were ina Vulture and

a Parrakeet respectively : in the former in liver, pericardium, and

heart, and in the latter in the kidney.

15. The diseases grouped under the term ‘“ Malaria” were due

in the greater number of instances to the Hamoproteus danilewskt,

in the others to the Plasmodium precox.

16. This occurred in an Alligator after a compound fracture

of the thigh.

No. 108.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

‘ February 6th, 1912.

E. G. B. Meapr-Watpo, Esq., Vice-President,

in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The Secretary read a Report on the Additions that had been

made to the Society’s Menagerie during the months of November

and December 1911 and January 1912.

Mr. E. G. B. Meapse-Watpo, V.P.Z.S., exhibited a mounted

specimen of a pale fulvous variety of the common wild Polecat

(Putorius putorius), belonging to the Rev. D. Edmondes Owen,

of Llandovery, 8. Wales. ‘Ihe animal resembled in colour a well-

known variety of the Badger, and was interesting because the

Polecats in that particular district were frequently of this

peculiar variety and had been well known locally for years.

The Srcrerary read an account of some observations made at

the Philadelphia Zoological Garden by Mr. R. D. Carson,

C.M.Z.S., on an instance of retarded development of the foetus

in a Red Kangaroo (Macropus rufus).

* 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 ot extra charge.

to all Fellows who subseribe to tke 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.

Mrs. Rost Haig Tuomas, F.Z.8., read a paper dealing with a

breeding experiment with Pheasants undertaken to confirm a

previous one, in which it had been observed that a male Pheasant

had transmitted to his F 2 9 offspring the female plumage of his

species. ‘The following cross produced the same result :—

Phasianus Jormosus Q X Phasianus versicolor SG.

x Ph. versicolor 3.

The five hens hatched in F 2 had grown up and were all

versicolor in pattern, colour, dimensions, and moral character.

One of these hens had been kept to breed with to test her purity,

and the skins of the other four were exhibited together with the

skins of a Ph. formosus Q and a Ph. versicolor Q for comparison.

The results of these two experiments did not appear to conform

to the theory that the cock was homozygous for sex ¢ ¢.

These experiments had also shown that the male had not

transmitted to his female F 2 offspring such constant purity of

male plumage. In the first experiment, Silver x Swinhoe, out of

four males three were “Si. Sw.,” one only pure “Sw.,” and the

only two males that lived in the second experiment, formosws

x versicolor, were both “ Fo, Ve.”

Mr. J. T. Cunnincuam, M.A., F.Z.8., read a paper on Men-

delian Experiments on Fowls. The paper described the characters

of ten individuals of the F 2 generation reared from a pair of F 1’s

produced by a cross between Silky @ x Bankiva g made by

Mr. D. Seth-Smith. The characters recognised were seven in

number—namely, colour of plumage, char. acter of plumage (whether

silky or normal), comb, pigmentation of skin and internal tissues,

toes (i.¢. presence or ‘absence of extra toe), feathering of legs,

crest on head. The Silky of the original cross had white ‘ silky ”

plumage, rose comb, crest on head, double hallux, feathered legs,

and black pigmented skin. The Bankiva had black-red plumage

of normal structure, single comb, unfeathered shanks, normal

toes, and normal unpigmented skin, no crest. The dominant

characters i in the F 1 were coloured plumage of normal structure,

rose comb with crest, pigmented skin, feathered legs, and dewiblie

hallux; but several of these characters showed irregular domi-

nance or intermediate condition. The most important results

recorded in the paper were imperfect segregation in the F2

generation in at least two of the characters—namely, the absence

of pigmentation in the plumage and the absence of pigmentation

in the skin and tissues. The recessive white of the plumage

occurred in four individuals, in all of which, but especially in one,

some pigment was present. Only one specimen apparently re-

cessive with regard to the absence of pigmentation in the skin

had been examined post mortem, and in this unmistakable traces

of pigmentation in the skin and peritoneum were observed.

Mr. Cunningham criticised the recent paper by Profs. Bateson

and Punnett, in the ‘Journal of Genetics,’ on the Inheritance of

pigmentation of the Silky Fowl,and maintained that the evidence

of these authors was not sufficient to prove that segregation in

this character was complete in their experiments, and that there-

fore their results were not necessarily contradictory to his own.

In the absence of Mr. H. G. Primer, F.RB.S., E.LS., F.Z.8.

Pathologist to the Society, the Sucrerary read a Report on the

Deaths which had occurred in the Society’s Gardens during the

past year.

Mr. J. Lewis Bonnors, M.A., F.L.S., F.Z.S., read a paper on

a further collection of Mammals from Kgypt, which he had

received from Capt. 8. 8. Flower. The communication dealt

with some twenty species, of which the following two were

described as new :—

Meriones crassus pallidus, from Atbara, Sudan, similar to

M. crassus sellysii, from which it differs in its larger size, paler

coloration, and more pointed snout.

Acomys russatus egyptiacus, from the Desert near Cairo. A

race of A. russatus, from which it differs in its smaller size and

much yellower coloration.

Mr. H. Wats Kew, F.Z.S., contributed a paper ‘On the

Pairing of Pseudoscorpiones,” based on observations made by him

on living specimens of Chelifer (Chelifer) latreillii Leach and

Chelifer (Chernes) cyrneus LL. Koch. Fertilization was found to

be effected, without intromission of a copulatory organ, by means

of a spermatophore. The male and female faced one another in

walking position, the male seizing with one or both palps one or

both palps of the female. There was a period of courtship

during which Chelifer 3 protruded at intervals long ‘‘ram’s-horn

organs”; and Chernes 8, not possessing these organs, made

peculiar movements with one of the palps and with certain of

the legs. At length, the spermatophore—which differed in

character in the two species—was extruded, and was affixed to

the floor, in front of the female, where it stood erect or obliquely.

The male then retired by taking a few steps backwards, the

female at the same moment coming forwards and receiving

the male-product from the spermatophore without delay. Con-

trary to opinions previously held, the “ ram’s-horn organs ” took

‘no part in the act of fertilization; their function appeared to

be excitatory. The modified legs of the first pair of Chelifer g

were found to be used for seizing the genital-opening of the

female at the moment when the spermatophore was approached ;

there was no corresponding action in the case of Chernes 3, in

which the legs exhibited no marked modification.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, February 20th, 1912, at half-past Hight

o'clock P.m., when the following communications will be made :—

1, A. T. Masrerman, M.A., D.Sc., F.Z8.

Notes on Age-determination in Scales of Salmonoids, with

special reference to Wye Salmon. (With lantern illustrations.)

_H. Tyson Jamuson, M.A., D.Sc. Ph.D. F.Z8.

Studies on Pearl-Oysters.—I. The Structure of the Shell

and Pearls of the Ceylon Pearl-Oyster (Margaritifera vulgaris

Schumacher), with an examination of the Cestode Theory of

Pearl Production. (With lantern illustrations.)

2 Se Eon ee

Mimicry amongst the Blattide; with a Revision of the

Genus Prosoplecta Sauss.

4. The Rev. O. Pickarp-CAmprines, F.R.S., C.M.Z.S.

Contributions to the Knowledge of the Spiders and other

Arachnids of Switzerland.

The following papers have been received :—

1, Herpert L. Hawkins, M.Sc., F.G.S.

The Classification, Morphology, and Evolution of the

Kchinoidea Holectypoida.

2. H. G. Puimuer, F.R.S., F.L.8., F.Z.8.

Blood-Parasites found in the Zoological Gardens during the

four Years 1908-1911.

3. Dr. G. O. Sars, C.M.Z.8.

Zoological Results of the Third Tanganyika Expedition,

conducted by Dr. W. A. Cunnington, 1904-1906. Report on

some Larval and Young Stages of Prawns from Lake Tan-

ganyika.

4, Mrs. E. W. Sexron.

Amphipoda from Bremerhaven.

5. Rospert Broom, M.D., D.Sc., C.M.Z.S.

On the Structure of the Internal Ear, and the Relation of

the Basi-cranial Nerves in Dicynodon, and on the Homology of

the Mammalian Auditory Ossicles.

6. Epwarp W. SHAnn, B.Sc.

Observations on some Alcyonaria from Singapore, with a

brief Discussion on the Classification of the Family Nephthyide.

7. Georae H. Kenricx, F.ZS.

A List of Moths of the Family Pyralide collected by Felix B.

Pratt and Charles B. Pratt in Dutch New Guinea in 1909-10,

with Descriptions of new Species.

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

Some early Fossil Cirripedes of the Genus Scalpellum.

9, JuLIAN S. Huxury.

A First Account of the Courtship of the Redshank (Zotanus

calidris).

10. Frank E. Bepparp, M.A., F.R.S., F.Z.S.

Contributions to the Anatomy and Systematic Arrangement

of the Cestoidea.—IV. On Species of /nermicapsifer from the

Hyrax and on the Genera Zschokkeella, Thysanotenia, and

Hyracotenia.

11. Dr. BAsurorp DEAN.

Additional Notes on the Living Specimens of the Australian

Lung-fish (Ceratodus forsteri) in the Collection of the Zoo-

logical Society of London,

Communications intended for the Scientitic Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Socruty or Lonpon,

Recent’s Park, Lonpon, N.W.

February 13th, 1912.

No. 104.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON®

February 20th, 1912.

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

in the Chair,

The Minutes of the last Scientific Meeting were confirmed.

Dr. A. T. Masterman, M.A., F.Z.S., gave a demonstration,

illustrated by a large number of lantern-slides, of recent investi-

gations on Age-determination in the Scales of Salmonoids, with

special reference to Wye Salmon,

Dr. H. Lysrer Jameson, M.A., D.Se., Ph.D., F.Z.8., read a

paper “Qn the Structure of the Shell and Pearls of the Ceylon

Pearl-Oyster (Margaritifera vulgaris, Schum.); with an Kx-

amination of the Cestode Theory of Pearl Production.” ‘The

author began by reviewing the work on the subject of Pearl

Production carried out in Ceylon by Prof. Herdman, F.R.S., and

his successors. He examined the theory, enunciated by Prof,

Herdman, that most Ceylon “fine” pearls had for their nuclei

the remains of cestode larve, and that these larvae, which are

abundant in the liver and connective tissues of the Pearl-Oyster

in Ceylon, were the “cause” of the most valuable pearls.

Dr. Jameson maintained that the evidence adduced in support

of this theory by Prof. Herdman and Mr. Hornell was insutfticient,

and that the only drawings in Prof. Herdman’s ‘“ Report on

the Ceylon Pearl-Oyster Fisheries,” published by the Royal

* 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,

Society, that purported to show the remains of cestodes in

the centres of pearls, were capable of other interpretation.

Moreover, he could not find in any of the pearls sectioned by

Prof. Herdman, numbering some twenty-five or more, which

Prof. Herdman had kindly allowed him to examine, a single

instance of a cestode larva forming the nucleus. This observation

was borne out by the results of the examination of between 300

and 400 pearls from Margaritifera vulgaris, mostly from Ceylon,

but also comprising examples from the Persian Gulf, the Gulf

of Cutch, the Mediterranean, Madagascar, New Caledonia, etc.,

none of which, on decalcification, showed cestode or other platy-

helminthian larve as nuclei. The centre of such a pearl was

(where it consisted of material other than the nacre or other

normal constituents of the shell) nearly always composed of an

abnormal form of shell-substance, analogous to that formed to

repair an injury to the shell, which, owing to its opacity, might

easily be mistaken for a foreign body.

These repair-substances were sometimes associated with granular

matter, the origin of which was obscure; this matter might

perhaps be derived from the tissues, or might possibly be of

parasitic origin, but Dr. Jameson saw no reason for regarding it

as derived from a cestode larva.

Dr. Jameson regarded the presence of these cestodes as a

disease parallel to, but independent of, “ margarosis”; and

compared the case of a pearl-oyster, containing both cestodes

and pearls, to that of a man suffering simultaneously from

echinococci and scabies, or of a dog infested at the same time

with tapeworms and mange.

The author then discussed the systematic position of these

cestode larve. Prof. Herdman regarded them as a younger

stage of a Tetrarhynchus larva, which also occurred in the pearl-

oyster, and which, acting on this assumption, Shipley named

Tetrarhynchus unionifactor. The adult of this worm occurred

in the Ray (Rhinoptera javanica). Here, again, the author

maintained that two parallel forms were confused, and that the

larva which Herdman thought to be the cause of pearls was

not a younger stage of Tetrarhynchus wnionifactor, but a species

referable to T'ylocephalum or some allied genus. He had found

a worm which might be the adult of this larva in the Ray

(Aétobatis narinari). This supposed pearl-producing larva, and

a similar smaller one, also found in the Ceylon Pearl]-Oyster, were

described as new species under the names 7'ylocephalum ludificans

and 7’. minus.

Preliminary Descriptions.

TYLOCEPHALUM LUDIFICANS, sp. 0.

The worm supposed by Prof. Herdman to be the cause of

“fine” pearls in the Ceylon Pearl-Oyster. Globular resting

scolex, with conical, flattened, or concave myzorhynchus,

usually retracted within a muscular collar, which possesses

a denticulated annulus. Average diameter 0°78 mm. ;

musculature of the myzorhynchus showing no trace of

division into bundles; collar-sheath usually widely open in

the resting stage.

Resting in fibrous connective-tissue cysts in the tissues of

Margaritifera vulgaris from the Gulf of Manaar.

TYLOCEPHALUM MINUS, Sp. n.

Similar to the above but smaller, average diameter 0°14

mm. As a rule the myzorhynchus appears more conical

than in 7’. ludificans, and the collar-sheath more constricted,

with smaller aperture, in the resting stage. In some

examples the musculature of the myzorhynchus shows a

tendency to break up into bundles of fibres.

Habitat as in 7’, ludificans.

Dr. Jameson claimed that he had found 7'ylocephalum ludificans

in specimens of the Ceylon Pearl-Oyster in Dr. Kellart’s collection

in the British Museum, and had considered the possibility of

their being concerned with pearl production, and dismissed the

theory as untenable, previously to Prof. Herdman’s original

departure for Ceylon.

The second part of the paper dealt with the structure and

formation of the shell and of pearls. The various repair-

stbstances, which replace the ordinary shell substances under

abnormal or pathological conditions, were described, their relations

to the normal substances of the shell were discussed, and their

occurrence in the pseudo-nuclei of pearls dealt with. The

“caleospherules” which Herdman regarded as free concretions,

and as the cause of “ Muscle Pearls,’ were considered to be in

fact minute pearls, composed of the hypostracum, or special shell-

substance to which the muscles are attached. This conclusion

had been reached independently by Rubbel, in Marburg.

The author said that his observations on the real cause and

mechanism which led to the formation of pearls in the Ceylon

Pearl-Oyster were still too incomplete to communicate; but he

maintained that, as he had already laid down in his 1902 paper,

the real cause of pearl production would have to be sought, not

in the nuclei or pseudo-nuclei of pearls, but rather in the patho-

logical conditions under which the tissues of the mollusc gave

vise to the pearl-sac. It was only in a few cases, like the

Trematode pearls in the common mussel, that the cause of the

pearl-sac, i.e. in this case the trematode, frequently remained to

form the “nucleus” of the pearl and tell the tale of its origin.

The author had found that, as observed by Prof. Herdman, a

minority of Ceylon pearls may have foreign bodies, such as sand-

grains, in their centres.

Mr. R. Suetrorp, M.A., F.2Z.S., communicated a paper on

‘“Mimicry amongst the Blattide ; with a Revision of the Genus

Prosoplecta Sauss.,” in which he dealt with a number of exceptions

to this usually cryptically coloured type of Cockroach, and in

greater detail with the Prosoplecta, nearly all the members of

which presented a remarkably close and detailed resemblance to

other insects.

A paper entitled “A Contribution to the Knowledge of the

Spiders and other Arachnids of Switzerland,” was contributed by

the Rev. O. Pickarp-Campripcr, M.A., F.R.S., F.Z.8. It was

based on a number of specimens collected for the author by

various persons, at different times, and contained the description

of one new species.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, March 5th, 1912, at half-past Hight

o'clock p.m., when the following communications will be made :—

1. Exhibitions and Notices.

2. Hersert L. Hawxins, M.Sc., F.G.S.

The Classification, Morphology, and Evolution of the

Kehinoidea Holectypoida. (With lantern illustrations.)

Blood-Parasites found in the Zoological Gardens during the

four Years 1908-1911. (With lantern illustrations.)

4. Dr. G. O. Sars, C.M.Z.S.

Zoological Results of the Third Tanganyika Expeditior,

conducted by Dr. W. A. Cunnington, 1904-1906. Report on

some Larval and Young Stages of Prawns from Lake Tan-

ganyika.

5. Ropert Broom, M.D., D.Sc., C.M.ZS.

On the Structure of the Internal Ear, and the Relation of

the Basi-cranial Nerves in Dicynodon, and on the Homology of

the Mammalian Auditory Ossicles.

1. Epwarp W. Suann, B.Sc.

4. Jutian 8S. Huxtey.

The following papers have been received :—

Observations on some Alcyonaria from Singapore, with a

brief Discussion on the Classification of the Family Nephthyide.

Grorce H. Kenrick, F.Z.%.

Meera sa Mes T leet.

A List of Moths of the Family Pyralide collected by Felix B.

Pratt and Charles B. Pratt in Dutch New Guinea in 1909-10,

with Descriptions of new Species.

Tuomas H. Wiruers, F.G.S.

Some early Fossil Cirri pedes of the Genus Scalpellum.

A First Account of the Courtship of the Redshank (Z'otanus

calidris).

5. Frank E. Bepparp, M.A., F.R.S, F.Z.8.

Contributions to the Anatomy and Systematic Arrangement

of the Cestoidea.—lV. On Species of /nermicapsifer from the

Hyrax and on the Genera Zschokkeella, Thysanotunia, and

Hyracotenia.

Dr. BAsurorp DEAN.

Additional Notes on the Living Specimens of the Australian

Lung-Fish (Ceratodus forsteri) in the Collection ot the Zoo-

logical Society of London.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZooLoGicaL Socrety or Lonpon,

ReEGENT’s Park, Lonpon, N.W.

February 27th, 1912.

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No. 105.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.’

March 5th, 1912.

Sir Joun Rost Braprorp, M.D., D.Sc., F.R.S.,

Vice-President, in the Chair,

The Minutes of the last Scientific Meeting were confirmed.

Mr. OuprieLp Tuomas, F.R.S., F.Z.S., exhibited the skull of a

German Wild Boar from Baden and of a Hungarian Wild Boar

from Kolozsvar, the latter recently presented to the British

Museum by Friiulein Sarolta von Wertheimstein.

The difference in size between these was so great that

Mr. Thomas considered that the Hungarian Boar should be

separated as a distinct species, which he proposed to call Sus

attila.

He also stated that the North and South Spanish Wild Boars

were, on the other hand, so much smaller in different degrees

than the German animal as each to deserve subspecific distinction

from the latter.

The three forms now described were therefore as follows :—

SUS ATTILA, sp. n.

Upper length of skull 452 mm.; height of skull on lower

jaw 271.

Hab. Hungary and Russia, Type locality Kolozsvar.

Type. Male. B.M. No. 12.1.23.1

SUS SCROFA CASTILIANUS, subsp. n.

Upper skull length 353 mm.; height 198.

Hab. North Spain. Type locality Burgos.

Type. Male. B.M. No. 11.10.5.3.

* 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,

Sus SCROFA B-ETICcUS, subsp. n.

Upper skull length 324 mm.; height 208.

Hab. South Spain. Type locality Coto Doinana.

Type. Male. B.M. No. 8.3.8.12.

Mr. H. L. Hawks, M.S8e., F.G.S., read a paper, communicated

by Dr. Henry W Saeed “RRS V. P.Z.S., on “The Classifi-

cation, Morphology, and Evolution of the Echinoidea Holecty-

poida,” illustrated with lantern-slides. The classification of the

Mesozoic Gnathostomatous Irregular Echinoids was revised, with

diagnoses of the families, subfamilies, and genera, and a new

genus and subgenus were introduced. The anatomy of the test

was described for the Holectypoida, and compared with that of

other Orders. The origin of the Irregular Echinoids was dis-

cussed, and the lines of evolution that they followed were indi-

cated and summarized in a genealogical table.

Mr. H. G. Puimmer, F.R.S., F.Z.S., Pathologist to the Society,

read a paper “On the Blood-Parasites found in the Zoological

Gardens during the four years 1908-1911,” illustrating his

remarks with a lar ge number of Jantern-slides. The paper con-

tained the results of examination of the blood of 6430 animals,

in about 7 per cent. of which parasites were found. Many of

these parasites were described for the first time, and in other

cases the hosts were newly recorded.

Prof. G. O. Sars, C.M.Z.S., presented a memoir entitled

* Zoological Results of the Third Tanganyika Expedition, con-

ducted by Dr. W. A. Cunnington, 1904-1906. Report on some

Larval and Young Stages of Prawns from Lake Tanganyik

Four forms were dealt with in this paper, two of which repre-

sented very early larval stages, and apparently belonged to two

quite different kinds of Prawns, but owing to the difficulty of

deciding with any certainty the species or even the genera to

which they were referable, they were not named, although a

detailed description was given and their probable origin sug-

gested. The remaining two forms represented a larva in the last

stage and a very young Prawn in the first post-larval stage, and

both were referred to a definite species.

Dr. Rosert Broom, D.Se., C.M.Z.8., communicated a paper

“On the Structure of the Internal Ear, and the Relation of the

Basi-cranial Nerves in Dicynodon, and on the Homology of the

Mammalian Auditory Ossicles,”’ the first part of which contained

an account, the first on record, of the bony labyrinth enclosing

the internal ear and the nerve foramina in relation thereto, in the

skull of the extinct Reptile, Dicynodon. The author had obtained

a cast of the internal ear of a specimen in which the matrix was

eo >

i Al ate

hardened by epidote and the bone mainly calcareous. By dis-

solving away the calcareous matter replacing the bone, he had

satisfactorily traced the structure of the ear. The author stated

that the vestibule was very remarkably elongated and that there

was no trace of a cochlea. The semicircular canals were of the

normal reptilian type.

The homologies of the auditory ossicles were discussed, and it

was shown that the bone which Dr. Broom had formerly regarded

as the tympanic was really the stapes. In the light of his new

observations the author stated that he was now prepared to accept

the view that the incus is the homologue of the reptilian quadrate,

the malleus the articular, and the tympanic the angular.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, March 19th, 1912, at half-past Eight

o'clock p.m., when the following communications will be made :—

1. A. Rapciyrre Duemore, F.R.G.S.

Lantern Exhibition of Studies of Wild Animals in Africa and

North America.

2. Epwarp W. Suany, B.Sc.

Observations on some Alcyonaria from Singapore, with a

brief Discussion on the Classification of the Family Nephtbyide.

3. Georce H. Kenrick, F.Z.8.

A List of Moths of the Family Pyralide collected by Felix B.

Pratt and Charles B. Pratt in Dutch New Guinea in 1909-10,

with Descriptions of new Species.

4. Tuomas H. Wiruers, F.G.S,

Some early Fossil Cirripedes of the Genus Scalpellum.

The following papers have been received :—

1. JuuiaAn 8. Hvuxuey.

A First Account of the Courtship of the Redshank (Totanus

calidris).

2h Frank KE. Bepparp, , M. A,, alle RS. 7 HZ: S.

Contributions to the Anatomy ad Sy stematic Arrangement

of the Cestoidea.—IV. On Species of Jnermicapsifer from the

Hyrax and on the Genera Zschokkeella, Thysanotenia, and

Hyracotenia.

3. Dr. BasHrorD DEAN.

Additional Notes on the Living Specimens of the Australian

Lung-Fish (Ceratodus forsteri) in the Collection of the Zoo-

logical Society of London.

4. Mrs. KE. W. Sexton.

Amphipoda from Bremerhaven.

5. W. M. Smatiwoop and Euizasets G. CLARK.

Chromodoris zebra Heilprin, a distinct Species.

6. C. Tate Racan, M.A., E.ZS.

Descriptions of new Fishes of the Family Loricarnde in the

British Museum Collection.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society or Lonpon,

ReGent’s Park, Lonpon, N.W.

March 12th, 1912.

-I

10.

11.

13.

16.

_ On the Milk-Dentition of the Ratel. By R. Lyprxxer. (Text-figs. 31 & 32.)

PAPERS.

. Distant Orientation in Amphibia. By Brucu F. Cummines. (Text-figs. 4,5.) ..-.+-

Some Remarks on the Habits of British Frogs and Toads, with reference to

Mr. Cummings’ recent communication on Distant Orientation in Amphibia. by

G. A. Boutmnamr, F.R.S., F.Z.8. «1 ee ee eee ee eee ee nies haeet at teats ;

. Game Sanctuaries and Game Protection in India. i) E. P. Srzszine, F.L.S., F.Z.8.,

BR GSe ses eae er alenaneg Ih vapatans reerealr a ccral Ave share loa situ aittalclsreye ois Paradies Aiehierece Ga tsi

. On the Moulting of an Arctic Fox (Vulpes lagopus) in the Society’s Gardens. By R. I.

Pocock, F.R.S., F.L.S., F.Z.S., Superintendent of the Gardens, (Text-figs. 6-13.)

. On the Moulting of the King Penguin (Aptenodytes pennanti) in the Society’s Gardens.

By Davin Sera-Surru, F.Z.S., M.B.O.U., Curator of Birds. (Pl. I.)

_ On the Presence of two Ovaries in certain British Birds, more especially the Falconide.

By T. E. Gunn, BLS. (Pls. TLV.) 2 cence cciee ses cteten see ee eres pe Cee re

On some Collembola from India, Burma, and Ceylon; with a Catalogue of the

Oriental Species of the Order. By A. D. Iums, D.Sce., B.A., Forest Zoologist to the

Government of India; late Professor of Biology, Muir College, and Fellow of the

University of Allahabad. (Pls. VI.-XIf. and Text-figs. 14, 15.) ....++-- :

. Ontogenetical Transformations of the Bill in the Heron (Ardea cinerea). By Prof. P.

P. Susnxry, C.M.Z.S., Kharkov, Russia. (Pl. XIIL.)...... PINS Aine Regiee ‘

_ The Duke of Bedford’s Zoological Exploration of Eastern Asia.—XV. On Mammals

from the Provinces of Sze-chwan and Yunnan, Western China. By O.prienp

THomas, F.R.S., F.Z.8.....0+..05--: RTPI Wheels isrevaret there ate ee ISAC soe cover at Aarau oN ets

The Freshwater Crayfishes of Australia. By Guorrrry Suirn, M.A., Fellow of New

College, Oxford. (Pls. XIV.-XXVII. and Text-fig. 18.)

Structure of the Alimentary Canal of the Stick-Insect, Bacillus rossii Fabr.; with a

Note on the Parthenogenesis of this Species. By Aurrup E. Cameron, M.A, BiSe;,

Fullerton Scholar of the University of Aberdeen and Research Student in the

University of Manchester. (Pls. XXVIIL.—-XXX.)....-..+-+4-- Ber peek by. tes

. Diagnoses of new Species of Terrestrial and Fluviatile Shells from British and

German East Africa, with the Description of a new Genus (/wssoia) from Eusso

Nyiro River, B.E. Africa. By H. B, Pruston, F.Z.8. (Pls. »[email protected] tgp. 0.0.4] aie Br

Contributions to the Anatomy and Systematic Arrangement of the Cestoidea.—-III. On

a New Genus of Tapeworms (Otiditenia) from the Bustard (Hupodotis kori). By

Frank E. Bepparp, M.A., F.RB.S., F.Z.S., Prosector 'to the Society. (Lext-figs. 19-30.)

_ On a Further Collection of Mammals from Egypt and Sinai. By J. Lewis Bonnotr,

MV A\,, Hn Sue ZSiicsrs aiecae Sis ye ae eiPetal eve eel er toter sakes Pel at earner nash hater ataa vet sare

Report on the Deaths which occurred in the Zoological Gardens during 1911. By

H. G. Pumner, F.R.S., F.Z.S., Pathologist to the Society .

Pa ge

LIST OF PLATES.

1912, Part I. (pp. 1-240).

Plate Page

I. Moulting of Aptenodytes pennanti ........065 ts eyes Monee 60

II. Paired Ovaries of Accipiter nisus and Falco tinnunculus .. \

III. Paired Ovaries of Circus, Falco, etc. -........06+ Sein Stas

IV. Ovaries of : 1. Accipiter nisus. 2. Circus cyaneus. 3. Podi- & 63

COEDS CLISTATUS (5 weve 2 o's Wid SHON aad BOON TRL 5.0 : |

V. Ovaries of Circus cyanews ss. ..% SE Se mee MCA )

VI.)

VII. |

VETS s

TX. }'Onientiali@ollembola cies owe le cece ck ede ee eels wh cee, ere

xX, |

XI. |

X11. /

XIII. Development of the Bill in Ardea cinerea .veveeeesseees 125

RV ASHACOMSIS! 7772000277205.) oie seru lose tetsnetuloneheteraio vel Nic)sirs cl sim ereieieneis \

OVC eAStRCORSES FRURICLLIIG, P\n sucletain wig aiioetetnrG atic nines Bre aie GACT

OVS: VAstadopsis Sanaa: (i cnis shes tie cies ee eRe etelo gle iste inna

MOVER, > Astacopsis serratus, \O tic sci tiers she duwtaeetebe s sieievss sie hake

XVIII. Astacopsis serratis, J se ceeeeeeacectaes BPA Weie/elciits) sien stare l

ene \ Astacopsis hershawt; S oe. .cee cece alehefeie ctvalaveeeieu aie ines |

XXI. Paracheraps bicarinatus, Gs... cece ee cee eee eee ewes

XI. . Charaps tonuimants, Bi) vee s dese ieee be case sete we ria

XXILL. Cheraps guinguecarinatus, Si. seees scene cece cede cant |

XXIV. 1. Cheraps quadricarinatus, G. 2. C. intermedius, 3 ...- |

XXV._ Astacopsis f rankl WA Peaern ete ee erm eer fer cutenaene

XXVI. 7-11. Astacopsis franklinii, 12-14. A. serratus. 15-24. |

Paracheraps RGR SAA GN IBISS AOS ASI 8

XXVIII. 25-29. Cheraps quinguecarinatus. 380). CL tenuimanus.

31-33. C. quadricarinatus. 34. C. intermedius ....2.., )

XXVIII. | :

XXIX, | Alimentary Canal of Bacillus rosstt 2. ...6. cece eee eee 172

XXX.

XXXT. \ New African Terrestrial and Pluviatile Shells............ 183

NOTICE.

‘The ‘ Proceedings’ for the year are issued in fowr parts, paged consecutively,

so that the complete reference is now P. Z. 8.1912, p.... The Distribution

is as follows :— ‘

Part I, issued in Mareh.

Petra E18 Ud + June.

et Re oe September.

BA ee Bye December.

« Proceedings,’ 1911, Part IV. (pp. 869-1213), were published on

‘December 15th, 1911.

‘The Abstracts of the Proceedings,’ Nos. 103 to 105, are

contained in this Part.

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIBTY

OF LONDON,

1912.

PART If.

CONTAINING Paces 241 to 504, with 28 Puares

AND 31 TExt-FIGURES.

n | -

/ “ASti¢, :

AUG 9 1917

+“)

JUNE 1912.

PRINTED FOR THE SOCIETY,

SOLD AT THETR HOUSE IN REGENT’S PARK.

LONDON :

MESSRS. LONGMANS, GREEN, AND CoO.,

PATERNOSTER ROW.

[Price Twelve Shillings. |

LEST. OF WC ONE Nes:

1912, Parr II. (pp. 241-504).

EXHIBITIONS AND NOTICES.

Page

Dr, A. T. Masrerway, M.A., F.Z.S. Demonstration of recent investigations on Age-

determination in the Scales of Salmonoids .......... SF tigie te ob ote cncta hs eae 390

Mr. Ouprreip Tuomas, E.R.S., F.Z.S. The Races of the European Wild Swine: ........ 390

Mr. A. Rapcuyrre Duenworz, F.R.G.S. Lantern exhibition of photographie studies of wild

animals in British East Africa and Newfoundland ................. Een 25: . ¥498

~ Messrs. B. Gurrarp & Sons. Exhibition of the head of a cow Sable-Antelope, the skulls

of an Isabelline Bear, a Leopard, and a Lion, all of which showed some abnormality.. 498

Mr. Orprietp Tuomas, F.R.S., F.Z.S. Two new Genera and a new Species of Viverrine

Carnivora. (Text-figs. 61-63.) ....-- Ge NON ee aS! Tali Soke Steps keene Sonate ees 498

Mo. Guy Donuman. A new Snub-nosed Monkey. oc 000500. e uu slew «cle os eevee oleae Owes

PAPERS.

17. Mendelian Experiments on Fowls. By J. T. CunninauaM, M.A., BZ.8.7 ca eeioee eee

18. Studies on Pearl-Oysters and Pearls.—I. The Structure of the Shell and Pearls of the

Ceylon Pearl-Oyster (Mangaritifera vulgaris Schumacher): with an Examination of the

Cestode Theory of Pearl-Production. By H. Lysrzr Jamuson, M.A., D.Sc., Ph.D,

F.Z.S. (Pls. XXXITI--XLVII. and Text-figs. 83-41.) 0.0.06 ce pe enn ee eee ee eT)

19. Mimicry amongst the Blattide; with a Revision of the Genus Prosoplecta Sauss., and

the Description of a new Genus. By Roserr Suenrorp, M.A., B.ZS. (Pl. XLVI.

and Text-figs. 42-46.) 2. ca ceea cece ees e ce neces cece ccs cece nue em ates alge aoe 398

Contents continued on page 3 of Wrapper

THE ZOOLOGICAL SOCIETY OF LONDON,

Turs Society was founded in 1826 by Sir Sramrorp Rarruus,

Mr. J. Sasine, Mr. N. A. Vicors,

and other eminent Naturalists,

for the advancement of Zoology and Animal Physiology, and for the

introduction of new and curious subjects of the Animal Kingdom,

and was incorporated by Royal Charter in 1829.

Patron.

HIS MAJESTY

THE KING.

COUNCIL.

HIS GRACE TH" DUKE OF BEDFORD, K.G., President,

Tue Kant or Atramont, F.S.A.

Sire Jonn Rose Braprorp,

K.C.M.G., M.D., D.Sc., F.B.S.,

Vice-President.

Ricuarp H. Burns, Ese., M.A.

Lr.-Cotr. Sire R. Havetock

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

AtrreD Henrace Cocks, Esa.,

M.A.

F. G. Dawrrey Drewitr, Ese.,

M.A., M.D.

Crartes Drummonp, Ese,

Treasurer.

Sir Epwarp Duranp, Br., C.B.

F. Do Cana Gopmay, Kse.,D.C.L.,

F.R.S.

Srtr Epmunp G. Lover, Br., Vice-

President.

| Epuunp G. B. Merapz-Watpo,

Ese., Vice-President.

Prorrssor Epwarp A. Mrincuiy,

M.A., F.RAS., Vice-President.

Cuatmers Mrirenent, Esea.,

Ms,” B8e.,- LED. BRS:

Secretary.

W. R. Oeitvin-Grant, Ese.

ALBERT Pam, Esa.

iE:

Aprran D. W. Pottocg, Esa.

Oxprietp Tuomas, Ese., F.R.S.

AntHony H. Wryeriexp, Ese.

A.Suira Woopwarp, Esa.,LL.D.,

F.R.S., Vice-President.

Henry Woopwarp, Ese., LL.D.,

F.RAS., Vice-President.

The Society consists of Fellows, and Honorary, Foreign, and

Corresponding Members, elected according to the By-Laws. It

carries out the objects of its foundation by means of the collection

of living animals, by its Library, and by its Scientific Publications.

The Office of the Society, Regent’s Park, N.W., where all com-

_ munications should be sent, addressed to ‘The Secretary,” is open

from Ten till Five, except on Saturdays, when it closes at Two p.m.

The Library, under the superintendence of Mr. F. H. Waterhouse,

is open daily at the above hours, except in 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 Oeteber, 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.Se., F.R.S., Prosector, assisted by Mr. H. G. Pliimmer, F.R.S.,

M.R.C.8., Pathologist to the Society.

TERMS FOR THE ADMISSION OF FELLOWS.

Frrrows 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 Frtiow until the Admission Fee and

first Annual Subscription have been paid, or the annual payments

have been compounded for.

Fritows elected in November and December are not liable for

the Subscription for the vear in which they are elected.

PRIVILEGES OF FELLOWS.

Fettows have Personal Admission to the Gardens upon signing

their names in the book at the entrance gate, and may introduce

‘'wo Companions daily.

The Wire or Husnanp of a Frttow can exercise these privileges

in the absence of the Fellow.

Tivery Fertrow is at present entitled to receive annually 60

undated Green Cards, and, when no specific instructions are

received, the supply will be sent in this form. 20 Green Cards

may be exchanged for a book containing 2 Orders for each

Saturday * throughout the year. A similar book of Sunday Orders

may also be obtained in lieu of 20 Green Cards. A Green Card

may also be exchanged for 2 Buff Cards for the use of Children

under 12 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.

The annual supply of Tickets will be sent to each Frtrow on the

Ist. of January in every year, upon filling up and returning the form

of Standing Order supplied to Fellows.

Frttows are not allowed to pass in friends on their written

order or on presentation of their visiting cards.

FrLtows are exempt from payment of the fee for Painting,

Sketching, and Photographing in. the Society’s Gardens.

Frttows have the privilege of receiving the Society’s ordinary

Publications issued during the year upon payment of the additional

Subscription of One Guinea. This Subscription is due upon the

1st. of January, and must be paid before the day of the Anniversary

Meeting, after which the privilege lapses. Frrtows are likewise

entitled to purchase these Publications at 25 per cent. less than

the price charged to the public. A further reduction of 25 per

cent. is also made upon all purchases of Publications issued prior

to 1881, if above the value of Five Pounds.

Fetxows also have the privilege of subscribing to the Annual

Volume of ‘ The Zoological Record,’ which gives a list of the Works

and Publications relating to Zoology in each year, for the sum of

One Pound Ten Shillings. Separate divisions of volumes 39 to

42 can also be supplied. Full particulars of these publications can

be had on application to the Secretary.

* The Saturday Orders are not available if the Fellow introduces friends

personally on that day.

Frertows may obtain a TRANSFERABLE Ivory Ticker admitting

two persons, available throughout the whole period of Fellowship,

on payment of Ten Pounds in one sum. A second similar ticket

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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 Frtiow, 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, 1912.

MEETINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON

FOR

SCIENTIFIC BUSINESS.

1912,

OES DA. OCTOBER nea e 29

a NovEMBER ...... 12 & 26

The Chair will te taken at half-past Light o'clock in the Evening

precisely.

ZOOLOGICAL SOCIETY OF LONDON.

LIST OF PUBLICATIONS.

Tue scientific publications of the Zoological Society of London

are of two kinds—“ Proceedings,” published in an octavo

form, and ‘‘ Transactions,” in quarto.

According to the present arrangements, the “ Proceedings

contain not only notices of all business transacted at the scien-

tific meetings, but also all the papers read at such meetings

and recommended to be published in the “‘ Proceedings ”’ by

the Committee of Publication. <A large number of coloured

plates and engravings are issued in the volumes of the

“ Proceedings,” to illustrate the new or otherwise remark-

able species of animals described therein. Amongst such

illustrations, figures of the new or rare species acquired in a

living state for the Society’s Gardens are often given.

The “ Proceedings” for each year are issued in four

parts, paged consecutively, in the months of March, June,

September, and December. From January 1901 they have

been issued as two half-yearly volumes, inilexed separately.

An “ Abstract of the Proceedings ” is published by the

Society on the Tuesday following the date of Meeting to

which it refers. It is 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 publi-

cation at the price of Sixpence, or, if desired, sent post free

for the sum of Six Shillings per annum, payable in advance.

The ‘ Transactions ” contain such of the communications

made to the scientific meetings of the Society as, on account of

the nature of the plates required to illustrate them, are better

adapted for publication in the quarto form. They are issued

at irregular intervals.

Fellows and Corresponding Members, upon payment of

a Subscription of One Guinea before the day of the Anni-

versary Meeting in each year, are entitled to receive the

Society’s Publications for the year. They are likewise

entitled to purchase the Publications of the Society at 25 per

cent. less than the price charged for them to the Public. <A

further reduction of 25 per cent. is made upon purchases of

Publications issued prior to 1881, if they exceed the value of

five pounds.

Fellows also have the privilege of subscribing to the

Annual Volume of the Zoological Record for a sum of 30s,

(which includes cost of delivery), payable on the 1st. of July

in each year; but this privilege is forfeited unless the

subscription be paid before the Ist. of December following.

The foilowing is a complete list of the publications of the

Society already issued.

TRANSACTIONS* OF THE ZOOLOGICAL SOCIETY OF LONDON.

4to. 19 vols. and Index. Be Brice toe

Vol. I., containing 59 Plates.... (1833-35) .... £318 6.... £418 OF

, IL, iy TAs yp oh we ASS OR4T) ss a OO 5 6 6f

op oI, as Gola oh, (1842249) 0.2 oe 8 Vas! 411 O*

ae (ins 55 (USI) se00 OD 4.0), 8 2 6f

s V., 3 OY as .. (1862-66) .os. 0614 3°. 61950

ap A $ OE a5 soy Cl) coca JU & ©; 16) 6 ©

oy LOGS . dour. SOIT). 5.610140: 13 12 0

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ose IBAA GE) 5, soo (ISTO=10) sooo HA i GO, Loa ae

xe Jom GSi(E719)- ee, c LO MOR eS a iIgeg @

Index, Nola RoN nen an eee (S338=/9)) ag Oe Oe ae POR ORG

Vol xe icontaiming; 97 lates. (s80589)) fac) ON Oey elem

9 XIN, i 5 GD py 56 (IBSESO) oye G 8 O | (ir an

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ee OXI. wa A oy on GICBE“8IS) oben OO O.. Co O

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yi DNS a CL oo (IVE CG) ey meek sees IS. ©

my OM WHS ieee CUE a Ones BS O

poe PNulEXe Deore, UECPaINO) acd) 45 Oseoo IS le o

» XX.—Part 1. (Pls, l= Vay (Heb soi?) ae OS Ole 1 4 0

Pe CXC hd: (Bis VA. Say) GAYoml NUE) 5. 2 Oe 3.0 0

PROCEEDINGS OF THE COMMITTEE OF SCIENCE AND

CORRESPONDENCE OF THE ZOOLOGICAL

LONDON. 8vo. 2 vols. (Letterpress only).

Part I. 1830-31. lvol. 8vo. .

Ii. 1832.

” 9

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PROCEEDINGS OF THE ZOOLOGICAL SOCIETY OF LONDON.

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» LIL 1835. 99 Als Gak oo (OS 9) XI. 1843. 90 ASNGGa ROSS

pe LV 1886s 6 4s. 6d. .. 6s. » II. 1844. es 4s. Gd. .. 6s:

., We USB. 9 As. 6d. .. 6s. Se ACUI RSA: - 4s. 6d. ... 6s.

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ee XVIII. 1850. 39 As, 6d. OSA5 tetera i 8 @ 4 SO

as XIX. 1851. 9 As. 6d. OSH Paes Oils O . Or

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Index 1848-1860. 90 As. 6d. 6s.

iP Out of print.

* In consequence of a re-arrangement of the stock of the ‘Transactions, the Society is

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ON MENDELIAN EXPERIMENTS WITH FOWLS. 241

17. Mendelian Experiments on Fowls.

By J. T. Cunnincuam, M.A., F.Z.S.

[Received October 25, 1911: Read February 6, 1912.]

I. Experiment 1.—Bankiva 3 x Silky 9Q.

Crosses between Silky fowls and other breeds have frequently

been tried, both before and since the formulation of the Mendelian

doctrine. The silky character of the feathers has always been

found to be recessive, and this was noticed before the recently

awakened interest in Mendel’s theory. Thus Tegetmeier (1867,

p- 224) found that the F, hybrids of Silkies with other breeds had

normal plumage, and Darwin (1876, chap. vii.) noted the same

result in the offspring of a white Silky hen mated to a Spanish

cock,

Bateson (1909, pp. 34, 35, 181) mentions the heredity of the

comb, of the silky character, and of the pigmentation of the skin,

describing experiments on the latter character in detail. He

also discusses (ibid. p. 103) the heredity of the white colour of

the Silky. ~Bateson’s results and conclusions will be discussed in

reference to the results of the experiment here recorded. CC. B.

Davenport (1906, p. 55) describes a cross between Silkies and

Frizzled fowls, but he reared only the direct offspring of the

cross, the F, generation.

My experiment was made with a pair of F,’s kindly given to

me by Mr. D.Seth-Smith. These had been reared in the Society's

Gardens from a cross between a male of the Jungle Fowl (Gallus

bankiva) and Silky hens, and my attention was called to the

interest of this cross by Mr. Frank Finn, F.Z.S. The Jungle

cock may not have been perfectly pure bred, but there is no

evidence that its pedigree was impure. The characters of these

original parents are as follows :—

Jungle Fowl ¢.

Comb, single.

Colour of plumage, black-red: breast and tail-feathers black,

neck and saddle hackles yellowish red.

Skin, unpigmented : comb and wattles red.

Legs, clean, not feathered.

Feet, normal, four toes on each.

No erest on head.

Silky Fowl 9°.

Comb, small, rose.

Colour of plumage, white.

Skin, black; comb and wattles black-red, ear-lobes blue.

Legs, feathered.

Feet, with double hallux on both.

Crest of feathers on head behind comb.

Proc. Zoot. Soc.—1912, No. XVI. 16

249, MR. J. T. CUNNINGHAM ON

Tous, including the silky character of the feathers, contrasted

with the normal, we have seven pairs of characters the heredity:

of which is to be investigated.

In the F, specimens the following characters are dominant :—

‘oloured plumage.

Normal structure of plumage.

Rose comb.

Black pigmentation of skin.

Double hallux,

Feathering of legs,

Crest on head.

But the dominance is somewhat irregular. Thus, one hen of

F, at the Gardens had a single comb. Several of the F,’s had a

normal foot on one side and a double hallux on the other. The

majority had the double hallux on both sides. In my pair the

cock has both feet normal and the hen has both feet abnormal,

with double hallux. This irr egularity of dominance of the extr

toe has been noticed by previous experimenters. The crest is

dominant, but much smaller in the F,’s than in the Silky. The

skin in my pair is pigmented, but not so dark as in the Silky ; it

looks blue instead of black. The feathered leg is dominant, the

feathering being present in all the F,’s but the feathering is less

than in the Silky. These facts on the whole agree with the idea

of dominance as the presence of a positive character (presence

and absence theory of dominant and recessive), which in the F,

is present only in one half of the zygote, 7. e., m one of the gametes

which are wnited in the cross, and therefore naturally does not

produce so much effect as when it is carried by both gametes.

Thus the crest, the feathering of the legs, and the pigmentation

of the skin may be said to “be intermediate in F, between the

conditions in the two parents. The dominance, however, seems

sometimes to fail, as in the case of the specimen with a single

comb, and the specimens with double hallux on one foot only | or

on neither. The failure of dominance of the rose comb is not

mentioned by Bateson, who states that the dominance of the rose

comb is very definite and that it is generally quite impossible to

distinguish pure rose from the heterozygous type containing

single. On the other hand, exceptions to “the dominance of the

extra toe are common, and are mentioned by Bateson, who states

that it occurs in all cases yet studied in birds and mammals.

In all F, specimens the coloured plumage and the normal

structure of the feathers are present; these characters are

completely dominant.

In other crosses which have been described in the experiments

above referred to much greater irregularity of dominance has

heen observed. Thus Bateson found, with regard to the pig-

mentation of the skin, that when the Silky hen was crossed with

a Brown Leghorn cock all the F, offspring, both male and female,

MENDELIAN EXPERIMENTS WITH FOWLS, 243

were either destitute of pigmentation or showed only a small

amount of it in certain parts. But when the opposite cross was

made, Silky cock with Brown Leghorn hen, the F, males had

unpigmented skins, while the females were almost as much pig-

mented as the pure Silky hens. Bateson explains these results

by the assumption that there are three pairs of characters or

allelomorphs affecting the results, namely :—

Presence and absence of femaleness.

Presence and absence of the black pigmentation.

Presence and absence of a factor which can suppress or

mask the development of the pigmentation.

He assumes that this last factor is double or homozygous in the

Brown Leghorn cock, single or heterozygous in the Brown Leghorn

hen, and absent in the Silky. There is no need for such assump-

tions in the case of the Bankiva cock, but it must be noted that I

have no experience of the opposite cross, Bankiva hen with Silky

cock.

In the experiment here considered the white colour of the

Silky plumage is perfectly recessive. This result also differs

somewhat from that of previous experiments. Davenport crossed

a Silky cock with Frizzled hens, but he states that the colour of

Frizzles is not constant but variable. His hens were of various

colours—dark brown, yellowish, light brown, and mixed black

yellowandred. Some of them may have been carrying white asa

recessive. Of his F, hybrids 7 were white and 25 coloured.

Davenport considers that the results do not agree w ith either the

hypothesis that white is recessive or that itis dominant. He con-

cludes that the white is not always dominant, but this seems to be

a slip: he probably means that the colour is not always recessive.

Bateson found that the white of the Silky, though recessive to

colour, gave coloured offspring when crossed with certain other

recessive whites obtained from crossings. He concludes, there-

fore, that the whiteness of the Silky is due to the absence of one

of two elements which together produce colour. We shall have

to consider this point again in discussing the colours of the F,’s

which form the point of chief interest in my results.

We now come to the consideration of the F, generation, that

is of the chicks which I have hatched and reared from the pair

of F,’s abovementioned. The first brood was hatched on May 15th,

1911, and consisted of nine chicks, one of which died within the

first ae days; the other eight are still alive and vigorous. The

hen afterwards sat again, but this time hatched only two chicks,

probably in consequence of the very dry weather; these two

were hatched in August. The following table (p. 244) shows the

characters of these chickens.

The numbers of these specimens are, of course, too small to

afford evidence of any importance with regard to the numerical

proportions in which the characters or combinations of characters

16*

NNINGHAM ON

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occur, and it is not the object of this paper to draw conclus

as to factors or segregation from the numerical proportions.

is assumed that characters united in F, tend to segregate in F,,

and that the explanat

s the separation in

ion 1

gat

this segre

1on of

the gametes of F, of the pairs of factors, so that half the gametes

MENDELIAN EXPERIMENTS WITH FOWLS. 9AD

contain one character and the other half its allelomorph, as it is

called. My object here is merely to draw attention to certain

points in my results which indicate that the segregation is not

necessarily complete. If we consider single characters and take

only the eight chicks of the first brood, omitting the one that

died, and the two of the second brood, we see that the Mendelian

proportion of 3 dominants to 1 recessive is, in many of the

characters, either exactly or approximately exhibited. ‘Thus we

have in the colour of the plumage 6 coloured to 2 white, in the

structure of the feathers 6 normal to 2 silky, in the combs 5 rose

to 3 single, in the colour of the skin 6 black to 2 normal, in the

crest 6 with crest to 2 without. The double hallux, however,

oceurs only once, instead of six times, and as the male F, had

normal feet although the extra toe character was present in the

cross, it is by no means certain that this character was segregated

in his gametes. The feathering of the legs and the crest on the

head exhibit a condition which does not harmonise exactly with

the Mendelian theory. The leg-feathering is present in all the

specimens except No. 10 in the second brood, but it exhibits very

different degrees of development. One explanation of this may

be that such a character is naturally more developed in the

dominant homozygote, where the character was present in both

gametes, than in the heterozygote, where it was present only in

one, and this is probably the case. But there is one specimen,

No. 5, in which the feathering is the merest trace, and Mendelism

affords no explanation of such variations within a character,

it merely takes account of its presence or absence. Similar

remarks may be made concerning the crest: of the 8 chicks of

the first brood there are 6 with crest and 2 without, the Mende-

lian proportion, and there are 4 with well-developed crest and

2 with a verv slight crest. The latter may be the hetero-

zygotes for this character, although there should be four of these

to two dominant homozygotes.

I have noticed an interesting correlation between the crest

and the character of the comb. As Bateson points out, the rose

comb of the Silky has its posterior end divided into three

irregular points. He describes the comb as a rose plus a trifid

element which is the cause of this condition of the posterior end,

and states that in F, from Silky x Single, regular rose combs are

produced in those individuals which have the rose factor without

this trifid element. In other words, the comb of the Silky is re-

presented not by one factor but by two. The regular or ordinary

rose comb, as it occurs in Hamburg fowls, has a flat upper

surface which is triangular in shape, the apex of the triangle,

called the peak or pike, being free from the head and _ projecting

backwards. Bateson’s statements are for the most part confirmed

by my results, but I have a somewhat different interpretation to

offer. The rose comb of the Silky is not only trifid posteriorly

but abbreviated. This character oecurs in both the F,’s in my

experiment; it is dominant. In the F,’s the same character

246 MR. J. T. CUNNINGHAM ON

occurs in all those that have a crest, both males and females,

although the comb is much smaller in the hens. The two

specimens without a crest, on the other hand, No. 1 ¢ and

No. 7 2, have the normal rose comb with a projecting point

hee The crest makes no difference to the shape of the single

comb. There is thus a correlation between the crest of feathers

and the truncated trifid character of the rose comb: these two

characters either occur together or are absent together. I believe,

however, that this is not a coupling of two distinct. characters

but that they are both manifestations of one character. What

Bateson calls a trifid element is in my opinion nothing but the

crest; the factor which causes the development of the crest of

the feathers also causes the posterior truncation of the comb.

The most remarkable and interesting result in my experiment

is the condition of the recessive whites in the first brood of F,’s.

Both of these whites show some colour, while according to the

Mendelian theory of segregation they ought to show none. The

specimen No. 1, now (Sept. 3 30, 1911) 44 months old and in

mature plumage, has a band of colour Agios the back behind the

shoulder; when I first noticed this colour it was a very pale

yellowish tinge, so slight that it was difficult to be sure that it

was actually pigment in the feathers and not accidental staiming.

Now it is darker and more orange, and there is also a slight

yellow tinge in the neck hackles. The other white, No. 2, has also

some shght colouring, a faintly tinted region on the back like

the other, and some very pale patches on the neck ventrally

and over the eye on each side. These white specimens were

yellowish white in the down when first hatched, but this is no

evidence of colour, for the down of the chicks of white fowls, as

of white ducks, is always yellowish. The two whites of the

second brood do not yet show any colour.

Now this fact shows that the Mendelian doctrine of complete

segregation and gametic purity is not always true. The fact

that these recessive whites do appear in the F, generation shows

clearly that gametic segregation does take place in the F,

individuals, but in my experiment the recessive white is not

exactly the same as the original white of the Silky: it is no

longer pure, but is mixed with a little of the colour factor with

which it has been mixedin F,, We must consider whether the

result obtained by me is explained in any degree by Bateson’s

conclusion concerning the white of the Silky. As alr eady men-

tioned, he concludes that this white is due not to the absence of

all colour factors, but to the absence of one out of two which

together produce colour, Now this partial colour factor cannot

explain my result, for it cannot by itself produce any colour, and

the essential point of the Mendelian doctrine is that the recessive

individual in F, is homozygous, 7. e. due to the union of two

gametes each of which carries the factor for the recessive

character. The recessive character in the case under considera-

tion is the white of the Silky, and the white in the individuals

L MENDELIAN EXPERIMENTS WITH FOWLS. 247

of F, which I have described is not the same as the white of the

original Silky. Bateson states that the adult Silky is pure white

as a rule, but the down of the chickens has some colour usually

though perhaps not always. The colour in this case is buff, not

blue or black, and it occurs on the sides of the head and on the

rump. This again has nothing to do with the result I have

described, for the colour in my F,, whites was not observed in the

chicks, or in the mature plumage when first developed, but

appeared later, and gradually increased in intensity. I have

not examined pure Silky chicks, as I have not reared any.

It is, I think, certain that in this particular case the recessive

character derived from a cross is not pure, and this naturally

suggests that no recessives derived from crosses are perfectly pure.

The determinants of these characters in the hybrid individuals

have been mixed in the F, individuals with the determinants of

other characters. Whatever the determinants may be, whether

special chemical compounds such as ferments, or parts of chromo-

somes, it is possible that the mixing in F, is such that each

determinant is affected, tainted as it were by the other. If this

were the case it would tend to throw doubt on the whole

conception of species and varieties reared on the Mendelian

phenomena. It may be that segregation occurs not because the

determinants or factors in the gametes are indivisible units, but

because the determinants of certain pairs of characters are too

different to mix completely, while others less different would

show more mixture. In any case the recessive white in my

experiment is not pure, and it is improbable that it is the only

case of its kind. I hope next season to breed from these whites

and study the heredity of the colour character in the F,

generation.

Prof. Punnett, in conversation with me on this subject, made

two different suggestions in explanation of the colour in the

white recessive. The first was that the Silky is not a perfectly

white breed, but normally carries a little yellow colour, which is

usually present in the cocks but not in the hens. As the stock

of Silkies from which my cross came were all hens, and I have

no experience of Silky cocks, I cannot disprove this suggestion,

but can only point out that it does not agree with the account of

Silkies given by Davenport, nor with that given in Wright’s

‘Book of Poultry,’ which says the plumage should be pure white.

The second suggestion was that the yellow or yellow to red colour

of the Bankiva might be a distinct factor from that of the black

pigment, and therefore be segregated independently. This seems

to me an untenable hypothesis, as, if it were true, we ought to get

wholly black and wholly yellow or red birds in F,, and no such

cases have occurred. Moreover, the F, bird No. | in my experi-

ment has produced one or two single feathers on the back which

contain black pigment as well as yellow. I have plucked out one

such feather and preserved it,

QA8 MR. J. T. CUNNINGHAM ON

EXxPEriImMENnt 2.—Japanese Long-tail § x Dorking @.

The second experiment I have to describe was chronologically

the earlier, but is less complete and less important than that

already described. It consisted in crossing a Japanese Long-

tailed cock with a Dorking hen. Crosses between the Long-

tailed fowls and other breeds have been previously made by

C. B. Davenport at the Carnegie Station for Experimental

Evolution at Cold Spring Harbour, New York, U.S.A. (‘ Inher-

itance in Poultry, Washington, 1906). In his first experiment

Davenport crossed a cock of the Long-tailed breed, which he

refers to by its proper Japanese name of Tosa fowl, with a White

Cochin Bantam hen. Six F,’s were reared, 3 cocks and 3 hens.

The males were all of the male Tosa-fowl coloration except that

every feather was repeatedly barred with white. The females

were all of the female Tosa-fow] coloration except that the light

shafting was very much broadened. The female Tosa in this case

is described as ‘black mossed with rusty with a straw-coloured

shaft to each feather.” All three males developed abnormally

long middle tail-feathers, in other words the tail character of

the Tosa was dominant.

Of the F, generation 57 individuals were hatched : of these 16

were white and 41 pigmented, which approximates to the theo-

retical proportion of one in four. But of the 16 whites only 5 were

without trace of reddish pigment, which occurred on the breast,

top of head, and remiges. Davenport draws from this the same

conclusion that I have suggested in the case of my cross between

Silky and Jungle fowl, namely that segregation is not always

perfect, and the gametes from which these whites arose were not

pure for the white character. As the F,’s were immature at the ~

time of publication, the inheritance of the long-tailed character

in this generation could not be described.

With regard to the F,’s, Davenport points out that the two

Sexes, except for the admixture of the white colour, resemble the

male and female Tosa fowl respectively, and remarks that from

a germ-cell of the male Tosa either a bird coloured like a male

Tosa or one coloured like a female Tosa may arise, that the male

germ-cells contain the Anlagen not only of the male characteristic

but also of the female characteristic. The same result is equally

evident in the results of my cross between the male Jungle fowl

and the White Silky, the hens in F, and the coloured ones in

F,, showing the coloration and markings of the female Jungle

fowl. This conclusion is in opposition to the Mendelian view

that the female is heterozygous but the male homozygous, the

latter not carrying the female character. Mendelians argue

that the effects of castration in male vertebrates generally can

be explained as merely the non-appearance of male characters,

and they would probably maintain that the coloration of the

hens referred to in these cases was merely the absence of the

male characters. But it is necessary to distinguish between

MENDELIAN EXPERIMENTS WITH FOWLS. 249

somatic or secondary characters and the actual sex. It is certain

that the somatic characters of the female of a particular breed

are actually transmitted by the male, and there are no facts

known which support the assumption that positive characters,

if the female possessed them, would not be transmitted in the

same way,

In a second experiment Davenport crossed the Tosa male

with a female dark Brahma. In this case only the F, generation

was reared, namely 16 males and 5 females. In some of the

male hybrids the tail-feathers showed greater length than those

of the adult Brahma, but as they were only six months old at

the time the Report was written, the inheritance of the tail

character was not completely known.

In my cross between the Tosa cock and Dorkinz hen only three

F, chicks were hatched and reared in 1905. They were all hens.

Two of them had the double hallux which occurs in the Dorking,

on both sides, the third on one side only. The combs were larger

than those of the Japanese hens; the coloration was speckled

much like that of the Japanese hens, each feather with a lght

shaft, but the coloration of the Dorking hen is not very different,

and I did not pay much attention to this character.

In 1906 the three F, hens were kept with a pure-bred

Japanese cock. The eggs of the hybrids were brown in shell-

colour, while those of the old Dorking hen were white. ‘The

egg-colour of the Japanese was therefore dominant. This is

another case for the Mendelians to consider. Surely the colour

of the egg-shell must be a female character, and here it is trans-

ferred to the female hybrids by the male. The eggs were also

smaller than those of the Dorking, another egg character derived

from the Japanese cock.

Two broods of F,’s, i.e. F, 2 x Jap. d, were reared. In the

first brood 5 chicks were hatched and 3 died i in the egg: two of

the 5 had the double hallux, three had normal toes, all those in the

egg had normal toes. Thus the number with double hallux was

2 out of 8, whereas the expectation, as the character is dominant,

was equal numbers of each character. Ultimately three of these

chicks were reared, and all were cocks, one with double hallux,

two with normal toes.

The second brood hatched consisted of 11 chicks, of which

3 had the double hallux and 8 had normal toes. The two broods

together therefore give 5 with double hallux to 14 with normal

toes, where the expectation was equal numbers. Of the second

brood 5 were cocks and 6 hens. Thus of the F,’s there were in

all 8 cocks, and if the long tail was dominant every one would

show it, while if it were recessive half would have the long tail

and half the short tail of the Dorking. The result showed

that the character was dominant, as it was recognised in all the

birds except one which died before the character was definitely

visible.

One cock and five hens of these F,’s were kept to breed from,

250 MR. J. T. CUNNINGHAM ON

all of them with normal toes, all those with the extra toe having

been killed off. From these were hatched and reared in 1907

5 F,’s (2 cocks and 3 hens), and in 1908 8 F,’s (3 cocks and 5 hens).

In all these cocks the J apanese character of the tail was observ ed,

although they were not all kept long enough to develop any

great length of feathers. The result, lowers? is not conclusive

with regard to segregation or absences of segregation in the tail

dherewier, for the E, taal bred from F, 2 and Japanese ¢ might

itself hare been a pure dominant in this character, 1 cane

case all the F,’s would have been either Momezse one or hetero-

zygous dominants. Similarly, the F, ¢ kept for breeding might

have been a homozygous dominant, and segregation would then

not be evident in F,. A complete experiment ‘to test the segre-

gation of the J apanese Long-tail character is therefore yet to be

male,

The double hallux did not reappear in any of the F, or F,

generations, which supports the view that the normal toe in the

ie S 1S a pure recessive.

Il. Incomplete Segregation of the Pigmentation of the

Silky Foul.

In Part I. of this paper, when describing the characters of the F’,

generation froma cross between Bankiva male and Silky female, 1

treated the inheritance of the peculiar pigmentation of the Silky

as a case of simple segregation. When I wrote that part, I had

only examined the F, birds in the living condition, in which there 1s

a conspicuous contrast between the sooty colour of the combs and

wattles in the majority and the blood-red colour of the others

which seemed obviously recessives. Afterwards it occurred to me

that if there were incomplete segregation in the colour of the

plumage the recessives in other characters also might be impure.

I was led to consider the possibility of slight degrees of body

pigmentation by the description of ene degvees given by

Bateson and Punnett in their recent paper in the ‘Journal

of Genetics,’ vol. i. no. 3. Accordingly 1 killed No. 10 of the

F,’s, wie had the white colour and silky character in its

plunage, and apparently normal colour in comb and skin, that

is to say the comb etc. was blood-red, and the skin, so far as

could be seen between the feathers, unpigmented. Before the

bird was plucked, however, I noticed a ring of dark colour on the

lips of the cloacal aperture. On further examination I found

considerable pigmentation of the subcutaneous tissue of the

abdomen behind the sternum, which was the cause of the

darkening, above mentioned, of the lips of the cloaca. There was

subcutaneous pigment on the tarsal joint and all down the tarso-

metatarsus, but none on the knee-joint. It occurred also on the

elbow-joint of the wing and on the wrist-joint, but not at the

shoulder-joint. In all these cases the pigment was visible

externally as a darkening of the skin, and under the microscope

MENDELIAN EXPERIMENTS WITH FOWLS. 251

as a reticulum of pigment cells in the connective tissue. On the

peritoneum the pigmentation was slight but distinct, not only on

the outer wall but also over the intestines, for example over the

gizzard ; the surface of the testes was also black over the

anterior three fourths of the organs, while the posterior fourth

was white.

The next thing to be done was to make a thorough examination

of a normal specimen of Gallus bankiva in order to compare it

with the recessive just described. Fortunately, the cock of the

original cross being now rather old and blind of one eye, was

available for the purpose and was placed at my disposal and killed

and plucked. I found a little pigment in the subcutaneous tissue

about the ankle-joint posteriorly, but none in the periosteum :

this pigment was merely a slight extension of the pigment on

the inner surface of the skin, which extended down the tarso-

metatarsus as in the F, recessive. In both cases the shanks

appeared externally olive-green. In the Bankiva there was no

pigmentation anywhere else, either in the subcutaneous tissue of

the abdomen or of the wing-joints, nor in the peritoneum, and

the surface of the testes was white.

It is certain, then, that the pigmentation present in the F,

recessive 1s not derived from the Bankiva, but must have come

from the Silky, with the exception of that of the shanks, which

occurs in both breeds. The next question is whether the recessives

are all pigmented to a similar degree or whether some are desti-

tute of pigmentation. Of such recessives, as seen in the list given

in Part I. of this paper, there are three. One of these, No. 2

of the first brood, is still alive, beg kept for breeding ; so far as

can be seen it is in the same condition as the one which was

killed, as it shows the dark colour round the cloaca. The third,

No. 4 of the first brood, also a male, was killed before I thought of

looking for slight degrees of pigmentation, and therefore I can say

nothing about it. In any case, we have the fact that one of the

recessives is not pure, and therefore segregation is not complete

and perfect in the gametes, as the Mendelian theory assumes.

It may perhaps be suggested that slightly pigmented F,’s are not

recessives at all but heterozygotes, but this in the ordinary

sense is impossible, because such birds do not occur in F,, in which

all are heterozygotes. |The only possible conclusion is that there

is a tendency to segregation, but that the recessive character

when it separates is no longer pure but is, at any rate in some

individuals, modified bya slight degree of the opposite character ;

or if we adopt the presence and absence theory, we may say that

the positive character is not entirely absent from the recessive

individuals, but that distinct traces of it are present in them.

It is necessary now to make a detailed comparison between my

experiment and those described by Bateson and Punnett in the

paper to which reference has already been made. It will be

seen that the inheritance of the pigmentation of the Silky in

different crosses is exceedingly curious and complicated. Bateson

ayy MR. J. T. CUNNINGHAM ON

and Punnett crossed the Silky with specimens of a certain strain

of Brown Leghorns in their possession. When the Silky hen was

mated with the Brown Leghorn cock, the F, chicks, instead of

bemg strongly pigmented as in the experiment I have described,

were scarcely pigmented at all. It appears at first sight as if in

the Silky x Bankiva cross the pigmentation is dominant, while

in the Silky-Leghorn the same character is recessive. Many of

the F, chicks were reared, and in the adult state were almost

indistinguishable in general appearance from pure unpigmented

birds. ‘Thus these F, birds were in appearance sunilar to the F,

recessives described ‘by me, though in constitution they were

very different. I shall have to return to this. point later.

Careful examination of the F,’s in Bateson and Punnett’s

experiment showed traces of pigmentation. ‘These are described

as sometimes patches on the wattles, skin, or shank ; pigment

was also often found in the periosteum of the femoro-tibial or

tarso-metatarsal joints and frequently in the peritoneum. No

mention is, however, made of pigmentation under the skin of

the abdomen or on the wing-joints as in my F, recessives.

On the other hand, when the reciprocal cross was made, namely

Silky cock with Brown Leghorn hen, the results were markedly

different: the males were like those of the first cross, but the

females were all deeply pigmented. It might be said that in this

case the pigmentation was dominant in the females, recessive in

the males, while when the Brown Leghorn was the male parent

the pigmentation was recessive in both sexes. It is certainly

remarkable to meet with such a sexual difference as this in a

character which showed no signs before the crosses were made of

being in any way sexual. No ysuch sexual difference in the Fs has

been found to occur in the cross with which I have exper imented,

or in crosses with the Silky described by other naturalists. Tn

all other crosses known the F,’s were deeply pigmented; and

Bateson and Punnett point out that the almost complete absence

of pigmentation in one or both sexes of the F,’s in their ease is

related to the fact that the Brown Leghorns used by them had

unpigmented shanks, while most breeds normally have pigment

in the skin of the shanks, as in the Gallus bankiva described

above.

Bateson and Punnett explain their results as due to the

segregation of three pairs of hypothetical factors, namely, P, p,

presence and absence of pigmentation, I, i, presence and absence

of a factor which inhibits the development of the pigmentation,

and I, f, presence and absence of femaleness. It may be

noted here as an instance of the difficulty of keeping pace with

Mendelian theories, that Bateson seems to have already

abandoned his former theory of sex according to which maleness

and femaleness were alternative to each other, for he now

suggests that there may be another pair of characters, presence

and absence of maleness. The inhibition factor I is supposed to

MENDELIAN EXPERIMENTS WITH FOWLS. 253

be homozygous in the male Brown Leghorn, heterozygous in the

female, and the pigmentation factor to be homozygous in both

sexesin the Silky. With regard to sex a further assumption is

made, namely, that when an individual is heterozygous for both F

and I, these two in segregation cannot pass into the same gamete,

as there is a repulsion between them. The female, moreover, is

considered to be always heterozygous for femaleness, while the

male is homozygous for the absence of this factor. All these

factors being assumed, whether any real meaning can be attached

to them or not, their segregation according to the Mendelian

theory is shown to produce results which agree approximately

with those observed. But the approximation is not very close,

there are many serious discrepancies between the expected and

the actual results.

The next point to be considered is the nature of the inhibition

factor in the Brown Leghorn. We have seen that the difference

between this breed and others, such as Gallus bankiva, with

regard to pigmentation, is that in the latter the skin of the

shanks is pigmented, while in the Leghorn they are unpigmented.

The difference, then, is in the degree of pigmentation. Why

should the less degree of pigmentation be called an inhibition

factor? It may be said it does not much matter what term is

used so long as we agree as to the facts. But there is more

here than a mere difference of terms. Bateson and Punnett

assume that the inhibition factor forms a pair of factors segre-

gating independently of the pigmentation factor and its absence.

1 see no justification for this. It seems more probabie that the

unpigmented character is alternative or allelomorphic to the

pigmented, and in F, is dominant in the case of the Brown

Leghorn because it is of a higher degree than in breeds with

pigmented shanks. It is interesting to note that in Reptiles,

which represent the ancestors of Birds, pigmentation in the skin

and interna! connective tissue is of general occurrence. It would

seem, then, that in birds generally the pigment was transferred to

the feathers and disappeared from the skin and interior of the

body, but remained in the skin of the shanks, which is still,

excepting those few species and breeds which have feathered legs,

in the reptilian condition covered with epidermiec scales. In the

Brown Leghorn fowl this last remnant of the internal pigmen-

tation has disappeared. But there is no evidence of a distinct

factor inhibiting the development of pigment. There is no more

reason for assuming the existence of such a factor in a bird with

unpigmented shanks than in one in which the shanks, but not the

rest of the body, are pigmented. In the latter case Bateson and

Punnett admit that the unpigmented condition is recessive to the

pigmentation of the Silky, and it is obviously more reasonable

and more scientific to regard the condition of the Brown Leg-

horn as merely a higher degree of the unpigmented character.

Such a higher degree would naturally imply that there is a

954 MR. J. T. CUNNINGHAM ON

stronger hereditary tendency in the Brown Leghorn to the

absence of pigmentation, and therefore it is not surprising that

this character should be dominant in the cross with the Silky. It

seems to me, then, more probable that the unpigmented character

in Bateson and Punnett’s experiments is directly alternative or

allelomorphic to the pigmented ; and I shall endeavour to show

that their results can be as well, if not better, explained on this

hypothesis as on their own. It will be evident from this paper

that I am far from denying the occurrence of segregation, which

is an obvious fact, although my own results show that it is not

necessarily absolute or complete. It is further evident that a

sexual difference could not appear in the F,’s if there were not

already some sexual difference in one or both of the two breeds

crossed. The pigmentation tends to appear in the female sex

more than in the male: this not only occurs in the F,’s of the

cross Silky ¢ x Brown Leghorn 9, but a similar tendency is

seen, according to Bateson and Punnett, in the shanks of other

breeds. It is not, then, an unreasonable hypothesis that the

female Brown Leghorn has a greater tendency to the develop-

ment of pigmentation, or that the hereditary tendency towards

the unpigmented character is weaker in this sex and therefore

not dominant. We may also assume that when fertilisation

takes place, whether we regard femaleness as segregated from

maleness or each of them as alternatively dominant, the tendency

to pigmentation accompanies the female character. To express

these assumptions in Mendelian fashion, we may write W for the

unpigmented character which is dominant, and Wp for the

weaker unpigmented character, retaining P for the pigmentation

of the Silky. Then, instead of the complicated formule of Bateson

and Punnett we should have the two reciprocal crosses expressed

as follows :—

CAG) Brown Leghorn ¢ x Silky 9.

WW GC IPP ©,

Gametes W + W Pg+PQ.

Tees Pies WP ¢ + WP @.

(B.) Silky ¢ x Brown Leghorn 9.

IPE 6S Wipe 2s

Gametes P+ P WS + Wp.

Lan aay WP ¢ +. Wp Po.

The above formule are not to be taken as implying that sex is

determined entirely by the female, or that either sex does not

contain the other ina latent condition, but merely that, as half

the ova in fertilisation must actually give rise to females, the

tendency to pigmentation passes only into these females.

We have next to see how this works out for the F.’s.

According to Bateson and Punnett’s formule, the F,’s from cross

A are for the males 6 slightly pigmented to 2 unpigmented,

MENDELIAN EXPERIMENTS WITH FOWLS. A535)

for the females 3 slightly pigmented, 3 fully pigmented, and

2 unpigmented. According to my hypothesis the results would

be as follows :—

Ee seungacte. eat 65 Xv, Welt Sa

Gametes W-+P Wi P.

Bie ie. WW + 2WP + PP.

That is 1 unpigmented, 2 slightly pigmented, and 1 fully

pigmented in both males and females.

Bateson and Punnett point out that according to their formule

no fully pigmented males would be expected from this mating,

while as a matter of fact a considerable proportion of fully

pigmented males actually occurred, namely 12 out of a total

number of 75. They regard these as of the constitution ffP PII

or ffP Pi, and suppose that i in the presence of a double dose of the

pigmentation factor the effects of the inhibitor are in considerable

measure overcome, in corroboration of which view they found

that these specimens became less fully pigmented when adult.

But here they have made a mistake in reading their own formule,

for the combination PP occurs only once among the males

(namely ffPPIi, fig. 4, p. 190 of the paper), and therefore the

expectation on their own interpretation would be 1 in 8, not 1 in

4 as they assume. The actual numbers are :—

Males. Females.

Full. Some. None. Full, Some. None.

Me 42, Dill 28 28 13

Correct l

expectation 94 46:8 18°75 26 26 117/

ace. to B. & P. j

Expectation

acc. to my }18°75 37:5 18:75 17°25 34:5 17°25

hypothesis.

It will be seen that the numbers obtained by experiment. of

fully pigmented males are too large for Bateson and Punnett’s

hypothesis and too small for mine, while it must be admitted that

the numbers of females agree more closely with their assumptions

than with mine. If we add together the numbers for the two

sexes we have : —

Full. Some. None.

40, 70 34,

which is a close approximation to the expectation on my hypothesis,

36 72 36.

The differences, therefore, between the actual results and the

256 MR. J. T. CUNNINGHAM ON

expectation on my hypothesis are an excess of fully pigmented

females and of partially pigmented and unpigmented males, with

a deficiency of fully pigmented males. These differences would

be explained if segregation were, as I have shown it to be in my

experiment, incomplete, andat the same time pigmentation tended

to develop more in the females. Thus in the females the WW’s

would be partially contaminated by P, and some of them would

be partially pigmented instead of unpigmented, while some of the

WP’s would appear as fully pigmented instead of partially so.

In the males, on the other hand, the PP’s would be affected by W,

and this would reduce the number of fully pigmented in that sex

and increase the number of partially pigmented.

We have now to consider the F, generation from the mating

B in which the Silky is the male and the Brown Leghorn the

female. According to the formula of Bateson and Punnett the

expectation is 3 fully pigmented, 3 partially pigmented, and

2 unpigmented in each sex. This expectation is fairly well

fulfilled in the males, but in the females there is a great excess of

fully pigmented over the partially pigmented: the former are

nearly half as many again as the latter, and this the authors call

a slight excess. ‘The complete numbers are :—

Males. Females.

Full, Some. None. Full. Some. None.

56 5) 34 74 55 38

Expectation 652°5 52°5 35 62°5 62°5 42,

According to my hypothesis the F,’s are

AWE 6h Wp Pe,

and the gametes

W+P, Wp+P.

The zygotes therefore will be

WoW, W Pos Work, RP:

The first and last of these are the same as those which occurred

in the original parents, while the second and third are the zygotes

of the male and female F,’s. This is the ordinary result of

Mendelian segregation in the F,: one pure dominant, one pure

recessive, and two heterozygotes. The difficulty is to decide what

will be the distribution of the sexes among these zygotes. If we

suppose that Wp is always coupled with the female character,

then ail the W Wp’s and Wp) P’s will be females, and all the rest

males. But this would make not only the W P’s but also all the

P P’s male, and it seems improbable that the pure Silky character

in the F, should be confined to one sex. We may conclude that

the P P’s will be of both sexes and likewise the W Wp’s. We

may assume that the heterozygotes W P and Wp P will be in the

MENDELIAN EXPERIMENTS WITH FOWLS. 251

same condition as in the F,, the former all males, the latter all

females. Thus we shall have for the males

W Wp, mW Py 12

and for the females

W Wp, ZeWy pil. PE.

In the males, therefore, the ratio should be one unpigmented, two

partially pigmented, and one fully pigmented. It must be ad-

mitted that this does not agree w ith the actual results obtained

by Bateson and Punnett; but if segregation is not always complete,

some of the W P’s might appear in Fas fully pigmented and

would then be added to the P Ps, and the number of partially

pigmented and fully pigmented might be equal.

In the females on my hypothesis there should be one unpig-

mented to three fully pigmented, and none partially pigmented.

This again does not agree with the actual figures; but if segre-

gation is incomplete, ‘the P in the Wp P may be affected by

the W , and so a certain number of females might appear as

partially pigmented. In any case there is actually a great excess

of fully pigmented females over the partially pigmented, whereas

according to Bateson and Punnett the numbers should be equal.

There is another point in Bateson and Punnett’s figures of the

F,’s from this cross which may be of great importance, although

it is not noticed by the authors themselves. According to their

hypothesis the total numbers of the sexes should be equal, whereas

they are actually 140 males to 167 females. ‘Thus there 1s not only

an excess of fully pigmented females over the partially pigmented,

but a large excess, more than 19 per cent., of females over males.

This excess occurs in almost every family, and it seems improbable

that it should be accidental. In the F,’s from the reciprocal cross

Leghorn ¢ x Silky 2, on the other hand, there is an excess of

males, 75 to 69 females. It would appear that the Leghorn was not

only dominant as to character, but prepotent asto sex, I cannot

attempt to explain this, but it is not in accordance with Mendelian

theory.

In any case we are justified in saying that the actual results

obtained by Bateson and Punnett in the F, generation do not

agree in either of the reciprocal crosses with their assumptions,

and that there is nothing in their results to prove that imperfect

segregation may not occur in the cross of Silky with Brown

Leghorn. It would searcely be profitable to compare in detail

the results of all the other matings recorded by Bateson and

Punnett with those which would be “expected on my hypothesis ;

the comparisons would be similar to those already given as

examples. It is sufficient to point out that all their explanations

are founded on the assumption that the pigmentation of the Silky

is inherited as an indivisible unit, and that the result of my

experiment shows that this assumption is contrary to fact.

Proc. Zoou. Soc.—1912, No. X VII. Le

258 MR. J. T. CUNNINGHAM ON

In view of the subdivision of the pigmentation character proved

by my results, the remarksof Bateson and Punnett on p. 201 of their

paper are somewhat curious. They state that the F, birds, either

of both sexes or of the male sex only, according to the mating,

in respect of the intensity of their pigmentation might belong to

either of their three classes, slight, some, or moderate, and that

their experience has been that these classes grade very much into

one another. Further on they remark that it would be possible

to choose birds of the F, generation from Silky 2 x Brown Leg-

horn ¢,and to arrange them in a series exhibiting continuous

gradation from full pigmentation to none at all. ‘“ Yet we now

know,” they assert, ‘‘that such a series is due to the interaction

of three definite factors, inclusive of the sex factor, and that the

continuity of variation manifested is in reality founded upon a

discontinuous basis.” It seems to me that this is a misuse of

words: we know nothing of the kind. What we know is the

continuous gradation; and even apart from the results of my ex-

periment, the discontinuous basis to which the authors refer is

purely hypothetical. Moreover, one cannot help asking, in the

light of the above quoted remarks, what value is to be attached

to the distinction of the three degrees of pigmentation, full, some,

and none, on the numbers placed under which degrees the validity

of the factors assumed by the authors entirely depends.

In view of the failure of complete segregation in the two

characters dealt with in this and the preceding paper, 1t 1s natural

to consider whether any of the other characters present in the two

forms crossed also fail to segregate. ‘This point is more difficult

to decide in the other characters. There could be no doubt that,

the white plumage and the unpigmented skin were recessive with

a trace of the dominant character, but in other characters any

condition short of complete absence might possibly belong to the

heterozygote.

The double hallux. This character obviously does not behave in

the normal Mendelian fashion. In F, the double hallux is

dominant but not completely so: in my F, cock both feet were

normal, in the hen both feet had the double hallux. In F, there

are nine normals to one with double hallux, but no intermediates.

Feathering of legs. There appear in the F,’s to be two homozygous

dominants, six heterozygous, and two recessives. If this is the

correct interpretation, one of the recessives contained a trace of the

dominant character, the degree of feathering being infinitesimal,

only a few very minute feathers between the large scales. I am

inclined to think that the only specimen recorded as having quite

clean legs had really a trace of the character, for although no actual

feathers were visible, the lower edges of the scales showed a slight

ridge.

Crest. There are two specimens, Nos. 1 and 7, one cock and one

hen, with no trace of crest and the rose comb pointed behind.

These seem to be pure recessives showing complete segregation.

MENDELIAN EXPERIMENTS WITH FOWLS. 259

But there are also two, Nos. 2 and 5, in which the crest is very

slight, less than in the F,. No decided conclusions can be drawn

in this case,

Comb. I have not yet observed any intermediates between rose

comb and single.

Silky Plumage. Ihave not yet observed incomplete segregation

in this character.

Many eases in which the results of crossing experiments are not

completely in accordance with the Mendelian “theory are mentioned

by Bateson in his ‘ Mendel’s Principles of Heredity’ (1909), among

them being the double hallux, the leg-feathering, and the crest

of fowls.

In the light of the facts which I have recorded, the doctrine

known as Mendelism cannot be said to be based on sufficient

evidence to justify the notorious dogmatism of its supporters.

Mendelians assume that heredity is the transmission of unit

characters, which may sometimes combine to form a new character

which is transmitted as a whole, but that the units cannot be

subdivided. If subdivision occurs it is assumed to be due to the

fact that a supposed unit was really compound ; if, however, the

process of segregation is at least in some cases not complete, but

each character after a cross is mingled with some fraction of the

opposite character, the assumption of units and all that is founded

upon it fallsto the ground. It is possible that segregation is only

incomplete between characters which differ from one another in

degree and not in kind, for example between pigmentation and

absence of pigmentation in fowls. But if the doctrine now gene-

rally adopted by Mendelians of presence and absence is valid, it is

difficult to see how this difference could exist. Hitherto, Mende-

lian experiments have been too exclusively carried out on the

assumption of indivisible characters, whether real or imaginary,

and the object kept in view has been to find, or invent, factors

whose complete segregation would agree with the observed results.

There is a promising ‘field for research in the inv estigation of the

divisibility of real characters.

References.

TrecerMeieR, W. B.—1867. The Poultry Book. London: Rout-

ledge.

Darwin, C.—1876. Variation of Animals and Plants under

Domestication. 2nd Edition, Vols. I. & II.

Bateson, W.—1909. Mendel’s Principles of Heredity. Cam-

bridge University Press.

Davenport, C. B.—1906. Inheritance in Poultry. Publications

of the Carnegie Institution of Washington, No. 52. Papers

of Station for Experimental Evolution, No. 7.

Bateson, W., & Punnett, R. C.—1911. The Inheritance of the

peculiar Pigmentation of the Silky Fowl. ‘Journal of

Genetics,’ Vol. I. No. 3, p. 185.

260 : DR. H. LYSLTER JAMESON ON

18. Studies on Pearl-Oysters and Pearls.—I. The Structure

of the Shell and Pearls of the Ceylon Pearl-Oyster

(Margaritifera vulgaris Schumacher): with an Hxami-

nation of the Cestode Theory of Pearl-Production. By

H. Lysrer Jamuson, M.A., D.Se., Ph.D., F.Z.8.

[Received November 20, 1911: Read February 20, 1912. |

(Plates XXXIIT-XLVII. and Text-figures 33-41.)

CoNTEN'S.

Page

(@) @intinodietvom ieee i, dace Bins Qala ee aetna OU)

(2) Outline of the recent ler aciionitons 4 in 1 Case « Pee ene ditcataa’ ZAoe

(3) Professor Herdman’s Conclusions on Pearl- Meeearion Reena aocnte 00)

(4) Examination of the Cestode Theory of Pearl-Production ......... 271

(5) Description of the two Globular Cestode Larve from the Clan Beat

Oyster .. si 290

(6) Obicaionen on wate Str etuite! o the Hibrows) Cas suounane he

Cestode Larve: in Margaritifera vulgaris .......00.00.ccccc teers eee 298

(7) Materials available for the Present Investigations ........................, 294

(8) Methods ........ Maa ENCE Reece accasa oe | ZO)

(9) Structure of the Sialkspinienaas. ele ae Rais te eRe MMe OAAcboo. LU)

(lO) RhewShrelleseeretime WBpitheliar ste. ta-- sree sees eens races eee cr eee eee

(11) Shell-Secretion ........ Baraat Ole)

(12) Abnormal and Bat Hologteal Phases: aif the shar isuneeaeey Bee psasticssc 310

(13) Varieties of Ceylon Pearls.. Bee REA eis Miia aE meats CEE seeeneds.. Sle)

A. Muscle-Pearls ............. RRO EASRR oat oadiead , oral

B. Parenchyma-Pearls (« Cre Bpeamlee e “Blardhaen) Peeeneanicciodsa! So)

(14) Descriptions of the Centres of Individual Pearls .........................:. 3381

AN, Wierrortenl tiramn tvs 1eeesviemn NUE Seccsc cu ccoccussnocabapcenccataseossns = Dall

1B, Jetenyellss jompelnesexsl sour (Cas OM sencaasocasoucunesadatondendrecosodcconeesnes = GB

C. Professor Herdman’s Shides.. oe Peaceesoeansnes Ooo

D. Unlabelled Pearls (dry) in ‘ie Baer ikea cooatet ee eee

E. Collection of Pearls given to me by Mr. Max Mayer We peecesters Sols.

¥. Collection of Ceylon Pearls given to me by Mr. E.

op kms ieee eee 339

G. Pearls from Moraes afer. a apace is = thanin ie Gulf of Kana 341

H. Pearls from M. vulgaris trom the Mediterranean ............... 343

I. Pearls from M. vulgaris from New Caledonia .................. 343

J. Pearl from MW. vulgaris from Nossi-bé, Madagascar ......... 344,

K. Pearls from the Lapi shell (WZ. eae is) trom the eopeiena

Islands, Papua .......... 344

L. Pearls from Placuna pelle how iLaibe e Tampalakamam,

Ceylonteeeesertas ete Rar a! 345

M. Pearls from J. cee fee a var. cumingii Tears (the

Black-edged Mother-of-Pearl ees from ae Gam-

bier Archipelago shee ake 346

(5) Geneval Simamiany; pss) eater aoe ee Acer ch pel ee eee

(16) Works referred tO) 5o...6. 0. usterseieee nnnccembee-pasejane nee Son ep thnjansiens psn eS ed

(17) Explanation of the Plates...............ceecceesesesceecee cee teeeetenereeterscnseeeee GOR

(1) Ixrropucrion.

The studies which are embodied in this paper have been deferred

and interrupted. When Professor Herdman published his Report

,Lith® London.

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THE CEYLON PEARL-OYSTER, 261

on the Pearl-Oyster fisheries of the Gulf of Manaar (16) I was in

South Africa, and material for the examination of some of his

conclusions, which I was unable to harmonise with my own

observations made prior to my departure from England, was not

available,

Since my return J] have made use of all available material

which is somewhat scanty ; and while it is insutlicient to enable

me to propound, at the present moment, a working hypothesis as

to the true cause or causes of the formation of Ceylon pearls,

J think I have been able to show that the Cestode theory

enunciated by Herdman, which has formed the basis of the

somewhat meagre experiments which have been prosecuted in

Ceylon since his return to England, and which even seems to

have led to definite regulations slsowhere® , rests on quite

insufticient evidence, and that, if the problem of the cause of

Pearl-Production in Margaritifera vulgaris is to be solved, and a.

scheme for increasing the productivity of the Oysters evolved, a

fresh start will have to be made.

The work that still remains to be done will centre around the

causes which lead to the development of the epidermal sacs in which

all pearls are formed—fine pearls and seed-pearls, “‘ muscle ”-pearls

and ‘“eyst”-pearls (or, as 1 prefer to call them, “parenchyma”-

pearls), and the mechanism which controls the secretion of

conchyolin and the deposition therein of carbonate of lime. In

fact, Iam led back to the principles enunciated by me in 1902

(25), that the essential element in pearl-formation is the pearl-sac,

aud not the nucleus, and that it is by a study of the causes which

lead to the development of the former that the problem of the

origin of pearls is to be solved.

Some material for these further investigations has just reached

me as | write this introduction, and more is promised, but, owing

to the difficulties and delays which may occur, | now publish

my researches on the structure of the shell, and of pearls and their

pseudo-nuclei and nuclei, without attempting to deal with the

origin of the pearl-sac.

