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PROCEEDINGS

OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY

OF LONDON.

1912, pp. 505-913,

witH 37 Puates and 63 TEXx’r-FIGURES.

223826

PRINTED FOR THE SOCIETY,
AND SOLD AT THEIR HOUSE IN REGENT’S PARK.
LONDON:
MESSRS. LONGMANS, GREEN, AND Co,
PATERNOSTER ROW,

ky JES) ae

OF THE

COUNCIL AND OFFICERS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

WS

COUNCIL.

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

THe Haru or Anramont, F.S.A.

Str Joun Rose Braprorp,
KC MEG MaDe SDiScs
F.R.S., Vice-President.

RicnarD H. Burne, Esq,
M.A.
Lt.-Col. Str R. Havetock

Cuaruss, G.C.V.O., M.D.
ALFRED H. Cocks, Esq., M.A.
F. G. Dawrrey Drewirt, Hsq.,

M.A., M.D.
CuHarLes DrumMonp,

Treasurer.

Sir Epwarp Duranp, Bt., C.B.
F. Du Cant Gopmayn, Ksq.,

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

Sir Epmunp G. Lopsr, Bt.,

Vice- President.

Ksq.,

E.G. B. Meapr-Watpo, Esq.,
Vice-President.

Prof. Epwarp <A. MuINcHIN,
M.A., F.R.S., Vice-President.

P. CHatmers MircnHent, Esq.,
Wiles IDisesy elon, Ibib. JD).
ELRS., Secretary.

W. R. Ocitvir-Grant, Esq.

ALBERT Pam, Hsq.

ApriAN D. W. Potiock, Esq.

OLDFIELD THomas, Hsq., F.R.S.

AntHony H. WINGFIELD,
Ksq.

A. Smith Woopwarpb, Ksq.,
LL.D.,F.R.S., Vice-President.

Henry Woopwarp,Ksq., LL.D.,
F.R.S., Vice-President.

PRINCIPAL OFFICERS.

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

Secretary.

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

R. I. Pocock, F.RS., F.L.8., Curator of Mammals and
Resident Superintendent of the Gardens.

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

Hewry G. Purmer, F.R.S., M.R.C.8., Pathologist.

Epwarp G. Boutencmr, Curator of Reptiles.

F. H. Wareruouss, A.L.S., Librarian.

JOHN Barrow, Accountant.
W. H. Coun, Chief Clerk.

NV H

LIST OF CONTENTS.

1912, pp. 505-913.

EXHIBITIONS AND NOTICES.

The Secretary. Report on the Additions to the Society's
Menagerie during the months of February and March

Mr R. 1. Pocotk, FURS, PALS. F:Z.S: Exhibition of
lantern-slide of two Polar Bear cubs (Ursus maritimus)
born; im-thes Gardens? .oiaje.0. eee, See Pata ne, 5D6

Mr. C. Tats Reean, M.A., F.Z.S. Exhibition of lantern-
slides of a Flatfish (Platophrys podas), showing its
power of changing its colour and markings................ 556

Dr. R. W. Suurenpr, C.M.Z.S. Exhibition of skins of two
Virginia Opossums (Didelphis virginiana) .......0..6.... 556

Mr. D. Seru-Suirn, F.Z.8. Exhibition of lantern-slides of
the display of the male Peacock Pheasant (Polyplectron
chinguis). Also of photographs of the young Cariama
cristata hatched and reared in the Gardens. (Pl. LXIX.) 557

The Sucrerary. Exhibition of a living specimen of a young
female Dorsal Hyrax (Dendrohyrax dorsalis) ............ 671

1V

The Secretary. Exhibition of photographs of an Hlephant
Lee Thal SVEN aaseoode sonopEeanOORUDARAAteGsd024J0540554cd000nc

Mr. R. I. Pocock, F.R.S., F.LS., F.Z.S. Exhibition of a
skin and a living specimen of a fawn variety of the
Brown Rat (Hpimys norvegicus) ........-00....:.6er es. -es

My. D. Seru-Smiru, F.Z.S. Exhibition of epidermal sheaths
shed from the base of the lower mandible of the King
Penguin (Aptenodytes pennanti) ............ce+csssseers

Dr. Francts Warp, F.Z.S. Demonstration, illustrated by
lantern-slides and the cinematograph, of a method of
observation of fishes, birds, and mammals under
AWE NIZIO! cuonodsonanagsaoasoUsoddeoonoboneooanddpuDoudovbugdueDusascd

The Secretary. Report on the Additions to the Society’s
Menagerie during the month of April 1912...............

Mr. A. Buayney Percivat, F.Z.S. Exhibition of photographs
and lantern-slides of Game Animals from British East
JANA is dockets < Rlinidcot s Sak ena arc EEE eR REE oR eR CREE

Mr. D. Seru-Smire, F.Z.S. Exhibition of two living
specimens of the Lory (Calliptilus solitarius), from
TRY s9c.ansd09se8 adotge 39099900 sodoad2acanqosq0050 sbonIRDOcHasdoBcd

Mr. G. A. Boutencer, F.R.S., F.Z.8. Notice of a paper
entitled “Second Contribution to our Knowledge of
ThenViaricties on the \\Velllelnizands dene) ees see

Mr. E. G. Boutencer, F.Z.S. Exhibition of a clay-ball
containing a cocoon of the African Lung-fish (Proto-
PlEVUS GMNECLENMS)....--- 0 ..c0te veer nsec ncen see eetnesteene cere.

Mr. E. G. B. Meape-Watpo and Others. Discussion on the
Preservation of the Native Fauna of Great Britain ..

The Secrerary. Report on the Additions to the Society’s
Menagerie during the months of May, June, July,
August, and September 1912 .............eseeeeeeeee sees

Mr. R. Lypexner, F.RS., F.Z.S. Note stating that
“ Gazella hayi” = Gazella fuscifrons ........0.. weceeeere ees

Page

671

671

806

806

806

807

807

. 807

908

eI ik

Vv

The Rev. T. R. R. Srespine, M.A., F.R.S., F.Z.S. Notice
of a memoir on the Crustacea Isopoda collected by the
‘Porcupine’ Expedition in 1869-1870 ...............05

Mrs. Rosz Hare Tuomas, F.Z.S. Exhibition of, and remarks
upon, the Eggs of Phasianus formosanus, P. versicolor,
and their F. 1 and F. 2 offspring. (Text-fig. 126.) ...

Sir Epmunp G. Lopmr, Bt., V.P.Z.S. Demonstration of the
capacity of the electric lantern presented by him to the
SOCICle megmememtes ite eae ren teste n aes coal TaMt,\. Ss cts

PAPERS.

26. Observations on some Alcyonaria from Singapore, with
a brief Discussion on the Classification of the Family
Nephthyide. By Epwarp W. Swann, B.Sc., De-
monstrator and Assistant Lecturer in Zoology in the
Victoria University of Manchester. (Pls. LXI.-
DN cere asia? caper evi ornnge esisieiaiewee Can acs c.ciinelcigenavea anaes wus

27. Some early Fossil Cirripedes of the Genus Scalpellum.
By Tuomas H, Wiruers, F.G.8. (Text-figs. 64 & 65.)

28. Experimental Pheasant-breeding. By Rosz Hate
MEOM AS, BZi.Se0 (isa eXeiV. INOW A ee eet

29. 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.
By Sir Grorce H. Kenricx, F.Z.8. (Pl. LXVIII.) .

30. Ona rare Stag (Cervus wallichii) from Nepal recently
presented to the Zoological Society by His Majesty
King George. By R. 1. Pocock, F.R.S., F.L.S., F.Z.8.,
Superintendent of the Gardens. (Text-figs. 66-71.)...

Page

913

528

546

5D8

dl.

32.

34.

36.

Sie

38.

39.

vi
Page
Contributions to the Anatomy and Systematic Arrange-
ment of the Cestoidea.—IV. On a Species of Jnermi-
capsifer from the Hyrax, and on the Genera Zschokkeella,
Thysanotenia, and Hyracotenia. By Frank K.
Bepparp, M.A., D.Se., F.R.S., F.Z.S., Proseetor to
thegooctebye ((lext-igs. (2a) eaten eee ene 576

Additional Notes on the Living Specimens of the
Australian Lung-fish (Ceratodus forster’) in the Col-
lection of the Zoological Society of London. By
Basurorp Dean. (Text-figs. 84 & 85.) ...............-6 607

. The Circulatory System of the Common Grass-Snake

(Tropidonotus natrix). By Cuas. H. O’ Donocuus,
BSc., F.Z.S., Assistant to the Jodrell Professor of
Zoology, University College, London. (Pls. LXX.—
TOROS, eral DUepe sti, (K=OIls)), so oonenoceassc0snssocecansnes 612

A First Account of the Courtship of the Redshank
(Totanus calidris). By Jurtan S. Huxuey, Lecturer
ofsballvol CollesevOxtond: a: ace Re ee eee renee ee 647

. Some Brackish-water Amphipoda from the Mouths of

the Weser and the Elbe, and from the Baltic. By
HE. W. Sexron, Marine Biological Laboratory, Ply-
moubh. =(CPis: WexeXN, dy NGXOXTRV 3) eee aes 656

Descriptions of new Fishes of the Family Loricariide in
the British Museum Collection. By C. Tarr Ruean,
WiloaNey J8o/AjS | (Uelise JU2OR WE IDO WINE) Ssonovcos030200 666

On a Collection of Fishes made by Mr. A. Blayney
Percival in British East Africa to the East of Lake
Baringo. By G. A. Bounencer, F.R.S., F.Z.S. (Pls.
AIOE TSTMS STE ONGC CN Es foe Cr ee 672

Contributions to the Anatomy and Systematic Arrange-
ment of the Cestoidea.— V. On a new Genus (Dasyuro-
tenia) from the Tasmanian Devil (Dasyurus ursinus),
the type of a new Family. By Frank EK. Bepparp,
M.A., D.Se., F.R.S., F.Z.S., Prosector to the Society.
(Rext-figs:(92— TOs) aie tel aescye be anGh « Juan) sc eee 677

Studies mm the Fossorial Wasps of the Family Scoliide,
Subfamilies Elidinze and Anthoboseine. By RowLanp
E. Turner, F.Z.8.,F.E.8. (Pls. LX XXI.-LXXXIIT.) 696

40.

41.

42.

43.

44,

46.

47.

48,

49,

vil
Notes on the Spanish Ibex. By ABEL CuApMAN, F.Z.S,

The Local Races of Burchell’s Zebra. By Major J.
STEVENSON-Hamrutron, C.M.Z.S., Game Warden of the
Mransvaails, (MextenesuO2— NO eecsceeemac sm necesdeesans

On Dipteropeltis, a new Genus of the Crustacean Order
Branchiura. By W. T. Cauman, D.Sc., F.Z.8. (PI.
TE EOCTN GA eee pete corso ested ss seid tsttnciser Sela el sl shite.

On two new Larval Trematodes from the Striped Snake
(Tropidonotus ordinatus sirtalis). By Wi1tu1AmM Nico,
M.A., D.Sec., M.D., F.Z.8., Lister Institute of Pre-
ventive Medicine, London. (Text-fig. 107.)

A Note on the rare British Nudibranch Wancochkia
eudactylota Gosse. By Sir Cuarues Exvior, K.C.M.G.,
C.B:, FA:S: (Pl UX XXyV-.)

Cece emer reer ee cesses eeeeresees

). The North Rhodesian Giraffe. By R. Lypekxkerr,

E.RS., F.Z.8. (Pl. LXXXV1)

On Antler-Growth in the Cervide, with special reference
to Hlaphurus and Odocoileus (Dorcelaphus). By R. I.
Pocock, F.R.S., F.L.S., F.Z.8., Superintendent of the
Gardens and Curator of Mammals. (Text-figs. 108-
| Iclizs) hes epi ipre aides ett OR Gt SR oe eae ane

Polycheta from the Pacific Coast of North America.—
Part I. SerPuLID#, with a Revised Table of Classi-
fication of the Genus Spirorbis. By Heten L. M.
PIxeLL, B.Sc., F.Z.S., Demonstrator of Zoology and
Reid Fellow, Bedford College, University of London.
(CRIS: XX VET EPNONGNGIGNG reece sate satclin vain ee

The One-sided Reduction of the Ovaries and Oviducts
in the Amniota, with Remarks on Mammalian Evo-

lution. By Hans Gavow, M.A., Ph.D., F.R.S., F.Z.8.

Contributions to the Anatomy and Systematic Arrange-
ment 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. By Frank E. Bepparp, M.A., D.Sc., F.R.S.,

765

767

770

“I
“
ww)

784

808

¥.Z.S., Prosector to the Society. (Text-figs. 113-121.) 822

vill
Page
50. On two new 'Trematode Parasites from the Indian Cobra.
By Wivxiam Nicout, M.A., D.Sc., M.D., F.Z.8., Lister
Institute of Preventive Medicine, London. (Text-
Pie M22 Vick bee « ceeeslenhiia lbs beds ood sabe ESE ae hea ER 851

Dl. Statistical Note on the Worm Parasites collected from
the Animals dying in the Zoological Gardens from
December 1910 till April 1912. By Witiiam Nico,
M.A., D.Sc., M.D., F.Z.S., Lister Institute of Pre-
ventive Mediemme ond om, tees -- cee esse eee Eee 856

52. On some new Fossil Reptiles from the Permian and
Triassic Beds of South Africa. By R. Broom, D.Sc.,
CAMEZ.S (PIs 2 XeC Xu NL ie aig. erase ee ae ee 859

53. On the Hydrocoralline Genus, Hrrina. By Professor
S. J. Hrcoxson, F.R.S., F.Z.8., The University of
Manchester, ails 72x liV/ xe (CiValle)) ens ee eeeee eee 876

D4. Descriptions of new Butterflies of the Genus Vhecla
from S.H. Brazil. By E. Duxryrretp Jonus, F.ZS.,
SES eGR OC NADL) 5 Cea eee eae a o aite sen ee cis Menem 896

Or
On

. The Bornean Bantin. By R. LypexKmr, F.RS., F.Z.S.
(Desst mies! Sh23 Ss): Jems a eae eee eee 902

56. Notes on the Breeding of the “ Millions” Fish (Gir-
ardinus pociloides). By Epwarp G. BouLENGER,

IB Ads, OUNENOIR OE IRONS Seo5--coscosney seceooudd suoesccos 906
Alphabetical List of Contributors ..............ccccccecececseee 1x
INGw Generic Herm (scucecak ieee eee Xvi

Aly HA Bir iC A Ly hast

OF THE

CON ERIE UTO Rs;

With References to the several Articles contributed by each.

(1912, pp. 505-913.)

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

to the Society.

Contributions to the Anatomy and Systematic Arrange-
ment of the Cestoidea.—IV. On a Species of Jnermi-
capsifer from the Hyrax, and on the Genera Zschokkeella,
Thysanotenia, and Hyracotenia. (Text-figs. 72-83.) ...

Contributions to the Anatomy and Systematic Arrange-
ment of the Cestoidea.x—V. On a New Genus (Dasyuro-
tenia) from the Tasmanian Devil (Dasyurus ursinus),
the type of anew Family. (Text-figs. 92-101.)............

Contributions to the Anatomy and Systematic Arrange-
ment of the Cestoideax—VI. On an Asexual Tapeworm
trom the Rodent, /iber zibethicus, showing a new form of
Asexual Propagation, and on the supposed Sexual Form.
Milessesfioe, 1) Si me ansebede adictamesid ated poasteeees ve atdesys

Page

Or
~I
=r)

x

Page
Bou.encsr, Epwarp G., F.Z.8., Curator of Reptiles.

Exhibition of a clay-ball containing a cocoon of the
African Lung-fish (Protopterws annecténs) .......1..cceseeee 807

Notes on the Breeding of the ‘“ Millions” Fish

(Girandinus pecciloides) Vem: terete ae. ee eee 906

BovuLENGER, GEorGcE A., F.R.S., F.Z.S.
On a Collection of Fishes made by Mr. A. Blayney
Percival in British East Africa to the Hast of Lake
Jeevan.) (Letise IbpO.C\7 INTL =U C50. Ce sac seaacoadceccec anon: 67

bo

: Notice of a paper entitled ‘‘Second Contribution to
our Knowledge of the Varieties of the Wall-Lizard” ... 807

Broom, Rospert, D.S8c., C.M.Z.8.
On some new Fossil Reptiles from the Permian and
Triassic Beds of South Africa. (Pls. XC.—-XCIIL.) ...... B59
Cauman, WiuiiAM T., D.Sc., F.Z.S.

On Dipteropeltis, a new Genus of the Crustacean Order

Isieenovolamtunieyn, (edly IUp-O-O-C EW caoasonnacnedossa=succode scone. 763

CHAPMAN, ABEL, F.Z.S.

INotessonkt hen Spam shill bexa merasseeea:aeceee eshte ee ee Rees 754

Dean, Basurorp, Ph.D.

Additional Notes on the Living Specimens of the
Australian Lung-fish (Ceratodus forsteri) in the Col-
lection of the Zoological Society of London. (Text-

HI GS18 40s OO) rnerreee ocean eer palpithaties duet seat Sone Bee 607

Exior, Sir Cuarues, K.C.M.G., C.B., F.Z.S.

A Note on the rare British Nudibranch Hancochia
eudactylota, Gosse.,, (Pl wa, \eccetr: (ucts oecenee etme 770

X1

Gapow, Hans, M.A., Ph.D., F.R.S., ¥.Z.S.

The One-sided Reduction of the Ovaries and Oviducts

in the Amniota, with Remarks on Mammalian Evolution.

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

TON, J.

Hickson, Professor SypNEY J.. D.Sc., F.R.S., F.Z.8., The
University of Manchester.

On the Hydrocoralline Genus, Hrrina. (Pls. XCIV.—
ECV EL ak creas tat anal ars ene setice te wae ence ae ate ce cine sea tel okt

Houxtey, Juuian S., Lecturer of Balliol College, Oxford.

First Account of the Courtship of the Redshank

LSTA TORS GTO CHATS) XS OS Re ACT SESE HOO MCG SEER MET SHE

Jones, E, DUKINFIELD, F.Z.S8., F.E.S.

Descriptions of new Butterflies of the Genus Thecla
Goren wie tore ziley g(t eG VA) ie ok vad. vinsola viene nue

KeENRICK, Sir GEorGE H., F.Z.S.

A List of Moths of the Family Pyralidee collected by
Felix B. Pratt and Charles B. Pratt in Dutch New
Guinea in 1909-10; with Descriptions of new Species.

(PEER VILE MMe ont) beeen veut coe) -)

Lover, Sir Epmunp G., Bart., V.P.Z.S.

Demonstration of the capacity of the electric lantern

presented by him to themsociety:* 17.21 0..0. oes. sce-e scsetaees

LybEKKER, RicHArp, F.R.S., F.Z.8.
The North Rhodesian Giraffe. (Pl. LXXXVI.) ......
The Bornean Bantin. (Text-figs. 123-125.) ............

?

Note stating that “ Gazella hayi” = Gazella fusci-

PARTON 3 5. os 3 x dame eetee nee ne ak Add a2 SaSb ES oan e ak tach?

Page

808

876

647

546

913

xii
Mrapvr-Watpo, Epmunp G. B., V.P.Z.S., and Others.

Discussion on the Preservation of the Native Fauna
ol Greatubribaniy t.c.6.caecees coe oben Soe See EERE ECE

Mrreueti, P. CHatmers, M.A., D.Sc., Hon. LL.D., F.RB.S.,
F.Z.S., Secretary to the Society.

Report on the Additions to the Society’s Menagerie
during the months of February and Mareh 1912 .........

Exhibition of a living specimen of a young female

Dorsal Hyrax (Dendrohyrax dorsalis) .........0..c10e0eeseees

Exhibition of photographs of an Elephant Kraal in
TIES 0 IRR ater RCPS Oe AIST Ie Oe SE, NN NR a
Report on the Additions to the Society’s Menagerie

Glureraey tla mnerTe Cie Ayorel WO - op ncasocessondasdesessoseoho-

Report on the Additions to the Society’s Menagerie
during the months of May, June, July, August, and
Septemiber WOW 2 ex. cc.cacc.os- yogeeanecenee tie occ cries eiaeetee

Nrcott, Witiam, M.A., D.Sc, M.D., EF.Z.S., Lister
Institute of Preventive Medicine, London.

On two new Larval Trematodes from the Striped
Snake (Z'ropidonotus ordinatus sirtalis). (Text-fig. 107.).
On two new Trematode Parasites from the Indian
Cobra. \(Mextatie V122)) 5 O00. erste oases cneere eee ene pee
Statistical Note on the Worm Parasites collected from

the Animals dying in the Zoological Gardens, from
December aI O tall Ayr GON Zens. o- eee eee cree ee eee

O’DonocHusE, Cuartrs H., B.Sc., F.Z.S., Assistant to the
Jodrell Professor of Zoology, University College,
London,

The Circulatory System of the Common Grass-Snake
(Tropidonotus natrix). (Pls. LXX-LXXIT. and Text-
HIgS, SO=911.)) 2. Sica sats a eeeeeceee belo eeee cs iaoece seer eee ee

Page

807

5D5

671

Giles

806

908

767

851

856

612

xiii
Percivat, A. Buayney, F.Z.S.

Exhibition of photographs and Jantern-slides of Game

Animals from British Hast Africa, ........ccccccsscscccccccce

Prxett, Miss Heten L. M., B.Sc., F.Z.S., Demonstrator
of Zoology and Reid Fellow, Bedford College,

University of London.
Polychzeta from the Pacific Coast of North America.—

Part I. SrrpuLip®, with a Revised Table of Classification
of the Genus Spirorbis. (Pls, LXXXVII.-LXXXIX.) .

Pocock, Reeimatp I., F.R.S., F.L.S., F.Z.8., Superintendent
of the Gardens and Curator of Mammals.

Exhibition of lantern-slide of two Polar Bear cubs

(Ursus maritimus) born in the Gardens ...............0s008

On a rare Stag (Cervus wallichii) from Nepal recently
presented to the Zoological Society by His Majesty King
Geurmomns (lextfs, GOs) oo cco. soensaciediasugvaonsocecsesas

Exhibition of a skin and a living specimen of a fawn
variety of the Brown Rat (Hpimus norvegicus) ............

On Antler-Growth in the Cervide, with special reference
to Hlaphurus and Odocoileus (Dorcelaphus). (Text-figs.

HOGE19.) 5 9. eae Se ese ee AG

Reaay, C. Tate, M.A., F.Z.5,

Exhibition of lantern-slides of a Flatfish (Platophrys
podas), showing its power of changing its colour and
FEAT CS ¢ «5 nntas ote ete meiner cafes Nantayioes Seniesa ee

Descriptions of new Fishes of the Family Loricariide
in the British Museum Collection. (Pls. LXXV.-
BRO SVL) «aie Oe eet See aa teres sass a unes sen yat tes oa,

Page

806

784

671

773

XIV

Page
Sera-Smrrn, Davin, F.Z.8., Curator of Birds.

Exhibition of lantern-slides of the display of the male
Peacock Pheasant (Polyplectron chinguis). Also of
photographs of the young Cariama cristata hatched and
reared siaythex(Crardens ay iG: aU XU1SXe) ene) eee ee 557

Exhibition of epidermal sheaths shed from the base of
the lower mandible of the King Penguin (Aptenodytes
VR LUTCULIO) WNG Betts SONAR AS GNSTRE On Gt CAC eee Bec tone ok 671

Exhibition of two living specimens of the Lory,
Cav atiaiens Sellonannks, 1OROTE TEN 6 gsenaoodsnoodonsecsobonono0a65pe 806

Sexton, Mrs. E. W., Marine Biological Laboratory,
Plymouth.

Some Brackish-water Amphipoda from the mouths

of the Weser and the Elbe, and from the Baltic.
(CEI) ©). Ma Bp. Gl ES) eB ace nciodngsdonbbaonode onROnton 656
SHANN, Epwarp W., B.Sc., Demonstrator and Assistant
Lecturer in Zoology in the Victoria University of
Manchester.

Observations on some Alcyonaria from Singapore, with
a brief Discussion on the Classification of the Family
INeplithyrdce se (Piss sXe XG) eee eee eee eee 505

SuHuFretpt, Roperr W., M.D., C.M.Z.S8.

Exhibitions of skins of, and remarks upon, two Virginia

Opessums | (ide) ohiseinoe tana) \....0e eee reee eee eee eee 556

SmitH, Davip Sera-. See SerH-Smrru, D.

Sreppine, The Rev. Tuomas R. R., M.A., F.RS., F.Z.8.

Notice of a Memoir on the Crustacea Isopoda collected
by the ‘ Porcupine’ Expedition in 1869-1870............64 912

xV

Page
Srevenson-Haminron, Major J., C.M.Z.8., Game Warden
of the Transvaal.
The Local Races of Burchell’s Zebra. (‘Text-figs. 102-
IRONS) 5 vv Sn Scolene Stace Me REN eater caisie Sect awreldnarectieesews nein’ 757
Tuomas, Mrs. Rose Hate, F.Z.S,
Experimental Pheasant ~- breeding. (Pls. LXIV.-
LDA BS) 8B te corn Seta ogc O Sip Bee ReOB EME OCOD Coca Rec ceEOUCIoC 539
Exhibition of the Eggs of Phasianus formosanus,
P. versicolor, and their F.1 and F-. 2 offspring. (‘Text-
ier MBLT)) aid sccnée hododebbe abypcagee GoupanbeennRdereasteneaceespccr 912
TuRNER, Rowand E., F.Z.8., F.ES.
Studies in the Fossorial Wasps of the Family Scoliide,
Subfamilies Elidine and Anthoboscine. (Pls. LX XXI.—
1 OU OFCTIO I) See heri sets socribo Abb 6. sue OBOE Ae EC Qo CER cach core Roe or cork 696
Watpo, E. G. B. Meaps-. See Meapr-WaAtpo, E. G. B.
Warp, Francis, M.D., F.Z.S.
Demonstration, illustrated by lantern-slides and the
cinematograph, of a method of observation of fishes, birds,
and maminals und erwabecy | cade. esac > dence aa eee 672

Wiruers, Tuomas H., F.G.S.

Some early Fossil Cirripedes of the Genus Scalpellum.
(Text-figs. 64 & 65.)

NEW

XVI

GENERIC TERMS

PROPOSED IN THE PRESENT VOLUME (pp. 505-913).

Page

Alopecorhinus (Reptilia) ...... 864
Braunsomeria (Insecta)......... 697
Brevicella (Insecta) ...........- 549
Dasyurotenia (Vermes) ......... 694
Dipteropeltis (Crustacea) ...... 768
Emydochampsa (Reptilia)...... 875
Emydops (Reptilia) ............ 871
860

aoesceteocsecce

Galeops (Reptilia)

Hyracotenia (Vermes)

Ictidepsis (Reptilia)

Protolzospira (Vermes)

Seymnognathus (Reptilia)......
Taurops (Reptilia) .............--

Urocystidium (Vermes)

Xenopharynx (Vermes)

INDEX

OF

SCIENTIFIG NAMES,

Acomus
erythrophthalmus, 909.
Acrochordus
Javanicus, 909.
Actinia, 519.
/®lurosaurus
Jelinus, 863.
striatidens,
875.
Agrotera
pyrosticta, 51,
Alce, 774, 778, 779.
Aleyonium, 518, 527.
Allopora, 892, 894.
Alopecorhinus, gen. noy.,
859, 864, 865.
parvidens, 859,
(2 oe
Aloposaurus, 862.
Ambia
novaguinensis, 546, 549,
5455)
Amia, 608.
Ammothea, 511.
virescens, 527.
Ammothée, 510, 511.
virescens, O11.

859, 863,

864

’

Amphilius
grandis, 675.
oxyrhinus, 672, 675,
676.
Ancistrodon

piscivorus, 633.
Andrya, 587.
Anoplocephala

hyracis hepatica, 593,

605.
Anoplotznia, 584, 603,

677.
dasyuri, 694, 846, 847.
Anthobosca, 696, 714,

724,

@ethiops, 745, TA8.

Anthobosea
albomacula, 725.
albomaculata, 728, 742, |

746, 753.
alhopilosa, 727, 732.
antennata, 728, 7438,
744, 746.
anthracina, 727,
754.
apicalis, 745.
arabica, 720, 728, 739, |
740.

733, |

argenteocincta, 727,
732. |
australasie, 725, T45— |
749. |
australis, 727-729, 745, |
754.
bidentata, 746, 752, |
TRB, |

bipunctata, 728, 742. |
bipustulata, 742, 746, |
753. |
carbonaria, (28, 7A4l, |
742, 745.
chilensis, 728, 741, 746.
elypeata, 727, 735, 751,
753, 754.
cognata, "727, 731.
crabroniformis, 746.
erassicornis, 736, 745,
(il, (Be |
donaldsoni, 728, 737.
errans, 746, 751.
erythronota, 728, 737,
739, 746, 755, 754.
erythropyga, 728, 744,
Fi £

Fasciculata, 736.

Jastuosa, 727, 729, 7538.

flavicornis, 727, 730,
731, 732.

flavopicta, 746, 752.

Anthobosea
Frenchi, '745, 749, 750.
gilest, 745, 747.
inornata, 736.
insularis, %28, ‘37,
754,
jhering?, 744.
lacteipennis, "746, 752,
753.
levifrons, 727,
734,
lagardei, 745, 750.
leucospila, 739.
longipalpa, 745, 750.
melanaria, 728, 736.
minima, 728, TAO.
moderata, ‘Ad,
753.
natalica, 728,738,739.
nigra, T45, TA8.
nigripennis, 745.
nubilipennis, 727, 730,
(30.
occipitalis, 696, 727,
734,
sauakinensis, 728,
signata, 727, 729.
strandi, 727, 731.
torresensts, 745, 747.
unicolor, 727, 733, 734.
varipes, 745, 747, 748.
Apenesia, 699.
Aphrocallistes
whiteavesianus, 787.
Aphytoceros
grosalis, Hd.
Apomatus, 788, 793.
timsii, 784, 787, 805.
Aporina, 849,
Aptenodytes
pennanti, 671.
Ara
severa, 806,

733,

749,

740.

xvlll

Araucaria, 726.
Arboricola

charltoni, 909.
Arcosealpellum, 588, 539.
Arctictis

binturong, 909.
Arctogalidia

stigmatica, 909.
Arges

cirratus, 666, 670.
Argulus, 765.
Argusianus

argus, 909.
Arvicola

arvalis, 859.
Asellus

aquaticus, 660.
Astacus, 519.
Aulacodes

brunnealis, 548.

metaloxalis, 548.
Austrotiphia

kirbyi, 730.
Avitellina, 848.

Bagrus
urostigma, 675.

Baleeniceps
rex, 909.

Balanus
nubilus, 798, 800.

Barbus
argyrotenia, 672, 674,

676.
erlangeri, 674.
mimus, 672, 674, 676."
nairobiensis, 674. i
neglectus, 675.
percivali, 674.
zanzibaricus, 674.

Belemnitella
mucronata, 530.

Bertiella, 677, 694, 844.

Bison
bonasus, 908.

Boa, 621, 643.
constrictor, 628, 634.
diviniloqua, 628, 638,

634.
reticulata, 644.

Bos

sondaicus, 902, 903,906.
— birmanicus, 906.

— lowi, 904-906.
— porteri, 906.

Braunsomeria, gen. nov.,

696, 697, 700,
atriceps, 696, 699.
quadraticeps, 696, 699-

758, 754,

Brevicella, gen. nov., 546,
549.
emarginata, 546, 549,
55D.
Bubalus
caffer, 910.

Calamachrous
albipunctalis, 546, 554,
555.
Calliobothrium, 693.
Callionymus
lyra, 908.
Calliptilus
solitarius, 806.
Callosila, 726.
Camelus
dromedarius, © 06.
Canis
Jamiliaris, 556.
Capra
pyrenaica, T54.
— victoria, 756.
Capreolus, 774, 778.
capreolus, 910.
Caprinia
diaphanalis, 551.
Cariacus, 774.
Cariama
cristata, 557, 558.
Carijoa
rupicola, 509.
Catreus
wallichi, 908.
Causus
rhombeatus, 634.
Cerastes
vipera, 622.
Ceratodus
forsteri, 607-612.
Cercaria
ordinata, 767, 768, 769.
Cercopithecus, 606.
Cervus, 778.
affinis, 564-575, 781.
albirostris, 558, 574.
aristotelis, 908.
canadensis, 574, 774,
CUD, TOH
— wardi, 575.
cashmeriensis, 565, 566.
— macneilli, 570.
duvaucelli, 783, 908.
elaphus, 574.
eldi, 776, '780.
hanglu, 565-575, 775,
aitite

kansuensis, 574.
macnetlli, 571-574.
porcinus, 908.

INDEX OF SCIENTIFIC NAMES.

Cervus
pygargus, 559.
thoroldi, 574.
wallichi, 556-575.
canthopygus, 575.
Cheetostomus
Jischeri, 667.
lepturus, 666, 667, 670.
marginatus, 667.
palmeri, 666, 667, 670.
paucispinis, 666, 667,
670.
Chapmannia, 600.
lapica, 603.
Chelys
Jimbriata, 911.
Chimarrhornis
leucocephala, 556.
Chironephthya
variabilis, 523, 524.
Chitinopoma
fabricii, 790.
greenlandica, 790, 805.
Chloéphaga
poltocephala, 556.
Chonopeltis, 765.
inermis, 766.
Chrysomitris
uropygialis, 910.
Circeis
armoricana, 796, 797.
Cissopis
leveriana, 911.
Cistecephalus
arctatus, S71.
nicrorhinus, 871.
Cittoteenia, 604, 677.
Clarias
lazera, 675.
Clarotes
laticeps, 675.
Clupeosoma
polusalis, 548.
Ceelopeltis
monspessulana, 617.
Colobosila, 726.
Coiobus
abyssinicus occidentalis,

vellerosus, 910.
Coluber
esculapit,
634.
corais, 638.
natrix, 613.
Connocheetes
gnu, 911.
Corallus
caninus, 645.
cookit, 628, 648.
madagascariensis, 643,

633,

622,

Coris
Julis, 908.
Coronella
catenifer, 626,
getula, 617.
levis, 622.
Corophium
acutum, 664.

lacustre, 656, 658, 664,

665.
triaenonyx, 664.
volutator, 658.
Cosila, 724, 725.
apicalis, 745.
argenteocincta, 732.
australis, "728.
biguitata, 729.
carbonaria, 741.
chilensis, 725, 741.
donaldsonti, 705, 737.
erythropygia, 744.
inornata, 736.
énsularis, 737.
Jjheringi, 743.
(Callosila) australis,
728.
(—) fasciculata, 786.
—) flavicornis, 730.
(—) minuta, 734.
(—) signata, 729.
Crocidolomia
binotalis, 553.
Crucigera
irreqularis, 786, 805.
zygophora, 786, 805.
Cryptohelia, 885.
Curena
externalis, 548,
Curicta
opposttalis, 554.
Cyanocorax
affinis, 806.
Cyclorchis, 690.
Cynognathus, 874.
Cysticercus
Jasciolaris, 838, 850.
longicollis, 839, 840.