I take this opportunity of expressing my thanks to the Ceylon

‘ompany of Pearl Fishers, Ltd., for purchasing, on my behalf,

pearls in Colombo to be used in these investigations ; to Professor

Raphael Dubois, for pearls from Magaritifera vulgaris trom the

Mediterranean; to Prof. W. R. Dunstan, F.R.S., for allowing me

to make use of material in the Imperial Institute; to Mr. J.

Caleott Gaskin, Assistant Political Agent at Bahrein, Persian

Gulf, for sending me, in 1903, a number of preserved specimens

of the Lingah shell (JZ. vulgaris), some of which contained pears ;

* Rules for Lower Burma under the Burma Fisheries Act, 1905. Rules 64 and 67,

which prohibited the capture of Balistes and Trygon inthe Pearl Fishery districts,

and required them, if accidentally caught, to be returned to the sea, appear to have

been inspived by the Cestode theory. These rules were cancelled in 1909, It may

be remarked that the Pearl-Oyster of Burma (the Mergui shell of commerce) is not

the same species as the Ceylon Pearl-Oyster, I. vulgaris Schumacher, but is the

great Mother-of-Pearl Oyster, IZ. maxima Jameson.

262 DR. H. LYSTER JAMESON ON

- to Mr. E. Hopkins, of Hatton Garden, for specimens of Ceylon

Pearls; to Professor W. A. Herdman, F.R.S., for specimens of

Pearl-Oysters from Ceylon, and for allowing me to examine bis

preparations ; to Mr. Max Mayer, of Hatton Garden, for specimens

of pearls for my work; to H.H. the Jam Saheb of Nawanagar,

for specimens of Pearl-Oysters from the Gulf of Kutch ; to Prof.

L. G. Seurat, for pearls from New Caledonia, Madagascar, and the

Gambier Archipelago; to Mr. A. E. Shipley, F.R.S., for allowing

me to examine and make use of his specimens of 7'ylocephala and

allied genera of Cestodes from Ceylon fishes; to Mr. KE. A. Smith,

1.8.0., for permitting me to make use of material in the British

Museum (Natural History), including specimens from Dr. Kelaart’s

collection; and to Mr. A. Van Noorden (of the firm M. Myers,

Mother-of-Pearl Merchants) for specimens of Lingah and other

shells.

After this paper was handed in I received a copy of Rubbel’s

paper (34a) setting forth in greater detail the results already

published by him (33 and 34). Herr Rubbel and I have arrived,

working independently on widely different molluscs, at identical

conclusions on several important points, such as the nature of

Herdman’s ‘“calcospherules.” Where practicable, I have inter-

polated references to his work in the text, and my only regret is

that it is not possible to discuss his valuable work more fully m

the present paper and to dwell at length on the many points where,

adopting a slightly different terminology and interpretation of the

phenomena, our respective works lead to the same conclusions.

(2) OUTLINE OF THE RECENT INVESTIGATIONS IN CEYLON.

In January 1902 Professor W. A. Herdman went to Ceylon, at

the request of the Colonial Office (who availed themselves of his

services on the recommendations of the Council of the Royal

Society and of Professor Ray Lankester), to investigate the

condition of the Pearl-Banks. Professor Herdman took with him

as his assistant Mr. James Hornell, who remained in Ceylon to

carry on the work after the former’s return to England in April

of the same year, and who collaborated in the preparation of

Professor Herdman’s reports. Professor Herdman’s visit in 1902

gave him seventy-eight days in Ceylon, and was largely spent in

an extensive biological and faunistic survey of the pearl-banks,

carried out on two successive dredging-cruises, each of several

weeks’ duration ; and he credits Mr. Hornell with the major part

of the observations on Pearl-production (Royal Institution

Lecture of March 27th, 1903) (14). Prof. Herdman himself

always seems to have regarded the condition and welfare of the

natural beds of oysters as a more important problem than the

question of pearl-production (Report on the Ceylon Pearl

Fisheries, Part I. p. 5, and Part V. p. 29; also Report of the

Annual Meeting of the Ceylon Company of Pearl Fishers, Ltd,

THE CEYLON PEAKL-OYSTER. 263

for 1908*). In fact, he contends on p. 30 of Part V. of his Report,

and in his address to the Linnean Society on 24th May, 1906,

that

“to reverse the popular saying, if we attend to the prosperity

of the bed as a whole, the individual oysters may be left to

take care of themselves, both in regard to health and pearl-

production.”

In January 1904 Mr. Hornell was appointed Marine Biologist

to the Government, to which post were subsequently added ‘the

administrative duties of Inspector of Pearl-Banks. While holding

these Government appointments Mr. Hornell continued to colla-

borate with Prof. Herdman, though it is clear that the executive

and administrative duties attached to his post interfered not a

little with the more strictly scientific observations. Thus, in his

Report on the Inspection of the Ceylon Pearl-Banks, November

1905, Mr. Hornell says (23), p. 6 :—

“The working out of this material must of necessity await

the long deferred time when a pause shall occur in the

field work in which I have been engaged for the past

eighteen months, and which permits me no leisure for the

correlation and marshalling of biological data.”

And, again, in Reports from the Ceylon Marine Biological

Laboratory, } No. 1, p. 23, 1905, he says :—

“The Marine Biologist should be given opportunity to

further investigate the Mite of the spherical Cestode so abun-

dant in the Pearl-Oy ator and which is the inducing agency

in the formation of ‘cyst-pearls’ (‘fine pearls’ . The

problem is far from solution, and will entail much unpleasant

and trying labour before a satisfactory conclusion can be

hoped for.”

The observations of Prof. Herdman and Mr. Hornell on the

spot were corrected and correlated by laboratory work in Liverpool,

carried out by Prof. Herdman and his staff at the University, on

the material sent home for investigation. » Prof. Herdman has

courteously allowed me to examine the slides made during these

investigations, showing sections of Pearls im sitw in the tissues,

and of ‘the Gestode larvee which he asociates with pearl-formation.

In March 1906 the Ceylon Company of Pearl Fishers, Ltd., was

formed and the pearl-fisheries were leased to the Company by the

Government, at a yearly rental of Rs. 310,000, the lease carrying

the obligation to spend, in addition to the above rent, a sum of

from Rs. 50,000 to Rs. 150,000 yearly ‘on the experimental or

* Financial Times, Dec. 19,1908. Sir West Ridgeway, Chairman of the Company,

on ae occasion said that

“with regard to biological research, Prof. Herdman was of opinion that in

the present condition of the Company’s pearl-banks accurate navigation,

careful and exhaustive inspection of the ground, and wise administration are

more important than the purely scientific side of the business.”

264 DR. H. LYSTER JAMESON ON

practical culture of the pearl-oyster and on the improvement of

the pearl-banks ” (50).

On the formation of the Company Mr. Hornell was transferred

to its service as local General Manager, Prof. Herdman being

made Scientific Adviser.

In April 1908 Prof. Herdman, at the request of the Company,

paid another short visit to Ceylon, to enquire into the question of

the inspection of the banks and other branches of the business.

As a result of Prof. Herdman’s inquiries, the post of General

Manager was abolished, being merged in that of Managing

Director, and Mr. Hornell resigned, being succeeded by Mr. T.

Southwell, A.R.C.Se. (Lond.), who since 1907 had been acting as

Mr. Hornell’s assistant, and previously to that had assisted

Prof. Herdman in his laboratory at Liverpool in the preparation

of the material sent home for investigation. Mr. Southwell

was made Scientific Adviser, a post which he still holds. Pro-

fessor Herdman continued to be retained in an advisory capacity.

Capt. J. Kerkham was appointed Superintendent of Fisheries*.

Besides the work of the Company’s scientific employees, Dr. A.

Willey, in his capacity as Marine Biologist to the Government

(a post which he held along with the Directorship of the Ceylon

Museum), has published some observations in the Ceylon

Administrative reports and in ‘ Spolia Zeylanica.’

Particulars of the work done, and of the conclusions arrived at,

by these several naturalists will be given in the course of the paper.

In considering the incompleteness of the observations, despite

the eight and a half years that have been devoted to the study of

the Ceylon pearl-banks and the very large sums of money that

have been expended, it must, of course, be borne in mind that for

the last three or four years the banks are stated to have been

practically bare of oysters t, and the prosecution of the investi-

gations initiated by Prof. Herdman has thus been seriously

hampered. But it is amazing that a Company whose prospects

were so largely dependent on scientific work should have failed

to set by an adequate stock of properly preserved material for

scientific investigations and to establish at a suitable station a

reserve of live oysters when the oysters were passing through

their hands by the million. Had this been done, the barren

years that have now come might have been devoted to the

examination and amplification of Prof. Herdman’s observations,

* Since the above was written the operations of the Company have ceased. It was

announced in the ‘Times’ of April 4th, 1912, that the lease had been terminated,

a deposit of £10,000 together with the property of the Company being forfeited to

the Government. An examination of the causes of the failure of this short-lived

Company, which started with a capital of £165,000, has lately been published by the

present writer (26 a).

+ Not entirely ; for it was possible to obtain 12,000 oysters in Feb. 1910 for Mr.

Southwell’s feeding experiment described in Part V. of the Ceylon Marine Biological

Reports, p. 213, and no less than 35,000 oysters ranging from 8 months to 25 years

old were obtained for the experiment described in Part IV. of the same publication,

p- 169. Mr. Southwell, in a paper published in May 1911 (42), says: “The only bed

which now exists is confined to an inshore area, and the oysters found thereon only

rarely contain the pearl-inducing parasite.”

THE CEYLON PEARL-OYSTER,. 265

which of necessity were somewhat cursory and superficial. The

result of this lack of foresight has been that the energy that ought

to have been concentrated on an intensive study of the pearl=

oyster and the mechanism of pearl-formation appears to have

been largely dissipated on general faunistic work, such as the

description of new species of crabs and tapeworms, matters which,

valuable as they are from the purely scientific standpoint, have

only a secondary bearing on the problem of increasing and

rendering more reliable the supply of pearl-oysters and pearls.

(3) Proresson Herpman’s ConcLustons oN PEArL-Formarion.

Professor Herdman distinguishes several causes of pearl-

formation, though only two of these are regarded as of sufficient

frequency to have economic importance, viz. Cestodes, causing the

majority of “eyst-pearls,” and ‘“ calcospherules,” causing “muscle-

pearls.” I will pass over the pearly excrescences or “blisters ” on

the inside of the shell, due to the irritation of boring animals or

intruding particles of foreign matter, as these should be kept in

a category entirely distinct from true pearls. The latter term,

following my paper published in 1902 (25), I shall confine strictly

to bodies developed independently of the shell, which are not in

any way continuous with the shell, except where, owing to the

rupture or absorption of the intervening tissues, they may become

secondarily covered over with nacre continuous with the lining of

the shell. When this happens to a pearl it becomes an “attached

pearl,” a body quite other than a blister. Attached pearls are

valued for the true pearl that can often be dissected out of them,

whereas blisters are used as substitutes for pearls where the

imperfect side can be concealed in the setting, e. g. in cheap

jewellery, rings, pins, brooches, etc. Prof. Herdman (Report 1.

p- 10) apparently applies the name “ ampullar pearls ” to blisters,

that is to say to bodies “which are not formed within closed

epithelial sacs like the others, but lie in pockets or ampulle of the

epidermis,” and on p. 146 of the same part speaks of blisters as

‘pearls of an inferior quality,” but I cannot help feeling that, in

scientific terminology at least, it is undesirable to apply the term

pearl” to these bodies at all.

Professor Herdman recognises the following causes of pearl-

formation in the Ceylon pearl-oyster :—

(i.) Grains of Sand and other Foreign Particles.

These, in the experience of Professor Herdman and Mr. Hornell,

only form the nuclei of pearls under exceptional circumstances.

In the whole of their observations they have only records of three

such cases out of hundreds of pearls examined (Report V. pp. 4

& 127). They say (V. p. 28) :—

‘* Probably it is only when the shell is injured, e. g., by the

breaking of the ‘ears,’ thus enabling sand to get into the

interior, that such particles supply the irritation that gives

266 DR. H. LYSTER JAMESON ON

vise to pearl-formation. The ectoderm, in such cases,

would probably also be damaged, and cells may be carried in

with the inorganic particles.”

As shown below, the presence of grains of sand and other foreign

econ: ovens 8 ae

particles in the nuclei of some Ceylon pearls has been confirmed

by the present writer.

(i.) Boring Animals.

While recognising that pearly excrescences or “ blisters ” are

mainly due to borers such as Zewcodore and Clione, Herdman and

Hornell say (Report V. p. 28) that

“in exceptional cases a free pearl may be formed in this

way.”

No specific instances, however, are cited, nor is any explanation

suggested as to what would be the mechanism in such cases.

(iu.) Parasites other than Cestodes.

In his lecture at the Royal Institution, referred to above,

Prof. Herdman said :

“¢ We shall I think be able to show in our final report that

Cestodes, Trematodes and Nematodes are all concerned in

pearl-formation.”

At the same time he recognised the ‘“ larval Cestode of the

Tetrarhynchus form” as the most important cause. Again, in the

Report (V. p. 29), Herdman and Hornell say :

‘“‘ A fuller experience is causing us to incline to the view

that various parasites may act as pearl nuclei, even in the same

mollusc. Some pearls are certainly formed around intrusive

Nematodes. We have a complete cyst pearl, free and

unattached, of which the nucleus is a coiled Cheiracanthus

unctnatus, on which the pearl deposit is not sufficiently thick

and opaque to obscure the coils so as to render identification

difficult.”

This pearl does not appear to be in Prof. Herdman’s collection.

(iv.) Pearls without a Nucleus.

Prof. Herdman points out that both in the case of the Ceylon

Pearl-Oyster and J/ytilus some pearls have no trace of a nucleus

(Report V. p. 18). He figures one such pearl from Mytilus,

magnified 100 times. The existence of pearls without nuclei

was recorded by Harley (11) in 1889. The observations set

out below show that in the Ceylon Pearl-Oyster, both in muscle-

pearls and in a great number of parenchyma-pearls, the presence

of a nucleus of foreign origin is quite unnecessary, and point to

the conclusion that the origin of the pearl-sac is usually due to

stimulation other than that caused mechanically by such bodies.

So far as J/ytilus is concerned, I attribute the conditions where

THE CEYLON PEARL-OYSTER,. 267

a nucleus is absent to two alternative causes. In the first case, the

Trematode described in my paper on the Origin of Pearls in

the P. Z.8. for 1902 (25) may migrate out of the sac, in which a

pearl without any nucleus, or with a nucleus consisting of a few

residual granules, may subsequently be formed. In the second

case, in certain localities, Wytilus edulis produces pearls through

an agency (which I am at present trying to investigate) other than

Vermian. These pearls may have as nuclei either a few dark

granules or nothing that can be detected without the use of much

higher magnification than 100 diameters. One of them is shown

at Cin text- -figure 33 (p. 277).

(v.) Musele-Pearls.

The discovery of “‘muscle-pearls” was, I believe, first announced

at the British Association Meeting in September 1903.

Under this name (Brit. Assoc. Report, Southport, 1903, p. 695)

Prof. Herdman distinguishes pearls formed “around minute

calcareous concretions, the ‘ calcospherules, which are produced

in the tissues and form centres of irritation.” They occur ‘“ most

abundantly in the muscular tissue near the insertions of the

levator and pallial muscles.” (Report, Part V. p. 27.) Herdman

and Hornell say, speaking of muscle-pearls :

“it seems probable that these have been formed by the

deposition of calcareous matter around a minute calculus in

the tissues” .... ‘The Muscle pearls when present are

usually abundant, and when examining under the microscope

a young pearl of this kind, im situ, it is common to find a

large number of minute calcareous depositions or calco-

spherules scattered in the neighbouring tissue. It is probable

that the muscle pearls are formed around these microscopic

calcospherules as centres of irritation, and as these [? their}

positions are invariably in our experience close to the surface

of the muscle or the mantle, there is no difficulty in under-

standing that there, if anywhere, ectoderm cells might

migrate to the source of irritation and thus be responsible

for the deposition of a pearl.” (Report V. p. 27.)

Muscle-pearls are especially numerous in certain localities ;

Prof. Herdman instances (Report V. pp. 30-31) that they were

particularly numerous on the 8.E. Cheval Paar in 1902 and 1903,

and

“that the vigorous and healthy oysters of the Eastern Cheval

and Periya Paar Karai produce practically all the examples of

this class of pearls,”

the numbers produced on other banks being insignificant.

Mr. Hornell, speaking of an examination of a number of pearls

attached to the shell, states that ‘ decalcification of the pearls

attached to muscle-scars reveals no organic nuclei, whereas the

Jattached| pearls irregularly disposed have Cestode embryos as

nuclei, exactly as ‘ fine’ pearls have.” (19, p. 12.)

268 DR. H. LYSTER JAMESON ON

Mr. Southwell (40, p. 194), referring to the caleospherules

causing ‘* muscle-pearls,” says :

“The origin of the latter bodies is quite unknown,

although it seems almost certain that they are depositions

from the blood.”

Again, in a later paper (42, p. 128), Mr. Southwell says

“‘ Other pearls are also found in the Oyster, but they have

no organic nucleus. Such pearls are termed muscle or seed

pearls. ‘Their origin is obscure, but they are always found

near the muscle insertions, and are believed to be formed

round a calcospherule of excretory origin, or by the sheer of

muscles moving in different planes.”

In the first mentioned of the above papers (40), he goes on to

say

‘“‘ Considerably more pearls are formed round calcospherules

than round parasites, the ratio being about 13 to 1. They

are therefore of considerable commercial importance.”

Unfortunately, Mr. Southwell does not give the number or

sources of the pearls on which this statement is based, although

it is quite clear, from the observations of Prof. Herdman and

Mr. Hornell, that ‘‘ muscle-pearls ” are characteristic of certain

localities, and ‘‘cyst-pearls ” of others, and that the ratio may vary

greatly on different banks.

My own observations on “* Muscle-pearls” and on the nature of

the so-called “ calcospherules” are given in a later part of this

paper.

(vi.) Cestode Larve.

Of fine or ‘ Orient” pearls Prof. Herdman and Mr. Hornell

claimed that the most frequent nucleus is a Cestode larva. In

their ‘ Conclusions on Pearl-Formation” (Report V. p. 29) they

maintain that their investigations have shown “that in Margariti-

fera vulgaris, at Ceylon, the production of the ‘ Orient’ pearl is

dependent upon Cestode infection and that the species mainly

concerned is Tetrarhynchus unionifactor,” and in the General

Summary of their Ceylon Report (V. p. 127) they say:

“The majority of these fine pearls contain as their nuclei

the more or less easily recognisable remains of certain

Platyhelminthian parasites, which we identify as the larval

condition of Cestodes belonging to the genus Vetrarhynchus.”

It is stated several times that this supposed identification was

made during Professor Herdman’s second cruise in March 1902,

but its elaboration must have been, in great part, the work of

Mr. Hornell at a later date. The narrative of the Cruise

(Report I. p. 70), published in 1908, states, in a paragraph

apparently inserted between the records of Mar ch 6th and 7th:

‘“TIn the intervals of dredging and when moving from

place to place, we were now continuously engaged in

THE CEYLON PEARL-OYSTER. 269

examining the parasites of the pearl-oyster and their

influence on pearl-formation. We also decalcified such pearls

as were found. This work was continued as time permitted

during the next few weeks, and also by Mr. Hornell after I

left. We found various parasites, in the liver especially,

some of which were Platyhelminthian and others Sporozoan

in their nature, and some of which were enclosed in

caleareous capsules. Mr. Hornell afterwards determined

that these were Tetrarhynchus larvee of Cestodes, and we have

no doubt that they are in many cases the nucleus of the

pearl, and the irritating cause of its formation,”

Again, in the preface to Part IT. of his Report, p. vi, dated July

1904, Prof. Herdman says :

“On the Cheval Paar, in March 1902, we satisfied ourselves

that the ‘Orient’ pearl, free in the tissues of the pearl-

oyster, is deposited around a cyst containing a Cestode larva,

and preliminary notices to this effect were published in my

Royal Institution Lecture of March 27, 1903, and at the

Southport Meeting of the British Association in September

1903.”

On p. 6 of Part V. of the Report (Pearl-Production), Herdiman

and Hornell say :

“One of the first facts that we were able to determine

in connection with the Ceylon Pearl-Oyster, in the spring of

1902, was that the Orient pearl in the Gulf of Manaar is

deposited around the young larva of a Cestode.”

And on p. 15:

“We found the Cestode larvee in association with pearls in

the tissues during our cruises in the ‘ Lady Havelock’ in the

Gulf of Manaar, in February and March, 1902. It was about

March 6th (see Narrative, p.70,in Part 1.), when cutting up

Oysters from the western part of the Cheval Paar, that we

first became convinced that the opaque white globular larvee

we were finding encysted in the liver belonged to Cestode

worms.”

On the other hand, Shipley and Hornell, in their paper on the

Parasites of the Pearl-Oyster (Herdman’s Report, II.), seem to

imply that at least the elaboration of these observations was

carried out subsequently to Professor Herdman’s departure from

Ceylon. Thus, they say (p. 79):

“These larvee first attracted attention during the second

cruise of the ‘ Lady Havelock,’ on March 6th, 1902, when

numbers of the early globular stage were dissected out from

the livers of oysters dredged from the West Cheval Paar.

Subsequently, during the investigation carried out at the

Galle Biological Laboratory, a second and more advanced

stage of a Tetrarhynchus larva was found in the same

270 DR. H. LYSTER JAMESON ON

material. Details of the morphology and histology were

then worked out, and the relationship which the larve bear

to pearl-formation was investigated.”

Strange to say, Professor Herdman’s Preliminary Report to

the Gover nment, dated J uly Ist, 1902 (18), makes absolutely no

reference to the Hiscome: y in the previous March of this important

aspect of the parasites of the Pearl-Oyster.

In the Preliminary Report referred to, Prof. Herdman says

(p. 2):

‘Samples of all the oysters obtained by us were examined

for parasites and for any diseases or abnormal conditions, and

although a considerable number of minute parasites, both

Protozoan and Vermean, were found, still that is by no

means unusual amongst molluscs, and we do not consider

that we saw anything which gave evidence of any epidemic

disease or widespread and injurious prevalence of parasites.”

And again in the same Report (p. 4), in his summary of

conclusions, the Professor says:

‘‘ A considerable number of parasites, both external and

internal, both Protozoan and Vermean, were met with, but

that is not unusual in molluses, and we do not regard tt as

affecting seriously the oyster population.” (The italics are

mine.)

In view of the last three quotations, if it were not for the very

definite assertions in Part II. p. vi, and Part V. p. 6, of Professor

Herdman’s full report, quoted above, I should be almost inclined

to think that, while the Cestode larvee were no doubt discovered

‘in association with pearls” during Professor Herdman’s cruise,

the Cestode theory of Pearl-formation might have been evolved

after Professor Herdman’s return to England, and after the above-

mentioned preliminary report had been submitted. In that case

Mr. Hornell might well have been misled by the false analogy of

the case of the Trematode origin of pearls in Mytilus, which was

dealt with at length in my paper (25). This paper appeared

in August 1902; that is to say shortly after the Professor’s pre-

liminary report of July 1st, 1902, containing no reference to the

Cestode theory, was submitted. The view that my paper might

have misled Mr. Hornell and Professor Herdman would also derive

support from the fact that the real point of my paper had apparently

been missed, viz. that it is not the presence of any parasite, but

the specific stimulation of a narticular kind of parasite that

causes the growth of the pearl-sac. I consider this point is by far

the most important contribution I have so far made to the subject,

and I believe it will be the basis upon which a rational system of

artificial pearl-production will ultimately be built.

The first announcement of Prof. Herdman’s theory of Pearl-

formation seems to have been made at a lecture delivered before

the Royal Institution on March 27th, 1903, an abstract of which

appeared in ‘ Nature’ for April 30th of the same year (14).

THE CEYLON PEARL-OYSTER. 271

(4) Examination or tHe Crsrope THEORY oF Peart-PRodUCTION.

It is unfortunate that more figures of pearls containing as nuclei

supposed Cestodes are not given in Prof. Herdman’s account of

Pearl-formation. The only figures that represent the nuclei of

decalcified pearls examined entire * as transparent objects appear

to be those on plate ii. in the Section on Pearl-Production in

Part V. of the Report, figures 5 and 7, figure 6 representing a

dead Cestode in a partially calcified cyst (not, however, a pear).

On p. 22 it is stated that these drawings, which are reproduced

from Shipley and Hornell’s article upon the parasites of the Pearl-

Oyster in Part II. of the Report, are the work of Mr. Hornell,

and it is not evident from the text that Prof. Herdman had ever

seen the specimens from which they were made. Turning to these

same figures on plate i. of the article by Shipley and Hornell on

the parasites of the Pearl-Oyster (Part Il. of the Report, figs. 5

(A) and 8 (B), (C), (D)), we find them described in the Explana-

tion of the Plates as the nuclei of decalcified pearls; but the same

figures are referred to in the text, p. 80, as representing the

Cestode larva enveloped in its “ tough elastic and fibrous capsule

of spherical form, derived from the adjacent connective tissue

cells.”

It is, I think, hazardous to identify these figures as the remains

of Cestode larvee without examination of sections, and I cannot

help feeling that each of these figures is capable of comparison

with the non-Cestodian centres of pearls described by me below,

It is a remarkable fact that nowhere throughout the Report is

there figured a section of a decalcified pearl showing the Cestode

remains in the nucleus, and to this fact I may add my own

observation that of all the pearls sectioned in situ by Prof. Herd-

man, numbering about 25 (not counting some minute clustered

muscle-pearls), which he kindly sent me to examine, I could not

find a single nucleus that I was able to accept as being a Cestode

or other Vermian parasite. The characters of the nuclei in these

preparations are described in the part of the paper which deals

with my own researches.

The superficial resemblance of the pearl figured on plate ii.

fig. 4A, in Part II. (Parasites) of the Report, and again in Part V.

(Pearl- Production), pl.i. fig. 5k, ¢, d, & e, to the globular Cestode

larvee found in the Oyster is har ‘dly enough to go upon. If such

a pearl consisted of a parasite thinly coated with nacre it would

probably be dark and valueless and not a “fine pearl,” for the

yellowish-brown dead tissue of the Cestode should be clearly

visible through the nacreous coat. It is not stated whether this

pearl was decalcified and sectioned to test whether the resemblance

was more than “ skin deep.”

* These figures are referred to by Southwell (42), p. 128, as “figures of sections

of decalcified pears,” but they are not described as ‘such in the text; and they

certainly appear to be no more than drawings of the centres of pearls ex xamined as

transparent objects.

DR DR. H. LYSTER JAMESON ON

With regard to the mechanism by which the Cestode is supposed

to cause pearl-formation, Prof. Herdman is unable to contribute

much. He seems to recognise that the particular conditions

necessary to transform the Cestode into a pearl-nucleus are not by

any means universally present, and that it is only, so to speak,

under exceptional circumstances that the Cestode, which is very

abundant in the Ceylon Pearl-Oyster, becomes the centre of a pearl.

The larva is surrounded by a connective-tissue cyst, and has not

been satisfactorily demonstrated in any instance with an epithelial

‘* pearl-sac” (such as I described for the Pearl-inducing Trema-

tode in Mytilus), though supposed proliferations of cells inside the

connective-tissue cyst are figured in the Report (Part V. Pearl-

Production, pl. iii. fig. 7). These, being inside a thick fibrous

connective-tissue capsule, are difficult to accept as being equivalent

to a pearl-sac, which I generally find to be surrounded by the

spongy subepidermal parenchymatous tissue, except in the case

of those parts of a ‘ muscle-pearl” into which muscle-fibres are

inserted. From my own observations [ am rather inclined to

regard these “cells” as granules excreted by the parasite itself,

with possibly an admixture of wandering leucocytes. In any ease,

if this 7s an epithelial pearl-sac, what becomes of the thick fibrous

cyst outside it, which is certainly not present around the pearls @

Professor Herdman himself (see below) does not think the Cestodes

enveloped in thick connective-tissue cysts are destined to become

nuclei of pearis.

The supposed migration of ectoderm-cells into the wall of a

pearl-sac already formed and already containing a pearl, as figured

in Part V. (Pearl-Production), pl. i. figs. 18-20, seems to be a

matter quite apart from the question of the primary origin of the

pearl-sac.

On p. 23 of Part V. Prof. Herdman says :—

“Tt is quite evident from the examination of a large series

of sections, such as we have worked through, that the

majority of these encysted parasites do not become encased

in pearls. Probably none of those in thick connective-tissue

cysts are destined to form nuclei. They are awaiting their

legitimate further development in the next host, after their

sheltering mollusc has been devoured by a fish. In such

cysts and around such parasites we find no epithelial sac, and,

as a consequence, there can be no pearl. Whether or not it

is the case that only dead parasites supply the stimulus

necessary to induce pearl-formation, and whether, as Giard

has suggested, the parasites may be infested and killed by a

species of Glugea, so that that Sporozoan comes to be even-

tually responsible for the pearl, we are not prepared to say

—we have found no fresh evidence in the Ceylon material

bearing upon that point. It seems clear to us, however, that

the epithelium is always associated with pearl-formation, and

that in the absence of the epithelium only a thick-walled

connective-tissue cyst is produced. If we adopt the view (see

THE CEYLON PEARL-OYSTER. Dike

below) that this epithelium is genetically related to the

ectoderm, then a possible explanation of the difference in

behaviour in the encysted condition would be that those larvee

that carried in ectodermal cells become covered (when dead or

while still alive) by a pearl-sac and embedded in a pearl,

while those that were free from ectoderm become surrounded

by the connective-tissue cyst.”

No satisfactory instance, however, is recorded of the Cestode

parasite being observed surrounded by an epidermal sac.

Again in Part III. of the Report, p. 32, Professor Herdman,

quoting a Report furnished to him by Mr. Hornell, says the

abundance or otherwise of cyst-pearls “ is connected with the

factors which control the relative abundance of the pearl-inducing

Cestode and those which conduce to its death during encystment

in suitable localities within the tissues—problems as yet obscure ” ;

while in Part V.(Pearl-Production), p. 15, he says “it is appar-

ently very difficult indeed to hit upon a stage showing the

commencement of the pearl-formation.”

And again, in an address delivered at the Anniversary Meeting

of the Linnean Society of London in 1906 (17), Prof. Herdman,

speaking of his Ceylon work, says that it is probably only those

Cestodes that are provided with an ectodermal covering forming

a pearl-sac that become the nuclei of pearls. But, as stated above,

such an ectodermal pearl-sac has not yet been found to occur

around the parasite, and its occurrence is purely theoretical.

The investigations made subsequently to the publication of

Prof. Herdman’s Reports have added little to our knowledge of

the subject. In 1905 Mr. Hornell published a Report on the

Placuna placenta pear|-fishery of Lake Tampalakamam, dated

June 15th, 1905 (21). In this he states (p. 5) that he dissolved

two Placuna pearls (out of five in his possession) and found that

“in each case the nucleus proved to be the dead remains of a

minute Platyhelminthian larva of the same stage and species as

that which forms the nucleus of cyst-pearls in Margaritifera vul-

garis.” Further study revealed the presence of Cestode larve in

the dorsal portion of the visceral mass. He considers that these

are identical in details of form and structure with those of Mar-

garitifera vulgaris, but expresses some doubt as to their specific

identity, and adds that if they prove distinct the Placuna

parasite will need a fresh name. Mr. Hornell further observed

that these larve multiply asexually by a process of endogenous

budding, which he wrongly refers to as parthenogenesis.

Mr. Hornell then goes on to make the following extraordinary

statement, which I quote as giving some indication of the con-

fusion of ideas which existed as to the identity of the supposed

pearl-producing larve even in 1905, 7. e. nearly three and a half

years after the beginning cf the observations :—

“The discovery of a stage in the life-history of this

parasite, which I am confident is homologous with the

Proc. Zoou, Soc.—1912, No. XVIII. 18

274 DR. H. LYSTER JAMESON ON

fedia-stage of a Trematode, confirms my original idea of it

being a larval Trematode—an idea formed when first I saw

it in March 1902” [7.e. the Cestode larva in Margaritifera

vulgaris|. “* Other facts point to the same conclusion, and 1

have now no doubt on the subject. The genus to which it

belongs is still doubtful, but as it is inconvenient not to have

a name whereby to make mention of it, I shall henceforth

refer to it under the cognomen of Distomum (¢) margariti-

factor, n. sp., the specific name having reference to the fact

that it is the inducing cause in the production of ‘ fine’

pearls.”

Mr. Hornell anticipated that asexual reproduction would be

found to occur in the Cestode parasites of Margaritifera vulgaris

also, and this has since proved to be the case.

An expurgated edition of the above report was published in

1906 (22), which, however, bore the same date, June 15, 1905, as

the Sessional Paper. In this the statements as to the ‘supposed

Trematode nature of the parasite were suppressed (though not

formally withdrawn), the name “Distomum margaritifactor, n.sp.,

cancelled with a stroke, still figuring in the hthographed plate

(Annexure IT.).

As explained below (p. 345) I have been unable to confirm the

presence of Cestodes in the centres of Placuna pearls from Ceylon,

any more than I can find them in the pearls of Margaritifera

v ulgari ws.

In 1907 Dr. A. Willey (48) confirmed and extended Mr. Hornell’s

observations on the endogenous reproduction of the Placuna

Cestode.

Mr. Southwell discovered (39, p. 173) that endogenous asexual

reproduction or budding, similar to that described by Hornell and

Willey in the parasite of Place. occurs occasionally im the Cestode

parasites of the Pearl-Oyster. He only observed the occurrence

twice, in November 1906 and January 1909 —in each case a single

endogen was found.

In 1903 the late Professor A. Giard (10) announced that

M. L. G. Seurat believed that in the black-lipped Pearl-Oyster

of the Gambier Archipelago (Margaritifera margaritifera var.

cumingit Reeve) pearl-formation was due to the presence of a

parasite, figures of which were given, and which Prof. Giard

vereererl i, a genus near to C ‘yathophyllus | Cyathocephatus ?|

Kessl. or Acrobothrium Olsson.

Subsequently Seurat found the adult of this worm in the

Eagle Ray (4étobatis narinari Kuphr.) in the spiral intestine, and

named it Vylocephalum margaritifere (36). The adult, which is

figured by Seurat (37), is quite a minute worm, not exceeding

4 mm.

The scolex occurs in cysts, similar to those occurring in the

Ceylon Pearl-Oyster, and multiple cysts (perhaps formed by bud-

ding of the larva as in the Placwna parasites) occur also. Seurat

THE CEYLON PEARL-OYSTER. ‘ 275

states that these cysts which form around the parasites become

the nuclei of pearls, and a decalcified pearl shows an organic

nucleus in the centre surrounded by concentric layers of conchyolin,

the whole having a diameter of about a millimetre, and the nucleus

being a scolex 225 mm. long and easily recognisable as that of

Tylocephalum.

It appears that in this case also the parasites may be present

in great numbers without pearls being found (Seurat (35), 1904,

p. 295).

Here, again, examination of the scanty material available (see

p- 346) has yielded no confirmation of the presence of Cestodes in

the pearls of MZ. margaritifera var. cumingit. I am endeavouring

to obtain further material from the Eastern Pacific, in order to

extend my observations on this species.

I may say that from the first time I read Professor Herdman’s

Reports and papers on the subject I was sceptical as to the

relationship of the Cestode to pearl-formation.

Indeed, before Prof. Herdman’s departure for Ceylon, on

examining Dr. Kelaart’s material at the British Museum, which

Mr. E. A. Smith kindly placed at my disposal, I had detected the

existence of these Cestode jarvee (which Kelaart seems to have

regardedas “eggs of Entozoa”) in their connective-tissue cysts in

the Ceylon pearl-oyster, and after having examined the larve, and

also having decalcified pearls from the same oyster, dismissed the

Cestode as probably not concerned in pearl-formation.

My chief grounds for doubting the Cestode theory were the

following :—

(a) The absence of evidence that the Cestode ever occurred in

an epidermal sac, and the fact that it was almost invariably sur-

rounded by a fibrous capsule or cyst which does'not occur around

the pearl. ui

(6) The large proportion of the Cestodes that showed no sign

of becoming pearl-nuclei, pointing to the conclusion that pearl-

formation does not necessarily, or even normally, follow from

infection. Thus, in a footnote to p. 12 of Part V. of his Report,

Prof. Herdman says :

‘“* In comparing these statistics [7. e. of numbers of parasites

and of pearls in Mytilus] with those of the Ceylon pearl-

oyster, one is struck by the wholly different ratio borne

by pearls to parasites in thetwo cases. In the mussels,

pearls are far more numerous than the living parasites. In

our Ceylon oyster, parasites may be exceedingly abundant * ;

while pearls (cyst-pearls) are relatively very rare, probably

not more than one to a hundred parasites.”

* Mr. Southwell (42), p. 128, says: ‘‘ As many as 120 have been counted in a single

oyster”; and further down on the same page: ‘‘ Occasionally several hundred oysters

can be examined, each containing 20 or 30 cysts, and not a single pearl is to be

found.” ‘

cS"

276 DR. H. LYSTER JAMESON ON

In this connection an observation made by Dr. Willey (49)

is very significant. Dr. Willey says :—

‘« A remarkable fact, indicating the subtle dependence of

the pearl-producing molluscs upon their environment, is that

whereas the most valuable pearls, called cyst-pearls by Prof.

Herdman, are formed about a parasite as their centre or

nucleus, yet the presence of these parasites in great numbers

does not necessarily predetermine the formation of pearls.

Pearl-oysters at Trincomalee may be heavily infected with

the parasites without yielding pearls. It may be said that

the parasitic infection and the pearl-disease are two separate

phenomena, the latter proceeding from the former under

certain conditions which are realised in the Gulf of Manaar.

Whether these exact conditions can be reproduced elsewhere

is one of the main problems before the Company. In the

same way the cultivation of the oysters and the multipli-

cation of pearls are two separate operations, the latter

proceeding from the former in response to certain conditions

affording the suitable stimulus. Conditions may favour the

bivalves, but not their parasites; or they may favour both

hosts and parasites, but not the production of pearls.”

Report that Mytilus pearls (which he examined in order to be

able to correlate his work with mine) differed from Ceylon pearls

in “the large size of the nucleus in the pearl (where a nucleus is

present) and its characters, which are quite different from those of

the encysted parasites in the Ceylon Pearl-Oyster.” Now the

nucleus of a A/ytilus pearl is generally about 0°5 mm. in diameter—

the size of the Trematode when contracted into a sphere. As the

Ceylon Cestode-parasite measures roughly from 0:5 to 1 mm. in

diameter, the nucleus of a Ceylon pearl, if composed of its

calcified remains, should if anything be larger, rather than

smaller, than that of a Mytilus pearl. And the characters of the

nucleus should not differ greatly—the dark opaque yellowish or

brownish substance formed by the decomposition and subsequent

calcification of the parenchymatous and muscular tissues of

a Trematode should not differ materially in appearance from the

analogous remains of a dead Cestode.

For purposes of comparison I figure below (text-fig. 33, A & B)

a Trematode pearl from J/ytilus, from Foulney, Lancashire

(Preparation CIII). A shows the pearl decalcified and examined

entire in oil of cloves, B shows the nucleus in section. In both

cases the foreign nature of the nucleus is obvious, quite apart

from the fact that in this preparation its Trematode character is

quite clear (which would, of course, not be the case where

decomposition had advanced considerably before calcification

commenced). The characters of this nucleus are quite different

from those of the pseudo-nuclei of Ceylon pearls figured in the

THE CEYLON PEARL-OYSTER. PART (

plates, the concentric stratification of the majority of which

never occurs in a ‘Trematode nucleus, and could hardly be expected

in a Cestode.

Text-fig. 33.

A Mytilus pearl, from Foulney, near Piel, Lancashire, with a Trematode nucleus :

A, examined entire in oil of cloves, after decalcification : B, the same in section.

In A the oral sucker and digestive cxca of the worm are distinctly visible.

In B the internal anatomy is still preserved. cu., cuticle; sp., spines on

same; dig., digestive cecum ; skr., ventral sucker; at dand q ave dark masses,

which may well represent the remains of the yolk-glands and gonads;

nac., nacre. C,a Mytilus pearl of non-Trematode origin, from Plymouth.

Here the nucleus is, as in many Ceylon pearls, a minute group of granules.

A X 20; B X 70; C x 20.

As Mr. Cyril Crossland, Marine Biologist to the Sudan Govern-

ment, is quoted by Professor Herdman (Report Ceylon Pearl

Fisheries, Pt. V. Pearl- Production, p. 3) as supporting the Cestode

theory, so far as MW. vulgaris in the Red Sea is concerned, I wrote

to him to ask him for further information. He replied, in

a letter dated December 9th, 1911 :—*I never published any state-

ment that Cestode larvee caused pearl-formation in the Red Sea.

The evidence to my mind is in need of revision. In all cases the

first result of excessive stimulation of the secretory epidermis of

the mantle is the formation of a dark brown horny material

[?.e. my amorphous substance.—H. L. J.]. How would this stain

in sections, and is it cellular like the horny material of the prismatic

layer? If so, would not a shrunken nucleus of such material

resemble the dry remains of a Cestode? This is a criticism which

I have had in mind several years, and have never put it to

the test.” From this it is clear that Mr. Crossland, though cut

off from the possibility of applying modern laboratory technique,

has arrived at much the same conclusion as that which I am

elaborating in this paper.

To summarise the supposed relation between Cestodes and

278 DR. H. LYSTER JAMESON ON

Pearls, as described by Professor Herdman and Mr. Hornell, the

position is briefly this :—

‘1) Ceylon Pearl-Oysters were found to contain large

numbers of Cestode parasites which occurred simul-

taneously with pearls, but which did not necessarily

result in the formation of pearls.

(2) The nuclei of the majority of “ cyst-pearls” were

thought to be identified as consisting of the remains of

these parasites, though Mr. Hornell’s figures of such

nuclei are capable of other interpretation.

(3) No satisfactory evidence was adduced of the Cestode

having acquired a surrounding epidermal sac, such as is

normally formed around the pearl-producing Trematode

in Mytilus, though Prof. Herdman admits that this sac

is essential for pearl-production. ‘The first stages in the

supposed process are therefore purely hypothetical and

unsupported by observation, besides pre-supposing an

abnormal departure from the parasite’s usual habit.

The evidence in favour of the theory is mainly that the more

highly infected the oysters are with these particular Cestodes, the

richer they are in pearls. Thus it was observed in 1904 (Report

III. p. 32) that the oysters from the North-West Cheval, besides

being the most extensively infected with Cestode-cysts were also

the richest in cyst-pearls. And, again, Mr, Southwell records

(40, p. 194), that

“the infection of the very old oysters | with tapeworm-cysts |

found on the Kondatchi Paar in 1908 was remarkably low,

and, as bearing practical proof that infection and pearl yield

are intimately connected, it is interesting to note that the

pearl yield also was remarkably low, the valuation only

working out at about Rs. 18 per 1,000 oysters.”

These facts might be explained, however, on the assumption

that the conditions favourable to pearl-production are also favour-

able to Cestode infection.

Characters, Identity, and Life-Histories of the Cestode Parasites

of Margaritifera vulgaris.

Apparently the first announcement of the supposed relation

between the Cestode and pearl-production was made by Prof.

Herdman at his Royal Institution lecture on March 27th, 1903

(14). Inthis he says that he and Mr. Hornell have proved so

far ‘that in Ceylon the most important cause is a larval Cestode

of the. Zetrarhynchus form.” Again, in the Report of the British

Association, Southport, 1903, p. 695, Prof. Herdman says: ‘‘ The

parasite in the case of the majority of the cyst-pearls of Ceylon

is the larva of a Cestode which appears to be new, and will

be described under the name of ‘ Tetrarhynchus unionifactor’

and the pearl-inducing parasite is referred to throughout Prot.

Herdman’s Report under this name.

THE CEYLON PEARL-OYSTER, 279

But the worm specifically described by Shipley and Hornell as

'etrarhynchus unionifactor ou p. 88 of Part IT. ‘of Prof, Herdman’s

a (Parasites of the Pearl-Oyster) and figured in plate il.

figs. 19 & 20, is a well-advanced Tetrarhynchus 65 to 7 mm.

long, which occurs in and around the intestine of the Pearl-

Oyster; and, to say the least of it, it is doubtful whether this worm

isa later stage of the globular cysts, which Prof. Herdman identi-

fied as the nuclei of pearls, and not an entirely distinct organism.

In order to avoid confusion of terms I am therefore giving

separate names to the larger and smaller globular Cestode larvee

which Herdman recognises, as it is caleulated to lead to much

confusion of issues if Frese are referred to by the name of T'etra-

rhynchus unionifactor before their identity with it can be demon-

strated more satisfactorily. The arguments for regarding the

supposed pearl-producing parasites as distinct from TVetrarhynchus

unionifactor are set forth below. In the absence of satisfactory

evidence of their relation to the genus 7etrarhynchus, | propose,

following Seurat (36), to whose larval Cestode, mentioned above,

they bear a close resemblance, to refer them to the genus 7'yloce-

phalum* and to describe the larger and smaller forms respectively

as T'ylocephalum ludificans, sp. n., and T'ylocephalum minus, sp. 0.

Two well-marked sizes occur in these globular larve and they

are regarded by Herdman as distinct organisms (Report V. p. Bay.

On the other hand, Southwell coneiders that the asexual repro-

duction, which he has occasionally observed, accounts for the

varying sizes of the larve in the Ceylon Pearl- -Oyster, and says

‘‘T am now convinced that these different sizes merely represent the

same species in different grades of development.” | am inclined

to share Prof. Herdman’s view that these two sizes are distinct

organisms. It may even prove that there are more than two

species represented. Jndeed, I should not be surprised if further

research on fresh material were to show that both 7'ylocephalum

ludificans and 7’. minus are composite species.

Professor Herdman regards 7’, ludificans as the earlier stage

of Tetrarhynchus unionifactor, and calls attention to its re-

semblance to Van Beneden’s ideal figures of the young of

Tetrarhynchus, while he treats 7. minus as another species of

Tetrarhynchus in its earlier stages. Nevertheless, he seems to

have had suspicions that some, ‘at least, of these larvee might be

Tylocephala, though he appears in the end to have decided that

they—and presumably with them Seurat’s larva—are a hitherto

unknown stage in the life-history of the genus Tetrarhynchus.

In the Preface to Part II. of his Report he says (p. vi) :—

“Tt is possible that some of our Ceylon Pearl-Oyster

Lape may also belong to the genus Acrobothrium’

.e. the genus to which Sans at’s larva was then referred |,

a ‘although. the more advanced ones are certainly Tate

rhynchids ” ;

* J,inton (27 a). pp. 805-9, pl. ix. figs. 5-9. Type 7. pingue, from spiral valve

of Rhinoplera quadriloba.

280 DR. H. LYSTER JAMESON ON

while in Part V. of the Report, p. 14, he and Hornell say :—

‘“‘Some of our Ceylon Pearl-Oyster parasites very closely

resemble the figures given by Giard” [7. e. of Seurat’s larva]

‘“and possibly may also belong to the genus Cyathocephalus

| Tylocephalum|, although most of them are certainly Tetra-

rhynchids ” ;

and on pp. 16-17:

‘““Tt is possible, however, that more than one species of

Cestode is represented—one is certainly a species of Tetra-

rhynchus (Rhynchobothrius), and another is probably the

same genus, or may possibly belong to Cyathocephalus ... .”

Later on, however (p. 20), Herdman and Hornell reject the

idea that the globular larvee may be Tylocephala or allied genera,

and, in discussing the opinions of Giard and Seurat on the

systematic position of Seurat’s larva, they say that they regard

the terminal invagination, not as a sucker with a papilla on its

floor, but as

“the opening in a hood or depression formed by the

sinking of the scolex into the front of its vesicle. The

changes of shape which we observed in this larva in the living

state, the protrusion and retraction of the papilla-like part

which we regard as the anterior end of the scolex, agree with

this interpretation. Consequently, we are of opinion that this

larval Cestode is not one of the Monobothria—that it belongs

to neither the Pseudophyllidea nor the Tetraphyllidea, but is

a young Tetrarhynchid belonging to the Trypanorhyncha, and

we give here (fig. 4) a series of diagrams in order to show

the positions that we suppose our stages to occupy in the

development oi,such a form.”

Shipley and Hornell (Herdman’s Report IT. p. 80) call attention

to the resemblance of ‘older examples of the larger larva (7'ylo-

cephalum ludificans) to Seurat’s form, and think there is little

doubt that they are at least generically the same (p. 82). Again,

Southwell says (39, p. 169) :

“Tt would certainly appear more probable as well as

simpler, for this larva to develop into a Tylocephalum (as is

believed by Seurat) than into a Tetrarhynchus.”

Again, Southwell, speaking of the great scarcity of the adult

of Tetrarhynchus unionifactor in Klasmobranchis taken by trawling,

says (42 p. 130):

‘“‘ Tt would almost appear that this fact in itself is sufficient

proof that the adult of the pearl-inducing worm is not

Tetrarhynchus unionifactor.”

But at the foot of the same page he reverts to the position

that it isa Tetrarhynchus.

THE CEYLON PEARL-OYSTER. 28]

It is strange how the Tetrarhynchus unionifactor theory,

once enunciated, has prevailed :—-indeed nobody seems to have

seriously followed up the obvious clue given by Seurat’s discovery

of the supposed adult of his worm.

Prof. Herdman says in his Report, Part V. pp. 20-21:

“Shipley and Hornell in Part IL. (p. 77) described and

figured various stages of the Cestode larve both from the

centre of decalcified pearls and also free in the tissues of the

pearl-oyster, but left it an open question whether the sub-

globular younger larvee [i. e. Z'ylocephalum ludificans, n. sp.|

belong to the same life-history as the elongated older forms,

which are young Tetrarhynchids. If our arrangement of

the stages observed in the tissues of the pearl-oyster is

correct, and if all these larve belong to the same species,

then the interpretation we have given above brings us to

the conclusion that the larger of our two globular larve

belongs to the worm which Shipley and Hornell described as

Tetrarhynchus unionifactor in 1904.”

And, referring to some figures of 7'ylocephalum ludificans on

pl. i. (Report, Part V. Pearl-Production) figs. 1-8, he says

(p. 21):

“there can scarcely be any doubt (1) that they all belong

to the same life-history, and (2) that they are young Tetra-

rhynchids leading on to the stages shown in figs. 10 and 11.”

Prof. Herdman gives, in support of his theory, a series of

figures showing the hypothetical transition from 7'ylocephalum

ludificans to Tetrarhynchus unionifactor (Report, Part V. p. 21).

Later, a younger Tetrarhynchus, 1 mm. in length, was found

ia the stomach and alimentary canal of the o ster, which Shipley

and Hornell (Report V. pp. 87-88) regarded as probably an

earlier stage of Tetrarhynchus unionifacior, though possibly a

distinct form. A single example of a still younger form, which

is figured in the Report on Pearl-Production (V., pl. iii. fig. 10),

and is described in the text (p. 22) as occurring “ encysted in the

liver” and in the explanation of the plates as “ from cyst between

stomach and liver,” appears to go still further towards linking

the two forms. This larva is referred to in the text as °53 mm,

long. T'ylocephalum ludificans grows to a much greater size than

this without changing its characters; thus the one figured by

Shipley and Hornell in the “Parasites of the Pearl-Oyster”

(Report, Part IT. pl. i. fig. 12), and described as x40, appears

from the size of the figure to be over 1:5 mm. long, and still

shows no sign of becoming a TJ'etrarhynchus. And T. ludificans

quite frequently measures 1 mm. in diameter. This discrepancy

in dimensions makes it hard to believe that they are the same

organism, and the gap between the 7Z'ylocephalum form, with its

round Balanoglossus-like scolex or ‘ myzorhynchus,” and the

282 DR, H. LYSTER JAMESON ON

Tetrarhynchus form, with its complex proboscides, cannot be said

to have been satisfactorily bridged.

Herdman found the Tetrarhynchus form much scarcer than

ae E ylocephalum form, and it appears from his Report, Part V.

p. 22, that the ratio of the latter to the former is about 200: 6.

Shipley and Hornell (Report, Part II. p. 79) give the ratio of

globular larve to undoubted Tetrarhynchi as 100: 1.

Prof. Herdman’s suggested explanation of this, namely, that the

globular parasite only occasionally advances to the Tetrarhynchus

stage, requires, it seems to me, a greater effort of the imagination

than the hypothesis that the two worms are distinct forms.

It is hard to conceive of conditions that would lead a small

minority of Zylocephalum ludificans or 7’. minus to leave their

tough fibrous cysts in the peripheral tissues, and migrate to the

sities dine, there to take on the Vetrar Hneios form. It seems to

me much easier to regard these as two (or three) distinct species,

and their simultaneous presence in one host as a case of parallel

infection.

In his latest paper (42, p. 129), Southwell, speaking of these

undoubted Tetrarhynchids, says :—

“These are by no means rare, and are in almost every

case confined to a particular part of the wail of the gut, about

one inch from the anus and on the terminal part of the gut.

They often occur in clusters of three or four. They are

small (about 1 mm.), but appear to be adult in every way,

save that strobilization has not commenced. This encysted

young ‘Tetrarhynchid is quite dissimilar to the globular cysts

found in the same oyster. In the latter case, the larvee are

so young that the Cestodian characters are te ill defined.

Tn the ronnen case, a normal and full-grown Tetrarhynchid

head is present. No stage or stages have been found inter-

mediate between them, and the evidence that they are both

stages in the life- history of the same parasite rests on cireum-

stantial evidence and on evidence obtained by feeding

experiments.”

And with reference to these feeding experiments, which are

referred to below (p. 287), and in which Zetrarhynchi were found

in Sharks that had been fed on pearl-oysters, Mr. Southwell says

(Ga, XO) e—

‘““The mere fact that the adults were obtained by feeding

is in itself almost sufficient to prove that they are the

adult of the pearl-inducing worm, for it is difficult to believe

that their occurrence in the (Ginglymostoma was a mere

coincidence each year.”

I think there is very good reason to believe that Southwell

did, in his feeding experiments, actually transmit Tetrarhynchus

unionifactor from. the Oyster to the ‘Elasmobranch, but it is

THE CEYLON PEARL-OYSTER. 283

difficult to escape the conclusion that the worms found in Gingly-

mostoma were derived from the Vetrarhynchus larve in or around

the alimentary canal of the Oysters, and not from the globular

Tylocephala in the other tissues, to which Southwell refers when

he speaks of the “ pearl-inducing worm.” ‘To dispute this view,

it would be necessary to demonstrate that the Zetrarhynchus-stage

was not present in the Oysters used.

Shipley remained throughout sceptical about the identity of

the supposed pearl-forming larvee with Zetrarhynchus unionifactor.

In Part II. of Herdman’s Report, p. 86, he says it is most

improbable that the young larve grow into the Zetrarhynchus

larva. In their report on the Cestode and Nematode Parasites

from the Marine Fishes of Ceylon, Shipley and Hornell say

(Pt. V. p. 66) :—

“It seems increasingly probable that the pearl-forming

Cestode is a 7. unionifactor, but this has not yet been

proved.”

Shipley and Hornell, in Herdman’s Report, Part V. p. 98, offer

the following hypothetical life-history :—

“Of the given number of larvae which enter at a very

early stage into the body of the Oyster a certain number arrive

in the mantle and other tissues, acquire an ectodermal sac and

there encyst, and find a costly grave in the developing pearl.”

[The ectodermal sac around these parasites is so far purely

hypothetical and has never been demonstrated.—H. L. J.]

“The remainder, however, reach the alimentary canal and

grow and flourish there. When they attain the dimensions of

thestages described in Part IT. they leave the alimentary canal

and encyst, usually upon the outer surface of the intestine.

Now they are too big for enclosure in a pearl, and they can

wait without anxiety for the advent of their second host

(Khinoptera javanica), within whose intestine they rapidly

become sexually mature.”

It would seem to the present writer much simpler to set aside,

for a while, the hypothesis that Tylocephalum ludificans and

7’. minus are younger stages of a Tetrarhynchus, and to seek for

their adult stages among the members of the genus Tylocephalum,

or allied types described as new genera, occurring in oyster-eating

Elasmobranchs. Shipley and Hornell have already described a

number of these, which I give below :—

*tTylocephalum (Tetragonocephalum) trygonis (Report, Part

III. p. 51 and Part V. pp. 48 & 83). Habitat: intestine of

Trygon walga and Aétobatis narinari. Diameter of head

0-03 mm.

*Tylocephalum (Tetragonocephalum) aétobatidis (Report, Part

III. p. 52 and Part V. p. 48). Intestine of <Aétobatis

narinari, Diameter of head 1°5 mm,

284 DR. H. LYSTER JAMESON ON

TTylocephalum dierama (Report V. p. 59). Intestine of J/ylio-

batis maculata. Diameter of head °6 mm.

Tylocephalum kuhli (Report V. p. 72). Intestine of Zrygon

kuhli. Head apparently about *5 mm. in diameter,

+T'ylocephalum uarnak (Report V.p. 76). Intestine of Z'rygon

uarnak and Trygon walga. Head apparently about 1 min.

in diameter.

*+Cephalobothrium aétobatidis (Report V. p. 44). Spiral valve

ot Aétobatis narinart. Diameter of head °5 mm.

*tAystocephalus translucens (Report V. p. 46). Intestine of

Aétobatis narinart. Diameter of head -4 mm.

*Hniochobothrium gracile (Report V. p. 64). Intestine of

Rhinoptera javanica. Head appears to measure about

‘4 mm. in diameter.

*tTiarabothrium javanicum (Report V. p. 67). Intestine of

Rhinoptera javanica. Head 1 mm. broad.

To these may be added two forms described by Mr. Southwell

(41 a), viz.:—

Cephalobothrium abruptum, from the spiral valve of Pteroplatea

micrura. Head 1:2 mm. broad.

Cephalobothrium variabile, from the intestine of Pristes cuspi-

datus. Head ] mm. broad.

The forms marked * occur in hosts which are known to feed

on pearl-oysters. Those marked f+ I have been able to examine

through the courtesy of Dr. Shipley. It is impossible not to be

struck with the resemblance between the heads of some of the

above species and the parasites which Herdman associates with

pearl-formation. Compare, for example, the head of Cephalo-

tothrium aétobatidis, figured in Part V.(Cestoda) plate i. figs. 1-4,

with some of the figures of Zylocephalwm ludificans in the chapter

on Pearl-Production (Part V. Pearl-Production, plate ii.) or

the heads of Tylocephala (Tetragonocephala) as figured on the plate

of Part IIT. (Parasites), and in the article on Cestodes in Part V.

(pl. v. figs. 76-7), with some of the figures in the article on the

parasites of the Pearl-Oyster in Part II. Or, again, compare the

section of Tylocephalum ludificans from the pearl-oyster shown

on Pl]. XLVI. fig. 58, with the head of 7. warnak, Pl. XLVI.

fig. 65, drawn from one of Dr. Shipley’s slides, asa representative

of the genus Z'ylocephalum, taking into account the difference

that in the former the myzorhynchus is withdrawn within the

collar, whereas in the latter it is fully protruded. Or compare

Pl. XLVI. fig. 59, a section of 7. ludificans from the pearl-

oyster, with the head of Cephalobothriwm aétobatidis, from one of

Dr. Shipley’s slides shown on Pl. XLVII. fig. 66. Similarly,

compare figs. 58 & 59 with the worm shown in figs. 61 & 62 on

Pl. XLVIL., also from 4 étobatis narinari (the final host of Seurat’s

larva). Ithink the worm shown in figs. 60-64 is quite probably

t It is strange that the oyster-eating habits of this Ray do not scem to have

been recorded in these Ceylon researches.

THE CEYLON PEARL-OYSTER, 285

the adult of Tylocephalum ludificans—in fact, the only difference

appears to be the presence of the four lateral suckers, which are

absent in the larva, but which may well not develop till its

transference to the final host.

Comparison of these figures makes one feel doubtful whether

the generic distinction between 7'ylocephalum and Cephalobothrium

is a valid one, or whether the conditions shown in figs. 58 & 65,

and in figs. 59 & 66, are not merely the expression of a uniform

type of myzorhynchus in different stages of contraction, as

suggested diagrammatically in the following text-figure.

9 f

« Oe

Diagram illustrating the relationship between the condition of the myzorhynchus in

Tylocephalum and Cephalolothrium. A, a generalised scheme of a scolex

such as the Tylocephala of the pearl-oyster. B, by protrusion of the

myzorhynchus, the outer surface of the same being tense, the inner surface

thrown into folds, the Tylocephalum-form may be produced (compare

Pl. XLVII. fig. 65). C, the partial retraction of ‘the myzorhynchus to form

a sucker-like disk gives the Cephalobothrium condition (compare Pl, XLVI.

fig. 59 and Pi. XLVII. fig. 66). D, the myzorhynchus retracted within

its collar, with its outer surface thrown into folds, as is characteristic of

many of the larve of Tylocephalum ludificans found in the pearl-oyster, and

of the adult worm figured on Pl. XLVII. figs. 61-62.

It does not necessarily follow that any of the above mentioned

worms actually represents the final stage of 7’ylocephalum ludificans

or 7. minus, though I think there are considerable grounds for

regarding the worm shown at figs. 60-64 as the former; but it

certainly appears more probable that these final stages will be

found among this class of parasites rather than among the

Tetrarhynchi, and it is strange that the position that the Tetra-

rhynchus unionifactor hypothesis may be wrong has never been

seriously faced and a fresh start made on the above lines.

It is not known how J'ylocephalum ludificans and Tylocephalum

minus enter the Pearl-Oyster; but Hornell found, in tow-netting

on the Muttuvarattu Paar on the 19th November, 1902, a free-

swimming larva, *37 mm. long when extended, which is figured

in Prof. Herdman’s Report (Part II. Parasites of the Pearl-

Oyster, plate i. fig. la-h). This larva certainly suggests an earlier

stage of T'ylocephaluwm ludificans, and one of the chief difficulties

in the way of accepting it disappears if this worm is dissociated

from the genus Zetrarhynchus, the normal habit of which is to

enter the digestive canal with the food while still in the egg-

286 DR, H. LYSTER JAMESON ON

stage. (The fact that the undoubted Zetrarhynchi in the pearl-

oyster occur in and around the digestive canal suggests that

they follow the normal course and are swallowed in the egg-stage,

and first hatch out in the intestine of the oyster.) Mr. South-

well states (Ceylon Marine Biological Reports, Part IV. No. 6,

p. 169, 1910) that this free-swimming larva has not been seen

since it was first discovered (see also 42, p. 127).

IT may here mention that one of Prof. Herdman’s slides which

IT examined shows an interesting phase in the biology of these

supposed pearl-inducing Cestodes which may have escaped him.

It shows a small Cestode, 0°12 mm. long, with myzorhynchus and

collar fully developed, clearly in the act of passing through the

tissues. This may possibly represent the young of either form,

when first entering the oyster, or it may be a case of voluntary

or accidental migration by 7'ylocephalum minus (Pl. XX XITI.

fig. 1). Dr. Willey (48, p. 50) records a similarly free larva seen

moving about in the liver of a species of Vens.

To return to the life-history of the true Tetrarhynchus unront-

factor, Shipley and Hornell have shown without doubt that the

adult sexually mature worm occurs in the Ray, Rhinoptera javanica

(Herdman’s Report V. Cestodes, pp. 65-66). The identification

of the final host as Rhinoptera javanica is announced by Mr.

Hornell in a postscript to his Report on the November Inspection

of the Pearl Banks, 1904 (20, p. 8). Mr. Southwell (42, p. 130)

gives Zeeniura melanospilos as another host.

Professor Herdman in his Royal Institution Lecture, and in

Pt. I. p. 12 of his Report, claimed the File-fish, Balistes, as an

intermediate host ; but Shipley and Hornell, in Herdman’s Report,

Part II. p. 83, say that “a more minute examination, however,

renders the connection between the parasites of the pearl-oyster

and those of the file-fish a doubtful one”; and the immature

Tetrarhynchi found in this fish are described as distinct species

under the names of Zetrarhynchus balistidis and 7’. pinne. Prof.

Herdman sums up the position in the article on Pearl-Production

(Report V. p. 24) by saying :—

“No fresh light has been thrown upon the possible

occurrence of an immature stage in Balistes (which is eaten

by the large rays), and although that intermediate host may

not be necessary to the life-history, since the rays also feed

upon pearl-oysters, still there is nothing in the observed facts

to forbid the existence of such a stage, and it is not unusual

in Tetrarhynchids to have two fish-hosts, an intermediate

Teleostean which is devoured by a final Elasmobranch.”

Mr. Southwell’s subsequent investigations confirm Prof. Herd-

man’s view that Balistes occurs as a collateral intermediate host

or “carrier”; he says (42, p. 132):—“ It is certain that my

eneysted Tetrarhynchus unionifactor from Balistes is not the

same species as those described” (2, e. by Shipley and Hornell

THE CEYLON PEARL-OYSTER, 287

from Balistes); and he claims that they are “ exactly similar to

the encysted Tetrarhynchid found in the oyster ”—~. ¢. the true

Tetrarhynchus unionifactor, except that they are slightly larger.

Mr. Southwell is satisfied that they are derived from the oysters

eaten by Balistes, and thinks that they are derived from

both the genuine Tetrarhynchids in the Oyster’s intestine and

from the globular cysts in its tissues. He considers that if

Balistes is eaten by an Elasmobranch, the young worms become

alult ; but Balistes is not « necessary host, it is merely a “ carrier.”

Johnstone (26 b) confirms the view that the Teleostean host is

a collateral one, and not a normal stage in the life-history, by his

recent researches on the European Tetrarhynchus erinaceus.

The adult stage of this worm occurs in various species of Skates

and Rays, and the first host is probably some Invertebrate.

Johnstone regards the frequent occurrence of a larval form of

this worm in Teleosts as a “ cul-de-sac” stage, due to the first

host being eaten by the wrong fish; and as normally leading no

farther, but ending in degeneration.

Mr. Southwell has further shown (38) that the adult Tetra-

rhynchus unionifactor occurs also in the Shark, Ginglymostoma

concolor. This fish was doctored with Male-fern and castor-oil,

and subsequently fed on pearl-oysters ; but Mr. Southwell does not

claim that the Vetrarhynchi were actually derived from these

pearl-oysters, though he is inclined to think they were. The

same experiment was subsequently repeated [Southwell (41) |

and Tetrarhynchus unionifactor was again found in Ginglymostoma

concolor ; and while Mr. Southwell admits that his results are not

altogether conclusive, it seems highly probable that the infection

was in fact induced by his feeding experiment. There is nothing,

however, to show that the adult tapeworms in Ginglymostoma

were derived from the globular cysts in the oysters; it seems

more probable that they were derived from the Tetrarhynchi

in the oysters’ intestines.

These elaborate experiments are, of course, chiefly of academic

interest, in the absence of proof (1) that the Vetrarhynchi are a

later stage of the globular cysts, and (2) that the latter are

concerned in pearl-formation.

To sum up, then, the gap between (a) the resting scolex

enclosed in its tough fibrous cyst in the connective tissues of the

Pearl-Oyster and strongly suggesting by analogy with other

forms—notably Seurat’s larva—a young Tylocephalum, awaiting

and ready for its final host to devour the tissues which contain it,

and (b) the equally expectant, but much larger, Tetrarhynchus

unionifactor in the wall of the oyster’s intestine, has not yet been

bridged.

I set out below diagrammatically the conclusions of Herdman,

Hornell, Shipley, and Southwell, as to the probable life-history

of these parasites, and also, for purposes of comparison, my own

attempt at an interpretation of the facts.

Diagram illustrating Prof. Herdman’s conclusions as to the

life-history of the parasites.

Species I. Tetrarhynchus unionifactor.

B C

P Globular cysts in

Hornell’s free- liver and connective

Se tissues of Pearl-Oyster

swimming

(=Tylocephalum ludi-

larva. acres) €

A Wir oes This transition is

\ doubted by Shipley.

Sate tap

Bees Larval Teérarhyn-

Sa chus wunionifactor in

pane _* neighbourhood of ali-

sea. mentary canal of Pearl-

\ Oyster.

\ /

‘ | /

\ , /

Adult Tetrarhyn- Immature Tetra-

chus unionifactor im rhynchus in Balistes.

alimentary canal of J _ Thisstage is rejected by

Rhinoptera javanica Shipley, and is regarded

and Ginglymostoma as collateral or acci-

concolor. dental by Southwell.

Species IT. ‘ Tetrarhynchus sp.”

Smaller globular cysts

in connective tissues of

Pearl-Oyster (= Tylo-

cephalum minus).

Southwell considers

this the same species

as “©” above. XK

a Possibly the smaller

Tetrarhynchus found

in the intestine of the

Pearl-Oyster, if it is

distinct from 7’. wnio-

nifactor.

Li?)

}

DrAGram illustrating my hypothesis of the relations of the several

forms and stages of Cestode larye described in Herdman’s

Report.

(The letters correspond to those in the above diagrams. Those

marked A’, A’, B’, and J’ are regarded as parallel stages to

A, B, and J.)

Species I. Tetrarhynchus unionifactor.

A D

Larval 9 Vetrarhyn-

Egg, (2) swallowed by yee i

chus unionifactor, in

Pearl-Oyster ; hatch-

: ree : RT and around the ali-

P 2

Ing out in intestine to mentary canal of the

become \ Pearl-Oyster.

x ¥

Adult Tetrarhynchus

unionifactor in ali-

mentary canal of Rhi-

noptera javaniea and

Ginglymostoma con-

color.

Species LI. Z'ylocephalium ludificans.

(?)

Hornell’s free-

swimming

J\arva.

ati

A! "sf ake CG

MESO? 2 SS Maa

Egg, (?) hatching in Globular cysts, Tylo-

sea-water to become B , cephalum tudificans,

or (?) swallowed ~by-————_—>-— in liver and connective

Pearl-Oyster to be- , tissues of the Pearl-

come C. \ Oyster,

x ¥

An adult Tyloce-

phalum-like form in

one of the oyster-eat-

ing Elasmobranchs,

probably — Aétobatis

narinari.

Proc. Zoou, Soc,—1912, No. XIX. 19

Dye

290 DR. H. LYSTHR JAMESON ON

Ql = .

Species IIT. Zylocephalwm minus.

B’

(?)

A free-swimming larva,

hke B above, only

smaller.

A’ i

Egg, (?) hatching in , The smaller globular

5 _ / cyst, Lylocephalum

sea-water to become b, Ta a minus, mM connective

or (2) swallowed by tissues of the Pearl-

oyster to become G. ‘ Oyster.

X ¥

An adult Tyloce-

phalum-like form, in

one ot the oyster-eat-

ing Klasmobranchs.

Scolex probably about

‘2 mim. in diameter.

(5) Description OF THE TWO GLOBULAR CEsToDE LaRVa FROM

tHE CEYLON PEARL-OysTER.

I append the following descriptions. They are certainly

incomplete, being based on examination of preserved material

and on the descriptions of previous writers :—

Tylocephalum ludificans, sp.n. (The larger globular larva ; the

supposed pearl-producing worm.) (Pl. XLVI. figs. 98 (type)

& 59.)

The type, fig. 58, is in Slide 94 in Prof. Herdman’s Collection.

Tetrarhynchus unionifactor (young) Herdman. (For other figures

see Herdman’s Report, Part II. Parasites of the Pearl-Oyster,

pl. i. figs. 12 & 13; Part V. Pearl-Production, pl. u. figs. 11, 12,

& 176, pl. iii. figs. 1-5, 6, 7, 8, & 9; also Part V. p. 21.)

Globular Cestode larvee, with rostrum or myzorhynchus (Linton)

retractile within a denticulated collar. Form more elongated

when liberated from capsule; length 0°5 to 1-5 mm, Average

diameter of seven specimens sectioned on Prof. Herdman’s slides

and examined by the writer, 0°78 mm.

THE CEYLON PEARL-OYSTER. 291

Myzorhynchus uniformly muscular, without obvious division

into muscular tracts; retractile within an annular collar; in

section it may appear either conical, lenticular, or flattened,

concave and sucker-like ; protrudes as a conical papilla when in

locomotion, This anterior muscular region, including the collar,

is about one-third of the total length of the larva when extended.

The whole myzorhynchus can be protruded, the collar then

forming an annulus around it.

Collar or cephalic sheath muscular with denticulated cuticle,

the denticles tricuspid. (Herdman’s Report, Part II. (Shipley &

Hornell, Parasites of the Pearl-Oyster) pl. i. figs. 10, 11, & 14,

pl. ii. fig. 18; also Pt. V. (Pearl- Production) pl. iii. fig. 9.)

The denticles measure from 3 p to 5 u in diameter.

Hinder part of the larva centrally parenchymatous, the

parenchyma containing the calcareous corpuscles characteristic

of Cestode larvee, peripherally more muscular.

The hinder part of the body is covered by a thick, radially

marked epicuticle, permeated by numerous closely-set tubuli,

and suggesting on superficial examination a coat of cilia. This

epicuticle varies in thickness but is generally about 0:03 mm.

thick, and the true cuticle lies under it.

This form is distinguished from the next described worm by its

larger size (Herdman gives the size as about six times that of the

smaller form), the undivided musculature of the myzorhynchus,

and the wider and more open character of the collar-sheath of the

myzorhynchus in the resting-stage.

Habit. Resting in spherical fibrous cysts, derived from the

connective tissue of the host, in the Ceylon Pearl-Oyster,

Margaritifera vulgaris. Most frequent in the visceral mass,

notably the liver.

Habitat. Gulf of Manaar (Herdman & Hornell). Trincomalee

( Willey).

The following is a description of a worm which I regard as

in all probability the adult of this larva. The single specimen

was obtained from the spiral intestine of Aétobatis narinari, by

Mr. Hornell, on 4th January 1905, and had apparently been

overlooked by Mr. Shipley among some duplicate specimens of

Kystocephalus translucens, along with which I found it when

examining Dr. Shipley’s material. After it had been cleared and

examined as a transparent object, Dr. Shipley very kindly allowed

me to have sections cut from it to compare with those of the

larva in the pearl-oyster.

(2) Adult of Z'ylocephalum ludificans, sp.n. (Pls. XLVI. &

XLVII. figs. 60-64.) Length 12mm. Head 0:6 mm. long by

0°5 mm. broad ; pyriform, slightly broader in front than behind ;

transition from head to neck not very sharply defined. The

myzorhynchus in this specimen is retracted within its sheath, as

is usually the case with the larva in the pearl-oyster; it is about

‘3 mm. in diameter. Around the head are four marginal suckers

about ‘125 mm. in diameter. Proglottides about 140 in number,

increasing but little in breadth from before backwards; they

19*

292 DR. WH. LYSTER JAMESON ON

begin to increase notably in length from about the 85th back-

wards. The largest hindmost segments are about *5 mm. long,

and slightly longer than broad (fig. 64). The armature of the

collar @ (in figs. 61-63) is similar to that of the larva. In section

(fig. 62) the myzorhynchus is seen to be retracted in such a way

that its anterior surface is thrown into folds, thus resembling the

condition of the larva shown in fig. 59. The only point in which

the head of this worm appears to differ from the larva in the

pearl-oyster is im the presence of the four marginal suckers, which

may well be a feature first acquired in the final host.

Tylocephalum minus, sp. n. (Plate XXXIIT. fig. 2.) (The

smaller globular larva, which Prof. Herdman thinks may

also be concerned in pearl-formation.)

(Tetrarhynchus sp., Herdman.)

For other figures see Herdman’s Report (Pearl Production),

Part V. pl. ii. figs. 1-3, 174, 18-22; also text-figure 3, p. 19.

Diameter of resting parasite in cyst from 0-07 to 0-2 mm.

Average diameter of 40 examples shown on Prof. Herdman’s

slides and measured by the present writer, 0-14 mm. Body sub-

globular, consisting, as in 7’. ludificans, of an anterior muscular

and a posterior parenchymatous part, the anterior muscular

portion (myzorhynchus) consisting of a conical papilla in a ecup-

or flask-shaped depression formed by the surrounding muscular

collar or sheath. Asa rule, in preserved specimens, the opening

of this depression seems relatively narrower, and the papilla more

conical and less flattened than in the previous species. The

musculature of the myzorhynchus shows, in some examples,

a tendency to break up into four longitudinal tracts. In

young examples the myzorhynchus may be barely differentiated.

Cuticular spines are present on the collar, but they are smaller

and relatively finer than in 7’. ludificans. The epicuticle is about

‘Ol mm. thick.

This form is distinguished from 7" ludificans by its smaller

size and finer armature of the collar, and by the tendency of the

myzorhynebus musculature to break up into four strands. It is

regarded by Southwell as the same species as 7’. ludificans.

Habit. Resting in spherical fibrous cysts in the connective

tissues of Margaritifera vulgaris, occurring in the visceral mass,

mantle, gills, ete.

Habitat. Gulf of Manaar.

Prof. Herdman, while he regards the form here named

T. ludificans as the pearl-producer ‘par excellence, considers that

the present species too ‘‘may occasionally form the nuclei of

pearls” (Report V. p. 22).

Particulars of the structure of both these forms are given on

pp. 79-82 of Part II. of Prof. Herdman’s Report.

THE CEYLON PEARL-OYSTER. 29

(6) Oxservations oN tHE SrRucTURE oF THE FrBrous Cysts

SURROUNDING THE CrsToDE LArvV® IN J/4RIGARTIFERA

VULGARIS.

T may here add a few notes on the structure of the investing

eysts of Tylocephalum ludificans and 7’. ninus, based upon my

examination of Professor Herdman’s slides and Dr. Kelaart’s

material,

In the earliest stages the cyst may be scarcely differentiated

from the surrounding tissue, and about 0°01 or 0-02 mm. thick.

This condition I find specially associated with a little-developed

phase of the smaller larva 7’. minus, measuring 0:08 mm. in

diameter, and resembling B in figure 4 on p. 21 of Part V. of

Herdman’s Report. In such thin cysts the nuclei of the fibres

are distinct, though hardly more so than those of the general

connective tissue of the oyster. One or two examples at this

and later stages were found in the muscular tissue without any

surrounding cysts at all, beyond a little of the interstitial tissue

of the muscle-bundles (PI. XXXIIT. fig. 2). As the fibrous

capsule becomes thicker the nuclei appear at first to become more

abundant, and this may well be associated with the growth and

multiplication of the fibres. As a rule, the thicker cysts (0-2 mm.

thick and over) seem to be less densely nucleated, and may even

show very few nuclei, especially when they become highly areolar

and oedematous. The outer part of the cyst is usually ordinary

areolar connective tissue, with branched and anastomosing fibres

passing over, often quite imperceptibly, into the general con-

nective tissue of the body, such as occurs between the tubules

of the liver. The cyst is, however, typically lined with several

layers of more regular parallel fibres, with abundant nuclei. In

some cases the fibres seem to coalesce to form a dense almost

gristly substance, without obvious nuclei.

The nuclei of the fibres are long and narrow, and are situated

on their outside edges.

In some cases the cyst appears to be entirely without nuclei,

and in such cases the fibres are often very thick, measuring as

much as 10 or 15 yx in diameter.

There is never, so far as I can see, any trace of a lining

epithelium, though the cyst may be lined with a layer of granules,

possibly derived from the Cestode or from the leucoey tes of the

blood.

In certain cases large dark-staining bodies were seen in the

aveole of the fibrous connective tissue of the cyst, which suggested

parasitic Sporozoa, but the preparations did not allow of detailed

examination.

A typical section of the cyst of the smaller parasite,

Tylocephalum minus, is shown on Pl. XX XIII. fig. 3.

This condition of things is very different to that which is found

in the case of the pearl- inducing Trematode of Mytilus described

in my paper (25). Here the parasite is surrounded by an

294 DR. H. LYSTER JAMESON ON

epidermal sac (d. ¢. pl. xv. fig. 5) of the same nature as the outer

shell-secreting epidermis. In such a case it is easy to understand

how the parasite, when it dies, hecomes encased in a pearl, laid

down, layer upon layer, by this epithelium *.

(7) MATERIALS AVAILABLE FOR THE PRESENT INVESTIGATIONS.

I have throughout these investigations been seriously handi-

capped by the extreme difficulty of obtaining material. Many of

the points which remain obscure could probably be cleared up if

I could obtain properly preserved specimens of pearl-bearing

oysters from the Gulf of Manaar. Unfortunately, I have been

quite unable to obtain these.

I endeavoured to do so through the Ceylon Company of Pearl

Fishers, Ltd., and Mr. Southwell, but without success, Mr. South-

well replying that there were no oysters on the banks and that

his own preserved material was finished. The Company, however,

kindly forwarded to me a suggestion made by Mr. Southwell to

the following effect :—

“As it is probable Dr. Jameson requires Ceylon pearls

(with the particular parasite giving rise to same), I would

suggest that in order to ensure that the pearls are from

Ceylon that they be bought here. I shall be glad to pur-

chase pearls for Dr. Jameson, if he will give me some idea

what to get and how much to spend.”

I gladly availed myself of this offer, and asked Mr. Southwell

to spend five pounds in the purchase of ‘“‘cyst-pearls.” For this

sum he procured from a local jeweller a parcel of 21 small “ fine”

pearls, which I received in February 1911.

* I must here incidentally refer to a quite erroneous interpretation which was

placed upon the expression of my views as to the origin of the sac in Mytilus in my

paper above referred to.

My account of the development of this sac on p. 149 appears to have been taken

by Herdman and by Boutan (3 & 4) to imply that I thought the sac arose from the

mesoblastic connective-tissue elements of the mantle. As I explained in a letter to

Prof. Herdman, which he was good enough to publish, as showing my views, on

p. 9 of Part V. of his Report, I never had any doubt that the sac was a true

epidermis. What I wished in my paper to emphasize was that mm Mytilus it

appeared to arise independently of, and not in continuity with, the outer epidermal

epithelium, perhaps from in-wandering epidermal cells, perhaps from more deeply

seated elements of epiblastic origin, some of which (e.g. certain flask-shaped glands in

Margaritifera, see P|. XI. fig. 33) appear to project below the basement-membrane.

Had I dreamed that I should have been suspected of attempting to promulgate

heretical views on the doctrine of the immutability of the three primary germinal

layers, I would have been more cautious in the choice of my phrases. But even if

my wording in that paper was unintentionally somewhat ambiguous, my 7éswmé of

my work in ‘ Nature’ (26) should have cleared away any misconception, for in that

paper I definitely stated (p. 280) that “a true pearl is laid down in a closed sac of

the shell-secreting epithelium, embedded in the subepidermal tissue of the mantle

and completely cut off from the outer epithelium itself... .. Such a sac, with its

contained pearl, may be compared to a human atheroma cyst.” I have not yet

reached the stage at which I can add to what I said in 1902 about the actual mode

of origin of the epidermal sac in Mytilus, but I hope before long to be able to

contribute some more facts on the subject.

THE CEYLON PEARL-OYSTER. 295

I then tried the Colombo Museum in the hope of getting some

oysters with pearls in situ, but Dr. Pearson had no preserved

material to spare. I tried to obtain material from Madras from

Mr. Hornell, but he wrote me, in January 1911, that his own

material was exhausted, and that he would not be able to obtain

any more till the next inspection, a year later. However,

H.H. the Jam Saheb of Nawanagar most kindly sent me some

preserved specimens of this species with pearls in situ from the

Gulf of Kutch, and I hope, in a later publication, to be able to

put forward some observations on the actual process of pearl-

production, based on these.

The following material was available for these investiga-

tions :—

(i.) Twenty-one pearls bought in Ceylon. It is, of course,

possible, though not probable, that some of these originally came

from elsewhere, e.g. the Persian Gulf wid Bombay, but they, or

at least the great majority of them, were certainly derived

from MW. vulgaris, the pearls of which have a characteristic

colour and lustre quite different from that of the pearls found in

M. margaritifera and M. maxima.

They were small “ fine pearls,” mostly spherical, a few oval or

slightly lenticular. One wasa brown pearl formed in the mantle-

margin from the prismatic substance. They were all decalcitied

and examined whole, cleared in oil of cloves, and drawn. They

were then sectioned (except in the case of three examples which

were preserved whole). Their nuclei were in no cases Cestodes ;

they usually contained a cavity with a few granules surrounded

by spheevocrystal-like matter, allied to or identical with the

“ repair-substances ” described below. In several cases, however,

the actual nucleus was a grain of sand. These specimens are

preserved as preparations XL, XLII, XLIII, XLIV, XLV,

Oe Vb, at VEEL, Jal DV Any,.A. LEV. By V dL Vp,

iN ey, ives ULI a, Lives, Livery, LEV s, and) LEV «

(Pls. XLI.-XLITI. figs. 35-45 and Pls. XLV., XLVI. figs. 50-57).

(ii.) Dr. Kelaart’s Material in the British Museum.

In 1901, when JI was investigating the origin of pearls in

Mytilus, Mr. EK. A. Smith, I.8.0., allowed me to examine five old

specimens of the Ceylon Pearl-Oyster from Dr. Kelaart’s

collections in the British Museum. Mr. Smith very kindly

allowed me to make further use of some of this material for the

present investigations.

The specimens are labelled :—

“] specimen of pearls in ovaria,

3 specimens of pearls in mantle,

1 specimen of ova of Entozoa in liver of Meleagrina

margaritifera.”

The specimen with ‘‘ pearls in ovaria” was a pearl-oyster with

296 DR. H. LYSTER JAMESON ON

a quantity of clustered pearls, mostly of the baroque and seed-

pearl classes, in the visceral mass. The three examples with

‘“‘pearls in mantle” showed pearls, chiefly in the regions of the

levator muscles, though some were in the non-muscular parts of

the visceral body-wall. The specimen with ‘ ova of Entozoa in

liver” is interesting, as showing the Cestode, Tylocephalum

ludificans, in 1ts whitish fibrous pearl-like cysts, which on super-

ficial examination suggest “eggs.” It was in this specimen,

in 1901, prior to Prof. Herdman’s departure for Ceylony that I

first became acquainted with these larve, which Prof. Herdman

identified as the cause of cyst-pearls. The few observations that

I was able to make at that time led me to the conclusion that

there was no evidence that this parasite was concerned in pearl-

formation *, a conclusion that I have, so far, seen no sufticient

cause for modifying.

Owing to their age, the state of preservation of these specimens

was naturally somewhat defective, and for real detailed histological

work upon the pearl-producing tissues they were quite useless.

Many of the pearls in these specimens had fallen out of their

sacs and lay in the bottom of the jar. Some of these, along with

others picked out of the tissue, numbering 22 in all, were

decalcified and examined in oil of cloves, and six of them were

then sectioned and further examined. [Preparations XIV, XV,

EXCV Ly XOXGT co, XOX G, XGXeT (GR IS.Y XOXO VAIS SXOXONG Vane

figs. 16-18 and Pl. XLIV. figs. 46—48).]

A large piece of tissue in the wall of the visceral mass,

measuring about 5x5x4 mm., and containing no less than

16 pearls, was cut out from the specimen with “ pearls in ovary,”

decalcified, stained with borax carmine and indigo carmine, and

sectioned [Preparation XXVIII (Pl. XXXVII. figs. 14, 15)].

These pearls were all of the class which I refer to provisionally

(see below) as muscle-pearls, and were mostly formed around

central cavities.

(111.) Onlabelled Material in the British Museum.

Mr. Smith also allowed me to examine two unlabelled speci-

mens of Margaritifera vulgaris in the British Museum, the

history of which is unknown. One of them was with the example

in spirit now on show in the Museum, and was accompanied by

its shell. It is from this example that preparations XX VII and

XXIX were cut. It contained a large number of muscle-pearls

and what Prof. Herdman calls ‘“caleospherules” in the left

mantle-lobe, and in the region of the adductor muscle on the left

side. The other example had been removed from its shell.

It contained a large number of clustered pearls, of all sizes, in

the right mantle-lobe. The tubes in which the specimens were

preserved contained also a lot of loose pearls which had dropped

out of both these specimens.

* Tn 1902 (25), p. 149, I poimted out that Cestode larvee were not surrounded by

a pearl-sac.

THE CEYLON PEARL-OYSTER. 297

From the first of these specimens the following preparations

were made :—

a) Preparation XX VII, a piece of tissue cut from the border-

land between the mantle and the adductor in the first of the

above specimens. This piece measured about 7x5x4mm., and

contained 36 small pearlsand numerous so-called “ealeospherules.”

The preservation was so bad that no differential staining of

the soft tissues could be obtained, all parts reacted alike to the

stains used, [Preparation XXVII (Pls. XXXVIII., XXXIX.

figs. 19-21, Pl. XL. figs. 24—27.)|

(b) Preparation XXIX was a piece of the mantle of the same

individual near the margin, containing 17 so-called ‘‘ caleospherules,”

one of which is becoming coated over with nacre (Pl. XX XIX.

figs. 22 & 23). The figures were made from the whole object—

tig. 22 representing it as it was before decalcification, fig. 23 after

it had been decalcified ; both as seen when cleared with oil of

cloves. The preparation was then sectioned, but the state of pre-

servation did not allow of the relations of the “ calcospherules ”

to the tisstes being investigated in this instance. A piece of

tissue was also cut from the second of these specimens, decalcified,

examined entire, and sectioned. It contained about 20 small

earls and numerous so-called “ calcospherules.” [Preparation

XXIV .|

Forty pearls, of varying sizes, some lying loose in the bottoms of

the jars containing the specimens, others taken from the tissues,

were decalcified and examined in oil of cloves. Of these, six were

sectioned and further examined. [Preparations XXIII, LXTII

(A, B, & L), and LXVI (4&1) (Pls. XXXIX—-XLI. & XLIV.

figs. 21 a, 28, 31, 32, 49).]

All the pearls from these specimens I refer to the class called

by Herdman “ Muscle-Pearls.”

(iv.) Three Specimens of the Pearl-Oyster collected by Professor

Herdman in 1902.

Prof. Herdman, at the request of the Ceylon Company of

Pearl Fishers, Ltd., very kindly allowed me to examine his

material (see (v.) below) and handed me three specimens of the

Pearl-Oyster, each of which contained a small “ muscle-pearl” at

the point of insertion of one of the levators of the foot. The pieces

of tissue containing these three pearls were cut out, decalcitied,

and sectioned [ Preparations IV, VI, and VII (Pl. XXXYV. fig. 8)}.

Although these specimens had, apparently, been preserved

in formalin, which is not the most satisfactory preservative for

histological purposes, they showed quite a lot of histological

detail, and enabled me to form some idea of the mode of origin of

muscle-pearls, and of the curious cyst-like bodies which precede

them.

(v.) Professor Herdman’s Slides.

As stated above, Prof. Herdman very kindly allowed me

to examine his slides, which he sent to me a few at a time.

298 DR. H. LYSTER JAMESON ON

Most of these slides were preparations showing the parasitic

Cestodes in the tissues of the oyster, but there were also a number

of preparations of pearls, sectioned im situ in the tissues, showing

in all about 25 pearls, 21 of which showed their nuclei more or

less distinctly. The nuclei of these pearls were of very different

characters, but in no case could I identify a Cestode larva in the

centre of a pearl.

(vi.) Three Specimens from the Persian Gulf.

In September 1903, Mr. J. Calcott Gaskin, Assistant Political

Agent at Bahrein, Persian Gulf, sent me 32 specimens of

Margaritifera vulgaris from Bahrein preserved in alcohol. Out

of 20 of these that I opened, 3 contained pearls, which, from their

position, could obviously be classified as ‘‘ cyst-pearls ” (Herdman).