Dama, 777.
dama, 781.

mesopotamica, 781.

Dasyurotenia, gen. nov.,
677, 680, 683, 694,

695, 823.

robusta, 677, 678, 679,

684, 694, 854.
Dasyurus
ursinus, 677.
Dausara
amethystina, 553.

INDEX OF SCIENTIFIC NAMES.

Davainea, 591, 592,
606.
Dendrohyrax |
dorsalis, 671.
Dendrolagus
ursinus, 806.
Dendronephthya,
510, 518, 521
disciformis, 506, 519,
520, 527.
Diadectes, 860.
Dizlurodon
whaitsi, 868.
Dicrocelium, 854.
infidum, 834, 856.
Dicrogenium, 725.
Dicynodon, 871, 872
laticeps, 859, 876.
leoniceps, 869.
lutriceps, 859, 868, 870,
876.

505,

psittacops, 859, 869,
876.

tigriceps, 868.
Didelphis
virginiana, 556.
Dilepis, 847.
Dimorphoptera, 724,
25.
anthracina, 733.
clypeata, 735.
cognata, 731.
Jastuosa, 729.
levifrons, 734.
morosa, 729.
nigripennis, 728.
sabulosa, 733.
scoliiformis, 728.
signata, 729.
unicolor, 734.
Dioicocestus, 844, 848.
acotylus, 831.
Diplostomum
sirtale, 767, 768, 769.
Dipteropeltis, gen. nov.,
765.

hirundo, 763, 766.
Dipylidium, O87.
Discognathus

dembeensis, 673.
Dissemurus

paradiseus, 556.
Distichopora, 892, 894.
Distoma, 856.

Dolops

longicauda, '768, 765.

Dorcephalus, 773, 778.

Hecasaurus
priscus, 875. }

Proc. Zoot, Soc.—1912, No. LXI.

| Kehidna, 814.

Kehinococeus, 837.
Eirone, 699, 725.
Elaphurus, 773, 778, 780,
783.
davidianus,
7177, 779.
Elapbus
indicus, 909,
Elis, 725.
alicia, 737.
apimacula, 721.
capitata, 711.
combusta, 724, 754,
dimidiata, 716.
dubia, 721.
ephippium, 724.
Jedtschenkvi, 721.
major, 696, 723.
sellowt, 696, 722.
tricolor, 721.
xanthocera, 709.
(Mesa) abdominalis,
703, 707.
(— 7 Ee, 703,

113, 774,

a ee 696,
704, 753, 754.

(—) ametalla,
713.

703,
704,

(—) apicipennis, 696,
703, 70

(—) apimacula, 714,
719.

(—) asmarensis, 704,

(—) auriflua, 696, 703,
705, 706

(—) bengalensis,
716.

(—) burmanica, 714,

(22.

714,

(—) capensis, 709.
(—) capitatu, 704,
710

(—) carbonaria, 718.

(—) Pa: 714,
TG 7

(—) Hie 713.

(—) diapherogamia,
706

(—) dimidiata, 713,
714, 715.

(—) dimidiaticornis,
714, 720, 753.

(—) dubia, 714, 719.

(— oY erythrapeda, 703,

as. pba 714, 721.
(— ) fedtschenkoi, 714,
719.

61

XX

Elis
(Mesa) fuscipennis, 713,
717.

(—) heterogamia, 704,
706

(—) ee 703,
708.
ioe) porn, 704, 7

Si incerta 696, 704,

(— ee, 713.
fre ee 703,

me a 714, 721.

(—) longiventris, 696,
704, 712, 753, 754.

(—) madalensis, 718,
714, 715, 721.

(—) mandibulari is, 713,
714, 7

ee ee 704, 707,

713.

(—) nursei, 714, 721.

(—) opacifrons, 714,
719.

cea Gs ingueyt, 703,

Ss oat 713,
714, 717-719, 721.

(—) picticollis, 714,

(—) pyxidata, "103,
707.

(—) reticulata, 713.

(—) rothneyi, 714,

rlyfe
(—) ruficeps, 704, 706,
754.

=) = agers,
704, 706
eae — diapher ogamia,

(— ae = ruficeps, 704.

(—) rufo- <femorata, 713.
(—) sausswrei, 703,
(=

ee
) spoliata, 696, 704,

rials

(—) torrida, 703, 708:

(—) tricolor, 714, 720,

753, 754.

(—) ustulata, 714, 718.

(—) xanthocera, 703,

709.

Emydochampsa, gen.
noy., 875.

platyceps, 875.

- Emydops, gen. noy., 859,

871.

7
minor, 859, 871, 876. ,

INDEX OF SCIENTIFIC NAMES.

Endothiodon, 871.
bathystoma, 867, 875.
platyceps, 859, 867,

75, 876

Endothiodon
whaitst, 859, 866, 867,
875, 876.
Entephria
glauctas, 550.
grisealis, 546, 550, €

Epimys
norvegicus, 671.
Equus
burchellt, 757.
— chapmanni, 757.
— selousi, 757.

760.
— wahlbergi, 757.
kiang, 908.
Krithacus
komadori, 910.
Errina
antarctica, 894.
capensis, 894.
carinata. 878, 880,
893.
cochleata, 880, 898,
894.

echinata, 880, 881,
893.

fissurata, 878, 880,
887, 894.

ee 878, 880, 895,
894.

gracilis, 878, 881, 894.
horrida, 878, 880,

labiata, 878, 880,
888, 893.

macrogastra, 878, 879,

— moseleyi, 894.

nove zealandie, 876,
877, 894.

pourtalesti, 880, 881,
893.

ranosa, 878-881, 893.

{Labiopora) antare-
tica, 881, 8&2, 887,
890.

(—) aspera, 881, 888-
891, 894, 895.

(—) capensis, 878, 886,
890-893. 895, 896. |

(—) gracilis, 889, 890,

(—) moseleyi, 881, 887.
888, 890.

wniseries, 866, 867, 875.

555.
mioswari, 546, 550, 555.

— transvaalensis, 757,

dabneyt, 880 893. 894.

Errina
(Labiopora) nove ze-
landie, 882-884,
889-893, 895, 896.
(Spinipora) echinata,
881, 895
Erythrura
pealei, 911.
Hryx
conicus, 634.
Jaculus, 633, 634.
Johni, 633, 634.
Hsoterodon
uniseries, 875.
whaitsi, 875.
Eunectes
marina, 854, 856.
murinus, 628, 631,
634.
noteus, 634.
Eupodotis
edwardst, 911.
Eurytrema, 854.
Kutropius
depressirostris, 6795.

Felis
nebulosus, 909.
onea, 555, 806.
pardus, 909.
tigris, 909.
uncia, 509.

Fiber
etbethicus, 822, 83

840.

Galechirus, 861.
Galeops, gen. nov., 859,
860, 861.
whaitsi, 859, 860, 87d.
Galepus, 861.
Galesaurus, 874.
Gallicrex
cinerea, 909.
Gammarus
duebenti, 657, 661.
locusta, 657, 661-665,
pulex, 657, 658, 660,
661.
zaddachi, 656-658,
660, 661, 663, 663.
Gavialis
gangeticus, 909.
Gazella
fuscifrons, 911, 912.
hayt, 911, 912.
tsabella, 912.
pelzelni, 911.
Geocichla
cyanonotus, 910.

Geotiphia, 724, 726.
Giratta
camelopardalis
quorum, 911.
— vrothschildi, ‘771,
GI
— schillingsi, 772.
— thornicrofti, 771,
772, 773.
—- tippelskirchi, 771,
772.
Girardinus
peciloides, 906.
Gly phodes
cesalis, 552, 53.
magnificalis, 046, 552,

anti-

599.

nigricincta, 546, 582,
555.

pheiffere, 546, 554,
599.

polyzonalis, 552.

pseudocesalis, 546, 552, |

598, 900.

Gorgonia
suberosa, 524.
Gorgonops

torvus, 861.
Govia, 770.,
Graphiurus

spurrelli, 555,
Gypaétus

barbatus, 806.

Hancockia
eudactylota, 770.
Haplochilus
sexfasciatus, 911,
Hemidactylus
bowringti, 909.
trenatus, 909.
leschenaultii, 909.
Hemistomum, 767, 770.
Hemitragus
Jemlaicus, 556.
Hesperornis, 819,
Holothuria, 519.
Hyena
hyena, 555.
Hyalopomatopsis, 792.
langerhansi, 791.
marenzlleeri, 791.
occidentalis, 791.
Hydrocheerus
hydrocherus, 911.
Hydropotes, 774.
Hymenolepis
acuta, 687.
murina, 822.
reticulata, 603.
villasus, 831.

INDEX OF SCIENTIFIC NAMES,

Hyracotienia, gen. noy.,
576, 579, 595, 604.
capensis, 576.
hyracis, 576, 594, 596,
597, 601, 605.
procavie, 604, 605.
Hyroides
dianthus, 804.
gronlandica, 790.
norvegica gronlandica,
790.

Ieterus
vanthornus, 806.
Tetidvgnathus
hemburyi, 859, 865,
875.
parvidens, 865, 866.
Ictidopsis, gen. nov., 859,
872, 874.
elegans, 859, 872, 873,
376.

Ictidosuchus, 861.
Inermicapsifer, 581, 582,
590, 59-4, 595, 599,
603, 607, 688, 692,
847.
sp., 593, 605.
capensis, 576, 577, 580,
586, 588, 591, 592,
593, 600, 602, 606.
criticus, 598.
hyracis, 577, 578, 579,
585, 586, 587, 588,
591, 592, 6U6.
interpositus, 592.
pagenstecheri, 593.
settii, DTT, 578, 592,
606.
Iswara, 715.

Jassa, 656,

Kachuga
dhongoka, 909.

Kistecephalus
arctatus, 871.

Labeo
cylindricus, 673.
neumanni, 673.
percivali, 672, 673,
676,

| Labiopora, 876, 879, 88),

881. 884, 88d.

antarctica, 887, 890, |

803, 8U4.

Labiopora
moseleyi, 878, 888, 893,
894.

Lacerta, 614, 626, 627,
631, 644.

muralis, 643, 807.
Lachesis

gramineus, 626.
Leospira, 793, 799.
Lama

pacos, bab.
Lanius

cristatus, 910.
Larus

hemprichi, 910.
Leptocheirus

pulosus, 66, 658.
Ligula, 824,
Linstowia, 677.
Lioheterodon

madagascariensis, 634,
Lithophyton

arboreum, 511,
Lithophytum, 511, 513.

arboreum, 511, 517,

a27.

Lobophytum, 527.
Lomanotus

genet, 770,
Lophophorus

impeyanus, 908.
Lophortyx

douglasi, 911.
Lycosuchus, 862.
Lystrosaurus, 873, 874.

Macealla

arruensis, 546, 547,
55d.

mioswart, 546, 547,

olivalis, 546, 547, 555,

syrichtusalis, 547.
Madrepora, 519.
Margarosticha

plumbealis, 546, 549,

O00.

Maurillus, 725.
Mazama, 778.

bricenti, 783,
Meleagris

ocellata, 909.
Melitodes

albitincta, 506, 525.
Mera

pusilla, 797,

Meria
abdominalis, 703.
Merops

apiaster, 911,

XXil

Millepora
aspera, 879, 881.
Moniezia, 604, 677, 692,
695.
Mormyrops
deliciosus, 673.
Mormyrus
kannume, 673.
Motacilla
citreola, 910.
Mungos
brachyurus, 909.
ichneumon, 909.
Myiarchus
tyrannulus, 911.
Myzine, 697, 725.
abdominalis, 696, 703,
754.
albomaculata. 742.
anthracina, 714, 733.
apimacula, 719.
bipunctata, 742.
bengalensis, 716,
braunsi, 696, 700, 753,
754.
burmanica, 722.
capitatd, 710.
carbonaria, TA1.
ceylonica, 717.
claripennis, 718.
clavata, 713.

combusta, 696, 714,
724.
constrictiventris, 696,
701, 754.

dimidiata, 715.
dimidiaticornis, 720.
erythropyga, 744.
Jfastwosa, M29:
flavicornis, 741.
fuscipennis, 717.
hortata, 718.
insularis, 737.
leta, 721.
lecointei, 742.
levifrons, 734.
madraspatana, 7109.
nodosa, 713.
orientalis, 714.
pallida, 714.
petiolata, 717.
rothneyt, 717.
ruficeps, T06.
rufifrons, 702.
sabulosa, 138.
sauakinensis,
740.
signata, 729.
stigma, 696, 699, 753,
TA.
swalei, 700, 701.

696,

Myzine
tricolor, 720.
umbratica, 696, 702.
unicolor, 734.
violaceipennis, 115.
xanthocera, 709.

Nacoleia

perdentalis, 551.
Naia

melanoleuca, 910.
Naja

tripudians, 851.
Neochmia

phacton, 911.
Neogensis

flavoplagialis, 548.
Nephthée, 510, 511.
Nephthya, 510, 512, 513,

519

albida, 516.
bed fordi, 505, 506, 514,
516, 517, 527.
celosa, D7.
chabrolii, 511, 512, 517,
518, 527.
cordierv, 511.
elongata, 516.
Noorda
arfakensis, 546, 553,
555.
nigripunctalis, 553.
Nototragus
melanotis, 909.
Numida
ptilorhyncha,
605.
Nursea, 725.
Nymphula
chryseis, 548.
polystictalis, 548.
Nythosaurus, 873.
brownt, 859, 874,
876.
larvatus, 872, 874.

576,

Odocoileus
sp., 773, 774, 778, 780,
2.
americanus, 783.
hemionus, 788.
savannarum, 781.
Odontothynnus, 696,
724.
bidentatus, 752.
lacteipennis, 752.
Omphisa
repetitalis, 593.

INDEX OF SCIENTIFIC NAMES.

Omphisa
variegata,
555.
Oochoristica, 677, 847.
Ophiocephalus
punctatus, 911.
Ophiophagus
bungarus, 617, 625.
Opisthoctenodon, 859,
868.
Ornithorhynchus,

546, 553,

814,

Otiditenia, 534, 590,
591

eupodotidis, 583.
Otocinclus
maculipinmis, 666, 668,
670.
nigricauda, 668.
perforatus, 668.
Oudenodon, 859, 872.
bolorhinus, 868.
magnus, 866.
Ovis
burrhel, 556.
Oxyloricaria
leightont,
670.
tamané, 666, 669, 670.

666, 669,

Pagyda
botydalis, 551.
jumosa, 546, 5o1,
555.

Paradexiospira, 793.
Paralospira, 799.
Paraspongodes
crassa, 506, 523.
Pareiasaurus, 860, 861,
863, 869.
Pelea
capreolus, 910.
Pelias
berus, 643.
Pelophilus
madagascariensis, 612,
633.
Peltidocotyle, 694.
Perameles, 819.
Perisyntrocha
suffusa, 546, 548, 555.
Petrophila
erythrogastra, 909.
Phasianus
Jormosus, 540,541,545,
912, 913.
genneus, 540.
torquatus, 540.
versicolor, 540-542,
545, 546, 912, 913.

Phryganodes
analis, 551.
centrabalis, 551.
erebusalis, 551.
tetraplagalis, 551.
usitalis, d51.
Phyllobothrium, 694,
Piletocera
diplatyla, 548.
ayimisalis, 548.
torsicutalis, 548.
Pilocrocis
angulifera, 546, 553,
990.
Platophrys
podas, 056.
Platynosomum, 854.
Plecostomus
honde, 666, 670.
Plerocercus, 839.
Plesia
abdominalis, TOT.
asmarensis, 712.
carbonaria, 710, 713.
continua, 696, 703.
dubia, 719.
erythronota, 696, 738.
fedtschenkoi, 719.
incisa, 712, 713.
leucospila, 696, 739.
mandalensis, 716.
melanaria, 696, 736.
nurset, 721.
picticollis, 720.
reticulata, 713.
rufo-femorata, 713.
tartara, 719.
(Mesa) abdominalis,
707.
(—) adelogamia, 706.
(—) atopogamia, 706.
(—) diapheroyamia,
706.
(—) disjuncta, 706.
(—) erythrepoda, 709.
(—) heterogamia, 706.
(—) hottentota, 708.
(—) hova, 707.
(-—) innotata, 709.
(—) mandibularis, 715.
(—) opacifrons, 719.
(—) peringueyi, 708.
(—) purpureipennis,
715.
(—) saussurei, 709.
Pecilotiphia
albomaculata, 719.
Pollicipes, 530, 537.
Polyplectron
bicalcaratum, 909,
chinguis, 557.

{ Polypterus
senegalus, 910.
Porella
antarctica, 881, 887.
Pristerodon, 859, 868.
Pristerognathus
platyrhinus, 859, 863, |
864, 875.
polyodon, 864.
Procayvia
capensis, 576, 593.
Procolophon, 874.
Prodicynodon
beaufortensis, 859, 867,
376.
pearstonensis, 868.
Proptopterus
annectens, 807.
Prorodes
mimica, 551.
Prospirorbis, 792.
Proterogynotenia, 685.
Protolwospira, subgen.
noy., 784, 792, 7ys.
Protula
atypha, 790.
capensis, 790.
diomedee, (90.

superba, T9U.
Pseudomeria, 699.
Pseudotantalus

ibis, 556.
Psilacabaria

gracillima, 506, 529.
Pygospila

tyres, 53.
Pyrausta

alentialis, 554.

deductalis, 554.

flammealis, 546, 554,

5d5.

occultilinea, 554,
Python

molurus, 638.

reticulatus, 806.

regius, 628.

sebe@, 633. 634,

spilotes, 626.

Rangifer, 774, 778.
Rehimena

cissophora, 551.
Rhinoceros

unicornis, 908.
Romanchella, 793.
Rucervus, 778.

duvaucelli, 776, 780.
Rusa, 776, 778, 78.

aristotelis, 775, 78.

INDEX OF SCIENTIFIC NAMES.

|

XXill

Sameodes
polythliptalis, 553.
Sarcophytum, 527.
Scalpellum
accumulatum, 528-530,
537-539.
arcuatum, 528-531.
comptum, 537, 538.
darwinianum, 530,
535.
Jossula, 531, 583, 537,
538, 589.
gallicum, 538.
hastatum, 530, 589.
maximum, d31.
— ciylindraceum,
538, 539.
simplex,
588, 539.
solidulum,
538, 539.
trilineatum, 528,
531, 538, 539.

530,
528, 530,

530,

. Sceliodes

grisealis,
505.
Sclerogorgia
suberosa, 524.

546,

pacifica, 784, 785, 805. | Sclerophytum, 509.

palinatum, 507, 508.
pinnulatum, 505, 506,
507, 508, 527.
viride, 508.
Scylacosaurus, 861, 864.
Scymnognathus, gen.
noy., 859, 861, 862.
whaitsi, 859, 861, 875.
Seymnosaurus, 863.
Semnopithecus
pileatus, 806.
Serpula
columbiana, 784, '785,
786, 801, 802, 805.
jukesui, 786.
spirillum, '796.
splendens, 784,
786.
triqueter, 780.
vitrea, 799.
zygophora, 786.
Sierolomorpha, 725.
Siphonogorgia
variabilis, 506, 528.
Sphenocercus
sphenurus, 908.
Sphenodon, 6438.
Spinipora, 876, 878, 879,
880, 881, 891.
Spirorbis
affinis, 802.
aggregatus, 795.

785,

XX1V

Spirorbis

ambilateralis, 784,792,
795, 798, 799, 805.

argutus, 795.

armoricanus, %94,
197.

bellulus, 794.

beneti, 795.

bernard, 795.

borealis, 796, 804.

cancellatus, 792, 794,
796.

carinatus, 802.

claparedei, 795.

comptus, 794.

cornu-arietis, 795, 798.

corrugatus, 794,

evolutus, 793.

Jfabricii, 802.

Joraminosus, 794.

Sformosus, 794.

granulatus, 802.

hoehleri, 795.

levis, 795.

langerhansi, 795, 801, |
KR

805.
lebrunt, 795.
levinsent, 795,
lucidus, (96.
malardi, 795,
marioni, 794.
mediterrancus, 795.
medius, 784, 793, 795,
800, 805.
militaris, 795.
morcht, 795, 801, 805.
pagenstecheri, 794.
patagonicus, 795.
perriert, 795.
pseudocorrugatus, 794.
pusilla, 794, 797.
pusilloides, 794, 797,
800.
quadrangularis, %95,
802,

racemosus, 784, 795, |

798, 799, 805.
rugatus, 794.
semidentatus, 794.
similis, 795.
spirillum, 794, 796,

797, 805.
sptirorbis, 795.
validus, 795, 800, 805.
variabilis, 795, 802.
verruca, 793, 795,

800.
violaceus, 794.
vitreus, 793, 794, 805.
(Dexiospira) spirilluiz,

796.

€pirorbis

(Leospira) asperatus,
800.

(—) langerhansi, 801.
(—) medius, 800.
(—) morchi, 801.

(—) quadrangularis,
02

(—) validus, 800.

(—) variabilis, 802.

(Paradexiospira)
vitreus, 793.

(Protoleospira) ambi-

lateralis, 798.
Spizaétus
limnaétus, 910.

Spongodes, 510, 511,

518, 519, 520, 521.
celosia, 512, 513, 518.

nephthyeformis, 612,
418

Sponodia, 518, 520, 521.

_ Stereonephthya, 510, 518,

519, 520.

lutea, 505, 506, 521, |

523, 527.
Stericta

rurealis, 546, 6548,

555.

Styphlodora
bascaniensis, 855, 856.
condita, 855, 856.
horrida, 851, 854.
naje, 851, 852, 854.
serrata, 854, 85d.
similis, 855.

Suberogorgia
suberosa, 506, 524.

Sylepta

dinawa, 546, 551, 555.

leucodonta, 551.
polydonta, 551.

| Synodontis

geledensis, 675,

schall, 679.
Syrnium

uralense, 909.

Tachypterus
crassicornis, 751.

Tenia
crassiceps, 839.
crassicollis, 838.
ghondhorensis, 593.
paronat, 593.
spatula, 593.
(Anoplocephala),

gondokorensis, 605.

(—) spatula, 605,

INDEX OF SCIENTIFIC NAMES,

‘Taognathus —

megalodon, 867.
Tapinocephalus
atherstonei, 859.
Tapirus
indicus, 909.
Taurops, gen. nov., 859.
macrodon, 859, 875.
Taurotragus
oryx, 806.
Telesto
prolifera, 509, 510.
riser, 509.
rupicola, 506, 509,
510.
(Carijoa) rupicola,
509.

Tetrabothrium, 58d,
Thamnobia
cambaiensis, 909.
Thaumalea
amhersti, 544.
obscwra, 544.
picta, 544.
Thecla
bertha, 897, 902.
biblia, 901.
campa, 901, 902.
castrena, 900, 902.
datitia, 901, 902.
esmeralda, 900, 902.
fancia, 897, 902.
JSernanda, 899, 902.
guina, 898.
gispa, 896.
hamita, 896, 902.
janias, 900.
Japola, 898, 902.
lemona, 899.
molena, 899, 902.
nora, 899, 902.
nubilum, 899.
phrosine, 900.
punctatum, 897.
schausa, 898, 902.
sicrana, 897, 902.
tadita, 901.
Thelognathus, 874.
Thrasaétus
harpyia, 806.
Thynnus, 725.
antennatus, 743.
cathreinii, 746.
jischeri, 747.
reischit, 748,
stolzii, 748.
Thysanotenia, 576, 584,
607.
gambiana, 579, 581,
582, 583, 585, 586,
d87, 590, 602, 606.

INDEX OF SCIENTIFIC NAMES,

Thysanotenia
lemuris, 583, 585, 606,
686.
Tilapia
alealiva, 676.
grahami, 676,
nilotica, O75.
percivali, 672, 676.
Tiphia
capensis, 709.
bipunctata, 742.
Totanus
calidris, 647.
Toxostoma
cinereum, 910.
Triplotenia, 677.
Trochalopterum
morrisonianum, 910.
Tropidonotus
Jasciatus, 626,
633.
natirix, 612, 613, 632,
634, 640, 643, 646,
647.
ordinatus sirtalis, 767.

624,

| Tropidonotus

tesselatus, 622.
Tympanuchus
americanus, 556,

Uroeystidium, gen. nov.,
840,
gemmiparum, 822, 824,
826, 828, 833, 840,
841.
Ursus
maritimus, 556.

Vipera
berus, 622.
russelli, 910.
Viverra
civetta, 909.

Wrightella, 526.
robusta 505, 506, 525,
527.

XXV
Xanthomelxna
schematias, 550.
Xenia, 509,
Xenocara
bufonium, 668.
heterorhynchus, 666,

668, 670.
multispinis, 666, 6638,
670.
stigmaticum, 668.
Xenopharynx, gen. noy.,
351.
solus, 851, 852.

Zatnenis
Fasciolatus, 909.
gemonensis, 681, 633,
634.
Zinckenia
lophoceralis, 551.
Zschokkea, 590.
Zschokkeella, 576, 591,
602, 603, 606, 607,
688, 692.
linstowi, 590,

INDEX

OF

ILLUSTRATIONS

Allurosaurus striatidens, Pl. XCI.
859.

Alopecorhinus parvidens, Pl. XCI.

Ambia novaguinensis, Pl. LXVITII.

p. 546.

Ammothea virescens, Pl. LXI. p. 505.

MTB TES oxyrhinus, Pl. TEXXIX.
p. 672.

A ipehesr albomaculata, Pl. LXXXI.

p. 696.

anthracina, Pl, UXXXII. p. 696.

australasie, Pl. LXXXTII. p. 696.

— australis, Pls. LUXXXIL.,
LXXXIIL., p. 696.

clypeata, Pls. LXXXI.-LXXXIII.,

p. 696.
-—— cerassicornis, P]. LXXXI. p. 696.
erythronota, Pls. LXXXI.,
LXXXIL., p. 696.
fastuosa, Pl. XXXL p . 696.
-—— insularis, Pls. LXXXIL,
wOuuan, p- 696.
moderata, P\. UXXXT. p. 696.

Apomatus timsii, Pl. LXXXVII. p. 784.

Barbus argyrotenia, P|, UXXIX., p, 672.
mimus, Pl. LX XIX. p. 672.
Bos sondaicus, Fig. 123, p. 903.
-— lowit, Figs. 124, 125, pp. 904,
905.
Braunsomeria quadraticeps,
Pls. LXXXI., LXXXIL., p. 696.

Brevicella emarginata, Pl. LXVIII.
p- 546
Calamachrous albipunctalis (var.),

Pl. LXVIII. p. 546.
Cariama cristata, Pl. LXIX. p. 546.

|

Ree: forsteri, Figs. 84, 85, pp. 610,

61

Cercaria ordinata, Fig. 107, p. 768.

Cervus, Fig. 110, p 778,

-—— canadensis, Hie. OSS, jos 7G:

hanglu, Figs. 70, 108, pp. 566, 775

—— macneilli, Fig. 71, p. 571.

-— waillichii, Figs. 66-69, pp. 559,

561, 563, 565.

Chetostomus lepturus,
p- 666.

—— palmeri, Pl. LXXV. p. 666.

—— paucispinis, P]. LXXV. p. 666.

Chitinopoma greenlandica, PA.
LXXXVIII. p. 784.

Corophium lacustre, Pl. LXXIV. p. 656.

Crucigera irreguaris, Pl. LXXXVII.

784.
zs zygophora, Pl. LXXXVII. p. 784.

Pl. LXXYV.

Dasyurotenia robusta, Figs. 92-101,
pp. 677-679, 681, 682, 684, 686, 688,
630, 691.
Dendronephthya disci formis, Pl. LXITT.
p. 505.

Dicynodon laticeps, Pl. XCII, p

lutriceps, Pl. XCII. p. & 859,

psittacops, P|. XCII. p. 859.

Diplostomum sirtale, Fig. 107, p. 768.

Dipteropeltis hirundo, Pl. LXXXIV.
p- 763.

. 859.

Eccasaurus priscus, Pl. XC. p. 859.
Elaphurus, Fig. 110, p. 778.
davidianus, Fig. 111, p. 779
Elis combusta, Pl. LXXXIL. p
(Mesa) alicie, Pls.
LXXXIII., p. 696.

696.
TXXXL. >

INDEX OF ILLUSTRATIONS.

Elis (Mesa) dimidiaticornis, P|, LXXX1.
p- 696.
) longiventris, Pls. LXXXL., |
LXXXIII., p. 696.
) ruficeps, Pls. LXXXIL.,
LXXXIIL., p. 696.
— (——) tricolor, Pls.
LXXXIL., p. 696.
Emydops minor, P|. XCITI. p. 859.
Endothiodon platyceps, Pl. XCIII.
p- 859.
—- whaitsi, Pl. XCIII. p. 859.
Entephria grisealis, Pl. LXVIIL., p. 546.
mioswari, Pl. LXVIILI. p. 546.
Equus burchelli, Figs. 102-106, pp. 758-
762.
Errina (Labiopora) aspera, Pl. XCY.
p- 876.
—— ( ) capensis, Pl. XCV. p. 876.
—— (—— nove zelandie, Pls. XCIV.—
XCVL., p. 876.

LXXXL., |

(Spinipora) echinata, Pl. XCOV.
p- 876.

Galeops whaitsi, Pl. XCI. p. 859.

Gammarus locusta, Pl. LXXIILI. p. 656. |

zaddachi, Pls. LXXIII., LXXIV.,
p. 656.

Guaffa camelopardalis
Pl. LXXXVI. p. 771.

Glyphodes magnificalis, P|, LXVIIL. |
| Pristerognathus platyrhinus, Pl. XCI.

p. 546.

nigricincta, P|. LXVITI. p. 546.

pfeiffere (var.), Pl. LXVIIT.
p. 546.

pseudocesalis, Pl. LXVIII. p. 546.

Hancockia eudactylota, Pl. LXXXV.
p. 770.

Hyracotenia hyracis,
pp. 594, 596, 597, 601.

Figs.

Letidognathus hemburyt, Pl. XCI, p. 859.

Ictidopsis elegans, Pl. XCILL. p. 859.

Inermicapsifer capensis, Figs. 72-78,
pp. 577, 480, 586, 588, 592.

Labeo percivali, Pl. LXXVIII. p. 672.
Lithophytum arboreum, P\. LXI. p. 505.

Macalia arruensis, Pl. LXVIITI. p. 546.
- mioswari, Pl. LX VIII. p. 546.
olivalis, Pl. LX VIII. p. 546.
Margarosticha plumbealis, P\, LXVIII.
546.

p:
Myzine abdominalis, Pls. LXXXIL.,

LXXXIII. p. 696.
—— braunsi, Pls. LXXXI., LXXXTLI.,
p. 696.
—— constrictiventris, Pls. LXXXTII.,
LXXXIIL., p. 696.

Proc. Zoou. Soc.—1912, No. LXII.

thornicrofti, |

FEB

XXV1l

Myzine stigma, Pls, LXXX1., LXXXIL.,
p- 696.

Nephthya bedfordi, P\s. LXI1., LXIILI.,
p. 505.

chabrolii, Pls. LX1., LXIT., p. 505.

Noorda arfukensis, P\, LXVITT. p. 546.

Nythosaurus browni, Pl. XCIII. p. 859.

Odocoileus, Fig. 110, p. 778.

sp-, Fig, 112, p. 782.

Omphisa variegata, Pl. LXVIII. p. 546.
Otocinelus maculipinnis, Pl, LXXVL1.

p- 666.
Oxyloricaria leightoni, Pl. LXXVII.
p- 666.

tamane, P|, LXXVII. p. 666.

| Pagyda fumosa, Pl. LXVIIL. p. 546.

Perisyntrocha suffusa, Pl. LUXVIII.
p. 546.
Phasianus formosanus, Pls. LX1V.—
LXVIL, p. 5389; Fig. 126, p. 912.
x Phasianus versicolor,
Pls. LXIV.-LXVII., p. 539;
Fig. 126, p. 912.

——— versicolor, Pls. LXIV.-LXVIL.,
p. 589; Fig. 126, p. 912.
Pilocrocis angulifera, Pl. LXVIII.

p. 546.
Plecostomus honde, Pl. LXXVI. p. 666.

p- 859.

Prodicynodon beaufortensis, Pl, XCIII.
p. 859.

Protula pacifica, Pl. LXXXVII. p. 784.

Pyrausta flammealis, Pl. UXVAIL.
p. 546.

Rucervus duvaucelli, Fig. 109, p. 776.
Rusa, Fig. 110, p. 778.
aristotelis, Fig. 108, p. 775.

Scalpellum accumulatum, Fig. 64,

p. 529.

arcuatum, Figs. 64, 65, pp. 529,
534.

—— hastatum, Fig. 64, p. 529.

—— simplex, Fig. 64, p. 529.

trilineatum, Fig. 64, p. 529.

Sceliodes grisealis, Pl. LXVITI. p. 546.

| Sclerophytum pinnulatum, Pls. LXII.,

LXITII., p. 505.
Scymnognathus whaitsi, Pl. XC. p. 859.
Serpula columbiana, Pl. UXXXVII.
84

p. 784.
Spirorbis ambilateralis, Pl. LXXXVILII.
p. 784.
asperatus, Pl. LXXXIX. p. 784.
—— langerhansi, P1. LXXXIX. p. 784.
medius, Pl. LXXXIX. p. 784.

62

XXV111

Spirorbis morchi, Pl. LXXXIX. p. 784.
—— pusilloides, Pl. LUXXXVIII.
p. 784.
racemosus, Pl, LXXXIX. p. 784.
spirillum, Pl. LXXXVIII. p. 784.
validus, Pl. LXXXIX. p. 784.
vitreus, Pl. LXXXVIII. p. 784.
Stereonephthya lutea, Pl. LXIII. p. 505.
Stericta rurealis, Pl. LXVITI. p. 546.
Styphlodora naje, Fig. 122, p. 852.
Sylepta dinawa, Pl. LXVIIT. p. 546.

Taurops macrodon, Pl. XC. p. 859.
Thecla bertha, Pl. XCVII. p. 896.
campa, Pl. XCVII. p. 896.
castrena, Pl, XOVII. p. 896.
datitia, Pl. XCVII. p. 896.
esmeralda, Pl. XCVII. p. 896.
— fancia, Pl. XCVII. p. 896.
Jfernanda, Pl. XCVIL. p. 896.
hamila, Pl. XCVITI. p. 896.
japola, Pl. XCVII. p. 896.

INDEX OF ILLUSTRATIONS.

Thecla molena, Pl. XCVII. p. 896.

nora, Pl. XOVII. p. 896.

schausa, Pl. XOVIL. p. 896.

sicrand, Pl. XCVII. p. 896.

Tilapia percivali, Pl, LXXIX. p. 672.

Tropidonotus natrix, Pls, LXX.—-LXXII.
p. 612; Figs. 86-91, pp. 616, 617,
619, 620, 627, 629.