Two contained a single pearl each, that in the first beg about

2°5 mm. in diameter, situate in the left mantle-lobe, above the

anterior end of the attachment of the gills [ Preparation LXIV 4],

that in the second {Preparation LXIV B] being about 1 mm. in

diameter and situate in the body-wall over the stomach. The

third specimen had two small pearls, about 2 mm. apart, in the

wall of the visceral mass, away from all muscle-impressions

|LXIVc}. All these pearls were decalcified in situ in the tissues

and sectioned. Their centres are described below.

In addition to the above the following pearls were decalcified

and examined. More detailed particulars are given under the

descriptions of the centres of individual pearls, given below :—

(vu.) Dry unlabelled pearls, probably from Ceylon, in the

British Museum, three examples were decalcified.

(viii.) Mixed lot of pearls; given to me by Mr. Max Mayer,

mostly from Margaritifera vulgaris ; 115 were decalcified, of these

8 were sectioned.

(ix.) A collection of pearls from the last Ceylon pearl fishery,

given to me by Mr. EH. Hopkins. Fifteen were decalcified,

and four of these were sectioned.

x.) A collection of pearl-oysters, with pearls im setw, from the

Gulf of Kutch; these are referred to above. Up to the time of

writing, 18 pearls from these specimens have been decalcified,

and 13 of them sectioned.

(xi.) Two pearls from Margaritifera vulgaris, from the

Mediterranean, given to me by Professor Raphael Dubois.

(xii.) Five pearls from Margaritifera vulgaris, from New

Caledonia, given to me by Professor L. G. Seurat.

(xii.) A pearl from Margaritifera vulg Sr, from Madagascar,

gtven to me by Professor Seurat.

(xiv.) Two pearls from Margaritifera vulgaris, from Papua,

from the Imperial Institute.

THE CEYLON PEARL-OYSTER. 299

(xv.) Twenty pearls from Placuna placenta, from Lake Tam-

palakamam, Ceylon, from the Imperial Institute.

(xvi.) About a dozen pearls from Murgaritifera margaritifera

var. cumingii, from the Gambier Archipelago.

In the course of these observations, apart from studies on the

structure and formation of pearls in other forms, 356 pearls

derived, with perhaps a few exceptions, from Margaritifera

vulgaris, chiefly from Ceylon, have been decalcified and examined,

175 of these having been studied in sections.

(8) Mernops.

For decalcification, whether the pearls were free or in situ,

preserved in alcohol or dry, I found alcohol of about 50 per cent.

strength, to which a few drops of nitric acid had been added, was

the best. Of course, a preserved pearl never decalcifies as well as

a fresh one. In the case of dry pearls, where only the nucleus

and central parts are required for examination, it is often

an advantage, during decalcification, to strip off the outer layers

of conchyolin, thus facilitating the penetration of the reagent.

Bubbles of carbon dioxide, generated in the process of decalci-

fication between the conchyolin-layers, cause a great deal of

trouble, especially in old dry pearls, where the conchyolin seems to

be particularly leathery and impermeable. Such bubbles often

greatly distort the normal structure of the pearl, as seen in section,

by tearing the conchyolin-layers apart and causing great spaces

betweenthem. ‘The most suitablereagent for expelling the bubbles

is absolute alcohol, in which the decalcified pearl is placed for

a few days. In some cases, however, it was necessary to extract

the gas under an air-pump.

For staining decalcified pearls hematoxylin was used, also

borax carmine. Sections of the pearls in the tissues were

stained sometimes with borax carmine and picro-indigo-carmine*,

sometimes with hematoxylin and eosin or orange, occasionally

with other reagents.

(9) SrRUCTURE OF THE SHELL-SuBSTANCES.

The shell of Margaritifera consists of the following parts :—

(i.) the outermost layer or so-called Periostracum ;

(ii.) the prismatic layer, forming with (1) the ‘“ back” of the

shell, the fragile, dark-coloured “lip,” and the lappet-

like processes of the margin ;

* Borax carmine (Grenacher’s) in bulk :—

Picro-indigo-carmine as under, on the slide.

A. Saturated solution of picric acid in 90 per cent. alcohol.

b. Saturated solution of indigo-carmine (Grubler) in 70 per cent.

alcohol.

1 part of A, 2 parts of 5, 6 parts of 70 per cent. alcohol.

300 DR. H. LYSTER JAMESON ON

(iu.) the Nacre or Mother-of-Pearl, forming the lining and

the bulk of the shell ;

(iv.) the Hypostracum, the substance to which the muscles

are attached by a specialised epithelium ;

(v.) the Hinge Ligament.

The mass of the shell is further divisible chemically and

microscopically into an albuminoid substance called ‘ conchyolin ”

and erystallme carbonate of lime deposited therein. Romer’s

careful observations (32) have shown beyond a doubt that there is

a sharp separation between these two substances: the conchyolin

forming an alveolar framework, in the chambers of which the

salts are deposited ; the structure of the calcium carbonate being

crystalline, its form being determined by that of the spaces in

which it is deposited.

The ratio of conchyolin to calcareous salts differs in different

parts of the shell. Thus Rémer (32) has found in Margaritana,

the fresh-water pearl-mussel, that the organic substance con-

stitutes 1:47 per cent. by weight of the prismatic substance, but

only *64 per cent. of the nacre. This 1s most interesting as

giving support to the theory of the present writer, enunciated

below, that the different structures of the different forms of

shell-building substances, normal and pathological, are in part

a function of the proportions in which these two constituents are

secreted by the tissues of the mollusc.

To turn now to the details of the structure of the several

constituents of the shell.

(1.) The Periostracum.

The origin of the Pertostracum can best be understood if we

consider first those forms which live in fresh or estuarine water,

or are otherwise subjected to conditions which render necessary

a thick cuticle-like layer to defend them from the erosive action

of organic acids derived from decomposing animal and vegetable

ranentor (e.g. the Unionide and J/ytilus). The periostracum

in such cases has been described fully by several authors,

e.g. Biedermann (1), Moynier de Villepoix (28), Tullberg (47),

Khrenbaum (9), Felix Muller (29), Stempell (44), List (27 5),

etc.

In these cases the periostracum 1s composed of two constituents.

The outermost layer is probably formed as a true cuticle directly

by transformation or cuticularisation of the outer surfaces of the

cells of a specialised epithelium in the inner (axial) face of a deep

groove which runs along the mantle-margin, and which has been

called by Moynier de Villepoix (28, p. 18) the ‘“‘fente marginale.”

This mar ginal eroove divides the mantle-margin into an inner

and an outer lobe, the former being piemented and sensory, the

latter being a part of the shell- -secreting apparatus. This outer

THE CEYLON PEARL-OYSTER, 301

layer of the periostracum is closely adherent to the specialised

epithelium, so that secondary thickening, if it takes place (and

my own observations on Mytilus and Modiola lead me to think

that it does so to some extent, a view which is held also by

List, 27 6, p. 55), differs from that of all other parts of the shell

(including the inner layers of the periostracum) in that it

is secreted from outside or centrifugally with respect to the

body of the animal and the shell, instead of from imside or

centripetally. It would, indeed, seem as though there were

morphological grounds for restricting the name periostracum to

this particular lay er of the outer cuticle-like substance, or, failing

that, for introducing a term which would separate it more shar ply

from the more bulky inner layers. The structural distinctions of

this layer are well shown by Romer (32) fig. 25 ( Margaritana), by

Moynier (28) fig. 50 (Mytilus), by Tullberg (47) Taf. iv. fig. 3,

fig. 4d (Mytilus), and by List (27 6) in the “My tilidee generally.

The greater part of the mi hes rs however, 1s laid down

centripetally, layer upon layer, by the epithelium on the outer side

of the marginal groove (7.¢e. on the inner side of the outer of

the two lobes-of the mantle-mar ein, Tullberg (47) p. 27). It is

stratified, and in Mytilus contains, near its outer limit, a charac-

teristic layer of largealveol. The stratifications of this substance

have been shown by Romer to correspond to layers of minute

alveoli (32, fig. 25).

Internally the inner layers of the periostracum pass over into

the conchyolin framework of the prismatic layer.

In some forms, e. g. Anodonta, the distinction between the two

constituents of the periostracum are emphasized by the outer

layer being much greater in area than the inner ones, and being

thrown into folds upon which the ner layers le unconformably.

The Mother-of-Pearl Oysters (together with such forms as

Ostrea and Pecten) differ from the types to which the above

description refers in the fineness of the periostracum and in having

much more freely retractile mantle-margins. In sections of the

decalcified shells of the Mother-of-Pearl Oysters it is diftieult—

indeed, I might say impossible—to differentiate the periostracum

from the outer layer of the organic basis of the prismatic sub-

stance (text-fig. 35, p. 302; see also Pl. XXXIV. fig. 6) ew Hiéne

marginal gr owth proceeds by a series of steps and retreats, the free

mantle-margin being retractile to the edge of the nacre, and being

so retracted when the shell closes. ie after a nee process

of the lip has been formed, the mantle-margin is withdrawn,

and forms a fresh attachment on the inner surface of the last-

formed lip, from whicha fresh lip is produced. So the periostracum

of lip no. 2 is attached to and apparently a direct continuation of

the inner surface of the conchyolin of the prismatic layer of lip

no. 1, and constitutes the outer layer of the conchyolin of the

prismatic layer of lip no, 2. But Herdman has shown that,

difficult as it is to distinguish a separate periostracum in sections

of the shell, such a layer, of extreme delicacy, does exist at the

302 DR. H. LYSTER JAMESON ON

margin, and arises in the normal manner in a marginal groove

ae Report, Part II. Anatomy of Pearl- Oyster, plate vill.

fig.

Text-fig. 35.

eTTTTATTT

mu i ial oe i ee

iii

Margaritifera vulgaris, Persian Gulf. Section through the lip of the shell, after

decalcification, showing the successive lappet- like processes of the prismatic

substance (J., 7.’, 1.’”, 1. my pr., prismatic substance ; Sér., stratification of same.

A, A’, A", AM points marking the successive retreats of the secreting margin,

which takes place when new lappets are to be formed. At these points the

** periostracum ” of the new lappet is continuous with, and indistinguishable

from, the inner conchyolin-layer of the prismatic substance of the last-formed

lappet. Preparation X (X 35), see also Pl. XXXIV. fig. 6

(ii.) The Prismatic Substance.

Reduced to its simplest terms the prismatic layer of the

Mother-of-Pearl shell consists of prisms of calcium carbonate

perpendicular to the surface of the shell, bounded externally and

internally by membranes of conchyolin, which are connected by

vertical membranes forming the septa between the prisms

(text-figs. 35 and 36; see also Pl. XXXIV. figs. 6,6a; Pl. XL.

fig. 29a). Where interruptions in the continuity of the growth

of this layer have occurred, the layer of prisms may be divided

by one or more horizontal sills of conchy olin (text-figs. 35, sér.,

and 38, str.), which break up the prisms into seoments, or which

divide the layer into two or more series of prisms. The individual

ends of the prisms in one layer do not of necessity coincide with

those of the prisms in the next layer, though they frequently do

coincide. The septa between the prisms may also show annular

thickenings, corresponding to zones of constriction around the

prisms (Pl. XXXIV. fig. 6 a, ann.). Romer has shown (82, p. 35)

that the prisms of the pearl-shell (like those of Pinna, described

by Biedermann (1), p. 9) behave between crossed nicols in the

same manner as single crystals. The prisms differ enormously in

size, according to the age of the oyster and the conditions under

which they are secreted.

There is a sharp line of demarcation between the prismatic

and the nacreous layers in Vargaritifera, the innermost layer of

the conchyolin of the former being connected to the outermost

layer of that of the latter by a series of fine connectives of

conchyolin (Pl. XXXIV. fig. 6 a, con.), forming a curious alveolar

layer.

THE CEYLON PEARL-OYSTER. 3038

The prismatic substance is secreted by that part of the epidermis

apposed to the inner surface of the shell which is nearest to the

margin of the mantle, and it is clear from the rapidity with

which the columnar lip and its lappet-like processes are re-

generated when injured and are added to in growing young shells

that the characters of this layer are associated with relatively

rapid secretion. This is significant, in view of the resemblance

of this layer to some of the rapidly secreted repair-substances

which replace the nacre under certain abnormal conditions

(see below).

Roémer’s work (p. 18) gives an interpretation of the nature of

the prisms, which 1 have found most useful in helping me to

interpret my own observations on pearl-formation. According to

this hypothesis, which was suggested to Romer by my illustrious

teacher, Prof. Biitschli, each prism is an incomplete spherocrystal,

the growth of which has been arrested in all directions but one,

viz. the direction from which the new shell-substance is secreted.

Romer says (p. 18) :—

“Dass diese Spharokristalle der einzelnen Prismen so

unvollstiindig ausgebildet sind, rithrt daher, dass gleichzeitig

und dicht nebeneinander die Anfinge der einzelnen Prismen

oder Spirokristalle gebildet wurden, die bald seitlich

aufeinander stiessen und sich so gegenseitig in der weiteren

Ausbildung hemmten; nur an ihren inneren Enden

vermochten sie einseitig weiter zu wachsen ” *,

If I may be allowed to state the proposition in slightly

different terms, the prismatic shell-substance (and, indeed, if my

interpretation of the variations of the nacre, normal and patho-

logical, are correct, the whole of the shell-substance) agrees with

a spherocrystal in that it is composed of crystalline or crystallised

substance which can only grow by the apposition of fresh layers

deposited on a single surface, owing to the matter in solution

only having access to one surface of the crystalline mass. When

this surface is the outer surface of a sphere, a body with more or

less of the characters of a spherocrystal results (e.g. Harting’s

bodies and Pearls); where it is approximately a plane surface, as

in the growth of the Molluscan shell, a structure such as the

* Biitschli in 1908 (6, p. 26) explained his definition of ‘‘spherocrystals,”’ more

especially with reference to the crystal-like prisms of Pinna (and by analogy

of Margaritifera), as follows :—

“ Wie aus den Darlegungen in meinem Werk von 1898 hervorgeht, verstehe

ich unter einen solchen nicht ein Aggregat zentrisch angeordneter Kristallnadeln

oder Einzelkristalle, was zwar die tbliche Anschauung ist, sondern ein

einheitliches Kristallgebilde, in welchem die besonderen feinsten Struktur-

verhiltnisse, die auch den gewéhnlichen Kristallen ihre charakteristichen

Kigenschaften verleihen, nicht entsprechend einer Axe, sondern um ein Zentrum

radiar orientiert sind. Wenn daher der Radius eines solchen Spharokristalls

sehr gross wird, und man ein radiales Stiick desselben, weit entfernt von dem

Zentrum, herausschneidet—und_ so verhalten sich etwa die Pinnaprismen—so

muss dieses Stiick sich natiirlich wie ein gewéhnlicher Kristal verhalten ;

obgleich die von mir gegebene Zuriickfiihrung auf einen Sphirokristall mit

erossem Radius ganz zutreffend ist.”

304 DR. H. LYSTER JAMESON ON

Molluscan shell results, the axes of its constituent elements being

approximately parallel.

(iu.) Zhe Nacre.

This substance, which forms the bulk of the shell, and gives

the shells of the genus Margaritifera their commercial value as

Mother-of-Pearl, and the pearls their beauty, is stratified, and

in it the calcium carbonate is divided into extremely minute

bodies in the organic network. It is secreted by the outer surface

of the mantle and body-wall.

I will not attempt here to review the many writings on the

structure of this layer. I can at present add little to the recent

work of Romer (32), who has studied its structure and that of its

decalcified conchyolin framework very thoroughly. The organic

basis which gives it its form, and which retains its iridescence

after the caleareous salts have been extracted, consists of a series

of parallel lamelle, of extreme fineness, united to one another

at intervals by radial connections, so as to form a series of minute

flat or lenticular chambers, separated by organic walls of extreme

delicacy. The calcium carbonate appears to be enclosed m

these chambers in the form of little polygonal plates or lozenges.

This structure is difficult to observe, owing to the distorting

effect of the decalcification process, which, owing to the evolution

of gas-bubbles, tears some lamellze apart and forces others

tightly together. It becomes much more obvious in some of

the abnormal and pathological varieties of nacre described as

“repair-substance ” below, notably in “ granular repair-nacre.”

I believe that the lustre (not the iridescence) of mother-of-pearl,

and of pearls, is in great measure due to the fact that each of

these tiny plates is a minute, biconvex lens; and that the

extraordinary and indescribable character of the light reflected

from the surface of a fine pearl is in part the cumulative expression

of the action of these myriads of little lenses upon the light

reflected from the surfaces of calcium carbonate and of conehyolin

which underlie them.

(iv.) Hypostracum, or Muscle-Attachment Substance.

This curious substance has not secured all the attention it

deserves. It has been described by several writers under the

names Hypostracum (Thiele, 46), Stabchenschicht (Ff. Miiller,

29), durchsichtige Substanz (Ehrenbaum, 9; Tullberg, 47).

I retain the name hypostracum, as emphasizing the distinct

origin and characters of this layer, and as shorter and more

convenient than ‘‘durchsichtige Substanz.”

This hypostracum is a fine columnar layer forming the surfaces

where the muscles are inserted into the shell (Pl. XXXIV. figs. 5

& 7, hy.; text-figs. 36 & 37).

It is more transparent than the nacre—indeed, the iridescence

and lustre of the muscle-scar is due to the nacre lying below and

THE CEYLON PEARL-OYSTER. 305

shining through this “durchsichtige Substanz,” the substance

itself not possessing the structure to which these optical properties

are due,

It is composed of columnar or fibrocrystalline needles of

carbonate of lime (Stiibchenschicht, Miiller), but shows in

places, in addition to its columnar structure, a distinct strati-

fication parallel to the surface; this is seen also in the basis

which remains on decalcification (Pl. XXXIV. fig. 5), I attri-

bute this stratification to variations in the organic basis, which

are probably independent of the form and structure of the

crystalline needles. Hypostracum only oceurs where the

specialised muscle-attachment epithelium is inserted into the

shell, and, as the muscles move away from the umbonal region

with the growth of the shell, it is quickly covered over by ordinary

nacre which is deposited in the wake of the advancing muscle.

In a section of the shell from the umbo through the adductor

scar the hypostracum layer can be traced across the shell through

the nacre from the scar to the umbo, the thickness of the over-

lying nacre increasing as the umbo is approached. By means of

this hypostracum layer, the wandering of the adductor muscle is

recorded in the shell-substance (text-fig. 36, hy.).

Text-fig. 36.

AANA NU NAAN

UNUU)LLA TOLD PRAY Ua a

Margaritifera maxima Jameson. A young shell or “chicken shell” from Port

Darwin, Northern Territory of Australia (London markets). Section from

the umbo to the shell-margin passing through the middle of the adductor

impression. p7., prismatic layer; A-—B, muscle-scar, covered with “ Hypo-

stracum”; hy., the hypostracum layer, by means of which the migration of

the muscle, from what is now the umbonal region, with the growth of the

shell ean be traced; nae., nacre of the shell-margin, formed external to the

muscle-sear; nac.’, nacre of the thick subumbonal region, deposited internally

to the hypostracum. ‘Two-thirds of natural size.

In the shell figured, a young example of the large white

Australian Mother-of-Pearl shell (17. maxima Jameson), the

hypostracum is 18-20 » thick over the muscle-scar, thinning

out to 10 » and then to 4 or 5 w at the extreme outer edge of

the scar, where the muscle has most recently made attachment.

As this layer is traced backwards towards the umbo, through the

nacre, it is found to get gradually thinner, just as the prismatic

substance (which in this shell is about 1 mm. thick in the region

of the adductor scar, and in the lip of very old examples of the

Same species may be 2 or 3 mm. in thickness) is found to get

thinner towards the umbonal region. These differences are no

doubt associated with the relative ages and sizes of the animal at

Proc. Zoot, Soc.—1912, No. XX. 20

306 DR. H. LYSTER JAMESON ON

the respective periods, and with the relative rapidity of peri-

pheral growth in young and older oysters. Text-fig. 37 shows the

hypostracum of the same example (J/. maxima) enlarged forty

times.

Text-fig. 37.

The inner limit of the adductor scar, in the same shell as that shown in text-fig. 36.

hy., hypostracum; ac., nacre external to same; nac.’, nacre internal to

same. X 40. .

Exactly the same relations occur in J/. vulgaris, a section

through the umbonal side of the adductor scar of which is shown

in Pl. XXXIV. fig. 7. But in this example, an old thick Lingah

shell from the Persian Gulf, 11 which, in all probability, peri-

pheral growth, and consequently the wandering of the muscle,

had ceased, the hypostracum is thicker, measuring 130 w in

thickness.

The hypostracum undergoes but little secondary thickening

compared with the nacre. Hence, in thick massive shells ake

M. maxima, where the newly forming nacre in the umbonal

region and also towards the lip and around the muscle-scar out-

strips the hypostracum in development, the muscle-scar is the

thinnest part of the entire shell, except the extreme edge. This

is well seen in text-fig. 86, in which figure the area between A

and B represents the muscle-impression, but it is even more

obvious in older thicker shells. Some interstratification of

hypostracum and nacre occurs at the borders of the muscle-scars,

where changes in the outline of the muscle ae taken place.

This is figured by Tullberg in 1; ytilus (47, Taf. v. fig. 2). The same

is shown for Margar itifer a vulgaris at hy.', hy”, in Pl. XXXIV.

fig. 7. Felix Miiller (29, Taf. xxix. fig. 13 6) shows the lateral

transition of this substance into nacre. I have observed the

same thing in some of the “ Muscle-Pearls” described below.

When decalcified the hypostracum leaves behind it an organic

basis, which is somewhat different from the conchyolin of the

rest of the shell in its reaction towards stains. This is of interest

in connection with the view generally held that this layer,

unlike the other calcareous parts of the shell, which are probably

THE CEYLON PEARL-OYSTER,. 307

due to simple secretion, arises by a gradual transformation into

shell-substance of the outer regions of the specialised epidermal

cells which underlie it, 7. e. in the same manner as the Crustacean

carapace and the outermost layer of the periostracum. It shows

a well-marked striation perpendicular to the surface, the strix

no doubt corresponding to the outlines of the spaces which were

occupied by the needle-like fibrocrystalline bodies of caleium

carbonate, and also at times indistinct lines parallel to the surface

(Pl. XX XTV, fig. 5). In sections in the plane parallel to the

surface this substance shows an alveolar structure. It sometimes

shows a tendency to break up into segments, corresponding to

the underlying epithelium-cells (Pl. XXXIV. fig. 5, hy.’).

(v.) Hinge-Ligument.

I do not propose to discuss the hinge-ligament here, as it has

not the same direct bearing on the question of pearl-formation

as the above layers, though leathery pearls, composed of this

substance, are sometimes found (e. g. in JZ. maxima in Australia),

(10) Tae SHeLi-sEecrerine Eprra ens.

The ordinary shell-secreting epidermis of I. vulgaris, so far

as I have been able to study it in the unsatisfactory material

available, consists of columnar or tesselated cells (Pl. XXXV.

a0;.6p:, igs 9, 0,ep.;.Pl. XXXVL fig) 11, oep.;, Pl XU.

lig. 33, ep.), brick-shaped or palisade-like, according to the

degree of contraction, in sections perpendicular to the surface,

with a certain and variable number of goblet- and gland-cells.

The nuclei of the epidermal cells are oval or spindle-shaped.

These cells are attached to the subjacent tissues by a basement-

membrane of delicate fibrille which distinctly marks the

boundary between the epidermal epithelium and the subjacent

tissues. Beneath this epidermis is a characteristic granular

parenchyma (Pl. XXXV, figs.8&9; Pl. XXXVI. figs. 10 & 11;

Pl. XXXVIT. fig. 14; Pl. XLI. fig. 33, par.), which contains a

great variety of elements, some being comparable to the

“ Rundzellen” and “ Langer’schen Blasen” described by List

(27) for the Mytilidee, some being dark-staining, apparently

glandular elements that open out between the epithelial cells

(fig. 33, gl.). As observed by List (27), this epithelium and the

underlying tissues are excessively variable in their characters.

Over the surface of the muscle-attachment the epidermis is

different (Pl. XX XIII, figs. 4, 4a; PL XXXV. fig. 8, m-ep.).

Here it consists of columnar cells, usually about 10-12 w long

and 2-4 » broad, which pass over basally without any distinct

dividing-line into the muscle-fibres. Whether the transition

is direct, or whether in fact a connective-tissue junction is

present, cannot be determined from the available preparations

of Margaritifera vulgaris; but in Mytilus edulis there is a

distinct connective-tissue layer (fig. 5, c.t.), the fibres of which,

20*

308 DR. H. LYSTER JAMESON ON

continuous with the bases of the epidermal cells, are attached to

the ends of the muscle-fibres (musc.), which may be produced out

into tails. Tullberg recognised that such a junction was present.

In Margaritifera vulgaris the attachment epithelium-cells may

have one or several tails, probably also of connective-tissue

character, passing over into as many muscle-fibres.

Distally these epidermal cells broaden out somewhat, ending

in a clean-cut surface, which may be represented in section by

a clear zone forming a slightly acute angle with the sides. The

nuclei, which are oval and about 3-4 p long, are situated in

the middle of their length.

It is possible to make out, in some cases, a striation of fhiene

cells in the direction of their ‘long AXES. Occasionally the distal

surface is raised into processes and papille, but this may well

be a result of imperfect fixation of the tissues.

These cells stand out as stiff, independent, almost bristle-like

entities, and are probably hard and tendinous in character.

They are frequently preserved in old preparations in which

all traces of the structure of the ordinary epithelia have

disappeared. In some cases they seem to have been drawn out

in the fixing process; thus the longest cell shown in Pl. XX XIII.

fig. 4a measured 26». It seems possible that in the shrinkage

consequent upon fixation the majority of the elements here

had broken away from the shell, but that this particular cell

had remained attached and was consequently fixed in a state

of extension. This figure shows that the connective-tissue

elements extend up between the bases of these cells.

This epidermis is very closely adherent to the specialised shell-

layer (hypostracum) to which it is attached, and the connection

seems to be between the cells and the organic basis of the shell.

Thus, in decalcifying a piece of the shell of Mytilus with the

adductor muscle attached, the hypostracum remained adherent

to the epithelium and tore away from the rest of the shell

(Pl. XXXIV. fig. 5).

(11) SHELL-SECRETION.

I will not attempt to survey the writings of previous inves-

tigators on this subject. This has been ably done by Stempell

(45), whose review contains a full and lucid discussion of the

question.

The general trend of opinion now seems to favour the theory

dating back to Reaumer, 1709 (31), and held by Tullberg,

Ehrenbaum, Moynier de Villepoix, and the majority of recent

French and German investigators, that the shell (except the

outermost layer of the periostracum and the hypostracum) is

formed from a fluid secretion, rather than the theory specially

associated with Huxley’s name (24) that the shell is derived

from a succession of fully developed skins or cuticles, shed as

membranes by the underlying epidermis.

THE CEYLON PEARL-OYSTER. 309

' The outermost layer of the periostracum (which in Margariti-

Jera is a negligible quantity) and the hypostracum probably

arise by direct transformation of the outermost portions of

specialised epidermal cells, and on this account it may prove

necessary to draw a sharper morphological distinction between

them and the rest of the shell than has hitherto been done *.

The prismatic layer and the nacre, together with the inner

layers of the periostracum, more probably arise as a secretion

which first hardens into a membrane 7d situ, and then forms

the delicate skin which Huxley observed between the mantle

and the shell in the freshwater mussels.

It would appear that the lime-salts and albuminous fluid

which hardens to form the conchyolin are independent of each

other, and may be secreted in varying proportions. Where

these two constituents are secreted under circumstances which

inhibit the control of the shell-secreting epidermis, or where

the secretion takes place so copiously and rapidly that the

epidermis is unable to regulate the deposition (as in the patho-

logical cases described below), lime-salts are precipitated in a

columnar form, much as in Harting’s bodies, and, concurrently

with this, the albuminous fluid is transformed into an insoluble

substance resembling conchyolin. The process of shell-secretion

at the rapidly growing edge of the shell resulting in the

formation of the prismatic layer—which in Margaritifera

vulgaris measures as much as 1 mm. or more in thickness—is

probably in some degree analogous to the process of secretion of

repair-substance, the epithelium exercising comparatively little

control over the arrangement of the elements.

But in the case of the nacre it is different. Here the epithe-

lium seems to exert a definite and very strict selective influence

resulting in the finely stratified and chambered structure which

ean, I think, best be interpreted as arising from rhythmically

intermittent secretory action on the part of the controiling

epidermis. Any disturbance of the normal rhythm of this

secretion, ¢. g. the stimulation of an intrusive particle between

shell and epidermis, results in the formation of the irregular

substances described below, such as granular repair-nacre, the

several varieties of columnar repair-substance, or the amorphous

non-calcified substance.

It would thus seem as though the structure of the shell-

substance, and its variations, normal and pathological, could

be expressed in terms of the proportions of lime-salts and organic

* The difference between the outermost layer of the periostracum and the hypo-

stracum on the one hand, and the remainder of the shell on the other, the former parts

arising by direct cell-transformation or cuticularisation of cell-protoplasm, the latter

as a secretion poured out by the cells, suggests a line of inquiry that might yield

interesting results. Can these two constituents of the shell be separated morpho-

logically and phylogenetically, and, if so, can the former be regarded as in any sense

homologous with the cuticular exoskeleton of an ancestor common to Mollusca and

Arthropoda, the latter being a subsequent addition peculiar to the Mollusea.

associated with their more sedentary modes of life, which has now, for all practical

purposes, replaced the more strictly cuticular element as an exoskeleton ?

310 DR. H. LYSTER JAMESON ON

salts secreted and of the periodicity of the secretion as determined

by the control, or loss of control, of the secreting epidermis,

Tn fact, if my interpretation is correct, the processes involved

in the building of the shell are the usual chemico-physical ones

which govern crystallisation in colloidal media* controlled and

limited by the time-factor which is a function of the activity

of the living cells.

It is less easy to imagine the conditions which determine

the transformation of the fluid albuminous secretion mto the

leathery conchyolin. One is naturally tempted to postulate

a chemical transformation asa direct or indirect result of the

action of nascent CaCO,, as in the case of the calcoglobin in

Harting’s bodies (12); but the formation of this substance

apart from the lime-salts, e.g. in the inner layers of the perio-

stracum and in amorphous repair-substance, and in the case

of shells grown in lime-free media (Moynier de Villepoix, 28,

p- 122), seems to negative this; and it may well be that this

change to an insoluble albuminoid is directly brought about

by the action of the secreting cells themselves, or follows from

the chemical composition of the secretion as shed.

(12) ABNorMAL AND ParnoLocicAL PHAsEs OF THE

SHELL-SUBSTANCE,

For a study of the beginnings of Ceylon pearls, a consideration

of the variations in the shell-substance, when it is secreted

under abnormal conditions, either on the surface of the shell

or of a growing pearl, is of importance.

Where the normal rhythm of the process of shell-secretion 1s

interrupted, e.g. by injury to the shell, or the intrusion between

the epithelium and the nacre of a foreign particle or by other

disturbances less easy to explain, certain irregularities in the

process of secretion occur, resulting in an altered product.

Tn the simplest case such a disturbance results in a modification

producing a granular appearance of the conchyolin-layers of

the nacre. This modified substance I propose to call “ granular

vepair-nacre.” In sections made through this substance, after

decalcification, the normal stratification is obscured by a highly

granular appearance which seems to be due to an_ infinite

number of connections between the successive conchyolin-layers

resulting in a distinctly alveolar membrane. This is shown

in text-fig. 38 (rep.nac.), which is taken from an artificial

“Dlister ” produced by the writer in Margaritifera margaritifera

after the “ Linneus” method, in British New Guinea in 1899.

The foreign body was inserted near the mantle-margin, and the

mantle secreted first a double layer of the prismatic subtance,

* Biedermann (2), p. 171, recognises that the structure of the shell is essentially

reducible to crystallisation processes, the influence of the cells being limited to the

composition of the fluid, and perhaps the orientation of the primary centres of

crystallisation, But I would add to these influences the periodicity of their

action. :

THE CEYLON PEARL-OYSTER, 311

and then nacre, which, at places, showed the characters of

“ granular repair-nacre.”

Text-tig. 38.

Margaritifera margaritifera Linneus (Black-lipped Mother-of-Pearl Oyster).

Part of an artificially produced blister. pr., prismatic layer ; Sér., horizontal

dividing membrane of conchyolin in same; nac., nacre; rep.nac., granular

repair-nacre. (Preparation VIII.)

Pl. XL. fig. 29, from the “repair-membrane” formed by

M. vulgaris over a hole in the shell (umbonal region), shows the

same substance at ér.’ passing over on the one hand into columnar

repair-substance, on the other into nacre. The same substance

is seen at ér.

Pl. XLI. fig. 30 (rep.nac.) shows the same substance, secreted

at the point of junction of two pearls (from one of Dr. Kelaart’s

specimens of Jf, vulgaris). Here it was secreted as a result

of disturbances following upon the fusion of the two pearls and

the absorption or calcification of the intervening tissues. The

granular repair-nacre in the preparation shows in places a

distinctly columnar structure, indicating a transition to the

columnar repair-substance ; such a transition is still more obvious

in Pl. XL. fig. 29.

The same granular repair-nacre is seen in Pl. XX XV. fig. 9,

Pl. XX XVII. fig. 15, and Pl. XX XVIII. fig. 18, surrounding the

central cavities of “‘ muscle-pearls,” where unduly rapid secretion

might well be expected, and in Pl. XXXIX. fig. 23, where a

hypostracum-pearl (“ calcospherule,” Herdman) is in process

312 DR. H. LYSTER JAMESON ON

of being coated over with nacre. The same substance is well

shown in Pl. XLI. fig. 35 and Pl. XLITI. fig. 43 (gr.). In the

last-named case it is seen to pass over on the one hand into

nacre, on the other into columnar and amorphous repair-

substances.

The next form of repair-substance is much more variable,

and occurs in several distinct, though intergrading forms. I

propose to call this “ columnar repair-substance,” in view of the

calcium carbonate being crystallised in columns.

Columnar substance resembles, more or less, the prismatic

layer of the shell—indeed, it is probable that Rubbel (33, p. 171)

had a substance analogous to this columnar substance before

him when he stated that the outer epithelium of the mantle

of Margaritana is capable, in repairing the shell, of producing

the prismatic substance which is normally only the product of

the mantle-margin. (In the same way, he treats as “ peri-

ostracum” the non-calcified material secreted under similar

conditions, which I describe below as “‘amorphous repair-

substance ”.) *

In its simplest form columnar repair-substance consists of

parallel needle-like rods of carbonate of lime (which Steinmann

(43), speaking of Harting’s bodies, has aptly called “ fibro-

crystalline’) deposited in an organic conchyolin-matrix, which,

when the calcium carbonate is removed by acids, and a section 1s

cut at right angles to the surface, presents a palisade-like appear-

ance, due to the septa of conchyolin between the calcareous rods

(Pl. XL. fig. 29; Pl. XLI. fig. 30, col.). In horizontal section

this conchyolin has a honeycomb-like structure.

All kinds of variations occur in the coarseness or fineness

of the calcareous elements and the organic framework.

This substance is frequently formed on the surface of the

shell or of a pearl when disturbances arise in the rhythm of shell-

secretion. In Pl. XL. fig. 29 it is seen in the repair-membrane

formed over an injury caused to the shell by a boring parasite.

In PI. XLI. fig. 30 it. is seen (col.) in the angle between the

surfaces of two “pearls which have become oy attached

together.

120 cb fig. 31 shows the same substance developed under

conditions similar to those existing in fig. 30. This figure is

a drawing of a section through the suture between two pearls

which have become secondarily fused together. The pearls

themselves, with the intervening suture, are shown in Pl. XLIV.

fig. 49; the end of the suture, where the curvatures of the two

pearls diverge, in fig. 31. In the entire object, examined in

oil of cloves (fig. 49), the suture was represented by a yellowish-

brown line, the colour being due to the dead remains of the

cellular membrane which originally separated the two pearls.

* While these substances are perhaps not strictly separable aye ede on

chemical and physiological grounds, I think it is well on morphological and patho-

logical grounds to emphasize the distinction.

THE CEYLON PEARL-OYSTER. alte

The membrane consisted of the lining epithelia of the two sacs,

and a layer of parenchymatous tissue between these two

epithelia.

The epithelia, and even the individual cells of the parenchyma,

can be detected in some places (Pl. XLI. fig. 32).

If we try to trace the steps resulting in the condition figured

on Pl. XLIV. fig. 49 and on Pl. XLI. fig. 31 (¢.e. to survey the

story of the formation of a double pearl), we may assume that

they were as follows. As the two neighbouring pearls, each

enclosed in a sac, grew in size, by the addition of fresh layers,

they exerted a pressure on the intervening tissues, resulting in

reduced circulation and consequent malnutrition which began

at the first point of contact and extended outwards. Thus the

contiguous surfaces tended to become flattened (fig. 49), and the

intervening tissue, consisting of the epithelia of the two pearl-

sacs and a small amount of connective-tissue between them,

finally ceased to be functional, died, and was preserved as a

yellow membrane (Pl. XLI. fig. 32). At the periphery of the

area of contact, where the curvatures of the two pearls diverged

and were separated by a wedge-shaped plug of tissue, nacre-

secretion continued longer, the last efforts of the epithelia

being represented by mac. and nac.' in fig. 31. Finally, the

epithelium ceased to control the deposition of its secretion, and,

with the shrinkage of the atrophied tissues a space occurred

on each side between the nacre and the epithelium, into which

an extravasation of organic matter and salts occurred. The

salts precipitated themselves in the form of columns or raphides

with their bases apparently in or on the epithelia, and con-

currently with this precipitation the soluble organic substance

became converted into the conchyolin framework between the

prisms, analogous to the “ calcoglobin” framework of Harting’s

bodies, derived from egg-albumen when calcium carbonate is

precipitated in it. Lastly, the epithelia and intervening con-

nective-tissue died and probably underwent irregular calcification,

breaking away from the still functional tissues and becoming

incorporated in the substance of the pearl. The still functional

tissues now formed a single sac surrounding the two pearls, and

quickly enveloped them both in a common nacreous covering.

Similar processes can be postulated to account for the condition

shown in Pl. XLI. fig. 30. Here, between the curvatures of the

surfaces of the two contiguous pearls, there was a triangular

plug of tissue, which for some time remained attached to the

degenerated membrane which separated the pearls. Its epithe-

lium gave rise before it broke away to granular repair-nacre

(rep.nac.) on the right, where the disturbance was presumably

least, and to a small amount of columnar repair-substance (col.)

on the left. Then it broke away from the degenerated and dead

membrane between the pearls and retreated rapidly, exuding

as it went the albuminous fluid, which, being secreted at a much

greater rate than the lime-salts, was practically devoid of lime

314 DR. H. LYSTER JAMESON ON

and formed coarsely stratified amorphous substance (am.), broken

by cleft-like cavities. Later on, when the retreat of the plug

of tissue was Jess rapid, this aiorphous substance passed over

into columnar substance (é.) and granular repair-nacre (¢.'), and

finally gave place to the nacre (nac.’) of the common investment

of the compound pearl.

The columnar repair-substance varies enormously, and passes

over imperceptibly into ‘“‘amorphous substance” or lime-free

conchyolin, granular repair-nacre, ordinary nacre, and the

prismatic substance of the shell. For example, the repair-

membrane, a part of which is shown in Pl. XL. fig. 29, showed

an immense number of variations from place to place. In some

parts a second layer of amorphous substance was interpolated

between the columnar layers; in others the columnar substance

passed over into a coarsely alveolar substance with irregular

cavities, some of which penetrated into the amorphous substance.

In yet other spots the amorphous substance passed over through

granular repair-substance into nacre.

Columnar substance is frequently stratified, consisting of a

number of consecutive layers. This is seen at col.’ in the repair-

membrane figured at fig. 29. It is also shown in the pseudo-

nucleus of the pearl shown on Pl. XL. fig. 28 and Pl. XLIV.

fig. 49, and in the pearls from the Persian Gulf in Pl. XII.

figs. 33 & 34. In the former of these last-named instances it

occurs immediately around the central cavity, in the latter case

interstratified and intergrading with the nacre.

In Pl. XLIIT. fig. 43 (col.) it is seen passing over on the one

hand into granular repair-nacre, on the other into amorphous

repair-substance. The same stratified columnar substance is well

shown in Pl. XLV. fig. 51, where it forms a curious flaw running

through the substance of a pearl.

Apart from this direct stratification, the columnar repair-

substance may have an internal alveolar structure such as is

shown in Pl. XLIT. figs. 36, 37, & 38. Figs. 40-42 on the same

Plate, taken from the pearl shown in Pl. XLVI. fig. 57 (a brown

pearl composed of prismatic shell-substance), show the transition

from amorphous repair-substance to columnar repair-substance’

(figs. 41, 42, col.), and from the latter to the prismatic layer of

the shell (fig. 42, pr.).

The third variety of repair-substance I call amorphous repair-

substance. In its typical form this substance is seen at am. in

Pl. XL. fig. 29, where it is obviously the result of the first effort

of the molluse to close the injury to the shell, and in Pl. XLI.

fig. 30, where it is the product of a fully functional epithelium,

retreating rapidly and leaving its secretion in itswake. It shows

little or no structure under ordinary magnifications, but is usually

faintly stratified. It may contain cavities, arranged in rows

parallel to the secreting-surface, and with at times also a radial

arrangement. These cavities typically contain carbonate of lime.

Pl. XL. fig. 29 @ (Margaritifera vulgaris, Lingah Shell, Persian

THE CEYLON PEARL-OYSTER, 315

Gulf) shows this substance formed as the first step in the

development of a new layer of prismatic substance to cover over

the tube of the worm Leucodore, which has entered between the

mantle-margin and the shell, as is its wont. Here the mantle-

margin, reacting to the stimulation of the parasite, has retreated

and secreted a new “lip” to exclude it. This lip, like the

normal lip, consists of the prismatic layer of the shell, but

the irregularly secreted first layers of it consist of amorphous

substance, containing alveoli in which a scanty supply of calcium

carbonate was deposited.

The amorphous substance frequently occurs in the centres

and around the central cavities of pearls, where it doubtless

represents the first matter which the mollusc shed into the cavity.

It probably corresponds to the “ Theile des Schalenepidermis ’ si

recognised by von Hessling (18, p. 313) in the nuclei of

pearls, and the “‘ Kern von Chitinsubstanz” referred to by

Pagenstecher (30, p. 502), and perhaps to the ‘“ Gelbbrauner

Substanz” of Rubbel (34, p. 412).

The amorphous substance shows little receptivity to stains.

It passes over sometimes into columnar substance (Pl. XLI.

fig. 30, é.), sometimes into granular repair-substance (Pl. XL.

fig. 29, tr., tr.'; Pl. XLI. fig. 30,¢r.'). It also sometimes inter-

grades with a substance resembling the prismatic layer (Pl. XLII.

fig. 41). Similar intergradations with prismatic substance were

shown in some parts ‘of the preparation from which Pl. XL.

fig, 29a is drawn. Pl. XLII. figs. 40-42 are of interest as

showing all manners of intergradations between amorphous,

columnar, and prismatic substances, the different structures

shown being apparently mainly dependent upon the proportions

of calcium cabénuta present. Thus we have in this pearl,

which, owing to the impermeability of the amorphous substance,

was imperfectly decalcified, tracing the layers from inside out-

wards: (1) a plug of nuclear matter of doubtful origin con-

taining well-marked crystals (fig. 40, mw.)—these are true

crystals (rhombohedra); (2) a layer of amorphous substance,

passing over into typical simple columnar repair-substance (fig.

40, col.) ; (3) numerous layers of amorphous substance (figs. 40,

41, am.), some layers being quite lime-free, some having scattered

elect containing calcium carbonate, some showing thew cavities

in radial rows, leading up, by transitions, to regular columnar

substance (figs. 41, 42, col.), which differs from the prismatic

substance proper (fig. 42, pr.) only in the smaller diameters

of its constituent elements, a difference which, in view of the

variability of the sizes of the prisms in the shell itself, is com-

paratively unimportant.

Again, the transition from the abnormal repair-substances to

nacre in the pearl shown in Pl. XLV. fig. 52, col., and Pl. XLII.

fig. 56, col., is equally striking. This is shown in detail in

Pl. XLII. fig. 43. At nz. is the outer wall of the sphero-

crystalline or columnar pseudo-nucleus of the pearl. At rac. is

316 DR. H. LYSTER JAMESON ON

shown the normal nacre of the pearl. The first-formed layers of

nacre are incomplete, passing over into this area of repair-

substance, and all stages of transition may be seen, corresponding

to the gradually increasing control exercised by the secreting

epithelium.

At first, amorphous substance (am.), alveolar in places, was

secreted, no doubt with irregular crystallised bodies in the alveoli,

some of which are actually preserved in the preparation, owing

to incomplete decalcification. Peripherally this gave place to

columnar substance (col.), which acquired a finely alveolar structure,

and passed over, through granular repair-nacre (g7.), into normal

nacre (nac.'), the layers of the conchyolin of which gradually merge

into the horizontal markings of the granular substance. The

amorphous substance in this preparation varies from layer to

layer in the degree to which it is alveolar; at some places it

might better be described as coarsely columnar substance.

Amorphous substance seems to be the first product where the

shell is perforated and the mantle makes a sudden effort to close

an opening to the exterior. In such cases it may be secreted so

copiously that a tough leathery skin results, with little or no lime-

salts in it (Pl. XL. fig. 29, am.). It is likewise secreted in layers

when a break occurs in the nacre-secretion of a pearl or of the

shell, owing to a pathological extravasation of cellular matter

(Pl. XXXVIIL. fig. 17, am., am.'). These facts suggest that the

organic basis of the shell is the constituent the secretion of which

varies in quantity, the secreting-tissues (perhaps the granular

subepithelial parenchyma in WMargaritifera) contaiming a

reserve of this material which can be poured out profusely when

the shell is injured. It would seem that the lime-salts, on the

other hand, are secreted more regularly, so that the mechanism

for furnishing these cannot keep pace with that which yields the

organic substance when the latter is called upon to make a

special effort to repair damage. The resemblance of the inner

layers of the periostracum (in forms with a thick periostracum)

to amorphous repair-substance may perhaps be explained by

postulating the absence or inhibition of the lime-secreting

mechanism in the underlying tissues.

That the secretion of calcium carbonate could not keep pace

with that of the organic substance, when the latter is produced

in large quantities, is easy to understand in view of the very

small proportion of CaO in the blood of Mollusca, and indeed of

all invertebrates that have been investigated. According to

Griffiths (quoted by Biitschli, 6, p. 62), the CaO in the blood of

a number of bivalves examined varied from 0°032 per cent. in

Anodonta to 0:067 per cent. in Jytilus.

Amorphous substance is seen in the pseudo-nuclei of pearls in

Pl. XX XIX. figs. 20 & 21, and Pl. XL. figs. 24, 26, & 27; figs. 20,

24, & 27 showing particularly well its continuity and intergra-

dation with the organic basis of the columnar repair-substance.

In the centre of a pearl it may contain, in addition to the

THE CEYLON PEARL-OYSTER. S17

central cavity, secondary cavities in its substance, in which

organic particles are lodged (fig. 20).

The variations of coarsely alveolar structure which amorphous

substance shows (e.g. Pl. XL. fig. 29 @ and Pl. XLII. figs. 40 & 41)

recall those structures which Biitschli (6, Taf. iii. figg. 20-33)

describes in the spherocrystals of (?) Trydimite formed when the

siliceous concretionary substance of the Bamboo (known as Tabaxir

or Tabasheer) is heated; this structure is probably in great measure

the expression of the physical conditions (surface tension, etc.)

which prevail when two substances in solution or in a colloidal

state separate from one another to form a spherocrystalline

mass.

Pl. XLII. figs. 37 & 39 are of interest as showing another

variation of the nacre, in the direction of columnar substance.

In this variety of nacre, the conchyolin-layers are connected

by a number of thickened junctions, which tend to occur

in groups and which are arranged in radial rows. In surface

view these junctions appear as groups of dark spots on the

conchyolin-layers ; in radial section they are as shown in fig. 37,

and can also be seen in Pl. XXXVI. fig. 13. These junctions

seem to be thickenings of the walls which normally connect

the several conchyolin-layers of the nacre to one another; they

may, in fact, be regarded as local exaggerations of the condition

described as “ granular repair-nacre.”

It is interesting to note the peculiar manner in which these

repair-substances occur in Japanese ‘Culture Pearls.” This

name was given by the late Professor Mitsukuri (27 ¢, pp. 283-4,

pl. xi. fig. 1) to pearl-like bodies—‘“ blisters,” as they would be

called on the Australian fisheries—which are artificially produced

in the Japanese Pearl-Oyster, Margaritifera martensit * Dunker.

The production of these “Culture Pearls” is an extensive

industry supporting about 100 persons, and is carried on by

Mr. Mikimoto on leased areas of sea-bottom in the Bay of Agu,

Shima Province, on lines originally suggested by Prof. Mitsukuri

in 1890. It has been going as a commercial success since 1898,

when the first crop of ‘* Culture Pearls” was marketed. In 1905

the number of oysters operated on per year was from 250,000 to

300,000.

The process, which is protected by patents, is analogous to that

adopted by the Chinese in the production of ‘ Buddha Pearls”

in the fresh-water mussel, Dipsas plicatus, and to the method

discovered by Linnzus in the 18th century (see Herdman, 16 a),

and consists in the introduction between the shell and the mantle +

of a bead of nacre, which in due course (the time allowed in

Japan is four years) becomes thickly coated over with nacre,

* This mollusc is regarded by some naturalists as a local race of M. vulgaris, to

which it is undoubtedly very closely related. Whether it be called M@. martensii or

M. vulgaris var. martensii is largely a matter of individual taste.

+ This is apparently done vid the edge of the shell and not by drilling as in the

Linnzeus process.

318 DR. H. LYSTER JAMESON ON

forming a hemispherical, or sometimes rather more than

hemispherical pearl-like excrescence, attached to the shell by its

base. These “Culture Pearls” are produced in large numbers,

and find a ready market for purposes for which “ half-pearls” are

used. ‘They are now familiar objects in Europe *.

Text-figure 39 is a section of a Japanese “Culture Pearl,”

which [ purchased in London, while still attached to the shell,

and decalcified.

Text-fig. 59.

TW.

“

TUES ES

SE SS eee

eee Za LF SSS

—— ; TATA

a AANA ITT MN ATTAIN

pr col.

cot! Fnac. G7-

Section through a decalcified Japanese “Culture Pearl” still attached to the shell.

nu., the artificial “ nucleus,’ a bead of nacre, the Jamine of the nacre being

cut transversely ; pr., prismatic layer; nac., origimal nacreous lining, which

existed before the nucleus was introduced ; nac.', more recent nacre, lining the

shell and extending over the “nucleus” to form the “Culture Pearl,”

secreted after the introduction of the nucleus; nac.’’, nacreous layers where

the lining of the shell is carried over the nucleus; col., col.’, repair-substance

secreted im a zone around the point of contact between nucleus and shell,

where the deposition of the shell-substance was not controlled by the

mantle; g7., granular matter, perhaps of foreign origin or of the nature of

amorphous substance. X 10.

The ‘‘ nucleus” has been very skilfully introduced, so that there

is practically no trace of “dirt” between it and the nacreous

layer with which it is invested, as is so often the case in the

“« blisters ” which have been produced by naturalists and experi-

menters from time to time. Moreover, the disturbance of the

normal functions of the mantle has been so slight that, in the

* Needless to say, these bodies are not “ Pearls,” biologically speaking, but

belong to the class of structures to which I have applied the name “ blisters,”

familiar on the Australian Fisheries and in the Trade. Various naturalists have

produced such bedies from time to time. Ihave recently seen some very beautiful

‘ones produced in Margaritifera maxima, and I myself produced some presentable

ones in Margaritifera margaritifera in Papua in 1899-1900. But although

attempts have been made, and are still being made, to do this on a commercial scale,

T am not aware that commercial success has yet been achieved anywhere else than

in Japan; indeed, I think that the combination of circumstances which has led to

the success of the Japanese enterprise—viz., skill, patience, and intelligence, backed by

the best scientific advice and supported by cheap labour—has generally been lacking

in other ventures. I may add that the price that could be obtained for the best of

these gems is insignificant compared with the value of a real pear] of like size. No

THE CEYLON PEARL-OYSTER. 319

particular sections that I examined, there was a marked absence

even of the repair-substances. But in the zone immediately

around the point of contact between nucleus and shell, where,

when the nucleus was introduced, the epithelium of the mantle

was presumably unable to fit closely against the surfaces, it is

otherwise. Here, on examining the whole “ Pearl” as a trans-

parent object after decalcification, an opaque ring or zone was

distinctly visible. This was due to the presence of granular

matter, perhaps derived from the exterior, perhaps from the

tissues of the animal (text-figs. 39 & 40 B, gr.), and to very

irregular columnar and amorphous repair-substance (col., col.’),

This columnar substance is shown in greater detail in text-fig. 40,

A & B, corresponding respectively to col. and col.’ in text-fig. 39.

In text-fig. 40, A, in the niche between the nucleus and the shell,

where the mantle-epithelium could not reach, we see the product

of its secretion consolidated away from the influence of the

epithelium. Here the columns, instead of forming the character-

istic palisade-like structure, with their long axes perpendicular

to the secreting surface, are arranged in groups suggesting

incomplete spheerocrystals.

The curious fan-like arrangement which the columns take on in

text-fig. 40, B, suggests that the repair-substance arose through an

extravasation of the shell-forming fluids at the point a, the layer

nac.’ representing the first normal nacre, secreted by the mantle

when it occupied that position, the irregular columnar and

amorphous substance being due to the consolidation of the

secretion which filled the space, triangular in section, which lay

between the nucleus (nz.), the shell (nac.), and the mantle; the

position occupied by the last named being represented by the layer

of nacre marked nac’'.

(13) Varreties or Ceyton Pears.

Pending a classification based on the nature of the causes

which give rise to the formation of the pearl-sac, I propose,

following Herdman, to separate the pearls which I have examined

in or from the Ceylon Pearl-Oyster into two main groups,

accordingly as they typically occur clustered in the neighbourhoods

really satisfactory proof has ever been given that free spherical “pearls” can be

produced in this way, though Prof. Mitsukuri (7. c.) says that there are some hopes

that this will be done. There is no theoretical reason why a modification of the

Japanese or Linnean operation should not be devised which would achieve this end—

indeed, there is some reason to think that Linneus actually did produce some round

“pearls” and not only “blisters.” But such bodies, if produced, would not be

“pearls ” in the strict biological sense, though it is quite likely that they would be

marketed as such in quantities before the difference was detected.

Since writing the above, I have been informed by Mr. Toyozo Kobayashi, Professor

at the Tokyo Higher Technological College, who is associated with Mr. Mikimoto

in his enterprise, that perfectly free “ pearls” have been produced by these methods

within the last two years, but so far only exceptionally, and on a scale so small as

not to be applicable commercially,

320 DR. H. LYSTER JAMESON ON

of the muscular insertions or singly in the non-muscular parts of

the body-wall and mantle.

Text-fig. 40.

A. The irregular columnar and amorphous substances, shown at col. in text-figure 39.

nac., nacreous lining of the shell; xac.’, nacre deposited shortly after imtro-

duction of nucleus, passing over into the repair-substance (é7.) ; nac.’’, nacre

continuous with the layers investing the nucleus; col., col.’, columnar repair-

substance; am., amorphous repair-substance. XX 35.

B. The fan-shaped mass of repair-substance, shown at col.’ in text-figure 39.

nu., the introduced “nucleus”; naec., the original nacreous lining of the

shell; wac.’, the first layers of nacre, separated after the introduction of the

nucleus; col., columnar repair-substance; am., amorphous repair-substance ;

gv., granular matter, perhaps of extraneous origin, 100.

THE CEYLON PEARL-OYSTER. BP all

LT adopt Professor Herdman’s term ‘“ Muscle-Pearls” for the

former class, while for the latter category I propose the name

‘“* Parenchyma-Pearls ” *, because they occur typically in the

parenchymatous subepidermal tissues of the non-muscular parts

of the body-wall and mantle, or, by secondary displacement, in

the more deeply seated soft tissues.

This group corresponds, I think, to Herdman’s “ Cyst-Pear]s,”

but I prefer not to adopt the latter name, as, if the word “ cyst”

refers to the encysted Cestode, which Herdman associated with

pearl-production, I have been unable to trace the connection

between it and the pearl; while if it refers to the pearl-sac or

“cyst,” this is found around all pearls, including muscle-pear's.

There is some reason for believing that some parenchyma-pearls

arise from causes different from those that lead to the formation

of muscle-pearls, and, indeed, it is quite possible that parenchyma-

pearis have several modes of origin, as Herdman believes ; but,

on the other hand, their differences may be due in great measure

to the different parts of the tissues in which they originate,

and it is certainly quite impossible, in many cases, to say, from

the structure of a pearl and of its nucleus and pseudo-nucleus,

whether it is a “ muscle-pearl” or a “ parenchyma-pearl.” With

regard to Herdman’s “ Ampullar pearls,” I cannot regard this

group as of equal value to the above two classes, as, in my

experience, so far as it goes, the “Ampulla” is of secondary origin,

due to the absorption of the tissues intervening between the pearl

and the shell, and to the epithelium of the pearl-sac and that of

the outer face of the mantle thus becoming continuous.

Before going further I had better explain a term that I am

introducing into this paper. I am restricting the word

‘“‘Nucleus,” as applied to the body found in the centre of a

pearl, to those bodies which appear to be either of foreign

origin or derived from the pearl-oyster otherwise than through

the agency of the shell-secreting mechanism. To the~ bodies

formed by the shell- and pearl-secreting mechanism, composed,

asa rule, of different kinds of repair-substance (bodies which

have no doubt often been wrongly mistaken for objects of foreign

origin), I propose to apply the name “ Pseudo-nucleus.” I have

endeavoured to be consistent in the use of these two terms, but,

as is so often the case in biological matters, there is at times a

difficulty in defining a sharp boundary-line between the objects

to which they are respectively applied.

A. Musele-Pearls.

I have set out above (p. 267) Professor Herdman’s views on

the nature and origin of these. Briefly recapitulated, they are

the following. From some unknown cause, minute calcareous

* Rubbel (34 a) applies the term “ Mantelperlen” to th se bodies, a term which

I prefer to mine, though it is too late to alter the nomencla ure in this paper.

Proc. Zoou. Soc.—1912, No. XXI, 21

322 DR. H. LYSTER JAMESON ON

depositions or caleospherules * arise in the tissues, close to the

attachments of the muscles to the shell. Ectoderm-cells may

“migrate to the source of irritation, and thus be responsible for

the deposition of a pearl.” No explanation of the origin of these

calcospherules is given, but Mr. Southwell thinks it is ‘almost

eertain that they are depositions from the blood,” and refers to

them elsewhere as ‘‘ of excretory origin ” (42).

I have been led by my observations to take a quite different

view of these “ caleospherules” 7, and as their origin is so closely

related to that of Muscle-Pearls, I cannot do better than begin

the present section of my paper with an account of their struc-

ture and origin.

According to my view, Prof. Herdman’s “ caleospherules ” are

not free concretions at all, but are minute pearls, composed of

hypostracum ; and I propose, therefore, to call them “ hypostra-

cum muscle-pearls,” to separate them from ‘“ nacreous muscle-

pearls.” As stated by Herdman, these bodies occur close under

the epidermis (unless secondarily displaced, e. g. by the addition

of new ones), and I usually find them in the region where the

muscle-attachment epithelium passes over into the ordinary shell-

secreting epidermis of the mantle. A group of these hypo-

stracum-pearls is shown on Pl. XX XIX. fig. 22, which represents

a portion of the mantle-musculature of one of the unlabelled

specimens in the British Museum, examined entire in oil of

cloves. The same pearls, decalcified, are seen in fig. 23. These

little bodies measured from 0°02 to 0°5 mm. in diameter. In

Pl. XXXVIII. fig. 19 similar bodies, hy.p., are seen in a section

along with ordinary nacreous muscle-pearls; while single indi-

viduals are shown in Pl. XXXIX. figs. 21 & 21a@and Pl. XL.

fig. 25. Sections ground from these bodies, or cut from the organic

residues left when they are decalcified, show them to be composed

of the same substance as the hypostracum of the shell. They

consist of caleium carbonate, in fine fibrocrystalline form, showing

radial and also concentric markings, with a small central cavity

(Pl. XX XIX. fig. 21a). Decalcified they also resemble hypo-

stracum in all details of structure and reaction to stains

(fig. 2D) cost Ulver organic basis stains more blue with hematoxylin

than the organic parts of the other shell-substances, and takes up

carmine more deeply. Their alveolar structure is also much finer

than that usually found in the columnar varieties of repair-

substance, so fine, in fact, that in surface-sections the reticular

structure seems almost like that of the protoplasm itself. As has

been observed in the hypostracum of the shell, this substance

sometimes passes over into nacreous conchyolin laterally. The

* This word is presumably intended to convey the same idea as the word

“ eoneretion’’ adopted by me (25) in 1902, 7. e. a sphrocrystal-like body arising in

the tissues otherwise than by epidermal secretion; and therefore analogous to

cholesterin calculi, etc. (ef. Harting’s “ Calcospherites,” 12).

+1] find that Rubbel (34 a), working on the freshwater Pearl-Mussel,

Margaritana, has arrived independently at the same view of the nature of these

bodies as that here propounded.

THE CEYLON PEARL-OYSTER. 323

central cavity of a hypostracum-pearl may contain granules of

doubtful origin, as in the case shown in fig. 21, but it is frequently

quite empty. At times the organic basis of one of these hypo-

stracum-pearls, when decalcified, shows a tendency to break up

into segments, especially at its inner surface; the segments in

such cases probably correspond to the outlines of the original

secreting-cells ; indeed, in such cases the whole body may have

an almost cellular appearance, which is not surprising in view of

the generally accepted theory that the hypostracum arises by

direct transformation of the muscle-attachment epiderimis.

These hypostracum-pearls shrink, on decalcification, to about

one-third of their original diameters (Pl. XX XIX. figs. 22 & 23),

When the tissue is old and defectively preserved, as in this

preparation and in that shown in fig. 19, they come away from

the wall of the enclosing sac during decalcification ; but in better-

preserved material, where the connection between the muscle-

attachment epithelium and the pearl is maintained, the organic

basis of the decalcified hypostracum-pearl remains attached to the

wall of the sac (Pl. XXXYV. fig. 8). Nacreous pearls, on the

other hand, almost always shrink away from the sac on decalci-

fication.

The smallest of these hypostracum-pearls that I observed

measured about 0°02 mm. in diameter.

As the muscle-attachment epithelium, in Margaritifera at any

rate, takes at most a very small part in shell-thickening, the size

of these hypostracum-pearls is limited by the maximum thickness

to which hypostracum normally attains. For further growth to

occur, resulting in the formation of a nacreous muscle-pearl, it is

necessary for some of the nacre-secreting epidermis to be present

also (Pl. XXXYV. figs. 8, s., & 9; Pl. XXXVI. fig. 10, sac.).

Fig. 8, from a specimen given to me by Prof. Herdman, shows

above a nacreous muscle-pearl and below a hypostracum muscle-

pearl. Here we have a cyst, which is more or less spherical, and

contains a large central cavity lined with a substance which is

indistinguishable from the organic basis of hypostracum. Where

an epithelium can be detected in the wal] of the cyst (m.ep.') it

possesses all the characters of muscle-attachment epithelium, its

cells being continuous with the muscle-fibres, m., on the one

hand, and with the hypostracum, hy., on the other. In some

cases the muscle-attachment epithelium can be traced on all sides

of the sac; in others, as in fig. 8 and fig. 10, ¢., only at certain

parts. In still others no such epidermis is recognisable. I think,

however, it is safe to assume, whether the hypostracum-pearl is

surrounded by a sac of attachment-epidermis or not, that such a

‘pearl can only arise where such a sac is present ; and it is easy

to detect the epithelium in most of the better-preserved examples

(figs. 8, 9, & 10, m.ep.). Still, in some of the fairly well-preserved

preparations I can identify no such epithelium. ‘This is the case

in Pl. XXXVI. fig. 11. In this example, which is on one of

Prof. Herdman’s slides, the hypostracum-pearl, which measures

21*

324 DR. H. LYSTER JAMESON ON

80 p in diameter, and has a wall about 10 y» thick, lies close to a

nacreous muscle-pearl, about 1 mm. in diameter, the sac of which

is shown at ep.p.s. The cyst is embedded in a strand of muscle

traversing the mantle-parenchyma obliquely, and ending in muscle-

attachment epidermis which was attached to the shell. (Such

connections between the general musculature of the mantle and

the shell occur here and there quite apart from the more regular

muscle-scars. For examples of this in Mytilus see List, 27},

Pl. 8. fig. 1.) The cyst contains at one point a little granular

mass. The muscle-fibres here appear to be in direct contact with

the hypostracum. The easiest explanation of this condition would

seem to be the hypothesis that the original epithelium has dis-

appeared. It is not difficult to suppose that a highly specialised

“¢endinous” epithelium, like the attachment-epidermis, whose

fate seems to be to become a part of the shell, is incapable of

regenerating itself, and, therefore, destined to die and disappear

on ceasing to be functional. If we take this view, the typical

hypostracum-pearl is not so much a stage in the development of a

nacreous pearl as a phase parallel with it; the latter arising when

the original sac contains some of the ordinary nacre-secreting

epidermis, or cells capable of giving rise thereto, the former when

it is composed of attachment-epithelium alone. The hypostracum-

pearl would thus have a limited growth, the nacreous pearl an

unlimited growth. However, in considering these cases where

there does not appear to be any attachment-epithelinm, it must

be remembered that this particular epithelium is often very difficult

to see, so that some workers have even failed to detect its existence

on the regular muscle-insertions. Much light can no doubt be

thrown on these questions by a really thorough study of the

behaviour of the cells at the places where the muscle-attachment

epithelium goes over into the ordinary epidermis of the mantle,

and of the histological phenomena associated with the wandering

of the muscle-attachment. The material of the pearl-oyster that

[ have examined so far is not sufticiently well preserved to allow

of such study. So far as I know, this important matter has never

been properly investigated in any mollusc.

T will now pass from the hypostracum muscle-pearls to the

nacreous muscle-pearls. Typical instances of these are shown in

Pl. XXXV. figs. 8 & 9 and Pl. XXXVI. fig. 10. These three

examples are all explicable as derivatives of the hypostracum-

pearl. Figs. 8 & 10 obviously lie im the borderland between

one of the regular muscles and the parenchyma (fig. 8 is at the

insertion of one of the pedal levators). Fig. 9, from one of

Prof. Herdmaun’s slides, is in a place in the free mantle where a

few small muscle-strands (mzsc.) are attached to the shell. ‘The

sac of each of these pearls is lined in part by ordinary nacre-

secreting epithelium, underlying which is the typical granular

parenchyma, in part by muscle-attachment epithelium, continuous

with the musculature. As the former is much more active than

the latter, these pearls are all eccentric in shape, having a hilum

THE CEYLON PEARL-OYSTER. 325

of hypostracum at one side, which, unlike the nacre, does not

increase appreciably in thickness. The centre of each is a cavity,

which in figs. 8 & 10 is obviously lined with hypostracum, and this

hypostracum is connected by a plug of the same substance with the

remaining muscle-attachment epithelium. In fig. 9 the growth

of the nacre has pulled down the plug of muscle-attachment

epithelium into the hilum, and produced quite a long strand of

hypostracum-like substance. The presence of these hila, together

with the effect of the mutual pressure of muscle-pearls when

crowded together, has much to do with the generally irregular

shape of commercial seed-pearls. Fig. 10, also from one of

Prof. Herdman’s slides, shows a very early stage in such a muscle-

pearl, with a small cyst-like hypostracum-pearl alongside it. Here

the ordinary epithelium of the sae seems to be gaining on the

muscle-attachment epithelium.

These muscle-pearls always contain a central cavity, which may

be broken up by trabecule of hypostracum-like substance or of

conchyolin, this substance being continuous with that forming

the lining of the cavity. The cavity, like that of the pure

hypostracum-pearl, may be empty or may contain more or less

granular matter.

Muscle-pearls are often clustered and may be very numerous,

Thus the old unlabelled material in the British Museum has

dense clusters of these pearls in some places, and so has some of

Dr. Kelaart’s material.

It is by no means the case that muscle-attachment epithelium

always persists in the sac of a muscle-pearl. The whole sac

may pass over at an early stage into nacre-secreting epithelium,

a process which is, perhaps, analogous to what occurs in the wake

of an advancing muscle in the growing shell. This was apparently

the case with the pearl that occupied the sac adjoining the body

shown on Pl. XXXVI. fig. 11. The nucleus of this pearl is

shown at fig. 12 on the same plate. The central portion of this

pearl is composed of irregular conchyolin-like substance, which

cannot be identified as hypostracum, and which quickly

gives place to ordinary nacreous substance (n.). In the neigh-

bourhood of this pearl is another, not figured here, the centre of

which was comparable to the pearl shown on Pl, XXXV. fig. 8;

this pearl had become more spherical secondarily by the dis-

appearance, in the course of its growth, of the muscle-attachment

epithelium. It is hard to conceive that these two pearls, and the

hypostracum-pearl associated with them, are not all of similar

origin.

Pl. XXXVI. fig. 13 shows the centre of another pearl, perhaps

a muscle-pearl, from the mantle-margin, in one of Prof. Herdman’s

slides. This pearl appears to have measured about 2 mm. in

diameter. The central cavity is about 0°1 mm. in its greatest

diameter and is lined by abnormally thick conchyolin-like

substance. Outside this are layers of ordinary nacre, which pass

over into a form of repair-nacre showing radial markings,

€

326 DR. H. LYSTER JAMESON ON

probably due to variations in the rate of secretion of the con-

stituent substances. This zone is 0°03 mm. thick. The central

cavity is empty, except for a few granules.

In this case the muscle-pearl, 1f such it is, does not contain a

pseudo-nucleus composed of hypostracum or a spherocrystal-like

body such as those shown in figs. 19 & 20; and the real

“nucleus” of such a pearl might be said to be a cavity which may

or may not contain a few indistinct granules, perhaps of foreign

origin.

The same condition is also typical of those pearls which I have

examined from Dr. Kelaart’s material. I have decalcified 58 of

these in all (Pl. XXXVI. figs. 14, 15,& 16; Pl. XX XVIII.

figs. 17 & 18; Pl. XLIV. figs. 46, 46 a, 47,47 a, & 48). That these

pearls are of the same nature as the other muscle-pearls seems

probable from the fact that a few hypostracum-pearls occur mixed

with the other pearlsin Dr. Kelaart’s specimens, and from the com-

plete series of intergradations between the various forms described

above, which is shown by the unlabelled specimens in the British

Museum, described below. Pl. XX XVII. fig. 14 is a section of

Dr. Kelaart’s specimen showing “pearls in ovary.” Hach of

these pearls lies in a cavity which doubtiess was originally lined

with an epidermal epithelium, though this can no longer be

recognised owing to the state of preservation. The cavity is

surrounded in every case bya layer of the granular subepidermal

parenchyma (par.). Some of the pearls have been forced ont of

the subepidermal layer, and now lie embedded in the deeper

connective-tissue, in which are seen muscle-bundles and tubules

of the ovary.

Tn each case the centre of the pearl is a small cavity, containing

a few granules or strands of what appears to be conchyolin ; but

the pearl in the lower right-hand corner contains also some

columnar substance. The irregular conchyolin-like matter is well

seen in the centre of the pearl in the top left-hand corner of the

sketch, which is shown enlarged in fig. 15. It is interesting to

note that the series of sections from which these drawings were

made contained an example of the smaller Cestode larva, Z'ylo-

cephalum minus.

Plate XLIV. figs. 46, 46a, & 47, 47a show two pearls

picked from one of Dr. Kelaart’s specimens, decalcified, and

examined whole in oil of cloves (46 & 47) and after being sec-

tioned (46a & 47a). Fig. 46 shows a dense central mass, of

closely laminated nacreous substance, which on _ superficial

examination might be taken for the remains of a dead parasite,

but a section shows that the whole pearl is composed of nacreous

substance around a small central cavity.

Fig. 47, examined whole, was extremely suggestive of a dead

parasite ; indeed, the concentric lamination of the pseudo-nucleus

was not disclosed till sections were cut. ‘These (fig. 47 a),

however, furnished the explanation. The real centre of the

pearl was, as in the rest of Dr. Kelaart’s material, a nacreous

THE CEYLON PEARL-OYSTER,. 327

pearly mass, with a central cavity, showing at one side a plug of

conchyolin-like substance. External to the normal central nacre

were some irregular layers, such as one gets on the inner surface

of the shell when a dark blotch or blister is caused by derange-

ment of the secreting epithelium (compare the ‘ Olflecken ” in

Margaritana, Rubbel, 34a). The opaque character of these

layers, some of which were brown through the immigration or

infiltration of what appeared to be cellular matter, others

distinctly columnar (repair-substance), rendered the real nature

of the pseudo-nucleus obscure till sections were cut. Outside

these abnormal and pathological layers typical nacre was sub-

sequently produced, thus giving a normal pearl with a dark

centre.

The same characters are shown on Pl. XLIV. fig. 48, where

the centre of the pearl appears opaque and granular for a similar

reason. In this case the pseudo-nucleus measured about 5 mm.

in diameter, and, examined entire, might have been taken for a

dead parasite. It was such a nucleus, coupled with the presence

of Trematodes, probably Muttwa margaritijere Shipley & Hornell,

in the tissues of Dr. Kelaart’s pearl-oysters, that, in 1901, led me

to the probably mistaken conclusion that a Trematode might be

one of the organisms which afford the stimulus for the formation

of the pearl-sac in MJargaritifera vulgaris, as the Trematode

Gymnophallus does in Mytilus * (25, p. 162).

But examination of sections (Pl. XXXVII. fig. 16 and

Pl. XX XVIII. fig. 17) showed that the opaque pseudo-nucleus

was due to a break in the continuity of the nacre; a layer of

granular substance (gr.), apparently dead cells (perhaps of the

nature of the “oil-spots” in JJargaritana, or derived from an

immigration of leucocytes such as Moynier de Villepoix observed

(28, p. 112) or from Protozoan parasites), being followed by the

secretion of two horny layers of amorphous repair-substance

(am., am.'), after which normal nacre resumed its development.

The real centre of this pearl, as in the rest of Kelaart’s material,

consists (PI. XX XVIII. fig. 18) of shreds of conchyolin-like

material, and a few obscure granules, in a cavity which is sur-

rounded first by granular repair-nacre and then by ordinary

nacre.

In the two unlabelled specimens in the British Museum, from

which Pls. XXXVIII.-XL. figs. 19-28 and Pl. XLIV. fig. 49

are drawn, while the pearls agree with those described above in

their clustered habit, occurrence in the muscular regions, and

association with hypostracum-pearls, we have more frequently as

nuclei either hypostracum or special sphzrocrystal-like bodies,

which I regard as formed of columnar repair-substance.

* Fuller knowledge and closer study lead me to doubt the accuracy of my own

observations as to the occurrence of the remains of 'rematodes in the pearls produced

by the other species of molluscs referred to on p. 162 of my 1902 paper with the

exception, of course, of Mytilus edulis, in which the relation between Trematodes

and pearls is beyond question.

328 DR. H. LYSTER JAMESON ON

Pl. XX XVIII. fig. 19 shows a group of such pearls, scattered

among which are hypostracum-pearls (hy.p.). The nucleus of

the eccentric pearl at mw.’ is clearly composed, like that of the

incipient pearl shown on Pl. XXXIX. fig. 22, nac., of hypo-

stracum, and resembles the hypostracum-pearl shown at fig. 21,

which has a small quantity of granular contents. One of the

columnar nuclei is shown enlarged at fig. 20. This is the nucleus

of the large pearl shown in the upper part of fig. 19

(Pl. XXXVIIT.). Centrally there is a small cavity, in which a

granular mass of doubtful origin is noticeable (gr.). This cavity

is surrounded by an irregular zone of substance which seems to

be the same as the amorphous repair-substance of the shell

(of. Pl. XL. fig. 29). Externally to this there is columnar

substance, forming a sort of spheerocrystal-like mass, the fibro-

crystalline calcium carbonate being deposited in a conchyolin-like

basis, which, on decalcification of the pearl, remains as a

framework. This substance is seen cut tangentially at one end

of the pseudo-nucleus. Outside this is the normal nacre of the

pearl. In one or two cases the amount of granular matter in the

centre of the pearl was considerable ; thus Pl. XL. fig. 24, which

is taken from the same series as the preceding figures, shows the

central portion of a pearl which has a dumbbell-shaped double

nucleus, the larger half being about 17 mm. in diameter, and

each half containing an opaque brownish mass of dead animal

matter, perhaps of cellular origin, but quite unrecognisable. The

contents of the two cavities were continuous at one point. There

is nothing to suggest that either of these bodies represented a

dead Cestode; I could find no denser portion, such as would

naturally occur at the point representing the myzorhynchus, nor

was there anything that could be safely identified as the cuticle

of the parasite. If the contents suggested anything, it was rather

the remains of a large Protozoan parasite, containing spore-like

bodies in a plasma (like the Sporozoan spores which Dubois claims

to have identified in the nucleus of a pearl from J/. vulgaris from

the Mediterranean, 7, p. 311, and 8, p. 104); or, perhaps, a mass

of mucus containing a few cells. Hach of these centres was

surrounded immediately by the amorphous substance, this being

followed by, and continuous with, a radially calcified layer, the

bases of the calcareous prisms being evidently embedded in the

amorphous substance, which extended up between them to form

the organic framework of the columnar substance. This condition

is comparable to that seen at tr. on Pl. XLI. fig. 30, which shows

the amorphous substance secreted at the junction of two pearls

in a compound pearl, passing over into columnar repair-

substance.

Externally to the columnar layer of the pseudo-nucleus the

normal nacreous layers of the pearl are formed,

The other extreme is shown on Pl. XL. fig. 25,a hypostracum-

pearl ‘08 mm. in diameter, the central cavity of which has no

contents whatever.

THE CEYLON PEARL-OYSTER. 329

Pl. XL. fig. 26 shows a pearl having for its nucleus a double

hypostracum-pearl, the two constituents being apparently

separated by a mass of amorphous substance. Other examples

were examined which showed intergradations between double

and single hypostracum-pearls.

In tig. 27, from the same preparation as the above, there is a

central cavity, containing a few granules, and surrounded by

amorphous and columnar substances followed by nacre and finally

by hypostracum. In another case in the same preparation the

same state of affairs was shown, but there were further layers of

nacre outside the hypostracum, resulting in a fair-sized pearl.

These cases are of interest as indicating that the attachment of

muscle-fibres to a pearl may arise secondarily, in the course of

its growth. Im some cases the columnar substance is stratified,

as in the pearl at the bottom left-hand corner of fig. 19,

Pl. XX XVIII. andin fig. 49, Pl. XLIV. In Pl. XL. fig. 28, which

is the left-hand pseudo-nucleus of the double pearl shown in

fig. 49, there is at one pole a distinct transition from the columnar

matter of the pseudo-nucleus to normal nacre (nac.'), just as

occurs in the substance of the pearls figured at Pl. X LI. fig. 34

and Pl. XLIIT. fig. 43. This pseudo-nucleus is followed by

normal nacre (xac.), after which there is a layer of brittle horny

matter (am.), which I regard as analogous to amorphous repair-

substance ; this is followed by another columnar layer (col.’'),

imperfect on one side, and this again by nacre.

A preparation from the second unlabelled specimen in the

British Museum showed, side by side, all kinds of intergradations

between the various kinds of nuclei and pseudo-nuclei of muscle-

pearls described above.

I regret that I am as yet unable to trace the origin of the

muscle-pearls beyond the cyst-like sac, lined with hypostracum

or another form of shell-substance. One or two preparations

show cavities lined with a few cells, which may represent the

first stages of pearl-sacs. Moreover, here and there, the relations

of the epidermis, at the place where the muscle-attachment

epithelium gives place to ordinary mantle-epithelium, suggest

that these cysts arise by direct invagination at this region. What

may be the mechanism which gives rise to this invagination I

cannot say, there are certain indications that suggest that it may

be associated with the presence of parasitic Protozoa, an idea

suggested by me in 1902 (25, p. 162); but until I have been able

to examine better-preserved material, it would be unwise to

hazard yet another theory of pearl-production. At any rate, it

is clear that, if pearls in J/. vulgaris are of parasitic origin (and

T adhere to the view that the distribution of pearl-producing

examples of this and most other molluscs can at present best be

explained on the parasitic hypothesis), the immediate cause of the

pearl is not the mechanical irritation caused by the body of the

parasite, but rather the toxic properties of its secretions, which

lead to the pathological changes (formation of the tumours that

330 DR. H. LYSTER JAMESON ON

we call pearl-sacs) in the tissues; and that consequently the

‘cause ” of the pearl is not to be looked for in the nucleus, which

in the Ceylon pearl-oyster frequently does not exist, but rather im

the tissues of the oyster. This is, after all, just what I said

at the bottom of p. 142 in my 1902 paper.

The characters of muscle-pearls may be summed up as

follows :—

(1) They usually and probably invariably arise in close associa-

tion with the epidermis at the point where muscle-attachment

epithelium passes over into normal nacre-secreting epithelium.

(2) They frequently occur several together or clustered in

numbers.

(8) They are typically formed around central cavities in which

granules may be, but are not necessarily, present, and which may

be lined in the first instance with hypostracum, ordinary nacre,

or repair-substances analogous to those which occur where the

normal shell-secreting processes are disturbed.

(4) They are often associated with great numbers of little

bodies, which Herdman calls “ calcospherules,” and which I regard

as minute pearls composed of hypostracum.

(5) They are, according to Herdman, characteristic of certain

of the Ceylon beds; and are, therefore, local in their occurrence,

which would give support to a parasitic theory of their origin.

B. Parenchyma-Pearls ( Cyst-Pearls,” Herdman).

This class contains a much more heterogeneous group of nuclei

and pseudo-nuclei than the last. The pearls which it comprises

may have arisen from more than one cause, and it is more than

likely that a great many of the pearls which I refer to it are, in

fact, of the same origin as muscle-pearls, but have been produced

singly at spots where two or three muscle-fibres are attached to

the shell, instead of in clusters at the regular muscle-insertions,

and have consequently assumed a spherical form *.

Parenchyma-pearls often show a distinct central nucleus con-

sisting of granules or masses of dark substance which might be

either of parasitic origin or derived from dead tissue-cells (e. g.

leucocytes). In some cases the nuclei of these pearls contain

or consist of grains of sand or other foreign particles. The

nucleus is typically succeeded by one or more layers of repair-

substance, which often intergrades with the normal shell-

substances (nacre etc.) of the pearl. In consequence of the

presence of this abnormal shell-substance the centre of a paren-

chyma-pearl frequently contains a dark, spherical, concentrically

laminated, radially striated pseudo-nucleus. This opaque pseudo-

nueleus has no doubt on many occasions been mistaken for the

remains of a parasite, in pearls decalcified and examined entire ;

but seetions generally reveal its real nature unmistakably.

* These occasional attachments of small muscles to the shell on the general

mantle-surface are well known. See e.g. List (27 0), pl. 8. fig. 1.

THE CEYLON PEARL-OYSTER. 331

Cestodes may possibly occur in the centres of these pearls at

times, as Professor Herdman maintains, just as grains of sand

and other organic and inorganic particles cer tainly do; but this

remains to be proved, and my material has afforded no evidence

in favour of this much-quoted theory. And, even if it should

be proved that Cestodes do occasionally occur as the “ nuclei” of

Ceylon pearls, the real causes which underlie the development

of the pearl-sac (which is the essential factor in pearl-production)

have still to be discovered. The characters of the centres of

parenchyma-pearls can best be understood by reference to the

next section of the paper, which is largely descriptive of them,

pion many of the pearls referred to therein are unquestionably

“ muscle- pearls,”

(14) Descriptions oF THE CENTRES OF INDIVIDUAL PEaRLs.

In the absence of evidence connecting them with muscle-

pearls I propose to treat the four pearls from the Persian Gulf,

and the twenty-one pearls purchased for me in Ceylon by the

Ceylon Company of Pearl Fishers, Ltd., as parenchyma-pearls,

and to describe along with them eight of the pearls in Prof.

Herdman’s collection of slides which are not obviously muscle-

pearls. These descriptions will be followed by an account of a

number of samples and collections of pearls from J. vulgaris

from Ceylon and other localities.

A. Material from the Persian Gulf.

It may be useful to begin with the material from the Persian

Gulf. ‘This, so far as it has been examined—for some of my

specimens still remain unopened,—comprised three pearl-bearing

examples, containing four small pearls. All these pearls

appeared to be of the kind classified by Prof. Herdman as “ cyst-

pearls” and by the present writer as ‘‘ parenchyma-pearls,” as

they seem to have arisen quite independently of, and away from,

the muscle-insertions.

I append a short description of the conditions in each of these

cases.

Preparation LXIV a contained a solitary pearl, about 2mm.

in diameter and slightly lenticular in shape, situate in the left

mantle-lobe, on a level with the anterior end of the attachment

of the outer gill-lamella. Unfortunately, owing to imperfect

penetration in the embedding process, this pearl did not presenti

a complete series in section. It lay wholly in the subepidermal

parenchyma, which was sharply cut off from the underlying

visceral connective-tissue by a layer of musculature, under which

were the tubules of the ovary. The pseudo- -nucleus was a radially

calcified concentrically laminated body, about 3 mm. in diameter,

with a few darkly staining granules in the centre.

The second specimen, Preparation LXIV 3, contained another

332, DR. HU. LYSTER JAMESON ON

solitary pearl, which before calcification measured somewhat over

a millimetre in diameter, but when decalcitied shrunk to ‘7 mm.

It was situated in the left side, in the wall of the visceral mass,

over the stomach. In section (Pl. XLI. fig. 33) the pearl is seen

to le in the subepidermal parenchyma (par. ), projecting a little

through the muscular coat (masc.) into the visceral connective-

tissue (ext. ), 1 which the tubules of the ovary (ov.) and muscular

strands to the wall of the stomach (musc.’) are seen. The pearl.

sac has been ruptured, presumably by the evolution of bubbles of

carbon dioxide during decalcification.

The pseudo-nucleus, which measures about *3 mm. in diameter,

consists of several layers of columnar substance (col.) formed

around a small central cavity. Probably such a pearl would, as

it increased in size, grow through the muscular layer and

work its way into the visceral mass; indeed, as soon as more

than half its bulk lay on the inner side of the superficial

musculature, the action of these muscles would tend to force it

into a deeper position, where, relieved from the pressure of the

shell, it would stand a better chance of growing into a perfectly

spher ical “‘ fine pearl.”

Preparation LXIV c, the third of these pearl-bearing examples

from the Persian Gulf, contained two pearls, of minute size, about

2mm. apart on the left side. Hach was rather less than 1 mm.

in diameter; one was round, the other elongated. Both lay in

the subepidermal parenchyma, separated from the ovary by the

usual muscular sheath of the visceral mass. Both had pseudo-

nuclei of columnar substance. That of the more elongated one

is shown in Pl. XLI. fig. 34. It has a central cavity, with

irregular granular contents (gr.), surrounded by a number of

layers of nacre (nac.), outside which is a complete layer of columnar

substance (col.) which is coated on two sides by nacre, passing

over at the two poles corresponding to the longest axis of the

pearl into further coats of columnar substance (col.'). The

transition from nacre to columnar substance is very well

seen.

The centre of the second pearl in this example was also

composed of columnar substance, formed around a central granular

mass. The granular mass measured about ‘02 mm. in diameter,

the whole centre or pseudo-nucleus °3 mm. These preparations,

apart from the difficulty of obtaining material from the Persian

Gulf, are of interest as showing that the nuclei or pseudo-nuclei

of the pearls produced by J. aul garis in these waters do not

differ from those of the pearls produced by the same species in

Ceylon.

B. Pearls purchased in Ceylon.

T will now proceed to a short account of the nuclei and pseudo-

nuclei of the tweuty-one pearls purchased in Ceylon. All of

these were decalcified, examined entire in oil of cloves, and

drawn. Eighteen of them were also sectioned. (The drawings

THE CEYLON PEARL-OYSTER. 333

of those not figured here are retained with the slides for

reference.)

(a) One (Preparation XLV) had no obvious nucleus or pseudo-

nucleus.

(>) One (Preparation XLITT) had a small spherocrystal-like

centre of columnar substance, with no foreign contents

(Pl. XLV. figs. 50 & 50a@ and Pl. XLI. fig. 35).

to be irregular masses of amorphous shell-substance, in

central cavities, passing over into repair-substance (PI.

XLV. figs. 53 & 53a; Pl. XLVI. fig. 57; and Pl. XLII.

fig. 40).

(2) Three (Preparations XLVIT, LIIT, & LIV c) had central

cavities containing more or less granular matter, the

character of which could not be determined, but which

suggested dead organic particles (Pl. XLV. fig. 52 and

Pl. XLII. fig. 36). This was surrounded by repair-

substance of columnar structure.

(e) One (Preparation LIV 3) had in its central cavity granular

matter in which diatoms, fragments of sponge-spicules,

(¢) minute sand-particles, and vegetable débris were present.

These were surrounded by columnar repair-substance

(Pl. X LITT. fig. 44).

(f) One (Preparation LIV) had, in the otherwise indis-

tinguishable granular contents of the central cavity, a

fragment of a Radiolarian shell (Pl. XLII. fig. 45).

(7) Four had undoubted sand-grains, forming the nuclei around

which the pearl was laid down (Preparations LIT,

LIV a, LIVu, & LIVt1). The largest diameters of

the sand-grains in these four cases measured, respectively,

‘8 mm. 3x'l5mm., ‘6mm., and ‘3mm. (Pl. XLV.

figs. 54 & 54a; Pl. XLVI. figs. 55 & 56).

(2) One (Preparation LIV x) had a large spherocrystal-like

pseudo-nucleus of repair-substance, intergrading with

nacre, and a central cavity which had been partly

obliterated in cutting the sections, but apparently con-

tained several small quartz-fragments.

(t) Six pearls (Preparations XLII, LI, LIVs, LIVp,

LIVae, & LIV F) had, in their centres, bodies which

were so hard that they broke away in sectioning, and were

unfortunately not returned to me with the sections. The

probable diameters of these bodies in the first five pearls

were respectively ‘45 mm., *2mm., ‘1 mm., -25mm., and

‘6mm. Preparation LIV F had a _ pseudo-nucleus of

amorphous substance so dense that it broke away. In the

cases of XLIT, LIV, and LIV a, I feel pretty sure

that the nuclei here too were grains of sand, the others

may have had either sand-grains or abnormally bard

amorphous substance in their centres (Pl. XLV. fig. 51 ;

Pl. XLII. figs. 37-39).

334 DR. H. LYSTER JAMESON ON

Turning now from the “ nuclei” of these pearls to the “ pseudo-

nuclei” which surround them, these do not appear to present any

characters which cannot be expressed in terms of the several

pathological varieties of shell-substance described above. Thus

P]. XLII. fig. 44 shows the nucleus (composed of diatoms, etc.)

surrounded by typical amorphous and simply columnar repair-

substance, followed by ordinary nacre. This appears to be

equally the case in Pl. XLVI. fig. 55, where the nucleus is a

sand-grain.

Pl. XLI. fig. 35 and Pl. XLV. figs. 50 & 50a, which had

no foreign nucleus, Pl. XLII. fig. 36, Pl. XLV. fig. 52 with

granular contents in a central cavity, and Pl. X LIT. figs. 37-39,

in which example the nucleus was a sand-grain, and Pl. XLY.

figs. 51 & 53 and Pl. XLVI. fig. 56, show these pseudo-nuclei to

be composed of varieties of the columnar substance. Fig. 57

(Pl. XLVI.) and figs. 40-42 (Pls. XLII. & XLII.) show it

highly stratified, and presenting every gradation from amorphous

to prismatic substance. This preparation is a “ brown pearl,”

vather over 3 mm. in diameter, composed of the prismatic shell-

substance, and probably produced in the mantle-margin. It

consists of an outer zone, clearly identical in characters with the

prismatic layer of the shell, and showing three distinct series of

prisms. The bases of the innermost layer of prisms are shown in

fig. 42, pr. Inside this normal prismatic substance is the “ pseudo-

nucleus,” which consists externally of a layer resembling in every

respect, except the diameters of its constituent elements, the

prismatic layers (figs. 41, 42, col.). Internally this passes over,

through a substance resembling irregular columnar repair-substance

(fig. 41, ¢.), into amorphous substance containing columns or

scattered depositions of carbonate of lime (figs. 40, 41,a@m.). In

places these bodies are ranged in radial rows, and the amorphous

substance nearly reaches a stage which could be called columnar

substance.

Just around the centre (fig. 40, col.) typical columnar substance

occurs, passing over internally into amorphous substance.

Owing to the impenetrability of the amorphous substance, the

ealeium carbonate has not been dissolved in the deeper layers of

the pseudo-nucleus, and it is seen in the nucleus itself to occur in

the form of perfect rhombohedra, a form J have never seen it

take on in the alveoli of the normal or abnormal shell-substance.

The resemblance of this pearl to the brown Scotch river-pearls

(e. g. that figured by me in ‘ Nature’ for Jan. 22nd, 1903, p. 281

[26]) is striking.

Preparation XLITI, from which figures 50 & 50a o0n Pl. XLV.

and fig. 35 on Pl. XLI. are drawn, showed an oval central body,

about °6 mm. long, forming a “ pseudo-nucleus.” On superficial

examination this nucleus is highly suggestive of a Trematode

(fig. 50). But sections showed it to consist of a small sphero-

crystal-like body (fig. 85) about ‘08 mm, in diameter, surrounded

by granular repair-nacre.

THE CEYLON PEARL-OYSTER, 335

Figure 51 shows a pearl, the ‘‘ pseudo-nucleus” of which is

produced out in one direction as a strand of columnar and

granular repair-substance extending through the pearl. This

column: ar substance is seen to pass over laterally into the ordinary

nacre, and no doubt corresponds to an area in the pearl-sac at which

the normal secretion-processes were disturbed. ‘This was one of the

examples in which the real nucleus broke away.

Fig. 36 on Pl. XLIL., which is the centre of the pearl shown

in fig. 52 (Pl. XLV.), enlarged, shows interesting transitional

substance at one pole. This is further enlarged at fig. 43, Pl.

XLII. The pseudo-nucleus is seen to be discontinuous with the

later-deposited shell-substance, a condition of things which is

sometimes found, and which seems to suggest a pause before the

epithelium becomes properly functional.

Plate XLV. figs. 53 & 53a show a pearl with a spherocrystal-

like pseudo-nucleus, the longest diameter of which slightly

exceeded 1mm. In this case the columnar substance was very

minutely reticulated and fell into three distinct zones. The

pearl is interesting, as showing at one side a patch of columnar

repair-substance apparently continuous with the nacreous layers,

secreted over some foreign matter (granules) that had found their

way into the pearl-sac during the course of the pearl’s growth ;

this columnar substance formed a small “ blister” in the substance

of the pearl.

Where a sand-grain is present as nucleus, it appears to be

followed, as a rule, by repair-substance ; but in Pl. XLV. fig. 54

this is hardly discer nible, except at one pole. This was a pearl

which was drawn entire, and sent to be sectioned, when it was

found that the nucleus broke away and resisted cutting. This

nucleus was returned to me, and proved to bea siliceous sand-

grain (fig. 54a), measuring about °8mm. in diameter. It was

surrounded by an opaque substance, probably “ amorphous repair-

substance.”

Fig. 56 (Pl. XLVI.) shows a sand-grain which measured

about *35 x ‘25 mm. surrounded first by amorphous substance,

then by stratified columnar substance.

Fig. 55 (Pl. XLVI.), taken from a pearl which I have

preserved entire, shows a pyriform sand-grain, ‘66 x ‘5 mm.,

surrounded by a thin layer of columnar substance, followed by

nacre.

Figs. 37-39 (Pl. XLII.) are from the capsule surrounding

a nucleus which broke away and was probably a sand-grain.

Here, again, we have first the amorphous substance, passing

over into “ columnar” substance of sphzerocrystal-like for m, with

regular alveoli, which are approximately equal in length, breadth,

and depth.

Vigures illustrating minor varieties of the repair-substance as

it occurs in the shell, in the pearl, and in the pearl pseudo-nucleus

might be multiplied indefinitely, but I think enough have heen

given to illustrate my contention that these dark spherical bodies,

336 DR. H. LYSTER JAMESON ON

which so often suggest parasites, are usually in the Ceylon Pearl-

Oyster pseudo-nuclei of spherocrystal-like structure, each of

which is reducible to a certain amount of amorphous non-

calcified matter forming the lning of the nuclear cavity

(figs. 35, 36, 37, 40, 41, & 44, am.), and the same substance, some-

times stratified, sometimes not stratified, containing depositions

of calcium carbonate. It may be necessary to differentiate these

pseudo-nuclei into two groups, typically represented by figs. 36

and 40-42 respectively—the one secreted in all probability before

the epithelium of the sac could control its deposition at all (and

thus comparable to a “ Harting’s Body”); the other secreted by

an already partly functional epithelium. When material with

pearls in various stages of formation a sitw in the tissues is

available, this distinetion may prove to have an important

relation to the actual processes by which the pearl-sac arises, and

to the difference between the nuclei of muscle-pearls and some

parenchyma-pearls.

The most remarkable thing about this lot of pearls purchased in

Ceylon is the high percentage which have as nuclei grains of

sand or other foreign bodies. These cases might certainly be held

to give support to the old theory that the stimulation of any in-

trusive particle, 1f it occurs at the right place, may give rise to a

proliferation of epidermis resulting in the formation of a pearl-sac

and a pearl. But, on the other hand, evidence is not lacking that

the matter is by no means so simple as this. I am instituting

some experiments on Mytilus (in connection with investigations

on the life-history of the pearl-inducing Trematode that I am

carrying on with the aid of a Gover nment Grant) which will, I

hope, throw some light on this question. It is idle to speculate,

without experiments on living molluses, whether these particles

enter the tissue (@) in connection with mechanical i injuries to the

shell, as Herdman suggests, or (6) from the intestine, by perfora-

tion of the wall, and escape of such bodies as are seen in figs. 44

& 45 (Pl. XLIII.) into the blood, or (c) through lesions of

parasitic origin.

With regard to the last-named figures, it is interesting to note

that so long ago as 1857 Kelaart (277) suggested that the : siliceous

skeletons of diatoms might become the nuclei of pearls.

3 C. Professor Herdman’s Slides.

Professor Herdman’s preparations, illustrating pearls, which he

very kindly lent me, consisted (omitting a few preparations of

clustered hypostracum-pearls) of 24 slides, showing about 25

pearls in section. Of these pearls 13, representing 7 preparations,

were obviously muscle-pearls, some of which are shown in figs. 9—

12(Pls. XXXV.& XXXVI.). In four of these preparations the

cyst-like hypostracum-pearls were also present.

Eight pearls which were not obviously muscle-pearls, repre-

senting six preparations, are briefly described below.

(a) A pearl described in the label of slide as a “ muscle-pearl.”

THE CEYLON PEARL-OYSTER. 337

It was about 1°3mm. in diameter, and situated in the body-

parenchyma over the ovary (as in the Persian Gulf examples

described above). The centre was a mass of concentrically

laminated columnar substance, passing over externally into nacre

‘and about 0°5 mm. in diameter. I treat it here provisionally as

a parenchyma-pearl, as there is nothing to Suggest that it is a

muscle-pearl.

(6) This was a nice little spherical pearl, well out in the

mantle. The centre was a spherocrystal-like body, of the same

nature as those shown in figs. 35 & 36 (Pls. XLI. & XLI [.). There

were no recognisable foreign contents,

in the Persian Gulf specimens, with, apparently, all the characters

of a “ fine pearl.” It had a large dark pseudo-nucleus, 0-4 mm. in

diameter, much like that shown on Pl. XLVI. fig.57. There was

a central cavity, 0°025 mm. in diameter, containing a small

granular mass; this was followed by nacre, then stratified

columnar substance, then nacre again.

(¢) Showed two pieces of free mantle, sectioned, one with one

and one with two pearls in situ. In the case of the former the

pseudo-nucleus was incomplete, but could be seen to be composed,

in part at least, of columnar repair-substance. In the other

specimen, both pearls had centres resemblin g those in Dr. Kelaart’s

material, and may well have been muscle-pearls,

(e) The central part of this pearl is shown on Pl]. XXXVI. fig..13,

The pearl is quite close to the mantle-margin, in the musculature.

In the centre there is a eavity with a few granular contents.

This is surrounded by what appears to be ordinary naere, then

nacre with radial reinforcements, then nacre again. It suggests

a muscle-pearl rather than a parenchyma-pear.

(7) Contained a pearl, near the mantle-margin, the pseudo-

nucleus of which was not complete. It consisted of alveolar-

columnar matter; its centre could not be made out.

This preparation also contained an example of Z'ylocephalum

minus.

The remainder of the pearls in Prof. Herdman’s collection, four

in number, appeared to be incomplete preparations, as no nuclei

or pseudo-nuclei could be observed.

D. Unlabelled Pearls (dry) in the British Museum.

In September 1911 Mr. E. A. Smith, 1.8.0., allowed me to

examine four dry pearls, from a small collection of pearls and

attached pearls and blisters, preserved in the Mollusca cabinets

at the British Museum.

These specimens were unlabelled and without history, but their

general characters (colour ete.) were those of If. vulgaris, and the

fragments of shell to which some of them were attached were

undoubtedly referable to this species, and probably to examples

from Ceylon (heavily incrusted and much corroded with Cliona).

Proc. Zoou, Soc.—1912, No, XXTI, 22

338 DR. H. LYSTER JAMESON ON

Three of these pearls were decalcified, and one of them, Preparation

LXXXVI 4, was also sectioned. All showed centres of the same

class as those of the pearls in Dr. Kelaart’s collection—that is to

say, central cavities containing irregular conchyolin-like strands

and a few granules. Two of these pearls were fair spherical

pearls, about 3mm. in diameter, which would have ranked as

‘fine’ pearls or ‘ cyst-pearls,” but for blemished surfaces. The

third was a multiple pearl, secondar ily attached te the shell, and

showed, by the presence of distinct traces of hypostracum in its

substance, that it had arisen in a muscular part.

KE. Collection of Pearls given to me by Mr. Max Mayer.

In April 1911, Mr. Max Mayer, of Hatton Garden, London,

and Paris, dealer in precious stones, most generously gaye

me for the purposes of my work a number of pearls, mostly

of the baroque and seed class (though some could better be de-

scribed as pearls that would have ranked as small “ fine” pearls,

but for defects of shape, colour, or surface). These included

a sample which, after eliminating a number of pearls that,

from their colour and other qualities, Mr. Mayer considered

were derived respectively from Australia and Panama (these will

be dealt with in another paper), left a mixed lot of 115 pearls,

most of which showed more or less of the characteristic and

indescribable creamy colour which is distinctive of the pearls

of Margaritifera vulgaris. Mr. Mayer had no hesitation in

saying that the majority of these were almost certainly Ceylon

pearls, though he emphasized the fact that it is not often that

pearls reach the European markets accompanied by “ certificates

of birth,” and that the sources of these specimens could only be

judged from their respective characters.

These 115 pearls were decalcified and eight of them were

sectioned, and several others were dissected to expose the

nature of the “nucleus” or pseudo-nucleus.

Of this collection not one had a nucleus that could be identified

as a Cestode, or indeed as any parasitic worm. Sixty of them had

spherocrystal-like pseudo-nuclei composed of columnar repair-

substance, sometimes simple, sometimes stratified, sometimes

interstratified with other repair- -substances. In some cases a few

granules could be detected in the centre, in others nothing of the

kind could be made out. These pseudo- nuclei were generally less

transparent than the residue of the pearl, often almost opaque

but in every case their nature was obvious. Forty-two ‘pearls

were of the kind described from Dr. Kelaart’s specimens in the

British Museum, ze. they had either no obvious nuclei or their

centres contained cavities in which were a few strands of

conchyolin-like substance. Six others were of the same class, but

contained some refractive granular matter. One double pearl

showed a columnar pseudo-nucleus in one constituent and a

centre of the “ Kelaart” type in the other, while another,

THE CEYLON PEARL-OYSTER. 339

a quadruple pearl, showed nuclei transitional between these two

types.

There remain the eight examples which had to be sectioned

to disclose the nature of their centres, owing to opaqueness. The

first, Preparation XCIIJ, had a dark pseudo-nucleus which

proved to be composed of stratified amorphous substance, in fact,

a sort of horny pearl or periostracum-pearl. It was too hard

to section properly, and its real nature was discovered only

by dissection of the pseudo-nucleus. Preparation XCIV was

a triple pearl, and contained pseudo-nuclei of three kinds, a

‘“* Kelaart” pseudo-nucleus, one composed of columnar substance,

and one composed of concentrically stratified amorphous substance.

Preparation XCV had an irregular dark nucleus which in seetion

proved also to consist of stratified repair-substance, interstratified

with granules. In Preparation XCVI the centre was a hard

mass of columnar substance, coutaining a body that broke away

and may have been either amorphous substanee or a hard body of

foreign origin. Preparation XCVII contained a typical columnar

body, masked by a thick outer coat of amorphous substance.

The remaining three had pseudo-nuelei of ordinary columnar

substance.

I cannot say whether any of the above pearls contained

minute quartz-grains or other foreign inorganic bodies, as

this cannot be determined, even with the aid of polarised light,

in a great many eases, until sections are cut, owing to the high

degree of double refraction possessed by the conchyolin, and the

distortion of its original arrangement in the decalcifying process.

But such foreign bodies were detected in two pearls given to me

at the same time by Mr, Mayer, which were, however, more

probably from Margaritifera margaritifera var. mazatlanica

(Panama Shell) than from JZ, vulgaris.

F. Collection of Ceylon Pearls given to me by

Mr, H. Hopkins.

In October last Mr. E. Hopkins, dealer in precious stones, of

Hatton Garden, kindly gave me a parcel of fifteen small pearls,

which he believed to be Ceylon Pearls. He wrote me (letter of

26th October, 1911): “The pearls were obtained from a dealer

whose son visited the Ceylon Fisheries on the last occasion when

they were open, and from what he has told me I have every

reason to believe that these were part of the goods which he

brought back.” These fifteen pearls were decalcified and examined

as transparent objects, and four of them, which could not be

satisfactorily made out otherwise, were subsequently sectioned.

Most of these pearls were of the “ muscle-pearl” class, small and

angular, but two or perhaps three, both from their more spherical

shape and different nuclei or pseudo-nuclei, may more properly be

treated as parenchyma-pearls.

‘Twelve of these pearls had centres of the kind characteristic of

22*

340 . DR. H. LYSTER JAMESON ON

‘the pearls in Dr. Kelaart’s specimens, described above, 7.e. had no

obvious nuclei, but simply central cavities containing either a few

irregular strands of conchyolin-lke substance or obscure granules.

Two out of these twelve were sectioned. One, Preparation

Text-fig. 41.

A. Centre of a Ceylon pearl given to me by Mr. E. Hopkins. Decalcified and

examined whole in oil of cloves. Examined thus the body might be taken

for the remains of a Platyhelminthian parasite.

B. The same in section. The body is seen to be a “ pseudo-nucleus” consisting in

great part of amorphous substance, am., along with which is some granular

matter, gr. In the centre are some strange spore-like bodies, «, of doubtful

origin (Preparation LXXXV 1). X 400.

THE CEYLON PEARL-OYSTER. 341

LXXXV 4, had its central cavity lined with hypostracum, as in

the pearls described from the unlabelled specimens in the British

Museum. The other had a central cavity lined on one side by

« hilum-like plug of hypostracum, giving the pearl a reniform

outline, and on the opposite side by columnar substance, going

over into nacre, ‘wo of these muscle-pearls were composed of

several fused units.

There remain three pearls which have some claims to be treated

as parenchyma-pearls. One of these, which was not sectioned, had

in its centre a string of brownish granular matter, *66 mm. long

by ‘17 mm. wide, of doubtful origin (Preparation LXXXV 0).

Another (text-figure 41, A & B, Preparation LXXXV L) had

a pseudo-nucleus +1 mm. in diameter, which, examined entire,

might have been taken for the remains of a minute Platy-

helminthian parasite, but which on section proved to be made

up in great part of amorphous repair-substance. Finally, one

pearl (Preparation LXXXV x), a round pearl just under 2 mm.

in diameter, possessing apparently all the external qualities

of a parenchyma-pearl, had for its centre a pseudo-nucleus of

stratified columnar substance, 1 mm. in diameter, surrounding

a sand-grain *35 mm. in diameter.

G. Pearls from Margaritifera vulgaris from the

Gulf of Kutch.

Being for the moment unable to obtain freshly preserved

material from Ceylon or Madras, I endeavoured to obtain examples

of the same species, preserved with pearls im situ, from Northern

India, and through the courtesy of His Highness the Jam Saheb

of Nawanagar, better known to most people in this country as

Prince Ranjitsinhji, I was supplied in October last with a valuable

collection of preserved oysters, containing pearls, from the

Nawanagar fishery. I wish here to record my indebtedness to His

Highness, and to Mr. Merwanji Pestonji, Dewan of Nawanagar,

for their help. My thanks are also due to Dr. G, A. Grierson,

C.1.E., Director of the Linguistic Survey of India, and to Mr. R.

E. Enthoven, I.C.S., for putting me in communication with the

Authorities of the State.

The detailed histological examination of this material will take

some time, but I have decalcified and sectioned a number of pearls,

free and iv situ, for purposes of comparison with the material from

other sources.

As the Nawanagar fishery is little known to biologists, the

following information, supplied to me by the Dewan, may be of

interest.

The fishery, which belongs to H.H. the Jam Saheb, is carried

on on the southern shore of the Gulf of Kutch, during the South-

West Monsoon season, from June onwards. The ground on which

the pearl-oyster is found is * usually the sloping bank of a reef,

facing east or south-east, consisting of shingly sand and small

342 DR. H. LYSTER JAMESON ON

rocks and stones, clear of all mud.” Mr. Pestonji says that the

bottom of the sea around the reefs is muddy, and that it is

believed that the heavy rains and seas wash away the mud from the

Oyster-beds, and so make it easier to see and collect the oysters.

“‘ At high tide there is usually about ten to twelve feet of water

over the Oysters. The oyster-beds are never dry, even at low

water, there always being an inch or two of water draining off ” *.

The fishery is carried on ky wading, at low tide. ‘ During the

Monsoon season four or five hundred men are sent out toa certain

reef, where they tramp about and pick up the oysters as they come

upon them. They stay three or four days on each reef, and when

one reef has been searched they move on to another. Hach man

is rewarded according to the number and nature of the pearls found

in the oysters he brings in.”

The oysters are not “rotted,” as in the ease of the Ceylon pearl-

fisheries, but are opened one by one, and the pearls removed “ by

scraping the flesh gently with a blade of a knife.”

The number of oysters fished annually is about 150,000, on

an average. ‘he value is uncertain. The number of pearls

extracted from these oysters comes to about 20,000 and over.

They vary in size from seed-pearls to those weighing 20 grainsand

over. The lustre and colour are of first-class order, but the shape

in over 60 per cent. of the pearls is poor.

Mr. Pestonji estimates that if care is taken to pick up oysters

which are about four years old, about 10 to 15 per cent. of them

contain pearls. He says, in a letter dated November 14th, 1911:

“‘ Lately we opened oysters three times. The first time we opened

643 oysters and got 452 pearls. The second time we opened

770 oysters, and got 537 pearls. The third time we opened 845

oysters, and got only 379 pearls.” Mr. Pestonji includes the

minute “ dust ” pearls as well as pearls of different sizes and shapes

in this statement, and explains that there are often as many as

fifteen or more minute pearls in a single oyster.

Highteen pearls from this collection have so far been decalcified

and examined, and thirteen of these have been sectioned. Some

of these were sectioned 77 szdw in the tissues, others were pearls

which had fallen out of the tissues in the preserving process.

They differed in no recognisable microscopic features from the

pearls produced by the same species in Ceylon and the Persian

Gulf. Thirteen of these pearls, six of which were from one

specimen and three from another, were of the same character as

those described from Dr. Kelaart’s material in the British Museum,

that is to say, they had a small central cavity, surrounded by

ordinary nacre. The remainder had more or less obvious

pseudo-nuclei, composed of columnar or alveolar substance, in some

cases intérstratified with nacre. Two of these (Preparations

N. VIII. and N. 1X.) had naere inside the columnar substance, with

* Spring tides rise 18 ft., and neaps 14 ft., at Rojhi, an island near the town of

Nawanagar, according to the Admiralty Sailing Directions, “West coast of

Hindustan Pilot ” (1898 edition).

THE CEYLON PEARL-OYSTER. 343

a central cavity, and their difference from the “ Kelaart” class of

pearls was therefore probably due to a difference in their secondary

growth, rather than in their primary origin. One of those

sectioned (Preparation N. IV.) had a pseudo-nucleus composed of

stratified columnar substance, with internally some nacre sur-

rounding a small mass of brownish granules.

It cannot, I think, be argued in comparing these pearls with

Ceylon pearls that they are all seed-pearls or ‘“ muscle-pearls.”

They occurred in all the usual parts of the external body-wall and

mantle, and some of them. had all the properties of small “ fine

pearls.” Preparation N. VI., for example, was a beautiful little

spherical pearl, of high quality, measuring about 2°5 mm. in

diameter, which I was extremely reluctant to sacrifice in the

interests of science, and a number of the others were solitary

* parenchyma-pearls ” of small size.

H. Z'wo Pearls from Margaritifera vulgaris from the Mediterranean,

given to me by Professor Raphael Dubois.

Professor Raphael Dubois most generously placed at my

disposal for comparison two small pearls taken from J/. vulgaris

from the coast of Tunis. This species has firmly established

itself in several parts of the Mediterranean, since the opening of

the Suez Canal, having no doubt come thr ough from the Red Sea,

either in the free-swimming larval stage or as young spat att ached

to the bottoms of vessels. I can add nothing to what Dr. Dubois

has already said about the structure of these pearls, (8) pp. 103-—

105. One contained a central body, rather less than *5 mm. in

diameter, which on being sectioned proved to be composed of very

homogeneous granular matter surrounded on one side by columnar

substance and on the other side by “‘amorphous substance.” The

other contained a small yellowish spherical body, ‘08 mm. in

diameter, which, examined in oil of cloves, showed no recognisable

structure. In section it appeared to consist of a very small amount

of granular matter, surrounded by what looked like stratified

amorphous repair-substance. Dr. Dubois’s observations have

shown, without doubt, that the pearls produced by this species in

the Mediterranean (and their single occurrence, in positions away

from the muscle-insertions, renders it necessary to treat them in

part at least as Parenchyma-pearls and not as Muscle-pearls) have

not Cestodes in their centres, but have nuclei and pseudo-nuclei

similar to those which I have described above, for pearls from the

same species from other localities.

I. Pearls from M. vulgaris from New Caledonig, given

to me by Professor Seurat.

Professor L. G. Seurat, at my request, very kindly sent me four

small seed-pearls from Margaritifera vulgaris, from New Caledonia,

and also a slide, showing a larger pearl from the same species

S44 DR. H. LYSTER JAMESON ON

decalcified 2nd mounted entire in Canada Balsam. In none of

these could I find Cestode nuclei.

The example mounted as a slide was a pearl about 2 mm. in

diameter, of oval shape, from the Isle of Pines. It had two

pseudo-nuclei, side by side, both composed of columnar substance,

coneentrically laminated. One measured ‘0 mm., the other

‘3mm. This pearl was subsequently sectioned, and no foreign

matter could be detected in the nuclei. I decalcified and examined

the other four pearls. The characters of three of them were

nearest to those of the pearls in Dr. Kelaart’s collection in the

British Museum described above, but one had several centres,

which were surrounded by columnar substance. The fourth had

a little brown body about -2 mm. in diameter, which sections

showed to be composed of unrecognisable granular matter

surrounded byamorphous substance. No Cestode characters could

be identified in it.

M. Seurat also sent me five slides, with sections of pearls from

M. vulgaris from New Caledonia, which had been mounted in parafiin

wax, but had not been stained or made into finished preparations.

I stained and completed these, and found that they comprised

about seven pearls, in all of which, so far as could be determined

(for the series were not complete), the pseudo-nuclei were either

simple cavities containing a few granules or strands of conchyolin-

like matter, or masses of concentrically deposited, stratified

columnar substance. There was no trace of anything that I could

accept as a Cestode.

M. Seurat stated (37), p. 24, that these pearls had for nuclei

Cestode Jarve, but I am unable to confirm this., In a recent

letter he has informed me that this assertion was based on the

appearance of the nucleus of a decalcified pearl, which was not,

apparently, examined in section.

J. Pearl from M. vulgaris from Nossi-bé, Madagascar.

M. Seurat also sent me an unmounted slide with sections of a

pearl from this species from Nossi-Bé. The centre of this pearl

was a mass of concentrically deposited columnar substance,

0:35 mm. in diameter, containing a cavity about 0°l mm. in

diameter.

K. Pearls from the Lapi Shell (M. vulgaris), from the

Trobriand Islands, Papua.

Prof. W. R. Dunstan, F.R.S., Director of the Imperial

Institute, very kindly allowed me to decalcify and examine two

pearls from this locality, which were taken from among those on

exhibit in the Papua Court at the Imperial Institute. These

specimens had recently been sent home, by direction of the

Administrator, the Hon. M. Staniforth Smith.

The pearl-banks occur on the western side of Kiriwina Island,

THE CEYLON PEARL-OYSTER. 345

and the fishery is carried on by the natives from whom the pearls

are bought by licensed traders. The annual yield is between £3000

and £4500 worth of pearls, as valued locally. The first of the two

pearls examined was decalcified by me, but was not sectioned.

he centre was a double pseudo-nucleus, slightly over a millimetre

in length, and a little under a millimetre in breadth. It was

obviously composed of stratified columnar substance. External

to this the pearl, which was quite a good one, was composed of

normal nacre.

The second example was decalcified and examined entire, and

then sectioned. The centre was a_ spherical pseudo-nucleus,

rather less than half a millimetre in diameter, also composed of

stratified columnar substance, with a minute central cavity, about

0-02 mm. in diameter.

These were certainly not “ Muscle” pearls, but small “fine”

pearls. The preparations have been returned to the Imperial

Institute.

L. Pearls from Placuna placenta, from Lake

Tampalakamam, Ceylon.

In view of Mr. Hornell’s statement, referred to in the account of

the work in Ceylon, to the effect that he had determined that a

Cestode larva, similar to or identical with that found in the pearl-

oyster, caused the pearls produced in such quantities by Placuna

placenta, the Window-pane Oyster, I thought that a study of this

form would throw light on the problem in J/. vulgaris. I made

several fruitless efforts to obtain material from Ceylonandelsewhere.

I examined one small pearl, decalcified and mounted whole, in

Professor Herdman’s collection (No. 78), labelled “ Pearl Shelled

from Mantle of Placuna.” This pearl measures about half a

millimetre in diameter, and the “nucleus” is a little yellowish

body, about ‘05 mm. in diameter, and thus too small to be Hornell’s

larva (which is said to be from -2 to +4 mm. in diameter), even if

it is of foreign origin. However, it was necessary to have

material to section before the matter could be investigated. As

I could not get other material Professor Dunstan very kindly

allowed me to take a few Placuna pearls from the collection on

exhibit in the Ceylon Court at the Imperial Institute. Two very

well marked size-groups occur in the pearls in this collection,

there being a majority of minute irregular pearls, 1 mm. in

diameter and under, and a minority of larger ones, averaging from

2 to 3 mm. in diameter. Prof. Dunstan allowed me to take

four of the latter and sixteen of the former. All these were

decalcified, and two of the larger and four of the smaller were

sectioned.

Of the larger pearls, Preparation CXLVI (sectioned) had for

its centre a cavity, ‘05 mm. in diameter, containing a little

irregular columnar repair-substance. Preparation CXLVIT had

a central cavity containing a small amount of yellow refractive

346 DR. H. LYSTER JAMESON ON

granular matter. The remaining two, which were decalcified but

not sectioned, had no obvious nuclei, the centres being, apparently,

like those of CXLVI and CXLVII. Of the sixteen minute

pearls four were sectioned. Preparation CXXXV had in its

centre a simple cavity, like the pearls in Dr. Kelaart’s material

of A. vulgaris. Preparation CX X XVI showed a central cavity,

with around it some columnar substance, or perhaps hypostracum.

Preparation CXXXVII had a large dark pseudo-nucleus, over

half a millimetre long, which might easily have been taken for a

foreign body, when examined entire, but which on being sectioned

proved to be composed of concentrically laminated amorphous

substance, passing over in places into columnar repair-substance,

and containing a minute central cavity. Preparation CX XX VIII

had in its centre a little brown body, probably composed of

amorphous substance, but this specimen did not prove satisfactory

on being sectioned. The twelve remaining pearls were decalcified

and examined entire in oil of cloves, but were not sectioned. Of

these four had either a small central cavity, surrounded by

ordinary nacreous conchyolin, or no obvious nucleus or pseudo-

nucleus. Six had such cavities, partly or entirely surrounded by

repair substances or hypostracum, one had a few yellowish

granules in its cavity, and one had a dark body, about *3 mm. in

diameter, of doubtful nature, but showing no characters that

would warrant its identification as a Cestode.

The absence of evidence in support of Mr. Hornell’s Cestode

theory of the origin of Placuna Pearls is an additional argument

in favour of the opinion that the supposed observation of these

worms by Mr. Hornell and Professor Herdman in the pearls of

M. vulgaris was a mistake.

M. Pearls from Margaritifera margaritifera var. cumingi Heeve

(the Black-edged Mother-of-Pearl Oyster), from vrkitea,

Gambier Archipelago.

In order to enable me to compare the pearls of M. vulgaris with

those of this species, in which Seurat claims that Cestodes occur as

nuclei, M. Seurat most generously placed at my disposal some

material. This consisted of three slides on which sections of

pearls from this species had been mounted, but which had not

been stained or completed. J stained and finished these three

slides. In two of them the centre of the pearl could not be found,

in the third there was not a section through the exact centre, but

there was one that cannot have been far from it. This showed no

Cestode remains, but a pseudo-nucleus rather less than half a

millimetre in diameter, consisting in its more central part of

concentrically deposited columnar repair-substance, passing over

peripherally into alternate layers of amorphous and columnar

substance, which in their turn passed over into nacre, through

catenulated and granular transitional substance, such as I have

described in MM. vulgaris.

THE CEYLON PEARL-OYSTER. 347

M. Seurat also gave me a piece of the latero-dorsal region of

the body of this species, with pearls in situ. This I decalcified

and examined in oil of cloves, and afterwards sent to be sectioned.

At the time of correcting proof the sections had not been returned.

Examined entire in oil of cloves this specimen showed a cluster of

about ten pearls. Most of these were like those described from

Dr. Kelaart’s material of M. vulgaris, i. e., they had no obvious

nuclei; one, however, had in its centre a minute hypostracum

pearl or columnar pseudo-nucleus, about ‘1 mm, in diameter ;

one had a tiny refractive body, about ‘03 mm. in diameter, which

may have been composed of amorphous substance ; and one had

some more opaque matter which, however, contained nothing that

could be identified as a Cestode.

It would seem possible, in view of these observations, that

M. Seurat may also have been led into the error of arguing that

because the Trematode which is associated with pearls in Mytilus

furnishes the stimulation necessary for pearl production, therefore

the Cestode in Margaritifera plays thesame role. I hope shortly

to receive further material from the French Pacific that will

enable me to go into this question more fully.

I may add that my studies on pearls from M. mawina, and

from the other varieties of I. margaritifera from other localities,

so far as these studies have gone up to the present, afford no

evidence of the occurrence of Cestodes in the centres of pearls.

(15) GENERAL SuMMARY.

The following are the principal conclusions to which these

investigations have led me :—

(1) The evidence that the globular Cestode larve, which Prof.

Herdman regards as the cause of the formation of * fine pearls” in

the Ceylon Pearl-Oyster, are a young stage of the worm described

by Shipley and Hornell as Tetrarhynchus wnionifactor is quite

inconclusive. I consider these worms to be more probably

referable to the genus 7ylocephalwm (or an allied form), and have

provisionally described them under the name of Tylocephalum

ludificans and 7’. minus, spp. nn.

(2) The theory that these Tapeworms are the cause or a cause of

the formation of pearls in the Ceylon Pearl-Oyster (in the sense

in which the Trematode is the “cause” of pearls in Mytilus,

where the pearl-sac is normally formed as a result of the specific

stimulation of the worm) is supported by quite insufficient

evidence, and even their occasional occurrence in the nuclei of

Ceylon pearls has yet to be demonstrated *.

* Sir West Ridgeway, formerly Governor of Ceylon, Chairman of Directors of

the Ceylon Company of Pearl Fishers, Ltd., in reply to a question put at the annual

meeting of Shareholders in 1909 (reprinted in the ‘ Financial News > on December

21st, 1909), as to what was known with regard to the real cause and mechanism of

pearl-formation, implied that the Directors were in possession of valuable informa-

tion, of a secret nature, on the subject. “It was most undesirable at this moment

that they [i. e. the Directors] should reveal the progress which had been made by

348 DR. H. LYSLTER JAMESON ON

It appears, in fact, as though the simultaneous presence of pearls

and Cestodes in the Ceylon Pearl-Oyster were a case of two parallel

diseases, comparable to the case of a dog infected simultaneously

with tapeworms and mange, or of a man suffering at the same time

from echinococci and scabies. And even should it be found that

tapeworms do sometimes form the nuclei of Ceylon pearls, an

explanation of the reason why this occurs in some cases only is

necessary. It is, of course, possible that in certain of the Ceylon

banks conditions may exist which cause Tylocephalum ludificans

to depart from its normal habit, and acquire an ectodermal instead

of a fibrous cyst; or it might even be found that in certain banks

another species of Tylocephalum (or other Cestode) occurs which,

like the Trematode in Wytilus, normally and habitually gives rise

to a pearl-sac in the tissues, and which has been confused with

Tylocephalum ludificans. But, in any case, pearls formed around

tapeworms, if such ever occur (and this still remains to be demon-

strated), must be so comparatively scarce that, from the economic

standpoint, the réle of the tapeworm in pearl-production in

Margaritifera vulgaris must be unimportant. Examination of

such pearls from Placuna placenta and Margaritifera margaritifera

var. cumingit as were obtainable failed to confirm the supposed

occurrence of Cestodes as their nuclei.

(3) The shell of Margaritifera comprises in addition to the

hinge-ligament the same layers as those of other typical Lamelli-

branchs, viz. a periostracum (the outermost layer of which 1s

secreted in a different manner to the remainder and is very

much reduced in Margaritifera), prismatic substance, nacre, and

hypostracum or muscle-attachment substance.

In addition to these, certain pathological varieties of shell-

substance arise when the normal rhythm of secretion is disturbed,

the chief of which are described as amorphous repair-substance

(which is probably simply uncaicified conchyolin), columnar repair-

substance, and granular repair-nacre. These substances intergrade

with normal nacre and prismatic substance, and with each other.

The peculiar characters of these substances are the chief cause of

the distinctive appearance of the ‘‘ pseudo-nuclei” of pearls. The

shell-substance, except the hypostracum and the outer layer of the.

periostracum (and probably the ligament), is secreted in liquid

form, and its structure and variation may be interpreted as the

expression of the normal processes of the crystallisation of CaCO,

in a colloidal medium, modified by the periodicity of the action

of the shell-secreting tissues of the mantle.

(4) The “Calcospherules,” which Herdman identifies as the

nuclei of muscle-pearls, are not free concretions or ‘‘ depositions

Mr. Southwell in his researches on this question; but he thought that gentleman

would support him in saying that those researches had not been unsuccessful up to

the present, and promised to be most interesting as well as satisfactory.” Mr.

Southwell’s thoroughly frank statements of the case in his subsequent publications

do not appear to me to reveal anything sufficiently epoch-making, from the com-

mercial standpoint, to render such mystery necessary !

‘THE CEYLON PEARL-OYSTER. 349

from the blood” (Southwell), but are minute pearls formed

of the hypostracum or muscle-attachment substance. They are

therefore not the cause of the nacreous muscle-pearls, but a phase

parallel to them. There is some reason to believe that the origin

of muscle-pearls is associated with pathological invaginations

or immigrations of the epidermis at the points where the

muscle-attachment epithelium passes over into the ordinary

outer mantle-epithelium.

(5) Parenchyma-pearls (which name I apply to Prof. Herdman’s

cyst-pearls) may be formed around grains of sand or other foreign

particles, organic granular matter of doubtful origin, or bodies

composed of varieties of the shell-substance which arise when

the normal rhythm of secretion is disturbed (repair-substance).

A foreign nucleus is probably rather exceptional. The ultimate

factors which give rise to the epidermal sacs in which they are

formed have yet to be discovered. Many of them are probably

of the same origin as muscle-pearls, except that they arise singly

at points where a few muscle-fibres are inserted into the shell,

instead of in clusters at the regular muscle-insertions. The

dark pseudo-nuclei of these pearls, which may easily be mistaken

for the remains of parasites, are usually composed of the repair-

substances.

(16) Works REFERRED 0.

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Molluskenschalen. Jenaische Zeitschr. f. Naturwiss. Bd. xxxvi. Hft. 1.

pp. 1-164, Taf.i-vi. 1901.

(2) Brepermann, W. 1902.—Ueber die Bedeutung von Krystallisationsprozessen

bei der Bildung der Skelette wirbelloser Thiere, namentlich der Mol-

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(8) Dusois, R. 1909.—Contribution 4 Etude des Perles fines. Annales de

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(Séance du 31 Octobre, 1903), t. lv. p. 1222. 1903.

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pp. 612-614.

(12) Hartine, P. 1872.—Recherches de morphologie synthétique sur la production

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Kon. Akad. d. Vetenskapp. Amsterdam, Deel 14, 1872, 85 pp., 4 pls.

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March 27th, 1903; Abstract in ‘ Nature,’ vol. 67, 1903, pp. 620-622.

350 DR. H. LYSTER JAMESON ON

(15) Henpman, W. A., & Hornett, J. 1903.—Note on Pearl-formation in the

Ceylon Pearl Oyster. British Association Report, Southport, 1903,

p. 695.

(16) Herpman, W. A. 1903-6.—Report to the Government of Ceylon on the

Pearl Oyster Fisheries of the Gulf of Manaar. London: Published by the

Royal Society. 1903-6.

Part I. 1908.

Herpman, W. A.—Introduction and Narrative, pp. 1-98.

ks A Description of the Pearl Banks, pp. 99-121.

re 5 Observations and Experiments on the Pearl Oyster,

pp. 125-146.

Part IT. 1904. Preface, pp. v—vii.

Hurrpman, W. A.—History of the Principal Pearl Banks, pp. 1-36.

5 A Anatomy of the Pearl Oyster, pp. 37—76, pls. ix.

Surpbtey, A. E., & Horney, J.—Parasites of the Pearl Oyster,

pp. 77-106, pls. i.-1v.

Part III. 1905. Preface, pp. v—viil.

Herpman, W. A. (based on Report and Letters of Mr. Hornett).

The Pearl Fishery of 1904, pp. 1-36.

Be >, (based on Report and Letters of Mr. Hornetr).

The Present Condition of the Pearl Banks, pp. 37-48.

Surpiey, A. E., & Hornet, J—FKurther Report on Parasites, pp. 49-

55, plate.

Part IV. 1905. Preface, pp. v—vi.

Herpmayn, W. A.—The Great Pearl Fishery of 1905, pp. vii-xvi.

Part V. 1906. Preface, pp. -vii.

Herpman, W. A., & Horney, J.—Pearl Production, pp. 1-42,

pls. i-iil.

Surety, A. E., & Hornet, J—Cestode and Nematode Parasites

from the Marine Fishes of Ceylon, pp. 43-96, pls. i.—v1.

Herpman, W. A.—General Summary and Recommendations, pp. 109—

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(16 a) Herpman, W.A. 1905.—Presidential Address delivered at the Anniversary

Meeting of the Linnean Society of London on 24th May, 1905. London:

Taylor & Francis, 1905.

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the Linnean Society of London on 24th May, 1906. London: Taylor &

Francis, 1906.

(18) Hesstine, I. 1859.—Die Perlmuscheln und ihre Perlen. Leipzig, 1859.

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1904. Reports from the Ceylon Marine Biological Laboratory, No. L.,

1905, pp. 1-39.

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Pearl Banks, 1904. Ceylon Sessional Papers, xi1., 1905.

(21) Hornett, J. 1905.—Report on the Placuna Placenta Pearl Fishery of Lake

Tampalakamam. Ceylon Sessional Papers, xiv., 1905. ;

(22) Hornets, J. 1906.—Report on the Placuna Placenta Pearl Fishery of Lake

Tampalakamam. Ceylon Marine Biological Reports, Part II., June 1906,

pp. 41-54, plate.

(23) Hornett, J. 1906.—Report on the Inspection of the Ceylon Pearl Banks,

November 1905. Ceyion Sessional Pavers, xvii., 1906.

(24) Huxiry, T. H. 1859.—Tegumentary Organs. Todd’s Cyclopedia of Anat.

& Physiol. vol. 5 (Suppl. Vol.). 1859.

(25) Jamuson, H. L. 1902.—Gn the Origin of Pearls. Proceedings of the

Zoological Society, 1902, vol. i. pp. 140-166, pls. xiv.—xvil.

(26) Jameson, H. L. 1903.—The Formation of Pearls. Nature, vol. 67, pp. 280-2,

Jan. 22nd, 1903.

(26 a) Jameson, H. L. 1912.—An Examination of the Causes ah aye led to

the Failure of the Biological work recently undertaken on the Ceylon

Pearl] Fisheries. Journal of Economic Biology, vol. vu. pt. 1, pp. 10-22.

Feb. 1912. \

(261) Jounstene, J. 1912.—Tetrarhynchus erinaceus van Beneden.—i. Strue-

ture of the Larva and Adult Worm. Parasitology, vol. iv. no. 4,

January 8, 1912, pp. 364-415, pls. xix.—xxiv.

(27) Kexaart, EK. HL. 1857. —Introductory Report on the Natural History of the

Pearl Oyster of Ceylon. Trincomalee, 1857.

(27a) Linton, E. 1891. —Notes on Entozoa of Marine Fishes of New England,

with Deseriptions of several New Species—Pt. II. Report U.S.

Commission of Fisheries for 1887 [1891], pp. 719-899, pls. 1.—xv.

THE CEYLON PEARL-OYSTER. 351

(276) List, T. 1902.—Die Mytiliden des Golfes von Neapel. Fauna und Flora

des Golfes von Neapel: 27. Monographie. Berlin, 1902.

(27c) Mirsuxort, K. 1902.—The Cultivation of Marine and Fresh-water Animals

in Japan. Bulletin of the U.S. Bureau of Fisheries for 1904, vol. xxiv.

1905, pp. 259-289, pls. i.—xi.

(28) Moynrer pe Virtieporx, R. 1892.—Recherches sur la formation et l’accrois-

sement de la Coquille des Mollusques. Journ. de l’Anat. et dela Physiol.

t. Ixxviil., 1892.

(29) Minter, F. 1885.—Ueber die Schalenbildung bei Lamellibranchiaten.

Schneider, Zoolog. Beitriige, Bd. i. Hft. 11. pp. 206-246, Taf. xxvili—xxx.

(80) Pacrnstecner, H. A. 1858.—Uber Perlenbildung. Zeitschr. f. wiss. Zool.

Bad. ix. pp. 496-506, Taf. xx, 1858.

(81) ReaumeR. 1709.—De la formation et de Vaccroissement des Coquilles des

animaux. Hist. de Acad. Roy. des Sciences; Paris 1711. Mém.

Anneé, 1709.

(32) Romer, O. 1903.—Untersuchungen tiber den feineren Bau einiger Muschel-

schalen, Zeitschr. fiir wiss. Zool. Bd. lxxv. Hft. iii. 1903.

(33) Rupeer, A. 1911—Zur Kenntniss der Schalenregeneration bei der Fluss-

perlmuschel. Zoologischer Anzeiger, 7. Marz, 1911, Bd. xxxvii. Nr. 8/9,

pp. 169-172,

(84) Russet, A. 1911—Die Entstehung der Perlen bei Margaritana margari-

tifera. Zoologischer Anzeiger, 25. April, 1911, Bd. xxxvii. Nr. 19/20,

pp. 411-416.

(84a) Russet, A. 1911.—Ueber Perlen und Perlbildung bei Margaritana

margaritifera nebst Beitragen zur Kenntniss ihrer Schalenstruktur.

Marburg, 1911. Pp. 80, two pls., and numerous text-figures.

(35) Srurart, G. 1904.—Sur la Biologie des Huitres perliéres et nacriéres des

Iles Gambier. C.R. Soc. Biol. tom. lvi. 1904, No. 7 (26 Février), pp. 294-295.

(86) Srurar, G. 1906.—Sur un Cestode parasite des Huitres perliéres déterminant

la production des perles fines aux Iles Gambier. Paris, C.R. Acad. Sci.

tom. 142, 1906, pp. 801-803.

(87) Seurar, G. 1906.—La Nacre et la Perle en Océanie.—Péche.—Origine et Mode

de Formation des Perles. Bull. Mus. Oceanogr. Monaco, No. 75, 1906.

(88) Souruwett, T. 1910.—On the Determination of the Adult of the Pearl-

inducing Worm. Ceylon Marine Biological Reports, Part IV. pp. 169-172.

(89) Sournwert, T. 1910.—A Note on Endogenous Reproduction discovered

in the Larve of Tetrarhynchus unionifactor inhabiting the Tissues of

the Pearl Oyster. Ceylon Marine Biological Reports, Part IV., May

1910, No. 7, pp. 173-174.

(40) Souruwett, T. 1911.—Physical and Biological Conditions on the Pearl

Banks. Ceylon Marine Biological Reports, Part V., March 1911,

pp. 191-194.

(41) Souruwett, T. 1911.—Further Notes on the Determination of the Adult

of the Pearl-inducing Worm. Ceylon Marine Biological Reports, Part V.

pp. 213-215.

(41a) Sourmwett, T. 1911.—Description of Nine new Species of Cestode Para-

sites, including Two new Genera, from Marine Fishes of Ceylon. Ceylon

Marine Biological Reports, Part v. No. 18, pp. 216-225, plates. 1911.

(42) Souruwett, T. 1911.—Some Notes on the Ceylon Pearl-inducing Worm.

Spolia Zeylanica, vol. vii. Part xxvii., May 1911, pp. 124-134.

(43) Steinman, G. 1901.—Ueber die Bildungsweise des dunkeln Pigments bei den

Mollusken, nebst Bemerkungen tiber die Entstehung von Kalkkarbonat.

Ber. naturf. Ges. Freiburg, Bd. xi. 1899-1901, pp. 40-45.

(44) Srempevy, W. 1898.-—Beitrage zur Kenntniss der Nuculiden. Zool. Jahrb.,

Supplement-Band iv. : Fauna Chilensis, Bd. i. pp. 339-430, Taf. 22, 25. |

(45) SrempPett, W. 1900.—Ueber die Bildungsweise und das Wachstum der

Muschel- und Schneckenschalen. Biol. Centralblatt, Bd. xx. Nos. 18-

22. 1900. ik

(46) Tutere, J. 1893.—Beitrage zur Kenntnis der Mollusken.—II. Uber die Mol-

luskenschalen. Zeitschr. f. wiss. Zool. Bd. lv. 1893, pp. 220-250, Taf. xi.

(47) Tuxtpere, T. 1882.—Studien tiber den Bau und das Wachsthum des

Hummerparzers und der Molluskenschalen. Kongliga Svenska Vetensk.-

Akad. Handlingar, Bd. xix., No. 3, 57 pp., 12 Taf. 1884.

(48) Wirtry, A. 1907.—Report on the Window-Pane Oyster (Placuna placenta,

“Muttuchchippi”) in the Backwaters of the Eastern Province (June

1907). Spolia Zeylanica, vol. v. part xvii., Nov. 1907, pp. 33-56.

(49) Wriuirey, A. 1909.—Report of the Marine Biologist for 1908. Ceylon

Administration Reports for 1908, Colombo, p. G. 1.

(50) Lease of the Ceylon Pearl Fisheries. Ceylon Sessional Papers, x1., 1907.

Fig.

Fic.

DR. H. LYSTER JAMESON ON

(17) EXPLANATION OF THE PLATES.

(The magnifications quoted are only approximate.)

PrateE XXXIII.

1. Cestode larva (Tylocephalum ludificans or T. minus) in the act of migrating

through the tissues of Margarivifera vulgaris. Drawn trom one of Prof.

Herdmman’s slides. ‘The larva measured 12 mm, pa., parenchymatous

subepidermal tissue ; ¢.¢,f., connective-tissue fibres.

2. Small larva, Tylocephalum minus, measuring ‘08 mm., in the muscular

tissue of the mantle of MW. vulgaris, without a surrounding connective-

tissue cyst. Fyrom one of Prot. Herdman’s slides. m-.f., muscle-fibres in

cross-section ; ep., epidermis.

3. Part of the fibrous cyst surrounding an example of the smaller parasite

(Lylocephalum minus). In this case the parasite measured ‘08 mm. m

diameter. J.¢., liver-tubules; 7./., inner, highly nucleated layer of the cyst.

< 800.

4. Margaritifera vulgaris. pithelial cells from the muscle-attachment,

showing the connection between the cells and the muscle-fibres.

< 1000.

.4a. Margaritifera vulgaris. Muscle-attachment epithelium, showing cells

drawn out in fixation, and connective-tissue elements between their bases.

X 1200.

Poate XXXIV.

5. Mytilus edulis. Section through the posterior adductor muscle-insertion

and decalcitied hypostracum. mus., muscle-fibres; hy., hypostracum,

Which remains adherent to the muscle-attachment: epithelium, ep. ;

hy.’, segment of hypostracum, corresponding to single epidermal cell;

nac., nacreous layers of the shell, which have torn away from the hypo-

stracum; ct., connective-tissue fibres, joining the muscle-fibres to the

attachment-epithelium. Preparation XXXIX. X 500.

6. Margaritifera vulgaris, Persian Gulf. The part represented at A in

text-figure 35, showing the point of origin of a new lip-lappet of the shell.

1., 1.’, two successively formed lappets or marginal processes; pr., pr,

prisms of same; s.pr., s.pr.’, conchyolin-septa of the prisms; b.m.pr.,

conchyolin basal membrane of the prismatic substance of the first lappet ;

per., periostracum of the second lappet, which appears to be nothing

more than the outer conchyolin-membrane of the prismatic substance ;

x, point at which the second lappet arises from the first, the peri-

ostracum of the seeond lappet being continuous with the basal

membrane of the first; y, irregular prismatic substance secreted at the

junction of the two lappets. Preparation X. X 300.

Fic. 6a. Margaritifera vulgaris, Persian Gulf. Decalcitied shell showing junction

g g J J

Fig.

between prismatic substance and nacre. pr., prisms; sep., conchyolin-

septa between same; sep.’, the same in surface view; anuw., annular

thickenings of the septa; p7r.b., basal conchyolin of the prismatic layer ;

con., connecting layer of alveolar conchyolin between the prisms and the

nacre; nac., nacre. Prepayvation XI. X 400.

7. Margaritifera vulgaris, Persian Gulf. “ Lingah Shell” from London

mariet. Section ground from an old-heavy shell. Portion of the adductor-

scar, showing the hypostracum, /y., covered over in the direction of the

umbo by nacre, zac.’ ; wac., nacre underlying the muscle-scar, and secreted

before the muscle-attachment had moved to this place ; hy.’, hy.'’, extensions

of the hypostracum interstratified. with nacre. Preparation LXXV.

X 120. -

Pratt XXXV.

8. Muscle-pearl and hypostracum “cyst,” from the insertion of the posterior

pedal levator muscle of Margaritifera vulgaris. Froma Ceylon specimen

given to me by Professor Herdman. The pearl is enclosed in a sac (which

has been ruptured by gas-bubbles in the decalcifying process), consisting

mainly of ordinary shell-secreting epithelium, but lined at one pole with

muscle-attachment epithelium (m.ep.’). Opposite the ordinary epitheliam

THE CEYLON PEARL-OYSTER. 353

the pearl consists of typical coats of nacre; but at the point where the sac

is lined with attachment-epithelium the pearl consists of hypostracum,

which also (or a substance analogous to it) lines the central cavity. Beside

the pearl, at ¢. is a cyst-like hypostracum-pearl, comparable to the lining

of the adjoining nacreous pearl. This hypostracum-pearl consistsof a cavity,

lined with hypostracum-like substance (hyp.’), and on two sides it shows

muscle-attachment epithelium, m.ep.’, but it has not acquired a lining

of ordinary nacre-secreting epithelium, and hence no nacreous coats

have arisen. c., cyst-like cavity of hypostracum-pearl ; ep., shell-secreting

epidermis ; ep.’, epidermis of pearl-sac ; m.ep., muscle-attachment epidermis ;

m.ep.’, do. of pearl-sac; m.ep.’’, do. of hypostracum-pearl ; m., muscle-

fibres; par., parenchyma; hyp., hypostracum which lines the muscle

pearl; hyp.’, do. of the hypostracum-pearl. Preparation IV,9. XX 100.

Fig. 9. A muscle-pearl in the free mantle, froma slide in Prof. Herdman’s collection.

Fig.

ie. 10.

ig. 13.

ig. 14.

15.

17.

The area where the muscle-attachment epidermis is attached to the pearl

is relatively very small. The greater part of the pearl-sac is composed of

ordinary nacre-secreting epidermis, and consequently the pearl is composed

of nacre at all parts but one. o.ep., outer shell-secreting epidermis ;

i.ep., inner ciliated epidermis of the mantle-cavity ; par., subepidermal

parenchyma-cells, surrounding the pearl-sac ; m.ep., muscle-attachment

epithelium, where hilum of pearl was attached ; musc., muscles of mantle ;

sac., pearl-sac ; nae., nacre; rep.nac., granular repair-nacre, around central

part of pearl; hyp., hypostracum. X 70.

PuatE XXXVI.

An early stage of a muscle-pearl, from one of Prof. Herdman’s slides.

sac., pearl-sac; ., nacre; par., parenchyma; m., muscle-fibres; hyp.

hypostracum-like cyst, around which the pearl is formed ; m.ep., muscle-

attachment epithelium; c., a hypostracum-pearl. X 65.

. A cyst-like hypostracum-pearl in the muscle-bundle, without any apparent

epithelial sac. From one of Professor Herdman’s slides. ‘The cyst adjoins

the wall of a sac, ep.p.s., in which a muscle-pearl was contained. hy.,

hypostracum ; m., muscles ; par., parenchyma ; 0.ep., outer shell-secreting

epidermis; ep.p.s., epidermis of pearl-sac. X< 400.

. Centre of the decalcified muscle-pearl which occupied the sac adjoining the

hypostracum-pearl shown in fig. 11. ¢., central cavity; g7., granules in

central cavity ; 2., nacre. > 500.

Centre of a decalcified pearl, close to the mantle-margin, in one of Prof.

Herdman’s slides. ¢., central cavity ; »., nacre ; rep.m., Yepair-nacre with

radial reinforcements ; /.c., lining of the central cavity. > 300.

PLatTE XXXVII.

Four pearls “in ovary” of Margaritifera vulgaris, from Dr. Kelaart’s

material in the British Museum. The pearls are close together as shown,

but their centres have been brought into the same plane in the drawing to

show the “nuclei.” Each pearl is surrounded by a mass of subepidermal

parenchyma, par. ep., outer epidermis; ov., ovarian tubules ; m., muscle-

bundles. Preparation XXVII, 8. ~X 30.

The centre of the pearl shown in the top left-hand corner of fig. 14.

e., central cavities ; nw., nucleus-like matter, probably of the nature of

conchyolin, contained in same; rep.n., repair-nacre, with granular structure,

tollowing upon the cavity; »., ordinary nacre of the pearl. Preparation

XXVII, 8. x 400.

. The central portion of the pearl shown on Plate XLIV. fig. 48. 2.. central

nacre; m.’, peripheral nacre; c¢., central cavity; g7.l., granular layer,

causing the opaque yellowish appearance of the central mass.

Preparation LXXI sg. xX 70.

PuatE XXXVIII.

A portion of the granular layer of the pearl shown in fig. 16, more highly

magnified. ., central nacre; m.’, peripheral nacre; g7., granular layer,

suggesting dead cells ; am.,am.’, two layers of amorphous horny substance.

Preparation LXXI gs. xX 400,

Proc, Zoou. Soc.—1912, No. XXIII. 23

3504. DR. H. LYSTER JAMESON ON”

Fig. 18. The centre of the same pearl, consisting of a cavity, about *08 mm. in

diameter, surrounded by granular repair-nacre; the cavity contains a few

nregular strands or lamine of conchyolin-like substance, together with

certain granular bodies, possibly of cellular origin. Preparation LXXI vg.

x 450.

Fig. 19. Group of muscle-pearls and hypostracum-pearls, from an old unlabelled

example of Margaritifera vulgaris in the British Museum. ‘The centres

of the pearls have been brought into the same plane in the drawing.

The pseudo-nuclei range from ‘08 mm. to °15 mm. in diameter. At the

bottom and on the left-hand sides of the sketch are two pearls with

eccentric pseudo-nuclei, in these cases composed of hypostracum,

showing that the nacre-secreting epithelium arose on one side of

the sac only. The preservation does not allow of any histological

details of the tissues being shown. Four hypostracum-pearls are also

shown. nae., nacre; nw., columnar pseudo-nucleus; nz.’, hypostracum

pseudo-nucleus, only partly surrounded by nacre; /y.p., hypostracum-

pearls: muse., muscle-bundles. Preparation XXVII, 4. X 30.

HAUTE) NOXGNGIEXS

Fig. 20. Centre of the pearl shown on the upper side of fig. 19. mac., nacre; col.,

columnar substance ; am., amorphous substance; g7., granules in central

cavity. Preparation XX VII, 4. > 350.

Fig. 21. A hypostracum-pearl, decalcified. gi., granules in central cavity; am.,

amorphous substance; /y., organic residue of the hypostracum.

Preparation XXVII,1. > 50.

Fig.21a.Part of a section ground through the middle of a hypostracum-pearl,

showing the columnar and stratified nature of its substance, and the small

central cavity. From an unlabelled specimen in the British Museum.

Preparation LXXVIII c. xX 70.

Fig, 22. Portion of the mantle of Margaritifera vulgaris, near the margin, from a

second unlabelled specimen in the British Museum. Cleared in oil of

cloves, aud examined entire before decalcification. ‘The specimen shows a

number of hypostracum-pearls iz situ. At nac.is seen a large hypostracum-

pearl in course of transformation into a nacreous pearl by the deposition

of nacreous layers around it. Preparation XXIX. X 20.

Fig. 23. The same decalcified and examined in oil of cloves. mac., as above. X 20.

Prave XL.

Fiz. 24. A double nucleus, with abundant contents, from a pearl in the same

series of sections as figs. 19-21. gi, granular substance, apparently

derived from dead organic matter; a@m., amorphous cuticle-like layers ;

am.’, a more deeply staining portion of the same substance, immediately

surrounding the granular contents of the nucleus ; col., columnar substance ;

nac., nacre. Preparation XXVII,7. X 250.

Fig. 25. A small hypostracum-pearl. Central cavity without contents. Preparation

XXVIII, 11. x 100.

Fig. 26. A double hypostracum-pearl, forming the nucleus of a nacreous pearl.

am., (2) amorphous substance; hy., hypostracum ; hy.’, do. at periphery of

amorphous substance; nac., nacre; nac.’, columnar repair-substance.

Preparation XXVII,11. xX 75.

Vig. 27. Small pearl, with nacreous substance overlain hy hypostracum. ¢., central

cavity, containing afew granules; am., amorphous substance which passes

over into col., columnar substance; nae., nacre, which is succeeded by

hy., a layer of hypostracum, forming the outermost layer of the pearl.

This specimen shows well the difference between the reaction to stain

of hypostracum and the other substances, e. g. columnar substance.

Preparation XXVIJI,7. X 350.

Fig. 28. Pseudo-nucleus of the left-hand constituent of the double pearl shown in

fic. 49 (Plate XLIV.). From one of the unlabelled specimens of

Margaritifera vulgaris in the British Museum. c., central mass composed

of (?) amorphous substance and granules; col., columnar substance ;

col.’, outer layers of the same, which pass over imperceptibly at one point

into nae.’, nacreous substance; nac., complete layers of nacre around

nucleus; am., horny layer, probably composed of amorphous substance :

THE CEYLON PEARL-OYSTER,

col.”, incomplete columnar layer; nac.'’, nacreous substance of the pearl.

Preparation XLIII B. xX 75.

Margaritifera vulgaris, Persian Gulf. Portion of the repair-substance

formed inside the shell to close a large hole made by a borer of some kind

in the lateral region, below the umbo. am., amorphous substance;

y, outer surface of repair-membrane ; a, a’, cracks in the outer face of the

amorphous substance; col., columnar repair-substance; col.’, stratified

do.; tr., ér.’, transitional substance (granular repair-nacre) from columnar

repair-substance to nacre; mac., nacre. Preparation LXXVI. x 400.

Fig. 29a. Margaritifera vulgaris, Persian Gulf. From a section through a blister

Fig. 30,

Fig. 31.

Fig. 32.

Fig. 33.

Fig. 34.

Fig. 35.

Fig. 36.

Fig. 37.

on the shell-margin, formed over a Lewcodore tube. am., amorphous

repair-substance; a@/v., the same with alveoli containing carbonate of

lime; pr., prismatic substance. Preparation XI. X 500,

Prats XLI.

Margaritifera vulgaris; Dr. Kelaart’s material. Sections showing junc-

tion between two of the constituents of a “multiple pearl,’ with the

various abnormal products which arose in association with the disturbances

caused by the fusion of the two pearl-sacs, and the retreat of the wedge-

shaped plug of tissue when its connection with its fellow of the opposite

side was broken down by the absorption or degeneration of the membrane

separating the two pearls. p., p.’, outer nacreous layers of the two pearls ;

rep.nac., granular repair-nacre; col., simple columnar substance, analo-

gous to that found in the nuclei of some pearls; am., amorphous

substance showing rough stratifications, perhaps corresponding to pauses

in the retreat of the wedge of tissue; ¢., transition from amorphous

substance to columnar substance; ¢7.’, transition from amorphous sub-

stance to granular substance ; nac.’, nacre of the compound pearl embracing

both constituent pearls. Preparation LXXIc. X 300.

Margaritifera vulgaris. Unlabelled specimen, British Museum. One end

of the suture between the fused pearls shown in fig. 49, Plate XLIV.

nac., nac.’, the nacreous layers of the two pearls; col., col.’, columnar

substance, secreted after the epithelium had, through atrophy, lost its

power of producing nacre; ep., ep.’, the degenerated remains of the

epithelia of the pearl-sacs ; par., the remains of the intervening paren-

chyma. Preparation LXIII Bp. X 350.

From the middle of the same suture; in this case the outlines of the

parenchyma-cells, par., are still visible. ep., remains of the epithelia of

the pearl-sacs. XX 350.

Parenchyma-pearl in situ in the tissues of Margaritifera vulgaris, from

the Persian Gulf. _ep., outer shell-secreting epithelium ; ep.’, epithelium of

pearl-sac ; par., subepidermal parenchyma ; g/., gland-cells in same ; musc.,

muscular coat of the visceral mass ; mause.’, musculature to wall of stomach :

C.T., connective-tissue of visceral mass; ov., tubules of ovary; col., columnar

substance forming the centre of the pearl; nae., nacreous portion of the

pearl. Preparation LXIV s. xX 80.

Centre of a pearl from another example from the Persian Gulf. gi., granular

matter; am., (?) amorphous substance ; nac., nacreous layer, immediately

surrounding the “ nucleus”; nac.’, outer nacre; col., columnar substance ;

col.’, imperfect layer of same, passing over laterally into nacre. Preparation

LXIV c. X 250.

The central, spherocrystal-like pseudo-nucleus of the pearl shown in figs.

50 & 50 a (Plate XLV.) ; from the sample of pearls purchased in Ceylon.

grm., granular repair-nacre ; am., amorphous substance. Preparation

XLII. 400.

ACD eal:

The pseudo-nucleus of the pearl shown in fig. 52 (Plate XLV.) ; from series

purchased in Ceylon. gr., granular substance in central cavity forming

the true nucleus; am., amorphous substance ; nw., pseudo-nucleus, which

here possesses a highly alveolar structure; col., columnar repair-substance

passing over laterally into mae., nacre. Preparation LIII. > 70.

Alveolar columnar repair-substance and radially reinforced nacre, from the

pseudo-nucleus surrounding a sand-grain (or other hard body that broke

away) in a pearl from the same series. am., amorphous substance, forming

23%

356

Fig.

Fi:

38.

39.

ig. 40.

ig. 4d.

ig. 42.

43.

44,

DR. H. LYSTER JAMESON ON

the innermost layer of the columnar pseudo-nucleus, col.; nac., nacre ;

nac.’, radially marked, catenuiated variety of nacre, characterised by

radial rows of junctions which occur in small groups between conchyolin-

layers (june.); nae.’’, the normal nacre, passing over into this specialised

variety of nacre. Preparation LIV G@. X 300.

Tangential section through the alveolar-columnar substance shown at

col, in fig. 87. X 300.

Tangential section through the radially catenulated repair-nacre, shown at

nac.’ in fig. 37. junc., the groups of junctions in radial rows, seen in

surface view. The section is near the inner surface of this layer. X 300.

Centre of the imperfcetly decalcified brown pearl, shown in Pl. XLVI.,

fig. 57. nu., nucleus with rhombohedra of calcite ; am.’, lining of

amorphous substance; col., columnar layer; col.tr., transition-layer from

alveolar-columnar substance to amorphous substance; am., amorphous

substance, with scattered alveoli, containing calcium carbonate. Pre-

paration XL. X 2650.

Part of the same pearl, external to that shown on fig. 40, showing the

transition from amorphous substance, am., with scattered alveoli, through

an intermediate substance, ¢7., to needle-like prismatic substance, col.

x 250.

Transition from the needle-like prismatic substance, col., shown in fig. 41,

to the ordinary prismatic shell-substance, pr., similar to that of the shell.

al., minute alveoli in the thickenings of the walls of the prisms, similar

to those observed by Romer (32). X 250.

PrarE XLIIL.

A portion of the repair-substance in the body of a pearl, shown at col.

in fig, 86, showing transitions from amorphous substance to nacre.

nu., outer part of the pseudo-nucleus; am., amorphous substance, with

alveoli, which passes over into finely columnar substance, col. This

in turn goes over through granular repair-nacre, gr., into normal nacre,

nac.’, and is followed by layers of normal nacre (nac.) externally. Pre-

paration LIIT. X 400. :

Portion of the contents of the central cavity of one of the pearls purchased

in Ceylon; showing also a portion of the columnar substance which

surrounded the centre. The contents drawn are selected from a number

of the sectioas in the series cut from this specimen. mac., nacreous

substance of the pearl; col., columnar repair-substance surrounding the

nucleus; am.,@m.’, amorphous substance; dia., diatoms; spic., fragments

of sponge-spicules ; veg., vegetable débris. Preparation LIV 5. X 500.

5. Fragment of a Radiolarian shell, from the centre of another of the same lot

of pearls. Preparation LIV £. X 600.

Prare XLIV.

s, 46, 46 a. Pearl from a cluster of pearls in one of Dr. Kelaart’s specimens.

Decalcitied and examined, 4.6, entire in oil of cloves, 46 a, in section. The

diameter of the pearl was just 2 mm., that of the central denser part was

13mm. There is a simple central cavity. Preparation XVI. X 27.

Vigs. 47, 47 a. Another pearl from Dr. Kelaart’s material, decalcified and examined,

47, entire, and, 47 a, in section. The diameter of the pearl was 2mm. ; that

of the dark, opaque, yellowish centre rather over 1 mm. Examined entire

the nucleus appears amorphous, and might well be mistaken for the dead

remains of a parasite; but this peculiarity was found, on sections being

made, to be due to a break in the continuity of the nacreous layers

and the interpolation of a dark layer, composed of irregularly calcified

conchyolin, and granules which are probably of extraneous origin. ‘There

is a small central cavity, as in the rest of Dr. Kelaart’s pearls, with a plug

of conchyolin-like substance at one side. g7., granular dead matter; col.,

columnar substance; , nacre. Preparation XV. X 27.

Vig. 48. A “double” pearl from Dr. Kelaart’s material, decalcified, cleared, and

examined entire. Sections through this pearl are shown on Plates

XXXVII. & XXXVIII. figs. 16-18. The pearl was 2°25 nm. in diameter.

The pseudo-nucleus was a dark opaque body, about 0°5 mm. in diameter,

similar to that shown in fig. 47, and might easily be taken, on examination

ot the entire pearl, for the shrivelled remains of a dead parasite. Sections

THE CEYLON PEARL-OYSTER. ao

showed that here again the real nucleus was a cavity containing strands

or sheets of conchyolin-like substance and a few granular bodies, perhaps

of cellular origin (Plate XXXVIII. fig. 18); while the dark appearance

of the central mass was due to the interpolation between the proper

nacreous layers ofan abnormal pathological product. Preparation LXX Iv.

xX 27.

Fig. 49, A double pearl from one of the unlabelled specimens in the British Museum;

decalcified, cleared, and examined entire. The nucleus of the left-hand

constituent is figured on Pl. XL. fig. 28, the characters of the substance

at the suture between the two pearls at figs. 31 & 32 on Plate XLI. In

the pseudo-nuclei of these pearls the columnar substance is stratified.

Preparation LXIII sp. X 27.

PLATE XLY.

Figs. 50, 50 a. The central portion of a white porcellanous spherical pearl, from the

sample of pearls purchased for me in Ceylon by the Ceylon Company of

Pearl Fishers, Ltd. Fig. 50, decaicitied and examined whole in oil of

cloves. Here the nucleus, when examined whole, might well be mistaken

for a small Trematode. Fig. 50 a. Section through the centre of the same

pearl. ‘The pseudo-nucleus is seen to cousist of an irregular nacreous body,

shown in fig. 35 (Plate XLI.) to be granular repair-nacre, surrounding a

minute sphwrocrystal-like mass; about 0°08 mm. in diameter. Pro-

paration XLIII. x 27.

51. Another pearl from the same collection. An oval pearl, of fine quality,

35 mm. X 3mm. in diameter. This pearl shows a tract of stratified

columnar and granular repair-substance, extending outwards from the

pseudo-nucleus and passing over laterally into the nacre, Preparation LI.

X 20.

Fig. 52. Another pseudo-nucleus, from a pearl in the same collection. This was a

small spherical pearl of fine quality, about 2 mm. in diameter. The

columnar substance here, col., is alveolar in structure, and surrounds a

central cavity containing afew granules. Fora section of the centre of this

pearl, see fig. 36 (Pl. XLII.) Preparation LIII. X 27.

Figs. 53, 53a. Centre of another pearl from the same series. The pearl was oval,

about 3 mm. in diameter, with a faint zonar constriction and a rather

coppery tint. Examined entire (fig. 50) it shows a large pseudo-nucleus,

over a millimetre in diameter, which falls into three layers. It also shows

a blemish in the nacre, coated over with repair-substance. Fig. 53 a

shows the same in section, The columnar repair-substance is finely

reticulated or alveolar. The blemish is caused by a few granules, which

have found their way into the pearl-sac and have been covered over with

columnar repair-substance, which passes over laterally into ordinary nacre,

Preparation LIV. xX 27.

Figs. 54,54 a. Fig. 54. Central portion of another pearl, which measured about

3mm. X 2°75 mm. _ A pearl of fine colour, but with slightly irregular

surface. he nucleus of this pearl is a sand-grain, enclosed in an opaque

yellowish coat, probably consisting of repair-substance. At one pole is

seen a process of repair-nacre, but, except for this, no columnar substance

appears to surround the grain. Fig. 54a. The sand-grain, dissected out

from the above preparation. Preparation LII. X 27.

PratE XLVI.

. 55. Fine spherical pearl, from the same series, about 2 mm. in diameter.

Nucleus a grain of sand surrounded by a thin layer of columnar substance,

thickened at one pole. Preparation LIV u. X 27.

Fig. 56. Another pearl from the same series, a small spherical pearl about 2 mm. in

diameter. ‘The nucleus is a grain of sand, forming the centre of a typical

pseudo-nucleus of columnar substance. Preparation LIV a. xX 27.

Fig. 57. A brown pearl, spherical, 3 :mm. in diameter, formed of the prismatic

substance. Fromthesame collection. nw., pseudo-nucleus ; co/., columnar

substance ; am., amorphous substance; col.’, fine columnar-prismatic sub-

stance ; pr., ordinary prismatic substance ; 6/., blister formed over intrusive

foreign matter. For the several substances of this pearl, more highly

magnified, see Plate XLII. figs. 40-42. Preparation XL. X 27.

358 _ MR. ROBERT SHELFORD ON

Fig. 58. Tylocephalum ludificans, sp.n. Type. Section through an example in the

tissues of the Ceylon Pearl-Oyster in Professor Herdman’s collection.

Slide 94 of Professor Herdman’s series. a@., armature of collar. This shows

the myzorhynchus of “ T'ylocephalum” form, due to the posterior face

(m.p.) being contracted and its surface thrown into folds and the anterior

tace (m.a.) stretched. Compare the adult Tylocephala im figs 61 & 62,

and also the species shown in fig. 65. X 70.

Fig. 59. Section through another example of the same species, showing the “ Cepha-

lobothrium’’-form of myzorhynchus; here the posterior surface of the

myzorhynchus (m.p.) is tense, and its anterior surface (m.a.) is thrown into

folds. a, armature of collar. (Compare theadult worm in fig. 66.) X 70.

Fig. 60. An adult worm, which may be the adult of TLylocephalum ludificans,

from the spiral intestine of Aétobatis narinari. X 6. From Dr. Shipley’s

collection.

Prare XLVII.

Fig. 61. The head of the worm shown in fig. 60, viewed as a transparent object.

m., myzorhynehus; s,s’, s’’, three of the four marginal suckers; a., armature

of collar ; seg., first proglottides. X 70.

Fig. 62. The same, in section; letters as above. X 70.

Fig. 638. Armature of collar of same, about point a in fig. 62. XX 1000.

Fig. 64. Penultimate segment of same.

Fig. 65. Head of Tylocephalum uarnak from Trygon warnak. From a slide in

Dr. Shipley’s collection. my., myzorhynchus, which here is fully protruded ;

ma., anterior face of same; mp., posterior face; s, s’, s’’, s’’’, the four

marginal suckers. X 46. [

Vig. 66. Head of Cephalobothrium aétobatides from Aétobatis narinari. From a

specimen in Dr. Shipley’s collection. Letters as above. X 70.

19. Mimicry amongst the Blattidee ; with a Revision of the

Genus Prosoplecta Sauss., and the Description of a new

Genus. By Roperr Saetrorp, M.A., F.Z.5.

[Received October 7, 1911: Read February 20, 1912. |

(Plate XLVIII.* and Text-figures 42-46.)

Speaking in general terms the Blattidee may be regarded as a

somewhat defenceless group of insects, preyed upon by numerous

enemies both vertebrate and invertebrate, to escape which they

must rely on their skulking, cryptic habits, and on a high degree

of speed when disturbed. In accordance with their cryptic mode

of life most cockroaches are obscurely coloured, harmonising

more or less perfectly with their inanimate surroundings, or, at

any rate, displaying no conspicuous markings to attract the

attention of potential foes.

Exceptions to this cryptically coloured type of cockroach, how-

ever, exist In some numbers. and are dealt with in the following

pages.

The Australian continent is the headquarters of a group of

Blattidee which presents all the features customarily associated

with extreme unpalatability. This group is the Polyzosteria

section of the sub-family Blattine. Nearly all the species are

apterous and expose themselves freely, and many are endowed with

a most repulsive odour. Mr. W. W. Froggatt informs:me that

Polyzosteria limbata Burm., P. cuprea Sauss., and others of this

* For explanation of the Plate see p. 376.

Horace Knight del.et lith

MIMETIC COCKROACHES AN]

D BEETLI

West, Newman chromo

EF MODELS.

MIMICRY AMONGST THE BLAT'TIDA, 359

genus and of Platyzosteria Br., ave fond of sunning themselves

on the tops of posts and tree-stumps. Commander J. J. Walker

when collecting in Australia frequently encountered a species,

Cosmozosteria lateralis Walk., which emitted so vile a smell that

he always refrained from touching it. ‘This species is russet-

brown in colour with some aviable yellow markings on the

thoracic and abdominal tergites, and on each posterior angle of the

ninth abdominal tergite is a brilliant orange-red spot ; when the

insect is at rest these two spots are almost concealed, the ninth

tergite being somewhat retracted within the preceding one, but

on the approach of an enemy the apex of the abdomen is elevated —

and slightly distended so that the orange spots become conspicu-

ously displayed to view. <A better example of a warning signal

associated with highly distasteful properties could not be found.

Prof. Baldwin Spencer, writing in ‘ Nature’ of July 28, 1892,

p. 309, says :—‘‘ One morning, when Mr. Frank Connelly and

myself were digging for worms, we accidentally cut in two a

cockroach. From between the segments in its back it poured

forth a milky-white fluid, possessing an odour so execrable ana

pungent that it drove us from the spot.” It is untortunate tl at

the species was not identified, but I expect that it was one of the

Polyzosteria group.

Commander J. J. Walker says of Periplaneta fortipes Walk., 2

synonym of Platyzosteria nove-zealandiw Br., that it is “ very

evil-smelling,’ but that its smell ‘ is quite mild in comparison

with several of the Australian species.” (Kntom. Monthly Mag.

(2) xv. p. 70 (1904).)

Dr. G. B. Longstaff recently took this species in some numbers

in New Zealand, finding it, however, not in exposed situations,

but under logs and the bark of dead trees. Quoting from his note-

book Dr. Longstaff tells me that one specimen had a “ moderate

cockroach odour, evanescent,” of another that it had “a strong

peculiar fetor.” The discrepancy may possibly be accounted for

by sexual differences (see remarks later on Hurycotis floridana

Walk.).

All the Australasian species of the Polyzosteria section appear

to be conspicuous insects. Many of them are shining black, a

colour which is noticeable enough in Nature when associated

with free exposure ; others are black edged with yellow, or with

red legs. The species of Anamesia Tepp., are chestnut-brown

banded or margined with yellow. Cosmozosteria zonata Walk.,

is black, banded with yellow or orange. Many of the species of

Polyzosteria Burm. ave bright with metallic colours, and Hu-

zosteria mitchelli Angas, with its bronzy dorsal surface, spotted

and barred with orange or yellow, its pale yellow ventral surface

and sky-blue tibize, is the most gaudy cockroach yet discovered.

In the New World the Polyzosteria section of the Blattine is

represented by the genera Hurycotis Stal, and Pelmatosilpha

Dohrn. The species do not appear to be so blatantly conspicuous

as their Australian relatives, but it seems likely that most, if not

360 MR. ROBERT SHELFORD ON

all of them, are endowed with very distasteful properties. Rehn

and Hebard (Proc. Acad. Nat. Sci. Philadelphia, 1905, p. 32)

write of Hurycotis floridanw Walk. :—‘ When seized these insects

emit a vile-smelling oily fluid. The females always produced

far more of this than the males.” This is a chestnut-brown

species, but the larve have the thoracic tergites margined with

pale yellow; it is found hiding under logs and stones. It is

evident, then, that amongst the Blattide a nauseous odour or

taste is not mvariably associated with aposematic habits,—the

insects themselves may be conspicuous enough when unearthed

from their hiding-places, but the point is, that they do not

voluntarily expose themselves, as do so many of the Australian

Polyzosteriae. Further confirmation of these rather puzzling

facts is afforded by observations made by Mr. G. A. K. Marshall

on two South-African cockroaches, Deropeltis erythrocephala Fah.

and Aptera fusca Thunb. The first of these belongs to the sub-

family Blattinzee and to a genus in which the males are winged

and the females apterous. In both sexes the posterior margin

of the fifth abdominal tergite is sinuate *, and beneath the tergite

are situated glands from which a sticky fluid exudes when the

insects are seized. Though it is reasonable to suppose that this

fluid is a distasteful secretion of a defensive nature, it must again

be noted that D. erythrocephala and probably most of the other

species of the genus hide beneath stones and do not expose them-

selves voluntarily. Most of the species of Deropeltis are bulky

insects, piceous in colour ; in D. erythrocephala the head and legs

are red. D. dichroa Gerst., from the Gold Coast, has a large

fulvous macula on each side of the sixth and seventh abdominal

tergites ; whilst D. paulinot Bol., from Angola, is equally conspi-

cuously marked with rufous fascie on the lateral margins of the

pronotum. Thefemale of Aptera fusca, one of the Perispheriine,

is a large, robust insect, piceous with conspicuous transverse

bands of ochreous or rufous ; when seized it exudes a violet fluid

which stains the fingers ; this species also has eryptic habits. This

is absolutely all the information that I have been able to gather

about unpalatable species of Blattide, and it certainly is little

enough. It is perhaps remarkable that the undoubtedly nauseous

Australian species are not mimicked either by other orders of

insects or by non-distasteful species of Blattidee, but it must be

remembered that the Orthoptera do not serve as models to other

orders of insects—not a single instance has ever been recorded,

and in Australia the paucity of cockroaches other than those of

the distasteful group, is quite exceptional.

Although no Blattidee are known to serve as models to mimick-

ing insects, there are several which mimic insects other than

Orthoptera, though in most instances the mimicry is of a very

generalised nature. I have no reason to suppose that any of

these mimetic Blattidz are other than palatable.

* A generic character.

MIMICRY AMONGST THE BLATTIDA, 361

In a very superficial sort of way cockroaches and beetles may

be said to be similarly constructed. In both the pronotum is

large whilst the other thoracic tergites (in the winged species)

are concealed; in both the membranous wings are covered by

elytra or tegmina of a coriaceous or corneous texture. In fact

only a slight modification of the cockroach-form is required to

produce a distinctly Coleopterous appearance. The names lycoides,

buprestoides, coccinelloides, dytiscoides, silphoides, given to species

of Blattidee by various authors, are sufficient evidence of their

resemblance to beetles. It is quite an open question whether

this generalised resemblance of certain Blattide to Coleoptera

can be legitimately classified under the heading of Mimicry. It

could well be argued that some of the species, at any rate, owe

their beetle-like form to convergence in development, or, to use

Sir Ray Lankester’s term, that cockroaches and beetles are homo-

plastic forms. On the other hand, as will be seen later, some of

the cases of resemblance are so detailed and close that it is

impossible to regard them as anything but examples of true

mimicry, and it becomes most difficult to draw the line between

the two classes of resemblance. For convenience’ sake, at any

rate, throughout this paper the Blattide which resemble insects

of other orders will be termed ‘“ mimics.”

Examples of generalised mimics of the Coleoptera are furnished

by species of Pachnepteryx By., Caloblatta Sauss., Paratropes

Serv., Phoraspis Serv., Hustegasta Gerst., Achroblatta Sauss.,

Corydia Serv., Areolaria Br., and Hypnorna Stil, whilst several

species in other less specialised genera might be quoted. Of not

one of these species can it be said that it is very like any definite

species of beetle. Hustegasta buprestoides Walk., from West

Africa, is a metallic green cockroach with round yellow spots on

the tegmina, and as its name implies, it is very like a Buprestid

beetle. But in spite of the most diligent search amongst col-

lections of Buprestide, I have never found a species which by

the greatest stretch of imagination could be regarded as even an

indifferent model for the cockroach.

Belt speaks of mimetic cockroaches in ‘The Naturalist in

Nicaragua’ as follows :—“ The phosphorescent species of Lam-

pyride, the fireflies, so numerous in Tropical America, are

equally * distasteful, and are also much mimicked by other

insects. I found different species of cockroaches so much like

them in shape and colour that they could not be distinguished

without examination. These cockroaches, instead of hiding in

crevices and under logs like their brethren, rest during the day

exposed on the surface of leaves, in the same manner as the

fireflies they mimic”?t. It was with much interest that I

found in the Hope Museum, Oxford, a specimen of the cock-

roach Achroblatta luteola Blanch., with the following note in

* I. e. with the non-phosphorescent species, by which Belt appears to mean the

beetles now known as Lycide.

+ I quote from the Everyman’s Library Edition (Dent & Sous, 1911), p. 243.

362 MR. ROBERT SHELFORD ON

Westwood’s handwriting attached to it :—‘“ This Blatta lives on

trees and closely resembles in its habits some of the large Lam-

pyvidee ['T. Belt].” This is doubtless one of the species mentioned

by Belt in his book, and I had great hopes of being able to match —

it with a definite species of Lampyrid beetle, but the most diligent

search through the Godman-Salvin collections of Central American

insects failed to reveal a ‘ model” to the cockroach. The Malaco-

dermatous appearance is undoubtedly but generalised, not specific.

The same may be said of Zypnorna amena Sauss. & Z., also from

Central America. This Blattid has all the appearance of a small

Longicorn, but it actually resembles no particular species of that

family. It may, of course, be argued that further collecting will

bring to light species of beetles which can legitimately be regarded

as models to the cockroaches, but I do not think that this is in

the least degree probable. Our knowledge of the Central American

Coleoptera must now be nearly complete, speaking from the point

of view of the systematist, and the West African Hustegasta bu-

prestoides is so very abundant that it might reasonably be expected

that its supposed model would be, if not abundant, at any rate in

sufficient numbers to permit of some specimens falling into the

hands of collectors. At the very end of this paper I describe

two new species of Blattide, belonging to a new genus, which

also must fall into the category os generalised ‘Coleopterous

mimics.

The two species of the Oriental genus Z'hyrsocera Burm. are

shining black cockroaches with large yellow spots on the tegmina,

a type of coloration frequently met with amongst the Endomy-

chide of the same region. These cockroaches, though far larger

than, and in other points quite unlike any Endomychide known

to science, may possibly be regarded as part of a “ convergent

group,” the dominant or “central” members of which are the

yellow-spotted Endomychide (genus Humorphus). A generalised

resemblance to certain families of Rhynchota is also shown by

some Blattide. For example, some species of Holocompsa Burm.

and Hypercompsa fiebert Br. are vather like small Capsidee ; the

illusory effect is produced by the tegmina, which are largely

membranous and hyaline, though opaque and coriaceous at the

base; the resemblance does not bear a very close examination.

Homopteroidea nigra Shelf. is not unlike some small Fulgorid or

Jassid. Mr. J. C. Kershaw found at Hong-Kong, under a stone,

several little black Pentatomids, and in company with these a

similarly coloured and shaped cockroach which appears to be

identical with Pseudophyllodromia parilis Walk., and he sug-

gested * that the cockroach mimics the bug. Having seen the

specimens I prefer to regard them as examples of syncryptism or

of homoplasy.

Linneus, deceived by the Coleopterous appearance of Coryd ia

petiverandé, placed it in his genus Cassida. My. T. Bainbrigge

gse

* Ina letter to Professor Poulton.

MIMICRY AMONGST THE BLATTID”, 363

Fletcher, who has watched this insect in a state of nature in

Ceylon, tells me that when it is flying it looks very like a con-

spicuous Agaristid moth, AZimeusemia ceylonica Hmpsn. The

resemblance is certainly not very striking when the dried insects

are seen side by side in a cabinet, but no field-naturalist will

attach very much importance to that, and in any case Mr. Fletcher

does not maintain that the resemblance is detailed and accurate,

but merely generalised,

The power which the females of species of Perisphaeria and

Pseudoglomeris have of rolling themselves up into spherical balls

when alarmed is well known, and on account of their convex

form and black shining colour, they undoubtedly bear an ex-

tremely close resemblance to the pill-millipedes which are so

abundant in the tropics. But here again I doubt if any pavrti-

cular species of millipedes are copied. It is certainly a fact that

whilst two species of Perisphaeria were not infrequently met with

in Sarawak, both rather small, black species, I never once found

a millipede corresponding in size or colour to them. It is by no

means certain that the pill-millipedes are distasteful animals—on

the contrary, it is quite probable that they are palatable but well

protected by their hard integuments and power of rolling up into

a ball. The same habit is shown by many terrestrial Isopoda,

but no one considers that the Isopods mimic the Muillipedes or

the Millipedes the Isopods. The similarity of habit and form is

attributed to homoplasy, and I see no reason why the same habit

of the cockroaches should not also have been quite independently

evolved.

Having now passed in rapid review the principal genera of

Blattidee which show a more or less generalised resemblance to

insects of other orders, it only remains to consider in greater

detail the genus Prosoplecta Sauss., nearly all the members of

which present a remarkably close al detailed resemblance to

definite specific models amongst the Coleoptera, so far as these

have been discovered. With but two exceptions the species of

Prosoplecta present an appearance which is conveniently sum-

marised as Coccinelliform ; that is to say, the outline of the body

is oval verging on spherical, the form is markedly convex, the

integuments are smooth and nitid, the tegmina are corneous

with obsolescent venation and do not extend beyond the apex of

the abdomen, the legs and antenne are short and, finally, the

insects are gaily coloured. It is scarcely necessary to point out

that the Coccinelliform type is found amongst other families of

Joleoptera besides the Coccinellide : it is found, for example,

amongst the Cassididee, Chrysomelide, and Gulerucides, whilst

many "of the Scutelleride, a family of Hemiptera, also present

much the same facies. The two species, P. coccinella Sauss. and

P. bipunctata Br., ave, in spite of the name of the first, far less

Coccinelliform than the other species of the genus, and may

certainly be regarded as more primitive. The form is more

depressed, and I am inclined to suppose that these two species,

364 MR. ROBERT SHELFORD ON

though distinctly Cole opterous in appearance, fall into the category

of generalised beetle-mimics.

Hemithyrsocera sp.

Wing expanded (a) and folded (6). The apical triangle is shaded.

In the genus Prosoplecta the wings have been modified in a

very remarkable manner, and as their structure has never been

properly described and is of particular interest when considered

together with the mimetic resemblances of the genus, it is

necessary to go now into some details of the cockroach wing-

structure. The wing of a cockroach such as blattella germanica

L., is divided longitudinally into an anterior and a posterior part

the anterior part is more or less rigid, being strengthened by

numerous longitudinal and transverse veins, while the larger

posterior part is supphed only with radiating longitudinal veins.

When the wing is closed the posterior part shuts up like a fan

and folds underneath the anterior part, which remains stiff and

unfolded. At the apex of the line of division between the

anterior and posterior parts of the wing is a quite inconspicuous

area Which, when the wing is closed, appears as a minute fold

lying on the top of the anterior part. ‘This insignificant area,

which belongs neither to the anterior nor to the posterion part

of the wing, is the forerunner of a part of the wing, which in

some genera of Blattidz assumes relatively enormous proportions.