Urocystidium gemmiparum, Bigs. 113—
121, pp. 824, 826, 828, 833, 835, 841,
843, 845, 846.

Wrightella robusta, Pls. LXIL., LXTIL.,
p. 50d.

Xenocara heterorhynchus, Pl. LXXVI.
p- 666.

multispinis, Pl. UXXVI. p. 666.

Aenophurynx solus, Fig. 122, p. 852.

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

se
08

PROCEEDINGS

OF THE

’ GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY
OF LONDON.

1912.

PART, TIL

CONTAINING Paces 505 To 756, witH 23 PuLaTEs
AND 38 TEXT-FIGURES.

PRINTED FOR THE SOCIETY,
SOLD AT THEIR HOUSE IN REGENT’S PARK,
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| MESSRS. LONGMANS, GREEN, AND CoO.,
| PATERNOSTER ROW.

) SEPTEMBER 1912.)
|

QR Price Roloc Shilins)

LIST OF CONTENTS.

1912, Parr III. (pp. 505-756).

EXHIBITIONS AND NOTICES. |
Page

The Srcretary. Report on the Additions to the Society’s Menagerie during the months
ol Hepruaryaand Marc lial OU reve alatete) <icjele aiein/averet=telelele’/a/ alicia oteteratare\tele eietetapatsen Pete 555

Mr. R.T. Pococs, F.R.S., F.L.8., F.Z.S. Exhibition of lantern-slides of two Polar Bear
cubs (Ursus maritimus) born in the Gardens ..... safe: baie stetolaenelavsleatele aieteeueelepelotenate 556

‘Mr. C. Tare Recan, M.A., F.Z.8. Exhibition of lantern-slides of a Flatfish (Platophrys
podas) showing its power of changing its colour and markings ...........+.. A piaie lets 556

Dr. R. W. Saurenpr, C.M.Z.8. Exhibition of skins of two Virginia Opossums (Didelphis
QINOLMEDIG) keierornvoin eralatdlicisicinjo\aia|eicjayisiotalaonias tai cle lolatlaterepntereferetelorelae ic yore sieroters ricee eto ter™ 556

Mr. D. Szru-Surru, F.Z.S. Exhibition of lantern-slides of the display of the male Pea-

cock Pheasant (Polyplectron chinquis). Also of photographs of the young Cariama
cristata hatched and reared in the Gardens. (Pl. LXIX.)..............--0-c08- 557

The Srcrerary. Exhibition of a living specimen of a young female Dorsal Hyrax
(Dendrohyrax, dorsalts) wa... es sice se = ea eee sm ooo HAGmaUa so oN dobonoon Sago son 671
The Secretary. Exhibition of photographs of an Elephant Kraal in Siam .......... 671

Mr. R. I. Pocock, F.R.S., F.L.S., F.Z.S. Exhibition of a skin and a living specimen of
a fawn variety of the Brown Rat (Hpimys norvegicus) ...01ececerecescccarcesces 671

Mr. D. Sern-Suitu, F.Z.S. Exhibition of epidermal sheaths shed from the base of the
lower mandible of the King Penguin (Aptenodytes penmanti) ..........000 siafelainte 671

Dr. Francts Warp, F.Z.S. Demonstration, illustrated by lantern-slides and the cine-

bo

matograph, of a method of observation of fishes, birds, and mammals under water. 67

PAPERS.

26. Observations on some Aleyonaria from Singapore, with a brief Discussion on the
Classification of the Family Nephthyide. By Epwarp W. Susann, B.Sc., Demonstrator
and Assistant Lecturer in Zoology in the Victoria University of Manchester, (Pls.
DPS PUT Eh Seago ano aoe Goes aadowead naa Heian (e's 9 ig ininin oda ete arene Re SN a 505

27. Some early Fossil Cirripedes of the Genus Scalpellum. By Tuomas H. Witners, F.G.S.
(Metborios, (G4 (ic (GON) ie k'e's ea! alaln inl hlels fein lclo(is seule aint votetia lai loleta alisha siaitehe BAMA Ata ao 528

Contents continued on page 3 of Wrapper,

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P.Z.5, A012.) Pls) tee:

Grout Intaglio et imp.

ALCYONARIA FROM SINGAPORE.

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Grout Intaglio et imp.

ALCYONARIA FROM SINGAPORE.

P.Z.9.. 1912. Pl exit.

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E.W.S. del. Grout Intaglio et imp.

ALCYONARIA FROM SINGAPORE.

PROCEEDINGS

OF TIIE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF TIE

ZOOLOGICAL SOCIETY OF LONDON.

PAPERS.

26. Observations on some Aleyonaria from Singapore, with
a brief Discussion on the Classification of the Family
Nephthyide. By Epwarp W. Suayy, B.Sc., Demon-
strator and Assistant Lecturer in Zoology in the
Victoria University of Manchester *.

{Received January 17, 1912 ; Read March 19, 1912.]

(Plates LXI.-LXIII.7)

INDEX.

Systematic : Page
Sclerophytum pinnulatum, Sp. DV. .cc.csececec serene ees 507
Wephthy@ Cedfor di spa lervesce ws see -cenee a soe ave 514
Stereonephthya luted, Sp. NM. ...3..2.2-4--0.0---e- -.-s 52D
Wrightella robusta, sp. 0. FORCE SSR ESCM BY)
Nephthya: Historical Summary of .................. 510
Dendronephthya do. peret edge OLS
Stereonephthya do. Basten ne esewerslOa0

A collection of Aleyonaria, made by Mr. W. F. Lanchester
and the late Mr. F. P. Bedford during their residence in Singa-
pore, was sent to the Zoological Department at the Victoria
University of Manchester, Professor 8. J. Hickson kindly

* Communicated by Prof. S. J, H1cxson, D.Sc., F.R.S., F.Z.S.
+ For explanation of the Plates see p. 527.

Proc, Zoot. Soc.—1912, No. XXXIV. 34

506 MR. E. W. SHANN ON

entrusted to me the task of identifying the specimens in the
collection. The collecting was carried on in shallow water around
the coast of Singapore; the precise localities will be cited under
the description of the several species. Several examples of
Nephthyide occur, and this necessitated facing the vexed ques-
tion of the classification of that family; I shall state below my
reasons for adopting the scheme of classification advanced by
Kiikenthal.

I wish to express my indebtedness to Prof. Hickson for his
constant advice as to my procedure and help in the pursuit of
literature bearing upon the subject. In the last-named branch
of the work I had frequent recourse to an exceedingly useful
catalogue of the Aleyonaria prepared by Dr. J. Stuart Thomson.
My thanks are also due to My. J. T. Wadsworth for preparing
the excellent photographs from which Plate LXII. figs. 7, 8, & 9
were made.

In all eleven species are described, of which four arenew. The
systematic positions of the several species are as follows :—

Order ALCYONACEA Verrill (pro parte).
Family ALCYONIIDA,

Sclerophytum pinnulatum, sp. n.

Family TELEstip”.
Telesto rupicola F. Miiller.

Family NEPHTHYID#.
Nephthya bedfordi, sp. n.
Dendronephthya disciformis Kiikenth.
Stereonephthya lutea, sp. n.
Paraspongodes crassa Kiikenth.
Family SIPHONOGORGIIDE.
Siphonogorgia variabilis Hickson.

Order GORGONACEA.
Suborder PSEUDAXONTA.

Family Scterocoreps.
Suberogorgia suberosa Pallas.

Family Mrtrropip®.
Melitodes albitincta Ridley.
Psilacabaria gracillima Ridley.
Wrightella robusta, sp. n.

ALCYONARIA FROM SINGAPORE. 5O7

Order ALCYONACEA.
Family ALCYONIIDA.
Genus SCLEROPHYTUM.

Sc. PINNULATUM, sp. n. (PI. LXII. fig. 7; Pl. LXIII. fig. 10.)

A single complete colony was taken just below low-tide mark
from Blakang Mati.

In its form and mode of branching the colony closely resembles
Se. palmatum Pratt (1903). As in the last-named species, the
colony (fig. 7) is erect, branched, and shows a marked lateral
compression. ‘The total height is 88 mm., of which the stalk
comprises 45 mm. The diameter of the stalk is 31x11 mm.,
and that of the capitulum is 60x 35mm. The stalk divides at its
distal end into two almost equal primary branches. From these
branches spring numerous secondary lobes which are arranged
in no definite order. ‘The secondary lobes frequently show
lateral compression in their proximal regions, but their termi-
nations are usually bluntly conical, while a few are reniform.
The length of these lobes is 12-20 mm., and their diameter
6—9 mm.

The colour of this specimen in alcohol is slate-grey. The
consistency is tough; the stem is hard and brittle, but the
secondary lobes are soft and fleshy.

The autozoids are of a brown colour and are uniformly scattered
over the surface of the secondary lobes. The average distance
between two adjacent polyps is*6 mm. Autozoids are also found
on the primary branches almost down to the level of the stalk,
but in this region the distance between them is very much greater,
viz. 3-4 mm. Many of the autozoids are more or less expanded,
but none of them are situated on raised rounded areas such as
occur in Se. palmatum. The anthocodie are of medium size
when compared with those of the genus Sclerophytum as a whole ;
the diameter of an expanded crown is 1 mm., but the polyp-heads
appear only half that size when contracted.

The tentacles (fig. 10) form the distinctive feature of this
species. They are ‘55 mm. in length, compressed, and expanded
at the distal end. There is a single row of free well-developed
pinnules on each side of a tentacle. The pinnules are -06 mm.
in average length, but the larger ones attain a length of :10 mm.
and a breadth of (035 mm. The larger pinnules are those which
are placed third, fourth, and fifth from the distal end; they
become shorter as they approach the base of the tentacle. There
are about twelve pinnules in each row.

The stomodzum is long and convoluted. The mesenteries
and mesenterial filaments are well marked. No sexual organs
were observed.

The siphonozoids, if present, are extremely degenerate. A
very few minute diverticula of the superficial canals, which point
in the direction of the surface, but never open to the exterior,

34*

508 MR. E. W. SHANN ON

may represent rudimentary siphonozoids. The superficial canals
form a dense network just beneath the surface of the colony ;
their diameter is (044-120 mm. The vessels of the internal
system are clearly defined and circular in cross-section, and are
fairly numerous.

Zoochlorelle are numerous in the superficial canal-system, and
occur in less abundance in the interior. They are also found in
the endoderm of the polyps, and extend into the tentacles,
occupying even the lumen of the pinnules.

The spicules are quite characteristic of the genus Selerophytum,
and indeed only differ from those of Se. palmatum in almost
insignificant detail. The little knotted clubs average "16 x ‘034 mm.,
and the spindles are sometimes only -058x‘015 mm. ‘These
minute spicules only occur immediately beneath the surface.
The spicules of the ccenenchyme are fairly numerous and are all
of the tuberculate warted type; they vary considerably, however,
in shape and size. The majority are spindles which narrow
rapidly towards their ends to rather acute points; their measure-
ments range from ‘4x‘1]1 mm. to 30x45 mm. Ivregularly
branched forms are by no means uncommon.

The following considerations are advanced as an apology for
the creation of a new species of Sclerophytwm based on the
examination of a single specimen.

This colony had been assigned in a preliminary investigation
to the species Sc. palmatwm Pratt (1903), and at first sight this
diagnosis appeared perfectly aceurate. The external appearance,
and indeed the actual measurements, agree closely with the
description of thetype specimen. The characters and distribution
of the spicules, the well-marked mesenteries and mesenterial
filaments, the rudimentary condition of the siphonozoids, the
orientation of the canal-systems, and the distribution of the
zoochlorelle, all tend to enhance the resemblance between the
two species. It is only when the autozoids are examined that
the true specific difference is realized; those of Se. palmatwm are
distinctly larger than those of Se. pinnulatwm. The characters
of the tentacles which form the essential divergence of the two
species are tabulated below :—

Sc. palmatunr Pratt. Se. purnulatum, sp. n.
‘7 mm. in length. -55 mm. in length.
Almost of uniform length. Expanded at distal end.
Possess a double rowof rudi- <A single row of free well-deve-
mentary pinnules down loped pinnules down each
each side. side.

Se. pinnulatwm could be confounded with no other species of
Sclerophytwm. At the same time, it is of interest to note that a
single row of free pinnules has been recorded on either side of
the tentacles of Se. viride by Thomson and Henderson (1906).
The possession of free pinnules, now recorded for two species of

ALCYONARIA FROM SINGAPORE. 509

Selerophytum, tends to strengthen the relationship of the genera
Sclerophytum and Xenia which has been suggested by Pratt

(1903).
Order ALCYONACEA.
Family TELESTID &.
Genus TELESTO.

T. rupicoLa F. Miiller.

Carijoa rupicola F, Miiller, Arch. Naturg., Jg. 33, 1867, p. 33,
tab. 9. figs. 56 & 57.

Telesto (Carijoa) rupicola Wright & Studer, ‘ Challenger’
teports, Zool. vol. xxxi. 1889, p. 262.

Telesto rupicola May, Jena Zeitschr. Naturw. vol. xxvi. 1900,
p. 58.

Telesto rupicola Hickson & Hiles, Willey’s Zoolog. Res. 1900,
p. 496, tab. 50. figs. 1 & 2.

Telesto rupicola Thomson & Henderson, Marine Fauna of
Zanzibar, 1906, p. 434.

Telesto rupicola H. Laackmann, Zoolog. Jahrb. Supp. 11,
Heft 1, 1908) pxSl shat eo ahos a, 251 Mal. 3y fig. 3.

Although the only species of Zelesto previously recorded from
Singapore is 7. prolifera v. Koch, the numerous small colonies in
this collection appear to bear a closer resemblance to 2’. rupicola,
and are therefore described under that name.

The largest of these colonies is of a grey hue. Its longest
axial polyp measures 85 mm. in length, it is 3 mm. in diameter
at the base and 1:25 mm. at the top. It bears six lateral polyps,
the longest of which measures 24 mm. Anthocodiz arise at
frequent, but pretty regular intervals, both from the axial and
lateral polyps; their average length is 3 mm., and breadth 1°5 mm.
The majority of the other specimens are pale yellow and of
smaller size. All these forms were obtained in shallow water
near Singapore. Some of the exact localities read as follows :
Pulo Brani, 6 fms.; Pulo Brani, 5-10 fms.; Blakang Mati, below ~
low tides ; Tanjong Pagar, 10 fms.

Previously recorded from Rio de Janeiro (F. Miller); Brazilian
coast (Munich Museum); Blanche Bay, New Britain (Hickson
& Hiles); Zanzibar (Thomson & Henderson); Bahia (‘ Chal-
lenger’).

The spicules show a very wide variety of forms. Not only is
this the case, but a different selection of spicules was found in
each of the five specimens examined.

One specimen contained spicules very much resembling those
of 7’. riisei. Indeed, the two species are probably very closely
related, for Laackman (op, cit. pp. 72 & 82) is at some pains to
distinguish between them. It should be remembered, however,
that 7. riiset has not yet been recorded from the Old World.

A very small specimen of a pale yellow colour was examined

510 MR. E. W. SHANN ON

and found to bear many points of resemblance with 7. prolifera ;
the spicules, for instance, agree well with those figured by
Laackmann for that species (op. cit. p. 87), and the stem walls
are very thin. At the same time, it was taken from the same
locality (Pulo Brani) as one of the larger specimens which un-
doubtedly belongs to the species 7’. rwpicola. Moreover, another
specimen was intermediate in every particular between the
above exceptional form and the larger examples, which had been
assigned without difficulty to 7. rupicola. As this small specimen
was evidently very young, one would not wish to attach too deep
a significance to the observations made or to draw any hasty
conclusions from them. At the same time, it is well within the
range of possibility that the accumulation of such knowledge
may lead eventually to a reduction in the number of species of
Telesto.

An Historical Summary of the Genera Vephthya, Dendronephthya,
and Stereonephthya, with reasons for retaining the definitions
of these genera of the family Nephthyide as enumerated
by Kiikenthal.

Genus NEeputTuya Savigny.

1817. Nephthée Savigny, Descr. de /Egypte, Hist. Nat. Suppl. i.
Atlas, Polypes, tab. 2. fig. 5

1828. Nephthea Audouin, Explication sommaire des Planches de
polypes de Egypte et de la Syrie, publiées par Jules-
César Savigny dans: Description de l’Egypte, vol. xxiii.
Paris.