In the genus Hemithyrsocera (text-fig. 42) this area has in-

creased in size and is conspicuous enough to have attracted the

attention of systematists, ever on the look out for characters

diagnostic of the difficult genera of the subfamily Pseudomopine.

The area is now known as the triangular apical area, or more

MIMICRY AMONGST THE BLATTIDA, 365

simply, the apical triangle, In the genera Hetobius, Thegan-

opteryx, and Chorisoneura, to select but three examples from

many, the apical triangle is still larger, has clearly defined

boundaries, and in the closed wing appears either rolled up in a

Text-fig. 43.

Chorisoneura taeniata Sauss. & Z.

Wing expanded (a) and folded (4). The apical triangle is shaded.

Text-fig. 44.

Anaplecta decipiens Sauss. & Z.

Wing expanded (a) and folded (4). The apical area is shaded.

spiral (Hetobius and Theganopteryz) or as a doubled fold (Choriso-

neura), lying on the top of the anterior part of the wing (text-

fig. 43). Between this stage of wing-development and that

366 MR. ROBERT SHELFORD ON

of the genera Anaplecta and Plectoptera there is every grade of

transition. Ina typical species of Anaplecta the apical triangle

has become a large parabolic area, now known as the apical area,

almost equal in size to the rest of the wing, over which it 1s

doubled back when the wing is folded (text-fig. 44). In the

species Anaplecta variipennis Shelf., the passage from an apical

triangle to an apical area can be traced; for some individuals

exhibit the OChorisoneuwra type of wing-structure ; in others is

seen the extension of the apical triangle by the widening out of

the angle made by the intercalation of this area between the

anterior and posterior parts of the wing; finally, some specimens

exhibit the complete development of the apical area, with its

straight base-line, cutting the wing transversely into two halves,

so that the longitudinal veins of the anterior part of the wing

and the first axillary veins of the posterior part, no longer impinge

on the outer margin of the wing, but on the base-line of the

apical area. The summit of wing-development is attained by

Diploptera dytiscoides Serv., for in this species the apical area is

closely veined, though the veins are not in organic continuity

with those of the other part of the wing.

Text-fig. 45.

Prosoplecta nigrovariegata.

Wing expanded (a) and folded (4). The pseudapical area is shaded.

A cursory glance at the closed wing of any species of Proso-

plecta (text-fig. 45) leads the observer to suppose that it 1s

constructed on the same principle as that of Hetobius and Thegan-

MIMICRY AMONGST THE BLATTID®, 367

optery, for there is visible a large curled-up spiral lying on the

dorsal face of the anterior part of the wing. But when the

wing is spread out it is seen that the method of wing-folding

is quite peculiar. The triangular apical area though | large is

not sharply defined as in Chorisoneura, Thegamopteryse, ete., and

the spiral fold involves not only this area but also the apical

portion of the anterior part of the wing, and a minute portion

of the apex of the posterior part. The part of the wing involved

in the spiral fold constitutes in fact a pseawd-apical area, and it is

suffused with a dark fuscous colour, just as the true apical area of

Anaplecta and the apical triangle of Chorisoneura ete. are coloured

more deeply than the other parts of the wing.

There can be little doubt that this type of wing-folding, unique

amongst the Blattide, is a modification brought abate by the

mimetic resemblance, ‘and it is certainly a very singular one.

The student of the mimetic resemblances which exist between

insects not genetically related, frequently comes across remarkable

modifications of structure, modifications which are alien, so to

speak, to the constitution of the insects; here a concentration of

pilosity to form a false spine, there a patch of colour, a shortening

of elytra, or a thickening of legs or antenne. But invariably fe

will find these are modifications of structure absolutely essential

to the success and perfection of the mimicry ; if he looks deeper

he will find that mimicry has not touched parts which are

concealed, and which therefore play no part in the mimetic

resemblance. In short, mimicry is essentially a superficial likeness

between organisms fundamentally different. Yet here in the

species of Prosoplecta we have an exception to this very g general

rule. The wing of the cockroach does not mimic the wing of the

beetle, but it has been modified because of the mimicry between

the two orders of insects, or in other words, in this case mimicry

has affected structures which play no part in a mimetic resem-

blance. To speak in the crudest of metaphors, it appears as if in

the production of these Coccinelliform cockroaches Nature had

been in a tremendous hurry. The immediate ancestors of

Prosoplecta were probably narrow and moderately elongate cock-

roaches with wings of the Hemithyrsocera type; passage from

this form to a short convex form might have proceeded along at

least two lines—either the wings might have become gradually

shorter part passu with the tegmina, as in certain § species

of Ceratinoptera, Allacta, etc., or the wings could have been

adapted to an ovate, abbreviated body-form by passing through

a Chorisaneura-like stage to the Anaplecta type. As a matter of

fact the evolution of Prosoplecta did not travel along either of

these lines, —the necessary shortening of the wing when in repose

has been produced by a rolling up of as much of the wing us was

necessary for the purpose, quite independently of the boundaries

between the intercalated apical triangle and the rest of the wing,

and so in defiance of the rule observed in all other species of the

family.

368 MR. ROBERT SHELFORD ON

The geographical distribution of Prosoplecta is as follows :—

Hight of the species are found in the Philippine Islands, one in

Celebes, the remaining three in Batchian and Ceram. None

has yet been discovered in the Great Sunda Islands, so that the

distribution is discontinuous and serves to emphasize the view

that if the Philippines are to be regarded as a part of the Indo-

Malayan region, their separation from adjacent land is of very

great antiquity. The Philippine Islands constitute an area in

which insect mimicry has attained great perfection. That is

shown not only by these wonderful cockroaches mimicking

Coleoptera, but Professor Poulton tells me that some of the most

wonderful examples of mimicry amongst butterflies are known

from these islands only, e.g. the distasteful Danaine genus Hestia

is mimicked very closely by a Satyrine and an Elymniine.

Again, the gorgeous little Curculionide of the genus Pachy-

rhynchus are mimicked by other weevils, by Longicorns, by

Cetoniids, and by a cricket *. A comparative study of mimetic

insects in geographically adjacent but zoologically distinct areas,

such as Borneo, the Philippines, and Celebes, is a piece of research

that would surely yield some very interesting results.

I now give a synoptical key to the genus Prosoplecta with

descriptions of all the species known to me. It will be observed

that I have not always been able to pair a species of Prosoplecta

with a definite Coleopterous model, but I am pretty confident

that these models will eventually be found. I have not had

access to comprehensive collections of Philippine Coleoptera, and

so it is chiefly the Philippine Prosoplectae which for the present

Tam not able to match. It is unfortunate that Semper’s collec-

tion of Philippine Coleoptera is broken up and dispersed ; Stal

purchased his Orthoptera for the Stockholm Museum, and here

it was that I found four new species of Prosoplecta, but the

Coleoptera captured at or about the same time and in the same

districts, I have not been able to trace.

Key to the Species of Prosoplecta.

1. Less convex species. ‘Tegmina with a smooth flat-

tened tubercle near the middle of the anal vein.

2. Pronotum piceous, margined all round with hya-

line testaceous ee ee me ecocciellamsaucss

2, BRAG LIAN [OMEN WWNGOWIS sess seccde09 s96 con o20 504 aco 008 P. bipunctata Br.

1’. Very convex species. ‘Tegmina without flattened

tubercles.

2. Ground colour of tegmina ochreous or rufous.

3. Tegmina maculate.

4. Macule of tegmina piceous.

5. Tegminadeeply punctate. (Pronotum with

Almmnacwlae})ieesescess P. trifaria Walk.

* Semper in his ‘ Animal Life,’ p. 390 (International Scientific Series, 1890)

gives figures of some of these mimetic insects, amongst them one with the legend

“ Phoraspis (grasshopper) mimics a Coccinella.” This is an error, for the Phoraspis

is a cockroach, apparently identical with Prosoplecta ligata Br. The species

described below as P. semperi is a much better mimic of the Coccinellid figured

by Semper than is P. ligata. Semper’s figures (and error) are reproduced by

Dr. Wallace in his ‘ Darwinism.’

MIMICRY AMONGST THE BLATTID®. 369

5’. Tegmina not deeply punctate.

ai Ground colour of tegmina ochreous ...... P. nigroplagiata, sp. n.

- Ground colour of tegmina rufous...:..... P. semperi, sp. n.

4’. M: acula of tegmina oc hreous.. ko . Perufa Kirby.

3’. Tegmina immaculate, margined with piceous . P.ligata Br.

2’. Ground-colour of tegmina piceous or dark cas-

taneous.

3. Tegmina with not more than 4 rufous or

ochreous maculz.

4. Tegmina distinctly punctate...........0......... P. quadriplagiata Walk.

ane Tezmina indistinetly punctate... Ea P. nigra, sp. n.

. Tegmina with more than 4 rufous or oc hreous

macule.

4, Pronotum ochreous with 5 piceous macula. P. gutticollis Walk.

4’, Pronotum piceous with the lateral margins

hyaline ochreous.

5. Macule on tegmina ochreous, more or less

GUMBO MUL AI as pnts cy P. mimas, sp. n.

. Macule on teemina ‘rufous, 1 more or less

Pande lcm e ees caer ee a eeeeeeae. tee PE ~ctelophorotdes) spe ie

PROSOPLECTA COCCINELLA Sauss. (Pl. XLVIII. fig. 5.)

Prosoplecta coccinella Saussure, Rey. Zool. (2) xvi. p. 324

(1864); Mém. Mexique, Blatt. p. 173 (1864).

2. Head castaneous, vertex paler. Antenne castaneous at

base, remainder fuscous. Pronotum transversely elliptical, with

the disc piceous, punctate, all the margins testaceo-hyaline.

Tegmina dark castaneous, seriate-punctate; mediastinal area

testaceo-hyaline; a minute, smooth, flattened tubercle at the

humeral angle, a smaller one near the base of the radial vein and

a larger one in the centre of the disc near the middle of the

obsolete anal vein, testaceous. Wings infuscated, veins cas-

taneous. Abdomen above fuscous, supra-anal lamina margined

with testaceous, sub-bilobate ; abdomen beneath piceous, nitid,

sub-genital lamina very large, semiorbicular. Cerci short. Legs

castaneous.

Total poneen 8-9 mm.; length of tegmina 6-2 mm.; pronotum

2°D mm. xX 3°D mm.

Pururppine Is. (Paris Mus., type; British Mus.; Oxford Mus.).

The species is not coccinelliform, but is more like a little

Chrysomelid.

PROSOPLECTA BIPUNCTATA Br. (Pl. XLVIII. fig. 2.)

Areolaria bipunctata Brunner von Wattenwyl, Nouv. Syst.

Blatt. p. 261 (1865).

2. Very similar to the preceding species, but the head is

bright rufous, the palpi and the bases of the antenne are piceous,

remainder of antenne clear testaceous. Pronotum rufous, lateral

margins testaceo-hyaline, posterior margin opaque testaceous,

with a fuscous line at the posterior angle. Tegmina piceous

with mediastinal area testaceous, a flattened smooth tubercle,

testaceous in colour, in the same position as the largest one in

P. coccinella. Abdomen piceous above and beneath, faintly mar-

gined with testaceous, Cox and femora rufous, tibie and tarsi

castaneous.

Proc. Zoou, Soc.—1912, No. XXIV, 24

370 MR. ROBERT SHELFORD ON

Total length 8-5 mm.; length of tegmina 6°5 mm.; pronotum

2mm. X93-d mm.

PururpPine Is. (Stettin Mus., type; Stockholm Mus,, coll.

Semper).

This species is also like a Chrysomelid.

PROSOPLECTA TRIFARIA Walk. (Pl. XLVITI. figs. 4 & 11.)

Prosoplecta trifaria Walker, Cat. Blatt. Brit. Mus. p. 190

(1868) scl:

Prosoplecta megaspila Walker, |. c. (1868) [= @ ].

Very convex, bright ochreous with piceous macule on pro-

notum and tegmina.

¢. Head and antenne ochreous, a blotch on the frons, the

labrum and apices of maxillary palpi, fuscous. Pronotum nitid,

unpunctate, transversely elliptical, with four small piceous maculze

on the disc. ‘Tegmina seriate-punctate, a line between the bases

of the radial and mediastinal veins, a short line above the

humeral angle (which is pronounced), the inner margin of the

left tegmen at the base, the portion of the right tegmen over-

lapped by the left and a round macula in the anal field, piceous.

Wings infuscated. Abdomen above and beneath and the legs

ochreous. Sub-genital Jamina symmetrical with a median plica-

tion, posteriorly emarginate, styles minute. Cerci moderate.

2. Similar, but the piceous markings on the tegmina heavier,

and a large additional macula on the discoidal field beyond the

middle. The abdomen beneath is castaneous, and the sub-genital

lamina is very large and semiorbicular.

Total length (¢) 9 mm., (2) 9°5 mm.; length of tegmina

(do) 75 mm., (2) 81 mm.; pronotum 3 mm. x4 mm.

Barcuran (Oxford Mus., types, coll. Wallace).

The male is an admirable mimic of Odes biplagiata, and the

female resembles a dark variety of the same insect (Chryso-

melide, subfam. Galerucine). (Pl. XLVITI. figs. 3 & 10.)

PROSOPLECTA NIGROPLAGIATA, sp. n. (Pl. XLVIII. fig. 9.)

®. Allied to P. trifaria, but distinguished by the obsolescent

puncturation of the tegmina. Head rufous, vertex ochreous.

Antenne very slender, ochreous, with the apex infuscated.

Pronotum ochreous, smooth, nitid, transversely elliptic, lateral

margins hyaline, eight piceous macule arranged in a circle on the

disc, the anterior pair almost fused, one pair minute. Tegmina

ochreous, extreme base of mediastinal area hvaline, a macula at

the base of the mediastinal area, a macula at the apex of the

same area, a line joing these, a round macula on the middle of

the anal vein, a curved line running from this over the humeral

angle to the base of the tegmen, where there is another spot, the

basal margin, a short line along the base of the sutural margin of

the right tegmen, a large macula in the discoidal field beyond

the middle, all piceous. Wings infuscated. Abdomen beneath

rufo-castaneous, sub-genital lamina very large, semiorbicular.

Cerei and legs rufous.

MIMICRY AMONGST THE BLATTID®. 371

Total length 9-5 mm. ; length of tegmina 7-9 mm. ; pronotum

3mm. x 5 mm.

Putipprne Is. (Stockholm Mus., type, coll. Semper).

This species is a good mimic of Prioptera sinwata Oliv. (Cassi-

dide), and it is highly probable that a Coccinellid and perhaps a

Chrysomelid also enter into mimetic relationship with these

forms. (Pl. XLVIII. fig. 8.)

PROSOPLECTA SEMPERI, sp. n. (PI. XLVIII. fig. 13.)

2. Differs from P. nigroplagiata in the following details :—

Size larger and broader; the general ground colour is rufous ;

there are only six macule on the dise of’ the pronotum, the

anterior pair being suppressed; on the tegmina there are no

lines joining any of the macule, the macule are as in nigro-

plagiata ; there is an ellipsoidal piceous marking on the part of

the right tegmen overlapped by the left; the dise of the sub-

genital lamina is suffused with castaneous,

Total length 10°5 mm. ; length of tegmina 7‘8 mm. ; pronotum

3 mm. xX 5°8 mm.

Puitrprine Is. (Stockholm Mus., type, coil. Semper),

Mimies Leis dunlopi Crotch (Coccinellide), (Pl. XLVITI. fig. 12.)

PROSOPLECTA LIGATA Br.

Cassidodes ligata Brunner von Wattenwyl, Nouv. Syst. Blatt.

p. 262, pl. vi. fig. 28 (1865).

The species is known to me only from the description, which

it is unnecessary to reproduce, since it occurs in the vade-mecum

of all students of the Blattide.

PuitiPPine Is. (Stettin Mus., type).

ProsopLecta RurA Kirby *, (Pl. XLVIITI. fig. 16.)

Prosoplecta rufa Kirby, Ann. Mag. Nat. Hist. () xi pr 379

(1903).

This species is known to me only from the description, which I

transcribe :—

“ Long. corp. cum tegm. 9 millim.; lat. 5 millim.

‘“ Female.—Light red, the tegmina somewhat darker. Pro-

notum smooth and shining, with the front border narrower than

the hind border, transversely oval, with broad borders of sub-

hyaline yellow covering the rounded off lateral angles, and the

hind border yellow. Tegmina with the costal margin subhyaline

yellow, a small yellow spot near the base, and another at one

third of the length, both near the costa ; between the second and

the inner margin is a much larger, transverse, oval, yellow spot.

Base of tegmina blackish. Wings, and a great portion of the

middle of the abdomen beneath, as far as the base of the terminal

segment, black.”

Puitippine Istanps, Mindanao (British Mus., type).

* It is probable that P. rufa is identical with P. bipunctata, a discovery made

long after this paper was in the press. The extreme difficulty under which the paper

was written must be my excuse.

24%

372 MR. ROBERT SHELFORD ON

PROsOPLECTA QUADRIPLAGIATA Walk. (P]. XLVIII. fig. 14.)

Prosoplecta quadriplagiata Walker, Cat. Blatt. Brit. Mus.

p. 189 (1868).

3. Head castaneous, antenne rufo-testaceous. Pronotum

piceous, smooth, nitid. Tegmina seriate-punctate, piceous, an

orange-rufous macula at the base, another in the centre of the

discoidal field. Wings fuscous. Abdomen above and beneath

rufous; supra-anal lamina transverse, carinate, apex emarginate ;

sub-genital lamina as in P. trifaria. Cerci and legs rufous.

Total length 9°5 mm.; length of tegmina 8 mm.; pronotum

2°38 mm. xX 4°5 mm.

BarcaiAn (Oxford Mus., type, coll. Wallace).

Somewhat resembles an EHrotylid of the genus Aulacocheilus or

Cyrtomorphus.

PROSOPLECTA NIGRA, sp.n. (Pl. XLVIII. fig. 6.)

S. Head rufo-testaceous, antenne testaceous. Pronotum cas-

taneous, impunctate, nitid, faintly striate transversely in the

middle of the anterior half. Tegmina very faimtly punctate,

piceous, an orange rufous macula at the apex. Wings infus-

cated. Abdomen above and beneath rufous; supra-anal lamina

transverse ; sub-genital lamina as in the preceding species.

Cerci and legs rufous.

Total length 9 mm.; length of tegmina 7-8 mm.; pronotum

2°9 mm. x 4°2 mm.

Purirprrne Is. (Stockholm Mus., type, coll. Semper).

The species is closely allied to P. ligata Br., and is probably a

mimic of some Chrysomelid beetle.

PROSOPLECTA GUTTICOLLIS Walk. (Pl. XLVIII. fig. 7.)

Prosoplecta gutticollis Walker, Cat. Blatt. Brit. Mus. p. 189

(1868).

@. Head rufo-castaneous, paler on the vertex; antenne tes-

taceous, fuscous towards apex. Pronotum smooth, nitid, ochre-

ous, lateral margins subhyaline, five piceous maculee on the disc.

Tegmina seriate-punctate, piceous, with the following large ochre-

ous macule on each tegmen :—one in the mediastinal area, one

in the middle of the marginal area, one at the base of the tegmen

almost divided into two by a short piceous line on the humeral

angle, and at its lower interior extremity just touching a macula

in the middle of the discoidal field, a fifth macula near the apex

of the tegmina. Abdomen above ochreous; supra-anal lamina

triangular, cucullate, apex not emarginate. Abdomen beneath

piceous, castaneous at base; sub-genital lamina semiorbicular,

ample. Cerci and legs rufous.

Total length 9°5 mm.; length of tegmina 8°5 mm. ; pronotum

3°] mm. x5 mm.

Ceram (Oxford Mus., type, coll. Wallace).

Very like a Coccinellid.

MIMICRY AMONGST THE BLATTIDA, 303

PROSOPLECTA MIMAS, sp. n. (Pl. XLVIII. fig. 15.)

3. Head rufous, vertex paler; antenne testaceous, infuscated

towards apex. Pronotum smooth, nitid, piceous, lateral margins

broadly hyaline, a large ochreous macula at the posterior angles,

enclosing three minute piceous spots, a minute V-shaped ochre-

ous mark in the middle near the posterior margin. Tegmina

minutely seriate-punctate, piceous, with the following ochreous

macule on each tegmen :—one in the mediastinal area, one in

the anal area extending on to the humeral angle, one in the

middle of the discoidal field, one outside this and touching the

radial vein, one at the apex, that on the left tegmen being

larger than that on the right. Wings infuscated. Abdomen

beneath, cerci and legs rufous, subgenital lamina as in the other

species of the genus.

2. Similar to the male but the head and abdomen darker, the

macula in the anal field of the tegmina almost divided into two,

two macule at the apex of the left tegmen. Supra-anal and sub-

genital lamine as in the preceding species.

Total length (¢) 8-9 mm., (9) 10°9 mm.; length of tegmina

(3) 7 mm., (2) 9 mm.; pronotum (¢) 2°9 mm. x 46 mm.,

(2) 3 mm.x5 mm.

PuitipPine Is. (Stockholm Mus., types, coll. Semper).

PROSOPLECTA CQSLOPHOROIDES, sp. n. (Pl. XLVITI. fig. 19.)

3. Head castanecus, vertex ochreous; antenne testaceous,

fuscous towards the apex. Pronotum impunctate, nitid, piceous

with the lateral margins broadly ochreous. ‘Tegmina seriate-

punctate, piceous, with the following rufous macule on each

tegmen:—a small circular one in the mediastinal area, a trans-

verse one extending along the base of the tegmen over the

humeral angle to the base of the radial vein, another broadly

transverse extending across the middle of the discoidal field from

near the sutural margin to the radial vein, a fourth circular in

shape, near the apex of the tegmen. Wings infuscated. Abdo-

men and legs rufous ; supra-anai lamina of usual sha pe.

Total length 9°2 mm.; length of tegmina 8 mm.; pronotum

3 mm. xX 4°9 mm.

Norra Cexepes, Toli-Toli (Oxford Mus., type, Fruhstorfer,

e coll. Van de Poll).

This is an admirably close mimic of the Coccinellid Calophora

formosa Cr. (Pl. XLVITI. fig. 21), a specimen of which was taken

by Fruhstorfer in the same locality and at the same time of year.

The colouring and arrangement of the macule on the pronotum

and wing-covers of the two insects cor respond very closely, but in

the beetles there is a transverse basal band on the elytra, whereas

in the cockroach the corresponding patch of colour is made up of

a band-like spot and an outer circular spot. The mimic is

smaller than its model, and that appears to be rather the rule

than the exception amongst this group of mimetic couples.

Anisolemnia distaura Muls. (Pl. XLVIII. fig. 20) is another

Coccinellid of nearly the same size and evidently also associated,

374 MR. ROBERT SHELFORD ON

Description of a new Genus and two new Species of Blattide.

1 1

Genus M&LyRoIDBA, nov.

Head semiglobose, eyes small, far apart, situated on the sides

of the head ; antenne with moniliform' joints, slightly incrassated,

not pilose, third joint twice as long as second. Pronotum

quadrate, angles rounded, margins somewhat reflexed, not cover-

ing the vertex of the head; disc with slight elevations and

depressions. ‘’egmina densely reticulated, obscuring the venation,

semi-corneous in texture, anal field scarcely distinguishable ;

scute]lum exposed. Wings with moderate apical field, which in

repose is doubled on itself longitudinally and tightly "rolled up,

not folded back over the rest of the wing; traces of venation in

the lower half of the apical field. Supra- ‘anal lamina of female

triangular or trigonal; sub-genital lamina divided by a sulcus;

cerci variable. Legs slender, all the femora unarmed, tibie very

sparsely spined, tarsi without arolia. Male unknown

Text-fig. 46.

oe) = Wine eae

hrntnseunesnwreeeebegs harm taearn

Melyroidea mimetica.

Expanded wing. The apical area is shaded,

This is quite one of the most remarkable genera of Blattidee

that has yet been discovered, ‘The two species comprised in it

present in their form and coloration a curious resemblance to

Malacoderm Coleoptera, whilst in their structural features they

appear to be intermediate between the Corydiine and Oxy-

haloine [= Plectopterine]. The head with its globose front

and vertex and widely separated eyes of small size, the cleft sub-

genital lamina of the female, and the naked tarsal claws are

characteristic features of the Corydiinee, On the other hand, the

wing-structure is totally unlike that of the Corydiine, in whiek

subfamily alone amongst the Blattide the anterior part of the

wing is greatly developed, the posterior part folding beneath not

ina fan-like manner. In Melyroidea the wing with its prominent

apical area and posterior part furnished with radiating veins is

MIMICRY AMONGST THE BLATTIDA. aaa

typically Plectopterine in structure, and this character, in con-

junction with the fan-like folding of the posterior part of the

wing, is overwhelming evidence in favour of the inclusion of the

genus in the Oxyhaloine [ = Plectopterine }.

MELYROIDEA MIMETICA, sp.n. (Pl. XLUVITI. fig. 17.)

2. Head bright rufous, last joint of maxillary palpi fuscous,

antenne fuscous, except the basal and last apical joints, which

are rufo-testaceous. Pronotum bright rufous, quadrate, anterior

and lateral margins slightly reflexed, sides only slightly deflexed ;

dise with slightly rounded elevations, two anterior, two antero-

lateral, and one, the largest, central. Tegmina dark casta-

neous, densely reticulated, mediastinal vein short, an oblique

sulcus on the right tegmen marking the outer limit of the area

overlapped by the left tegmen. Wings infuscated, radial vein

bifureated from the base, twelve costal veins, apices not incras-

sated, median vein simple, medio-discal area crossed by fourteen

irregular venules, thrice as broad as the medio-ulnar area, which is

crossed by eleven venule, ulnar vein bifurcate, transverse venulee

connecting it with the dividing vein, first axillary vein tri-

‘amose; apical area about one third of total wing-length, its base

obtusely angled, nearly equally divided longitudinally, some

obscure venulations in the lower half. Abdomen piceous above

and beneath, supra-anal lamina triangularly produced, sub-genital

lamina cleft by a sulcus; cerci short, acuminate, rufous. Coxe,

femora, and bases of tibia testaceous, rest of tibie and the tarsi

fuscous; formula of apical spines ,, xy > genicular spines on mid

and hind femora; front tibie unarmed except for three apical

spines, mid and hind tibie with one pair of basal and two apical

spines above, with ten spines in a double row and two apical

spines below.

Total length 13-5 mm.; length of body 11 mm.; length of

tegmina 9°5 mm. ; pronotum "4 mm. x 4 mm.; hind femora

4 mm.; hind tibize 4 mm.; hind tarsi 3 mm.

Hab. Uncertain, but probably near Rro DE JANEIRO.

One example (Miers collection, Oxford Museum).

The species 1s very like a Telephorid beetle.

MELYROIDEA MAGNIFICA, sp.n. (VI. XLVIII. fig. 18.)

2. Head bright rufous, maxillary palpi piceous, antenne

piceous, except for three joints beyond the middle, which are

testaceous, slightly incrassated in the middle. Pronotum bright

rufous, quadrate, very slightly broader anteriorly than post-

eriorly, all the borders somewhat reflexed, disc with two antero-

lateral crescentic depressions. Tegmina green, the humeral

angle and the part of the right tegmen overlapped by the left

dark shining blue, densely reticulated but the veins elevated,

mediastinal vein short, radial vein bifurcated from near the base,

six highly irregular and branched costal veins, ulnar vein with

three ramose branches, apex of anal vein attaining a point at

376 MR. H. WALLIS KEW ON THE

more than one third of the sutural margin. Wings very dark

fuscous. Abdomen, cerci, and legs dark blue with metallic

reflections, supra-anal lamina trigonal, sub-genital lamina cleft

and valvular in appearance; cerci very long, with sparse erect

pubescence, not acuminate. Formula of apical spines * e = no

genicular spine on front femora; front tibie with three apical

spines and one spine beneath, mid and hind tibie with two

widely separated spines above, four apical spines and a double

row of spines beneath.

Total length 14 mm.; length of body 11 mm.; length of

tegmina 11°5 mm. ; pronotum 2°8 mm. x 3 mm.

Hcuapor.

One example (Saunders collection, Oxford Museum).

This gorgeous little cockroach is also very like a Telephorid,

but I have not been able to match it with any particular species.

In the preparation of this paper I have been much indebted to

the kind assistance of my friend, Mr. G. J. Arrow, who has

diligently searched the rich collections of Coleoptera in the

British Museum for models to some of the remarkable mimetic

cockroaches described above. Mr. Arrow has also supervised the

preparation of the plate accompanying this paper.

EXPLANATION OF PLATE XLVIII.

Fig. 1. Megapyga eximia Boh. Fig. 12. Leis dunlopi Crotch.

2. Prosoplecta bipunctata Br. 13. Prosoplecta semperi, sp. 0.

3. Oides biplagiata Jac. 14. P. quadriplagiata Walk.

4. Prosoplecta trifaria Walk., 3. 15. P. mimas, sp. n.

5. P. coccinella Sauss. 16. P. rufa Karby.

6. P. nigra, sp. n. 17. Melyroidea mimetica, sp. n.

7. P. gutticollis Walk. 18. MW. magnifica, sp. n.

8. Prioptera sinuata Oliv. 19. Prosoplecta calophoroides,

9. Prosoplecta nigroplagiata,sp.n. sp. 0.

10. Oides biplagiata Jac., var. 20. Anisolemnia distaura Muls.

11. Prosoplecta trifaria Walk., 2. 21. Celophora formosa Crotch.

20. On the Pairing of Pseudoscorpiones.

By H. Watuis Kew, F.Z:8.

[Received January 20, 1912: Read February 6, 1912. ]

(Text-figures 47-50.)

1. Introduction.

The breeding habits of Arachnida are of special interest from

the fact that in no other Class do we find so great a diversity of

method *. Much attention has been paid to the subject, and

the main facts are established for most of the Orders. For Pseudo-

scorpiones, however, scarcely anything is known.

Résel von Rosenhof, so long ago as 1755 (1), tells us that he

kept these animals together for a long time in the hope of seeing

* Cf. Pocock (19), p. 2.

PATRING OF PSEUDOSCORPIONES. Sa

their pairing, yet without success. So also McIntire (5), who

had great experience with several species in captivity, watched

patiently for amiable traits, but all in vain. Schtschelkanowzeff,

in fact, appears to be the only author who has seen the pairing of

any animal of this Order. He states, in a memoir published in

1910 (17), that he saw this act frequently in a species of Chelifer

(Chernes) ; and he has established the fact that fertilization is

effected without intromission of a copulatory organ. But he did

not see exactly what took place ; and the remarkable details of the

process remain, it is believed, quite unknown.

Mr. R. I. Pocock called the writer’s attention to this subject

in 1903, since which time a careful watch has been kept on these

animals, both in the open and in captivity.

The captive individuals, it may be explained, were housed in

what are known as “ McIntire cells”; that is to say, in little cases

about three inches long, an inch or so broad, and a quarter of an

inch or less high. The}body was of sheet-cork, the floor of glass

covered with blotting-paper, and the roof of clear glass ; the whole

being held together by rubber-bands. The glass forming the roof

was transversely cut and hinged, so that food and moisture could

be readily administered *. In these abodes the animals lived in

health for a long time, and they were easily watched both under

low powers of the microscope and with a lens. Observation was

facilitated, it may be added, by the animals walking on the under

surface of the roof, so as to expose to view the genital area, which

occupies in this Order the usual position at the base of the

abdomen,

The sexes meet in ordinary walking position head to head, and,

after some preliminary fencing, the male manages to grasp with

one or both of the hands of the palps one or both of the hands of

the palps of the female. Early observations showed this to be the

case in Chelifer cimicoides Fabr.t, whose pairing, however, has

not yet been fully made out. Similar behaviour was afterwards

witnessed in the relatively gigantie Chelifer cyrneus L. Koch, and

it was on this species that most of my observations were made.

First of all, however, something must be said of Cheliter latreillii

Leach, an animal subgenerically distinct from those just named

and one of considerable interest in many respects.

Il. Pairing of Chelifer latreillii Leach.

Chelifer latreillii, always maritime in Britain, ranges with us

from Fifeshire to Sussex, and is excessively abundant on the

great sand-dunes of the coasts of Lincolnshire and Norfolk. In

such places it makes its home for the most part in the tussocks of

Ammophila arenaria; but it evidently moves about freely, being

found often under pieces of wood, etc., on the sandy ground.

It belongs to the subgenus Chelifer s. s., a small group

* Cf. McIntire (4), pp. 71-2.

+ For the nomenclature employed in this paper, cf. Kew (18).

378 MR. H. WALLIS KEW ON THE

remarkable for the specialization of the male, in which both primary

and secondary characters are unusually pronounced*. The

genital area of this sex is large and conspicuous, the first genital

plate being short and of peculiar character, while the second is

both long and broad, and under this latter are found two very

large structures, inappropriately called ram’s-horn organs, to which

it will be necessary to recur, ‘he fourth pair of coxe, which

bound the genital area anteriorly, differ much from those of the

female, being strongly concave behind and containing a peculiar

organ, the coxal sac of With (18). Of characters remote from the

genital area there is considerable diversity within the group; there

are generally remarkable modifications in the legs of the first pair,

and these modifications are unusually well-marked in the present

species. ‘The whole leg is greatly strengthened, with hump-backed

tarsus, and greatly enlarged claws, the anterior claw being of

peculiar shape with oddly turned extremity and with a process

along its anterior margin. Further, while the hand of the palp

is a little smaller than that of the female, the fingers when closed

at the tip have a wider gape.

The ram’s-horn organs already mentioned—supposed to be

tracheal in origin—have been studied in allied species by several

authors. They are concealed under the second genital plate, where

they he in a highly contracted condition. Preserved specimens,

in exceptional cases, have them protruded externally ; and they

have been figured thus protruded by Simon (6), and after him by

Tomosvary (7); and by With (16). It does not appear, however,

that they have been seen in action; and the suggestion of Menge

(2) and Simon (6) that they are sper m-transmitters, perhaps intro-

mittent organs tf, is erroneous, as also is that of Schtschelkanowzetf

(17), who thought they might be concerned in placing sperm-

masses on the ground.

With regard to the other characters, except that With (14) and

Schtschelkanowzeff (17) have supposed the coxal sac to be a sense-

organ with some sexual significance, no suggestions appear to have

been made. The gape of the fingers, however, is obviously con-

nected with the grasping of hands already mentioned ; and the

observations now recorded show what is done with the ram’s-horn

organs and with the legs of the first pair.

It was in May 1905, on the sand-dunes between Sandwich and

Deal. that I first saw the weeting of the sexes of this animal. On

turning over a piece of tin, I found on its under surface a male

and female which had approached each other head to head, and

were actively fencing with the palps. The male at length dis-

armed the female by getting a firm grasp of the hands, which

were held fast during the whole of these preliminary proceedings.

* With (14), pp. 132-3; With (16), p. 220; Kew (18), p. 47.

+ Menge (2), 1D Wr, Tole in, ines; WB) Hansen (8), pl. vil. fig. 24; Croneberg (9),

pp. 456- 7. pl. xia. figs. as AVP Bernard (LO), pp. 423-6 ; Oudemans (15), pp. 136-140;

Schtschelkanowzeff veupy pp. 6-14.

t¢ On the supposed occurrence of intromittent organs in this Order, cf. Lankester

(12), p. 256; and Croneberg (9), p. 39.

PAIRING OF PSEUDOSCORPIONES, 379

The animals moved considerably backwards and forwards and

round about; and the male, all the time in a state of great

activity, made at intervals determined advances towards the

female, but he was prevented from approaching closely. On the

occasion of each of these advances the male brought the ram’s-

horn organs into action, running them out rather rapidly to their

full extent ; they took at these times a forward lateral direction,

their extremities usually assuming an outward curve and passing

under or over the femora of the male’s palps. Occasionally they

touched some part of the palps of the female, but this seemed

accidental. At the full extension of the organs, the male vibrated

his body in a peculiar manner and had at this time a remarkable,

even villainous appearance ; but the organs were exposed only for

a few moments, after which they ran in again rather rapidly, the

male then taking a few steps backwards. Eventually the animals

separated — perhaps disturbed by the sunlight to which they were

exposed—without proceeding to any act of fertilization. In 1906,

also in May, similar observations were made on specimens collected

on the Lincolnshire coast, but again no act of fertilization was

seen ; and no more observations were made till the spring of 1911.

In the third week of April in that year, a good number of

specinens were collected on the Camber sand-hills in Sussex ; and

they were kept under daily observation in one of the cells above

described. During the remaining days of April and the early

days of May, the males, which had the abdomen rather full,

seemed to have difficulty in restraining the ram’s-horn organs,

the tips of which constantly appeared from under the great genital

plate, which was at such times raised anteriorly and depressed

into the abdomen posteriorly. Many fencing contests between male

and female were seen; but the male did not always obtain any

advantage in the grasp and was often in fact rather roughly used ;

he continued, however, even at these times, to display the ram’s-

horn organs with great energy. As before, I failed to observe the

acts of fertilization. It was evident, however, that such acts had

occurred, either before or after the animals came under observation,

for about a month later eggs began to appear externally on a

few of the females, and in the first week of July young broods

appeared. Shortly before this time, fortunately, that is to say

about mid-summer, there was a recurrence of sexual activity in

the cell; and on 22nd June the whole process of the passing of the

male product to the female was successfully witnessed.

The animals, male and female, had taken up a position, ventral

face uppermost, on the under surface of the glass; and the male,

who was firmly holding both hands of the female, was making

periodical advances towards her with display of the ram’s-horn

organs as above described. It soon became evident that the

female was offering but slight resistance; the male was allowed

to make a near approach, and at length he was even permitted to

caress with his chelicerze the chelicerze of the female. This done,

he quickly retired as before by taking a step or two backwards;

380 MR. H. WALLIS KEW ON THE

and it was now seen that as he did so the female was quite eager

to take the corresponding steps forwards. As this tendency

became more and more marked, the male released his grasp of the

hands of the female, who was now free, at least from corporal

Text-fig. 47.

Chelifer latreillii Leach.

Male and female; in ventral view, seen through a piece of glass, on the under-surface

of which they are standing. ‘The position is that of the last phase of the court-

ship; the male has released the hands of the female, and is about to extrude

the spermatophore; the ram’s-horn organs are fully extended or nearly so.

x 13.

(The bristles and tactile-hairs are omitted.)

control. The male continued, however, to make advances and to

display the ram’s-horn organs even with greater energy than

before; and finally, on the occasion of one of these advances—while

PAIRING OF PSEUDOSCORPIONES. 381

not in contact with the female in any way—the ram’s-horn organs

being fully extended (text-fig. 47), he extruded from the genital

opening between the bases of these organs a large elongated

structure—evidently a spermatophore *—one extremity of which

became at once attached to the glass by means of a foot-like pad

of quick-drying adhesive matter. The other extremity was still

in contact with the genital opening of the male; and when thus

freshly extruded this spermatophore bore externally, somewhat near

the middle, some clouded liquid in the form of a moderately large

surrounding globule. After a delay of a few moments, the male

stepped backwards, and thus released the spermatophore, leaving

it attached to the glass as just mentioned, but otherwise free, in

an oblique position, with the unattached extremity directed towards

the male. No sooner had the male thus stepped back, than the

female came quickly forward till the female genital opening was

in contact with the spermatophore. This forward movement of

the female was accompanied, I believe, by a slight but rapid

forward movement of the male; and, however this may be, the

head of the male had passed under that of the female; and at the

same moment, with great suddenness, the male threw forward the

stout legs of the first pair and seized with them the anterior

margin of the female genital opening, on which the enlarged

peculiarly formed claws became firmly hooked ; and now the male

commenced a rather long series of violent pulling movements with

these legs, by means of which the body of the female was moved

on the spermatophore, a part of which had, I believe, entered the

female genital opening. These movements brought the proceedings

to a close, and the animals now separated, leaving the sperma-

tophore, or at least the shell of it, still attached by its foot to the

glass. No repetition of these acts was observed. Another

spermatophore was found in the cell, however, towards the end

of August.

Text-fig. 50 A (p. 386) shows the spermatophore from above and

from below. It was of firm substance and somewhat complicated

in structure. Beyond the foot of attachment it was rather slender

but it gradually increased towards a neck-like constriction, beyond

which was a widened head with a small horn-like point on either

side ; and beyond this head was a rather long narrowed extremity

of definite construction. The total length was considerably more

than half that of the entire animal.

III. Pairing of Chelifer cyrneus LZ. Koch.

Chelifer cyrneus is known with us only in Sherwood Forest

(Nottinghamshire) and Richmond Park (Surrey), where it lives

under rather close-fitting bark of dead or partly dead oak-trees.

* The occurrence, in this Order, of a spermatophore, though not mentioned in the

text-books, is not entirely new, since McIntire (5) saw Chthonius @, in captivity,

extrude elongated structures in which were spermatozoa. No female appears to have

been associated with the male during this act, and it is thus doubtful whether the

extrusion was normal.

382 MR. H. WALLIS KEW ON THE

It belongs to the subgenus Chernes; and the male (text-fig. 48),

in sharp contrast with that of Chelifer latreilli, is but little

specialized. The genital area of this sex is only moderately con-

spicuous, the second genital plate being short; the fourth pair of

coxe differ only a little from those of the female, and there is no

coxal sac; further, there are no protrusible ram’s-horn organs, and

the legs of the first pair exhibit no marked modification. A

slight enlargement of the hands of the palp, in fact, is the only

secondary character of note.

Text-fig. 48.

Chelifer cyrneus Li. Koch.

Male in ventral view. X 20.

(The bristles and tactile-hairs are omitted.)

Compared with Chelifer latreillii, the present animal is much

larger and more heavily built, and unlike the former species, it is

PAIRING OF PSEUDOSCORPIONES. 383

eyeless. In the male, in addition to the important differences

above indicated, there is a deep-seated unlikeness in the internal

organs; and we shall find that the spermatophores are entirely

unlike, with corresponding differences in the pairing.

In the second week of April 1911, a supply of specimens was

obtained from Richmond Park *, and a cell was stocked with five

or six individuals, which were kept under daily observation tiil

the beginning of the second week of May. During this time,

fortunately, abundant opportunities were afforded for observing

the pairing, which was carried on with great persistence on ati

least eight occasions. The animals did not walk easily on the

under surface of the glass, and pairing was not observed in that

position ; it was well seen, however, both in dorsal and lateral

view, and under the latter condition the details of the process

were distinctly made out.

The male and female met, as already stated, in walking

position head to head (text-fig. 49, p. 384); and, as in the former

species, they engaged in some preliminary fencing with the palps.

During this fencing the male always obtained with one of the

hands a firm grasp of one of the hands of the female; and, con-

trary to what occurred in the former species, this grasp was

always maintained throughout, that is to say until the animals

parted company when the pairing for the time being was

complete. Moreover, while the former species held the female

with both hands, the present animal invariably employed one

hand only in this way, always keeping the other hand free.

After the grasp was made, the animals fenced with the free palp

and moved about a little backwards and forwards; the male—

all the time alert and eager—constantly attempting to approach

closely to the female. Wate hing the animals carefully at this

time, it was soon observed that the fencing had given place to a

regular system of display on the part of the male, whose actions

in this respect were quite unlike anything seen in the former

species. In the present case—in the absence of ram’s-horn

organs—the display was made with the free palp and with the

legs of the first pair. The free palp was brought round at

frequent intervals and the great hand rapidly shaken in the face

of the female in a remarkable threatening or perhaps beckoning

manner; and the first legs were rapidly “moved, that is to say

lifted and replaced, in most peculiar fashion. After a time the

female, apparently much impressed with these actions, offered

little or no resistance, the male having now no difficulty in

approaching closely. At length the male—head to head with

the female but not in contact ‘except for the continuous grasp of

one hand—deflected the base of the abdomen to the floorand atlixed

there the adhesive foot-like attachment of the spermatophore.

Almost immediately, the body was raised to its normal position,

* The writer is indebted to His Majesty’s Office of Works and to Mr. S. Pullman,

the Superintendent of the Park, for the permission and facilities necessary for the

taking of the animals in this place.

MR. Hf. WALLIS KEW ON THE

384

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PAIRING OF PSEUDOSCORPIONES. 385

and one then saw that a whitish filament was stretched from the

foot-like attachment to the genital opening ; ; and now the male

produced very quickly from the genital opening a large globule

of brilliant liquid; and this globule remained on the filament,

just below the genital opening, like a great bead on a thread.

Standing thus for some moments, perhaps awaiting some sign

from the female, and continuing with great energy the shaking

of the free hand, the male at last quickly raised his body and

took a step or two backwards so as to free the globule and filament ;

and it was now seen that the filament, which passed through

the globule, bore just above it a small irregular termination of

whitish substance. The remarkable spermatophore thus com-

pleted remained standing on the ground in an erect or suberect

position. At the moment when the male raised bis body and

stepped back, the female, still held by the hand by the male,

ran forward; the movement of the two animals being per fectly

mutual and just sufficient to bring the genital area of ‘the female

into the exact position occupied “the moment before by that of

the male. As the female thus ran forward the genital orifice was

widely open; and just as the spermatophore was reached a slight

forward and downward movement was made upon it; and, the

genital orifice being quickly closed, the globule together with the

whitish termination of the filament were taken in at a gulp.

Only the naked filament remained. At the moment of this rapid

gulping in, the female drew back ; and at the same moment the

male, with great eagerness and with the chelicere extended and

open, ran forward until the fore parts of male and female were in

contact; and the two animals remained thus in a state of quiescent

embrace for some time. During this embrace the head of the

male went under that of the female; but no part of the male was

at any time directed towards the female genital opening. The

coupling action of the legs of the first pair, which appears to be

an essential condition in the pairing of Chelifer latreillii, had no

counterpart in the present species. At the conclusion of this

period of repose, the male roused himself to activity again,

recommenced the rapid movements of the free palp and of the

legs of the first pair, and the whole process was repeated.

‘The whole process was, in fact, always repeated many times.

To this there was no exception in all the observations. The male

never once released, not even during the periods of repose, the

fixed grasp of the hand of the female; spermatophores were

regularly produced and received at intervals of from eight to ten

minutes; and this recurrent pairing was continued for two hours,

or even for three hours, or more.

By the beginning of the second week of May the activity of

the males showed signs of abating, and the colony was broken up.

Soon afterwards, however, a new colony was established, and

pairing was seen again in 5 uly and August.

Text-fig. 50 B shows the spermatophore—two of eae tale

lateral view. This object agrees with that of the former species in

its attachment to the floor, but is otherwise of different character,

Proc. Zoot, Soc,—1912, No. XXV. 25

386 MR. H. WALLIS KEW ON THE

being altogether more simple and smaller. Beyond the small

foot of attachment it consists, as we have seen, merely of asimple,

more or less rigid filament, which bears around it near the top a

large globule of liquid and has just above the globule an irregular

termination of whitish substance.

Text-fig. 50.

—

Y Ne

A. Spermatophore of Chelifer latreillii Leach, from above and from

below. X 50.

B. Spermatophores of Chelifer eyrneus L. Koch, from the side. X 50.

C. Spermatozoa of Chelifer cyrneus 1. Koch. (Drawing commu-

nicated to the writer by Mr. C. J. With.)

The globule has a diameter somewhat exceeding the depth of

the tibia of the legs of the animals. Unfortunately it was not

ascertained whether the spermatozoa (text-fig. 50 C) were contained

in the globule or above it at the termination of the filament—

from my experience in this direction the obtaining of a com-

plete spermatophore for examination will not be an easy task—

but however this may be, the amount of material transferred

from the male to the female is surprisingly large.

PAIRING OF PSEUDOSCORPIONES, 387

On the breaking up of the first colony, the animals were

examined them, and found spermatozoa in the females. These

formed a mass in the vagina ; and occurred also, placed in a single

row, in the two long narrow irregularly-coiled tubes which run

out from the vagina anteriorly. These tubes were figured in an

allied species by Croneberg (9). From their structure, and from

the presence of spermatozoa which would evidently be preserved

there, they must be regarded as receptacula seminis, not as glands

as Croneberg supposed *.

It was on the present species that Schtschelkanowzeff (17)

made the already published observations to which reference is

made at the commencement of this paper t. He found the animal

in the Russian Government of Tschernigoff, in a forest belonging

to his father, where certain pine-stumps were reserved for the

purposes of observation. By removing the bark from these

stumps he was able to learn much of the animal’s manner of life,

and frequently saw their pairing. He relates that after the

grasping of one hand and other preliminaries, the male made

some convulsive movements and depressed the abdomen to the

stump ; and thereupon the female took the place of the male and

depressed her abdomen exactly at the spot where the male had

done so; and these actions were repeated several times. The

male, he concluded, had deposited sperm and the female had

taken it up; and on subsequent dissection of the female, sperma-

tozoa were detected in the vagina and receptacula seminis. This

impression of the pairing, it will be seen, is roughly in agreement

with the account above given; it is, in fact, exactly what would

be obtained by a not very close observation of the animals in

dorsal view, in the open, with or without a lens. Presumably

they were not viewed laterally, for the sperm’ was supposed to

have been deposited in little heaps ; and thus there is no mention

of the characteristic spermatophore. Schtschelkanowzeff supposed,

finally, that the active rdle was taken throughout by the female;

but in this, doubtless, he was mistaken.

IV. Summary.

The two Pseudoscorpiones observed belong to the genus

Chelifer s.1., and represent respectively the subgenera Chelifer s.s.

and Chernes.

The males are differently equipped : the Chelifer has an elaborate

genital area, long ram’s-horn organs, and much modified legs of

the first pair; the Chernes has a less elaborate genital area, no

ram’s-horn organs, and no modified legs. There is agreement,

* This result appears to have been arrived at already by Schtschelkanowzeff

(17, p. 27), and ef. Lubbock (3, p. 615).

+ Schtschelkanowzeff (11) described his animal as C. multidentatus, Sp. n.3

but paratypes obligingly communicated to the writer establish its identity with

C. cyrneus.

25*

388 MR. H. WALLIS KEW ON THE

however, in a feature of prime importance: both are destitute of

intromittent organs of copulation.

Fertilization is effected in both by means of a spermatophore.

This structure in the Chelifer is large and somewhat compli-

cated ; in the Chernes relatively small and merely filiform

Correlated with the differences im equipment and in the

spermatophore are considerable differences in the pairing.

There is agreement, however, as follows. The male and female

face one another in walking position. ‘The male grasps with one

or both hands one or both hands of the female. There is a forced

courtship, during which the male makes display of definite

character. At length he extrudes the spermatophore, which is

attached to the floor in front of the female, where it stands erect

or obliquely. From this object the male retires backwards, and

the female at the same moment comes forwards. The movement is

just sufficient to bring the female genital opening into contact

with the spermatophore, and the male product is thus received

without delay.

The differences may thus be stated. The Chelifer male holds

the female with both hands; and makes display with the ram’s-

horn organs. He releases the hands previously to the extrusion of

the spermatophore. When the female comes forward, he seizes

her by the genital opening with the legs of the first pair, and then

executes a series of pulling movements by which presumably the

reception of the male product is facilitated. The animals now

separate; and it is unlikely that the process is repeated, except

perhaps at long intervals. The Chernes male holds the female

with one hand only. He makes his display with the other hand

and with the legs of the first pair. He does not release the

female previously to the extrusion of the spermatophore, but

continues to hold her by the nand throughout. When the female

reaches the spermatophore, she takes the male product quickly

and retires backwards. There is no seizing of the genital opening

by the male. As the female retires, however, the male quickly

follows; and there is a period of repose; after which the whole

process is repeated. Moreover, it is repeated, with similar periods

of repose, many times; and a large number of spermatophores

are thus produced and received in rather quick succession.

By way of conclusion it may be recalled that the genus

Ohelifer s.1. comprises, in addition to the subgenera Chelifer s.s

and Chernes, two others, Alemnus and Withius : ali distinguished

by remarkabie differences in the sexual equipment of the males—

even within the subgenera there are striking divergences in the

secondary characters ; and these facts, in view of the differences

above noted, certainly lead one to expect great variations in the

pairing processes. A review of the males of the whole Order,

moreover, serves greatly to increase this expectation—one may

refer for instance to the complete dissimilarity in the genital

areas of Obisium and Chihonius—so that there is here undoubtedly

PAIRING OF PSHVDOSCORPIONES. 389

a fertile field for investigation. Nevertheless it may perhaps be

predicted that the general lines now indicated are those on which

fertilization will be found to be effected in all Pseudoscorpiones.

V. List of References.

(1) Rost von Rosgnnor, A. J.-—Der monatlich-herausgegebenen

Insecten-Belustigung, iii. Niirnberg, 1747-1755.

(2) Mencr, A.— Ueber die Scheerenspinnen, Chernetide.

Neueste Schriften d. Naturforschenden Gesellschaft, v.

pp. 1-42, pls. i-v. Danzig, 1855.

(3) Lussockx, J.—Notes on the Generative Organs and on the

Formation of the Egg in the Annulosa. Philosophical

Transactions of the Royal Society, cli. pp. 595-627.

London, 1862.

(4) McIntire, 8. J.—Some cheap aids to Microscopical Studies.

Journal Quekett Microscopical Club, i. pp. 69-72. London

1868.

(5) McIntire, S. J.—Pseudoscorpions. Hardwicke’s Science-

Gossip, v. pp. 243-247. London, 1869.

(6) Stmon, E.—Les Arachnides de France, vii. Paris, 1879.

(7) TomosvAry, O.— Pseudoscorpiones Faune Hungarice.

Magyar Tudomanyos Akad. Math. és Természettud,

Kozlemények, xviii. pp. 135-256, pls. i-v. Budapest,

1882.

CRONEBERG, A.—Beitrag zur Kenntniss des Baues der

Pseudoscorpione. Bull. Soc. Imp. des Naturalistes (n. s.),

i. pp. 416-461, pls. x.—xi. a. Moscou, 1888.

(1G) Bernarp, H. M.—Notes on the Chernetide, with special

reference to the Vestigial Stigmata and to a new form of

Trachea. Journ. Linn. Soc., Zoology, xxiv. pp. 410-430,

pls. xxxi-xxxi. London, 1894.

(11) ScurscHELKanowzerr, J. P.—Chernes multidentatus, n. sp.,

nebst einem Beitrige zur Systematik der Chernes-Arten.

Zoologischer Anzeiger, xxv. pp. 350-355. Leipzig, 1902.

(12) Lankester, EK. R.—The Structure and Classification of the

Arachnida, Quart. Journ. Microscopical Science (n. s.),

xlvili. pp. 165-269. London, 1904.

(13) Wir, C. J.—On Chelonethi, chiefly from the Australian

Region, in the Collection of the British Museum, with

Observations on the ‘ Coxal sac” and on some cases of

Abnormal Segmentation.. Ann. & Mag. Nat. Hist. (7)

xv. pp. 94-143. London, 1905.

(14) Wirx, C. J.—The Danish Expedition to Siam, 1899-1900.

Chelonethi: an account of the Indian False-Scorpions

together with studies on the anatomy and classification

of the Order. Kgl. Danske Videnskabernes Selskabs

Skrifter (7), i. pp. 1-214. Copenhagen, 1906,

(8) Hansen, H. J.—Zoologia Danica, iv. Copenhagen, 1885.

390 MR. OLDFIELD THOMAS ON

(15) Oupemans, A. C.—Ueber Genitaltracheen bei Chernetiden

und Acari. Zoologischer Anzeiger, xxx. pp. 135-140.

Leipzig, 1906.

(16) Wir, C. J.—An Account of the South American Chell-

ferine in the Collections of the British and Copenhagen

Museums. Trans. Zool. Soc. xvill. pp. 217-340, pls. xxix.—

xxxl. London, 1908.

(17) ScutscHELKANOowzEFF, J. P.—Der Bau der miannlichen

Geschlechtsorgane von Chelifer und Chernes. Festschrift

zum sechzigsten Geburtstage Richard Hertwigs, i.

pp. 1-38. Jena, 1910.

(18) Kew, H. W.—A Synopsis of the False-Scorpions of Britain

and Ireland. Proc. Royal Ivish Academy, xxix. B.

pp. 38-64, pls. iv—vi. Dublin, 1911.

(19) Pocock, R. 1—A Monograph of the Terrestrial Carboni-

ferous Arachnida of Great Britain. (Palsontographical

Society.) London, 1911.

EXHIBITIONS AND NOTICES.

February 20, 1912.

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

in the Chair.

Dr. A. T. Masterman, M.A., F.Z.8., gave a demonstration,

illustrated by a large number of lantern-slides, of recent imvesti-

gations on Age-determination in the Scales of Salmonoids, with

special reference to Wye Salmon.

March 5, 1912.

Sir Joun Rose Braprorp, M.D., D.Sc., F.R.S.,

Vice-President, in the Chair.

The Races of the European Wild Swine*.

Mr. Ouprietp Tuomas, F.R.S., F.Z.8., exhibited the skull of a

Hungarian Wild Boar, recently presented to the National Museum

by Fraulein Sarolta von Wertheimstein, and that of an ordinary

German Wild Boar, representing the typical Sus scrofa Linn.

* [The complete account of the new forms described in this paper is given here,

but since the names and preliminary diagnoses were published in the * Abstract,’

they are distinguished by being underlined.—EpirTor. |

EUROPEAN WILD SWINE. 391

The difference in size between these two skulls was so enormous

that Mr. Thomas expressed the opinion that the Hungarian Boar

should be distinguished as a diflerent species, which he proposed

to call Sus attila. A fine stuffed specimen of the same species,

from Volhynia, 8. Russia, presented by Count Potocki, had been

on exhibition for some years in the Museum, under the old name

of Sus scrofa.

The difference between the two species was mainly in size, as

would appear from the skull-measurements given below; but

it might be noticed that both specimens of Sus attila were, on

the whole, lighter in colour, and had whiter muzzles, than the

available specimens of Sus scrofa. No series of skins, however,

was available to show how far these colour-differences were constant.

The character of the pelage of Sus attila was similar to that of

S. scrofa in the presence of a thick woolly underfur. The median

anterior hoofs in Sus attila were 65 mm. in length, as compared

with 48 mm. in S. scrofa.

The type locality of S. attila was Kolozsvar (= Klausenburg),

Transylvania, and the species no doubt extended through Russia

into Siberia.

The other names usually placed in the synonymy of Sus scrofa—

selosus Boddaert, aper Boddaert, ewropeus Palas, and celtica Strobel

—were all merely suggested alternatives for scrofa, and applied

solely to the German Wild Boar, this being the typical form of

Linneus’s species *.

Furthermore, Mr. Thomas pointed out that the Wild Boar of

Southern Spain was, on the other hand, so very much smaller

than the true Sus scrofa of Germany that it appeared worthy of

a special subspecific name, and he proposed to call it Sus scrofa

beticus. In addition to its small size, the two skins in the British

Museum (both of winter specimens) were remarkable for having

no woolly underfur whatever, that of Sus scrofa scrofa being thick

and abundant.

A female specimen of this small Wild Boar, from Seville, had

been presented to the Museum by the late Lord Lilford in 1895,

and a male skin and skull, and a separate skull, from the Coto

Dofiana, by Mr. Abel Chapman in 1908.

Finally, the Wild Boar of Nor oe n Spain was intermediate in

size between S. s. beeticus and S. s. scrofa, and possessed, at least

in winter, a woolly underfur as in teadinsy Wild Boars. Of this

form a fine male and female had been obtained for the Museum

by the Rev. Saturio Gonzalez at Quintanar de la Sierra near

Burgos.

Mr. Thomas proposed to call this race, whose skull-measure-

ments were included in the table below, Sws scrofa castilianus.

The following were the skull-dimensions in millimetres of (1)

the type of S. aétila, an adult but not old male, (2) of a rather

* Cf. Thomas, P.Z.5. 1911, p. 140.

Bi) ON EUROPEAN WILD SWINE.

older male of Sus scrofa from Baden, Germany, (3) of the type

skull of S. s. castilianus, and (4) that of S. s. beticus, intermediate

in age between the first two :—

Sus scrofa Sus scrofa Sus scrofa

Sus attila. scrofa. castilianus™. beticus.

3: 3. 3. 3.

Occipito-nasal length ............... 452 385 353 324,

Condylo-basal length ............... 407 (c.) 3855 335 (c.) 305

Zygomatic breadth .................. 174 146 135 142

Interorbital breadth .................. 102 73) 78 72

IWeyalls, WOVEN asbseensvscancdcaraseuce 250 208 189 173

weiperapy)| ORG evo fn ORive eamn aetna 53 36 30 33

Median occipital height ............ 159 ae 114

Bs f > skull on

lower jaw rere: ate e AA 271 208 198 208

Height at anteorbital foramen ... 84. 65 60 61

Upper cheek-tooth series ............ 134 121 125 117

Lower molar series .........,........ 84 69 79 72

Breadth of antero-internal face of

WOWIEE CAA® sjoshysosncooee Woo setae 25°5 215 23 20

IWS» Seacrest Aya ue ae ere Mcaconre sche tania AKO) Seo Pig 35X21 43 X 22 35°5 X21

Ve ees Aa o's ase us setaet ernst a pL LO, 36X17 42175 36 X17

The results arrived at were as follows :—

1. Sus scrora Linn.

Upper length of skull not exceeding about 410 mm.; height,

including lower jaw, at most 210 mm,

Subspecies :—

] A. SUS SCROFA SCROFA.

Synn. sefosus, aper, ewropeus, celtica.

Upper length of skull of male about 380-410mm, Woolly

underfur present.

Range, Central Europe, from Germany to the Pyrenees.

Type locality, Germany.

1 B. SUS SCROFA CASTILIANUS.

Thos, Abstract P. Z.8, 1912, p. 13 (March 12).

Upper length of skull of male 353 mm., of female 331.

Underfur present.

Range. Northern Spain.

Type locality. Quintanar de la Sierra, near Burgos.

Type. Adult male. B.M. No. 11,10.5.3. Collected by Rev.

-Saturio Gonzalez; presented by the Hon. N, Charles Roth-

sehild.

* An old female skull of 8. s, castilianus measured 331 mm. in occipito-nasal

length,

ON SWISS SPIDERS. 393

1c. Sus SCROFA BATICUS.

Thos. Abstract P. Z.S. 1912, p. 14 (March 12).

Upper length of skull of male 324 mm. No woolly underfur

present, even in winter.

Range. Southern Spain.

Type locality. Coto Dofiana, Huelva.

Type. Old male. B.M. No. 8.3.8.12. Collected 6 February,

1908, and presented by Abel Chapman, Esq.

2. Sus ATTILA.

Thos. Abstract P. Z. S. 1912, p. 13 (March 12).

Upper length of skull of male 452 mm.; height, including

lower jaw, 271 mm. Woolly underfur present.

Range. Hungary and 8. Russia, probably extending into

Siberia.

Type locality. Kolozsvar (= Klausenburg), Transylvania.

Type. Adult male. B.M. No. 12.1.23.1. Collected 8th Decem-

ber, 1911. Presented by Friiulein Sarolta von Wertheimstein.

PAPERS.

21. A Contribution towards the Knowledge of the Spiders

and other Arachnids of Switzerland. By the Rev.

O. Pickarp-CamBripcE, M.A., F.R.S., C.M.Z.S8., etc.

[Received October 13, 1911: Read February 20, 1912.]

(Text-figures 51 & 52.)

The materials for the accompanying List have been kindly got

together for me by those whose initials are appended to each

species. They have been collected for the most part during tours

through Switzerland, and in one or two cases during short

sojourns in some of the localities mentioned. It is not pretended

that the number of species enumerated represents more than a

small proportion* of those recorded by other authors, or likely to

be found by careful expert search in Swiss regions, but it shows

how very many objects in one special branch of Natural History

can be obtained even by observers who may be non-specialists, as

was the case with nearly all those whose initials are here appended.

Of course it must be taken into consideration that in the collect-

ing of Arachnids there is no necessity for the paraphernaha

absolutely required for the collecting and preserving of entomo-

logical specimens in general. A few small bottles of methylated

* The present known number of Swiss Spiders, according to Dr. Roger de Lessert,

amounts to 616, See Revue Suisse Zool, vol. xvi. p. 485, 1909.

394 THE REY. O. PICKARD-CAMBRIDGE ON

spirit of wine, a few glass tubes of different sizes, contained in

strong conveniently shaped tin boxes, are really all the materials

that are needed for the preservation of Arachnids; while the

‘“‘eye of faith and the finger of instinct” assisted by an empty

glass tube will suffice in the actual field. Not, of course, but that

a Strong entomological sweeping hoop-net, for the brushing and

sweeping of shrubs and herbage, would fill the pocket-bottles all

the faster, and probably with some species not obtainable by

other methods. The preponderance in the following List of

these groups of Spiders which at once and most obviously claim

the attention of the tourist as he toils along—viz., the Hpeiride,

Thomiside, Lycoside, and Salticidee—shows, it seems to me, how

rich a harvest would be reaped by a resident specialist in some of

the localities. The number of species here recorded, in these

four groups alone, amounts to 102 out of the total of 212 species

collected of the Araneidea (or true Spiders); while those of the

great group Theridiide (in its widest sense) are only 65 in

number: a group which, I may remark in passing, numbers 1

the British Islands alone upwards of 250 out of a total of the

whole Order of somewhere about 550! The greater number,

however, of the Theridiide require something more in the way of

wearisome search than even the most observant eye of the

ordinary passing tourist; and hence the tourist’s bottle is most

commonly filled by species of the other four groups mentioned,

and which, often in countless numbers, are obvious on all sides

on a fine day in the mountains.

I must here acknowledge my indebtedness to Mons. Kugéne

Simon of Paris,and Dr. Roger de Lessert, of the Natural History

Museum, Geneva, for their invaluable assistance in the identifica-

tion of many of the species in the following List with which I

was unacquainted.

List oF Swiss ARACHNIDS.

Collected, or sent to me, at various periods and in various

o) ’ p

localities by the following :—

Initials in List.

A.W.P.-C. (A. W. Pickard-Cambridge, Balliol College,

Oxford.)

R.J.P.-C. (Rev. R. J. Pickard-Cambridge, Warmwell

Rectory, Dorchester.)

H.A.P. (Jate Rev. H. A. Pickard, Airedale, Oxford.)

C.W. (Cecil Warburton, Christ’s College, Cambridge.)

F.P.S. (fF. P. Smith, 5 Gibson Square, London.)

R.G. (Robert Godfrey, late of 46 Cumberland Street,

Edinburgh.)

G.N. (late George Nicholson, Director, Royal Gardens,

Kew, and 37 Larkfield Road, Richmond.)

A.S.A. (late A. 8. Atkinson, Nelson, New Zealand.)

SWISS SPIDERS. 395

Lnitials in List.

E.S. (Eugéne Simon, 16 Villa Said, Avenue du bois

de Boulogne, Paris.)

A.E.E. (Rev. A. E. Eaton, Symondsbury, Bridport,

Dorset.)

H.S8. (Henry Speyer, Reigate.)

R.deL. (Dr. Roger de Lessert, Museum of Natural

History, Geneva.)

C.E.M.I. (C.K. M. Ince, per late F. O. Pickard-Cambridge.)

CU. (Dr. Collingwood, the late.)

Class ARACHNIDA.

Order ARANEIDEA.

Fam. DYSDERID 2.

Harpactes drassoides Sim. Chamounix. R. J. P.-C.

Segestria senoculata Linn. Simplon. A.S. A.

Fam. DRASSIDA.

Drassus hispanus L. Koch. Sulden-Tyrol. A. W. P.-C.

troglodytes C. Li. Koch. Zinal, Engstlen Alp, and Chalet

de Meléze, St. Gervais les Bains. A. W. P.-C.

Simplon. A.S. A. Zermatt. R. J. P.-C.

Switzerland. G. N.

Drassodes lapidosus Walck. Simplon. A.S.A. Switzerland. H.S.

Chamounix, R.J.P.-C. St. Gervais les

Bains. A. W.P.-C. Montreux. C.

% pubescens Thor. Arolla. A. W. P.-C.

Prosthesima apricorum L. Koch. Simplon. A.S8. A.

Es talpina L. Koch. Bel Alp. G.N.

ms petiwerti Scop. Montreux. C.

a nigrita Fabr. Chalet de Meléze, St. Gervais les

Bains. A. W. P.-C,

) latreillii Sim. St. Gervais les Bains. A. W. P.-C.

o prefica L. Koch. “Switzerland.” H.S.

Callilepis nocturna Linn. St. Gervais les Bains. A. W. P.-C.

Simplon, A.S.A. Switzerland. H.8.

Gnaphosa badia L. Koch. (@. molesta Cambr.) St. Gervais les

Bains, Arolla, and Sulden-Tyrol.

AW Pe.

+ petrobia L. Koch. Switzerland. G. N.and R. J. P.-C.

o tigring Sim. Switzerland. R.J. P.-C.

Micaria hospes Kulez. Simplon. A.S. A.

» breviuscula Sim. Zermatt. R. J. P.-C.

scenica Sim. Bel Alp. G.N. Zermatt. R. J. P.-C.

Zinal. A. W. P.-C.

» pulicaria Sund, St, Gervais les Bains. A. W. P.-C.

396 THE REY. 0. PICKARD-CAMBRIDGE ON

Clubiona hilaris Sim. Simplon. A.S. A.

Cheiracanthium italicum Can. & Pav. Zermatt. H.8.

Zora maculata Bl. Chamounix. A. W.P.-C. Montreux. C.

Fam. DICTYNIDSE.

Dictyna viridissima Walck. Alps. H.5.

» jlavescens Walck. Alps. E.S.

4 uncinata Westr. Switzerland. H.§.

Amaurobius fenestralis Stroem. Simplon, A.S. A. Zinal.

A.W.P.-C. Montreux. C.

a jferox Walck. Montreux. C.

_ 4-guttatus Hahn. Alps. E. 5.

Fam. AGELENID.

Crypheca silvicola C. L. Koch, Zinal. A. W. P.-C.

Celotes terrestris Wid. Switzerland. R.G. and A. W. P.C,

Interlaken. A. EK. EH.

» pickardii Cambr. Switzerland. H.A. P.

» atropos Walck. Chamounix. A. W. P.-C. Switzerland.

H.S8.

, Claustrarius Hahn. Switzerland. R.G., A. H.-E.,

R.J.P.-C., and G.N.

Tegenaria campestris C. L. Koch. Alps. E.S.

i; tridentina L. Koch. Simplon. A.S. A.

a derhamii Scop. Montreux. C.

‘ pusilla Sim. Chamounix. A. W. P.-C.

He domestica Clk. Montreux. C.

Cicurina cinerea Panz. Switzerland. R.G.

Textrix denticulata Oliv. Simplon. A.S.A. Bel Alp. G.N.

Montreux. C.

Agelena labyrinthica Clk. Simplon. A.S. A. Switzerland. H.5.,

R.J. P.-C.,and H. A. P. Montreux. C.

Fam. ZODARIIDS.

Zodarion gallicum Sim. Alps. H.S.

Fam. PHOLCID &.

Pholeus phalangioides Fuess. Montreux. C.

Fam. THERIDIID&.

EH pisinus truncatus Walck. Montreux. C.

Theridion riparium Bl. Switzerland. R.G.

+ bigibbum, sp. n. (text-fig. 51). Alps. H.8. (For

description see appendix, p. 403.)

SWISS SPIDERS. 397

Theridion sisyphium Clk, Simplon. A.S.A. Zinal and Engstlen

Alp. A. W.P.-C. Switzerland. H.8.

and H.A.P. Montreux. C.

* nigro-variegatum Sim. Geneva. R. de L.

i. nigro-punctatum Lae. Alps. ELS.

* blackwallii Cambr. Alps. E.S.

Phyllonethis lineata Clk. Switzerland. H. A. P.

Dipena braccata C. L. Koch. Alps. E.S.

Steatoda bipunctata Linn. Simplon. A.§8. A.

Lithyphantes corollatus Linn. Alps. E.8.

HY paykullianus Clk. Alps. ELS.

Teutana grossa C. L. Koch. Alps. ES.

Vuryopis flavomaculata C. L. Koch. Alps. E.S8.

Asagena phalerata Panz. Alps. E.S. Switzerland. G.N.

Chamounix. A. W. P.-C.

Enoplognatha thoracica Wid. Alps. B.S.

Robertus lividus Bl. Simplon. A.S. A. Arolla. A. W. P.-C.

Tapinopa longidens Wid. Alps. E.8.

Bolyphantes alticeps Sund, St. Gervais les Bains. A. W. P.-C.

Drapetisca socialis Sund. Switzerland. C. E. M. I.

Linyphia marginata C. L. Koch. Switzerland. G.N.

» triangularis Clk. Alps. E.8. Montreux. C.

» parygiana C. L. Koch. Alps. ELS.

» montana Clk. Simplon. A.S.A. Switzerland. H. A. P.

» pusilla Sund. St. Gervais les Bains. A. W. P.-C.

» frutetorum C. L. Koch. Alps. E.S.

» emphana Walck. Alps. E.S.

hortensis Sund. Alps. E.S.

Leptyphantes tenuis Bl. St. Gervais les Bains. <A. W. P.-C.

ns fragilis Thos. Zinal. A. W. P.-C.

# expuncta Cambr. Zinal. A. W. P.-C.

ae tenebricola Wid. Switzerland. G.N. Alps. E.S.

Sulden-Tyrol. A. W. P.-C.

3 pulcher Kuler. Simplon. A.S. A. Sulden-Tyrol.

A Wee

mengtt Kulez. Arolla, A. W. P.-C.

a minutus Bl. Chrissons. R. J. P.-C.

leprosus Ohl. Montreux. C.

Bath yphantes concolor Wid. Gorge of Massa. G.N. Chamounix.

R.J. P.-C.

variegatus Bl. Switzerland. G.N.

Hilaire montigena L. Koch. Eggishorn. G. N.

Tigellinus saxicola Camb. Geneva. R. de L.

Triconchus mebi Dahl-Bos. Chamounix. A. W. P.-C,

Centromerus subalpinus Lessert. Geneva. R. de L.

ay afinis Wid. (= C.pabulator Cambr.-Less.). Engstlen

Alp, or St. Gervais les Bains. A.W. P.-C.

Mieryphantes gulosa L. Koch. Eggishorn, A. W. P.-C,

Oreoneta fortunata Cambr, St. Gervais les Bains. A. W. P.-C.

398 THE REY. O. PICKARD-CAMBRIDGE ON

Microneta rurestris C. L. Koch. LEggishorn and St. Gervais les

Bains. A. W. P.-C.

A viaria Bl. Arolla. A. W. P.-C.

Gongylidium apicatum Bl. Bel Alp. G.N.

Erigone tirolensis L. Koch. Switzerland. G.N. Bel Alp. G. N.

» remota L. Koch. Bel Alp. G.N.

,» ara Bl. Alps. H.S.

» Jjugorum lL. Koch.

,, dentipalpis Wid. St. Gervais les Bains. A. W. P.-C.

Diplocephalus cristatus Bl. St. Gervais les Bains. A. W. P.-C.

Chamounix. R. J. P.-C.

a castaneipes Sim.-Lessert. Geneva. R. de L.

. kochii Lebert-Lessert. Geneva. R. de L.

a eboredunensis Cambr. Engstlen Alp. A. W. P.-C.

Lophocarenum nemorale Bl. Geneva. R.de L. (Sent to me as

LL. stramineum Menge.)

Nematogmus sanguinolentus Walck. Alps. E.S.

Araeoncus altissimus Sim. Bel Alp. G. N.

Styloctetor (sub Hilaira) carli Lessert (sent to me for examination

by Lessert). Geneva. R.de L. (Appears to

me to be rather a Styloctetor than a Hilaira.)

,, oroccha L. Koch. Geneva. R. de L.

Tapinocyba affinis Lessert. Geneva. R. de L.

"3 pallens Cambr. Geneva. R.de L.

OCeratinella brevis Wid. St. Gervais les Bains. A. W. P.-C.

Fam. ULOBORIDS.

Hyptiotes paradowus C. L. Koch. Simplon, A.S8. A.

Fam. HPEIRID®.

Subfam. TEYRAGNATHIN &.

Tetragnatha eatensa Linn, Switzerland. H.S8. Montreux. C.

Pachygnatha degeeriti Sund. Switzerland. C.K. M. I.

Meta meriane Scop. Simplon. A.S8.A. Switzerland. G.N.

Montreux. C.

» menardi Latr. Montreux. C.

» segmentata Clk. Simplon. A.S. A. St. Gervais les Bains.

A. W.P.-C. Switzerland. R.G. Montreux. C.

Subfam. ARGIOPINA.

Cyclosa conica Pall. Simplon. A.S. A.

Subfam. HPEIRIN«.

Zilla montana C. L. Koch. Switzerland. G.N. Simplon. A.S. A.

Alps. E.S.

» w-notata Clk. Montreux. C.

SWISS SPIDERS. 399

Zilla stroemii Thor. Switzerland. G.N.

Cercidia prominens Westr. Alps. E.S.

Singa albovittata Westr. Switzerland. G,N.

» hamata Clk. Alps. E.S.

Epeira nordmannit Thor. Zinal. A. W. P.-C.

carbonaria L. Koch. Switzerland. G.N.

alpica L. Koch. Switzerland. G. N.

» eirce Sav. Simplon. A.S. A.

cucurbitina Clk. Simplon. A.S.A. Switzerland. H. A. P.

Montreux. C,

diademata Clk. Switzerland. H.S., R.G., R. J. P.-C.,

G.N.,and H.A.P. Simplon. A.S8. A.

Montreux. C. ;

» ceropegia Walek. Simplon. A.S. A. Switzerland. G.N.

St. Gervais les Bains. A. W. P.-C.

Halkern Thal. A. E. E. Zermatt.

R. J. P.-C.

pyramidata Clk. Halkern Thal. A. E.E. Switzerland.

EAL P.

» marmorea Clk. St. Gervais les Bains. A. W. P.-C.

Switzerland. H. A. P.

» sclopetaria Clk. Montreux. C.

» quadrata Clik, St. Gervais les Bains. A. W. P.-C.

, dioidea Walck. Montreux. C.

» adianta Walck. Alps. E.S.

., triguttata Fabr. Geneva. R. de L.

,, sturmit Hahn (agalena Bl.). Geneva. R.de L.

,, redit Scop. Montreux. C.

» montana C. L. Koch. Alps. E.S.

Fam. THOMISID&.

Thowisus onustus Walck. Montreux. C.

Misumena vatia Cik. Simplon. A.S. A.

Diea dorsata Fabr. Montreux. C,

Xysticus cristatus Clk. Switzerland. R.G. Simplon. A.S. A.

St. Gervais les Bains. A. W. P.-C.

Montreux. C.

lanio C. L. Koch. Switzerland. R.G.

”

af gallicus Sim. St. Gervais les Bains, Arolla, and

Eggishorn. A. W. P.-C,

4 kochii Thor. Switzerland. R. J. P.-C.

Re pint C. L. Koch. Simplon. A.S.A. Montreux. C.

Fs lateralis Hahn. Switzerland. R.G.

glacialis L. Koch. Gorge of Massa and Bel Alp. G.N.

luctator L. Koch. Simplon. A.S. A.

bifasciatus C. L. Koch. Switzerland. R. J. P.-C.

erraticus Bl. Arolla. A. W. P.-C.

400 THE REY. 0. PICKARD-CAMBRIDGE ON

Philodromus alpestris C. L. Koch. Simplon. A.S.A. Zermatt.

R.J.P.-C. Arolla and St. Gervais

les Bains. A. W. P.-C. Switzer-

land. G. N.

lividus Simon. Switzerland. G.N.

laricium Simon. Switzerland. G.N.

a collinus C. L. Koch. Switzerland. C. W.

ce margaritatus Clk. Simplon. A.S,A. Switzerland.

G.N. Var.: jejunus Panz. Switzer-

land. C. E. M. I.

. emarginatus Schr. Simplon. A. S.A.

Thanatus arenarius Thor. Kggishorn. A. W. P.-C.

ee formicinus Clik. Simplon. A. S.A. Montreux. C.

Eggishorn and Sulden-Tyrol.

A.W.P.-C. Switzerland. G.N.

Fam. SPARASSID®

Micrommata virescens Clk. Switzerland. R.G. Simplon. A.S.A.

Montreux. C.

Fam. PrIsauRID™.

Pisaura mirabilis Clik. Switzerland. R.J. P.-C. and H. A. P.

Montreux. C.

Fam. LycosipD@.

Dolomedes fimbriatus Walck. Ualkern Thal. A. EK. E.

Trochosa terricola Thor. Chamounix. R.J.P.-C. Switzerland.

EvAcrias

leopardus Sund. St. Gervais les Bains. A. W. P.-C,

Switzerland. H. A. P.

Pirata latitans Bl. St.Gervais les Bains. A.W.P.-C. Montreux. C.

Tarentula andrenivora Walck. Arolla. A. W. P.-C. Switzerland.

G.N. Montreux. C.

lessertii, sp. n. (text-fig. 52). Switzerland. H.S. (For

description see appendix, p. 404.)

: pulverulenta Clk. Arolla and St. Gervais les Bains.

A.W.P.-C. Simplon. A.S. A.

Zermatt. R.J.P.-C. Switzerland.

G.N. and H.A. P.

ys trabalis Clk. Simplon. A.S.A. Switzerland. G.N.

Zermatt. R. J. P.-C.

a miniata C. L. Koch. Montreux. C.

a nemoralis Westr. Gorge of Massa and Bel Alp. G.N.

St. Gervais les Bains. A.W, P.-C.

Simplon, A.S, A. :

‘ barbipes Sund. Gorge of Massa, G. N.

SWISS SPIDERS. 40]

Tarentula aculeata Clk.-Lessert (= 7’. accentuata auctt., Lessert)-

Zermatt and Schiedeger. R. J. P.-C. Arolla.

A. W. P.-C.

Lycosa amentata Clk. Switzerland. R.J.P.-C., G. N,, and H.A.P.

Simplon. A.S.A. Engstlen Alp and

St. Gervais les’ Bains. A. W. P.-C.

» paludicola Clk. Switzerland. C.W.

» accentuata Walck.-Sim. Switzerland. F. P.S.

» pulverulenta Clk.-Sim. Switzerland. F.P.S. St. Gervais

les Bains. A. W. P.-C.

nigra O. L. Koch. Zermatt and Zinal. A.W.P.-C. Switz-

erland. G.N.and R.G, Zermatt.

BR,.d.)P.-C.

blanda C, Iu. Koch (& cursoria, auctt.). Sulden-Tyrol,

St. Gervais les Bains, Zinal, Cha-

mounix, and Arolla. A. W. P.-C.

Switzerland, Gorge of Massa, and

Bel Alp. G.N. Simplon. A.S. A.

Chamounix and Zermatt. R.J.P.-C.

» giebeli Pav. Switzerland and Zermatt. R.J.P.-C. Zinal.

A.W.P.-C. Switzerland, Bel Alp. G.N.

» riparia C. L. Koch. Switzerland. G.N. Zermatt.

R.J. P.-C. St. Gervais les Bains

and Switzerland. A.W. P.-C.

Jerruginea L. Koch. Zermatt and Sulden-Tyrol, A. W. P.-C.

. bifasciata C. L. Koch. Montreux. C.

» pedestris Sim. Switzerland. R. J. P.-C.

monticolaC, L. Koch. Aletsch Glacier, Zinal, and Switzer-

land. A. W. P.-C.: Bel Alp and

Switzerland. G.N. Eggishorn.

A.W P:2,

lugubris Walck. Rhone Glacier. R. J. P.-C. Switzerland,

G.N.

» torrentum Sim. Bel Alp and Switzerland. G.N.

» wagleri Hahn. Switzerland, Rhone Glacier, and Zermatt.

Ri. Ji. P.-C,

palustris Linn. Switzerland and St. Gervais les Bains.

AW .P eC,

», ludovica Dahl. Simplon. A.S. A. Engstlen Alp. A. W.

P.-C,

mixta Kulez, Simplon. A.S. A. Gorge of Massa. G.N.

tarsalis Thor. Simplon. A.S,A. St. Gervais les Bains

and Chamounix. A. W. P.-C.

albata L. Koch, Bel Alp. G.N. Switzerland. R.J.P.-C.

St. Gervais les Bains. A. W. P.-C.

agrestis Westr. Switzerland. H.S.

A saloiein albimana Walck. Montreux. C,

Proc. Zoou. Soc.—1912, No. XX VI. 26

402 THE REY. O, PICKARD-CAMBRIDGE ON

Fam. SALTICID4.

Epiblemum scenicum Clk. Simplon. A.S. A. Bel Alp. G.N. Arolla

and St. Gervais les Bains. A. W. P.-C.

Montreux. C.

Heliophanus metallicus Sim. Arolla and Eggishorn. A.W. P.-C.

eneus Hahn. Switzerland. G.N. Simplon. A.8.A.

Montreux. C. Arolla. A.W.P.-C.

Zermatt. R. J. P.-C.

Euophrys petrensis C.L. Koch. Arollaand Eggishorn. A.W.P.-C.

Switzerland. G.N.

if pictilis Sim. St. Gervais les Bains. A. W. P.-C.

» jrontalis Walck. Chamounix. A. W. P.-C.

* erratica Walck. Switzerland. R.G.

Sitticus longipes Can. Simplon. A.S. A.

Attulus histrio Sim. Simplon. A.S8. A.

Attus rupicola C. L. Koch. Rhone Glacier and Switzerland.

Rede. C sand iFieAgs:

5, pubescens C. L. Koch. Switzerland. G.N.

Dendryphantes encarpatus Walck. Montreux. C.

Phileus chrysops Poda. Simplon. A.S. A.

“Elurops v-insignitus Clk. EHggishorn and Arolla. A.W. P.-C.

Simplon. A.S. A.

Pellenes lapponicus Sund. Chamounix. R. J. P.-C.

Synageles venator Luc. Chamounix. A. W. P.-C.

Order PHALANGIDEA.

Homalenotus quadridentatus Cuv. Switzerland. C. W.

Phalangium opilio Linn. Switzerland. R.J. P.-C. Simplon.

A.S.A. Switzerland. H. A. P.

53 parietinum De Geer. Switzerland. H. A. P.

* brevicorne C. L. Koch. Zermatt. R.J. P.-C.

Liobunum limbatum L. Koch. Switzerland.

ss rotundum Latr. Switzerland. R.J. P.-C.

Platybunus pinetorum C. L. Koch. Chamounix. R. J. P.-C.

3 triangularis Herbst. Chamounix. A. W. P.-C.

Oligolophus morio Fabr.

Var. palliata C. L. Koch. Eggishorn and St. Gervais les

Bains. A.W. P.-C.

Var. glacialis C. L. Koch. Bel Alp and Zermatt. R.J. P.-C.

Var. alpinus Herbst. Simplon. A.S.A. Zinal, Arolla,

St. Gervais les Bains, Chamounix, and Sulden-

Tyrol. A. W. P.-C. Chamounix. R.J. P.-C.

Bel Alp. G.N.

Normal form. St. Gervais les Bains. A. W. P.-C. Switzer-

land. R. J. P.-C. and G.N.

Megabunus rhinoceros Can. Simplon. A.S. A. Switzerland.

G.N.

SWISS SPIDERS. 405

Nemastoma dentipalpe Auss. Chamounix. A. W. P.-C.

* quadripunctatum Perty. Interlaken. R.G.

Order CHERNETIDEA.

(Pseudo-scorpiones. )

Obisium jugorum L. Koch. Zermatt. R.J. P.-C.

Chthonius tetrachelatus Preyssl. Interlaken. R.G,

Order ACARIDEA.

Ceculus echinipes L. Dufour, Zermatt. R. J. P.-C.

APPENDIX.

Description of 7heridion bigihbum, sp. nu. (‘Text-fig. 51.)

Adult male, Length =; of an inch or 1°7 mm.

The cephalothorax is broad, rounded behind, Caput prominent.

Text-fig. 51.

Hh,

sal

k, "

Theridion bigibbum.

A. Cephalothorax and abdomen from above and behind.

(Nat. length =} of an inch or 17 mm.)

B. Left palpus (outer side).

C. Cephalothorax (in profile).

Clypeus high. Colour dull yellow, a wedge-shaped dark blackish

central band runs from, and includes, the ocular area to the

26*

404 THE REV. O. PICKARD-CAMBRIDGE ON

central impressions. The sides of the thorax are slightly

margined also with blackish.

The eyes are rather large, and occupy the whole of the fore

part of the caput. The four centrals form a large quadrangle

rather longer than broad and a little narrower behind. The

convexity of the curve of the hinder row is slightly directed back-

wards, and the interval between the hind-central pair of eyes is

rather greater than that between each and the lateral eye next

to it.

The legs (1, 4, 2, 8) are moderate in length and strength,

of a pale yellowish-white hue, with a slight appearance of

a dark but defective annulation at the extremities of the joints.

They are furnished with hairs and a few slender bristles only,

The palpi are short, similar to the legs in colour. The digital

joint very large, of an oval form; and the palpal organs, directed

outwards, are simple in form but completely encircled on the

margin by a long coiled black spine issuing from near the hinder

margin, from and in connection with a largish corneous process,

and ending 4 in a fine somewhat sinuous slender point underneath

the fore extremity of the joint.

Falces rather small and weak, and directed slightly forwards.

Maaille, labiwm, and sternwm normal, and, w ith the falces,

similar in colour to the cephalothorax.

The abdomen is large and of a general cretaceous white colour.

Its form (looked at from above) is somewhat subtriangular, and

at the shoulders (or each anterior fore corner) is a slight pro-

minent point or gibbosity. An obscure pattern of a dusky hue

may be traced on the hinder extremity, and the fore extremity is

also similarly suffused.

Two examples, both males, one adult the other immature, were

contained among other Swiss spiders kindly given to me some

years ago by Mons. Eugéne Simon. It is most probable that

a series of specimens would show a richer colouring, and a more

definite pattern than could be traced in these two examples, which

had apparently not yet attained their full development in these

respects. In general appearance the present species shows some

resemblance to Zheridion pallens Bl., but the decided difference

of the thoracic markings, as well as the form of the abdomen, and

the details of the palpal organs, will serve to distinguish them

without difficulty.

Description of Tarentula lessertii, sp. n. (Text-fig. 52.)

Adult female. Length 63 lines.

Of normal general form. The cephalothoraz is of a dark

brown colour, with a broad central longitudinal straight band of

a paler hue, without lateral constriction, and clothed densely

with short hoary hairs; the normal lateral converging in-

dentations are marked by rather darker lines or narrow str ipes.

SWISS SPIDERS. 405

The eye-area is dark blackish at the sides and in front; the

quadrangle of the four posterior eyes is rather broader than long,

and its anterior side is shortest; the two anterior eyes are

separated by less than a diameter’s interval, and the anterior

curved row of four eyes (of which the two centrals are much the

largest) are separated by about equal intervals.

Tarantula lessertii.

“Epigyne,” 2.

The leys are of moderate length and strength (4. 1. 2. 3),

their colour is a uniform yellow-brown without any annuli; the

normal spines are rather slender. ‘The palpi are similar in

colour to the legs. Falces normal and of a deep blackish-brown

hue; sternwm dark brown. Abdomen dark yellow-brown, its

fore-side above abundantly clothed with grey hairs, with the

characteristic central longitudinal marking on the fore part

of the upper side clothed with hoary hairs, margined with

a black line, and followed towards the spinners with a series of

indistinct angular lines, or chevrons. The under side of the

abdomen is of a uniform paler yellowish-brown colour than

the upper side, and clothed with short hoary hairs. ‘The abdo-

minal markings, however, were much obscured by the rather

shrunken condition of the abdomen, The epigyne is of a very

distinct and characteristic form (see text-fig. 52).

The example above described has been examined by Dr. Roger

de Lessert, and its species is unknown to him. I have not been

able to identify it with any known continental species, and am

therefore induced to describe it as new to science. It was

taken during a tour in Switzerland and kindly sent to me by

Mr. Henry Speyer of Reigate.

406 MR. H. G. PLIMMER ON

22 On the Blood-Parasites found in Animals in the Zoo-

logical Gardens during the four years 1908-1911. By

H. G. Pummer, F.R.S., F.Z.8., Pres.R.M.S., Pathologist

to the Society.

[Received March 2, 1912: Read March 5, 1912.}

(Plates X LIX.-LV.*)

During the last four years I have examined the blood of every

mammal, bird, reptile, and batrachian which has died in the

Zoological Gardens, 6430 in all. In 447 of these, that is about

7 per cent., 1 have found blood-parasites. These 447 affected

animals belonged to 256 different species, as in many cases the

same parasite was found in several animals of the same kind.

Many of these parasites are described or recorded here for the

first time; in other cases the hostsare new. In the tables which

follow I have indicated those which are new, or in new hosts,

by placing asterisks against the names of the animals in which

they were found. In spite of much generous help from Mr. R. I.

Pocock, F.R.S., Mr. Seth-Smith, and Dr, R. T. Leiper, to whom

my best thanks are due, it is impossible to be quite certain that

I have not given myself too much credit, partly on account of

the large amount of scattered literature on the subject, and partly

on account of the constant change of names of animals on the

part of zoologists.

Although the importance of the study of blood-parasites has

only been recognised in comparatively recent years—that iS

since 1880, when Laveran discovered the hematozoon of malaria

in the military hospital of Constantine—it should be remembered

that the first mention of a blood-parasite dates from 1841, when

Valentin described an ‘‘ entozoon ”—-as he called it— in the blood

of a fish, Salmo fario. Nowadays the importance of these

parasites is well recognised, as well as the necessity of studying

them comparatively, since we know that many of the gravest

diseases of man and of animals are caused by them. I need only

mention such diseases as malaria, syphilis, sleeping-sickness, and

kala-azar of man, and trypanosomiasis (in its various forms),

‘Texas fever, and the coccidioses of animals, to remind you of this.

I have not attempted to give names to any of those parasites

described here for the first time. In the present state of our

knowledge it seems better to tabulate the hosts and give the

general characteristics, or type, of the parasite. The custom of

naming as new—very often with very absurd names—every

similar parasite found in a new host is, I think, a very bad one.

We can only name them reasonably when we know a great deal

more about them than we do at present.

The parasitic organisms [ have found in the blood of animals

* Vor explanation of the Plates see p. 417.

PZ. 1912). SIX.

BLOOD PARASITES.

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BLOOD PARASITES.

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BLOOD PARASITES.

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BLOOD PARASITES.

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BLOOD PARASITES.

BLOOD-PARASITES, ; 407

in the Gardens belong to the two great groups of the Protozoa

and the Worms, and, as regards the elements of the blood, they

are either extra-corpuscular or intra-corpuscular,

I will begin with the extra-corpuscular parasites, those which

live in the serum, and these are the nematode worms, the spiro-

chetes, the trypanosomes, and other flagellates,

Firstly, the nematode worms —they all belong to the class

Filaria. The embryos only are found in the blood, as the adult

filarie could not pass through the capillaries. These microfilarie

are the least harmful to their hosts of any of the blood-parasites,

and all the pathological effects we know of filarize are due to the

adult worms, which have been found only in a very small pro-

portion of the cases here. The embryos are generally surrounded

by a fine capsule, which is cast off when they enter into the body

of the mosquito, when their further development takes place.

I have found microfilarie in the blood of 15 mammals of

13 different species, in 101 birds of 74 different species, and in

13 reptiles of 5 different species; and filarie have not been

found before in 88 out of these 92 species of animals, and are

new, at any rate as regards hosts. The animals in which they

have been found have come from all parts of the world, the largest

number from the Americas, but the following list will show how

wide-spread the infection is.

Emeryo FInarLe rounD IN tHE BLooD OF MAMMALS.

(All are either new hosts or new parasites, or both, except those

marked with an asterisk.)

Name. HaAprrar. Tyre or Frnaria.

Suricate (Suricata suricatta) .......... S. Africa. Long.

Black-handed Spider Monkey (Ateles C. America. Long.

geoffroyi).

Black-headed Lemur (Lemur mungos). Madagascar, Long and slender.

Crowned Lemur (Lemur coronatus)... do. Very long, striated.

White-fronted Capuchin (Cebus Ecuador. Medium, thick.

albifrons).

Golden Cat (Felis temmincki) ......... Sumatra. Long.

Bridled Wallaby (Onychogale fre- N.S. Wales. Long.

nata).

Rat-tailed Opossum (Didelphys nudi- S. America. Medium.

caudata).

Collared Peceary (Tayassu tajacu) ... do. Long.

3 Pinche Marmosets (Leontocebus Colombia. Long.

edipus).

Lion Marmoset (Leontocebus rosalia). Brazil. Long.

Javan Chevrotain (Tragulus Java. Long.

Javanicus).

Ant-eater (Myrmecophaga tridactyla) S. America, Medium.

Birps,

Himalayan Whistling Thrush (Myio- N. India. Short.

phoneus temmincki).

Lanceolated Jay (Garrulus lancco- do, Short.

latus).

408

Birps (continued).

NAME. Hasirar.

Spotted Emerald Tanager (Calliste

guttata).

2 Great-billed Touracous (Turacus

macrorhynchus).

Wood-Thrush (Lurdus mustelinus)...

S. America.

W. Africa.

N. America.

*3 Blue-faced Honey-eaters (Hntomyza Australia.

cyanotis),

Sooty Crow Shrike (Strepera fuli- do.

ginosa).

*2 Black-backed Piping Crows (Gymno- do.

rhina tibicen).

Ultramarine Jay (Aphelocoma ultra- Mexico.

marina).

Stanley Parrakeet (Platycercus W. Australia.

icterotis).

Black -headed Cat-Bird (4@lurcedus

melanocephalus).

New Guinea.

6 King Birds of Paradise (Cicinnurus do.

regius).

4 Lawes’ Birds of Paradise (Parotia do.

lawesi).

2 Count Raggi’s Birds of Paradise do.

(Paradisea raggiana).

Red-winged Francolin (Francolinus

gariepensis).

Common Cow-Bird (Molothrus pecoris.) N. America.

2 Orange-cheeked Waxbills (Hstrelda W. Africa.

melpoda).

Red-backed Shrike (Lanius collwrio).

Banded Aracari (Péteroglossus tor-

S. Africa.

Europe.

C, America.

quatus).

Cuban Mocking Thrush (Mimocichla N. America.

rubripes).

Crimson-eared Waxbill (Hstrelda W. Africa.

pheenicotis).

Black-crested Cardinal (Gubernatrix S. America.

cristata).

Violet Manucode (Phonygama chaly- New Guinea.

beata).

Fire-red Finch (Spermophila minuta). Venezuela.

American Robin (Turdus migratorius) N. America.

Red-capped Parrot (Pionopsittacus Brazil.

pileatus).

White-bellied Plumed Dove (Lopho-

phaps leucogaster).

Orange-headed Thrush (Geocichla

citrina).

2 Violet Tanagers (Huphonia violacea)

Blue-winged Tanager (Calliste cy-

anoptera).

Whydah Bird (Urobrachya albonotata) 8. Africa.

India.

C. America.

S. America.

*Garrulous Honey-eater (Myzantha Australia.

garrula).

2 Brazilian Hangnests (Icterus Brazil.

jamaicai).

2 Jay Thrushes (Garrulaax leuco- N. India.

lophus).

MR, H. G. PLIMMER ON

N. Australia.

TYPE.

Short, thick.

Short, straight.

do.

Short, thick.

Long, edges serrated.

Short, straight.

Short, thin.

Short, straight.

Short, thick.

Very short, thick.

2 kinds: one long and

thick, the other short.

Short.

Small.

Short.

Medium.

Long.

Short, straight.

do.

Very long.

Short.

2 kinds: one long and one

short.

Long.

Short, thick.

Long.

2 kinds: one long and one

short.

Medium, thick.

Long.

Short.

Long, thick.

Medium.

NAME.

6 Green-billed Toucans (Ramphastos

dicolorus).

BLOOD-PARASITES.,

Grass Finch (Hrythrura prasina) ...

Baillon’s Avacari (Andigena bailloni)

Fire Finch (Lagonosticta senegala).

Blue-cheeked Barbet

asiatica).

Shama (Ciétocinela macrura)

Dial Bird (Copsychus saularis)

(Megalema

White-collared Ouzel (Merula albo-

cincta).

Song Sparrow (Melospiza pusilla) ...

Cat-Bird ( Gialeoscoptes carolinensis).

Orange Weaver- Bird (Huplectes

Sranciseana).

Beautiful Wood-Hawk

orchis spectabilis).

(Dryotri-

Rose-breasted Grosbeak (Hedyimeles

ludovicianus).

Mexican

luxuosus).

Yellow-bellied Tanager

Slaviventris).

Blue Jay (Cyanocorax

(Calliste

Scarlet Cardinal (Cardinalis pheni-

ceus).

Baltimore Hangnest (Ieterus balti-

more).

3 Blue Tanagers (Tanagra episcopus).

Green - tailed Towhee (Oreospiza

chlorura).

Allied Saltator (Saltator senilis)

Thick-billed Violet ‘Tanager (Hu-

phonia laniirostris).

Naked-eyed Pigeon

nophthalma).

2 Yellow-winged Sugar-birds (Caveba

cyanea).

(Columba gym-

Lavender-backed Finch (Spermophila

castaneiventris).

Black Manucode (Manucodia atra)...

Black-throated Hangnest

gularis).

3 Blue Birds (Sialia sialis)

Brown-eared Bulbul (Hemixusflavala)

(Icterus

Chilian Starling (Cureus aterrimus).

White-throated Jay Thrush

lax albogularis).

(Garru-

Rainbow Bunting (Cyanospiza le-

clancheri).

Rufous-necked Tanager

cayana).

Levaillant’s Francolin (Francolinus

levaillanti).

Golden-eyed Babbler

sinense).

Scoter (demia nigra) ......

Grey-necked Serin Finch

canicollis).

(Calliste

(Chrysomma

(Serinus

HABITAT,

Guiana.

Sumatra.

Brazil.

W. Africa.

India.

do.

N. America.

do.

Africa.

W. Africa,

N. America.

Mexico.

S. America.

do.

N. America.

S. America.

N. America.

Brazil.

do.

Venezuela.

S. America.

Guiana,

Aru Islands.

Mexico.

N. America.

India.

Chili.

India.

Mexico.

S. America.

Cape Colony.

India.

British Isles,

S. Africa,

409

TYPE.

Short, very thick.

Long, thin.

Thick.

Long, thin.

Long.

Short, thin.

Short, thick.

Medium, thick.

Short, thick.

2 kinds: one short, and one

long with very pointed

ends.

Long.

Short.

Medium, straight.

Long.

Short to medium, thick.

Long.

Short, thick.

Short, straight.

Long.

Long, thick.

Long.

Short, thick.

Very long.

Short.

Short, straight,

Short.

Medium,

Long.

Short, thick, no vacuole.

Long, very pointed,

Long, thick.

Long.

Striated,

Long.

410 MR. H. G. PLIMMER ON

Birps (continued),

Name. Haprrar. Typr.

Capueira Partridge (Odontophorus S. America. Long.

capueira).

Occipital Blue Pie (Uvrocissa occi- India. Short.

pitalis).

Barred Dove (Geopelia striata) ...... do. Short, very thick.

White-backed Piping Crow (Gymno- Australia. Short, thick.

rhina leuconota).

ReEprriues AND BATRACHIANS.

Bearded Lizard (Aimphibolurus bar- Australia. Short, thick.

batus).

Spiny-tailed Iguana (Cyclura acan- C. America. do.

thura).

8 Giant Toads (Bufo marinus)......... 5. America. do.

Pine.Snake (Pitwophis sayi)............ N.America. Long.

*2 Edible Frogs (Rana esculenta) ...... Kurope. Short.

I now come to the Protozoa, and will take first a spirochete,

whieh belongs to the so-called Proflagellata. The spirochetes are

important as being the cause of such diseases as relapsing fever,

tick fever, and syphilis.

This particular one was found in the blood of a Monkey—Cer-

copithecus sabeus—and is of the type of Spirocheta recurrentis,

the cause of relapsing fever.

SPIROCH ELA FOUND IN THE BLOOD OF A

Namen. HaABirat. CHARACTER,

Green Monkey (Cercopithecus sabeus). Sierra Leone. Like Sp. recurrentis.

The next in order are the Flagellates, and of these 1 will take

first the Trypanosomes, which are flagellated organisms living in

the blood serum, ‘hey are the cause of many deadly diseases

in man and animals.

I have found Trypanosomes in 50 mammals, 7 birds, and

3 amphibia. 49 of the mammals were ordinary rats, out of 500

examined at the Gardens for another purpose, in which the

ordinary rat trypanosome, 7/rypanosoma lewist, was found. The

other mammal was a Fat Mouse (Steatomys pratensis) from

S. Africa, in which trypanosomes have not hitherto been described.

'The bird trypanosomes are all new in these particular birds ; but

they would appear to belong to the general type of Trypanosoma

avium. These bird trypanosomes are quite different to the mam-

malian variety: they are very large and thick, they stain densely

and indistinctly, and they move very slowly. The spleens of the

infected birds were large and their blood anemic, ‘The trypano-

somes found in reptiles were all of the type of Z'rypanosonia

rotatoriwm, which was the one described by Gruby in 1843. It

has not been described before in the Tree Frog (Hyla venulosa).

BLOOD-PARASITES,. Al]

'TRYPANOSOMES FOUND IN THE BLoop or MAMMALS.

NéMe.

Fat Mouse (Steatomys pratensis) ............

*49 Rats (Epimys norvegicus)

CHARACTER.

Of Nagana type.

T'. lewisi.

Hapirar.

S. Africa.

England.

Birps.

Red-winged Francolin (Francolinus S. Africa.

Lon]

gariepensis).

T’. avium type.

Chukar Partridge (Caccabis chukar) ...... India. do,

Marsh Bird (Leistes guianensis) ............ Demerara. do.

Waxbill (Hstvelda melpoda) Australia. do.

Levaillant’s Francolin (Francolinus levail- Cape Colony. do.

lanti).

Dial Bird (Copsychus saularis) India. do,

Bower Bird (Amblyornis subalaris)......... New Guinea. do.

BATRACHIANS.

*Edible Frog (Rana esculenta) ........ ...... Europe. T. rotatorium type.

*Tree Frog (Hyla arborea)..................... do. do.

Tree Frog (Hyla venulosa) S. America. do.

There are certain flagellated organisms which may appear in

the blood and live there not as actual parasites, but accidentally.

In reptiles there are certain kinds of inflammation of the intes-

tine, accompanied by ulceration, which cause the walls of the

intestine to become permeable, so that some of the flagellated

organisms which are often found there are able to get into the

blood and live there. The alteration of structure of the intestine

during the inanition of hibernation is also favourable to the

passage of organisms through the intestinal wall. I have found

these flagellates in the blood of 8 reptiles, and they have been

either varieties of Heaamitus, found first by Dujardin in 1841 in

the frog’s intestine, or of Trichomonas: the former in the frog

and tortoises, the latter in the snake.

The only other mention of these organisms in the blood is by

Danilewsky, in 1889, who found the Hexamitus in the blood of

Emys lutaria and Rana esculenta, so that those given in the

following list are all new hosts for this kind of infection. In the

Hewamitus infections there was general edema of the tissues and

ascites, and the organisms were found also in the transudation.

INTESTINAL ORGANISMS FOUND IN THE BLoop or REpriLes AND

BATRACHIANS,

NAME. Haspirat. TYPE oF PaRasiTE.

Bull Frog (Rana catesbiana) ............... N. America. Hexamitus.

Leopardine Snake (Coluber leopardinus). Europe. Trichomonas.

Indian Cobra (Naia tripudians) India. do.

Box Tortoises (Cistudo carolina) .........

Three-banded Terrapin (Cyclemys tri-

Sasciata).

Hog-nosed Snake (Heterodon simus) ......

West-African Python (Python sche)......

Angulated Tortoise (Testudo angulata)...

N. America.

I. Indies,

N. America.

W. Africa.

S. Africa.

Hexamitus.

do.

Trichomonas.

do.

Hexvamitus,

412 MR. H,

G. PLIMMBER ON

[ now come to the intracellular parasites, and will take first

the Hemogregarines, which are intracellular parasites occurring

chiefly in ‘the reptile group, and, so far as my experience here

goes, ‘entirely in that group.

I have found them here in 150

reptiles, of 67 different species, and in 41 of these for the first

time. The schizont is found generally in the erythrocyte of the

host, and is an elongated body with a well-marked nucleus, and

with no pigment. Se shizogony occurs

generally in the internal

organs, especially in. the lungs ; this stage, however, has not yet

been observed in all eases,

They vary much in their effect on

the cell and on the nucleus, and I have indicated in the following

table the most obvious of these

gregarines very little change takes place in the cell; with others

the cell gets smaller and out of shape; with others it gets

enormously enlarged

and

effects. With some hemo-

dehemoglobinised. Others again

destroy the nucleus, like the Aa@ryolysus of lizards and snakes.

Some infections are very severe,

scarcely a cell is unaffected,

and there may be sometimes as many as four parasites in a cell,

T have found a very remarkable one of a new type ina Bengal

Monitor. ‘This parasite divides the nucleus of the cell into two

parts which remain connected by a thread of nuclear material

which has a tiny enlargement at its central point, T have

found that the schizogony of this parasite takes place in the lung,

several stages of which are shown in the Plates.

H.®MOGREGARINES FOUND IN

Name.

¥3 Moorish Geckos (Tarentola meau-

ritanica).

Bengal Monitor (Varanus benga-

lensts).

%2 Rat-tailed Serpents (Zachesis

lanceolatus).

+ SS i een Bs i

2 Teguexins (Zupinambis tequevin).

3 Black- spotted Lizards (lgi-

reides nigropunctatus).

*9 Indigo Snakes (Coluber corais)...

Vivaceous Snake (Tardophis fallax)

¥3 Lace Monitors (Faranus varius).

*Horseshoe Snake (Zamenis hippo-

erepis).

*6 Indian Pythons (Python molwrus).

20 Diamond Rattlesnakes (Cre-

talus atrox).

*King Snake (Coronella getula)

HABITAT.

Mediterranean.

India.

S. America.

do.

Adniatic.

C. America.

. Europe.

Australia,

S. Europe.

India.

Texas,

N. America.

tHE Buoop oF Rerriwes AND BAYTRACHIANS.

CHARACTERS.

Parasites large and doubled over at one

end, sometimes at both.

Nucleus of cell divided into two by

the parasite, with connecting thread.

Schizogony in lungs.

Host-cells enlarged.

Parasites bulky and very full of granules ;

many with only remains of nucleus

attached,

Short to medium, some bottle-shaped ;

chromidia at both ends; free sporonts

in all.

Host-cells enlarged and dehemoglobi-

nised,

Medium ; cells unaltered.

Parasites large and contain granules ;

cells unaltered.

Bulky ; turned over at one end.

Host-cells often deformed; generally

diminished in size.

Host-cells enlarged and dehemoglobi-

nised,

Very thick, short forms; some free

sporonts which are long and pointed,

Name.

*Hoary Snake (Paeudaspia cana) ...

Puff Adder ( Bitia arictana)

*Mexican Snake (Coluber melano-

leuens).°

Senegal Chameleon

senegalensis).

#6 Cobras (Naia tripudiana)

Large Grieved Tortoise (Podo-

Cnemis ExXpansa).

*Alligator (Alligator mississippi-

(Chameleon

ensis).

2 Clarke’s Lizards (Sceloporus

clarkii).

*3 Diamond Pythons (Python

spilotes).

Gallot’s Lizard (Lacerta galloti)...

*5 Indian Rat Snakes (Zamenia

MUCOBA).

* Msculapian Snake (Coluber longis-

simus).

42 Common Boas (Boa constrictor).

Reeves’s Terrapin (Damonia recvesi)

* Reticulated Python (Python reticu-

latus.)

4 Chicken Snakes (Coluber obso-

letus).

2 Corn Snakes (Coluber quttatus) .

Panded - tailed Snake (Leptophis

liocercus).

4 Spiny-tailed Snakes (Cyclura

acanthura).

*3, Coach-whip Snakes (Zamenis

flagdliformis).

*3 Pond Tortoises (Emys orbhicu-

laria).

Painted Terrapin (Chrysemys

picts).

Smaller Rattlesnake (Sistrurus

miliarius).

Boddaert’s Snake (Drymohius hod-

daerti).

3 Black Snakes (Zamenis con-

strictor).

Hog - nosed Snake (Heterodon

simus).

Mocassin Snake (Tropidonotus

fasciatus).

Pine Snake (Pituophis sayi).........

*4 Giant Toads (Bufo marinus) ......

*2 Spanish Terrapins (Clemmys

leprosa).

Horrid Rattlesnake (Crotalus

horridus).

4 Bell's Cinyxes (Cinirys helliana),

8S. America.

N. America.

do.

Australia.

Canary Islands,

India.

Europe.

S. America.

China.

E. Indies.

N. America.

do.

S. America.

C. America.

N. America.

8. Europe.

N. America.

do.

C. America.

N. America.

do.

S. America.

Spain.

N. America.

C. Africa.

BLOOD-PARASITES. 43

HAeirat. CHARACTERS.

§. Africa. Short, bulky, and full of granules; howt-

cells deformed.

do. Medium ; host-celle unaltered.

Mexico. Host-cells very much enlarged.

W. Africa. Short, bulky.

India. Long parasites; cells unaltered.

Bulky and granular, with eccentric

nuclens.

Parasites small, granular; nucleus often

eccentric.

Long, with dense nuclei, some hooked ;

cella greatly altered.

Very long, doubled over.

Adherent to nuclens of host-cell ; granules

at both ends.

Medium sized; cells unaltered.

Medium «ized; adherent to nucleus of

host-cell.

Long and hooked ; some cells enlarged.

Short, thick. with eccentric nuclei.

Long, doubled over, cells enlarged.

Host-cells enlarged.

do.

Medium ; cells unaltered.

large and granular, cell nearly filled;

nucleus compressed and dense.

Host-cells deformed; parasite long and

hooked at one end, later donbled over.

Short and thick, with eccentric nuclei;

a few very thin and doubled over.

Short and thick, with granules: cell«

deformed.

Long and thick, with eccentric nuclei.

Bulky and hooked; cells deformed.

Large and hooked; cells deformed.

Long; cells enlarged and ultimately

dehamoglobinised.

Thin and long forms; cells unaltered.

Host-cells enlarged and deh#moglobi-

nised.

Short and thick; cells unaltered.

Bulky ; cells deformed,

Long and thin; cells unaltered.

Short, thick.

414

Name.

Black Sternothere (Sternotherus

niger).

Hasselquist’s Gecko (Ptyodactylus

lobatus).

2 Speckled Terrapins (Clemmys

guttata).

Green Tree Viper (Atheris chlor-

echis).

Three-keeled Terrapin (Stawo-

typus triporcatus).

Egyptian Gecko (Tarentola annu-

laris).

Hilaire’s Terrapin

hilarii).

*4 Kyed Lizards (Lacerta ocellata).

(Hydraspis

*Nilotic Monitor (Varanus niloticus).

*2 Cook’s Tree Boas (Corallus

cooki).

Green Tree Snake (Dendraspis

viridis).

*2 Russell's Vipers (Vipera russelli).

*2 Hryxs (Hryx johnt)................0

3 Dahl’s Snakes (Zamenis dahli)...

*Confluent Rattlesnake (Crotalus

confluentus).

Glass Snake (Ophiosaurus apus) ...

Long-nosed Viper (Vipera ammo-

dytes).

Tuberculated Iguana (Iguana

tuberculata).

Peloponnesian Lizard (Lacerta

peloponnesiaca).

Madagascar Boa (Boa madagas-

cariensis).

West African Python (Python

sebe).

2 Dark Green Snakes (Zamenis

gemonensis).

Square-marked Toad (Bufo regu-

lanis).

HABITAT.

W. Africa.

Egypt.

N. America.

W. Africa.

C. America.

Egypt.

Brazil.

8. Europe.

Africa.

W. Indies.

W. Africa.

India.

do.

S. Europe.

N. America.

S. Europe.

do.

C. America.

S. Europe.

Madagascar.

Tropieal Africa.

Europe.

Africa.

MR. H. G. PLIMMER ON

CHARACTERS.

Medium sized.

Short; cells deformed.

Short; bulky; nucleus eccentric.

Long and thin.

Medium.

Short, bulky; cells deformed.

Long, doubled over.

Host-cells enlarged and some dehemo-

globinised. Long, thin; some bottle-

shaped.

Medium, thick.

Host-cells deformed, short; nucleus not

central.

Large, often two in cell which is not

enlarged.

Bulky : cells deformed.

Long, with hooked end.

Long, thin; turned over at both ends.

Long, doubled over.

Large and granular; host-cells elongated.

Bulky ; cells not enlarged.

Short, stout, sometimes round.

Short.

Short, thick ; cells deformed. Schizo-

gony found in liver.

Medium sized.

Cells enlarged and dehwmoglobinised ;

parasites long and thin.

Short and very thick.

The next group is that of the Plasmodide, to which the various

malarias belong.

I have found only one mammal—a Green Monkey—infected

with, malaria, probably Plasmodium kocha.

It is uncertain,

because I could only get certain stages; the host-cells do not

seem to be greatly altered.

In the birds there are two kinds of intra-corpuscular parasites,

the Plasmodium precow and Hemoproteus danilewskyi, which

have been sometimes confused, but are really entirely distinct.

The Plasmodium precow has many points of similarity to the

parasites of human malaria, and in many stages is so like that it

can only be differentiated by the presence of the oval nucleus of

BLOOD- PARASITES. 415

the bird’s erythrocyte. I have found Plasmodium precor in 20

different species of birds, in none of which has it been recorded

before. Geographically the distribution of the 21 birds in which

I have found this parasite is as follows :—KEurope 1, Asia 4,

Africa 5, Americas 8, Australia 3. It is a deadly disease in

birds, and produces a marked anemia, and considerable enlarge-

ment of the spleen and changes in the bone-marrow.

[ have found Plasmodia in 6 reptiles of 5 different species, in

all for the first time. They all probably fall under the heading

of Hemocystidium. They all have pigment, and when large have

rather the appearance of Hamoproteus.

altered.

PLASMODIAL PARASITES FOUND IN THE

The cells are not

Bicop or

MammMats.— Plasmodiwm kochi.

NAME.

*Green Monkey (Cercopithecus sabeus) ..................

Hapirar.

Sierra Leone.

Birps.— Plasmodium precoc.

Long-tailed Glossy Starling (Lamprotornis eneus) .

Japanese Hawfinch (Coccothraustes melanura)

2 Crowned Cranes (Balearica regulorum)

Orange-cheeked Waxbill (Estvrelda melpoda) .........

Small Hill Mynah (Gracula religiosa)..................

Red-and-Blue Macaw (Ava macao) ee Reese VEN

Tambourine Pigeon (Tympanistria bicolor) 2.22.2...

Red-eared Bulbul (Pyenonotus jocosus) ba dib ess

Nutcracker (Nucifraga caryocatactes).................

Yellow-breasted Tanager (Calliste thoracica) .........

Blue Sugar-Bird (Daenis cayana) .......000..............

Brazilian Hangnest (Icterus jamaicdi)..................

Ox-Bird (Tenor alector) ... cerervees pep ell):

Flycatcher (Stoparola melanops) ........................

Jay-Thrush (Garrulax leucolophus) .......000...5. 0005.

Mexican Finch (Carpodacus mexicanus) .............

Rainbow Bunting (Cyanospiza leclancheri)

Blue-tailed Fruit Pigeon (Carpophaga concinna)

Cuban Finch (Phonipara canora) .....

Scaly Dove (Scardafella squamosa) .....

W. Africa.

Japan.

S. Africa.

W. Africa.

India.

C. America.

W. Africa.

India.

Europe.

Brazil.

S. America.

Brazil.

W. Africa.

Australia.

N. India.

Mexico.

Aru Islands.

Cuba.

S. America.

Reptites.—Hemocystidium.

Bell’s Cinixys (Cynivis helliana) wae

Three-tailed Terrapin (Staurotypus triporcatus)

Painted Terrapin (Chrysemys picta)

Home's Cinixys (Cinixys homeana) ....................

2 Eroded Cinixys (Ciniwys erosa) ...................

Africa.

B. Honduras.

N. America.

W. Africa.

The second parasite of birds is Hamoproteus danilewskyi, which

has been found in a large number of cases :

is unknown.

and its development

Tt is found first as a small irregular body in the

416 MR. H, G. PLIMMER ON

erythrocyte of the bird and then grows in the long axis of the

cell, and turns round the end of the nucleus, and it contains

pigment. It has been shown by the Sergents and by Aragao

that a species of fly—Lynchia maura—is the principal host.

The process of fertilization in these parasites has been described

by MacCallum, and it is believed that other stages are found in

the lungs.

I have found these parasites in 69 birds of 48 different species,

in 46 of the latter for the first time.

The following list shows that the parasite is very widely

distributed. 76 of the hosts came from Africa and 42 from the

Americas.

H_&MOPROTEUS DANILEWSKYI FOUND IN Brrpbs.

Name. Hapirar.

3 Blue-winged Tanagers (Calliste cyanoptera) ...... S. America.

Bouvier’s Owl (Scotopelia bouviert) ...... 2.0... Lagos.

3 Great-billed Touracous (Turacus macrorhynchus) W. Africa,

Black-shouldered Tanager (Calliste melanonota) ... Brazil.

Porphyrio (Porphyrio madagascariensis) ............ Madagascar.

2 Green-headed Tanagers (Calliste tricolor) ......... S. America.

Red-sided Kelectus (Heleetus pectoralis) ............ New Guinea.

White-backed Piping Crow (Gymnorhina lewconota) Australia.

Hanging Parrakeet (Loriculus galgulus) 0.0.0.0... Malacca.

3 Brown-necked Parrots CS Oa ge ce W. Africa.

Bower Bird (Amblyornis subalaris) ....... .... New Guinea.

5 Yellow-winged Sugar-birds (Cereba nea cee S. America.

5 Maequeen’s Bustards (Houbara macqueeni) ...... W. Asia.

3 King Birds of Paradise (Cicinnurus regius) ...... New Guinea.

Dial Bird (Copsychus saularis) ..c....cc.cc cece ecee ees India.

*African Barn Owl (Strix flammea) .........0000.... S. Africa.

Grenadier Weaver Bird (Huplectes oryx) ............ W. Africa.

Parrot Finch (Mrythrura psittacea) ......... 060.6... New Caledonia.

Australian Sheldrake (Tadorna tadornoides) ...... Australia.

2 White-crested Touracous (Luracus corythaix) ... S. Africa.

Mexican Hangnest (Cassiculus melanicterus) ..... Mexico.

Palm Tanager (Tanagra Bear) Sedona RAGE RE ny OSHA CTUCAs

Honey-eater (Péilotis fusca) ......ccn6cskeeecuseeseee Australia.

Schlegel’s Dove (Calopelia puella) .........cccc cece W. Africa.

Cut-throat Finch (Amadina fasciata) ......cc W. Africa.

Scops Owl (Scops leucotis) .........se.eseeseeeeeeeeeeeee Gambia.

Black-throated Lorrikeet (Trichoglossus nigrigu- New Guinea.

laris).

Grass Finch (Erythrura prasinad) v......ccec00.... Sumatra.

Baer’s Duck (Fuligula baert) .............sceceeeeess India.

2 Kestrels (Tinnunculus alaudarius) ....... British Isles.

Spotted-backed Weaver (Hyphantornis “ga TeaceTo) Africa.

Festive Tanager (Calliste festiva) ..........c0000. Brazil.

Rose-breasted Grosbeak (Hedymeles ludovicianus). N. America.

Senegal Touracou (Twracws persa) .......0000. ~~ W. Africa.

*Cape Hagle-Owl (Bubo capensis) .......c00.c eee Ss Africa.

Hooded Siskin (Chrysomitiris cue TEARS veces IN. America,

Fuscous Honey-eater (Ptilotis fusca) .........0.0.... Australia,

BLOOD-PARASITES. 417

Name. Hapirar.

Whydah Bird (Vidua paradisea) ..................... W. Africa,

Manchurian Crane (Grus japonensis) ................... N. China.

Lawes’ Bird of Paradise (Parotia lawesi) ............ New Guinea,

Eagle-Owl (Bubo maculosa) .............................. S, Africa,

Indian Roller (Coracias indica) ....................... Yndia.

Yellow-cheeked Tit (Machlolophus wanthogenys) ... do.

2 Blue Tanagers (Tanagra episcopus) ............... 8S. America,

Fraser’s Eagle-Owl (Bubo poensis) ...................... W. Africa.

Cape Sparrow (Passer arcuatus) ..... ................... ®. Africa.

2 Purple-capped Lories (Lovins damicella) ......... Moluceas.

Purple Sugar-Bird (Careba cyanea) ................... S. America.

One other parasite remains to be mentioned—the Leuco-

cytozoon: this is found in the blood of birds in the form of a

long, spindle-shaped, unpigmented body, about three or four

times the length of a normal erythrocyte, and about one third

of the breadth. Unstained they are colourless and are always

free from pigment. They are rare, and at present uncertain as

to position, and they occur only in the blood in the sexual forms.

The dull plasma staining may indicate that the forms in the blood

are macrogametes, The earlier observers of this parasite—

Danilewsky and Ziemann—believed that the host-cell was a

leucocyte, but Laveran has shown that it is an erythrocyte.

There would seem to be some analogy between the curious

enlargement of the host-cell and the enlargement and dehzemo-

globinisation of the erythrocytes of snakes, containing one of the

varieties of hamogregarines mentioned before.

I have found these parasites in two birds only, both of which

are new as hosts,

LEUcoCcYTOZzOA FOUND iN PHE BLOOD oF Birpa.

NAME. Haprrat,

Scaup’s Duck (Fuligula mavila) ............-0000....... Europe.

Levaillant’s Francolin (Fyancolinus levaillanti) ... Cape Colony,

EXPLANATION OF THE PLATES.

The microphotographs were made for me hy Dr. A. Norman, and the drawings

were made to scale by Mr. W, S. Berridge.

The scale divisions are ‘01 millimetre.

Pu, XLIX,

Fig. 1. Pilavia in the blood of a Wallaby. x 250. The eapsyle can be seen with

a lens gs a white line outside the filaria.

The blood in this case was very disorganised and there was a marked

-eycocytosis, In the ponch there was a very young foetus in the body-

cavity of which there was one adult filaria similar to others found in the

body-cayity of the mother, No embryo tilariz were found in the blood

ot the foetus.

Fig. 2, Filavie in the blood of a Giant Toad. > 300. The capsule is quite visible,

and also the pointed anterior end.

These were present in large numbers.

Proc. Zoot. Soc.—1912, No. XX VII. 27

Hic, 3

Fig. 4,

Fig. 5

Fig. 6

Vie. 7

Fie, 8

Fig. 9

Fig. 10.

Iie, Il.

Fig. 12

Fig. 13.

Fig. 14.

Fig. 15

Fig. 16

Fig. 17

Fig. 18.

Fig. 19.

ON BLOOD-PARASITES.

. Spivochetes in the blood of a Green Monkey. X 1000.

The monkey had also malaria (not shown).

. Trypanosomes in the blood of a Dial Bird. They are large and thick:

the one marked a is probably degenerating, and that marked 6 is of a

different shape and stained more easily.

. Intestinal organism of Trichomonas type in the blood of a Leopardine

Snake. 1000.

. Intestinal organisms of Hexamitus type in the blood of a Bull Frog,

Pits 1,

. Filariez in the blood of a Pinche Marmoset. X 135. Present in very large

numbers. The capsule is not distinguishable.

. Hemogregarines in the blood of a Hog-nosed Snake. X 500.

A very severe infection, nearly every erythrocyte contaimed one or more

parasites. At a parasites which have escaped from the erythrocytes and

are free in the blood are seen, and at 6 a cell containing so-called male

and female forms.

Veit, Mil.

. A drawing of various forms of the same Hewxamitus type of organism of

which a photograph is shown on Pl. XLIX. fig. 6.

A scale-drawing of the Hemogregarine found in the blood of a Hog-nosed

Snake, of which a photograph is given on PI. L. fig. 8.

a. An erythrocyte containing two parasites of so-called male and

female types. 6. Two parasites free in the blood, one hyaline, the other

very granular. ¢. Schizogony; two cysts from the lung. d. Also from

the lung; probably an earlier stage than ec.

A scale-drawing of Heemogregarines in the blood of a Black Snake. At

a are two parasites joined together by the remains of the nucleus of the

erythrocyte. 6 points to a number of probable odkinete forms found

in the stomach of a tick found upon the snake.

. Hemogregarines in the blood of a Rat-tailed Serpent. > 500. This isa

late stage of the infection, showing the parasites attached to the remains

of the nuclei of the erythrocytes.

Hemogregarines in the blood of a Mexican Snake. X 300. Ata can be

seen infected erythrocytes which are enormously enlarged, very much

thinned, and entirely dehamoglobinised.

Pru. LID.

Drawing of Hzemogregarines from the blood of a Black-spotted Lizard.

Showing enlargement of the corpuscles and the various forms assumed

by this parasite. At @ is an erythrocyte very much enlarged and altered

in shape and texture, and vacuolated. At 6 are two free forms, showing

considerable nuclear activity and very granular protoplasm.

. An erythroevte from the bone-marrow of the same Black Snake to which

Fie. 11 refers. X 750. It contains two hemogregarines of the so-called

male and female type.

. Drawing of Hemogregarines from the blood of a Madagascar Boa. En-

larged cells containing one and two parasites. At @ is a cyst from the

lung, showing commencing schizogony.

. Drawing of a leucocytozoon from the blood of a Scaup’s Duck. At a are

deformed erythrocytes containing the parasite; at 6 remains of nuclei of

erythrocytes; at c a macrogamete; and at d probably a microgamete.

Pr. LIII.

Blood of a Brazilian Hangnest, showing infection with Plasmodium

precox. X 450. Many of the infected cells show a multiple infection

as at a.

Blood of a Crowned Crane showing the same parasite as Fig. 18. The

blood is very anemic. There are single and multiple infections of the

erythrocytes, and at a is a parasite breaking up into a rosette.

PZ Ss ole el Eis

: A.S.C.

AR.Broom del. Huth,Lith? London.

THE AUDITORY REGION IN DIGYNODON.

ON THE AUDITORY REGION IN DICYNODON. 419

Pr LDV.

Fig. 20. Blood ofa Palm Tanager showing infection with Hemoproteus danilewskyi.

Three full-sized parasites are present, and some eariy forms are seen in

the photograph as dots in the erythrocytes.

Fig. 21. Blood of a Bengal Monitor showing an ordinary hemogregarine and the one

mentioned on p. 412 of text. 450. This is probably a double infection

as no intermediate stages between these parasites can be seen.

At a the ordinary form of hemogvegarine is seen; those erythrocytes

marked 6 contain the new type of parasite, to which reference has been

made in the text. This parasite stains badly, there is a very small

amount of nuclear material in it when stained by any modification of

the Romanowsky method or by iron-logwood, and the altered nucleus of

the erythrocyte stains very densely.

Fig, 22. Drawings of the same parasites. At a the ordinary hemogregarines are

shown, and at 6 the other parasite, in an early stage in the drawing to

the left, and in the later stage, with the cell-nucleus divided, to the

right.

12 ay, LV.

Fig. 23. Drawings of the new parasite.

The erythrocyte in the photograph and drawings is seen to he deformed,

and eventually dehemoglobinised (as in the cell marked a). At 6 is

seen the tiny nodule which is generally to be found ou the delicate con-

necting-thread between the pieces of the cell-nucleus. The cell marked

¢ contains what appears to be an abortive attempt to divide the cell-

nucleus equally: very few of these were found.

Fig. 24. Drawings of forms found in the lungs, showing various stages in schizogony.

None of these smal! forms have been found in the erythrocytes. No

stages were found in the other organs.

23. On the Structure of the Internal Har and_ the

Relations of the Basicranial Nerves in Dieynodon, and

on the Homology of the Mammalian Auditory Ossicles.

sy R. Broom, M.D., D.Se., C.M.Z.S.

{Received January 12, 1912: Read Mareh 5, 1912.]

(Plate LVI.* and Text-figure 53.)

Few questions connected with vertebrate morphology have

given rise to a larger amount of discussion than the homology

of the mammalian auditory ossicles and the connected problem

of the fate of the quadrate in mammals, Most embryologists

have been struck with the resemblance of the malleus and inecus

to an articular and quadrate ; many comparative anatomists have

been impressed rather with the resemblance of the mammalian

auditory chain with the auditory apparatus of the Reptilia;

while paleontologists, for the most part, have hesitated to

believe that the incus could be the quadrate, from the difficulty

of imagining the intermediate stages, and have inclined rather to

believe that the quadrate has become part of the zygomatic arch,

or the meniscus, or become lost, or converted into the tympanic

bone.

Until a few months ago I was of opinion that the quadrate

became either the meniscus or, in those few forms where a meniscus

is absent, was lost. It is unnecessary to repeat the arguments

* Por explanation of the Plate see p. 425,

27%

420) DR. R. BROOM ON THE

by which I have endeavoured to maintain this position; suttice it

to say that I now regard the position as untenable, since the

element in the Anomodonts which I have long regarded as the

tympanic now proves to be unquestionably the stapes.

In the type skull of Dicynodon latifrons there is a slender rod-

like bone lying in the cavity which is bounded above by the

exoccipital, and usually by a dumbbell-like bone below. The

stout dumbbell-like bone which stretches from the basioccipital

process to the quadrate I have hitherto believed to be the

tympanic, and the slender rod-like bone the stapes. New

evidence now shows that whatever the slender bone may be—

perhaps a displaced part of the hyoid—the supposed tympanic is

undoubtedly the stapes, and what I believed to be the fenestra

ovalis is the foramen for the vuth nerve.

Having the imperfect occiput of a moderate sized Dicynodon,

in which the matrix was much hardened by epidote and the

bone mainly calcareous, I saw it might be possible to dissolve the

bone and leaye a cast of the internal ear and the basicranial

nerve-passages. Though on one side much of the internal ear

was found to be occupied by calcite, the result of the decalcification

on the whole was so satisfactory that the structure of the ear in

Dicynodon is now almost as well known as in recent animals, and

as much is known of the cranial nerves as is ever likely to be

known.

The internal ear (Pl. LVI. figs. 1-3) is unlike that of any

animal hitherto described, so far as I am aware. While the

semicircular canals are apparently not unlike those of recent

reptiles or mammals, the vestibule is very remarkably elongated

and there is no trace of a cochlea.

The vestibule has an upper, somewhat triangular portion,

which lies between the upper end of the canal for the viith nerve

and the upper end of that for the ixth, xth, and xith nerves.

In close contact with the brain cavity there is an upward

extension, with which is united the canal common to the anterior

and posterior semicircular canals. On the anterior and posterior

sides are irregular excrescences, which are perhaps vascular

spaces filled with matrix. The broadest part of the vestibule is

where the ampulle of the anterior and posterior semicircular

canals are situated. From this point the vestibule passes down-

wards for a distance about twice as great as the maximum width.

To near the middle of this descending portion there is a steady

narrowing, but beyond the middle the vestibule again expands and

the fenestra ovalis is about as wide as the widest part of the vesti-

bule. The fenestra ovalis lies near the middle of the descending

basioccipital process, but looks outwards and slightly downwards.

This large descending process, so conspicuous in the Anomodont

skull, is formed posteriorly by the basioccipital bone, and in front

it is supported by the basisphenoid, but there seems considerable

reason for believing that the centre portion is partly prootic and

partly opisthotie. Down the descending part of the vestibule there

appears to be evidence of the suture between the two otic bones.

AUDITORY REGION IN DICYNODON, 491

The semicircular canals are lost from the left side of the

specimen, and though they are preserved on the right side, as they

are partly filled with calcite and the bone is much « epidotised, it 1s

impossible satisfactorily to decalcify the right side. Still, sufficient

has been successfully accomplished to show at least a part of each

anal.

The posterior semicircular canal has a distinct ampulla, which

hes very close to the canal for the ixth, xth, and xith nerves.

From here it passes upwards and forws ards, then inwards, and

lastly downwards, doubtless uniting with the upper end of the

anterior canal.

The horizontal canal, though well shown in its middle region,

has neither its anterior nor posterior ends displayed. The position

of the canal will be better understood from the figure than from

description.

The anterior semicircular canal is badly displayed. It runs

mainly upwards to unite with the descending part of the posterior

canal.

The cast of the posterior portion of the cranial cavity and of

the basicranial nerves is beautifully preserved.

Behind the lower end of the posterior semicircular canal is seen

the cast of the large canal for the ixth, xth, and xith nerves;

where it leaves the cranial avity if is shghtly extended antero-

posteriorly, but where it leaves the skull it is flattened a little

vertically.

The canal for the xiith nerve passes down a short distance

behind the jugular foramen, and unites with it before it leaves

the skull. The fact that the xiith nerve has only a single canal

is interesting in view of the fact that in the Cotylosauria, as

exemplified by Pareiasaurus, we have two foramina for the xiith

nerve, and a similar condition is found in the Cynodontia and in

the Marsupialia. In the Pelycosauria, as has been shown by Case,

and in the Eutheria, there is, as in the Anomodontia, only a

single foramen for the xiith nerve.

The canal for the viith nerve lies a little in front of the

vestibule. It is long and narrow and passes downwards and

slightly outwards, and ends in the small foramen seen at the

bottom of the deep groove between the back part of the basi-

sphenoid and the pterygoid. ;

In front of the specimen is seen the deep pituitary fossa, and

below it the carotid canal. The latter begins at a small rounded

foramen near the front of the posterior processes of the basi-

sphenoid. The main part of the canal passes upwards, forwards,

and slightly inwards, and unites with the canal of the opposite

side just in front of the hypophysis. From the lower end of the

canal is given off a smaller canal, which passes upwards and

somewhat more forwards. While the large canal is pretty

certainly that for the carotid artery, the smaller is possibly for

the arteria ophthalmica. Whatever be the condition of the

arteries within the cranial cavity, it must have been very unlike

that of Sphenodon oy other living reptiles, and though perhaps a

422 DR. R. BROOM ON THE

little more like the condition in mammals, the resemblance could

not have been very close.

The stapes has long been an element which gave rise to

difficulties. So far as | am aware, the first author who gave a

good figure of the bone was P. Fischer, in 1870, and he regarded

the element asthe pterygoid. From nearly all the British Museum

skulls it has been removed during development, but it still remains

and is well shown in the skull which forms the type of Aiste-

cephalus chelydroides. Unfortunately, unlike most of Griesbach’s

work, his figure of this skull in Owen’s Catalogue is rather poor,

and in the description Owen says nothing about the bone in

question. But in any case, as Owen mistakes a portion of the

squamosal for the quadrate (his “tympanic”), though the quadrate

itself is present in a fairly well-preserved condition, it is manifest

that he had not a very clear idea of the structure of this part

of the skull. Seeley, when studying the Anomodont skull in

1888, took the massive stapes to be a malleus, and the unquestion-

ably homologous bone in the Cynodonts he later described as

possibly a ‘straight cochlea.” Hitherto, I have always looked

upon it as the tympanic. Now we know that it is certainly the

stapes, though so very unlike the stapes of any living form.

Though a considerable number of Anomodonts are known

showing the stapes in position and in perfect condition, I shall

describe and figure it and the related bones from the beautiful

specimen which I have recently made the type of Oudenodon

kolber (Pl. LVI. figs. 4-6). The stapes is here a short dumbbell-

shaped bone with the ends flattened and shghtly oblique. Across

narrowest part of the middie measures 12 mm. across. When

viewed from behind the stapes is seen to be much flattened, the

narrowest part of the bone measuring only 4 mm. in thickness,

A considerable part of the inner end has probably been carti-

laginous. The outer end is much flattened and is closely articu-

lated to the inner side of the quadrate. There seems to be no

supra- or éxtra-columella, or if there has been one it must have

been relatively very small.

The quadrate, one would readily think, would be the most easy

bone in the skull to identify. Many years ago Owen figured what

he believed to be the quadrate, and nearly everyone has agreed

with his determination. There is a large articulation, about as

broad as long. Along its inner third there runs an antero-

posterior ridge; near the middle there is an antero-posterior

shallow groove ; and the outer part of the articulation is a convex

disc. This outer part forms the base of a bone which on passing

upwards becomes a wide fan-like squamous plate, which is closely

applied to the squamosal. If the whole articulation be quadrate

this squamous plate must also be quadrate, as has been the view

of Owen, Seeley, and myself. It would now appear that about

half of the articulation is formed by the quadrato-jugal, and that

the squamous plate is also thus quadrato-jugal.

The squamosal comes down behind the quadrate and quadrato-

AUDITORY REGION IN DICYNODON. 423

jugal almost to the articulation. A shallow but distinct groove

can be traced down, partly formed by the squamosal and partly

by the exoccipital. This is evidently for the external auditory

canal, but how the tympanic membrane was attached or whether

there was one at all, the evidence does not show. Not improbably

there was a membrane attached to the exoccipital, squamosal, and

to a small part of the quadrate.

Though the articular region is more perfectly known in the

Anomodonts than in other Therapsida, a good deal is known about

its condition in the other groups. In the Pelycosaurs the stapes

is large and of a peculiar shape, the quadrate is aiso large and

forms the whole of the articulation, and a small quadrato-jugal is

present. Little is known of the condition in the Dromasauria.

In the Dinocephalia the quadrate is large and forms the whole

articulation, and a quadrato-jugal is apparently always present.

It is known in Delphinognathus, Moschops, and Tapinocephalus.

The stapes is probably very similar to that in the Anomodontia.

In the Therocephalia the quadrate is small, but forms the whole

articulation. No quadrato-jugal is known in any genus. The

stapes is not well known, but appears to be large andlong. Inthe

Cynodonts a quadrato-jugal is never present and the quadrate is

usually small. In some genera (e. g. Cynognathus) a considerable

part of the articular surface for the lower jaw is formed by the

Squamosal, the quadrate forming only about three-fourths of the

joint. The stapesin Bawria and Cynognathus is a fairly strong cy-

lindrical rod : in 7'rirachodon it is also a long rod, but very slender.

Inal] these genera its outer end is firmly attached to the quadrate.

As there are very strong reasons for believing that the mammals

are descended from one of the mammal-like reptiles, probably a

primitive Cynodont, all the information that can be obtained on the

structure of the auditory apparatus is especially welcome, because

of the light it may throw on the nature of the mammalian auditory

ossicles. And the certain determination of the stapes seems to

settle the homologies of the various elements. Had the outer

end of the stapes been free and given attachment to a carti-

Jaginous extrastapedial, the dispute might have gone on. But

there is apparently no extrastapedial, and certainly the greater

part of the outer end of the stapes is firmly fixed to the

quadrate. The similar attachment of the stapes to the incus in

mammals is striking; and though I have long advocated another

view, I now feel forced to agree to the view that the incus is the

reptilian quadrate, as advocated so many years ago by Reichert.

The shipping out of the quadrate from the joint is not now a

very serious difficulty, for in Cynognathus we see it already partly

slipped out. Nor is there any very great difficulty in under-

standing how the new dentary-squamosal joint arose. In Cyno-

gnathus, the posterior end of the dentary nearly takes part in the

joint, and there is no difficulty in realising the next stage in the

evolution, where the dentary and articular together hinged on

the squamosal and the quadrate. As the direct articulation of the

dentary on the squamosal became more firmly established, the

2A DR. R. BROOM ON THE

A. Diagrammiatic representation of artictilation in Dicynodon.

B. Articulation in the Cynodont, Cynognathus.

C. Hypothetical diagram of the condition when the dentary articulates with the

squamosal.

D. Hypothetical diagram of the condition wher the articular and angular no longer

support the jaw.

E. Hypothetical diagram of the condition when the articular and angular take on

their auditory function.

F. Diagram representing the perfecting of the auditory function (the mammalian

condition).

AUDITORY REGION IN DICYNODON. 495

quadrate, articular, and angular degenerated, and might have

been lost had not the attachment of the stapes to the quadrate

compelled them to take on an auditory function.

The. tracing of the steps by which the articular became the

malleus and how the manubrium was formed is a matter of some

difficulty in our ignorance of the position of the tympanic

membrane, but we may assume as very probable that soon after

the dentary-squamosal joint was formed, the angular and articular

rapidly degenerated and ceased to move with the jaw. The

tympanic membrane, which possibly in the Cynodonts lay behind

the plane of the articulation, probably in the early mammals

became shifted more forwards and inwards, and the articular came

to support the membrane by the development of the manubrium,

while the angular also took part in the support of the membrane

and became the tympanic bone. Gaupp is probably correct in

regarding the goniale as not the homologue of the angular, but a

distinct membrane-bone closely associated with the articular.

In the series of six diagrams given (text-fig. 53, A-F), the

probable course of the evolution of the auditory ossicles is shown

trom the Cynodont type to the type found in the mammal.

Though the first figure (A) represents the Anomodont type, this

must not be looked upon as on the direct mammalian line, but as

a specialised modification of the ancestral type. In fig. B we have

represented the Cynodont type as seen in Cynognathus. The

articulation is formed by the articular hinging on the quadrate

and squamosal, The dentary alinost reaches the joint. Fig. C

represents the probable condition in the later Cynodont type

when the dentary came to take part in the articulation. Fig. D

represents the later stage when the quadrate has completely

slipped out from the joint and the articular and angular have

degenerated and no longer move with the dentary. In fig. E we

have represented what was the probable condition when the

articular and angular first became specialised for their auditory

function. This is almost the Monotreme condition. Fig. F

represents the condition in the typical higher mammal.

EXPLANATION OF PLATE LVI.

A.S.C. Ascending semicircular canal; B.O. Basioccipital; B.S. Basisphenoid ;

Car. Carotid artery; H.O. Exoccipital; E.P¢. Epipterygoid?; ¥.O. Fenestra

ovalis; F.M. Foramen magnum; H.S.C. Horizontal semicircular canal; Ju.

Jugal; O.C. Occipital condyle; P.S.C. Posterior semicircular canal; Pé. Ptery-

goid; Q.J. Quadrato-jugal; Qu. Quadrate; Sq. Squamosal; S¢. Stapes; 7y.M.

Tympanic membrane; Vesé. Vestibule.

Fig. 1. Left side of decalcified brain region of small Dicynodont. Nat. size.

. Right side of partly decalcified brain-region of small Dicynodont. Nat.

size.

3. Cast of osseous labyrinth of small Dicynodont, restored from the facts

revealed by the two sides. Nat. size.

4. Quadrate region of Oudenodon kolbei. Reduced about one half. Front

view.

. Quadrate region of Oudenodon kolbei. Reduced about one half. Lower

view.

6. Quadrate region of Oudenodon kolbei, Reduced about one half. Back

view.

496 PROF. G. O. SARS ON

24. Zoological Results of the Third Tanganyika Expedition,

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

Report on some,larval and young stages of Prawns from

Lake Tanganyika. By Prof. G. O. Sars, C.M.Z.S.

[| Received December 5, 1911: Read March 5, 1912.}

(Plates LVII.-LX.*)

INTRODUCTION.

In the tow-nettings taken by Dr. Cunnington during the Third

Tanganyika Expedition and placed in my hands for examination

of the Copepoda and Ostracoda, some larval and young stages of

Prawns were also found. Having submitted these stages to a

closer examination, I find that some notes about them may be of

interest. We owe to Prof. E. von Daday a rather elaborate

account of the postembryonal development of Caridina iilotica,

var. gracilipes (=C. wyckii de Man) from Lake Victoria Nyanaza,

and I have also myself had occasion to examine some larvee

from that lake, occurring in the samples taken by Dr. Cunnington.

They agreed on the whole very well with the descriptions and

figures given by Prof. Daday. On the other hand, the larvee found

in the samples from Lake Tanganyika have proved to be rather

different, apparently owing to the fact that they belong to genera

quite different from Caridina.

In the present paper I propose to describe three larval forms

and one young one, representing as many different stages of

development. Two of these forms undoubtedly fepresentl very

early larval stages (so- called Zoée); but they differ conspicuously

both in size ad in the development of the appendages, and

apparently belong to two quite different kinds of prawns. Of

course it is very difficult to decide with any claim of. certamty

as to the species or even genera to which these larve are

referable ; but I believe that they are in themselves interesting

enough to merit a detailed csserpnen: They are here simply

recorded as Zoéa Nos. 1 and 2, though some suggestions about

their probable origin will be sth forth. The 3rd form isa larva

in the last, so- called Mysis stage; and the 4th is a very young

prawn in the Ist postlarval stage. Both these forms admit of

being more certainly referred to a definite species. At the close

of this paper some general remarks will be given.

Zoiia No.1. (Pl. LVIL)

This larva undoubtedly represents the very first postembryonal

stage of some Tanganyika Prawn, having apparently just been

hatched. It is remarkable for its small size and the very simple

structure of the appendages, being indeed the most primitive

* For explanation of the Plates see pp. 489-440.

Peano. IOIACPE UVil,

London Stereoscopic Co. imp

G.0. Sars.

PRAWNS FROM LAKE TANGANYIKA.

Pees Vola. Play:

London Stereoscopic Ce. imp.

G.0.Sars.

PRAWNS FROM LAKE TANGANYIKA.

i ger lole. re Lia.

G.0. Sars.

PRAWNS FROM LAKE TANGANYIKA,

FZ. VOLE An ae

G.0. Sars. London Stereoscopic Co. imp.

PRAWNS FROM LAKE TANGANYIKA.

PRAWNS FROM LAKE TANGANYIKA, 427

Oaridean larva that [ have ever met with, and in so far is very

different from the earliest stage of Caridina wilotica described by

Prof. Daday under the name of Muzoéa.

The length of the specimen examined, measured from the

anterior edge of the eyes to the end of the caudal plate, is only

70 mm., and the present larva is accordingly not even half as

large as the earliest stage of Caridina wilotica, the length of

which, according to Prof. Daday, is 1‘70 mm. If we imagine the

larva curled up in its original embryonic attitude, we may

approximately determine the length of the ovum from which it

has escaped to have been 0°25 mim. by a width of O15 mm, This

agrees pretty well with the measurements of the ova in Limmno-

caridina spinipes given by Dr. Calman, and as the form of the

antennal scale in the present larva agrees better with that in this

species than with that in any other of the Tanganyika prawns,

I am led to the suggestion that it perhaps might be referable

to L. spinipes, of which indeed an adult specimen was found in

one of the four samples in which the present larvee occurred,

The general form of the body (Pl. LVII. figs. 1 & 2) is

moderately slender, with the anterior division somewhat tumid

and rounded oval in shape.

The carapace is evenly convex above and slightly emarginate

behind in the middle, leaving the dorsal part of the hindmost

segment of the trunk uncovered. At the antero-lateral corners a

well-defined spine is present pointing obliquely outwards (see

fig. 1). The anterior part of the carapace is badly defined, and

no true rostrum could be detected, though a narrow stripe is

seen exténding forwards in the middle, without, however, forming

any projection in front of the eyes. The latter ave rather im-

perfectly developed, as compared with these organs in most other

Caridean larve, They are apparently quite immobile, being

contiguous at the base inside, and do not project beyond the

lateral faces of the carapace. The corneal part is only faintly

defined and contains a comparatively small and irregular

mass of pigment, from which the visual elements radiate in the

form of slight stripes, no corneal facets being as yet visible.

Immediately beneath the front parts of the eyes a semilunar

lobe occurs (fig. 3, oc.), and within this lobe the simple eye

(ocellus) may be faintly traced, for which reason it is here

termed the ocellar lobe.

Of appendages only the two pairs of antennse and the oral

parts are as yet present, and all these appendages exhibit a very

simple structure, being filled up with indifferent cells. Especially

are the mandibles and maxille remarkable for their imperfect

development, and indeed these parts seem to be quite unfit for

any true mastication, the larva in all probability subsisting in

this stage exclusively on the nutritive yolk, of which a con-

siderable quantity is seen accumulated within the anterior part

of the body.

The relative position of the appendages is best seen on viewing

428 PROF. G. O. SARS ON

the animal from the ventral face (fig. 3). The two pairs of

antenne (a & a7) originate on each side of a median slightly

convex area, the hypostome (H), the one pair, the antennule (a’),

occurring in front and inside the other pair, the antenn (a).

From the hind part of the hypostome the flap-shaped anterior lip

(L) is seen to arise covering the inner parts of the mandibles.

The latter (M) are easily observed as two comparatively large,

transverse, bow-shaped eminences, and immediately behind them

in the middle the two rounded lobules of the posterior lip (/) are

clearly seen. Behind these parts the two pairs of maxille have

their place, the 1st pair, or anterior maxille (m'), lying inside and

slightly in front of the 2nd (m*), which are extended obliquely

outwards. The maxille are followed by three pairs of much larger

appendages (mp'—mp°) exhibiting a very similar appearance, each

consisting of a short basal part and two diverging unequal rami, the

inner one (endopodite) forming a short incurved stem, the outer

one (exopodite) a much larger, laterally extended natatory ramus.

These three pairs of appendages, which are separated in the middle

by a somewhat irregular flattened area (the sternal plastron),

represent the maxillipeds in the adult animal. Behind them not

even the slightest trace of any pedal buds is to be detected.

The posterior division of the body, or metasome (figs. 1 & 2),

is rather slender, being about twice as long as the anterior

division, and is composed of six well-defined segments, the last of

which is much the longest and very narrow. This segment passes

without any intervening suture into the caudal plate. The latter

is rather unlike that found in most other Caridean larvee and more

resembles in shape that part in the protozoéa stage of Penwus. As

in that form, it is cleft by a deep median incision into two slightly

diverging lobes obtusely rounded at the end. Each lobe carries

six finely ciliated setz of rather unequal length, the outermost

one and the innermost but one being considerably smaller than

the others. The 4th seta, counted from the outer side, is the

longest, equalling in length the last segment to the bottom of the

median incision. In some of the larve, which otherwise agreed

with that here figured, a very small additional seta occurred

inside the others on each lobe (fig. 12). On the anterior caudal

segments slight traces of the epimeral plates could be detected,

those of the 4th and 5th segments terminating in an acute corner.

The ventral faces of the segments are somewhat convex ; but not

the slightest trace of pleopoda is as yet observable.

Structure of the Appendages.

The antennule (fig. 4) are comparatively small, and each consists

of a simple biarticulate stem extended forwards. The proximal

joint is much the larger and represents the peduncle. It 1s,

however, quite simple, without any trace of a subdivision and

exhibits no armature whatever. The distal joint is much

narrower than the proximal one, and also shorter, of oblong-oval

PRAWNS FROM LAKE TANGANYIKA, 429

form, and movably articulated to the former. It carries on the

tip three or four very delicate bristles, one of which seems to be

sensory in character. This jot undoubtedly answers to the

outer flagellum in the adult animal. Of the inner flagellum not

the slightest trace is to be found in the present larva, whereas

in all other larvee known to me it is at least indicated by a strong

seta attached to the inner distal corner of the peduncle,

The antennee (fig. 5), like the antennule, are each composed of

two sharply defined joints, the proximal of which forms a

thickish basal part imperfectly subdivided in the middle, and

produced at the end inside to a short conical prominence indicating

the place where subsequently the flagellum will be developed.

The distal joint, representing the antennal scale, is considerably

longer than the basal part and of narrow linear form, Its outer

edge is perfectly smooth and terminates in a well-marked tooth-

like projection, whereas the inner edge carries in its outermost

part four strong ciliated sete attached to as many well-marked

ledges and cur ved somewhat outwards. In most other Caridea

known to me the antennal scale exhibits in the earlier stages of

the larval period a rather different structure, its distal part being

narrowly produced and divided by well-marked transverse sutures

into several successive joints. Of such a subdivision not the

slightest trace is found in the present larva, nor is the distal part

of the scale produced beyond the tooth of the outer corner, this

tooth in other Caridean larvee being formed only at a much ‘later

period.

The mandibles (fig. 6) have the inner, or masticatory, part

somewhat expanded and divided by a slight median incision

into two lobes; but neither of these lobes eehihis the slightest

trace of any armature.

The anterior mawille (fig. 7) ave trilobate, with the outermost

lobe (palp) well defined as a distinct joint. The middle, or

masticatory, lobe is somewhat more prominent than the innermost,

or basal lobe. All three lobes extend obliquely inwards and have

only very slight traces of marginal spines or setz.

The posterior mawxille (tig. 8) have the form of oblong oval

plates, the edges of which are irregularly indented or divided into

a number of rounded lobules. ‘The two outermost of these lobules

are separated by a somewhat deeper incision, and represent the

one the exopodal, the other the endopodal part, the remaining

three lobules of the inner edge representing the true masticatory

lobes in the adult animal. As in the anterior maxille, only very

slight rudiments of sete are present on the lobules.

The three pairs of mawillipeds (figs. 9, 10, & 11), as above stated,

are of a very similar structure and only differ in the relative size

of the endopodite. The basal part consists of two imperfectly

defined segments and is a little broader and more flattened in the

Ist pair (fig. 9) than in the other two, The endopodite forms

a slightly incurved cylindric stem and gradually increases in

length from the Ist to the 3rd pair. In the Ist pair it is

430 PROF. G. O. SARS ON

searcely longer than the basal part and divided only into three

joints. In the other two pairs (figs. 10, 11) the middle joint

appears faintly subdivided behind the middle. Very small sete in

process of formation are seen on the terminal joint, and partly

also on some of the other joints inside. The exopodite, which is

movably attached to the outer distal corner of the basal part, is .

in all three pairs of exactly the same appearance, forming a very

. flexible, somewhat flattened stem with four long cilated sete at

the end, two of them issuing from a minute apical joint. In the

two posterior pairs another rather small seta is seen issuing inside

at some distance from the end.

Occurrence.—Several specimens of this peculiar larva, all closely

agreeing both in size and structure, were obtained from four

different samples. Two of these samples were taken at Niamkolo

(S. end of lake), another at Mbete (likewise at the south end of

the lake), and the fourth at Sumbu (8.W. of lake).

ZOE Now 2e (Ele aiivain)

The differences between this and the preceding larva are so

conspicuous, that I think we are justified im assuming that

it belongs to a different kind of prawn. It is of much larger

size, and in some respects exhibits considerably more advanced

development. Yet the imperfect structure of the oral parts, in

connection with some other features, would seem to prove that

it in reality represents a very early larval stage.

The length of the specimen examined is about 1°60 mm., and

the size of this larva is accordingly more than twice that of the

preceding one, and nearly equals that indicated by Prof. Daday

for the earliest observed stage (Huzoéa) of Caridina nilotica. The

present larva cannot, however, by any means belong to that

species, which does not occur in Lake Tanganyika, but must

be derived from some other form of prawn, in which the ova

are of a similar size to those in the said species. Consulting

again the paper of Dr. Calman, we find that in this respect only

one of the Tanganyika prawns would seem to come under con-

sideration, viz., Caridella cunningtoni Calman, the ova of which

are in reality much larger than in any of the other forms

examined. Jam therefore led to the conclusion that the present

larva in all probability may be an early stage of that form.

As compared with the preceding larva, the body (PI. LVIII.

figs. | & 2) appears rather short and stout, with the anterior diy1-

sion considerably dilated and the posterior one less slender.

The carapace is somewhat gibbously vaulted in its anterior

part, and, as in the preceding larva, is slightly emarginated

behind in the middle. At the antero-lateral corners a well-marked

spine occurs; but this spine does not project laterally (fig. 1).

Anteriorly the carapace is produced into a well defined, though

quite simple narrow spiniform rostrum, which projects distinctly

in front.

PRAWNS FROM LAKE TANGANYIKA, 431

The eyes are of very large size and somewhat project beyond

the lateral faces of the carapace (fig. 1). They are, however,

like those in the preceding larva, still imperfectly separated in

the middle and apparently quite immobile. The corneal part is

well defined and contains a semi-oval assemblage of a dark pig-

ment, from which numerous distinctly developed visual elements

radiate. The ocellar lobe is partly visible in the dorsal aspect of

the animal immediately in front of the eye-bases, but appears

more distinctly on viewing the animal from the ventral face.

As to the appendages, fig. 3 will give a general view of their

form and arrangement in the present larva. On comparing this

figure with fig. 3 on Pl. LVIL., it is at once seen that both

the antennule and the antenne are considerably larger and also

more advanced in development. On the other hand, the oral

parts (anterior and posterior lips, mandibles and maxille) are

very little different. The same is also the case with the three

pairs of maxillipeds; but these appendages appear in the present

larva more densely crowded and more remote from the hind limit

of the mesosome, a considerable space being left behind them,

which is oceupied by three additional pairs of limbs not found in

the preceding larva. These limbs, representing the three anterior

pairs of legs are, however, still only in process of formation, being

quite immobile and folded beneath the mesosome.

The metasome (figs. 1 & 2) is not fully twice as long as the

anterior division of the body, and, as in the preceding larva,

consists of six segments, the last of which is very narrow in its

anterior part, but gradually expands distally, to form the caudal

plate. The latter is comparatively smaller than in most other

Caridean larvee, and looks rather different from that in the pre-

ceding larva, not being cleft into two lobes, but only slightly

emarginated behind in the middle (see also fig. 11). Each half

of the plate carries the usual number of marginal sete, viz. 7,

the outermost and innermost ones being rather small, the

others nearly equal-sized and comparatively shorter than in the

preceding larva. Of uropoda or pleopoda no traces are to be

found.

Structure of the Appendages.

The antennule (fig. 4) have the peduncle rather prolonged and

of cylindrical form, with a slight indication of subdivision into

three joints. The outer flagellum, as in the preceding larva,

consists of a single oblong oval joint carrying on the lip four

unequal bristles, the innermost one distinctly ciliated. The inner

flagellum, as in most other Caridean Zoée, is replaced by a strong

ciliated seta attached to the inner distal corner of the peduncle.

This seta, however, scarcely exceeds half the length of the

peduncle, whereas in the earliest stage of Curidina nilotica it is,

according to Prof. Daday, nearly twice its length.

The antenner (fig. 5) exhibit the three principal parts well

defined. ‘The basal part scarcely differs in structure from that in

432 PROF. G. 0. SARS ON

the preceding larva ; but at its distal inner corner, in place of the

simple conical projection found in that larva, a well-defined

cylindric stem has been formed representing the flagellum. This

stem is about the length of the basal part and exhibits a slight

trace of subdivision behind the middle. It terminates in a spini-

ocess from the base of which a rather long ciliated seta

originates. The antennal scale is rather unlike that in the

preceding larva, and agrees more in structure with that generally

found in Caridean Zoée. It exceeds the flagellum by about one

quarter of its length and is of narrow oblong form, slightly curved

outwards. Its distal part is narrowly produced and divided by

well-marked transverse sutures into three successive joints

gradually diminishing in size. The scale carries inside and at the

tip a regular row of eight ciliated sete and has, moreover, outside

the tip a very small hair-like bristle. Another similar bristle

is also seen originating from the outer distal corner of the

form pr

proximal joint.

The mandibles and mawxille do not seem to exhibit any essential

difference in their structure from those in the preceding larva,

and are scarcely as yet functionally developed.

The three pairs of mawillipeds (figs. 6-8) likewise exhibit much

the same structure, though the difference in size of the endopodite

is still more sharply marked than in the preceding larva, that of

the Ist pair (fig. 6) being extremely small as compared with those

of the two succeeding pairs, the size of which is also somewhat

unlike.

Of the three pairs of imperfectly developed appendages suc-

eceding the maxillipeds the two anterior ones (fig. 9), representing

the developing chelipeds, ave bifid or cleft into two nearly equal

digitiform rami, one of which is the endopodite, the other the

exopodite. ‘The posterior pair (fig. 10) are still undivided and

shorter than the two anterior ones. They represent the first

pair of pereiopoda, the last two pairs being not yet formed. All

these appendages exhibit a very simple structure and are filled

up with indifferent cells, no articulation or setous armature being

observable.

Oceurrence.—Of this larva also several specimens, exactly

agreeing with each other, were found. They occurred in two of

the samples, the one taken at Niamkolo, the other at Mbete,

both located at the south end of the lake,

Larva No. 3. (Pl. LIX.)

This larva represents a much more advanced stage than the

1g ones, and therefore can be determined with more

T think that I am right in considering it as the

the so-called Mysis-stage, of Limnocaridina

parvula Calman. The solitary specimen which has come

under my notice, was found in a sample which contained several

young specimens of that prawn, and both as to size and general

two precedir

certainty.

last larval stage,

PRAWNS FROM LAKE TANGANYIKA, 433

appearance there was a striking resemblance between them and

the present form, the larval character of which, however, is at once

apparent by the presence of well-developed natatory exopodites on

all the legs, except the last pair.

The length of the specimen examined was 2°60 mm., measured

from the tip of the rostrum to the end of the caudal fan.

The general form of the body (Pl. LIX. figs. 1 & 2) is rather

slender and exhibits the characteristic Caridean aspect, the meta-

some being well developed and distinctly bent in the middle,

The carapace has the dorsal face somewhat flattened, with a

faint cervical suleus in front of the middle and the posterior

edge emarginated dorsally, exposing a part of the last segment of

the trunk. At the antero-lateral corners a small spine is seen,

somewhat remote from the margin. The rostrum is very small

and still quite simple, without any trace of denticles.

The eyes are comparatively large and project considerably

beyond the lateral faces of the carapace. They ave distinctly

separated in the middle and freely mobile, being attached to each

side of a narrow transverse eminence located just beneath the

base of the rostrum. The corneal part is well defined from the

thickish pedicles, and has the visual elements very distinct.

The dimbs of the anterior division are complete in number, five

pairs of true legs being present. All these, except the last pair,

carry outside natatory exopodites of the same structure as those

on the maxillipeds, making in all seven pairs of such appendages.

The metasome is more than twice as long as the anterior

division and much more powerfully built than in the two pre-

ceding larve. On the five anterior segments the epimeral plates

are distinctly developed, though they do not as yet extend beyond

the ventral face of the segments. The 31d segment is more

vaulted than the others and has the posterior edge somewhat

bowed in the middle, advancing over the base of the succeeding

segment. The 6th segment is very narrow, and considerably

longer than the two preceding segments combined. It carries at

the end a well-developed caudal fan, consisting of a median piece

(telson) and two movable lateral appendages (uropoda), each

divided into two setiferous lamelle. he five preceding segments

earry each a pair of ventral appendages (pleopoda) exhibiting all

the chief parts found in the adult animal, but these appendages

seem not yet to have entered into function as swimming organs.

Structure of the Appendages,

The antennule (fig, 3) have the peduncle divided into three

well-defined joints, the first of which is much the largest and

exhibits outside near the base a lamellar expansion terminating

in a small anteriorly curving denticle, Some very delicate bristles

are seen issuing from the outer distal corner of this and the

succeeding joint, and from the inner corner of each of the two

outer joints a somewhat stronger plumose seta originates, pointing

Proc. Zoo. Soc,—1912, No, XXVIII, 28

434 PROF. G. 0. SARS ON

straight forwards. Both flagella are distinctly defined and of

about equal size. They are, however, still rather short, scarcely

exceeding half the length of the peduncle, and do not exhibit any

trace of subdivision. At the tip each flagellum carries a bundle

of small bristles, and on the outer flagellum, moreover, two bundles

of delicate sensory filaments have developed, attached to well-

marked ledges of the anterior margin.

On the antenne (fig. 4) the scale is rather large and oblong-

oval in form, with the outer edge straight and perfectly smooth,

terminating in an acute tooth-like projection. The distal part

of the scale is obliquely truncated and projects somewhat beyond

the tooth of the outer edge. It is, like the inner slightly convex

edge, fringed with a regular row of slender ciliated sete, about

sixteen in number. The flagellum considerably projects beyond

the scale and points straight forwards. At its base a short joint

has been cut off representing the peduncular part. The terminal

part of the flagellum is still quite simple, cylindric, and without

any trace of subdivision.

The mandibles (fig. 6) have the cutting-edge divided into a

number of small denticles, behind which some minute bristles are

attached. The molar tubercle may also be distinctly traced.

The anterior maaille (fig. 7) exhibit a structure nearly agreeing

with that found in the adult animal, the middle, or masticatory

lobe being rather strong and securiform in shape, with a row of

well-developed denticies along the straightinner edge. ‘The very

small terminal lobe, representing the palp, is attached outside

the masticatory lobe to a well-marked ledge, and carries on the

tip three rather slender bristles. The basal lobe is cordiform and

fringed with ciliated sete.

On the posterior maxille (fig. 8) the exopodal plate is now well

defined and mobile, its anterior part bemg edged with slender

finely ciliated sete, and the posterior part exserted to a narrow

lappet carrying on the tip a few sete. Of the lobes belonging to

the endopodal part the outermost one is cut off as a distinct

joint, representing the rudimentary palp or endopodite proper ;

the remaining three lobes are edged with well-developed spines

and setee, and constitute the masticatory part of the maxilla.

The three pairs of mamillipeds (figs. 9-11) are less conspicuously

transformed, though the 1st pair (fig. 9) by the greater size of

the basal part and the considerable reduction of the endopodite

somewhat approach the structure of these appendages in the

adult animal. The characteristic difference between the two

posterior pairs observed in the adult animal is, however, not

yet apparent, the endopodite being in both (figs. 10 & 11) of a

very similar appearance and agreeing with that found in earlier

larval stages, with the only exception that the sete clothing the

joints are more fully developed. The exopodites in all three

pairs are unchanged, acting still as natatory organs.

The two anterior pairs of legs, or the chelipeds (fig. 12), are of

exactly the same structure, the endopodite of both terminating in

PRAWNS FROM LAKE TANGANYIKA. 435

a didactylous hand. They are, however, still comparatively short,

and do not exhibit the full number of joints, the ischial joint

being not yet separated from the meral one, The hand js also

imperfect in so far as none of the fingers is as yet mobile, Both

fingers are clothed at the obtuse extremity with a few very slender

bristles, and similar bristles are also attached inside the other

joints. The natatory exopodite is about the Jength of the endo-

podite, and of exactly the same structure as in the maxillipeds.

The three succeeding pairs of legs, the pereiopoda (figs. 13 & 14),

are a little longer than the chelipeds, and have the endopodite of

cylindric form and more or less curved, being composed of four

well-defined joints, The last joint, or dactylus, is conical in

form and carries on the tip a slender spine accompanied by a few

bristles. Similar bristles, partly of considerable length, are also

attached to the inner side of the other joints. The two anterior

pairs (fig, 13) carry well-developed natatory exogpodites of the same

structure as those on the chelipeds, whereas the last pair (fig, 14)

do not exhibit any trace of such appendages.

The pleopoda (figs. 15 & 16), as in the adult animal, consist each

of a somewhat flattened, oblong quadrangular basal part and two

terminal, lanceolate lamellz, apparently answering to the endo-

podite and exopodite. ‘These lamellze, however, still only exhibit

very slight traces of marginal setz in process of formation. On

the Ist pair (fig. 15), as in the adult animal, the inner lamella is

rudimentary ; on the other pairs (fig. 16) it is well developed,

though somewhat shorter than the outer one, and carries inside,

at some distance from the base, a small digititorm appendage.

The telson (fig. 17) has the form of an oblong quadrangular

plate, well defined from the last caudal segment, and slightly

exceeding half its length. It is armed at the nearly transversely

truncated extremity with eight spines of unequal size, the two

outermost being much the longest and somewhat diverging, the

two innermost and the outermost but one on each side rather

small,

The uwropoda (fig, 17) have the outer plate of a shape very similar

to that of the antennal scale; the inner one is a little shorter and

lanceolate in form, Both plates are fringed with ciliated sete

and extend somewhat beyond the telson.

Occurrence,—The solitary specimen described above was found

in a sample taken at Sumbu (S.W. of lake). In the same sample

also occurred a few yery young specimens of Limnocaridina

parvula Calman,

Youne Prawn IN THE Ist PostLaARvAL Stace. (Pl. LX.)

There cannot, I think, be any doubt that the speeimen described

below and figured on the accompanying plate is referable to

the same species as the preceding larva, and accordingly is a

developing stage of Limnocaridina parvula Calman. The simi-

larity in the general aspect is very striking, and the difference in

28"

436 PROF. G. O. SARS ON

size is so slight that we have every reason to believe it to represent

an immediately succeeding stage. But, whereas in the preceding

stage the larval characters were very apparent, they are wholly

lost in the present specimen, which accordingly has entered the

postlarval period, representing here in all probability the very

first stage.

The length of the specimen examined measures 2°70 mm., and

thus very little exceeds that of the preceding larva. The adult

ovigerous female of the present species attams, according to

Dr. Calman, a length of 6°70 mm. ;

The general form of the body (figs. 1 & 2) is very slender, even

more so than in the preceding stage.

The carapace is of a quite similar shape, and the rostrum is

very short and spiniform; but on its upper edge three or four

small denticles have appeared (fig. 2).

The eyes considerably project laterally and exhibit a somewhat,

fusiform shape, their pedicles being conspicuously dilated in the

middle. The corneal part, on the other hand, appears somewhat

less expanded than in the preceding stage.

The metasome is more powerfully developed, and exhibits the

Caridean bend on the middle very distinctly (see fig. 2). The

epimeral plates of the five anterior segments are deeper, extending

slightly beyond the ventral faces of the segments.

On the appendages several changes have taken place, the most

conspicuous of which are the complete loss of the natatory

exopodites on the legs and the reduction of those attached to the

maxillipeds, the natatory function being now wholly transferred

to the metasome and more particularly to the pleopoda.

Structure of the Appendages.

The antennule (fig. 3) have the joints of the peduncle more

sharply defined and several additional plumous sets have appeared

inside the joints. The flagella are still rather short, but on both

a distinct subdivision has taken place, the inner one being com-

posed of three, the outer of four well-defined joints. On the

outer flagellum, moreover, a short uniarticulate appendicular

ramus has developed from the end of the second joint, carrying

on the tip a bundle of sensory filaments.

On the antenne (fig. 4) the scale is nearly unaltered, whereas

the flagellum has considerably increased in length, being now

fully twice as long as the scale. It is, moreover, divided into

several joints, the outer of which are very sharply marked off

from each other and provided with small hair-like bristles, whereas

the proximal ones are still imperfectly defined. The flagellum

also appears to be freely mobile in relation to the peduncular

part, as it is found in some cases extended forwards, in other

cases more or less obliquely backwards (figs. 1 & 2).

The mandibles (fig. 5) agree on the whole in their structure

with those in the preceding stage, though the molax prominence

PRAWNS FROM LAKE TANGANYIKA. b 43

appears more distinctly defined and exhibits a well-marked

triturating surface.

The maxilla (figs. 6 & 7) scarcely differ from those in the pre-

ceding stage, except in a somewhat richer supply of marginal

spines and sete.

The mamillipeds (figs. 8-10), on the other hand, have been con-

spicuously transformed, and now look very dissimilar. In the

Ist pair (fig. 8) only a slight rudiment of the endopodite is left,

and the exopodite also is considerably reduced and apparently

quite immobile, without any sete at the tip, and having the distal

part curved downwards. On the 2nd pair of maxillipeds (fig. 9)

the endopodite bas assumed the characteristic securiform shape

observed in the adult animal, the outer two joints forming to-

gether a broad plate folded upon the proximal part and armed at

the inner edge with a number of spiniform anteriorly curving

sete. On the 3rd pair of maxillipeds (fig. 10) finally the endo-

podite has preserved its pediform shape, but is considerably

prolonged, forming a nearly straight cylindric stem almost twice

as long «s the exopodite. The latter, as in the preceding pair, is

extended straight anteriorly, and appears to be very little mobile.

The chelipeds (fig. 11) have increased in length, and now

exhibit the full number of joints. On the hand, one of the

fingers (dactylus) has been well defined ab the base, and connected

with the hand by a very mobile articulation ; on the tip of each

finger a short claw-like spine has developed in addition to the

bristles, the number of which is considerably increased. The

sete on the other joints are, however, shorter and less numerous

than in the preceding stage. Outside the basal part a very small

and irregular protuberance is seen, indicating the place where the

exopodite has originally been attached.

The pereiopoda (figs. 12 & 13) are somewhat longer than the

chelipeds and quite normal in structure. The dactylar joint

nearly agrees both in shape and armature with that in the adult

animal, as described by Dr. Calman. Outside the basal joint of

the two anterior pairs (fig. 12) a minute protuberance quite

similar to that occurring in the chelipeds is seen, indicating the

original presence of exopodites on these legs. In the last pair

(fig. 13) no trace of such a protuberance is to be detected.

Above the bases of the legs, except the last pair, and covered

by the lateral parts of the carapace, the gills have developed,

forming on each side a regular series, as indicated in fig. 2.

The pleopoda (fig. 14) are now freely mobile and have the

terminal lamelle fringed with long ciliated sete. The number

of these sete is, however, as yet very limited, only eight being

counted on the outer lamella and six on the inner. All these

sete exhibit close to the base a very conspicuous joint.

The telson (fig. 15) does not differ much in shape from that

in the preceding stage; but the number of apical spines has

diminished, only five such spines being present. On the other

hand, three pairs of lateral denticles have appeared, the outer-

438 PROF. G. O. SARS ON

most one occupying the outer distal corner, the other two placed

at some distance from each other and more dorsally.

The wropoda only differ from those of the preceding stage in a

somewhat richer supply of marginal sete.

Occurrence.—The above-described specimen was found in the

same sample as the preceding larva, and was associated with three

other somewhat larger specimens. Moreover, young specimens of

the same species occurred in two other samples, the one taken at;

Mbete (S, end of lake), the other at Karema (KE. shore of lake).

In the latter sample numerous specimens were present.

GENERAL REMARKS.

Among the Macrura inhabiting fresh water, so far as I know,

the genus Caridina is the only one in which a true metamorphosis

has been found to occur. In the other forms the postembryonal

development is either direct, as in Astacus, or at least exhibits

only very slight traces of a metamorphosis. This difference in

the development, according to the medium, may even be found

in a single species, viz., in the case of Palemontes varians. ‘The

typical form of this species, found in salt or brackish water,

passes through a quite normal metamorphosis; whereas in the

variety inhabiting purely fresh water the larval development 1s

much abbreviated, the young leaving the egg with all the append-

ages present, except the uropoda. A similar suppression of the

metamorphosis is certainly also observed in some of the marine

forms; but these are, as a rule, only such as inhabit very deep

water. In shallow-water forms, on the other hand, a compli-

cated metamorphosis is always found to prevail. In most cases

the relative size of the ova, borne by the female beneath the

metasome, will be suggestive as to the decision whether the

metamorphosis in the observed form is complete or suppressed.

If the ova are very large and few in number, this is an infallible

sign that the metamorphosis is incomplete or quite absent. —

On the other hand, the presence of very small and numerous

ova points to the assumption that the young hatch in an

imperfect larval condition, and of course must pass through a

more or less complicated metamorphosis before reaching maturity.

In most of the prawns inhabiting Lake Tanganyika, as established

by the measurements given by Dr. Calman, the ova are of un-

usually small size, and from this fact alone it could be inferred

that a true larval development must be present. This has indeed

been fully confirmed by the investigations here published, and in

one case it has, moreover, been proved that the larvee, on escaping

from the ova, exhibit a still simpler and more primitive structure

than found in any other known form of this group.

A very peculiar feature, exhibited by the two Zoéx described in

this paper, remains to be mentioned, viz., the very rudimentary

aud as it were embryonic appearance of the oral parts (mandibles

and maxille). In all free marine larve belonging to the present

“PRAWNS FROM LAKE TANGANYIKA. 439

group, even immediately after hatching, I have found that these

parts, though more or less differing in structure from those in

the adult animal, yet at all events are functionally developed.

This cannot, however, by any means be said to be the case in the

two Z“oée here under consideration. On the contrary, it is

evident, both from the very imperfect structure of these parts

and their position in relation to the mouth, that they cannot

possibly act as true masticatory organs. The larve, of course, in

these early stages are quite unable to feed in the ordinary manner,

and must be assumed to subsist exclusively on the nutritive yolk

still left within the body. It seems to be very difficult to account

for this anomalous case; but a hypothesis may perhaps be set

forth. I suggest the possibility that in the lake there is scarcity

of that kind of food which would be suitable to the larve in this

early period, and perhaps also the absolute absence of Cladocera

in this lake may be explained in a similar manner. In the more

advanced stages, when the oral parts have been more fully de-

veloped, the larve in all probability feed upon the small Copepoda

which abound in the lake.

EXPLANATION OF THE PLATES,

Pr. LVI.

Recently hatched larva (Euzoéa) of ? Limnocaridina spinipes Clm.

1. Dorsal view of the larva.

2. Same larva, viewed from left side.

3. Anterior division of the body, viewed from the ventral face, and more

highly magnified. The three posterior limbs on left side are not fully

drawn, in order to show those occurring immediately in front of them

more distinctly. 41, antennule ; 47, antenne ; oc, ocellar lobe; H, hypo-

stome; DL, anterior lip; J£, mandibles; /!, posterior lip; m/, anterior

maxilla; m?, posterior maxilla; Jp!-Mp*, maxillipeds of 1st to 38rd

pairs.

4, Left antennula.

5. Left antenna.

6. Inner part of a mandible.

7. Anterior maxilla.

Fig.

. Posterior maxilla.

. Maxilliped of 1st pair.

. Maxilliped of 2nd pair.

11. Maxilliped of 3rd pair.

12. Left half of caudal plate.

Px. LVILI.

Early larval stage (Zoéa) of ? Caridella cunningtoni Clm.

. Dorsal view of the larva.

. Same larva, viewed from left side.

Anterior division of the body, viewed from the ventral face, and more

highly magnified. The three maxillipeds on left side are omitted in

order to show the maxillw# more distinctly.

. Antennula.

. Antenna.

. Maxilliped of 1st pair.

. Maxilliped of 2nd pair (exopodite not fully drawn).

. Maxilliped of 3rd pair.

9. Developing cheliped.

10. Developing pereiopod.

11. Caudal plate.

Cobo

DO OTHAS

440

Fig.

25. The Classification, Morphology, and Evolution of the

By Herpert L. Hawkins,

M.Sc., F.G.S. ; Lecturer in Geology, University College,

Reading *.

Within the limits of a single class, it would be difficult to find

greater contrasts than those which distinguish the various orders

OMAN WN

Echinoidea Holectypoida.

MR. HERBERT L. HAWKINS ON

Pu. LIX.

Last larval stage (Mysis stage) of Limnocaridina parvula Clm.

Dorsal view of the larva.

Same larva, viewed from left side.

. Antennula.

. Antenna.

. Anterior lip.

. Right mandible and masticatory part of left.

. Anterior maxilla.

. Posterior maxilla.

. Maxilliped of 1st pair.

10. Maxilliped of 2ud pair.

. Maxilliped of 31d pair.

2. Cheliped (exopodite not fully drawn).

13. Pereiopod of 1st pair (exopodite not fully drawn).

4, Pereiopod of last pair.

. Pleopod of 1st pair.

. Pleopod of 2nd pair.

. Extremity of last caudal segment, with telson and left uropod ;

Pu. LX.

First post-larval stage of Limnocaridina parvula Clu.

. Dorsal view of the specimen.

. Same specimen, viewed from left side.