1834. Mephthya Khrenberg, Die Corallenthiere des Rothen
Meeres, p. 284.

1846. Nephthya Dana, ‘ Zoophytes,’ Philadelphia, p. 610.

1857. Nephthya Milne-Edwards, Histoire Naturelle des Coral-
laires, vol. 1. p. 127.

1877. Nephthya Klunzinger, Die Korallthiere des Rothen Meeres,
AMG lio, Bris

1887. Nephthya Studer, in Arch. Naturg., Jg. 53, vol. i. pp. 19,
20.

1887. Nephihya (pars), Danielssen, in Norske Nordhavs Exp.
vol. v. p. 82.

1889. Nephthya and Spongodes (pars) Wright & Studer, ‘Chal-
lenger’ Reports, Zoology, vol. xxxi. p. xxv.

1895. Spongodes (pars) Holm, in Zool. Jahrb., Bd. 8, p. 24.

1895. Spongodes (pars) Kiikenthal, in Zool. Anz., Jg. 18, p. 428.

1896. Nephthya Kukenthal, in Abh. Senckenb. Ges. Frankfurt,
vol. xxii. pp. 89-91.

1899. Nephthya May, in Jena Zeitschr. Naturw. vol. xxxiii. p. 156.

1903. Nephthya Kikenthal, in Zool. Jahrb., Bd. 19, p. 141.

In 1817 Savigny described two genera of Nephthyide to which
he gave the names Ammothée and Nephthée. Audouin (1828),

ALCYONARIA FROM SINGAPORE. 511

to whom fell the task of describing Savigny’s plates, believed
that Savigny’s Tab. 1. fig. 8 represented Ammothée, and that
Tab. 2. figs. 5 & 6 represented Mephthée. The genus Nephthea,
as the author wrote it, was recognised by Ehrenberg (1834); but
at the same time he disputed the interpretation of Savigny’s
plates, maintaining that Audouin had given the name Vephthea
cordierti to the form, represented in Tab, 2. fig. 6, which Savigny
had intended to cail Ammothée. Ehrenberg’s view has been
accepted by all subsequent authors, and it is now generally agreed
that Tab. 2. fig. 5, correctly designated by Audouin Wephthea
chabrolii, represents the type of Savigny’s genus Wephthée, while
Tab. 2. fig. 6 represents his type of the genus Ammothée, namely
A. virescens. There can be no reasonable doubt with regard to
the authenticity of origin of the genus Vephthya, that is to say,
that the genus was based on the description of the species
NV. chabrolii, which is figured in Savigny’s Tab. 2. fig. 5. In the
ease of Ammothée, or Ammothea as the genus was known for
many years, the name was changed to Lithophytum by Kiikenthal
(1903), since that author found that Savigny’s type species,
A. virescens, is identical with a form described forty-two years
previously by Forskal under the name Lithophyton arboreum ; in
deference to the International Rules of Zoological Nomenclature
the older name must be retained. Thus it is of little moment
whether or not Ehrenberg was justified in disputing Audouin’s
interpretation of Tab. 2. fig. 6.

Copies of Savigny’s plates are extremely scarce, so that it is
not always possible for the research worker to examine the
original figures; many have probably been compelled to content
themselves with descriptions by other authors. With this
difficulty in view, Professor Bourne very kindly had photographs
taken for Professor Hickson from Savigny’s plates, Tab. 2.
figs. 5 & 6, in the Radcliffe Library at Oxford. Prof. Hickson
has given me permission to publish these figures in this paper,
so that they may be readily accessible to all workers on the
Nephthyide. They are reproduced in Pl. LXI. figs. 1-5 and
PI xe tie.6)

Ehrenberg (1834) distinguished Vephthya from Ammothea by
the prominence of the polyp-spicules in the former genus, for he
says of Vephthya:—“ polypis in verrucas inermes retractilibus.”
We see, then, that Ehrenberg recognised the distinction between
the genera Vephthya and Lithophytum (Ammothea) which obtains
at the present day, namely the presence and absence of armed
polyps (polyps with “ Stiitzbiindel ”) in these genera respectively.
Ehrenberg’s definition of Vephthya was recognised by Dana (1846),
Milne-Edwards (1857), Klunzinger (1877), Studer (1887), and
Danielssen (1887). The numerous new species described during
this period were distinguished by their authors, on the one hand
from Ammothea by the presence of armed verruce, and on
the other from Spongodes Less. by the comparatively slight
development of the spicules which formed the armament of

512 MR. BE. W. SHANN ON

the verruce. Wright and Studer (1889) described a specimen
under the name Spongodes nephthyeformis, concerning which
they observe :—‘‘The entire habit of the colony recalls much
more that of MWephthya than that of Spongodes, and this im-
pression is strengthened by the slight development of the spicules
surmounting the little heads, whence the colony does not appear
so prickly as other species.... The species must be referred to
the genus Spongodes, because the polyps are placed sideways
within a bundle of spicules, although these only project slightly.”
That is to say, Wright and Studer recognised on the anthocodiz
of Sp. nephthyeformis the presence of what is now known asa
“« Stiitzbiindel,” and here we have the starting-point of the
difficulty of discriminating between the genera Spongodes and
Nephthya. Wolm (1895) “Faced the problem of reconstructing
the genus Spongodes in the light of the knowledge which had
seemamnlevradl since Lesson fies described the genus in 1834,
This author pointed out that Spongodes nephthyceformis W.& St.
is identical with Nephthya chabroht Audouin, and added that
NV. chabrolii differs from Spongodes in many characters, such as
the branching of the colony and the arrangement of the polyps:
on these characters one can establish two genera, but it is
necessary then to add to the genus Wephihya many species
hitherto included in the genus Spongodes, including the type
Sp. celosia. Though Holm shrank from submitting a well-known
type like Spongodes celosia to such treatment, he proceeded
fearlessly to include all the species of Mephthya, including the
type WV. chabrolii, within the genus Spongodes; he retained
Nephthya, however, as a subgeneric title. Here was a step in
the direction of elucidation; Nephthya and Spongodes, as hitherto
defined, were shown to be synonymous; but Holm’s solution of
the problem threw too great a burden on Spongodes. Kiikenthal
(1895), writing during the year in which Holm’s paper was
published, accepted the genus Spongodes in its new distended
form; but the term Spon odes had become so obviously cumber-
some that this author (Kiikenthal), in a later paper *, reinstated
Nephthya with full generic honours. In this paper Kukenthal gave
a summary of the family Nephthyide, and divided the various
genera into two groups as they possessed or lacked a “ Stiitz-
bindel”; he summed up his remarks as follows :—‘“ Innerhalb
der Familie der Nephthyiden ist als wichtigstes Merkmal zu
betrachten, ob die Polypenképfchen terminal auf ihrem unteren
Teile, dem Stiele, sitzen oder seitlich davon. Letzterer Fall tritt
stets dann ein, wenn sich auf einer Seite, der oberen, ein Biindel
Spicula besonders stark entwickelt : das Stiitzbiindel.” The
genera possessing a “ Stiitzbiindel” were distinguished from one
another by the disposition of the polyps on the colony, as Holm
(1895) had already suggested; the name Vephthya was applied
to forms resembling the original type WV. chabrolit Audouin, in
which the polyps are collected on branchlets, the latter bemg
arranged in catkins or lappets, and the name Spongodes was

* Abh. Senckenb. Ges. Frankfurt, vol. xxiii. p. 88 (1896).

ALCYONARIA FROM SINGAPORE. 513

retained for forms in which the polyps are disposed sporadically
or in bundles. The net result of this reformation was the
inclusion in the genus Vephthya of all forms hitherto included in
the “ Spicatee ” group of the genus Spongodes.

In adopting this means of classification it became necessary
to include Sp. celosia, Lesson’s type of the genus Spongodes, in
the emended genus Vephthya; nevertheless, the name Spongodes
was retained by Kiikenthal for his other emended genus, since
the latter included a large number of forms which during many
years had been described under Spongodes. The definition of the
genus Vephthya given in the paper under consideration runs
thus :—‘‘ Nephthyiden mit ‘ Stiitzbiindel.’ Die Kolonie ist
buschig veriistelt, die meist Kurzen und nur vereinzelt sterilen
Stammteile sind durch einen abgeflachten, oft membranosen
Basalteil verbunden. Die Polypen stehen in grosser Zahl und
zeimlich gleichmiissig verteilt auf den Steinzweigen, die dadurch
die Form von iihrenférmigen Lappen oder ‘ Katzchen’ erhalten.
Hervorragende Spicula der Polypenképfchen fehlen.” May (1899)
accepted the foregoing definition of the genus Vephthya, aud in
his * Revision of the Nephthyide’ (1903) Kiikenthal has not had
occasion to modify it.

A * Stiitebiindel.”

The crucial point in Kiikenthal’s classification of the Neph-
thyidee lies in the definition of the term ‘“Stiitzbiindel.” Much
of the opposition to the above classification has arisen through
the different interpretations which various authors have attached
to the term. Kiikenthal, to whom we owe the word, uses it in
an extremely comprehensive sense, the range of which can be
circumscribed, however, by the following limits :—

A “Stiitzbiindel” is an aggregation of spicules disposed along
the abaxial aspect of an anthocodia and lying approximately
parallel to its axis. The spicules are usually spindle-shaped ;
they are not infrequently larger than those from any other
portion of a given specimen, and one or two of them commonly,
but not invariably, reach from the apex of the polyp-stalk into
the substance of the colony. A few of the spicules in a
characteristic example protrude beyond the polyp-head, but
such a condition is not essential.

A definite “Stiitzbiindel” may not be recognisable in every
polyp of a given colony; but if such is present it will appear
most obvious in the younger polyps near the distal ends of the
branches. A specimen in which a “ Stiitzbiindel” is demonstrable,
whether or not in every polyp, must be classified as possessing a
* Stiitzbiindel.”

In the genus Lithophytwm, which closely resembles Vephthya
in external appearance, the anthocodiz being massed on small
terminal lobes or lappets, there is no “Sttitzbiindel.” The
-anthocodiz contain very few spicules, some of which are loosely
arranged en chevron along the abaxial surface. Both the small
size and oblique position of the spicules so arranged prevent their
being deseribed as forming a “ Stiitzbiindel.”

514 MR. E. W. SHANN ON

NEPHTHYA BEDFORDI, sp. n. (Pl. LXII. fig. 8; Pl. LXIII.
fiers 2)

Two specimens which, while conforming with the characters
described above for the genus Vephihya, fail to agree in detail
with any of the large number of forms hitherto described, have
necessitated the creation of a new species of this genus.

The colony (fig. 8) is bilaterally compressed, the growth is
bushy, and the major diameter of the capitulum is approximately
equal to the total height, including the stem. The consistency
is tough and leathery. The stem, which shows signs of bilateral
compression, is Short, and gives rise at its distal extremity to a
variable number of main branches. These main branches are
again divided into unequal secondary branches. From both
main and secondary branches spring the short terminal lobes.
The latter are conical in shape, but rounded; on them the
anthocodiz are tolerably evenly distributed. As the polyps are
situated very close together, and the terminal lobes are ex-
ceedingly numerous, the whole capitulum appears to be covered
with anthocodiz. ‘The polyp-heads when at rest make an acute
angle with their stalks. The latter scarcely protrude from the
colony, with the result that the polyp-heads are very closely
apposed to the surface from which they arise.

Colour in alcohol cinder-grey, polyps brown.

Locality : below low-tide mark, Blakang Mati.

Detailed measurements :—

Region measured. Specimen I. Specimen II.
Motaleheigit ewes crease see 60 mm. 42 mm.
Height of stem ............... 17 mm. 14 mm.
Diameter of stem ............ 18 x 10 mm. 12 x 10 mm.
Height of capitulum......... 43 mm. 28 mm.
Breadth of capitulum ...... 52 x 20 mm. 45 x 20 mm.

In both specimens the length of the terminal lobes is about
4 mm., but the variation in this respect is between 3 and
6mm. The diameter of the terminal lobes is from 3 to 4mm.
The edges of adjacent polyps are seldom more than *175 mm.
apart. The polyp-heads have an average length of 1-1 mm. and
diameter of ‘6 mm. :

The spicules show a wide range in size and shape. The
smallest forms are found in the tentacles, where they are
arranged in no very definite order, but for the most part le at
an acute angle with the axis of the tentacle. The tentacle-
spicules are minute spiny spindles, some of which are straight,
others crescentic. The polyp-spicules (fig. 11, /) are very brittle,
longitudinally striated spindles. They are straight or curved,
and usually smooth, but sometimes bear a few minute spines.
It is difficult in most cases to discern an arrangement of the
polyp-spicules en chevron, but some of the polyps show such a
condition more clearly than others.

ALCYONARIA FROM SINGAPORE. ole

The “Stiitzbiindel,” though undoubtedly present, is very ill-
defined. From four to six spindles can usually be observed
along the abaxial surface of each anthocodia, but in no instance
do these protrude beyond the polyp-head. It is exceedingly
difficult to dissect out a single polyp with its supporting spicules
intact; some preparations were obtained, however, after soaking
portions of the branches for twelve hours or more in oil of
cloves. A more satisfactory method of observing the “ Stiitz-
biindel” is obtained by placing one of the terminal lobes,
previously cleared in oil of cloves, under a low-power binocular
microscope. By this means a stereoscopic view of the polyp and
its supporting spicules is obtained, such as is represented in
fig. 12. The cortex of the capitulum contains numerous hori-
zontally disposed spindles, some of which are among the largest
spicules found in this species; they resemble in type those
described from the polyps. In the outer wall of the stem are
found small spicules with broad rays (fig. 11, @) which are usually
numerous, but vary considerably in size and concentration.
These forms are covered with warts ‘015-:036 mm. long, and
interlock with one another, thus accounting for the tough
consistency of the stem. Among the forms described above
there are also in the outer wall of the stem some larger spindles
with remarkable spines (fig. 11, 6); the latter, which have an
average height of ‘044 mm. and basal breadth of -02 mm., are
often larger and sharper on one side of the spindle than on the
opposite side, and, since the spines all lie approximately in one
plane, give the spicule a comb-like appearance. The spicules of
the canal-walls (fig. 11, ¢, d) are not very plentiful, but are
distinguished by their stout appearance. They may be described
as very thick, longitudinally-striated spindles, somewhat flattened
and bearing low rounded warts. Their ends are either rounded
or bluntly pointed, the body of the spicule is straight or slightly
curved. Among these regular forms there are found a few
branched spicules. The latter resemble the regular forms in
structure, but are either triradiate or show short irregular
processes springing from the central region of a typical spindle. —

The following are characteristic measurements of the spicules,
length by breadth, in millimetres :—

(@EPolyp:(outer): -..y.seeuseese reece *3 X '025
(6) Bolypi(amnen)) 5. csespecmeeneeeeeee ae eee ei 112 X '023
07 X °02
(@) FRANTIC Gos casncsensopaccosooroocconsenssenpecs. IB SSO *6 X 05
‘4 xX 06
(Ai@ortex of capitulunameereeeeseeeceeene anata es 1:0 X ‘075 ‘9 X 08
(e) Cortex of stem, radiate forms ..................... “115 X ‘181 058 X “054
rib etetiess of branches *03-"015
Do. spindles (including spines) ... °85 X °125 22 X °125
(ayCanal-walls; spindleStaynccnttaeer neal nl ON Bay 2. “6 5<ohIe
“4X ‘08
Do. triradiate forms “41 X ‘4, thickness of ray ‘1

(measured by taking two ter-
minal points asa base line and
the third point as the extremity). ‘22 °18 thickness of ray ‘05

516 MR. E. W. SHANN ON

On applying the key to the species of Wephthya given by
Kukenthal (1903, p. 145) to the case of WV. bedfordi, it is seen
that this species falls within the group characterised by forms
in which the terminal lobes are conical but rounded. The
inner polyp-spicules are smaller than the outer; moreover,
they have the form of smooth rods. These characters taken
together indicate that WV. bedfordi resembles closely WV. pacifica
and WV. albida; and, since the spicules do not form a ring around
the base of the polyp, it may be inferred that the new species is
most nearly allied to IV. albida Holm, so far as the key to the
species can be relied upon. Excepting in the character of the
inner polyp-spicules, however, WV. bedfordi appears to be more
nearly related to WV. elongata Kiikenthal than to JV. albida.
This supposition is supported by the fact that NV. elongata is
reported from Ternate, while WV. albida is a Red Sea form. ‘The
points of difference and of resemblance between JV. bedfordi and its
two nearest allies may be seen at a glance in the appended table :—

Character. NV. elongata. N. bedfordi. N. albida.
Terminal lobes 1 ree oa 2 ve z
GONE sass00 °8 11 1

Polyp-heads 1 tiie Mb ‘9-6 6 7
Angle between Polyp and

Siballilcr ey Aeeeen ck Peeve a 45° Acute. Right.

Hager Outer ... 27xX— *3X< 025 3X03
Polyp-spicules { 70 fede glyco 112-023 “AX<-O15

(Spiny spindles.) | (Smooth rods.) | (Smooth rods.)

“Sttitzbtindel” ............) Projects slightly. | Does not project. | Projects slightly.
Spicules of “Sttitzbtindel”

WOVE coo oon 15 6 12

Thick ... ae *05 12
Spicules of Upper Cortex:

Long ...... 8 il 14

WE S00 ae “075 12
Spicules of Lower Cortex:

ONG ener 8 “35-22 "85
IMOVIE 505 (Compact.) 125 “22

(Comb-like and
radiate forms.)
Spicules of Canal-walls:
iomcueeeee 1:2 11-4 il

Thick ...! (Scattered.) ‘2-08 i

(Spindle and tri- |
radiate forms.) |

Colour (in alcohol) ...... Greyish yellow. Cinder-grey. Greyish white.
IDOE, sccconcranoncs stoncnons|| erauRnas, BS ston, Singapore. Red Sea.

All measurements are in millimetres.

Lest the use of an artificial key to the identification of species
should have led me to overlook a member of the genus not
included in the group to which the Singapore specimen apparently

ALCYONARIA FROM SINGAPORE. 517

belongs, but at the same time exhibiting a close resemblance to it
in structural detail, I have read descriptions of all the species of
Nephthya which might possibly have proved to be identical with
NV. bedfordi. In no instance is there a greater resemblance
between iV. bedfordi and another species of Vephthya than exists
between any two established species of that genus. Indeed, the
exceedingly feeble development of the “ Stiitzbiindel” and the
presence in the stem-cortex of exceptionally small, though
curiously comb-shaped, spicules are characters which serve clearly
to distinguish 1. bedfordi from any species previously recorded.

Among some unpublished notes by Miss Coward, which
Professor Hickson has kindly placed at my disposal, the following
paragraph occurs :—

‘*In his work on the family Nephthyide (1903) Kiikenthal
names a specimen Vephthya chabrolii. In doing this he refers
to Hickson and Hiles’ (1900) description of WV. chabrolii. These
writers, however, state that the spicules of their specimen are
just as described by Klunzinger (1877)—that is to say, they
do not form a ‘ Stiitzbiindel.’ In the description of his specimen
Kiikenthal says the spicules only rarely project beyond the
polyp-heads—and yet his diagnosis is the presence of a
‘ Stiitzbiindel.’”