. Antennula.

Antenna.

. Mandible.

. Anterior. maxilla.

. Posterior maxilla.

Maxilliped of 1st pair.

. Maxilliped of 2nd pair.

. Maxilliped of 3rd pair.

. Cheliped.

. Pereiopod of Ist parr.

. Pereiopod of last pair.

. Pleopod.

. Outer part of telson,

[Received November 16, 1911: Read March 5, 1912.]

(Text-figures 54-60.)

ContvTENTS.

I. Introduction ..... SPAR RA Sirs came sane 6 hie

II. History of Past Claseif cation. .c<...dteaod cates ene

III. The Revised Classification..

IV. Comparative Morphology ......

V. The Internal Evolution of the Group .. shendeaiaues eet

WileeihverExternalavelationshup samcieeeeeateet ere reee ee eeeece ere Eee

VII. Summary

VIII. List of Literature « consulted ..

I. Lytropvuction.

dorsal view.

* Communicated by Dr. Henry WoopwarpD, F.R.S., V.P.ZS.

SEA-URCHINS. 44}

of the Echinoidea. To a casual observer, unacquainted with the

anatomy and embryology of the forms, such genera as Cidaris and

Echinocardium would seem to represent two absolutely different

“kinds” of animals. Moreover, search as he might among the

Kchinoids of the present day, he could find no types that would

effectively bridge the gulf that separates the Regular from the

Irregular Sea-Urchins. All the Regularia have thick tests, built

on a radially symmetrical plan ; possess a strong jaw-apparatus ;

and are armed with stout, often very long, radioles. On the other

hand, most of the Irregularia have thin tests, bilaterally sym-

metrical; have no jaws; and are covered with small, almost

hair-like radioles. Even those forms, the ‘ Cake-Urchins,” which

show a radial symmetry and possess jaws, have other characters

which render them very different in appearance from the

Regular Echinoids,

And again, from an anatomical study of Jiving forms alone,

although a similarity of structure sufficient to warrant their

inclusion in the same order of the Echinoderma might be found,

no certain clues as to the relation of the two types to one another

could be discovered. Even ontogenetic evidence is lacking to a

considerable degree, for the peculiar larval life led by Echinoids

has tended to neutralise the effect of recapitulation.

At this point, where Zoology in its narrower sense fails,

Paleontology can supply the connecting links between such

different forms as a Cidarid and a Spatangid ; and of these links,

the majority are to be found in the group which forms the

subject of this research. The order Holectypoida arose soon after

the commencement of the Jurassic period, and became entirely

extinct before the close of the Mesozoic era. During its existence

it gave rise to forms which, by stages so gradual as to be hardly

distinguishable, laid the foundations of all the great groups of

Irregular Echinoids that are living to-day. It is the purpose of

this paper to indicate in outline the processes through which the

specialization of Clypeastroids, Spatangoids, and ‘ Cassidulids ”

was achieved.

When P. M. Duncan (44)* wrote his classic “ Revision of

the Genera and great Groups of the Echinoidea,” there existed

“much diversity of opinion regarding the nature of the peri-

gnathic girdles and jaws of some genera” (¢.c., p. 135) of the

Holectypoida, the most primitive group of the Irregular

Echinoids. As Duncan’s system of classification was based to a

very large extent on the varying characters of these structures, it

naturally resulted that in the case of the Holectypoida and their

allies the grouping of genera into families, or even into larger

groups, was somewhat tentative. It is only necessary to remark

that jaws are now known to have existed in two genera, in which

their presence was denied by Duncan, to show that a revision of

the classification of the group is required.

* Where a number in brackets follows the name of an author, the full title of the

paper referred to will be found under that number in the chronological list of

literature at the end of this paper.

449, MR. HERBERT L. HAWKINS ON

However, the difficulties of a systematic grouping of the

primitive gnathostomatous Irregularia are not removed, or even

lessened, by the additions that have been made to our knowledge

of their comparative anatomy. Rather are they increased, for

the establishment of affinities between genera leads to greater

complexity of classification than that of differences. The

Holectypoida are an annectant group, the history of whose

evolution is so intimately interwoven with that of the early

stages of most of the Irregular orders and suborders, that to

frame a purely natural classification would need an impracticable

plasticity of diagnoses. This systematic trouble is, however, more

than compensated by the phylogenetic evidence that it indicates.

In the course of the following work I have endeavoured, while

recasting the artificial classification of the systematist, to lay

emphasis on relationships rather than on contrasts, and to show

the position occupied by the Holectypoida at the foundation of

the varied structures of the Irregular Echinoids.

The present essay is the outcome of several years of study of

the group, and contains a summary and amplification of a series

of papers (see list at end) that have been published in the

‘Geological Magazine.’ I have thought it unnecessary to repeat

here many of the details described in those papers, so that, except

where corrections or additions have been possible, the results

arrived at in them are taken for granted. There are, however,

descriptions of a number of features that find a place here which

were not dealt with in the shorter papers given here.

After a brief sketch of the history of the classification of the

group, the revised scheme is put forward. This is followed by a

morphological comparison of the genera within, and of some

genera without, the boundaries of the order; and lastly, the

directions of evolution thus indicated are discussed.

II. History or Pasv CLassivicaTion.

This part of the paper does not pretend to be a complete

account of all the past work that has been done on the group,

but it is a summary of the chief systems of classification that have

been proposed up to the present time.

The history of the group may be said to date from 1734, when

Klein, in his ‘ Naturalis dispositio Echinodermatum,’ distin-

guished “ Sectio I, /ibula,” from the rest of the “ Hchini catocysti

circulares.” The section was diagnosed as follows :—“ Echinos

fibulares dicimus Catocystos circa Basis circularis peripheriam

Anum, Os in medio aperientes.” He included two genera only

in the section, Conuluws and Discoides, both of which are recog-

nized to-day, and have been associated with one another in most

of the systems of classification. Klein apparently did not

know of any of the other genera included among the Holectypoida

in the present paper.

SEA-URCHINS. 443

The classification adopted in 1840 by L. Agassiz, in his

‘Catalogus systematicus,’ was in many ways of less value than

that of Klein; for he grouped together, under the heading of

‘“Clypeastroidee,” all the non-Spatangid Irregular Echinoids.

However, Desor, in the sequel to that Catalogue (11) showed that

a more detailed system of subdivision was practicable, and founded

the first definite scheme of the classification of the group. The

“tribu” of the ‘“ Galérites” was regarded as a division of the

“ Clypéastroides.” It contained the following g ‘ara-

tomus, Discoidea, Echinoneus, Galerites, Globator, Holectypus (as

a subgenus of Discoidea), Hyboclypus, Nucleopygus, Pygaster, and

Pyrina. This list of genera contains a very natural grouping of

all those Irregular Echinoids which have simple, apetaloid

ambulacra. That, however, is almost the only trait that could

associate them, and in Car ee at least, the simplicity of the

ambulacra is not absolute.

In the ‘ Catalogue raisonnée’ (1847) Agassiz and Desor retained

this grouping of the genera in its entirety, but rearranged

the position of the “tribu” as a whole. It appears as the

family Echinoneidz, a section of the Cassidulide. This change

was at once an advance and a retrogression. It applied to the

family a name under which some of the genera have remained

since that time, but by associating the whole series with the

Cassidulids, it tended to obscure the importance of the gnatho-

stomatous char acter of many of the genera.

In the Synopsis, Desor (21) retained the division under the

name of Galeridées, and added large numbers of genera to the

list. He recognized, however, the fundamental importance of

the presence of jaws in determining the systematic position of a

genus, and so separated the Galeridées into two groups. Of these,

the fir st, or “Galeridées proprement dits,” contained Aeey

genera with j jaws (or rather, supposed to possess them), while the

second, the “‘ Echinonées,” included Hehinoneus only. The first

group contained all the genera of the ‘“Galérites ” of 1842, except

Caratomus and Echinoneus, and there were added the then newly

described genera dAnorthopygus, Asterostoma, Desorella, Galero-

pygus, Pachyclypus, and Pileus. It is curious that Desor should

have taken it for granted that all these fossil genera were gnatho-

stomatous, for he cannot have had any positive evidence to work

upon in the majority of cases.

In 1857 Pictet, in the second edition of his ‘ Paléontologie ’ (22),

reverted to the method of er ouping originated by Desor in 1842

changing the word “Galérites” to the subordinal term “ Galeri-

tiens,’ and adding the then recently described Desoria.

One year before the appearance of Desor’s Synopsis, Wright

(20) had grasped the essential differences which divided the

“ Galerites”” into two sections. He founded the family of the

Kchinoconidee, which contained Discoidea, Echinoconus (the

Galerites of most previous authors), L/olectypus, Hyboclypus, and

444 MR. WERBERT L. HAWKINS ON

Pygaster. He contrasted this family with the Echinoneide, in

which he placed Hehinoneus, Pyrina, and several other genera.

This was the first time that the presence of jaws was treated as

an essential feature in the classification of the group.

Wright's system of classification was adopted for many years

by almost all the Echinologists who dealt with the group, although

slight changes in the generic personnel of the Echinoconide were

introduced. Cotteau (28) removed Hyboclypus, with good reason,

from the family, and added the genera (unknown to Wright in

1856) Anorthopygus and Pieus.

The compact group thus determined, bound together by the

characters of a short and accurate diagnosis, became generally

accepted. Lovén (31) worked on this classification as a basis, and

Wright (82) agreed with Cotteau’s modifications. Perhaps no

gurer indication of the natural character of the grouping could

anywhere be found than in the fact that Pomel (87) was unable

to find any cause for more than internal changes in the family.

With a subordinal rank within the group of Gnathostomes

Clipéiformes, Pomel placed the section Galérides. This section

he subdivided into two chief families, the Echinoconidés and the

Piléidés. The latter family was further separated mto two sub-

families, the Discoidiens and the Pygasteriens. Although many

new ‘‘generic” terms were introduced, no forms were included

among the Galérides that were not previously classed with the

Echinoconide. The separation of the Hchinoconus-group from

the other genera was natural. In the Piléidés, the first group

was simply the original genus Discoidea of Agassiz in a dis-

membered state, while tlie second group included the same author’s

early conception of the genus Pygaster, The classification of

Pomel was therefore, in this group, quite orthodox, a condition of

affairs sufficiently surprising in view of the great changes he

roposed in the arrangement of many of the other groups.

Tn 1889, Duncan (44), who expressly dissociated himself from

Pomel’s views of the relative importance in classification of

various structures, published the invaluable Revision of the

renera, etc. of Echinoidea. In this work, he realised the great

importance of the Holectypoida as an annectant group. So

thoroughly intermediate in its characters was the group that he

definitely stated that his classification was artificial, and as such

tentative. The Holectypoida received the rank of an Order,

equivalent in importance to the Clypeastroida or the much larger

groups of the Diademoida and Spatangoida.

It was chiefly on the peristomial and jaw-structures that

Duncan classified the group, and on that account it was particularly

unfortunate that he should have had such a fixed belief in the

absence of jaws in some genera in which they have since been

discovered. Curiously enough, although, in the same year as the

publication of the Revision, he definitely stated his disbelief m

the existence of jaws in Discoidea (45), he allowed that genus to

SEA-URCHINS. 445

find a place among the Holectypoida, while Lehinoconus was

banished to the Echinoneidze among the Spatangoida. <A similar

fate befell Anorthopygus, while Conoclypeus, a genus till then

usually classed with either Hchinanthus or Clypeaster, was brought

into the Holectypoid group. (Conoclypeus was regarded as a

Galerites” by Grateloup, 5.)

Duncan divided the Holectypoida (whose brief diagnosis was

** Exocyclic, oligoporous Ectobranchiata”) into two unnamed

sections. The subdivision was made on the details of the

perignathiec girdle, and Discoidea and Conoclypeus, on account of

the supposed rudimentary state of their processes, were thus

separated from Holectypus and Pygaster. Galeropygus and

Pachyclypeus were regarded as being Holectypoids, but

as not sufficiently known to be definitely associated with, or

separated from, any of the other genera.

Duncan’s classification was followed absolutely by Sladen in

the “ Zittel-Eastman ” Text Book of Paleontology (58).

The only remaining classification of the group in which any

important changes are made is that devised by Gregory in 1896

(50), and published in Lankester’s ‘ Treatise on Zoology.’ Here

the “‘ Holectypina” (a group corresponding in part with the

Holectypoida) are regarded as a suborder of the Gnathostomata.

The chief contrast between Gregory's group and that of Duncan,

is that the former author so modifies the diagnosis of the

Holectypina as to admit Galerites (Hchinoconus), although it is

believed to be edentulous. The Holectypina are divided into

four families, the Pygasteridee, the Discoidide, the Galeritide, and

the Conoclypeide. Discounting the genera described since 1889,

the Pygasteridz correspond to section I. of the Holectypoida,

with the queried inclusion of Galeropygus and Pachyclypeus. The

Discoidiidee and Conoclypeidee together contain the members of

Dunean’s second section, while the Galeritide are the first sub-

family of the Echinoneide of the Revision.

The classification proposed by Gregory seems to accord better

with our knowledge than any of those previously suggested. In

view of the complexity of the relations of the Holectypoida, owing

to its primitive and annectant character, I prefer to regard it as

a group so much apart from the other Irregularia as to merit

its retention as an Order, as Duncan originally considered it.

After a thorough study of the comparative morphology of most

of the genera included in the order, I have attempted to revise

its internal classification in such a manner as to indicate the

affinities, both internal and external, which the study has made

manifest.

III. Tae Revisen* CLAssiFicaTIon.

The characters of an annectant group are inevitably plastic

and unstable. For this reason a natural classification of such a

446 MR. HERBERT L. HAWKINS ON

group becomes an almost impossible task. The features which

seem of essential importance in one genus may be quite absent or

profoundly modified in another. But in the case of the

Holectypoida, the length of time during which this plasticity of

structure was retained gives possibility for a classification that is

fairly im accord with the evolution of the group, and at the same

time is free from a confusing multiplicity of detail. However,

as will be seen on a comparison of the scheme submitted below

with the genealogical table given in a later part of the paper, the

two groupings do not agree in every particular, All of the

genera have a great phylogenetic significance, and would, if the

classification were to do justice to that importance, require each

a separate family. Probably, as our knowledge of the relations

of the group extends, the present genera will become the bases

of distinct families, and will be themselves divided into many

smaller sections. The great variety of species which are at

present grouped under the generic names Holectypus and Discoidea

seems to lend support to this belief. For convenience of reference,

it has seemed preferable to retain, as far as possible, a more

generalized system of grouping than, by comparison with other

orders of Echinoidea, the individual peculiarities of the genera

in reality demand,

The Holectypoida are an order intermediate in characters

between the Echinoidea Regularia and all the various orders of

the Irregularia. The features in their structure which are

naturally the most uniform in character are therefore those

relics of ‘ Regularity ” that they retain. They show a persistently

retarded progress in their evolution, and from them, at various

points, relatively accelerated offshoots break free, Throughout

the entire group two features remain constant in their presence,

although they undergo a gradual reduction in the perfection of

their development. These features are:—the existence of a

masticatory apparatus ; and the presence of external peristomial

branchiz. Both of these struetures are of essential importance

physiologically, and both fortunately leave traces of their existence

on the skeletal structures. It is a postulate (which probably

expresses a fact) that, whenever the peristome is centrally situated,

and at the same time circular in outline, jaws are present,

Certainly the existence of a well-developed perignathie girdle

indicates their presence, so that, when either of these characters

can be observed, the existence of a mastieatory apparatus can be

inferred, even if it has not been discovered, The branchial

incisions on the peristome margin are naturally easy to recognize

when that part of the test is preserved.

After the scheme of classification has been summarized, revised

diagnoses will be given forgthe order, families, and genera.

Subsequently, the reasons for the exclusion of some genera which

have hitherto been classed with the Holectypoida will be

discussed, and their positions in the other orders indicated.

SEA-URCHINS. 447

Order HOLECTYPOIDA.

Family I. PYGASTERIDAS.

Subfamily 1. PYGASTERINA,

Genus Pyaaster Agassiz.

Subgenus 1. Pygaster sens, str.

‘s 2, Megapygus, nom. nov.

Ws 3. Macropygus Cotteau,

Subfamily 2. PILEIN &,

Genus 1. Piteus Desor.

» 2. ANnortHopyaus Cotteau,

Family IT. DISCOIDITD As.

Subfamily 1. HOLECTYPINS.

Genus 1. Hotectypus Desor.

Subgenus 1. Holectypus Desor.

5 2. Canholectypus Pomel,

*e 3, Lanierta Duncan.

Genus 2, Copropiscus Cotteau & Gauthier.

Subfamily 2. DiscorpIIN®.

Genus Discomna Agassiz.

Family IIT, CONULIDA,

Genus Conuuus Leske.

Incertze sedis :—DiscHoLectyPus Pomel.

Order HOLECTYPOIDA Duncan (emend.).

Euechinoidea Irregularia with external peristomial branchie

(Ectobranchiata) and a central mouth armed with jaws and

surrounded by a perignathic girdle (Gnathostomata).

Ambitus circular, subpentagonal, posteriorly truncated, or

slightly elongated, Adapical surface acutely or bluntly conical,

apex practically central. Peristome central, usually circular, some-

times decagonal or even obliquely elliptical in outline. Perigna-

thic girdle discontinuous, composed of both processes and ridges.

Jaws more like those of the Regularia than of the Clypeastroida.

Periproct very variable in size and _ position, always posterior.

448 MR. HERBERT L. HAWKINS ON

Ambulacra straight, narrow, similar, and simple, with usually

larger podial pores on the adapical than on the adoral surtace.

Towards the peristome the pore-pairs may become grouped

into ares of three. Ambulacral plates small, simple or compound,

the latter always derived from three original primary plates.

Interambulacra broad; the plates usually concentric in arrange-

ment, rarely bent along their median line. Tubercles always in

recognizably vertical series. Apical system variable ; madreporite

usually large. Radioles short, and longitudinally striated.

Liassie to Uppermost Cretaceous.

Family J]. PYGASTERIDA.

Holectypoida with a circular or posteriorly truncated ambital

outline. Bluntly conical adapically, concave adorally. Peristome

large, with strong perignathic processes and feeble ridges.

Jaw-structure insufficiently known for diagnosis. Branchial

incisions deep. Periproct large, always on the adapical surface,

often oblique. Ambulacra slightly tumid, composed of primaries

to a point about midway between the ambitus and the peristome.

Outer members of pore-pairs often transversely elongated (to a

slight degree only) on the adapical surface. Interambulacra broad,

paucituberculate (for an. Irregular HEchinoid), the admedian

tubercles being in concentric series, the adradial oblique.

Granulation irregular, faintly scrobicular. Apical system with

four perforated genital plates; the fifth being usually represented

by several small plates. Ocular plates small and similar. No

internal buttresses to the test. Radioles short, longitudinally

striated.

Liassic to Lower Cretaceous.

“

Subfamily 1. PyGASTERIN4.

Pygasteridee with the periproct in contact with the apical

system. Ambulacral pore-pairs uniserial except near the peri-

stome.

Genus 1. PyGaster Agassiz.

With the characters of the subfamily.

Genotype, P. semisuleatus Phillips.

Subgenus 1. PyGAsrer sens. str. (= Plesiechinus Pomel).

Pygaster with the periproct inside, as well as outside, the

apical system, and with its greatest width in the adapical part.

No posterior genital plate, the remaining plates of the system

being arranged transversely. Tubercles with shallow scrobicules,

regular in their introduction.

Subgenotype. P. semisulcatus Phillips.

Liassic to Middle Oolitic.

SEA-URCHINS. 44.9

Subgenus 2. Mrcaryaus nov. (?ygaster restr. Pomel).

Pygaster with periproct pyriform, and constricted towards the

apex. Fifth genital plate present, or replaced by several small

plates. Tubercles with shallow serobicules and irregular in

their introduction.

Subgenotype. JZ. umbrella (auctt.).

Middle and Upper Oolitic.

Subgenus 3. Macroryeus Cotteau.

Pygaster with the periproct and apical system as in Megapyqus.

Posterior margin str ongly truncated. Tubercles with large, deep

scrobicules; regular in their introduction.

Subgenoty pe. MW. truncatus Agassiz.

Lower Oolitic to Lower Cretaceous.

Subfamily 2. PILEIN »#.

Pygasteride with the periproct midway between the apex and

the ambitus. Madreporite very large, occupying the centre of

the apical system.

Genus 1. PruEus Desor.

Pileine with the ambulacral pore-pairs biserial adapically.

Periproct very slightly oblique. Fifth genital plate small,

imperforate.

Genotype. P. pileus Agassiz.

Corallian.

Genus 2. ANorTHOPYGUS Cotteau.

Pileine with the ambulacral pore-pairs uniserial throughout.

Periproct markedly oblique. Apical system ethmolysian.

Tubercles of interambulacra in sloping lines on both sides of the

central series.

Genotype. A. orbicularis Grateloup.

Lower and Middle Cretaceous.

Family IT. DISCOIDITDAs.

Holectypoida with a circular or posteriorly elongated ambital

outline. Depressed or elevated, conical, adapically; flat or

slightly concave adorally. Peristome of moderate size, with

strong perignathie processes and well-developed ridges. Bran-

chial incisions well marked. Periproct of variable size, marginal

or adoral in position. Ambulacral plates primaries to the

ambitus, compound there and adorally. Pores usually equal and

circular. Interambulacra broad, with many vertical rows of

tubercles, often supplemented by hypertrophied granules.

Granulation usually transversely linear. Tubercles much larger

on the adoral than on the adapical surface ; arrangement as in

Proc. Zoou. Soc.—1912, No, XXIX. 29

450 MR. HERBERT L. HAWKINS ON

Pygasterine. Apical system composed of five genital plates, the

posterior one perforated or not. Madreporite central. Ocular

plates often very small. Radioles short, acuminate, longitudinally

striated.

Lower Oolitic to Upper Cretaceous.

Subfamily 1. HOLECTYPINE.

Discoidiidee with the periproct marginal or adoral in position.

Fifth genital plate smaller than the other four. Tuberculation

usually sparse. Perignathic ridges low. No internal buttresses

to the test.

Genus 1. Honecrypus Desor.

Holectypinz with the characters of the subfamily.

Genotype. //. depressus Leske.

Subgenus 1. HoLEectyPus sens. str.

FTolectypus with the posterior margin often elongated; witha

large periproct, marginal or adoral in position. Fifth genital

plate small and imperforate. Madreporite central and

prominent.

Subgenotype. H. depressus Leske.

Lower and Upper Oolitic.

Subgenus 2. Caznnotectypus Pomel.

Holectypus with a circular ambitus ; periproct of comparatively

small size, usually adoral in position. Fifth genital plate almost

as large as the others, and perforated.

Subgenotype. C. macropygus Desor.

Lower to Upper Cretaceous.

Genus 2. Copropiscus Cotteau & Gauthier.

Holectypine similar to Canholectypus, but with deep excava-

tions along the plate sutures on the adapical surface.

Genotype. C. nemice Cott. & Gauth.

Genus 3. LantertaA Duncan.

Holectypine similar to Cenholectypus, but globular in shape.

Genotype. LZ. laniert VOrbigny.

Upper Cretaceous.

Subfamily 2. DiscoIDIIN#®.

Discoidiidee with a circular ambitus; with the periproct on the

adoral surface. All five genital plates of approximately equal

size, the posterior plate perforated or not. Madreporite often

scattered over all five genitals. Perignathic midges high.

Tnternal buttresses present on the adoral surface.

SEA-URCHINS, 45]

Genus 1. DiscorpgA Agassiz.

With the characters of the subfamily.

Genotype. D. subucula Leske.

Lower and Upper Cretaceous.

Family III. CONULIDA.

Holectypoida with a posteriorly elongated or circular ambital

outline. Usually tall and conical adapically ; flat or subeconvex

adorally. Peristome small, slightly elliptical and sometimes

oblique, with the perignathic ridges of equal height with the

processes, the whole girdle leaning against an i nternally thickened

portion of the adoral surface. Branchial incisions very slight.

Periproct small, marginal. Ambulacra of compound plates

almost throughout, pore-pairs definitely triserial near the

peristome. Interambulacra multituberculate, with both ad-

median and adradial series sloping adorally, and often in linear

sets of three instead of two on each plate. Granulation irregular,

granules sunken on the adapical surface. Apical system with

four genital plates only. The two posterior oculars meet along

the middle line, and are larger than the other three. No

internal buttresses, but a marked thickening of the interradial

parts of the adoral surface. Radioles similar to those of Dis-

coidiide ; but in addition short ¢ pedicellaria-stumps arising from

the invaginated granules,

Lower to Upper Cretaceous.

Genus 1. Conunus Leske.

With the characters of the family.

Genotype. C. albogalerus Leske.

Lower and Upper Cretaceous.

INCERTA SEDIS.

DiscHo.tectypus Pomel.

Holectypoida with the characters of Canholectypus except in

the ambulacra, which are composed of compound plates through-

out, as in the Conulide.

Genotype. D. meslei Gauthier.

Lower Cretaceous.

Discussion of the Systematic Position of Genera JSormerly included

among the Holectypoida, but now removed Srom the Group.

PLeEstecHINuS Pomel.

This genus (or subgenus) included Pygaster semisulcatus

Phill. and other species related to it. As P. semisulcatus is the

29*

452, MR. HERBERT L. HAWKINS ON

type of Pygaster Agass. (6), the name Plesiechinus cannot be

retained. I have, therefore, renamed Pomel’s subgenus Pygaster

sens. str. The ‘‘ Pygaster” of Pomel thus requires a new sub-

generic name. As this group, which is typified by P. wmbrella,

is similar in the character of the periproct to Macropygqus, and

seems to mark a parallel though distinct line of evolution to that

subgenus, I have named it Megapygus.

PyGAStTRIDES Lovén. :

As was realised from the first, this small recent form possesses

all the essential features of a Pygaster, and the species

(P. relictus) was originally given that generic name by Lovén.

There is no direct evidence, so far as I have been able to gather

it, whereby this genus should be omitted from the Pygasteride.

But there is a serious doubt as to its being a “ genus,” in the

strict sense of the word. It is founded on one broken and

minute specimen. On first principles, the great lapse of time,

unbridged by any similar forms, which separates P. relictus from

even the latest members of the Holectypoida, renders it im-

probable that 1¢ can be a revived example of the group.

Moreover, there has recently come to light some indirect

evidence which seems thoroughly to undermine the foundations

of the “genus.” The presence of a generally Pygaster-like

facies, and of a complete lantern, in a small recent Hchinoid

known to belong to the genus Hchinonews (Agassiz, 58), in

addition to the extraordinary interest of its mere existence,

makes it practically certain, to my mind, that the specimen

described by Lovén was a similarly atavistic post-larval form.

For this reason, I have thought it best to ignore Pygastrides in

the diagnosis of the Holectypoida, and to omit it altogether from

the classification.

Gateropycus Cotteau (Desor).

Several well-marked features render it impossible, as well as

unnatural, to associate this genus with the Pygasteride. Two

striking differences are the strong curvature of the two posterior

ambulacra at their adapical extremities (and the extreme narrow-

ness of the areas generally), and the irregularly multituberculate

character of the interambulacra. Moreover, the peristome is

small, unnotched for branchiz, and placed anteriorly from the

centre. It must be regarded as the earliest known genus of the

Nucleolitide of Gregory (50), and its affinities will be more fully

discussed in Section VI. of the present paper.

Ecuinires Duncan (Protocyamus Gregory).

Bather has shown (55) that this “ genus ” (renamed by Gregory

in 1900), being founded on Discoidea subucula Leske, must be

considered a simple synonym of Discoidea. D. subucula is the

type of the genus.

SEA-URCHINS. 453

CoNnULopPsIs, gen, nov,.*

The following is a brief diagnosis :—Ambitus circular, or very

slightly elongated posteriorly ; adapical surface conical, not very

elevated; adoral surface flat or slightly concave. Peristome

small, slightly excentric, surrounded by interradial “ bourrelets.”

Periproct marginal or inframarginal, transversely expanded.

Ambulacra of simple primaries, large adorally ; pores almost sub-

petaloid adapically. Interambulacra multituberculate, tubercles

not in vertical series, deeply scrobiculate, imperforate. Granu-

lation coarse, closely packed.

Genotype. C. roemeri d’Orbigny, sub Galerites.

Upper Cretaceous.

This genus corresponds with the “ Echinoconus” of Desor (21),

but is certainly not a member of the Holectypoida. “ Galerites

roemeri” is not a “ Galerites” at all, but, like the ‘ Zehinoconus

abbreviatus” of our own uppermost Cretaceous (its probable con-

gener), seems to be a near ally of Caratomus (see Schlueter, 54),

but to be sufficiently distinct to demand a new generic name. A

further discussion of the affinities of Conwlopsis will be found on

p- 491.

ADELOPNEUSTES Gauthier.

This genus is founded on one specimen, A. lamberti, from the

Upper Chalk of Tunis. It is compared by Gauthier (46) with

Galerites roemert d’Orbigny, and the comparison, judging by the

figures, seems justified. It must therefore follow Conulopsis to

the Caratomus-group.

ConociyPeus Agassiz.

There are two noteworthy features which separate this genus

from the Holectypoida. ‘There are no branchial incisions on the

margin of the peristome, and the ambulacra are definitely sub-

petaloid, Moreover, the interambulacral tubercles are closely

packed, without any recognizable vertical arrangement. Cono-

clypeus, and with it probably Oviclypeus Dames, represents the

most primitive family of the Clypeastroida. The family name of

Conoclypeide used by Gregory (50) may be retained for these

two genera.

AMBLYPYGUS Agassiz.

This genus has not, so far as I am aware, been previously

associated with the Holectypoida, but its affinities with that order

are at least as well marked as those of the genus last mentioned.

It is certainly edentulous, and the tubercles are quite irregularly

packed together on the interambulacra. The ambulacra, how-

ever, in spite of an appreciably subpetaloid development on the

* A more detailed account of the characters of this genus will be given in a paper

that I hope to publish shortly.

454 MR. HERBERT L. HAWKINS ON

adapical surface, have exactly the plating-characters of Conulus.

Amblypygus seems to represent the simplest form of the Tertiary-

Recent section of the “ Cassidulide ” (see Hawkins, 66), which is

characterized by Hchinolampas and its allies. The genus will

receive fuller consideration in Section VI. of the present paper.

IV. ComparativE MorpHouoey.

Throughout this part of the paper references are made chiefly

to the structures of the four common British representives of the

group (Pygaster, Holectypus, Discoidea, and Conulus). Only when

peculiar characters or important contrasts occur in the less

abundant or foreign genera is a description of them inserted.

I have followed this principle advisedly, because, as this work is

largely one of generalization, it seemed preferable to use forms

where plenty of material was available, rather than to run the

risk of laying too strong an emphasis on a feature which,

occurring in an uncommon type, might be an individual

peculiarity,

A, The proportions of the Test.

1. The circumference.

All the forms which are included in the Holectypoida have

typically a radially symmetrical outline around the ambitus. In

some of the earlier forms, notably in varieties of Pygaster semi-

sulcatus, there is a tendency for the outline to be quite sharply

pentagonal by reason of the prominence of the ambulacra, but

outside the borders of Pyyaster sens. lat. this feature rarely

appears. It is perhaps worth noting, in this respect, that among

the markedly pentagonal forms in my collection of the species

just mentioned, there are many of quite small size. Although

thus apparently a constant feature throughout life in some

individuals, the angularity seems not to represent any phylo-

genetic stage, but to be merely an irregular, though frequent,

variation.

Among the Pygasters, when any departure from radial sym-

metry is encountered, it is found to result from a shortening of

the antero-posterior diameter in proportion to the width of the test.

This effect is appreciable in P. (Megapygus) umbrella, but reaches

its extreme in P. (Macropygus) truncatus, It is in all probability

due to an interference with the growth of the plates of the posterior

interambulacrum by the great size of the periproct. The trun-

cation of outline is rarely found outside the genus, but in Desorella

and G'aleropygus it reappears, often to an increased degree.

The slight groove which, in P. semisulcatus, passes from the

periproct to the posterior margin, is probably due to the same

cause as the shortening of the interradium. It is very interesting

and suggestive to find a trace of the sulcus in this genus, in view

ot the fact that its presence is a notable feature in Galeropygus,

SEA- URCHINS. 455

and most of the non-Holectypoid Jurassic Irregular genera, A

further reference to this feature will be found in the section on

the interambulacra (p. 465).

In Holectypus there is a tendency opposite to that of Pygaster,

but one probably caused by the same agent. In such a form as

H. depressus, where the periproct is of very large size and is

situated on the adoral surface, there is frequently a backward

projection of the posterior interambulacrum to accommodate it.

Even with this projection, there is often but.a thin rim of test

between the periproct and the peristome—a fact which shows the

necessity for some such arrangement. In many of the Jurassic

species which have a marginal periproct, a similar tendency is seen

(e. g., Holectypus oblongus Wright). Although the lengthening of

Text-fig. 54.

Diagrams of the adoral surface in some Holectypoida and their allies showing

the shape of the ambitus and characters of the peristome.

A. Pygaster semisulcatus. B. Galeropygus agariciformis. C. Pygaster (Macro-

pygus) lagaroides. VD. Holectypus depressus. EE. Discoidea cylindrica.

F. A Clypeastroid. G. Conulus albogalerus. H. A cretaceous Echinonoid.

the antero-posterior axis results in a bilateral symmetry compar-

able with that of many of the Echinoids which are more advanced in

“Trregularity,” it was not a feature retained by the Holectypoida

after Jurassic times. All the species of the subgenus Cenholec-

typus, and all of Discoidea, have an approximately circular outline.

The slightly indented character of the interradii on the ambitus

of Discoidea gives an alternating concavity and convexity to the

margin which may be compared with that of a Clypeaster. In

Conulus the tendency to elongation reappears, particularly in the

large, high-zonal forms of C. albogalerus, wheve the marginal

456 MR. HERBERT L. HAWKINS ON

periproct is often situated on a considerable projection of the

posterior interradius. The renewed appearance of bilateral sym-

metry under these conditions seems to confirm the belief that, so

far as the Holectypoida are concerned, deviations from radial

symmetry are connected with the migrations of the periproct, and

depend upon them, having no real significance of their own.

2. The Adapical Surface.

With the exception of one rare form (P. semisulcatus var.

conoideus), all the species of Pygaster are depressed. The conical

shape is not quite regular owing to the interference of the peri-

proct, and the apex is usually a little to the rear of the centre.

In Holectypus the cone is rarely less elevated than in an average

Pygaster, and is usually considerably higher. The extreme flat-

ness of Anorthopygus orbicularis is quite exceptional for the

group. The cylindrical form of Discoidea cylindrica is all the

more curious because of the normally conical shape of the other

and smaller species of the genus. The outline of Conulus sub-

rotundus sometimes resembles it, but the vertical character of the

sides is never so complete. The Upper Chalk Conwli tend, as a

rule, to assume an acutely conical shape, so that the area of the

adapical is sometimes twice as great as that of the adoral surface.

The apex of the test is practically central in all the genera except

Pygaster.

Among outside genera, almest the only forms to show the

sharply conical shape of the Holectypoida are Conoclypeus and its

allies. Most of the Clypeastroids are exceedingly flat in shape,

while the bilaterally symmetrical groups naturally cannot be

compared with the Holectypoida in this feature.

3. The Adoral Surface.

There is a very constant progressive change in the form of this

region of the test within the group, From Pygaster, with a base

so concave that specimens placed with the mouth downwards

rest on the ambitus alone, to Conalus, where the base is to a con-

siderable extent convex, every gradation may be traced. As this

feature is directly associated with some of the peristomial char-

acters, it will be better to postpone its discussion to that section

of the paper,

B. The Peristome and Associated Structures.

1, The Peristome.

The central position of the peristome is constant throughout

the group. In its size there is a progressive reduction traceable

through the Jurassic to the Cretaceous forms. In Pygaster the

peristome is of about the same size, relatively to the test diameter,

as in an average Diademoid (about one fifth), In Holectypus a

SEA-URCHINS. 457

reduction is initiated, which is maintained and even accelerated

in Discoidea, until in D. cylindrica, and also in Conulus, the peri-

stome has only about one ninth the diameter of the test.

In dealing with this character it is important to realise that the

size of the peristome is not of necessity directly connected with

the presence or absence of jaws. It is true that in Galeropygus

and Pyrina, where jaws were absent (at least, in adult forms), the

peristome is quite small ; but in the majority of the Clypeastroids

the peristome is smaller in proportion than in these genera, and

yet powerful jaws are present.

In the matter of the proportional representation of the ambu-

lacral and interambulacral areas on the peristome margin, a slight

but important change occurs in the course of the development of

the group. In Pygaster sens. str., the proportions are 1 to 14 in

favour of the interambulacra. In Conulus there is no appreciable

difference in the share taken by the two areas. This change is

partly due to the increased phyllodal tendency of the adoral parts

of the ambulacra in Conulus, but still more to an actual narrowing

of the interambulacra. In view of the extreme reduction which

is found in the latter areas of Clypeaster and its allies, the pro-

gressive change, though slight, is significant.

Apart from the characters of the branchial slits, which will be

considered in the next paragraph, the shape of the peristome

undergoes no important changes until Conulus is reached. In

that genus the circularity of its outline becomes slightly modified

into an elliptical shape, with a tendency for the long axis of the

ellipse to be oblique in its relation to the antero-posterior diameter.

The departure from the circular form is very insignificant in itself,

but when viewed in the light of the persistently elliptical, and

usually oblique, peristome of the Echinoneide, it becomes invested

with greater meaning.

2. The Branchial Slits.

All of the genera that I include among the Holectypoida

possessed external branchize. Within the boundaries of the group,

however, it is possible to trace the gradual reduction of these

structures (as indicated by the shallowing of the peristomial slits

through which they passed) until they become hardly appreciable

in size. In no ease are there signs of the smooth calcareous

developments of the interambulacra, for the support of the gills in

a recumbent position, which are characteristic of many of the

Regularia Ectobranchiata. The slight modifications of the plate

surface that exist are discussed in the section on the interambu-

lacral areas.

In Pygaster sens. str., the branchial slits are extremely well

marked (in Megapygus they are slightly shallower), and the depth

of their incision renders the outline of the peristome festooned

and decagonal. This stellate shape of the peristome is retained,

though toa reduced degree, in Holectypus. In Discoidéa the slits

458 MR. HERBERT L. HAWKINS ON

are so small that, in spite of the relative minuteness of the peri-

stome, the margin is actually less notched than in Pygaster. In

Conulus the slits are only just distinguishable on the thickened

rim of the peristome. In Pyrina, and in the Echinoneide gener-

ally, they seem to be altogether absent.

The concavity of the adoral surface is found to correspond

fairly closely with the development of the branchiew. In Pygaster,

Anorthopygus, and Holectypus, the adoral surface is markedly

concave, and the peristome is situated in an additional hollow in

the centre. In Discoidea the surface is almost flat, and yet the

peristome is deeply sunken. In Conulus, on the other hand, the

mouth is practically flush with the test-surface. In the Regular

Ketobranchiata the length and stoutness of the radioles are suf-

ficient to keep the test permanently raised above the rock surface.

The branchize are by this means kept free from the danger of

becoming bruised or fouled by contact with the ground. In

the Holectypoida the radioles were certainly not so strong as, and

probably of far less length than, those of an average Regular

Kehinoid. As a consequence, the adoral surface would be usually

very near to, if not in actual contact with, the rock surface. Such

a condition would have a disastrous effect on such delicate organs

as the external branchiz. It seems possible, therefore, that the

concavity of the adoral surface of the test is a device for sheltering

these structures. In the case of Discoidea, where, for purposes of

internal consolidation, the lower part of the test is flattened, the

region of the peristome is sunk to a proportionately great degree

to afford this shelter for the branchie. In Conulus, where the

branchiz were practically negligible in size, and probably in

function also, no such precautions were necessary. For gnatho-

stomatous forms, like the Holectypoida, which were presumably

not wholly, or even chiefly, microphagous, this depression of the

peristome would appear to be disadvantageous for the capture of

food; and only the safety of the equally essential process of

vespiration could warrant such a development. However, 1t must

be remembered in this connection that the Clypeastroids, in a

considerable number of cases, possess a re-entrant peristome

without any external branchie. They have grooves on the adoral

surface converging on the mouth, which may counteract what

seems to be an unprofitable structure. Moreover, among them

the indentation of the peristome is in all probability connected

with the accommodation of the large jaw-apparatus.

The Holectypoida offer no satisfactor y evidence as to the

velation between the peristomial and petaloid branchiew. In the

Upper Jurassic Pygasters (e. g., P. (Megapygus) macrocyphus)

there is a marked tendency towards petaloid structure in the

adapical parts of the ambulacra, but the branchial slits are as

well developed asin any of the earlier species. As the genera

are traced to the Upper Cretaceous, there is an irregular but

frequent tendency seen for the adapical ambulacral pores to

become dissimilar, but there is never any contrast sufficient to

SEA-URCHINS. 459

warrant a belief that ambulacral branchize were present. And

so in Conulus, in the almost complete absence of peristomial gills,

there seem to have been no special structures, either left or

developed, to perform the function of respiration. In the

Kchinoneide the saine condition obtains, but the Clypeastroida

show an ever increasing perfection of adapical petals to com-

pensate for the loss of the more primitive adoral branchie.

3. The Perignathic Girdle.

Detailed and valuable studies of this structure in Discoidea and

Conulus have been made by Duncan and Sladen (40 & 41) and

Lovén (43 & 48). These researches have the additional value

that they were pursued with different aims. The former authors

were intent upon demonstrating the absence of jaws in the two

genera, while Lovén predicted, and later realised, their discovery

in Discoidea. The structure of the perignathic girdles of Pygaster

and Holectypus is not so fully known, and in the case of

Anorthopygus there were no known traces of the girdle when

Duncan (44) placed the genus in the same family with Hchino-

neus. Lovén (48) knew of its existence in all the three genera,

but gave no details of its structure.

y Z Y

WOO

Diagrams of the perignathie girdles in

A. Pygaster. B. Anorthopyqus. C, Discoidea. D. Conulus.

In Pygaster the processes are very strongly developed, while

the ridges are hardly recognizable. There is no tendency for the

processes to form an arch over the ambulacra—in fact, they

slope away from one another. Thus there is initiated the

persistently disjunct girdle which characterizes all the gnatho-

stomatous Irregular Echinoids. In Holectypus the structure

seems to have been practically the same, but there are indications

that the ridges were slightly more pronounced, This was

certainly the case in Anorthopygus. In Discoidea the processes

460 MR. HERBERT L. HAWKINS ON

are proportionately similar to those of Pygaster, but their

prominence is almost masked by the extreme elevation of the

ridges. In Conulus this latter feature is carried so far that,

except for the suture-line showing that the process is present,

the only visible and free portion of the ambulacral part of the

girdle is a minute shining knob at each corner of the ridge.

It is, I think, a point of great interest to find that both

ingredients of the perignathic girdle are so strongly developed

in these later Holectypoid genera. One of the most obvious

contrasts between the divisions of the Clypeastroids is the

presence in some genera of one support for the jaws in each

interradius, and in others of two. It would seem that, when

they are double, these supports represent processes, and when

single, they are the degenerate relics of ridges. Both such

conditions could be obtained readily by the modification of a

perignathie girdle in which both portions were equally repre-

sented. All that is necessary is a simple process of the elimination

of one or the other of the parts.

Another feature of interest in the perignathic girdle of the

Holectypoida, and one connected intimately with the method of

use of the jaws, is the angle which the supports make with the

plane of the adoral surface of the test. In Pygaster the processes

are almost, though not quite, at right angles to that surface,

with a slight outward slope. This inclination is rather more

marked in Anorthopygus, and considerably so in Holectypus. In

Discoidea the angle between the girdle and the floor of the test is

quite acute, while in Conwlus it becomes, especially in thin-tested

forms, almost 45 degrees. The practically vertical girdle of

Pygaster would indicate a correspondingly vertical working of the

jaws, similar to that of the Regular Echinoids; while the highly

inclined system in Conwlus seems to show a tendency towards

the horizontal working of the jaws of Clypeastroids.

In Conoclypeus, according to the description and drawings

given by de Loriol (35), the two ambulacral processes are pr seats.

but the ridges have dwindled to insignificant proportions, both in

width and “height. This brings the processes closely together,

and there is consequently induced a markedly Glypeastroid

appearance in the perignathic girdle.

4. The Jaws.

Our knowledge of the jaws of the various genera of the

Holectypoida is very meagre and unequal. In fact, of the details

of the structure of the pyramids and teeth of Jurassic forms next

to no evidence is at present available. There are two reasons

why this condition of affairs should exist. Firstly, the jaws are

internal organs, and so, if they are preserved in the interior of a

specimen, it is necessary to break it up before they can be

studied. Moreover, it is usual to find the matrix that filtered

into the tests of Oolitic forms more compact and refractory in

SEA-URCHINS. 461

texture than the surrounding rock. Secondly, the large size of

the peristome in the earlier genera would be liable to let the

jaw-fragments slip through when their supporting muscles had

decayed. As all the species of Pygaster and Holectypus are more

or less conical in shape, the natural position that the test would

assume when allowed to settle on the sea-floor would be with the

oral surface downwards. After the jaws had slipped through

the peristome they would, on account of their relatively light

weight, become scattered by currents which were too gentle to

move the whole test.

Jaws are known to exist in Pygaster, but I have been unable

to find descriptions or specimens in which their structure was

adequately shown. From the characters of casts of the pyramids

preserved in an ironstone mould of P. ? semisuleatus that I have

seen, these parts of the lantern seem to have been large and

massive, and of a shape corresponding with that of the pyramids

of Cidaris. Wright (20) has figured a specimen of /olectypus

depressus in which the complete lantern is preserved. I have

examined the specimen (B.M., E. 1687), but it is impossible to

trace any of the ossicles to their extremities, so that no measure-

ments of any value can be taken. The general facies of the

pyramid is strikingly “ Regular.” Nothing seems to be known

of the jaws of Anor thopygue, but they must certainly have

existed.

For a long time the presence of jaws in Discoidea was doubted,

and sometimes, notably by Duncan (41 & 45), absolutely denied.

In 1892, Lovén, in the wonderful store of information as to the

perig ionathic structures of Echinoids contained in his Kchinologica

(Loven, 48), gave a description of the pyramids in D, cylindrica,

and recently I was able (Hawkins, 60) to confirm and amplify his

description with the additional features of the epiphyses and the

teeth. In this genus the pyramids have still a markedly

“Regular” appearance, although they were probably much more

closely attached to the processes of the perignathic girdle than

in any Regular Kechinoid. This shortening of the muscles of

attachment resulted in a far less vertical position for the lantern

as a whole, while the strong incurving of the adoral parts of the

pyramids will have increased the angle to one of about 45 degrees

at the peristome. The teeth are curved considerably to correspond

with this arrangement. They are strong, and built on the

Echinoid plan, in contrast to the Diademoid, with a pronounced

keel on the concave side.

In the case of Conulus, the long controversy as to the presence

or absence of jaws has been partly settled by the discovery of

teeth in a specimen of C. subrotundus (Hawkins, 65). There is

as yet no evidence as to the characters of the jaws; and the teeth

in themselves, beyond their similarity to those of Discoidea, show

no features of special interest. They are less curved than those

of that genus, and more sharply pointed, the latter character

being in contrast to what might be expected in view of the

462 MR. HERBERT L. HAWKINS ON

bluntness of Clypeastroid teeth. The peculiar structures in

C. albogalerus, usually known as the ‘“ buccal plates,” are probably

in some way derived from jaw-ossicles, as their anomalous

character separates them absolutely from the peristomial plates

which exist in many other genera. In the paper to which

reference has just been made, I have suggested a possible origin

and function for the buccal plates, but it must be confessed that

the theory advanced there has a very insecure foundation.

So far as 1s at present known, there is nothing in the structure

of the lantern of the Holectypoida which even foreshadows the

curiously expanded pyramids of the Clypeastroida. The probable

delicacy of texture of the pyramids in Conulus subrotundus may

indicate the incoming of a reticulate structure similar to that

of the corresponding parts in Clypeaster. With regard to the

manner of working, the angle of setting of the Jaws shows a

progressive tendency towards the Clypeastroid method. This

retention of the ‘‘ Regular” facies of jaw-structure throughout

the group is rendered the more remarkable by a comparison

with the fragmentary pyramids in Conoclypeus described by

de Loriol (35). That genus, with its Clypeastroid (almost

Kehinanthine) general build, seems to have possessed the compact

pyramids of a Discoidea. It is true that the only record of its

jaws is very imperfect, but this much seems obvious on a study

of de Loriol’s drawings. But in Conoclypeus, in spite of the

Holectypoid jaw-structure, the perignathic girdle is very like

that of Clypeaster.

The recently described teeth and lantern in a young specimen

of an Hchinoneus (Agassiz, 58) have a most important bearing on

the relation between the Holectypoida and the Kchinoneide.

The presence of the jaws is undoubtedly a vestigial character,

for they seem to be resorbed while the individual is still quite

immature. The jaws and teeth both have a Discoidea-like

appearance, rather than a Clypeaster-facies. It is probable that,

as they exist in the young stages of Hchinoneus (the most,

advanced member of its family), they will have been present in

such genera as Pyrina at a corresponding stage of development.

The likelihood of their discovery in fossil forms is extremely

remote, owing to their minute size and delicate texture, but

analogy tells strongly in favour of their existence. This dis-

covery 1s a remarkable instance of the completion by Ontogeny

of an unfinished chain of evidence supplied by Paleontology,

and removes any doubt which may have existed as to the

intimate relationship which links the Conulide with the early

Echinoneide.

In this connection it seems well to suggest the possibility that

Lovén’s genus Pygastrides (Loven, 43), a “ Pygaster” lingering so

long after the day of the Holectypoida was past, may be only

another example of the vestigial gnathostomatous stage of some,

probably Echinoneid, genus. So strongly am I of this opinion,

that I have omitted its name from the new classification.

SHA-URCHINS. 463

c. Che Periproct,

Since the excentric position of the periproct, outside the apical

cycles of plates, is a diagnostic feature of the Irregular Echinoids

as a whole, it is natural to find that in the Holectypoida, which

includes the most primitive of the ‘ Exocyclic” forms, its position

is very variable. When once the periproct has left the apex, its

chief tendency seems to be to assume a position as absolutely

posterior as possible, and in the course of its passage to such a

position, it undergoes many changes itself, and is the cause of

many others to the test. It always lies in the posterior inter-

ambulacrum.

1. The Position of the Periproct.

Practically the only distinguishing feature between a young

specimen of a Pygaster sens. str. and a primitive Diademoid is

the fact that in the former the periproct has broken through the

posterior part of the apical system. It cannot be said to lie

altogether outside the system, for to some extent it occupies the

position of the posterior genital plate, and extends well up to the

apex of the test. It is, im part, more nearly central in position

than in some of the Saleniide, although its large size causes it to

reach away from, as well as into, the apical system.

In Pygaster sens. lat., the periproct is always in contact with

the apical system,—in the earliest forms reaching to the inner

margins of the anterior and antero-lateral genitals, and in the

later ones touching only the outer margins of the redeveloped

posterior genital. When traced from Pygaster sens. str. to

Macropygus, however, the position of the widest part of the

periproct is found to pass gradually backwards, while the posterior

edge of the opening approaches the ambitus of the test. Pygaster,

then, shows a stage in which, although the periproct retains

its primitive association with the apical system, the posterior

tendency in its position is recognizable.

Pileus and Anorthopygus agree in having the periproct entirel

on the adapical surface of the test, but quite separated from

the apical system. In most cases, however, the posterior part

of the periproct is not so near to the ambitus as in Macropygus.

In Holectypus we find two groups, as regards the position of

the periproct. Both groups appear at almost the same stage of

the Lower Oolite, but one is more retarded in character than the

other. The former, which may be exemplified by H. hemi-

sphericus, has the periproct opening on the margin, that is, in

the posterior extremity, of the test. The latter, of which a

common representative is H. depressus, has the periproct entirely

on the adoral surface, and often very close to the peristome, It

would appear at first sight that in the former group the periproct

had reached the necessary limits of its retrogression, and in the

latter had, as it were, overshot the mark. But, in the light of

464 MR. HERBERT L. HAWKINS ON

the later genera, it is impossible to regard the adorally situated

periproct of H. depressus as a case of overspecialization. Most

of the species of Comnholectypus have the periproct in that

position, as have all the forms of Discoidea. The H. hemi-

sphericus character reappears in Conulus, and is retained in most

of the Cretaceous species of Pyrina.

Text-fig. 56.

neg

igi

Diagram showing the shape and position of the periproct in A. Pygaster sens. str.,

B. Megapygus or Macropygus; C. Anorthopygus; D. Holectypus (hemi-

sphericus); KE. Conulus; ¥. Holectypus (depressus) ; G. Discoidea. The

thick line represents the ambitus. The proportionate height of the inter-

radial plates is indicated.

Lovén (36) has indicated the correspondence in periproct-

migration that exists between the Holectypoida (his Echinoconide)

and both the Echinoneide and Cassidulide. G'aleropygus and

Clypeus both have the Pygaster-like periproct, in contact with

the apical system, while the Nucleolitide show an arrangement

more like that of Pilews. The Echinolampide include forms in

which the periproct may be marginal or adoral in position.

In connection with the position of the periproct, an interesting

feature of the posterior interradius may be discussed. Reference

SEA-URCHINS. 465

has been made already (p. 455) to the truncated form of many

of the Pygasters, and of Galeropygus, and the comparatively

elongated outline of some /olectypi and of Conulus. The

truncation of the posterior interambulacrum is associated with

the presence of a more or less defined posterior sulcus; and, in

fact, whenever the periproct is on the adapical surface, this

sulcus is developed. The functional value of a suleus below the

anus is obvious, as if would tend to restrict the passage of fecal

matter to a definite channel, and so to prevent it from coming

in contact with the podia, But the developmental meaning

of the structure would seem to concern the interference in the

growth of the interambulacral plates by the periproct. These

plates, formed at the apex and foreed downwards towards the

ambitus, have to separate along their median sutures to pass

round the periproct, and subsequently have to close together

below it. The irregularity thus caused results in a retardation

of their downward movement (and a consequent shortening of

the distance from the apex to the posterior margin of the test),

and in a sagging inwards of the reconstructed portions of the

plates to form a groove. The probability of this explanation of

the structure so characteristic of Pygaster, Galeropygus, and the

Nucleolitide, becomes increased when the opposite conditions

are considered, When the periproct is marginal or inframarginal

in position, the interambulacral plates can pass freely over the

adapical surface until the edge of the periproct is reached, Here,

in consequence of the lessened width of the divided halves of the

area at the sides of the periproct, a delay in the progress of the

plates occurs. Asa result, the oncoming plates become heaped

up against one another above the periproct, and give rise to the

elongated, carinate posterior shape which characterizes Holectypus

sens. str., Conalus, and also the Spatangoida.

Although it must be admitted that the two opposite conditions,

suleate and carinate, of the posterior interradius have, from a

teleological standpoint, an obvious and similar functional value,

the explanation given above seems natural in view of their

regular association with the position of the periproct. In the

case of the Spatangoida, there is the complication of a “posterior

surface” to the test, at the upper part of which the periproct is

situated. If this surface were curved in conformity with the

rest of the test, the periproct would open at a point about

midway between the apex and the ambitus, as in Pileus or

Anorthopygus. It seems to me to be a very striking fact that,

in the last-named genus, no trace of a posterior sulcus is developed,

but that the declivity of the test is appreciably increased as a

whole in the region behind the periprect. The Spatangoid

posterior surface might, then, be regarded as the product of a

retarded growth of the entire posterior interambulacrum owing

to the interference of the periproct, while the carina above it

would be caused by the same agent in its opposite intluence.

In the few cases where absolute circularity of outline is

Proc. Zoou. Soc.—1912, No. XXX. 30

466 MR. HERBERT L. HAWKINS ON

regained in the Irregular Echinoids, as in Discoidea and many

Clypeastroida, the periproct is so small as to necessitate very

little modification of the steady progress of the coronal plates

from the apex to the peristome.

2. The Shape of the Periproct.

In Pygaster sens. str., the periproct is roughly elliptical in

shape, and very large, often having the same width (in transverse

measurement) as theapicalsystem. In Megapygus and Macropygus

its outline becomes pyriform, owing to the partial closing in of

the interambulacral plates round its adapical extremity. ‘The

width never becomes greater than in Pygaster sens. str., but the

actual size is much larger in these later subgenera, owing to the

backward shifting of the posterior edge of the ‘periproct without

a corresponding retraction of the adapical margin. The pyriform

shape caused by this lagging behind of the upper part of the

periproct leaves its impression on the shape of the aperture in

later genera. In Holectypus, for example, the periproct has

its adoral margin rounded, but adapically it tapers to a point.

The same feature is seen in Pileus. In Holectypus sens. str., the

periproct is still large; in some species, e.g., H. depressus, it 1s

of an extraordinary size. But in Canholectypus it. has generally

decreased so as to be smaller than the peristome. In Anortho-

pygus the periproct is of moderate size, and has a characteristically

oblique position. Obliquity in the case of the peristome is not

uncommon among Irregular Echinoids (e. g., Pyrina and Tremato-

pygus), but this is practically the only form where such asym-

metry affects the shape of the periproct to a considerable degree.

In this connection it is interesting to find that in P. (Megapygus)

umbrella the large pyriform periproct shows a distinct inclination

towards the left side of the interradius, thus giving an indication

of potential obliquity.

In Discoidea the periproct is usually lanceolate in outline,

often equally pointed at both extremities, but it is always longer

than broad. In Conwlus the marginal periproct is similar in

shape to that of Holectypus hemisphericus, although smaller im

size. The pointed character of its adapical part is more pro-

nounced in young specimens than in adults. The size and

shape of the periproct in the Echinoneide compares well with

those in the Conulide. In the Clypeastroida the periproct is

always adorally situated, as in Discoidea, but it is very small, and

usually circular in shape.

3. The Anal Plates.

The plating of the periproct-membrane is at present unknown

in Pygaster, Pilews, and Conulus. In the case of the two genera

first named, this is probably due to the large size of the periproct,

SEA-URCHINS. 407

and the resulting flexibility of the membrane and weakness of

the plating. In the case of Conulus (and also of those Holectypi

which have the periproct marginal), the exposed position of the

anal plates on the ambitus may account for their non-preservation,

A thickly plated membrane occupies the periproct of Hchinoneus,

In Discoidea the plates of the periproct are not infrequently

found in situ, and they are known in Canholectypus, Anortho-

pygus, and Coptodiscus. ‘The last-named genus differs, as regards

this character, from all the others, in possessing a single ring of

almost equal- ‘sized plates around the inner margin of the

periproct, and in not having, so far as is known, any smaller

plates in the immediate surroundings of the anus.

Discoidea has one large anal plate, usually bearing a tubercle,

occupying most of the adoral half of the periproct-opening, and

a Series of fringing plates which decrease in size as they approach

the adapical part of the aperture. <A few, often only two, small

plates occur within this irregular ring, and they are always in

contact with the largest plate. The anus is thus situated quite near

to the adambital edge of the periproct, in a position far removed

from the mouth. In Canholectypus, to judge by a figure of

C. jullient from Algiers (Péron & Gauthier, 34), the arrange-

ment was on a similar plan. There, however, the adorally

situated plate is relatively small, and the fringing plates are also

smaller and more numerous than in Discoidea. The inner anal

plates are exceedingly minute, and are preserved in considerable

numbers.

In Anorthopygus | have been able to study only the outlines

of the anal plates, these being easily traceable on a siliceous

mould of 4, orbicularis in the British Museum. In this specimen

the arrangement of the plating is exactly the reverse of that

which obtains in the two genera just described. The largest of the

anal plates are adapically situated in the oblique periproct, and

a series of pentagonal and hexagonal plates, of approximately

equal size, covers all the remaining surface of the aperture

except fora very sinall area in its extreme adoral part. The

actual anus, which is represented in the mould by a prominent

unsutured portion of the infilling matrix, lies in the true antero-

posterior axis of the test, thus being unaffected by the asymmetry

of the periproct as a whole. There seems to have been no space

occupied by plates between the anus and the periproct margin.

The position of the anus, in its relation to the situation of the

periproct on the test, is interesting. When the aperture is on

the adoral surface the anus tends to open in its adambital corner,

while the same tendency, with an opposite effect, appears when

the periproct is supramarginal. A generalization, founded on

the somewhat slender evidence of only three generic types, may

be made that:—Wherever the periproct may be situated, the

anus assumes a position within its borders as near to the ambitus

(i.e., the most posterior part of the test) as possible.

30*

468 MR. HERBERT L. HAWKINS ON

D. The Ambulacra.

1. The Podial Pores.

Although one of the diagnostic characters of the Holectypoida

consists in the apetaloid nature of the ambulacra, it would be a

mistake to assume that the pore-pairs are therefore similar

throughout the group. The Nucleolitide, which are a group of

almost the same antiquity as the Holectypoida, early developed

a marked heteromorphy in the podial pores of the adapical

surface; anda similar character, continually recurring, but as

often held in check, is apparent among all the Jurassic members

of the order. Pygaster sens. lat., has uniformly larger pores on

the adapical than on the adoral surface, and the members of an

individual pore-pair are dissimilar in the former region of the test.

Even Pygaster semisulcatus sometimes shows this feature. The

outer pore of the pair is a little larger than the inner, although

both are somewhat ellipticalin shape. In P. (Jlegapygqus) umbrella

and still more in J. macrocyphus, the difference becomes increased.

The inner pore is circular, and the outer retains an elliptical

shape, often on quite an elongated plan. The long diameter of

the outer, elliptical pore never becomes more than Ewite as great

as the dinnagiez of the inner, circular one; so that. the whole

ambulacrum cannot be said to show even a subpetaloid structure.

In Pygaster, while this dimorphism of the adapical pore-pairs

increases, the size of the pores on the adoral surface steadily

decreases. ‘These latter pores are always circular, and the members

of each pore-pair are separated by a prominent granule. Their

small size renders them quite difficult to distinguish in the

Upper Jurassic forms.

In Holectypus a similar tendency is seen, although it is hardly

appreciable in the Cretaceous subgenus. The diversity of shape

and size in the adapical pore-pairs is rarely carried so far as in

Pygaster, but the reduction in the diameter of the ambital and

adoral pores is quite as well marked. In the case of Discoidea

the tendency is less noticeable. The pores of the adapical

surface are only very slightly larger than those of the adoral

(both series being minute), and are themselves always circular.

The outer member of a pore-pair is sometimes just distinguishable

from the inner one in point of size.

In Conulus, by way of contrast, the pores of the ambulacra are

everywhere exceedingly minute, those of the adapical surface

being even smaller than those of the adoral. The largest pores

in this genus are generally situated on or near the ambitus.

Discoidea, and to a further extent Conulus, may be regarded as

illustrating the triumph of simplicity of ambulacral structure over

the persistent tendency to complexity which induced variation in

the earlier genera. In the case of Discoidea, the simplicity would

seem to have heen short-lived as soon as its successor, Conoclypeus,

had emerged from the order Holectypoida, and initiated the

Clypeastroida, where often the petals are developed to a great

SEA-URCHINS, 469

degree, As regards Conulus, only the Echinoneide, among its

external relatives, retained the apetaloid cienieler: The

Kchinolampidee sad the Conulopsis group show a pronounced

subpetaloid development. Nevertheless, the fact remains that, in

the matter of their ambulacral pore- -structur e, the later genera of

the order conform more absolutely to the letter of the diagnosis

than the earlier forms.

Text-fig. 57,

ATER CERCA»

Diagram showing some characteristic plates of the ambulacra (from the adapical

surface) in A. Pygaster sens. str.; B. Galeropygus; C. Megapygus ;

D. Pileus; E. Discoidea; F. Conoelypeus; G. Conulus; H. Pyrina.

The pore-pairs in Pilews show an anomalous character in being

biserial on the adapical surface, without any corresponding in ter-

ference with the primary nature of the ambulacral plates. There

seems nothing among Irregular Echinoids to compare with such

a condition, which recalls the similarly inverted development of

biserial pore-pairs in the ambulacra of Diplopodia, 'Thesomewhat

analogous appearance of the pores in the anterior ambulacrum of

the Spatangoid Heteraster is accompanied by “ plate-cr ushing ” in

the structure of the area. Pilews, in this respect, as in some

A470 MR. HERBERT L. HAWKINS ON

others, must be regarded as a curiously specialized offshoot from

the Pygaster-stock, which only survived a short time, and left no

descendants.

2. The Ambulacral Plates.

Recently, inthe ‘Geological Magazine’ (Hawkins, 62), I indicated

in outline the principles of ambulacral structure which charac-

terize the Holectypoida. Later (66) I extended the line of

enquiry to the other Jurassic groups of Irregular Echinoids, and

showed the influence that plate-structure exerts on the features of

the phyllode. It will, therefore, be necessary only to summarize

the results of those studies here, for the sake of completeness.

All the Holectypoida show a crushing together of the primaries

to form compound plates in their ambulacra. The degree of

crushing is a progressive one. Most of the ambulacrum of

a Pygaster is composed of primaries, while hardly any unmodified

primaries remain in the ambulacrum of a Conulus. The building

of the compound plates is carried out on a perfectly uniform plan,

three original plates going to form one compound plate. The

significance of this triple arrangement will be discussed at the

beginning of section V. of this paper. Conulus differs from all the

other genera of the order (except the little-known Discoholectypus),

partly in the early stage at which the crushing commences,

and partly in the fact that two ont of the three plates concerned

retain their primary character (though modified in shape) for

a considerable distance beyond the first crushing point, often

right down to the ambitus. Two genera may be cited, re-

presenting two widely divergent groups, which show an exactly

similar ambulacral structure. These are Pyrina, of the Echino-

neidee, and Amblypygus, of the Kchinolampide. It is hard to

believe that so peculiar a structure can have been evolved four

times independently. .

Although there is no true phyllode-structure (nor appearance)

developed in the adoral parts of the ambulacra in any of the

Holectypoida, the nature of their plate-crushing inevitably

results in a “hypophyllodal” character (see Hawkins, 66) of that

region. It is not until Conulus is reached, however, that the

displacement of the plates drives the pore-pairs into a definitely

triserial order. In Pygaster the pores hardly deviate from a

straight line throughout the length of the ambulacrum, and no

regular displacement can be traced in the poriferous zones

of Holectypus. In Discoidea the pore-pairs become appreciably

triserial midway between the ambitus and the peristome, but

recover their linear arrangement before the peristome is reached.

In Conulus a triserial character appears practically at the ambitus,

and becomes more pronounced as the ambulacrum is traced

towards the mouth; until, near the peristome, the triads are

inclined at an angle of 45 degrees to the direction of the radius.

The greatest difficulty that appears when an attempt is made

to trace a phylogenetic sequence from the Holectypoida to the

SEA-URCHINS. 471

Clypeastroida is the presence in the latter group of a few large

ambulacrals in the adoral parts of the area, with no signs of plate-

crushing. Can a compound plate be resolved by evolution into

its constituent primaries, or is the simplicity of the Clypeastroid

ambulacral only apparent, being in reality the result of the

fusion of the components of a compound plate, followed by the

atrophy of two of the three pore-pairs? Bather (59) has

expressed his belief in the possibility of the former process in his

discussion of the ambulacrals of Orthopsis. In support of the

alternate suggestion it may be remarked that in the Clypeastroids

the pore-pair “of each large polygonal ambulacral is situated near

the adoral margin of the plate, leaving a high non- poriferous

region along the rest of the adradial margin. Moreover, in the

case of Discoidea just cited, the triserial ar rangement of the pore-

pairs is arrested soon after its inception, and the poriferous zones

again become straight. There seems to be no indication of

a corresponding reduction in the degree of compression of the

demi-plates towards the peristome, but rather an increase, until

the platelets become so minute that the small pore-pair can hardly

find room to pass through the test within its borders. The

presence of this feature of simplification in the sequence of the

pores, but not in the structure of the ambulacral plating, in

Discoidea seems especially significant ; for Discoidea is the nearest

ally of the Clypeastroids that is found among the Holectypoida.

However, I do not feel justified in expressing a positive opinion,

in one or the other direction, upon this question. Much must be

done in the study of the postlarval growth of the test in the

Clypeastroids before any proof of the origin of their ambulacral

structure can be expected.

EK. The Intsrambulacra.

l. The Interambulacral Plates.

The interambulacral are always much broader than the

ambulacral areas, and the proportionate width (about 3: 1 at the

ambitus) is retained almost unchanged from Pygaster to Conulus.

Owing to the absence of expanded petals and phyllodes in

the ambulacra, there is no compression of the adapical or adoral

extremities of the interambulacra such as occurs in most of the

Irregular Echinoids. The areas increase regularly in width from

the margins of the genital plates to the ambitus, and decrease

as regularly, though more rapidly, from the ambitus to the

eristome.

In Holectypus depressus there are shallow pits on the transverse

sutures at points directly above the branchial slits. I know of

no evidence which could ascribe a function to such features,

The interradial suture is usually only slightly zigzag in character,

and in some forms, notably among the Jurassic Holeckypi, it is

practically straight, so that the plates become roughly rectangular

A472 MR. HERBERT L, HAWKINS ON

in outline instead of being pentagonal. The plates of the adapical

surface are generally much broader than high, but on the adoral

surface this difference is lessened. In Holectypus sens. str., the

contrast in the height of the plates of the two surfaces results in

the presence of very few interambulacrals on the adoral surface.

In Pygaster the difference is not so strongly marked.

The only member of the group in which any striking difference

in the appearance of the interambulacral plates themselves occurs

is the peculiar genus Coptodiscus. Here, in a form otherwise

hardly to be distinguished from Cenholectypus, all the margins

of the plates are bevelled, so as to leave deep grooves along the

sutures. This feature, which recalls the Siler structures

in Goniocidaris and the Temnopleuride, is restricted to the

adapical surface. Whether it is a result, in this case, of a paucity

of carbonate of lime in the water, or of some physiological

peculiarity, it is impossible to judge. The feature seems to be

quite unique among the Irregular Kchinoids.

2. The Primary Tubdercles.

In their structure and proportions, the primary tubercles show

no more variety, when traced through the group, than do the

radioles that they support. The equality in size of those of the

adapical and adoral surfaces, which is marked in P2 ygaster, becomes

gradually replaced by a tendency towards an increase in size of

the adoral tubercles, with a corresponding decrease of those of the

adapical surface. In Discoidea, especially in D. subuculus, the

reduction of the adapical tubercles has proceeded so far that they

can hardly be distinguished im size from their attendant miliaries.

Apart from a tendency in Conulus for the boss to become wholly

convex in side view, and so fill the scrobicule more completely

than do the partly concave sides of the boss in Pygaster, there are

no changes of importance to be traced in the actual structure of

the tubercles.

In the arrangement of the tuberculation more variation is

found, and there becomes manifest a continual tendency towards

a progressive increase in its complexity. I have dealt with this

character (Hawkins, 67) in considerable detail, and give here

a summary of the results obtained in my recent paper.

As Saemann and Dollfuss (27) showed, the actual number

of tubercles present on each interambulacral plate depends largely

on the size, that is, on the age, of the individual. In all the

Holectypoida there is at least one plate, at each end of the half

interradius, which supports a single tubercle. This is obviously

a relic of the primitive, unituberculate character of the plates

of the earlier Regular Echinoids. The number of such plates

remaining decreases steadily as the group is traced from the

Lower Jurassic to the Upper Cretaceous. The median series of

tubercles persists in an unbroken line from the apex to the

peristome, but, except in Pygaster, is not readily distinguishable

SEA-URCIIINS. 473

from its associates in point of size. Each tubercle in this median

> the adoral than the adapical

series is placed slightly nearer

transverse margin of the plate. All the other tubercles, of which

Text-fig. 58.

Soo sa -

: Oe OC SU OVO) So ais

Pie tans pea eee

é 2 ae wo 8s 90009

showing the third plate above the ambitus in A. Pygaster sens. str.-

. Holectypus (hemisphericus) ; C. Holectypus (depressus) ; D. Discoidea ;

Diagram

—

E. Galeropygus ; F. Anorthopygus ; G. Conulus. Figs. H & K are side

views of primary tubercles in Pygaster and Conulus respectively.

great numbers may occur in the later genera, are developed in

recognizably vertical series. When their number becomes great,

a natural tendency to slight irregularity appears, but this is never

ATA MR. HERBERT L. HAWKINS ON

sufficient to mask the plan of the tuberculation. Ivrregularity

generally consists of either the suppression of a member of

a series on one plate (a feature often seen, even in the primary

row, in Hchinus), or its replacement by two tubercles. The

Holectypoida are peculiar among Irregular Echinoids in retaining

throughout life the unituberculate plates and the vertical arrange-

ment of the tubercles. Only the Echinoneidee show any similar

plan, and in them it is very much obscured by irregularity

of development. Some forms of Pygurus (e.g., P. blumenbachi)

have one or more unituberculate plates at the adapical extremities

of their interambulacra, but this retardation of development is

obviously due to the considerable narrowing of the plates caused

by the expansion of the ambulacral petals. However, most, if

not all, of the Irregular Echinoids whose post-larval development

has been studied, show a unituberculate stage. This is notably

the case in Hchinolampas (see Agassiz, 30).

_ The tubercles of the additional series* which develop on the

interradial tracts are at first situated each in the middle line (in

a vertical sense) of the plates. This results in a transverse line

of tubercles extending between the main series and the inter-

radial suture. The concentric arrangement thus caused charac-

terizes Pygaster sens. lat., Holectypus sens. lat., and Discoidea.

It seems somewhat irregular in the case of Pilews, and is definitely

absent in Anorthopygus and Conulus. In these two genera the

tubercle series of the interradial tracts appear near to the adapical

and adoral margins of the plates alternately, thus giving an

oblique arrangement (sloping interradially and adorally) to the

tubercles in the complete interambulacrum. A similar arrange-

ment to this affects the adradial tubercle-series in all the genera

of the group.

The oblique setting of the tubercles results in a much more

uniform and packed tuberculation over the whole area than

exists when the interradial series are transverse. The closeness

of the arrangement is increased by the doubling of many of the

tubercles in Conulus ; so that, instead of two tubercles, three or

even four are concerned in the composition of the oblique line on

each plate. The complexity of arrangement, coupled with a

homogeneity of character, of the tubercles, which was thus slowly

obtained during the course of evolution of the Holectypoida, was

rapidly developed, and carried to a further degree, by the earliest

of the non-Holectypoid Echinoids. The species of Galeropygus

from the Lias show typically the bewildering profusion of small

tubercles which characterize the interambulacra of all the

Nucleolitide, ‘‘ Cassidulidee,” Clypeastroida, and early Spatangide.

Only the Kchinoneide seem to preserve a Conulus-character in their

tuberculation, and in them it becomes so irregular as to be hardly

appreciable except in the newest formed plates.

The peculiar sunken supernumerary tubercle of some of the

adapical interambulacrals of Holectypus depressus from the Corn-

brash recently described (Hawkins, 67) is without a parallel

SEA-URCHINS, 475

among the other members of the order, Although suggesting a

comparison with the large interpetalous interambulacral tubercles

of such a genus as Hupatagus, it seems to be a specific character

of no genetic value.

3. The Miliary Granules.

In Pygaster, and to a less degree in Holectypus, the granulation

shows a “ Regular ” affinity in being grouped around the primary

tubercles to form scrobicular rings. This circular arrangement is

not retained after the Jurassic period. In Holectypus, especially

in H. depressus, the scrobicular miliaries on the adradial tracts of

the plates near the peristome are often guttate in form. In

Macropygus and Anorthopygus the large size of the scrobicules of

the primary tubercles leaves little room for miliaries, but those

which exist do not appreciably group themselves around the

primaries. In Discoidea and Cenholectypus, and to a slight

degree in Holectypus sens. str., rows of granules radiate from the

central primary tubercle, maintaining a more or less transverse

direction, to reach the adradial and interradial margins of the

plate. In the genus first named these linear rows of granules

become interspersed, near the ambitus, with additional tubercles.

In Conulus the granulation is apparently without a definite

system of arrangement. The granules are sunk slightly below the

level of the test on the adapical surface, being enclosed in minute

pits. On the adoral surface they regain their projecting character,

and often occupy broad bands, slightly elevated, which correspond

in position with the transverse sutures of the plates,

Ea ives Kia dia oullers:

Our knowledge of the acanthology of the Holectypoids is

fragmentary and inadequate. Enough is known, however, to

show that there exists a considerable uniformity in the character

of the radioles throughout the group. Wright (20) has described

the primary radioles of Pygaster, in the species semisulcatus and

(Macropygus) morrisii. For both he uses almost the same words—

short, needle-shaped bodies with fine longitudinal lines on the stem.

I have not seen any specimens in which they are preserved. In

the case of Holectypus there is a specimen of /. depressus (from

the Inferior Oolite of Cheltenham) in my collection which retains

a considerable number of radioles on both the adapieal and adoral

surfaces. Wright (¢.c.) describes them in the same species. The

primaries of the upper surface are very short and slender, with

blunt tips. Those of the adoral surface were apparently quite

long, and but slightly tapering. The collars of the adoral radioles

are prominent, and often very oblique. The shafts of both sets

of radioles are longitudinally fluted with closely-set ribs. The

miliary granules support spines of a similar character to the

adoral primaries, but far more minute, so that Wright’s descrip-

tion of them as “ hair-like” is accurate.

476 MR. HERBERT L. HAWKINS ON

I have been unable to find any record of the preservation of

radioles in Discoidea. In Conulus the primaries are very much

like those of the adoral surface of Holectypus in shape and

ornament. The miliaries support curiously. blunt prominences,

which are usually preserved im situ, but are very easily rubbed off

by too vigorous development of the specimen. These blunt spines

were figured by Forbes (14), who also gave a drawing of a curious

body that he regarded as a pedicellaria. The characters of the

miliary spines suggest a comparison with the calcareous supports

of pedicellariz ; but if they all had this function, the number of

those organs would be extraordinarily great in proportion to the

radioles.

Nothing seems to be known as yet of the microstructure of the

radioles of any genera of the group, as Hesse (51) did not choose

an Holectypoid for his researches.

The slight progressive change traceable in the primary radioles

seems to lead merely to an increase in the length, and perhaps in

the slenderness, of their shafts ; while,as would be expected from

the difference in size of the tubercles on the two surfaces of the

test, the adoral radioles become proportionately longer than those

of the adapical surface.

G. Internal Buttresses.

The difference in form which makes so violent a contrast between

a Scutella and an Hchinus must result ina corresponding difference

of resisting power against the pressure of the waves. As both

types of Echinoid may live between tide-marks, where the violence

of the waves is most felt, they both have the same forces to repel.

A spherical body such as that of an Hehinus, or even a hemi-

spherical one, like that of an Holectypus, could easily ward off the

blow of a breaking wave, in the same manner as a Patella does. But

a flat test, such as that of a Scutellid, would offer a blank resistance

to the waves, and, if hollow, would almost certainly be crushed.

For this reason, the few groups of the Irregular Echinoids that

frequent the exposed littoral habitat so characteristically occupied

by the Regular forms, strengthen the resisting power of their

tests by the development of massive calcareous buttresses within.

Practically the only Irregular forms which live openly on the shore

at the present day are the Clypeastroida. It becomes, therefore,

a point of especial interest to find the beginnings of internal

supports to the test among the Holectypoida, which is the only

other gnathostomatous (and therefore rock-dwelling) order.

Although the development of the buttresses in the two groups

might easily be regarded as an illustration of similar adaptation

to similar environment alone, yet, in the light of the other less

obviously utilitarian features of the two groups, it seems in this

case that a genetic explanation exists as well.

On the internal mould of a Pygaster, and yet more in one of a

Pileus, besides the deep pits left by the prominences of the

SEA-URCHINS. ATT

perignathic girdle on the adoral surface, there are grooves

(representing ridges in the test) that pass from the processes

alongside the ambulacra (but situated on the interambulacra) for

a short distance. The structure might be regarded as indicating

a gradual rise of the inner surface of the test to form a keel which

culminates in the perignathic process. As, among Regular

Kchinoids, the perignathic girdle rises quite abruptly from the

inner surface, this gradual rise of the test towards the processes

shows a new feature, the beginning of the inner buttressing of

the test.

In Holectypus, as the name implies, a diagnostic feature of the

genus as first tentatively suggested by Desor (11) is the absence

of grooves in the internal ‘moulds. This of course means the

absence of internal ridges passing radially outwards to a point

beyond the ambitus. An investigation of some siliceous moulds

of H. ? sarthacensis has shown me that, although there is nothing

in the interambulacra to compare with the strong ‘“ cloisons” of

the succeeding genus, there nevertheless exists a considerable

thickening of the adoral regions of those areas, even more than in

Pygaster. In Discoidea the first signs of a really efficient internal

butitressing appear. Down a line, rather to the adradial side of

each half-interradius, there passes a thickening of the test which

is rounded near the peristome and becomes carinate further out,

and which extends beyond the ambitus. The partitions do not

pass for any considerable distance up the adapical surface. The

perignathic girdle tends to lean against the adoral’ ends of the

supports.

In Conulus no such well-marked buttresses appear, but the

interambulacral areas undergo a great amount of thickening

towards the peristome. Indeed, the perignathic girdle, which is

itself well developed, is often less internally elevated than the

interambulacral plates against which it reclines. The ambulacra

pass in sunken grooves across the adoral surface. In one specimen,

on cutting a section through the interambulacrum at a point just

outside the perignathie girdle, I found a large hollow to be included

between an inner and an outer wall of calcite. I have not been

able to verify the occurrence of this feature im other sections. It

may, therefore, have been an abnormality or the result of an

accident ; but if it should be found to be a general tendency, or

even one of fairly frequent occurrence, it would be very significant

in the comparisons that might be drawn between it and the double

flooring of the test of many of the higher Clypeastroida.

It is only in Discoidea that the buttresses are in such a freely

projecting condition that they could be expected, by a growth in

their height and an accompanying depression of the adapical

surface, to form complete vertical partitions in the test. As it is,

these “ cloisons” of Discoidea are rather more strongly developed

than the corresponding structures of Hchinocyamus, which other-

wise they resemble very closely. In fact, Gregory (50), in renaming

the “ genus” called Hehinites by Duncan (44), which included only

478 MR. HERBERT L. HAWKINS ON

the species Discoidea subucula, used the name Protocyamus “ to

indicate the affinity of this Echinoid with the Hchinocyamus

series.” The name, on systematic grounds, must be abandoned,

but its significance remains.

In Conulus, but, so far as I am aware, in that genus only, a

definite “sand-canal,” similar to that of Hchinocorys and the

Spatangidee, is well developed on the inner surface of the madre-

poric genital. In the same genus, in adult specimens, a double

row of hemispherical prominences occurs, partly encircling the

inner part of the test a little above the ambitus. Klinghardt (68)

has recently discussed the relation of these thickenings to the

course of the alimentary canal, for the mesenteries of which they

seein to have given attachment. He has compared the course of

the gut thus indicated with that of several fossil and recent species

of Spatangide. In the present state of our knowledge, however,

but little of importance can be ascertained of the comparative

anatomy of the soft parts of fossil Echinoids.

H. The Apical System.

Much of the systematic work that has been done on the Irregular

Eehinoids has had as its basis the character of the apical system.

Notable cases where this feature has been utilized for the purposes

of classification are the works of Gaudry (‘ Enchainement du Monde

animal’)and Pomel. Froma purely morphological standpoint the

system has been carefully described by Lovén (81). In the case

of the Holectypoida, and of some of the near allies of that group,

I have recently summarized the state of our knowledge of the

apical system (Hawkins, 70) in a paper that was definitely a

preliminary note to the present work. In consequence, only the

comparative aspect of the subject need be dealt with here, and for

the description of details that paper may be consulted.

The apical system is at first thoroughly disorganized in its

composition and structure by the passage of the periproct through

its cycle of plates. In Pygaster sens. str., the first stage of dis-

ruption is still visible. The posterior genital plate is entirely

absent, and the remaining four genitals are grouped in a roughly

semicircular order around the anterior edge of the periproct. The

madreporic genital is not much larger than the other three.

The oculars are small, and show no features of special importance.

From a broken and open condition such as that shown by Pygaster

sens. str., the processes of evolution work along two definite

directions. ‘The first aims at a restoration of a cyclic, or at least

of a compact, character in the system as a whole, and the second

is concerned with the infilling of the centre of the system (when

the cycle is regained) to replace the absent periproct.

In the reconstruction of the cycle of genital plates, the posterior

(fifth) genital is not necessarily resuscitated. In fact, a very

large number of the great groups of the Irregular Echinoids are

permanently without this plate. Anorthopygus shows the simplest

SEA-URCHINS. 479

condition of the cycle-restoration on this plan. In that genus the

postero-lateral genitals close in, so as to bring the system to an

F. Cenholectypus.

E. Pygaster (Megapygus).

H. Clypeaster.

(Reduced to a common size, but correct in proportion.)

D. Conulus.

G. Discoidea (the posterior genital pore not invariable).

C. Spatangus.

The apical system of some Holectypoids and their allies.

B. Anorthopygus

A. Pygaster sens. str.

approximately circular shape, and the place of the posterior plate

is taken by a prolongation of the madreporic genital through the

480 MR. HERBERT L. HAWKINS ON

system. This great increase in the size and extent of the right

anterior genital achieves two results. Firstly, the terior of the

apical system is filled by it (with the madreporite), and secondly,

the posterior margin of the system is completed by its extension.

The resulting structure is an apical system of the ethmolysian

type (see Gregory, 50). The great importance of this character

in Anorthopygus becomes evident when it is realized that such a

system is found only in it and in a section of the Spatangide. It

may be stated at once that the method of infilling of the centre

of the system shown in this genus is characteristic of all those

Holectypoida in which the apical plates regain a genuinely cyclic

arrangement.

The second method whereby the system is rendered compact,

without the redevelopment of the posterior genital plate, is shown

by Conulus. Here the two posterior oculars become greatly

increased in size, and meet along the posterior margin of the

system. The postero-lateral genitals undergo a similar transverse

extension to a greater or less degree, and meet above them. The

madreporic genital, although large and partly occupying the centre,

is in this way separated from the posterior region of the system.

A slight antero-posterior lengthening of the whole system usually

accompanies this method of development, and, in a simple sequence

indicated in my recent paper, the markedly elongate apical systems

of a Pyrina and a Holaster can be readily derived. ‘The Conulus-

plan is characteristic of the apical systems of many Jurassic

Nucleolitide, although it is not the only type developed in that

complex series of forms.

Of the type of apical system in which the fifth genital is re-

developed (or perhaps replaced by a new but similar plate),

Pygaster (Megapygus) shows the first stage. Here one small plate,

perhaps more, imperforate and in all probability flexibly united to

the others, makes its appearance at the adapical extremity of the

periproct. It seems probable that this new genital plate is a

specialized member of the anal series which has become in-

corporated into the apical system. In Megapygus it is always

small and imperforate. The next stage in recovery is seen in

Holectypus sens. str. In this genus the fifth genital is present as

a recognizable unit of the genital cycle. It is always smaller

than its four associates, however; and of these, the madreporic

genital is very large, occupying all the central part of the system.

The posterior cena is still imperforate. In the succeeding series

of forms (Cwnholectypus) the relations of the genital plates are

similar to those in the earlier subgenus ; but a genital pore,

quite as large as those of the other plates, passes through the

posterior genital. Canholectypus shows, then, the perfect re-

storation of the apical system. All five genital glands will have

been functional, each with a separate pore; while the centre

of the apical system is filled by the madreporite, situated, as

usual, entirely on the right anterior genital plate.

The apical system of Discoidea is particularly interesting. The

SEA-URCHINS. 481

fifth genital becomes practically indistinguishable, in point of size,

from the others of the cycle, even the right anterior plate

being much reduced from its condition in Holectypus. The pos-

terior plate may or may not be perforated, this irregularity

affording in itself ample proof of the plastic condition (in a

variational sense) of the genus. The madreporite, instead of

being restricted to the right anterior genital, is more or less

uniformly distributed over all five of these plates in some species,

a feature never found in the preceding genera. (In the case of

an otherwise abnormal Conulus albogalerus, a similar development

exists : see Hawkins, 70.) The oculars have dwindled considerably

in proportional size.

The chief interest of this peculiar structure is seen when a

comparison is made between the apical systems of Discoidea and

Clypeaster. In the latter genus the madreporite is central

and prominent, but it is quite impossible to distinguish the

sutures of the genital plates, at least in adult forms. The

oculars are minute. Discoidea, then, shows the preliminary

stages of the assimilation of the genitals—a phenomenon that is

preparatory to their coalescence and fusion in the Clypeastroida,

V. Tse INTERNAL EvoLution OF THE ORDER.

1, Features of Phylogenetic Importance.

In paleontological attempts to trace a phylogenetic sequence

through any series of organisms, the first and essential feature to

be considered is the order in time in which the various forms

appear. Most of the serious errors that have marred the value

of some past work in this direction have resulted from an insufficient

reliance on the stratigraphical relations of the genera considered.

It is true that our knowledge of the occurrence of fossils at various

horizons is very inadequate: it is only necessary to consider the

number of cases where a gap exists in the sequence of forms that

are known to occur in widely separated horizons, to realize this

incompleteness of our knowledge. But it seems a fair postulate

to assume that the order in which various genera make their

appearance is approximately the true sequence of their evolution.

Especially is this the case in the Holectypoida. Not only are

they, in common with most Echinoidea, eminently adapted for

preservation in suitable deposits, but the periods of their exist-

ence, the Jurassic and the Cretaceous, were times when, at least

in this country, the conditions of deposition were exceptionally

favourable for the preservation of organicremains. In the scheme

of evolution put forward below, no apparent relationship has been

accepted unless the stratigraphical evidence confirmed it.

Asecond great principle from which reliable evidence of genetic

affinity can be deduced is that of Ontogeny. Here, unfortunately,

our knowledge of the Holectypoida ismeagre. In the Echinoidea

generally the process of recapitulation is always very much obscured

Proc. Zoo. Soc.—1912, No. XXXI, 3l

489 MR. HERBERT L. HAWKINS ON

by the existence of a free-swimming larval stage. Of post-larval

changes in the Class but httle is known, Agassiz (80), in the

Revision, summarized the state of knowledge of the “young stages

of Echini,’ and but little has been added since that date, at least

in the case of the Irregularia. Ontogenetic characters are always

difficult to observe and to appreciate among fossil forms, and far

more zonal collecting of young stages of the Jurassic and Cretaceous

Echinoids will be necessary before this line of evidence can be used

for their correlation.

Some slight details are available at present, such as the Hemi-

pedina-phase of Pygaster senisulcatus and the young stages of

Conulus with an adapical periproct. (Valette, 69, has described

a young specimen of C. swhconicus in which the periproct is

already in the adult position, although the individual has a dia-

meter of only 10 mm.). Unlike the Mollusca and Brachiopoda,

the Echinoidea do not retain the first-formed portions of the test

‘throughout life; so that, although new parts are continually being

developed, the acceleration by which these new portions assume

adult characters almost nullifies any recapitulatory features they

may possess. In the matter of the interambulacral tuberculation,

which at first seems a promising structure for ontogenetic study,

this feature of acceleration renders the characters of the new plates

practically worthless.