This observation came to hand after the account of the new
species WV. bedfordi had been written and the ‘ Stiitzbiindel”
described. As stated above, the ‘‘ Stiitzbindel” was very poorly
developed. TI then thought it advisable to examine JV. chabrolii
to satisfy myself as to the nature of the “ Sttitzbiindel” in that
species. ‘The specimen at my disposal was the identical one
described by Hickson and Hiles (1900), in Willey’s collection.
Preparations, which have been cleared in oil of cloves and
examined under the binocular microscope, reveal the presence of
distinct bundles of spindles supporting the polyps. One or two
of these spicules not infrequently project beyond the polyp-
heads. One must admit that WV. chabrolii is characterised by
the presence of a small but clearly defined “ Stiitzbiindel.” It
seemed possible that the “ Stiitzbiindel” was so degenerate in
NV. bedfordi that the polyps might be described as being without
this characteristic. Were this the case, the specimen in question
would come under Kiikenthal’s definition of the genus Litho-
phytwm Forsk. I accordingly made and examined preparations
of Lithophytwm arboreum ; but in this case there was no trace of
a bundle of spicules on the abaxial surface of the anthocodiz
in the least degree comparable with the condition which I have
described in JV. bedforde.

The genus Vephihya, as at the present time accepted, contains
a large series of species showing every gradation in the develop-
ment of a ‘ Stiitzbiindel.” At one end of the series is found
such a form as JV. celosia Lesson, in which one or two of the
“ Stiitzbiindel ” spicules reach 2 mm. beyond the polyp-heads ;
in the middle .V. chabrolii, in which they only project slightly ;
and at the other end WV. bedfordi, in which they do not project.

518 MR. E. W. SUANN ON

Genus Denpronerurayva Kiukenthal.

1791-97. Alcyoniwm Esper, Pflanzenthiere, pp. 49, 50, tab. 16.
1834. Nee Spongodes Lesson, Illustrations de Zoologie, vol. ii.
part 2, p. 89.

1834. WNephthya (pars) Ehrenberg.

1846. Spoggodia (pars) Dana, p. 625.

1857. Spoggodes (pars) Milne-Kdwards, p. 127.

1862. Spoggodes (pars), Morchellana Gray, in Proc. Zool. Soc.
London, p. 27.

1877. Spongodes (pars) Klunzinger, pp. 34, 35.

1889. Spongodes (pars) Wright & Studer.

1895. Spongodes (pars) Holm, p. 16.

1896. Spongodes (pars) Kiikenthal, p. 97.

1899. Spongodes (pars) May.

1905. Dendronephthya Kiikenthal, Zool. Jahrb. xxi. p. 526.

The genus Spongodes was founded by Lesson (1834) on the
type Sp. celosia, and for many years workers in the field of the
Nephthyidee found no difficulty im discriminating between
Spongodes Lesson and Wephthya Savigny. I have endeavoured
to show in my section on the genus WVephthya that as the number
of species of the above-mentioned genera increased it became
more and more difficult to draw a hard-and-fast line between
them; further, that this fact became manifest when, in 1889,
Wright and Studer described a specimen as Spongodes nephthyc-
formis, which was subsequently relegated to the genus Wephihya,
and, indeed, shown to be identical with WV. chabrolii, the type of
that genus. A new distinctive feature was required: such a
feature was discovered by Holm and applied in a practical form
by Kiikenthal. The feature in question is the arrangement of
the polyps upon the stem, and its application has already been
mentioned above. Kiikenthal’s (1896) definition of the emended
genus Spongodes is as follows :—“ Polypenstock baumartig veriis-
telt, unterer Stammteil nackt. Die Polypen sind in Biindeln
vereiniet oder stehen vereinzelt.” Under the genus Spongodes
thus defined, Kiikenthal included as subgenera Spongodia and
Spongodes ; he further subdivided the latter into three groups,
namely, Glomerate, Umbellatz, and Divaricate. May (1899)
and others adopted this classification ; and so the matter stood
until the-year 1905, when Kiikenthal published the second
part of his “‘ Revision of the Nephthyide.” In this paper was
introduced the division of the time-honoured genus Spongodes
Less. into the two new genera, Dendronephthya and Stereonephthya,
which has provoked such a storm of criticism. It must be
remembered, however, that the group Spongodes ‘ Spicatze,” and
with it the original type Sp. celosia, had already been relegated
to the genus Vephthya; so that the genus Spongodes, as then
accepted, no longer retained its ancient prestige ; moreover, since
the genera Vephthya and Spongodes were shown to be synony-
mous, Kiikenthal was justified by the International Rules of
Zoological Nomenclature (Art. 28) in retaining the older name—

ALCYONARIA FROM SINGAPORE. 519

Nephthya—to designate the merged genera. The genus Dendro-
nephthya is almost synonymous with Spongodes (Glomeratie,
Umbellate, and Divaricatie), as defined by Kiikenthal in 1896,
and Stereonephthya has a like relation with his Spongodes
(Spongodia).

There is no doubt that here, again, Kiikenthal was acting in
accordance with the International Rules of Nomenclature. Had
these Rules been extant at the time Kiikenthal published his
classification of the Nephthyide in 1896, the name Spongodes
would doubtless have been discarded by him then, for in the
second section of Art. 30 it is written: ‘‘In no case, however,
can the name of the original genus be transferred to a group
containing none of the species originally included in the genus.”
Thus Spongodes, as used by Kiikenthal in 1896, would not be
considered as a valid generic name by those who accept the
International Rules of Zoological Nomenclature, which appeared
in 1905. One cannot but feel the loss of a name which for
nearly a century has been familiar to students of the Alcyonaria,
and regret the inconvenience caused in Museums and Zoological
Laboratories throughout the world by its suppression. Nomen-
clature is a matter of convenience and cannot be regulated
absolutely by. arbitrary laws. Generally -speaking, of two
synonyms, that which is most familiar to the majority is to be
preferred ; a more practical reason should be required than a
mere regulation before supplanting an old familiar generic name
by anew one. If we are required to obey literally the Inter-
national Rules of Zoological Nomenclature, such everyday names
as Astacus, Holothuria, Actinia, Madrepora, and many others
will be forfeited, as has been Spongodes. While regretting that
Kiikenthal did not exercise his authority in retaining the name
Spongodes, in deference to his extensive knowledge of the Neph-
thyide and his able reorganisation of that family, I have adopted
the term Dendronephthya for a genus of Nephthyide defined as
follows by Kiikenthal (1905) :—

“ Nephthyiden von baumformig verzweigtem Aufbau, deren
Polypen stets in Biindeln vereinigt sind, Polypen mit ‘Stiitz- .
biindeln.’”

DENDRONEPHTHYA DIscIFoRMIS Kikenthal 1905, (Pl. LXIII.
fig. 13.)

Three specimens in the collection agree so nearly in all
essential features with the description given by Kiikenthal
(1905) for D. disciformis that there is little doubt that they
may be assigned to that species. Kiikenthal’s specimen came
from the China Sea.

The largest of the three colonies is 8 em. high, 7 cm. broad, and
3°5 em. thick. It was taken off Pulo Brani in 5 fms. of water.

Two other specimens were taken in 7 fms. of water between
Pulo Hautu and Blakang Mati. Their measurements, height by
breadth by thickness, are 4x 4x 2°5 cm., and 3°5x2°5x1:5 em.
respectively.

520 MR. E. W. SHANN ON

It may be noted that the two last-mentioned specimens are
much smaller than any recorded by Kikenthal. he colour
of the upper branches and “ Stiitzbundel” is pink, whereas the
typical colour for this region is deep red. Moreover, no radial
spicules were observed in the lower portion of the stem.

The exact systematic position of a fourth specimen is rather
doubtful; but, until further evidence is forthcoming, it has
been deemed advisable to describe the specimen under the specific
title D. disciformis.

This specimen shows externally the typical ‘ disciformis”
features. It is 9°5 em. in height and 8:5 x 3-0 em. in breadth,
thus exceeding somewhat in dimensions the largest specimen
described above. The characters and distribution of the spicules
are identical with those of typical examples, only excepting those
from the lower portion of the stem. The spicules found .in the
last-mentioned area are spiny clubs (fig. 13), among which are
scattered a few of the typical radiate forms. The clubs vary in
length from °70—27 mm., but the majority are ‘45 mm.; the
thickness is almost the same even in forms of different length,
the average width of the handle of the club is ‘05 mm., that
of the head *12 mm., excluding the spies, which sometimes
reach :044 mm. in height. The colour of the branches and
“ Sttitzbiindel ” is a dull yellowish brown, and there are no red
spicules such as are found in these areas in all the other recorded
specimens. Kiikenthal (1905) described a colony whose branches
were pale brown and whose polyps were white, but even in this
specimen red spicules were observed.

Doubtless one might be led to base a new variety of Dendro-
nephthya on the characters of the colony described above, a
colony possessing a very characteristic form of spicule which has
not been observed, either from the same region or from any
other region, in any previously recorded specimen of D. disci-
formis. Such a course might have been adopted had not this
aberrant form been taken at the same time and from the same
place as the two smaller of the three colonies described above,
namely, between Pulo Hautu and Blakang Mati, 8/xii/98, in
7 fms. It seems probable, then, that the species D. disciformis
is liable to considerable variation in the colour of the terminal
branches and “ Stiitzbiindel,” and in the character of the spicules
which are found in the lower region of the stem. The latter
phenomenon is very remarkable, and, it is hoped, may act asa
check to the frequency with which new species are created to
separate two or more specimens of Aleyonaria which only differ
from one another in the shape of their spicules.

Genus STEREONEPHTHYA Kukenthal.

1869. Spoggodia (pars) Gray, in Ann. & Mag. Nat. Hist. (4)
vol. 111. p. 128.

1877. Spoggodia (pars) Klunzinger.

1889. “ Divaricate ” (Spongodes) Wright & Studer.

ALCYONARIA FROM SINGAPORE. HOW

1895. Spongodia (pars) Holm, p. 25.

1896. Spongodia (pars) Kiikenthal.

1904. Spongodia (pars) Holm, Weiteres tiber Mephthya und
Spongodes.

1905. Stereonephthya Kikenthal, p. 694.

“ Spongodes”-like forms in which the polyps were scattered
irregularly over the stem were recognised at an early date, and
were classified sometimes as a distinct genus, at others as a sub-
genus of Spongodes, under the name Spoggodia or Spongodia.
Wright and Studer (1889) recognised this group, but included
its members in the genus Spongedes under the designation
“ Divaricate.” Later writers, such as Holm (1895) and Kiiken-
thal (1896), agreed that Spongodes and Spongodia were synony-
mous. Hence Spongodes dragged Spengodia with it in its
downfall in 1905. For similar reasons to those stated in the
section on Dendronephthya I have adopted the new genus Stereo-
nephthya which Kiikenthal (1905) defined thus: ‘Sehr starre
Nephthyiden, deren Polypen weder in Liippehen noch in Biindeln
angeordnet sind, sondern einzeln oder in kleinen Gruppen direkt
vom Stamm wie den nicht oder wenig verzweigten Hauptiisten
entspringen. Polypen mit ‘ Stiitzbiindeln.’”

SrEREONEPHTHYA LUTEA, sp. n. (PI. LXITT, fig, 14,)

The specimen, on the observation of which this new species is
based, agrees in every respect with the diagnosis given by
Kiikenthal (1905) for the genus Stereonephthya, At the same
time it possesses characters which serve clearly to distinguish it
from other species hitherto described.

The colony is upright and branched. Numerous tapering
branches leave the insignificant stalk either singly or in groups,
and spread in all directions, but with a distinct tendency to
assume an upright position. When they arise in groups, the
members forming a group are frequently united for a considerable
distance by a common cortex. The primaries sometimes, but not
invariably, bear a few stationary branches, which also show a
tendency to grow in a vertical direction. From both the primary
and secondary branches short lateral lobes are given off at
irregular intervals.

Measurements :—

Hieicixtiof colony 10, .ie.neeorrea. TS AUR, Fa T: Khe (bem.
Breagth'at base ¥.,. shot tocar seca Ae stat Tea
Maximum breadth of capitulum ........,.....0...ce eee ores
ength of primary branches 72. Bod tes os 3°5-5'0 ,,
Breadth of primary branches { vi ik : ide i
Length by breadth of secondary branches ............ Tro 0°5: 4,
Length by breadth of lateral lobes ................00005 OOD? %,

Proc. Zoo. Soc.—1912, No. XX XV. 35

O22 MR, E. W. SHANN ON

The polyps arise sporadically from all parts of the colony.
They are closely aggregated on the lateral lobes, and are most
scarce on the basal portion of the primary lobes. The anthocodiz
are supported by a “ Stiitzbiindel ” (fig. 14), three horny spindles
of which project ‘2--4 mm. beyond the polvp-head. The polyp-
heads make an acute angle with their stalks ; the stalk is 1:2 mm.
high, the head is 1 mm. in length and ‘7-85 mm. in maximum
breadth. The polyps contain thorny spindles of various sizes
and a few clubs. These are sometimes placed horizontally, but
are more usually oblique, and in the distal area are arranged
en chevron.. These spindles measure °45x°035, °27x:04, -2x
"025 mm. Among the above and extending into the tentacles
are numerous little spicules which are nearly rectangular in
outline; they measure 110-036, :095 x -015, :050x-015 mm.
The ‘Stiitazbindel” is composed of a number of thorny spindles
of a bright yellow colour (they resemble at first sight crystals of
sulphur), These spicules are ten in number round the base of
the polyp-stalk. but there are only three at the distal end. The
four spicules which underlie the point of origin of the polyp-
head are semi-lunar in shape, the concave sides facing the
polyp-head, The “ Stiitzbiindel ” spicules reach 1-8 mm. in length
and -12 mm.in breadth. The cortex of the colony derives its
characteristic colour from the enormous number of slim yellow
spicules which it harbours. There are also thorny spindles; they
lie parallel to and very near the surface, and vary very much in
size and shape. Some are twice, others only half, the size of the
“Sttitzbiindel” spicules. Few are straight, most are semi-lunar
or S-shaped, and very exceptionally a triradiate form occurs.
The ends of the spindles are usually bluntly pointed, but
occasionally forms were observed with their ends frayed out mto
a number of points. The largest measure 3:0 +28, 2:7 x °30,
2-1 x15, the smallest °67 x -07, °67 x 04, 55 x 05 mm., but every
conceivable intermediate stage can be found, A remarkable
feature of this species is the entire absence of spicules from the
canal-walls.

The colour of the colony in alcohol is pale yellow, the polyps
are cream-coloured.

The specimen was taken in Imbiab Bay, where it was left
exposed at the lowest tides,

Two other specimens taken in the littoral region at Teluk Ayer
have also been assigned to this species. At first sight they do
not resemble closely the type specimen, for they are shorter,
broader in proportion, and more bushy in appearance than the
colony described above. One of these colonies is 4:3 cm. in
height, its base is °8 cm,, and the maximum diameter of the
capitulum is 3°l em.; the measurements of the other, taken in
the above order, are 3x°7 cm.: 45cm. Theapparent difference
in the branching from that of the type is due to the fact that
the secondary branches are no larger than the lateral lobes, and
that both these kinds of offshoot are relatively numerous. The
polyps are arranged much more densely than in the type,

ALCYONARIA FROM SINGAPORE. be

especially at the tips of the branches. The colour of these
specimens, moreover, is of a duller yellow. ‘These slight differences
appear quite insignificant when one considers that these two
specimens resemble the type of Stereonephthya lutea in the
following important characters :—

The size and shape of the anthocodie. The size, distribution,
and remarkable sulphur-yellow colour of the spicules; the dis-
tribution of spicules agrees even to the disposition of those
forming the “ Stiitzbiindel ”; the large internal canals, with thin
walls devoid of spicules.

Genus ParaspongopEs Kiikenthal.

P. crassa Kiikenthal, 1896.

This species was first described by Kiikenthal in 1896, and
further reference to it is made by the same author in his
‘Revision der Aleyonarien’ (1907), where it is described under
“Species incerti generis.” Two small colonies of a pale brown
colour are placed without hesitation in this species.

One colony was taken in 5-9 fms, of water from the New
Harbour. It measures 20 mm. in height; the diameter of the
capitulum is 13 mm., that of the stem 7°5 mm,

The other colony was taken in 5 fms. of water, the precise
locality is not recorded. Jt measures 20 mm, in height; the
diameter of the capitulum is 11 mm., and the stem is broken.

The only important character in which these specimens differ
from the type is the size of the spicules in the stem-cortex.
Measurements, length by breadth in millimetres, for three of the
largest spicules from this region are 1°2 x°18, 1:0 x-15, -95 x -22.
Kiikenthal found spicules measuring as much as 18 x ‘24 mm. in
the stem-cortex. The spicules from the Singapore specimens
show, however, the typical warts (:02 mm, high).

Previously recorded from Ternate, at a depth of 30 fms,

Family SIPHONOGORGIIDZ.
Genus SIPHONOGORGIA.
S. vARIABILIS Hickson (olim Chironephthya variabilis Hickson),
Two beautiful little specimens taken in 10 fms, of water south
of Blakang Mati are referred to this very variable species, Each
possesses a very slender stem, from the apex of which two main
branches are given off, so that the appearance of the colony is
roughly ¥Y-shaped. These primary branches give rise te secondary
branches, which in one or two cases are again divided. The
polyps occur most frequently on the tips of the branches, but a
few are scattered on all portions of the colony except the stem.