In addition to their sequence in time, it is therefore necessary

to consider the adult characters of each genus separately. The

features of an adult are divisible into two kinds. The first group

is that of adaptation to circumstances ; and the characters due to

this tendency, though interesting from other standpoints, have

little phylogenetic meaning. The second group of characters are

those which are unaffected, or are not necessarily affected, by the

surroundings of the organism, and which must in consequence owe

any peculiarities they possess to the line of evolution of the group

to which the individual belongs. Such features, which include

atavistic and vestigial structures, are of first-rate importance for

showing the phylogeny of a group. In the Hchinoidea, the cha-

racters that would fall into the first category would be those

directly concerned with assimilation, respiration, reproduction,

and locomotion. The characters of the second type would consist

of apparently trifling variations in the ornament or structure of

the test—-variations of such a kind as not to affect the vital

processes to any serious degree, nor be affected by them. Such

characters are the details of the plating of the ambulacra and the

variations, within certain limits, in the structure of the apical

system. These two characters are regarded as essential indices

of relationship in the present paper.

There is, however, in the investigation of an extinct, annectant

group like the Holectypoida, an additional principle of evolution

that gives safe guidance. The two extremes of structure—those

of a Cidarid and of a Spatangid—are known. Generally speaking,

the Holectypoida should show a gradual tendency, in the course

SEA-URCHINS. 483

of their evolution, to depart from the characters of a Cidarid, and

to approximate to those of the Irregular types. A recognition of

this direction of evolution in the group renders the interpretation

of the various structures more intelligible by including them all

in one coherent scheme. <A complete. reliance on this principle

would probably result in a misinterpretation of degenerate or

retarded development, so that the trend of evolution must be

considered in direct connection with stratigraphical evidence.

To sum up, the characters used here as indices of phylogenetic

development are of two kinds. One series is available for tracing

the evolution of the group as a whole. Such features are (i.) the

gradual loss of masticatory structures and of peristomial branchie,

(ii.) the backward movement of the periproct, (i1.) the loss of

radial symmetry, and (iv.) the increase in density, and decrease

in coarseness, of the tuberculation. The other series is used to

indicate the intimate relations of the individual genera of the

group. These features are (i.) the plating structure of the am-

bulacral areas, (ii.) the composition of the apical system, and

(iii.) the stratigraphical sequence.

2. The Origin of the Group.

The oldest known member of the Holectypoida is Pygaster

reynesi, which occurs in the Middle Lias of France. It will

therefore be necessary to look for the ancestor of this typically

Holectypoid form among the Regular KEchinoids of the Liassic

Triassic periods. It is unfortunate that the origin of the

group should date from these periods, for, unlike the purer waters

of the Oolitic seas, the muddy shore-lines of the Liassic ocean,

and the saturated lagoons of the Triassic coral-reefs, were un-

favourable to the free development, as well as to the ultimate

preservation, of Echinoids. However, it is significant to find

that the earliest Irregular Echinoid appeared so soon after the

first stage of differentiation had begun among the Regular orders.

Its inception thus seems to have been an effect of that unrest in

structwre and habit that usually accompanies profound changes

in the course of the evolution of a Class.

The Liassic Regular Echinoidea seem to belong to two orders

only, the Cidaroida and the Diademoida, The former group had

become more or less stereotyped in character during the Permian

and Triassic periods, having been, as Bather (59) indicates, the

only surviving member of the varied Paleozoic types. The

Diademoida, as the same author has shown (¢. ¢c.), were beginning

to assume the typical features of the order in Triassic times, but

still retained features, such as a primary character of the am-

bulacral plates in the greater part of the area, anda shallowness

of the branchial clefts, which are reminiscent of their Cidaroid

ancestry.

From the Lias a considerable number of primitive Diademoida

are known, and they have been recently studied by Lambert (52),

31*

A84 MR. HERBERT L. HAWKINS ON

Tornquist (57), and Bather (59). A great part of their ambulacra

is still built of primaries, which show no signs of their subse-

quent modification except in the arrangement of the primary

tubercles, one to each group of three ambulacrals. A large

number of these Liassic forms are grouped under the generic

names of Diademopsis and Hemipedina. These genera and their

Diademoid allies have been so exhaustively studied by Bather

(é. c.) that no detailed discussion of their characters or affinities

is necessary here. One of the most obvious features which

separate these early Diademoids from their descendants is the

structure of the perignathic girdle. The processes, although well

developed, are rendered quite inconspicuous by the considerable

elevation of the ridges. The latter structures are, of course, a

relic of Cidarid characters. ‘This shows that the change from an

interradial to a radial position for the perignathic prominences

was a gradual one. The view that the increasing complexity of

ambulacral structure is connected with the growth of the peri-

gnathic processes, which hinder the passage of the ambulacrals on

to the peristomial membrane, is supported by Bather (é. c.) on

this evidence.

There are, then, two orders of Echinoidea from which, on

stratigraphical evidence alone, the Holectypoida may have been

evolved. Of these orders, the Cidaroida were well established,

with their special structures stereotyped, before there is any

evidence of the existence of Irregular Echinoids. This fact alone

would seem to render unlikely any hypothesis which regarded the

early Cidaride as directly ancestral to the Holectypoida,

When consideration is taken of the essential features of a

Pygaster, a notable correspondence between them and the

structures of the early Diademoida becomes apparent. The

ambulacra are chiefly composed of primaries (with a triple ar-

rangement of tubercles), and towards the peristome a partial

compression of the plates into triads is seen. Triad formation,

in the same part of the ambulacra, is characteristic of all the early

Diademoids, and is one of the diagnostic features of the whole

order. The perignathic girdle of Pygaster shows well-developed

processes, but hardly appreciable ridges. This character, the

absolute antithesis to that of the Cidar idee, is known to have been

gradually attained by the Diademoids thr ough their Triassic and

Liassic representatives. Again, the apical system of Hemipedina

often shows a prolongation backwards into the posterior inter-

ambulacrum.

Most significant of all is the indication of affinity between the

two orders by the slight ontogenetic evidence already available.

In discussing the affinities of Hemipedina boner, Wright (20)

admitted that he was uncertain as to the true generic relations

of the species. He was at one time inclined to class it with

Pygaster. Bather (59) has referred to this species, and is of the

opinion that A. bonei, if it is not a Hemipedina, should be asso-

ciated with Pygaster. The species is a small one, and the shape

SEA-URCHINS. 485

of the sear left by the apical system indicates a considerable back-

ward prolong: ition of that structure. J have before me a series

of ten specimens from the Pea Grit of Crickley Hill (near Chel-

tenham), which are presumably the young of Pygaster semisuleatus ;

but I am unable to find any satisfactory distinctions between

them and the type of H. bonei. If there is any appreciable

difference, it consists in the fact that. the periproct does not

project so far into the posterior interambulacrum in the Pygasters

as does the “scar of the apical disc” in the Hemipedina. It

seems hardly possible that, so early in the history of both orders,

heterogenetic homeomorphy could have reached such a degree of

perfection, and I ain therefore strongly of the opinion that

“ Hemipedina” bonet is a Pyyaster, and almost certainly a young

form of P. semisuleatus.

It thus seems established that Pygaster is intimately related to

some primitive, probably Liassic, Diademoid. It is impracticable,

in the present state of cur knowledge, to search for the actual

generic ancestor; but if the choice were to lie between Diade-

mopsis and Hemipedina, the former would seem to possess the

stronger claim to recognition. As defined by Lambert (52),

Diademopsis is distinguished from Hemipedina by the presence

of pronounced secondary tubercles in the interambulacra.

Bather (59) has shown that the distinction is not so absolute as

Lambert’s diagnosis would suggest, but the fact remains that,

among the earlier species of the genera, there is a more strongly

developed tendency to a multituberculate character in Lia-

demopsis. As Pygaster is also a multituberculate form, the

alliance with Diademopsis would seem natural, but 1 do not feel

justified in expressing a positive opinion on the matter, beyond

the statement that the immediate ancestor of the Holectypoida

must surely have been a Diademoid.

3. The Pygasteride and Conulide.

The three subgenera of Pygaster sens. lat. mark three stages

in the evolution of that genus. Pygaséer sens. str. is undoubtedly

the most primitive type. Megapygus shows an advance in two

directions. The periproct is undergoing a change of shape pre-

liminary to its actual separation from the apical system, and the

tuberculation is assuming slight irregularity of arrangement.

Both these features point towards “ Irregularity.” J/acropygus,

which appeared at about the same horizon as Megapygus, shows a

similar character in its periproct, but the tuberculation, instead

of becoming superficial and irregular, shows a deepening of the

scrobicules, “and a corr esponding reduction of the miliary surface,

The distinction from the Megapygus umbrella group is not very

great in appearance, but seems important in its results. I regard

the two subgenera as parallel lines springing from the common

ancestor Pygaster sens. str.

Pileus is undoubtedly a short-lived offshoot from the Pygaster-

486 MR. HERBERT L. HAWKINS ON

stock, for the anomalous biserial ambulacra are unlike any other

genera of the order. The periproct position shows an advance on

the Megapygus-condition, and, owing to certain irregularities of

the tuberculation, I am inclined to regard Pilews as a side-branch

of that line. Anorthopygus, which in the classification I have

associated with Pilews, seems to show a course of evolution

parallel with, although in many ways differing from, that of the

aberrant genus. The oblique position of the periproct does not

appear to be an important character, although peculiar. The

tuberculation is definitely like that of Macropygus in structure,

though not in arrangment, and therefore I haveregarded it as an

offshoot from that subgenus in Lower Cretaceous times, which

corresponded with the similar offshoot from the Megapygus-line

in the Upper Jurassic.

At about the same horizon in which Anorthopygus occurs,

Oonulus appears. The earlier species seem very difficult to dis-

tinguish from those of Pyrina with which they may be strati-

graphically associated. In the matter of the tuberculation the

adoral surface of Conaulus shows much the same characters as the

whole test of Anorthopygus. Moreover, the arrangement of the

tubercles is similar in both genera. The periproct has passed to

the posterior edge of the test, although in many young specimens

of C. subrotundus (some of which are almost globular), the aper-

ture is on the adoral surface quite near to the apex. ‘The feature

which marks off Conulus so sharply from the Pygasteridz is the

accelerated condition of the ambulacral plate-crushing. There

is no appreciable tendency to increase the number of demi-plates

in the Pygasteride, from the few adorally situated ones, which

were probably directly inherited from the Diademoid ancestor.

However, in many other features Conulus shows almost equal

acceleration. When the Upper Chalk is reached, the genus

disappears suddenly after a short existence, during which few

important specific modifications were evolved. Its relations to

the Pygasteride are not very easy to decide, but, on the character

of the tuberculation, | have connected it with the Anorthopygus-

line. An additional lnk between the genera is afforded by the

structure of the apical system, the fifth genital plate being per-

manently absent from both.

4. The Discoidiide.

Holectypus sens. str. appears in the Inferior Oolite in asyso-

ciation with Pygaster sens. str. It is only in the position of the

periproct that considerable acceleration is shown, but the dif-

ferentiation of the characters of the tubercles on the upper and

lower surfaces of the test is also a feature of advance. The

Holectypine are a perfectly homogeneous group, and must be

regarded as an unbroken series. Coptodiscus is apparently a

peculiarly specialized offshoot from Cemnholectypus, and the

suturing of the adapical surface may perhaps be ascribed to

SEA-URCHINS. 487

gerontic degeneration of armour (see Oswald, 61). Lanieria is

also allied to Cenholectypus, but, unlike most of the Holectypine,

is almost globular in shape. The position of Discholectypus 1s

more difficult to determine. In every obvious feature it is a true

Holectypine, but it shows an ambulacral structure closely resem-

bling that of Conulus. The absolute contrasts of tuberculation,

periproct-position, apical structure, and general form which

appear when Discholectypus and C ‘onulus are compared, preclude

any possibility of a genetic connection between the genera.

Discholectypus would seem, therefore, to be a branch of the

Holectypus-line, which developed complex ambulacral plating by

a process of acceleration, This parallelism of development

(heterogenetic homeeomorphy) of a feature in two distinct genera

is rendered particularly interesting by the correspondence in time

at which the specialization took place.

There can be no doubt as to the close relationship which exists

between Discoidea and Holectypus. On stratigraphical evidence,

and also because of the variable nature of the apical system (in

the matter of the perforation of the posterior genital plate), I

have considered the Discoidiine as descendants of Holectypus

sens. str., whose appearance coincided in time with the modi-

fication of the parent stock into Cenholectypus.

5. Summary of Internal Evolution.

The Holectypoida originated from a Diademoid ancestor in the

Triassic or early Liassic periods, and subsequently developed along

two definite lines. In one line (Pygasteride and Conulide) the

apical system never fully regained, and finally lost, the posterior

genital plate, while the whole system tended to become elongated ;

the tuberculation gradually became uniformly distributed over

the interambulacra, and irregular in its arrangement ; the shape

of the test showed various departures from radial symmetry ; and

the jaw-structures dwindled and ultimately almost disappeared in

adults. Inthe other line (the Discoidiide), the fifth genital plate

was early redeveloped, and later regained its function, while the

system as a whole became circular in shape; the tuberculation

retained its regularity of arrangement, but became insignificant

adapically and coarse adorally ; the shape of the test eventually

regained a radial symmetry ; and the jaws, though modified,

showed little or no decrease in power.

VI. Tue ExtTerNAL AFFINITIES OF THE ORDER.

The primitive character of the early Holectypoida (in an

Irregular sense) is so pronounced that it would naturally be

expected that the group existed for some time before any of the

more elaborate forms were evolved, and that these appeared at

subsequent intervals as offshoots from the Holectypoid stock.

Such, however, was not quite the case. The Holectypoids are

A488 MR. HERBERT L. HAWKINS ON

merely a retarded series of Irregular Echinoids, and some of the

orders of that subclass early became differentiated from the

Pygasteride by a relatively accelerated evolution. It is becoming

increasingly manifest that large groups of organisms, such as the

Irregular Kchinoids, are not often homogenetic in the strict sense

of the word. When a series of forms that have been regarded as

belonging to an individual genus can be shown (as Beecher and

others have proved for some Brachiopoda) to pass through widely

divergent lines of ontogenetic (and therefore phylogenetic)

development, the problem of the evolution of a class or subclass

must be considered more complex still. Indeed, at first sight, it

would seem that, without the evidence of Ontogeny, no reliable

clue to genetic relationship can be deduced from even the most

accurate correspondence of adult characters.

Stratigraphical paleontology, however, shows a kind of extended

ontogeny which, although fragmentary, is infallible so far as 16

can be understood. ‘The same phenomena which complicate the

study of recapitulation in recent species are as widely developed

among the families and orders of past periods. Acceleration and

retardation, adaptation and degeneration, tend to obscure the true

sequence of genetic affinity to such a degree that, in the present

state of knowledge, only the bare outlines of the evolution of the

larger groups can be indicated.

Tn this section of the paper, an attempt is made to show the

affinities (with persistent regard to stratigraphical relations)

which appear to link certain genera of the Holectypoida with

those of other orders. Little account is taken of the subsequent

changes which may have been developed in these other groups,

and no opinion is expressed as to their absolutely homogenetic

characters. The name of a fairly primitive member of each main

group is inserted in the diagram (text-fig. 60, p. 493) in its true

stratigraphical position, and by a thin vertical line each of these

names is connected with that of a characteristic genus now

living, which is usually regarded as belonging to the same group.

1. Pygaster and Galeropygus.

Galeropygus appears in the Upper Lias with at least two

species, one of which (G. dwmortieri Paris) is British, The genus

is thus contemporaneous with Pygaster sens. str. Gregory (50),

probably on account of its obviously primitive characters, included

it among his Pygasteridsx, although in almost every feature it

offers a violent contrast to the diagnosis of that family. Practi-

cally the only diagnostic character in which it resembles an

Holectypoid is the apetaloid nature of its ambulacra. A feature

which would tend to connect it with some of the later Pygasters

(e. g., Macropygus truncatus) is the shape of the test, which is

commonly rather broader than long. The deep anal sulcus finds

a shallow counterpart in the posterior interradius of Pygaster

semisulcatus, but 1 have indicated above (p. 465) that this sulcus

SEA-URCHINS. 489

is probably due to the presence of the periproct near the apex,

and has, in consequence, little direct phylogenetic meaning.

Galeropygus may be regarded as differing from Pygaster sens.

str, by a marked acceleration in the characters of its tuberculation

and peristome. The former feature is already in the uniform

condition, no definite order of appearance being traceable for

individual tubercle series. The peristome is quite small, and

slightly excentric anteriorly, with no visible adaptation for Jaws.

I regard the genusas a primitive member of the Vucleolites-group,

with all the characters of that group except the subpetaloid

ambulacra. As there is a marked tendency to develop this

feature even among the Holectypoida, it seems that its production

in the descendants of Galeropygus could be naturally postulated.

Owing to the stratigraphical appearance of Galeropygus, 1 should

consider it an offshoot from the Diademoida that hardly, if at all,

progressed along the Holectypoid line of descent before developing

striking acceleration in all its characters except the periproct and

the ambulacra. It is interesting to find that the position of the

periproct remained more or less constantly primitive in the

majority of the Jurassic descendants of Galeropygus (e. g., Hchino-

brissus and Clypeus), although the ambulacra early began to show

elaboration. In the periproct feature, indeed, the Pygasteride

show a greater acceleration than the Nucleolitide, although the

Holectypoida are, in most characters, a retarded group.

Even if the affinity between Galeropygus and Pygaster were to

be proved to be less close than I have indicated in the diagram,

the characters of the ambulacral plating would show that it was

derived, directly or indirectly, from a Diademoid ancestor. As

I interpret the relations of the genera at present, Galeropygus and

Pygaster stand together at the root of all the Irregular Echinoids,

in structure as well as in stratigraphical position. The subsequent

modifications of the Galeropygus stock I have briefly outlined in

a recent paper (Hawkins, 66), and I hope to amplify that re-

adjustment of the classification of the ‘“ Cassidulide” at some

future time.

2. Pygaster, Conulus, and the Echinoneide.

Since its first recognition by Desmoulins (4), the genus Pyrina

has been the occasion of great confusion. The extraordinary

similarities that appear when it is compared with Conulus make

the generic position of species ascribed to them more difficult to

determine than their specific distinctions. Such a form as

P, desmoulinsi, with its elongated ovoid ambitus, is easily dis-

tinguishable from a Conulus, the species of which are almost,

though rarely quite, circular in outline. To restrict the genus

Pyrina to such elongated forms would, however, result in a very

unnatural grouping of the species, and, unless details of the

anatomy can be traced, the distinction of a roughly circular

Pyrina from a Conulus becomes almost impossible. Theoretically

490 MR. HERBERT L. HAWKINS ON

a Conulus should possess vestiges of jaws, and a peculiar type of

perignathic girdle; but jaws are very rarely preserved in fossil

Kehinoids, and Pyrina has a somewhat similar series of structures

around the peristome. The similarities between the two genera

include the shape (generally), the oblique peristome, the position

and shape of the periproct, the structure and arrangement of

the tubercles, the ambulacral plating, and the composition of the

apical system. Added to these there would probably be the

presence of vestigial jaws in young individuals of Pyrina, since

these structures have been found in a small specimen of the more

highly specialized Hchinonews. An additional difficulty in the

separation of species belonging to the two genera results from the

fact that both were evolved at about the same time, and flourished

side by side during the Cretaceous period.

So many correspondences in important structures cannot point

otherwise than to a close genetic affinity between Pyrina and

Conulus, and the only feature that can be considered to exclude

the former genus from the Holectypoida is the absence of jaws in

the aduit state. The presence of these organs in young specimens

cannot be considered sufficient evidence for the inclusion of

Pyrina in the order; for, if vestigial characters are taken into

account in classification, by analogy the Mammalia, by reason of

their embryonic gills, would have to be classed with the Pisces.

The earliest members of the Echinoneide, such as Vucleopygus,

have the periproct in a supra-marginal position similar to that of

Anorthopygus. As Lovén (36) has shown, a gradual migration of

the periproct takes place in this family along an exactly parallel

line to that passed through in the Holectypoida, until in Hehinoneus

the anus is in a position similar to that of Discoidea. The features

of Desorella are so little known that it is unsafe to ascribe

a definite systematic position to it, and it has been ignored for the

purposes of the present work. I have regarded the Echinoneide

as offshoots from Pygaster (Macropygus) in Upper Jurassic times,

which for some distance followed the Anorthopygus branch, and

left it simultaneously with the Conulide. They were at first

distinguished from that family by the accelerated degeneration

of their jaw-structures.

3. Anorthopygus and the Spatangide.

The earliest members of the Spatangide proper appear in the

Lower Cretaceous, They are not very clearly distinguishable

from some other groups, especially from the Echinocorythide.

The structure of the apical system is, however, different in these

two families. The system of the Echinocorythide is elongate, and

has been compared with that of the Collyritide. (I have recently

shown (Hawkins, 70) that this structure could easily be evolved

from the Conulid type by acceleration.) The Spatangide have

a compact apical system, which often nearly resembles that of the

Conulide, but is sometimes ethmolysian—that is, with the

SEA-URCHINS. 491

madreporic genital extending right through the system to occupy

some part of the posterior border. In every case the fifth

genital is absent, and this feature alone serves to distinguish

a Spatangid from the great majority of the “ Cassidulide.” At

first sight there do not seem to be many points of resemblance

between Anorthopygus and the Spatangidee. However, the

stratigraphical appearance of the two types is the same, and,

in the structure of the apical system, Anorthopygus shows an

ethmolysian character in both the known species. No types of

Kehinoids other than Anorthopygus and some Spatangids have

this feature. The position of the periproct in the Holectypoid

genus is about midway between the apex and the ambitus, and

the same character holds in almost all the Spatangide. In

several small specimens of A. orbicularis that 1 have seen there

is an appreciable increase in the declivity of the test behind the

periproct, and I regard this as a rudimentary posterior surface.

The plating of the periproct-membrane also shows some similarity

in the two groups.

The evidence for their genetic affinity is very slight, but I have

ventured to connect the Spatangide with the Anorthopygus line

of descent on account of the apical structure and stratigraphical

correspondence.

4. Conulus and Amblypyygus.

Amblypygus is a Tertiary Echinolampid (Duncan (44) classed it

as an Hchinoneid) whose characters are best known from the

descriptions of Indian species given by Duncan and Sladen (89).

The shape of the test, the obliquity of the peristome, and more

especially the structure of the ambulacra, all show features of

similarity with those of the Conulide. The ambulacra are sub-

petaloid adapically in Amblypygus (a marked contrast to all the

Kchinoneide), but in that region, as much as on the adoral

surface, the inclusion of one demiplate between two primaries

is regularly shown. It is difficult to imagine that this peculiar

structure could be evolved independently in heterogenetic

genera. The only other form with which Amblypygus could

be associated by reason of this structure is Discholectypus, but,

apart from the contrasts which the two genera show in other

respects, the stratigraphical sequence is not favourable. As

Amblypygus is a very primitive type, and one of the oldest

known genera, of the Echinolampide, I have separated that

family from the other “ Cassidulide,” and derived it from the

Conulid stock. The later members of the family seem to have

reduced their ambulacral structure to a condition of simple

primaries ; a process that, outside the order of the Holectypoida,

seems to have been the usual one adopted.

5. Conulus and Conulopsis.

A group of Upper Cretaceous Echinoids, which was formerly

classed with “ Galerites,” was separated from that genus by

492, MR. HERBERT L. HAWKINS ON

Desor (21) under the name of “ Hehinoconus.” ‘‘Galerites”

roemert VOrb., on which the genus was founded, seems to be

congeneric with the “ Hchinoconus abbreviatus” and “‘ orbignyanus”

of the Upper Chalk of Norfolk. The differences between these

species and a typical Conulus are manifold, and, as the name

Kchinoconus cannot be retained for them, I have distinguished

them as Conulopsis. The tuberculation of Conulopsis is irregular,

and the tubercles of the adapical surface have deeply sunken

scrobicules. ‘The ambulacra are composed of primaries through-

out, the adapical pore-pairs being almost subpetaloid; while

round the peristome the interambulacra are raised into definite

‘‘bourrelets.” The general facies of Conulopsis is similar to

that of Caratonus (the latest discussion of this genus being by

Schlueter, 54). It is possible, however, that some real genetic

relation may exist between the later Conuli and Conulopsis, and

that the resemblance of the latter genus to Caratomus may be

deceptive. Even if Conulopsis is a descendant of Conulus, it

is certainly not an Holectypoid. It would show a development

which would have a peculiar interest when compared with the

development of Conoclypeus from Discoidea. The same loss of

regularity in the tuberculation is seen, and the ambulacral plates

have become restored to their primary state. (The large polygonal

ambulacrals of the adoral surface of Conulopsis ave strikingly

similar to those of a Clypeaster or of a Spatangid.) The develop-

ment of a subpetaloid character in the adapical parts of the

ambulacra would be comparable in the two genera, while a

similar correspondence is shown in the peristomial ‘“ bourrelets.”

Only the position of the periproct (almost marginal in Conulopsis),

and the presence of strong jaws in Conoclypeus, would tend

to separate the two genera. These last features would be

definitely due to the characters of the ancestor, Conwlopsis

agreeing in them with Conulus, and Conoclypeus with Discoidea.

Connlonsis and Conoclypeus would then mark parallel accelera-

tions from different branches of the Holectypoid stock. At

present, however, I do not feel satisfied that the genetic connection

between Conulus and Conulopsis exists, but I have connected the

two by a broken line in the table.

6. Discoidea and the Clypeastroida.

The similarities of structure that link the Discoidiine with

the Clypeastroida are many and of fundamental importance. A

circular outline ; an invaginated peristome; an infra-marginal

periproct ; a madreporite scattered over five genitals, all of which

may be perforated by a genital pore; and internal buttresses to

the test: are common to most genera of the Clypeastroids, and

are diagnostic features in Discoidea. The jaws in this genus

are strong, in view of its late appearance among the Holectypoida,

and, although conforming more to the ‘ Regular” than to the

Clypeastroid type, may well have assumed a more expanded

SEA-URCHINS. 493

shape when the height of the test diminished. The ambulacr

in all Clypeastroids are either petaloid or subpetaloid, but Cono-

elypeus, from the Upper Cretaceous, serves to link the simple

Text-fig. 60.

w us

2 w S <x 2 S

ra) s a S = fr

= es eee z = E

3 S s 3

a < So =

3 k 5 a >

S) — Zi

2 ra S rs) ee Seow

ECHINOLAMPAS ECHINOC YAMUS

NUCLEOLITES ECHINONLUS CLYPEASTER

SPATANGUS

AMBLY PYCU : 2

PYGURUS

aos Ob.

DISCHOLECTYPUS

————

ANORTHO MACROPYGUS

eae COLNHOLECT YPUS

HOLECTYPUS

am wu

ia,

"3 _[IECHINOBRISsU.

(Se)

4 0

CLYPEUS

Vf bees

Be OR GALEROPYCUS PYGASTER

pe ais

uw Jp}

3 i< DIADEMOID ANCESTOR (#DIADEMOPS!S)

Si)

Phylogenetic table of the Holectypoida and their allies. Names of Holectypoid

genera are connected by thick lines, those of other orders by thin lines. No

details are inserted in groups other than the Holectypoida, the names in the

external groups being those of the earliest-known genus that has been

satisfactorily described. Short, thick, horizontal lines above names indicate

that the genera became extinct at that horizon, and left no direct descendants.

ambulacral pores of Discoidea with the more elaborate structures

of the later genera. Perhaps, if the jJaw-structures of Cono-

clypeus were to be discovered in a more perfect condition than

494 MR. HERBERT L. HAWKINS ON

those described by de Loriol (35), they also would show an

intermediate character. The presence of ‘“ bourrelets” round

the peristome in this genus are the only features that seem

antagonistic to its being regarded as ancestral to the Clypeuster-

series.

The similarity between the small species of Discoidea and

Echinocyamus caused Gregory (50), when revising the unnecessary

generic division of “ Hcehinites” made by Dunean (44), to propose

the name Protocyamus. The name is inadmissible on systematic

grounds, but would be morphologically appropriate. Hchino-

cyamus occurs first in the Upper Cretaceous, and has developed

but few changes in structure from that time to the present day.

H. L. Clark (64) has recently suggested that the characters of

Echinocyamus are not primitive, but rather degenerate. On the

stratigraphical evidence I incline to regard them as truly

primitive, and to have retained ancestral traits by the retardation

of development consequent on their small size.

Echinocyamus (of the Fibulariide) was then directly evolved

from the smaller (typical) Discoideew, while Conoclypeus (of the

Clypeastride) appeared at the same period as a descendant of

(probably) the larger species of Discoidea (the “ Pithodia” of

Pomel, 37). The former group underwent little change in

subsequent periods, but the latter became rapidly differentiated

into the numerous and complex types that characterize the

other families of the Clypeastroida.

VII. Summary.

The Holectypoida are restored to the rank of an order of

the Echinoidea Irregularia. A classification, somewhat modified

from that proposed by Gregory (50), is given, and revised

diagnoses of the families, subfamilies, and genera are drawn up.

A comparative study of the morphology of the skeletal structures

of typical genera of the group is given; and, in the light of the

results of this study, the course of evolution both within and

beyond the limits of the order is indicated.

The Holectypoida are regarded as an annectant group of the

Trregular Echinoids, whose characters retain a considerable

uniformity owing to a persistent retardation of evolution. At

various periods offshoots from the Holectypoid stock appeared,

which, usually with a relatively accelerated differentiation,

developed into the various orders and families of the Irregularia.

The order commenced in the Liassic period, and became extinct

at the end of the Cretaceous. Two of the groups of Echinoids

now living retain many features that were characteristic of the

Holectypoida (the Echinoneide and the Fibulariide), The other

groups of Irregular HEchinoids show a much greater departure

from the primitive character, but they all possess some features

which indicate their Holectypoid ancestry.

SEA-URCHINS. 495

Two new names are introduced in the Systematic Part :—

Megapygus as a subgenus of Pygaster, corresponding with the

Pygaster (sens. sty.) of Pomel (37), with type WM. umbrella; and

Conulopsis, a genus including the “ Hchinoconus ” of Desor (21),

with type C. roemeri d’Orbigny. The latter group will be

studied in greater detail in a forthcoming paper.

VILI. List oF LireRATURE CONSULTED.

(The numbers of pages and plates in square brackets are those

which have a special bearing on the subject. Where no such

brackets are present, the whole paper has been consulted.)

1. 1734. J. T. Kuery.—Naturalis dispositio Echinodermatum ete. [p. 24;

Pls. xiii—xiv. |

2. 1824. C. Sroxes.—Extracts from a letter explanatory of three drawings of

Echini. ‘Trans. Geol. Soc., ser. 11, vol. 11, p. 406.

3. 1829. J. Puitires.—lIllustrations of the Geology of Yorkshire, ete. Part I:

[p. 127; Pl. iii. fig. 17.] e

4. 1835. C. DesmouLrns.—Etudes sur les Echinides. II. Généralités. Actes

Soc. Linn., Bordeaux, Tome vii, pp. 315-432.

5. 1836. E. Grarerour.—Mémoire de Géo-zoologie sur Jes Oursins fossiles

(Echinides) qui se rencontrent dans les Terrains caleaires des environs

de Dax. Actes Soc. Linn., Bordeaux, Tome vill, pp. 108-191 ;

Pls. i. & ii. [pp. 140-160; PI. i1.}

6. 1837. L. Agass1z.—Prodrome dune Monographie des Radiaires ou Echino-

dermes. Mém. Soc. Sci. Nat., Neufchatel, Tome i. (Also in) Ann.

Sci. Nat., Tome vii (and in) Annals Nat. Hist., vol i. [p. 302. |

7. 1837. C. Desmoutins.—Etudes sur les Echinides. III. Synonymie générale.

Actes Soc. Linn., Bordeaux, Tome ix, pp. 45-364. [pp. 92-105 ;

200-363. | ,

8. 1839. L. Acasstz.—Description des Fichinodermes fossiles de la Suisse.

Premiere partie, Spatangoides et Clypéastroides. [pp. 61-94;

Pls. vi. ; xxii bis. ]

9. 1840. L. Acassrz.—Catalogus systematicus _Ectyporum Echinodermatum

fossilium Musei Neocomensis ete. [pp. 3-7. |

10. 1841. L. AGassiz.—Monographies @Echinodermes, vivans et fossiles. Echin-

‘tes. Famille des Clypéastroides. Mon. II. Des Scutelles.

1812. E. Desor.—ld., Mon. III. Des Galérites.

12. 1847. L. Acassiz & EK. Drsor.—Catalogue raisonnée des familles, des genres,

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13. 1849. E. Forpes.—Mem. Geol. Surv., United Kingdom. Figures and

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14. 1850. KF. eT cit., Decade IIT. [ Pls. vi-viii. |

15. 1852. T. Wr1ent.—On the Cassidulidse of the Oolites, with descriptions of

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[pp. 329- 332. |

19. 1856. J. W. Sarrer.—Mem. Geol. Surv., United Kingdom. Figs. and Desc.,

Brit. Org. Rem., Decade V._ [ Pls. vii & viil. |

90. 1856. T. Wricur.—Monogr. Brit. Foss. Echinod., Ool. Form., Part II.

Paiwontogr. Soc. [pp. 258-290 ; Pls. xvili-xx. |

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496

22.

23.

34.

35.

36.

37.

38.

39.

40.

41.

42.

43.

44.

45.

46.

47.

48.

1857.

1859.

1860.

1861.

1889.

1891.

1892. §

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MR. HERBERT L. HAWKINS ON

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G. Corrrau & —. TrigeR.—Hchinides du département de la Sarthe,

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A. p’OrsiaNny [posthumous].-—Paléontologie francaise. Terrains

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Part vi. Paleontogr. Soc. [pp, 198-224; Pls. xlvi-li.]

Description des Kchinides Tertiaires de la Suisse.

(Echinologie Helvétique, Partie iii.) Mém. Soc. Pal. Suisse,

Tome ii. [np. 80.]

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P. pz Lortot.—Monographie des Echinides contenus dans les couches

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from the Khirthar Series of Nummulitic strata in Western Sind.

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_ P.M. Duncan.—On Galerites albogalerus Lamarck, syn. Echinoconus

. P.M. Duncan.—On the Perignathic Girdle of the Echinoidea. Journ.

Tinn. Soc., Zool., vol. xix, pp. 179-212; Pls. xxx—xxxi.

_ P.M. Duncan & W. P. Stapen.—On the Anatomy of the Perignathic

Girdle and of other parts of the test of Discoidea cylindrica. Loc.

cit., vol. xx, pp. 48-61.

_ G. Corrgau.—Paléont. frang., Terr. tert., Tome i, Echinides Kocénes.

Tp. 452. |

_S. Loviy.—On a recent form of the Echinoconide. Bihang K.

Svensk. Vet.-Akad. Handl., Bd. xiii, Afd.iv, No. 10.

_ P.M. Duncan.—Revision of the Genera and great Groups of the

Echinoidea. Journ. Linn. Soc., Zool., vol. xxii, pp. 1-311.

[pp. 25 ; 135-173.]

_ P.M. Duncan & W. P. Stapen.—Note on the Perignathic Girdle of

Discoidea cylindrica. Ann. Mag. Nat. Hist., ser. vi, vol. iv,

pp. 2384-239. ys

V. Gaururer.—Description des Echinides fossiles recueillis en 1885

et 1886 dans le région sud les Hauts-Plateaux de la Tunisie par

M. Phillipe Thomas. Exploration scientifique de la Tunisie.

G. Corrrav.—Paléont. franc., Terr. tert., Tome ii. [pp. 190-898 ;

Pls. 249-259. |

‘ Lovin.—Echinologica. Bihang K. Svensk. Vet.-Akad. Handl., Bd.

xviil, Afd, iv, No. 1.

49.

50.

51.

52.

53.

54.

55.

56.

57.

58.

59.

60.

61.

62.

63.

64.

65.

66.

67.

68.

69.

70.

Proc. Zoou. Soc.

SEA-URCHINS. 497

1895. G. Corrrau & V. GAurutrr.—Mission scientifique en Perse, par

J.de Morgan. ‘ome iii, Partie ii, Paléontologie. Pt. 1, Echinides

fossiles. [p. 76; Pl. xii.]

1900. J. W. Grecory.—(in) Lankester’s Treatise on Zoology. Vol. iii,

Echinoderma, Chap. xv, Echinoidea. [pp. 315-321: 328.]

1900, E. Hesse.—Die Mikrostructur der fossilen Echinoideenstacheln und

derer systematische Bedeutung. N. Jahrb. f. Min., Stuttgart,

Beilage-Band xiii, pp. 185-264; Taf. xii & xiii.

1900. J. Lamperr.—Etude sur quelques Echinides de !’Infra-Lias et du Lias.

Bull. Soc. Sci. hist. nat., Yonne. Semestre I, Partie 2, pp. 3-57;

Pl. i, Tab. A & B.

1900, W. P. SraprEn.—(in) Zittel-Eastman, Text Book of Paleontology,

Vol. I. (Echinoidea). [ pp. 233-239. |

1902. C. ScurvETER.—Zur Gattung Caratomus. Zs. D. Geol. Gesell., Bd. 54,

pp. 302-335 ; Pls. xi & xii.

1905. F. A. Batrwer.—The Echinoid name Discoidea subucula. Ann. Mag.

Nat. Hist., ser. vii, vol. xv, pp. 145-148,

1908. E, 'T. Parts.—Notes on some Echinoids from the Lias of Worcestershire,

Gloucestershire, and Somerset. Proc. Cottesw. Nat. F. Cl., vol. xvi,

part 2, pp. 143-150. [p. 149.]

1908-9. A. TorNquist.—Die Diadematoiden des wurttembergischen Lias.

Zs. D. Geol. Gesell., Bd. 60, pp. 378-384; Pls. xv—xix (1908) ;

pp. 385-430 (1909).

1909. A. AGAss1z.—On the existence of Teeth and of a Lantern in the genus

Echinoneus Van Phels. Amer. J. Sci., ser. iv, vol. xxviii,

pp. 490-492 ; PI. ii.

1909, F. A. Barner.—Triassic Echinoderms of Bakony. Result. Wissensch.

Erforsch. des Balatonsees. Bd. I, Teil i, Pal. Anhang. [pp. 108-

113. |

1909. H. L. Hawkrns.—On the Jaw-Apparatus of Discoidea cylindrica

(Lamarck). Geol. Mag., Dee. v, vol. vi, pp. 148-152; Pl. vi.

1909. F. Oswatp.—The Degeneration of Armour in Animals. Science

Progress, No. xiii, pp. 123-134.

1910. H. L. Hawkrns.—Some Ambulacral Structures in the Holectypoida.

Geol. Mag., Dee. v, vol. vii, No. 554, pp. 349-353.

1910. W. LorscuER.—Die westfalischen Galeritenschichten mit besonderer

Beriicksichtigung ihrer Seeigelfauna. N. Jahrb. f. Min., Beilage-

Band xxx, pp. 269-312.

1911. H. lL. CLarx.—The genera of Recent Clypeastroids. Ann. Mag. Nat.

Hist., ser. viii, vol. vii, pp. 593-605. [p. 601. ]

1911. H. L. Hawx1ns.—On the Teeth and Buccal Structures in the genus

Conulus Leske. Geol. Mag., Dec. v, vol. viii, pp. 70-74; PI. iii.

1911. H. lL. HawxKrys.—On the Structure and Evolution of the Phyllodes

in some fossil Echinoidea. Tom. cit., pp. 257-265; PI. xiii.

1911. H. L. HawxKrys.—On the Tuberculation of the Holectypoida. Tom. cit.,

pp. 442-454,

1911. Fr. Kurincuarpr.—Ueber die Organisation und Stammesgeschichte

einiger Irregularer Seeigel der Oberer Kreide. Jena. Pp. 1-27;

Pls. i-xili. ¥

1911. A. Vanerre.—Description de quelques Echinides nouveaux de la

Craie. (Supplement.) Bul. Soc. Sei. hist. nat., Yonne. Semestre IT

(1910) pp. 121-151. [pp. 147-149.]

1912. H. L. Hawkrns.—On the Evolution of the Apical System jn the

Holectypoida. Geol. Mag., Dec. v, vol. ix.

1912, No. XXXII. 32

498 MR. OLDFIELD THOMAS ON A

EXHIBITIONS AND NOTICES.

March 19, 1912.

. KF. Harmer, Esq., M.A., Sc.D., F.R.S.,

Vice-President, in the Chair.

Mr. A. RapctyrreE Duemore, F.R.G.S., gave a lantern exhi-

bition of a large number of photographic studies of wild animals

in British East Africa and Newfoundland. This ver y beautiful

series of pictures was briefly described by Mr. Dugmore, who

dwelt particularly on the advantage to be gained by hunting with

the camera, and showed that it was a sport as well as a means of

securing interesting and valuable material.

Messrs. EK. Gurrarp & Sons sent for exhibition :-—

(a) A very fine head of a cow Sable Antelope (Hippotraqus

niger), shot by Major EK. C. F. Garraway, C.M.G., which was of

interest in that the mane was white instead of the normal dark

brown, and in the horns also being near the record for length.

(5) The skull of an jlealve line Bear (Ursus isabellinus) with

eight well-marked incisors in the lower mandible instead of the

normal six.

nasals and premaxillary bones which had arrested the shedding

of the milk-canines and the growth of the permanent canines.

The skull was quite adult and the remaining teeth were fully

developed and normal.

(¢) The skull of a Lion (Felis leo) showing a severe injury or

disease of the left side of the palate and the upper mandible,

which had not affected the growth of the teeth.

Two new Genera and a new Species of Viverrine Carnivora *.

My. OuprieLD THomas, F.R.S., F.Z.S., exhibited an example

of a Viverrine Carnivore from Tonkin which had a striking

external resemblance to the common Hemigalus t hardwickei, but

which differed so much in the skull and dentition as to necessitate

its being considered a distinct genus. Reason was also given for

regarding the Bornean “ Hemigale” hosei as representing a third

genus of the group.

The following was Mr. Thomas’s account of the animals re-

ferred to :—

“Commencing with the known species, from Borneo, I propose

* |!The complete account of these new genera and the species is given | here, but

since the names and preliminary diagnoses were given in the ‘ Abstract,

(distinguished by being underlined.__Eprror. |

y Better, but erroneously, known as Hemigale.

they are

NEW VIVERRINE CARNIVORE. 499

to found the following genus for ‘ /emigale’ hosei, which 1

have long realized should be separated from the banded animals

forming the true genus /emigalus.

“¢ DIPLOGALE,

“Thomas, Abstract P. Z. 8. 1912, p. 18 (March 26).

* Pattern of coloration normal, not banded.

“Skull with pavallel-sided, square-fronted muzzle, the tooth-

rows nearly parallel. Septum between anterior palatine foramina

without mesial foramen. Meatus and bulla about as in

Hemigalus.

“Teeth :—Upper incisors forming a straight transverse line, the

outer pair scarcely set back behind the level. of the others. Middle

teeth (p*, m1?) of the cheek-tooth series proportionally small,

terminal ones (p', m°) comparatively large. Pb! as long antero-

posteriorly as p*, with two roots and both anterior and posterior

supplementary cusps. P* also with marked supplementary cusps,

and a postero-internal basal cusp, this being quite absent in

Hemigalus.

“Type, Diplogale hosei (Hemigale hosei Thos.). Animal and

skull described and figured P. Z. 8S. 1892, p. 222, pls. xviii. & xix.

“The different shape of the muzzle and palate, the double-rooted

p' and other differences in the teeth, and the essential difference in

the pattern of coloration are the chief reasons for distinguishing

this genus from //emigalus,

** CHROTOGALE.

“Thomas, Abstract P. Z. 8S. 1912, p. 17 (March 26).

“ Pattern of coloration as in Hemigalus, not as in Diplogale.

* Skull—at least in the young—shaped more or Jess as in

Eupleres, the muzzle long and slender and the premaxille drawn

out forwards, with an abnormally long gap between the last

incisor and the canine. Median septal foramen between anterior

palatine foramina excessively long, as broad as either of the lateral

foramina, and nearly twice their length. Auditory region with

the bulle small and narrow, and the meatus very large and open,

its diameter approximately equal to that of a cross-section of the

bulla itself.

‘Teeth (of a young specimen with the milk-teeth still in place),

—Upper incisors quite unique in shape among Carnivora, broad,

flattened, spatulate, with a straight cutting- ‘edge, the edges of

the six combined forming nearly a_half-circle, owing to the

position of i° partly behind the level of i', and of i° again quite

behind it—the general outline of the incisive dentition recalling

more that of some of the smaller Kangaroos than that of a

Carnivore. P! small, single-rooted. Milk-premolars 2, 3, and 4

small and delicate, much smaller than the corresponding teeth of

Hemigalus and Diplogale, the proportions more as in Lupleres.

Lower incisors again broadened terminally, spatulate, the third

with a broader blade than the other two. .

500 MR. OLDFIELD THOMAS ON A

*¢ Type -—

‘‘CHROTOGALE OWSToNI. (Text-figs. 61-63.)

“Thomas, Abstract P. Z. 8S. 1912, p. 17 (March 26).

‘Colour and other external characters as in Hemigalus hard-

wickei, except for the differences indicated in the following

description :—

“Size apparently about as in H. hardwickei, but the only

specimen immature. Fur longer and less sleek than in that

animal, the longer hairs of the back, in a half-grown specimen,

about 19 mm. in length. Fur of upper side of neck all smoothly

directed backwards, like that of the back, instead of being sloped

forwards from a whorl on the withers.

Text-fig. 61.

Chrotogale owstoni.

Upper view of skull. The opening in the frontal region is probably of

parasitic origin.

‘General colour-pattern in most respects as in A, hardwickei,

but the dark dorsal bands are continued rather lower down on

the sides of the belly, where also there are a certain number of

NEW VIVERRINE CARNIVORE, 501

small irregular dark spots; on the neck the two dark bands are

broad and sharply defined, and outside them, on the sides and

under surface of the neck, there are similarly a number of blackish

spots, such dark markings in this region being quite absent in

H. hardwickei. Light ground-colour of upper part lighter than

in H. hardwickei, whitish instead of buffy. Under surface pale

buffy. Limbs greyish white, the digits scarcely darker. Tail as

in H. hardwickei, except that the terminal dark-coloured half is

more bushy, the hairs attaining a length of over 30 mm,

Text-fig. 62,

TERAI~

Chrotogale owstont.

A. Palatal aspect of skull. | 3B. Anterior part of muzzle, enlarged.

“Skull and teeth as described above, the description being

based on a specimen with milk-dentition. Adult dentition un-

known.

“ Dimensions of the type, measured on the skin:—Head and

body (c.) 400 mm. ; tail 260 ; hind foot 70.

“Skull: greatest length 91 mm. ; condylo-basal length 85 ; zygo-

matic breadth 39:5; nasals 22; interorbital breadth 15; breadth

502 ON A NEW VIVERRINE CARNIVORE.

of brain-case 33°7; palatal length 46°5; breadth between outer

corners of mp* 21°73; palatal foramina 7-2; median septal

foramen 11-6; horizontal length of mi' 2, mi? 3, mi® 3°3, mp? 4-6,

mp* 5:4, mp* 3°9.

Text-fig. 63.

Chrotogale owstoni.

A. Side view of skull. | B. Front view of muzzle.

“ Hab. Yen-bay, on the Song-koi River, Tonkin.

“ Type. Young male. B.M. No. 12.4.21.3. Original number 11.

Collected 16 September, 1911, by Mr. Alan Owston’s collector

Oru.

“This remarkable animal would at first sight be regarded as

merely a local representative of the well-known Banded Civet

(Hemigalus hardwickei), but a study of its skull and dentition,

although only at present represented by a young specimen, shows

that it is a most peculiar form, parallel in some respects with the

anomalous genus Lupleres, while in others—notably in its almost

MR. GUY DOLLMAN ON A SNUB-NOSED MONKEY, 503

kangaroo-like incisors—itis quite unique among Carnivora. And,

again, granting its essential distinctness from //emigalus, its

really striking external resemblance to that animal is not the

least of its many peculiarities,

“Tt is to be hoped that adult specimens may soon be obtained,

so that we may learn the characters of the permanent dentition.

Observations on its habits and food will also be of the greatest

interest.

‘“‘T have named the species in honour of Mr. Alan Owston, of

Yokohama, to whose enterprise in sending an expedition to

Yunnan its discovery is due.”

A new Snub-nosed Monkey *.

Mr. Guy Dotuman exhibited a new species of Snub-nosed

Monkey, allied to Rhinopithecus bieti M.-Kdw., collected at Yen-

hay on the Song-koi River, Tonkin. He proposed to call this new

form Rhinopithecus avunculus, and described it as follows :—

RHINOPITHECUS AVUNCULUS.

Dollman, Abstract P.Z.S. 1912, p. 18 (March 26).

Size smaller than in Rhinopithecus bieti M.-Edw., with dorsal

surface of body black and under parts yellowish buff.

Size and general proportions of body much less than in diet? ;

tail considerably longer. Hair shorter than in any of the other

forms, the general appearance more that of a true Presbytis than

a Rhinopithecus. No well-marked crest on crown of head, the

hairs in this region only slightly longer than on the neck; in

bieti a conspicuous crest is present in both sexes. Ears not con-

cealed by dense tufts of hair, rather more marked than in the

Mekong species. General colour of dorsal surface deep black,

tinged with brownish on the crown and nape; the black color-

ation extends all over the dorsal surface, both of body and limbs.

Sides of face and forehead creamy-white tinted with buff, the

colour gradually darkening on the back of the forehead and

merging into the dark blackish brown of the crown. Supra-

orbital line of stiff black hairs well developed. Face around eyes

naked and apparently flesh-coloured ; a crescent-shaped row of

ereamy-white hairs extending from the temporal region on to and

across the cheek. Sides of neck orange-buff, the bright buft-

coloured hairs forming a most conspicuous border to the dark-

coloured nape. Ears with creamy-white tufts arising from their

inner sides. Rump with two prominent buffy-white patches on

either side of the tail, connecting with the light-coloured markings

on the backs of the thighs, but not forming the large conspicuous

light-coloured areas such as occur in biet?. Backs of hands and

feet covered with rather short black hairs, the hair not sufticiently

* |The complete account of this new species appears here, but since the name and

a preliminary diagnosis were published in the ‘Abstract’ it is distinguished by

being underlined.—Eprror.

504 MR. GUY DOLLMAN ON A SNUB-NOSED MONKEY.

long to conceal the digits nor extending over the nails. Entire

ventral surface of body yellowish buff, the buff tint most domi-

nant on the throat and anterior portion of the belly ; no trace of

any grey or dark-coloured hairs on the chest and neck. Inner

sides of arms yellowish white, the colour almost white at the

elbow and visible from above as a light cream-coloured border, the

contrast between the black hairs on the outer side and the bright

yellowish white of the inner side of the arm being most marked.

The light yellowish colour extends right down the inner sides of

the limbs as far as the hands and feet, where it is somewhat duller

and dirtier. Tail long, measuring 100 mm. longer than that of

bieti ; clothed with comparatively short hairs, except at the tip,

where the hairs are rather longer and form a distinct tuft.

Colour, above dark black mixed with creamy-white down the mid-

line with a creamy-buff border; terminal tuft almost pure white,

faintly washed with pale buff; ventral surface considerably

lighter.

Skull like that of bieti as regards the general shape, but smaller

and with smaller cheek-teeth. Supraorbital region rather more

like that of rowxellane than of bieti; muzzle formed as in the

latter species, without any well-marked concave depression of the

premaxillary region. Interorbital breadth less than in rowxellane.

Zygomatic arches almost parallel, not expanded laterally.

Dimensions of the type (measured in the flesh) :—

Head and body 520 mm.; tail 660; hind foot 165; ear 40.

Skull: greatest length 109 mm. ; basilar length 68 ; zygomatic

breadth 77 ; greatest breadth across orbital region 72°5 ; greatest

diameter of orbit 28; interorbital constriction 11; greatest width

of cranium 68°3; palatilar length 32:9; width of palate (inside

m!) 20:5; length of upper tooth-row, from front of first premolar

to back of last molar, 30. .

Hab. Yen-bay, Song-koi River, Tonkin.

Type. Adult female. Original number 8. Collected by

Mr. Alan Owston’s collector, Orii, on September 14, 1911.

This interesting Monkey was readily distinguished from its

nearest ally, Rhinopithecus bieti, by its dark black back, yellowish-

buff underparts, and light-coloured tail.

A young specimen, sent with the type, was coloured in much

the same manner as the adult, only very much paler, the crown

of the head and hind limbs being yellowish tinged with grey,

whilst the back was just commencing to become dark. The entire

underparts were, as in the adult, a rich yellowish-buff colour.

In the course of describing this new Snub-nosed Monkey,

Mr. Dollman had been in correspondence with Prof. E. L.

Trouessart, of the Paris Museum, concerning the specimens of

Rhinopithecus bietti under his charge, and was indebted to Prof,

Trouessart for much valuable assistance, most liberally given.

Without this assistance the description of this new Rhinopithecus

would have been a far more difficult matter.

No. 106.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

March 19th, 1912.

S. F. Harmer, Esq., M.A., Sc.D., F.R.S.,

Vice-President, in the Chair,

The Minutes of the last Scientific Meeting were confirmed,

Messrs. E. GerRARD & Sons sent fer exhibition :—

(a) A very fine head of a cow Sable Antelope, shot by Major

EK. C. F. Garraway, C.M.G., which was of interest in that the

mane was white instead of the normal dark brown, and in the

horns also being near the record for length.

(6) The skull of an Isabelline Bear with eight well-marked

incisors in the lower mandible instead of the normal six.

premaxillary bones which had arrested the shedding of the milk-

canines and the growth of the permanent canines. The skull

was quite adultand the remaining teeth were fully developed and

normal.

(Z) The skull of a Lion showing a severe injury or disease of

the left side of the palate and the upper mandible, which had not

affected the growth of the teeth.

Mr. OuprreLD THomas, F.R.S., F.Z.S., exhibited the skin and

skull of a Viverrine Carnivore from Tonkin, which, externally, was

remarkably like the Banded Civet (Hemigalus hardwickei), but

whose skull and teeth proved to be completely different. It was

proposed to be called

CHROTOGALE OWSTONI, gen. & sp. nn.

General coloration as in Hemigalus hardwicket. Skull with

long muzzle as in Hupleres. Incisor teeth broad, spatulate,

* 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,

with long level cutting-edges instead of points, their series

above forming nearly a half-circle. Cheek-teeth proportionally

small.

Hab. Tonkin.

Type. Young male. Original number 11.

Mr. Thomas at the same time proposed to distinguish ‘* Hemi-

gale” hoset, described by him in 1892, as a special genus to be

called

DIPLOGALE, gen. n.

segs of coloration normal, not banded.

kull with parallel-sided, square-fronted muzzle, the tooth-

rows nearly parallel. P* with double root.

Type. Diplogale hosei (Hemigale hosei, Thos.).

Mr. Guy DouumAn exhibited a Monkey, belonging to the rare

genus Lhinopithecus, trom Tonkin, which he considered to

represent a new species and proposed to call

RUHINOPITHECUS AVUNCULUS, Sp. Nn.

Size smaller than in R. bieti, M.-Hdw.; tail longer.

Colour of dorsal surface ible. under pai rts yellow ish buff; tail

black overlaid with yellowish white, terminal tuft white.

Dimensions of type :—

Head and body 520 mm.; tail 660; hind foot 165; ear 40.

Skull: greatest length 109 mm.

Hab. Yen-bay, Tonkin,

Type. Adult female, Original number 8.

Mr, A. Rapctyrre Duemore, F.R.G.S., gave a lantern exhi-

bition of a large number of photographic studies of wild animals

in British East Africa and Newfoundland. This very beautiful

series of pictures was briefly described by Mr. Dugmore, who

dwelt particularly on the advantage to be gained by hunting with

the camera, and showed that it was a sport as well as a means of

securing interesting and valuable material.

Mr. Epwarp W. Suann, B.Sc., read a paper, communicated by

Prof.S. J. Hickson, D.Sec., F.R.S., F.Z.8., entitled ‘‘ Observations

on some Alcyonaria from Singapore, with a brief Discussion on

the Classification of the Famiiy Nephthyide.” This collection

was made by Mr. W. F. Lanchester and the late Mr. F. P. Bedford

during their residence in Singapore. All the specimens had been

obtained in shallow water, from low water-mark to a depth of

about 10 fathoms, and of the eleven species dealt with in this

paper, representing six different families, four were described

as new. A brief historical summary of the genera Vephthya,

Dendronephthya, and Stereonephthya was given, together with

the author’s reasons for retaining the definitions of these genera

of Nephthyide as enunciated by Prof. W. Kiikenthal in his

‘Revision of the Alcyonaria.’ Among the illustrations were

reproductions of some of Savigny’s figures of his genera Ammothée

and Nephthée.

Sir Georcz H. Kenrick, F.Z.S., presented a paper entitled

“A List of Moths of the Family Pyralide collected by Felix B.

Pratt and Charles B. Pratt in Dutch New Guinea in 1909-10,

with Descriptions of new Species.”

Mr. T. H. Wrruers, F.G.S., read a paper, communicated by

Dr. W. T. Calman, F.Z.S., on “Some early Fossil Cirripedes of

the Genus Scalpellum.”

‘Attention was drawn to the form of the carina of the

geologically older species of Scalpellum, and it was shown that

the earliest forms known resembled more closely the carina of

Pollicipes, from which Scalpellum 1s considered to be derived.

An almost complete capitulum of the Albian Scalpellum arcuatum

was described, together with some scales of the peduncle, and a

restoration was given. This specimen was important because,

with the exception of a few detached valves found in the Aptian

(Lower Greensand), it was the oldest-known fossil Cirripede that

could with certainty be referred to the genus Scalpellum, sensu

lato. S. arcuatum was considered to be an ancestral form of the

group of almost exclusively deep-sea species, which Dr. P. P. C.

Hoek had separated as a subgenus under the name Arcoscalpellum,

and its relationship to other species was discussed. S. trilineatum

was also re-described.

The next Mecting of the Society for Scientific Business will

be held on Tuesday, April 2nd, 1912, at half-past Hight

o'clock P.M., when the following communications will be made :—

1. Exutpirions anp Noricns.

ES LE PET LE OTD

2. D. Seru-Surru, F.Z8.

Lantern Exhibition of Nestling Cariama, and the display of

the Peacock Pheasant.

3. R. I. Pococs, F.R.S., F.L.S., F.Z:8.

On a rare Stag (Cervus wallichii) from Nepal, presented to

the Society by H.M. The King. (With lantern illustrations.)

4, Frank EK. Bepparp, M.A., B.RS., F.Z.S.

Contributions to the Aue a Systematic Arrangement

of the Cestoidea.—IV. On Species of Znermicapsifer from the

Hyrax and on the Genera Zschokkeella, Thysanotenia, and

Hyracoteentt.

5. Dr. Basurorp Duran.

Additional Notes on the Living Specimens of the Australian

Lung-Fish (Ceratodus forsteri) in the Collection of the Zoo-

logical Society of London.

The following papers have been received :—

1. Junin S. Huxtey.

A First Account of the Courtship of the Redshank (Totanus

calidris).

2. Mrs. EH. W. Sexton.

Amphipoda from Bremerhaven.

3. C. Tare Rucan, M.A., F.ZS.

~ Descriptions of new Ae of the Family Loricariide in the

British Museum Collection.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Soctery or Lonpon,

Reegent’s Park, Lonpon, N.W,

March 26th, 1912.

No. 107.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON,*

April 2nd, 1912.

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

in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The Sxcrerary read a Report on the Additions that had been

made to the Society’s Menagerie during the months of February

and March 1912.

Dr. R. W. SxHurevot, C.M.Z.8., sent for exhibition the skins

of two young Virginia Opossums (Didelphis virginiana). These

specimens were each about ten weeks old and belonged to the

same litter.

The bones of the entire skeleton at this age were quite elemen-

tary in character, especially the terminal vertebre of the tail,

the bones of either carpus and those of the tarsi, and the epiphyses

of the long bones, ete. ‘he marsupial bones were well formed in

both sexes, and upon either side were nearly as long as the

corresponding ilium, and about one-fourth the size in bulk.

According to Flower, the number of vertebree in the spine of

the Virginia Opossum was 7 cervicals, 13 thoracic, 6 lumbar, and

26 caudals. This was probably correct for the adult animal of

this species, while in subaduits, of an age here considered, the

last three caudals were not developed, and the three or four

anterior to them were in the most rudimentary condition

possible.

* This Abstract is published by the Society at its offices, Zovlogical 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 ef publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payable in adyance.

In both these specimens the first dentition had perfectly erupted

and was as follows :—

; 1 2 3 9

tp €7, PMs, ma= 42,

the formula for the adult being

3 4

3, m y= 90.

: 1

a Cy, pm

The premolars were triangular, sharp-pointed, and flattened from

side to side; the molars had numerous sharp cusps, and the

canines were large and curved.

Mr. D. Sera-Smiru, F.Z.S., Curator of Birds, exhibited, by

means of lantern-slides, photographs of the male Peacock Pheasant

(Polyplectron chinquis) displaying to the female.

The typical display, as depicted in the photographs, resembled

very closely that of the Argus Pheasant, the bird facing the

female while he lowered the breast to the ground and expanded the

wings and tail like a shield, the head being held sideways against

one wing.

Mr. Seth-Smith also exhibited photographs of the young

Cariama cristata hatched and reared in the Gardens in 1911, and

remarked that although young of this species had been hatched in

the Menagerie on previous occasions, he believed that this was the

first occasion on which the young had been reared to maturity.

Mr. BR. I. Pococs, F.R.S8., F.L.S., F.Z.8., exhibited a lantern-

slide of two Polar Bear cubs born in the Gardens in November

1911, and made some remarks upon the causes of the difficulty

experienced in all Zoological Gardens in rearing the offspring of

this species.

Mr. C. Tare Rucan, M.A., F.Z.8., exhibited some lantern-

slides, prepared from photographs taken by Dr. F. B. Sumner,

of a Mediterranean Flatfish (Platophrys podas) on sand, gravel,

and various artificial backgrounds, showing its power of changing

its colour and markings to resemble the gr ‘ound on which it lies.

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

the Gardens, read a paper “ On a rare Stag o ( Cervus wallichit) from

Nepal, recently presented to the Zoological Society by His Majesty

Tone George.” The author pointed out the distinctive pecu-

liarities of this species, which, on account of its great scarcity,

had never been satisfactorily classified since it was described by

G. Cuvier in 1825 froma coloured illustration of a specimen living

at that time in the Barrackpoor Menagerie. Some authors had

supposed this Stag to be identical with the Kashmir species,

or Hangul (Cervus hanglw) ; others had referred it to the Chumbi

Valley species, or Shou (Cervus affinis). From both of these,

however, it differed in having the rump-patch of large size, ex-

tending, that is to say, nearly to the summit of the croup, and

showing no trace of a dark median line dividing it into a right

and left portion. Nevertheless, with respect to the size of the

rump-patch and the distinctness and width of this divisional line,

the Shou (C. affinis) seemed to be intermediate between the

Hangul (C. hanglu) and Wallich’s Stag (C’. wallichii). In certain

other respects, such as the colour of the lips, the length of the

head, and the coarseness of the hair, C. afinis and C. wallichii

were more like one another than either was like C. hanglu, and the

evidence tended to show that the two former were subspecies of

a species for which wallichii was the oldest available name.

Mr. F. E. Bepparp, M.A., F.R.S., F.L.S., Prosector to the

Society, read a paper on species of Tapeworms of the genus

Inermicapsifer obtained from the Hyrax, with notes on the

genera Zschokkeella and Thysanotenia. An account of the

structure and characters of the species was given, together with

the description of a new genus and two new species,

Dr. Basurorp DeAN contributed an account of the living

specimens of the Australian Lung-fish (Ceratodus forstert) in the

Society’s Collection. This paper contained some further observa-

tions made by the author in June 1911, supplementary to his

previous communication published in 1906, and dealt with the

coloration, size, and age of the specimens. Details of the rate of

growth of this species were also given, with notes on their method

of breathing, their food, and an account of the regeneration of a

portion of the left ventral fin which had suffered an injury.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, April 23rd, 1912, at half-past Hight

o'clock p.m., when the following communications will be made :—

1. Exurpirions anp Norices.

2. JULIAN S. Huxtey.

A First Account of the Courtship of the Redshank (Totanus

calidris).

3. Mrs. E. W. Sexton.

Amphipoda from Bremerhaven.

A. Cx ae Ruean, M. Noy | B.ZS

Descriptions of new ches “sf the Family Loricariide in the

British Museum Collection.

SCS LE CONGR Las IAs

“The Circulatory System of ie P Gator Grass-Snake (Zropi-

donotus natrix).

The following papers have been received :—

1. Major J. Svrvenson Haminton, C.M.Z.S.

The Local Races of Burchell’s Zebra.

. Rownanp HK. Turner, F.Z.S., FES.

Studies in the Fossorial Wasps of the Family Scoliide, Sub-

families Klidine and Anthoboscine.

3 G. A. Boutmnesr, F.R.S., ye ZS.

On z Gites of Fishes aa by Mr. A. Blayney Percival

in British East Africa to the East of Lake Baringo.

Dpane JB, Ennioien, Wig Wiles

“Contr ibutions to ne Ae and Sjstemati Arrangement

Devil ieee ursinis), the Type of a new Family.

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 9th, 1912.

No. 108.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

April 23rd, 1912.

Dr. 8S. F. Harmer, F.R.S., Vice-President, in the Chair,

The Minutes of the last Scientific Meeting were confirmed.

The Secretary exhibited a living specimen of a young female

Dorsal Hyrax (Dendrohyrax dorsalis) from Nigeria, recently

presented to the Society by Mr. J. L. McKellar.

The Secretary exhibited a number of photographs of an

Elephant Kraal in Siam which had been presented to the

Society by the Rt. Hon. Sir Cecil Clementi Smith, P.C.,

GOM.G:

Mr. C. H. O’Donoeuvsz, B.Se., F.Z.S., read a paper on the

Circulatory System of the Common Grass-Snake (7Z'ropidonotus

natrix). Several interesting features correlated with the loss of

limbs and the elongation of the body were stated to occur in the

blood-vessels. The vessels, like the viscera they supplied, were

asymmetrical ; not only were those on the right anterior to those

on the left, but they were also noticeably larger. No indication

of the descent of snakes from a limb-bearing ancestry was to be

found in the circulatory system, save perhaps a small pair of

veins which might correspond to the pelvic veins in Lacertilia.

There was a marked tendency for the vessels to form longitudinal

systems, ¢. g., the arteries supplying the gut and the fat-bodies ;

and each ovarian artery formed a longitudinal trunk along the

corresponding supra-renal body. The hepatic portal vein arose

by two roots, one from each renal portal vein, and ran the whole

length of the gut up to the liver. By the side of each oviduct

* 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,

was a conspicuous oviducal sinus, a vessel which had not been

described previously in snakes.

The right carotid artery was not present in the adult, and to

compensate for this the left side of the head received its arterial

blood by means of three anastomoses—one beneath the medulla

oblongata, one beneath the fore-brain, and one beneath the

symphysis of the lower jaw. The part of the anterior cardinal

vein in the head of the embryo was completely replaced during

development by a new vessel, the lateral cephalic vein.

Mr. Juuian S. Huxuery read a paper, communicated by Prof. G.

C. Bourne, D.Sc., F.R.S., F.Z.S., containing an account of the

Courtship of the Redshank (Z'otanus calidris).

The first purpose of this paper was to draw attention to the

many valuable results to be obtained by simple watching of very

common British birds ; and the second was to show how the facts

observed in the Redshank bore on the theory of Sexual Selection.

Tn this species there was no rival display between several males at

once: a single female was courted by a single male, as in Man.

The courtship started with a pursuit, the hen running in a

circuitous course, followed by the cock. The pursuit was followed

by a display, but only if the hen were willing that the courtship

should continue. During display the cock uttered a special note,

spread his tail, raised his wings above his back, and advanced with

a curious high-stepping action towards the now stationary female.

Tf the female so wished, pairing followed the display. But in quite

90 per cent. of observed courtships the female rejected the male,

either during the pursuit or during the display, by simply flying

away, when the cock was quite powerless to enforce his desires.

Thus the consent of the hen was absolutely necessary if pairing

were to take place, and this consent was usually withheld: in other

words, selection by the female was a reality in the Redshank,.

Other interesting points were as follows :—The plumage of the

two sexes was identical, and was decidedly cryptic when the birds

were at rest. During flight the white underside of the wings

and the white tail were conspicuously revealed, and probably

served as recognition marks. ‘The significance of the red legs

‘was unknown. During display the male drew attention to

the underside of the wings by raising and vibrating them, to the

tail by fanning it out, and to the red legs by his slow, high

steps; besides this he uttered a note heard at no other time.

Thus, since the actual colours and structures used in display were

found in both sexes, the only peculiarly male possession—the

only secondary sexual character of the Redshank—was a special

behaviour, devoted to showing off these common colours and

structures in a special way.

This seemed to show that secondary sexual differences in birds

were originally differences of behavionr, and that only when

these were established did differences of colour and structure

come to be developed.

Mrs. E. W. Sexton contributed a paper, communicated by the

Rev. T. R. R. Stebbing, M.A., F.R.S., F.Z.S., based on a small

collection of brackish- water Amphipoda from Bremerhaven.

Special reference was made to a new species of Gammarus, which

inhabited both fresh and brackish water, and was interesting as

showing in a marked manner the effects of environment on .

development,

Mr. C. Tare Recan, M.A., F.Z.S., read a paper containing

descriptions of ten new species of South American Fishes of the

Family Loricariide in the British Museum Collection.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, May 7th, 1912, at half-past Hight

o’clock p.m., when the following communications will be made :—

1. Norices AND EXHIBITIONS,

2. FrANcIS WaRD, M.D., F.Z.8.

Lantern and Kinematograph Demonstrations of Photographs

of Fishes and Aquatic Animals in Natural Hlumination.

3. G. A. Boutencer, F.R.S., F.Z.8.

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

in British East Africa to the East of Lake Baringo.

4. Rowianp E. Turner, F.Z.S8., F.H.S.

Studies in the Fossorial Wasps of the Family Scoliide, Sub-

families Hlidine and Anthoboscine.

5. Ape. CHApmay, F.Z.S.

Notes on the Spanish Ibex.

The following papers have been received :—

1. Major J. Stevenson Hamitton, C.M.Z.S8.

The Local Races of Burchell’s Zebra.

2 _ FRANK EK. BrpparD, : M. A., , -R. 8., et Z. 8.

~ Contributions to the ‘Anatomy and Systematic Arrangement

of the Cestoidea.—V. On a new Genus from the Tasmanian

Devil (Dasyurus ursinus), the Type of a new Family.

3. WuiuuiAM Nicott, D.Sc., F.Z:S.

On two new Larval Trematodes from the Striped Snake.

4. W.T. Cauman, D.Se., F.Z.8

On ies a new Ce of the Crustacean Order

Branchiura.

f, Sg Uno teen, ICG Oe ee

‘A Note on the rare Brush Nadia ror Baas eudactylota

Gosse.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

Zoo.tocicat Sociery or Lonpoy,

Recent’s Park, Lonpon, N.W.

April 30th, 1912.

No. 109.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON®*

May 7th, 1912.

Prof. E. A. Mincuty, F.R.S., Vice-President, in the Chair,

The Minutes of the last Scientific Meeting were confirmed,

Mr. R. I. Pocock, F.R.S., F.L.S8., F.Z.8., Superintendent of

the Gardens, exhibited a skin and a living specimen of a fawn

variety of the Brown Rat (Hpimys norvegicus), which had been

caught on an island in the middle of Loch Corrib, Co. Galway,

and presented to the Society by Lord Headley. Mr, Pocock

remarked that although similarly coloured varieties of this rat

had been caught now and again in different parts of England, it

was especially interesting to put on record Lord Headley’s state-

ment that it was quite common on the island, no fewer than

eleven having been trapped while others had been seen ; and that

it did not occur, so far as was known, on the mainland. Typically

coloured brown rats lived on the island as. well.

Mr. D. Sera-Smirx, Curator of Birds, exhibited two horn-like

sheaths which had been shed from the orange-coloured patch at:

the base of the lower mandible of the King Penguin (A ptenodytes

pennanti) living in the Society’s Gardens. Mr. W. E. de Winton

had observed the shedding of this epidermal sheath in a bird living

in the Gardens in 1898 (P. Z. 8. 1898, p. 900); but although the

present specimen had been carefully watched during two successive

moults in March and October 1911 (P. Z.S. 1912, p. 60), no sign of

* 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 pey annum, payable in advance,

this process wasobserved. The bird, however, went through another

complete moult in March to April of the present year (1912), and

shortly after this was completed the epidermal covering of these

orange-coloured patches became loose and finally fell off; the

pieces somewhat resembled the wing-cases of a large beetle,

being semi-transparent and of a clear orange-colour.

Dr. Francis Warp, F.Z.8., showed a number of photographs

and diagrams illustrating a method of observation of fishes, birds,

and mammals under the water, the principle being that the

subjects under consideration were illuminated by natural light,

and the observer being ina dark chamber in the water was not

seen. The appearance of black-feathered birds was shown;

these, by carrying down air-bubbles among the feathers, were

converted into reflectors : and a Water-Hen was shown bright red

and then green as it reflected the different surroundings in

which it had been placed. Otters and Seals were also shown as

seen under the water. The demonstration was illustrated by

numerous slides and by the cinematograph.

Mr. G. A. Boutenerr, F.R.S., F.Z.S., presented a paper on a

collection of Fishes made by Mr. A. Blayney Percival in British

Kast Africa to the east of Lake Baringo. This collection was of

special importance as coming from a district the fishes of which

had not been collected before, and contained examples of five new

species.

Dr. F. E. Bepparp, M.A., F.R.S., F.Z.S., Prosector to the

Society, gave an account of his paper on a new Genus of the

Cestoidea, founded on some specimens of Tapeworms which he

had discovered in the small intestine of an example of the

Tasmanian Devil (Dasyurus ursinus). In briefly describing the

most salient points of anatomical interest in this form, which

formed the type of a new Family, he remarked that in view of

the very considerable peculiarities of structure observed it was

remarkable that the generative organs did not show any marked

features of interest as compared with those of other Tapeworms.

Mr. R. E. Turner, F.Z.8., F.E.S., communicated a memoir

entitled ‘‘Studies in the Fossorial Wasps of the Family Scoliide,

Subfamilies Elidine and Anthoboscine.” Several new species of

Elidine from South Africa were described, including a new

genus in which the female was wingless, and the genus

Anthobosca was monographed. The geographical distribution of

Anthobosca, which was almost entirely confined to the Southern

Hemisphere, was discussed, and the conclusion was reached that

the distribution was due to survival from a wider range in the

past, and not to a southern origin.

31 -

Mr. Apex Cnapman, F.Z.S., contributed a short paper

containing some notes on the Spanish Ibex, with reference to

Prof. Angel Cabrera’s recent paper on this species,

The next Meeting of the Society for Scientific Business will

be held on Tuesday, May 21st, 1912, at half-past Hight

o'clock p.M., when the following Comiiunications will be made :—

1. A. Buayney Perctvat, F.Z.S.

Lantern Exhibition of Game Animals from British East

Africa.

2. Major J. Stevenson Hamiuton, C.M.ZS.

The Local Races of Burchell’s Zebra.

3. WiuuiAM Nicotu, D.Sc., F.Z.S.

On two new Larval Trematodes from the Striped Snake.

4. W.'T. Cautman, D.Sc., F.Z.8.

On Dipteropeltis, a new Genus of the Crustacean Order

Branchiura.

5. G. A. Bouencer, F.R.S., F.Z.8.

Second Contribution to our Knowledge of the Varieties of

the Wall-Lizard (Lacerta muralis).

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

————— an

A Note on the rare British Nudibranch Hancockia eudactylota

Gosse,

The following papers have been received :—

1. R. Lypexser, F.R.S., F.Z.8.

The North Rhodesian Giraffe.

2. Prof. 8. J. Hickson, F.R.S., F.Z.S.

On the Hydrocoralline Genus Hrrina.

32 -

3. FRANK E . Bepparp, |] M.A., D Se., aE RS qlee Ss.

Contributions to the renee ree Sy eoiatie Arrangement

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

Rodent Fiber xzibethicus, showing a new form of Asexual

Propagation, and on the supposed ‘Sexual form.

4. HELEN lie INL, LEibeaiiin, BSe., EZ

Poly cheeta from Pacific a North America.—Part I.

Ser pulidee, with a Revised Table of Classification of the Genus

Spirorbis.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society oF Lonpon,

ReEGENtT’s Park, Lonpon, N.W.

May Ath, 1912.

Wo. 110,

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

May 21st, 1912.

Sir Epmunp G. Loprr, Bt., Vice-President, in the Chair,

The Minutes of the last Scientific Meeting were confirmed.

_ The Secretary read a report on the Additions that had been

made to the Society’s Menagerie during the month of April 1912.

Mr. A. Buayney Percivan, F.Z.8., exhibited a number of

photographs and lantern-slides of Game Animals from British

East Africa, including a fine series of the Reticulated Giraffe.

Mr. D. Sera-Smiru, F.Z.S., Curator of Birds, exhibited two

living specimens of a rare Lory, Calliptilus solitarius, from Fiji,

and remarked that Dr. Philip H. Bahr had recently brought

home two specimens which had died. The specimens exhibited

were from a collection of eight brought home alive by Mr. Rood

Tarte, of Taviuni Island, one of the Fijian group, where this very

beautiful species was still abundant, its numbers having been

very considerably reduced in the other islands by the introduced

mongoose. The exhibitor referred to a recent note on the species

by Dr. Babr in ‘ The Ibis’ for April 1912, p. 293.

Major J. Stevenson Hamiuron, O.M.Z.S., the Game Warden of

the Transvaal, communicated a short paper, illustrated by photo-

graphs, on the local races of Burchell’s Zebra, and pointed out that

it was possible to shoot in one herd individuals presenting the

characters of various subspecies as described by systematists. In

the Transvaal, for example, he obtained skins exhibiting features

claimed to be distinctive of such races as LZ. burchelli wahlbergi,

E.b. transvaalensis, and £. b. chapmanni ; and from his experience

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent’s Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications; but it may be obtained on the

day of publication at the price of Siapence, or, if desired, sent post-free for

the sum of Sia Shillings per annum, payable in advance,

he expressed the opinion that these subspecies had been based

upon inadequate museum material.

Dr. WitiiAM Nico, F.Z.8., communicated some observations

on two new Trematode larve found encysted in enormous numbers

in the mesentery of several Striped Snakes ( 7ropidonotus ordinatus

sirtalis) which had died in the Society’s Gardens. He named these

forms, as neither could be referred to any adult species already

known. It could be safely predicted, however, that the second

species completed its life-cycle in the intestine of a bird, and from

this fact it could be inferred that the Striped Snake was eaten by

birds.

Dr. W.T. Catan, F.Z.S., read a short paper describing a new

genus and species of the Cructacean Order Branchiura.

A parasite of fishes collected by Spencer Moore, Esq., at

Corumba, Matto Grosso, Southern Brazil, was referred to a new

genus as follows :—

DIPTEROPELTIS, gen. n.

Differing from Argulws in having no spine on the preoral

papilla ; in having the antennules and antenne very minute and

imperfectly segmented ; in having no large spines or hooks on the

under surface of the carapace, body, or appendages ; in having no

furcal rami on the abdomen ; and in having the lateral wings of

the carapace greatly elongated.

Genotype, D. hirundo, sp. n., with the characters of the genus.

Mr. G. A. Boutencer, F.R.S., F.Z.8., read a paper entitled

“Second Contribution to our Knowledge of the Varieties of the

Wall-Lizard.” This paper was a continuation of one published

in the Society’s ‘ Transactions ’ in 1905, and dealt chiefly with the

variations of Lacerta muralis in South-Eastern Europe and South-

. Western Asia. It also contained a supplement to the first part,

thus completing an account of the varieties, of which about thirty

were regarded as more or less definable, the author endeavouring

to show the inconstancy of the characters adduced by some her-

petologists in assigning specific rank to a number of these forms,

connected by many gradations. Mr. Boulenger hoped to support

his statements by a number of photographic figures of specimens

selected out of the enormous material which had passed through

his hands in the course of his study of this polymorphic and widely

distributed lizard.

This paper will be published in the ‘Transactions’ in due

course.

A short note on the rare British Nudibranch Hancockia

eudactylota Gosse was received from Sir Charles Eliot, K.C.M.G.,

C.B., F.Z.S., with some coloured figures of a single specimen

captured at Plymouth. These were of special interest, as no

coloured figure of this species had yet been published.

The next Meeting of the Society for Scientific Business (closing

the Session 1911-1912) will be held on Tuesday, June 4th, 1912,

at half-past Eight o’clock p.m., when the following communications

will be made :—

1. E. G. B. MeapEe-Watpo, V.P.Z.S. will introduce a discussion

on the Preservation of the English Fauna.

2. R. Lypex«er, F.R.S., F.Z.S.

The North Rhodesian Giraffe.

3. Prof. 8. J. Hickson, F.R.S., F.Z.S.

On the Hydrocoralline Genus Zrrina.

4, Frank KE. Bepparp, M.A., D.Sce., F.R.S., F.Z.S8.

Contributions to the Anatomy and Systematic Arrangement

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

Rodent Fiber zibethicus, showing a new form of Asexual

Propagation, and on the supposed Sexual form.

5. Heten L. M. Prxett, B.Sc., F.Z.S8.

Polycheta from the Pacific Coastof North America.—Part I.

Serpulide, with a Revised Table of Classification of the Genus

Spirorbis.

6. R. Broom, D.Sc., C.M.ZS.

On some new Fossil Reptiles from the Permian and Triassic

Beds of South Africa.

The following papers have been received :—

1. E. Duxr1nFretp Jonss, F.Z.S., F.E.S.

Descriptions of new Butterflies of the Genus Thecla from

8.E. Brazil.

2 ee Bsns VAG BSc.

On the Development of the Pectoral Girdle in the Pipe-fish

(Syngnathus acus).

3. F. F. Larpraw, F.Z:S.

Some Notes on Bornean Dragonflies, with Descriptions of

New Species.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society or Lonpon,

ReEcENtT’s Park, Lonpon, N.W.

May 28th, 1912.

Nor dE.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

June 4th, 1912.

E. G. B. Meapr-Watpo, Esq., Vice-President, in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

Mr. E. G. Bourencer, Curator of Reptiles, exhibited a clay-

pall containing a cocoon of the African Lung-fish (Protopterus

annectens) presented to the Society by Capt. J. A.M. Vipan, F.Z:8.

He briefly alluded to the habits of the fish and the formation of

the cocoon, and gave an account of the method to be employed in

order to release the fish,

The Preservation of the Native Fauna of Great Britain.

Mr. E. G. B. Meapre-Wapo, Vice-President of the Society,

ntroduced a discussion on the Preservation of our Native Fauna,

in which Mr. A. Heneage Cocks, Dr. F. G. Dawtrey Drewitt, and

Mr. Stewart Blakeney (who sent a written contribution) joined.

‘Lhe necessity of creating public opinion on the matter was urged.

It was agreed that the laws with regard to birds were sufficient,

if administered strictly. With regard to mammals, it was the

opinion of those present that the use of steel traps, instead of

snares, for catching rabbits was chiefly responsible for the ex-

termination of wild cats, martens, and polecats in many parts of

the country, and ought to be suppressed.

Mr. R. Lypexkxer, F.R.S., F.Z.S., communicated a short paper

describing a new local race of Giraffe from the Petauke district

of North-east Rhodesia.

* 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.

Miss Heten L. M. Prxett, B.Sc., F.Z.S., read a memoir entitled

“ Polycheta from the Pacific Coast of North America. Part I.”

This paper contained a description of Serpulide from the Straits

of Georgia, chiefly the Departure Bay region of Vancouver Island,

together with some specimens from Victoria and Puget Sound,

eighteen species inall, of which five werenew. Serpula splendens

(Bush, 1905) was shown to be identical with Serpula columbiana

(Johnson, 1901); and Caullery and Mesnil’s classification of the

genus Spirorbis was revised and enlarged to contain three new

species and such other Pacific species as had been adequately

described. Further evidence was given in support of the sug-

gestion made by Caullery & Mesnilin 1897 that Spirorbis (Circeis)

armoricana (St. Joseph, 1894) was only a variety of Spirorbis

spirillum (Linné, 1760).

Mr. R. 1. Pocock, F.R.S8., F.Z.S., Curator of Mammals, read a

paper, illustrated by lantern-slides, on antler growth in the

Cervide with special reference to Hlaphurus and Dorcelaphus,

and pointed out that the growth of the individual antler in

Elaphurus, as shown by a series of sketches kindly supplied by

Lord Tavistock, proved that the anterior and posterior branches

of the antler of Hlaphurus were homologous with the brow-tine

and beam of the Sambar’s antler, and that in Dorcelaphus the sub-

basal snag was the homologue of the brow-tine in the Old World

deer as Sir Victor Brooke claimed.

Dr. Hans Gapow, F.R.S., F.Z.8., read a paper on ‘The One-

sided Reduction of Ovaries and Oviducts in the Amniota, with

remarks on Mammalian Evolution.” He stated that the

reduction began with the oviduct, and a first cause of the

invariably right-sided bias had to be looked for in the turning of

the embryo upon its left side, a position which influenced the

growth and relative position of the stomach and primary

intestinal loops, these being stowed in the abdomen in such a

way that they were less disturbed by an egg passing through the

left than through the right oviduct. In the Monotremes also

only the left ovary and duct were functional, although those of

the right side were structurally not affected. This was not a case

of reptilian inheritance. Proto- Meta- and Eutheria represented

a continuous, monophyletic line of evolution, with the Monotremes

and Marsupials as offshoots. The Metatherian stage was diphy-

odont, marsupiate and placental. It was the parting of the ways.

Those which developed a corpus callosum, correlated with higher

mental faculties by further improving the placentation, and losing

the marsupium (not the bursz), became Placentalia; whilst the

remaining stock, being driven to arboreal life, intensified the

marsupial and thereby diminished the uterine gestation. Their

arborealism implied the necessity of taking their young with them.

When Eutheria took to arboreal life the chance of reviving the

lost marsupium was gone, nor did their higher mental and

placental organization require it.

Dr. F. E. Bepparp, M.A., F.R.S., F.Z.S8., gave an account

of an asexual Tapeworm, obtained from the Musquash (Fiber

zibethicus), showing a new form of asexual propagation, and also

described a sexual worm which he believed to be the mature form

of the same Tapeworm,

Dr. Wiuur1AM Nico, F.Z.8., described two new Trematode

parasites from the Indian Cobra (Naja tripudians). The first

was found in the gall-bladder and was made the type of a new

genus of the family Dicrocoeliide. The second was found in the

ureters and represented a new species of the genus Styphlodora.

Dr. Nicoll also gave statistics of the Worm Parasites collected from

the animals which had died in the Society’s Gardens during the

period from December 1910 till April 1912.

Dr. R. Broom, C.M.Z.8., presented a paper based on some new

Fossil Reptiles from the Permian and Triassic Beds of South

Africa.

Prof. 8. J. Hickson, F.R.S., F.Z.S., communicated a paper on

the Hydrocoralline genus Hrrina. This genus was founded by

Gray in 1835, and since that date two other genera (Labiopora

and Spinipora) closely related to Hrrina had been described. An

analysis of the characters of these three genera was given.

An examination of Gray’s type of Errina proved that it was

more closely related to the species referred by Moseley and others

to the genus Labiopora, than to the species referred by them to

the genus Hrrina. The author proposed, therefore, to merge the

three genera into one with the common generic name L7rrina,

dividing the species into three subgeneric groups. Two new

species were described, one from New Zealand waters and the

other from the Cape of Good Hope.

This Meeting closes the Session 1911-1912. The next Meeting

of the Society for Scientific Business will be held on Tuesday,

October 29th, 1912, at half-past Hight o’clock p.m.

The following papers have been received :—

1. E. DuxinrieLp Jonss, F.Z.8., F.E.S.

Descriptions of new Butterflies of the Genus Thecla from

S.E. Brazil.

2. F. F, Larpuaw, F.Z.S.

Some Notes on Bornean Dragonflies, with Descriptions of

New Species.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society or Lonpon,

Recent’s Park, Lonpon, N.W.

June 11th, 1912.

Papers (continued).

Pie

Page

20. On the Pairing of Pseudoscorpiones. By H. Waxuts Kew, F.Z:S. (Text-figs. 47-50.) 376

21. A Contribution towards the Knowledge of the Spiders and other Arachnids of

Switzerland. By the Rev. O. Prckarp-Camsripeas, M.A., F.RS., C.M.ZS.

(ext-fies: 51h 52.) es eee ose Bish CON TREN CT BOE DEI a Sarde S ck wode 398

22. On the Blood-parasites found in Animals in the Zeological Gardens during the four

years 1908-1911. By H. G. Puruer, F.R.S., F.Z.S., Pres.R.M.S., Pathologist to

GEL tC Ee Stee IE TAN Yat alc sbecehn'e a9 aja: ease a +0 pace, « don talsnn deat as ate eas 406

23. On the Structure of the Internal Ear and the Relations of the Basicranial Nerves in

Dicynodon, and on the Homology of the Mammalian Auditory Ossicles. By R. Broom,

MP sey OOM Z Sir. (bis aV Land Text-fig, 53.) 0 co. ee as oe aleleleveime a ale tune state 419

24. Zoological Results of the Third Tanganyika Expedition, conducted by Dr. W.

A. Cunnineron, 1904-1905. Report on some larval and young stages.of Prawns from

Lake Tanganyika, By Prof. G. O. Sars, C.M.Z.S. (Pls. LVIL-LX.)............. 426

25. The Classification, Morphology, and Evolution of the Echinoidea Holectypoida. By

Hersret L. Hawgriys, MSc, F.G.S8.; Lecturer in Geology, University College,

ume ( MieC BIO, B—-GU.) cv ie aye av sess « cnoy « viisde. # 60 0, 6 #inié do ve vin la view nithe a) Selle a eA

Titlepage i

List of Council and Officers ii

List of Contents........ ili

Alphabetical List of Contributors ix

New Generic Terms ........... xvi

SMPTE ERM eA eS CLC IROL ETT ONL IELEN I *solvivr ee. cose artlaverepeheta +6, 0 Wid mye, uals oie» oslcahe alan som ale AT ee Pade tea xvii

Tndex of Bisaeeuisone Bd oor

LIST OF PLATES.

1912, Part IL. (pp. 241-504).

Plate Page

XXXII)

XXXIV.

XXXV.

XXXVI. |

KXRWII! |

XXXVIIL.

DOOD |

XL. Peper a veg Gd gate ok ARN ln eyelet hel

XLVIII. Mimetic Cockroaches and Beetle Models ............... 858

XLIX.y

L. |

LL. | :

HeLa SPOOR ATABLLESE 50 wolaye thee ermal a lllerainer aN eh ae reeY a eek te eee

LI.

LIV. |

Ly. :

LVI. The Auditory Regiom in Dicynodon ........0.......++2. 419

LVII. \

ee Cee from: ake. Tanganyika +/7).\s0) Jens ose vase aoe

LX. )

f NOTICE.

The ‘ Proceedings’ for the year‘are issued in fowr parts, paged consecutively,

so that the complete reference is now P. Z. 8. 1912, p.... The Distribution

is as follows :—

Part I. issued in Mareh.

asi Ned ls 3 June.

» LIL. 4, September:

Cad BN - December.

‘ Proceedings,’ 1912, Part I. (pp. 1-240), were published on

March 13th, 1912.

‘The Abstracts of the Proceedings,’ Nos.106 to 111 are

contained in this Part. 43)

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ih yi i i i ) / f )

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