Measurements :-—
Specimen A. Specimen B.
mm, mm,
ileroht Of Sten. 27. seme eee ; 18 li
Diameter of; Stem cases eee ees 6 3
Length of primary branches ...... 23 and 23 24 and 18

524 MR. E. W. SHANN ON

The spicules are of the usual shape and of medium size, and
assume the colour of the part of the colony in which they are
embedded. ‘The large warty spindles of the ccoenenchym are
tinted with mauve and do not exceed 3:0 x°3 mm. In the polyp-
area the spindles are more frequent than the clubs ; all the polyp-
spicules are more or less bent, they are either bright red or bright
yellow in colour, and the larger ones measure *6x°05 mm,,
“47 x °065 mm. (the red spicules mainly compose the “ crown,”
the yellow ones the “ points”).

The colour of the colony in alcohol is a delicate pinkish mauve.
The polyps are deep red, and the tentacles bright yellow. It is
difficult to conceive how the colouring could have been more
vivid or the blended tints more pleasing even before these little
colonies were removed from their natural habitat,

For a list of the colour-variations to which this species is liable
to run, see Hickson’s ‘“‘ Aleyonaria of the Maldives,” Part I.
p. 488 (under Chironephthya variabilis), and Thomson and
Simpson (1909, p. 125).

Previously recorded from Mahlos Atoll, S. Nilandu, Persian
Gulf, Andamans, and the Arakan Coast.

Order GORGONACHA.
Suborder PSHUDAXONTA.

Family Sc LEROGORGIID&,

Genus SUBEROGORGIA.

S. SUBEROSA Pallas.

Gorgonia suberosa Pallas, Elench. Zooph. p. 191.

Suberogorgia suberosa Gray, Proc. Zool. Soc. 1857, p. 159.

Sclerogorgia suberosa Kolliker, Icon. Histiol. p. 142, pl. xix.
fig. 13 (2).

Sclerogorgia suberosa Studer, Monatsber. K. Akad. Wiss.
Berlin, 1878, p. 666.

Suberogorgia subercsa Ridley, Journal Linn. Soc. Zoology,
vol. xxi. p. 243.

Suberogorgia suberosa Wright & Studer, ‘Challenger’ Re-
ports, vol. xxxi. p. 166.

Suberogorgia suberosa Brundin, “ Aleyonarien aus dem Samm-
lung der Zool. Mus. in Upsala,” Svenska Vet.-Akad, Handl.
XXli. pt. iv.

There is in the collection a bottle which contains a number of
fragments, apparently the component parts of a single colony of
S. suberosa. The basal portion is present and has a diameter
of 6 mm.; the diameter of the terminal branches is 2°5 mm.
The specimen was taken in about 14 fms. of water off Pasir
Panjang.

ALCYONARIA FROM SINGAPORE. 625

Previously recorded from the coasts of Western Africa and the
West Indies (Pallas and Esper); the Mermaid Straits, Dampier
Archipelago, and N.W. Coast of Australia (Studer); Port
Denison, Queensland, and Torres Straits (‘ Alert’ Coll.); Mauritius
(Coll. Brit. Mus.); Admiralty Islands (‘Challenger ’ Coll.) ; and
Sumatra (Brundin).

Family MELITODID4.

Genus MELITODEs.

M. Avsitincta Ridley.

This species is represented by numerous fragments, but none
of these represents an entire colony. The specimens were taken
in 15 fms. of water from Blakang Mati.

Previously recorded from Port Molle, Queensland.

Genus PSILACABARIA.

P. GRACILLIMA Ridley.

Psilacabaria gracillima, n. gen. et sp., Ridley, Rep. Zool. Coll,
H.M.S. ‘ Alert,’ Aleyonaria, p. 363.

After a lapse of nearly three decades this delicate little species
is to be recorded once more. It is represented in the collection
by four portions of colonies, which show the mode of branching
and length of the internodes, and by sundry fragments. None
of the colonies are complete, but the largest intact portion
measures 45 mm. in height. The internodes vary in length, the
majority, however, fall within the range given by Ridley, viz.
12-16 mm. The branches are much more delicate than those of
the type, measuring only 1-2 mm. in diameter, as compared with
3-7 mm. Inthe mode of branching, size of polyps, and details
of spicule-distribution the Singapore specimens resemble the
type very closely. The spicules show the typical shapes; but
they are in each instance somewhat smaller than their prototypes.
The colour of the ccenosarc and polyps is fawn, that of the axis
white.

Locality. Salat Sinki, in 4-5 fms.

Previously recorded from Port Molle, Queensland, 12-20 fms. ;
Port Darwin, 8-12 fms.; H. Australia, 42 fms. (Ridley).

Genus WRIGHTELLA.

W. ropusta, sp.n. (Pl. LXIT. fig. 9; Pl. LXIII. fig. 15.)

This species has been formed to include a single well-developed
colony, which agrees more closely with Wrightella Gray (1870)
than with any of the other genera of the family Melitodide.
The form of the colony differs, in its stouter dimensions and in
its erect position, from the members of the four existing species
of Wrightella ; but it was thought expedient to associate it with

526 MR. E. W. SHANN ON

these, rather than to create a new genus from the observation of
a single specimen.

Wright and Studer (1889, p. xxxvi) give the following de-
finition of the genus Wrightella:—“The branches and twigs are
compressed ; the projecting polyp calyces occur especially on the
sides. In the cortex there are foliaceous clubs. There are no
nutritive canals in the axis.” The specimen fulfils all these
conditions.

The single main stem arises abruptly from a strong reticulate
base (fig. 9). The base, which is broken at the edges, measures
15 mm. in diameter, and the gaps in its meshes average | mm.
in diameter. The colony is 120 mm. in height and 45 mm. in
breadth (some of the lateral branches are broken, so it is probable
that the true breadth exceeded 45 mm.). The branching of the
colony is dichotomous, and takes place at the swollen nodes. The
nodes are less prominent in the distal branches. All the branching
takes place in one plane; the terminal twigs are markedly flattened
in the plane of branching. Anastomosis of the upper branches
takes place at infrequent intervals. The nodes near the base are
globular, having a diameter of 4 mm.; the internodes in this
region are circular in section, their diameter is 2°5 mm., which is
slightly exceeded by their length. The internodes beyond the
lowest three become more elongated and show an average length
of 10 mm. The termmal twigs are only 1 mm. wide. The
varruce, which measure *75 x75 mm., are not densely crowded,
and show a tendency to arrange themselves on the lateral aspects
even of many of the lower branches, but more especially in the
terminal twigs. Both cortex and verrtice are yellow in colour.
The axis 1s white, and is not traversed by nutrient canals.

The precise locality is not recorded, but, like the other speci-
mens in this collection, it was taken im shallow water near
Singapore.

The spicules attain all manner of shapes (fig. 15); they
are quite colourless, The foliaceous clubs characteristic of the
genus Wrightelia ave present in Jarge numbers, and show the
following range in measurement : length by breadth -27 x -12 mm.,
-15x-06 mm. ; handles of clubs ‘02 mm.in diameter. Numerous
spindles occur, some are foliaceous and measure *27 x:07 mm.,
°20x 08 mm., -12x-06 mm.; others are spiny, the spines fre-
quently being confined to the central region, and measure
°22x°05 mm, ‘17:03 mm, 12:03 mm. A few stellate
forms are found which have a diameter of -10-15 mm. Minute
scales abound; they measure 05-025, ‘036-030 mm.,
028 x:014 min.

Literature.

1870. Gray, J. E.—Cat. Lithophytes Brit. Mus. p. 31.
1900. Hicxson, 8. J., and Hines, Isa L.—‘‘ The Stolonifera and
Aleyonacea collected by Dr. Willey in New Britain,
” Willey’s Zool. Results, pt. iv; pps 493-508.

ete.”

1877.

1903.

1905.
1907.
1903.

1906.

IKON),

1889.

*Hio, 3
*Wig, 4
*Wio, 5
*Kig. 6
Wise, 7,
Fig. 8
Fig. 9
Fig. 10
Fig. 11
Fig. 12
Fie. 13
Fig. 14
Fig. 15

ALCYONARIA FROM SINGAPORE. By 7

Kuunzincer, C. B.—Die Koralithiere des Rothen Meeres,
Part I. p. 33, tab. 2. fig. 5

KiKenrHat, W.—“ Versuch einer Revision der Aleyo-
navien.” IT, Die Familie der Nephthyiden: 1 Theil.
Zool. Jahrb. Syst. xix.

Do. do. 24 Eheil.) .Op- cit. xo.

Do. dos W335 Uhenliss Opmetie xox

Prarr, Eorra M.—‘ The Alcyonaria of the Maldives.
Part Il. Sarcophytum, Lobophytum, Sclerophytwm, and
Alcyonium.” Fauna and Geog. of the Maldive and
Laccadive Archipelagoes. (J. Stanley Gardiner), Vol, 1.
Part I.

THomson, J. ARTHUR, and HenpErson, W. D.—‘‘ Alcyo-
naria of Zanzibar and British Hast Africa.” Proce.
Zool. Soe. pp. 393-443,

THomson, J. ARTHUR, and Srmpson, J. J.—‘‘ An Account of
the Aleyonarians collected by the Royal Indian Marine
Survey Ship ‘Investigator’ in the Indian Ocean.”
Part IT.

Wricut, E. P., and Sruper, TH.—* Seiten on the
Scientific Results of the Voyage of H.M.S. ‘ Challenger.’ ”
Vol. xxxi. Aleyonaria,

EXPLANATION OF THE PLATES.
Pruate LXI.

. Ammothea uineee ens Sav. (= Lithophytum arboreum Korsk.), entire colony ;
A th cens S 4 5 J

Savieny, Tab. 2. fig. 1. 1.

; Pee de chabrolit. Audouin, terminal lobe magnified; Savigny, Tab. 2.

fig. 5.

WE ‘chabrolii, single polyp magnified, lateral aspect; Savigny, Tab. 2.
fig. 5. 3.

: Same, abaxial axis; Savigny, Tab. 2. fig. 5. 4.

. Same, more highly maguitied; Savigny, Tab. 2. fig. 5. 5.

Prate LXII.
. N. chabrolié, entire colony ; Savigny, Tab. 2. fig. 5. 1.

. Sclerophytum pinnilatum, sp. u., entire colony ; from a photograph,

slightly reduced.

. Nephthya bedfordi, sp. n., entire colony; from a photograph, natural

S1Ze.

. Wrightella robusta, sp. n., entire colony; from a photograph, X 4.

Prate LXIIT.

. Sclerophytum pinnulatwn, sp. n., tentacle bearing pinnules, X 245.

. Nephthya bedfordi, sp. n., spicules, X 72.

2. NV. bedfordi, terminal lobe bearing polyps, X 25.

; Dendronephthya disciformis Kiukth. (aberrant specimen), spicules from

the stem-cortex, X 72.

. Stereonephthya lutea, sp. n., polyp and “Sttitzbtindel,’” x 72.
. Wrightella robusta, sp. n., stem-spicules, X 72.

* Figures marked thus are reproduced from photographs procured by Professor

Bourne,

and kindly given by him to me for the purpose.

528 MR. T. H. WITHERS ON

27. Some early Fossil Cirripedes of the Genus Svalpellum.
By Tuomas H. Wrruers, F.G.S.*

[Received January 31, 1912: Read March 19, 1912.]
(Text-figs. 64 & 65.)

INDEX.
Page
Structure of carina of Scalpellam ........0000...00.--. 528
Evolution of early fossil forms of Sealpellum ...... 530
Distribution (Geological) of Cretaceous Scalpellum 537
Scalpellum arewatwm Darwin ............0esssceeeseeeee 582
Sh WRU IDETAWAIN (oponas0s000de3ax0dsIGecaasaa0003009 SBI
Scalpellum spp.—discussion of affinities ............ 538

Among the Cirripede remains from the Albian (Gault) of Folke-
stone in the collection of the British Museum (Natural History),
three detached valves were noticed, which, for certain reasons to
be explained later (see p. 532), appear to be the original valves
upon which Darwin founded the species Scalpellwm arcuatum *.
A portion of one of the valves, a carina t, happened to be broken
away from its matrix, showing that the intraparietes, said by
Darwin to be absent in this species, were really present. These
parts, however, are developed in such a way that they can be
seen only by cleaning the valve free from matrix. The finding
of the intraparietes in Darwin’s type of 8. arcuatwm led to the
examination of further examples of the carina, and the fact was
established that intraparietes are developed in all carine of this
species.

S. trilineatum Darwin (1851, p. 38, pl. i. fig. 5), from the
Cenomanian (Grey Chalk) of Dover, was said by Darwin to come
nearest to S. arcuatum, owing to the absence of intraparietes.
The carina, which is the holotype of S. trilineatwm, is in the
British Museum (Natural History), registered 38461. On freeing
this specimen from the gum and matrix which obscured the inner
portion of the valve, it was at once apparent that not only were
the intraparietes present, but that they were developed almost
exactly as in S. arcuatum.

Other Lower Cretaceous species of Scalpellwm in which the
intraparietes of the carina are said to be absent are S. simplex
Darwin (1851, p. 39, pl. 1. fig. 9) and 8. accumulatum Withers §,
both of which come from the Aptian (Lower Greensand). The
unique carina of S. simplex cannot at present be traced, but since
the parietes do not reach to the basal margin of the valve (see
text-fig. 64, 7) as in those carine which have intraparietes, it
does not appear probable that intraparietes could be present. In

* Communicated by Dr. W. T. Caiman, F.Z.S.

+ C. R. Darwin, 1851, Pal. Soc. Monogr. Foss. Lepadide, p. 40, pl. i. fig. 7.
t+ For the names of the various valves see text-fig. 65.

§ 'T. H. Withers, 1910, Geol. Mag. dee. v. vol. vii. p. 152, figs. 1-4.

FOSSIL CIRRIPEDES. 529

S. arcuatum (text-fig. 64, 3) and S. trilineatum (text-fig. 64, 4)
the parietes reach to the basal margin, and in my opinion the
intraparietes will be found to be developed in S. accumulatwm
(text-fig. 64, 2) as in those species, since S. accuwmulatwm is
similar in its general external form. The only carina known

Text-fig. 64.

\

\\
.
MN
HX,
\\
\\

>
zg

Early types of the Carina of Scalpellum.
1a-5a, side view; 36, 46, 5 6, inner view; 1 c-5 c, transverse section near apex.
t., tectum ; p., parietes; 2.p., intraparietes.

of S. accumulatum is embedded in such a hard matrix that it is
impossible to expose its inner surface without danger to the speci-
men, so that the matter must here remain until further specimens
are forthcoming.

So far, then, we have proved the existence of intraparietes in

530 MR. T. If. WITILERS ON

the carinz of the two species S. arcuatum and WS. trilineatwm, and
pointed out the possibility of their presence in S. acewmulatum.
The intraparietes in these species, instead of forming a thin wall
on each side of the carina as in the form represented by text-
fig. 64, 5, are bent inwards almost at right angles, and the upper
regions of their inner margins meet to a greater or less extent
(see text-fig. 64, 3b, 4b); the upper part of the valve is solid, and
must have projected freely to the extent indicated by the meeting
of the intraparietes*. The peculiar development of the intra-
parietes in the caring of these species is therefore of importance
as showing a development of the carina distinct from that in
which the intraparietes form a thin wall on each side of the
carina.

Three types of carina, all having an apical umbo, were there-
fore already developed amongst these early forms of Scalpellwm,
and the geologically oldest of these more closely resemble the
carina of Pollicipes, from which Scalpellum is considered to be
derived. These are (1) represented by S. simples (text-fig. 64, 7)
from the Aptian (Lower Greensand), which has no intraparietes,
the tectum being flatly-arched transversely, the parietes bent
almost at right angles to the tectum and not extending to the
basal margin ; this type is distinguished from the Pollicipes type
of carina only in the parietes being separated from the tectum
by a distinct angle; (2) represented by S. acewmulatum (text-
fig. 64, 2) (Aptian, Lower Greensand), S. arcwatum (text-
fig. 64, 3) (Albian, Gault), and S. trilineatwm (text-fig. 64, 4)
(Cenomanian, Grey Chalk), which have intraparietes, these parts
being bent inwards almost at right angles and joining, the upper
part of the valve being solid and projecting freely to a consider-
able extent ; in the solidity and free projection of the upper part
of the valve this type is allied to Pollicipes ; (3) represented by
S. hastatum (text-fig. 64, 5) and other species from the Ceno-
manian which have intraparietes also, but are characterized by
these parts forming a thin wall on each side of the carina, the
apex of which projects freely, slightly, or not at all; this latter
type is more typical of the genus Scalpellum, and, owing to the
upward growth of the intraparietes in some forms, subsequently
gave rise to the species with an angularly bent carina having the
umbo in a subcentral position, a type which is not known below the
Upper Senonian. The only species with an angularly bent carina
known from the English Chalk is S. darwinianwm Bosquet tT, but
this has an early specialized form of carina in which the upward

* A somewhat similar development of the intraparietes can be seen in the carinze
of S. maximum var. cylindracewm Darwin (1851, p. 38, pl. ii. fig. 2) from the Upper
Senonian (Belemnitella mucronata-zone) of Norwich, Norfolk, and S. solidulum
Steenstrup (1839, Kr@yer, Naturhist. Tidsskrift, Bd. 11. p. 412, pl. v. figs. 14, 14*;
Darwin, 1851, Pal. Soc. Monogr. Foss. Lepadide, p. 42, pl. i. fig. 8) from the Upper
Senonian of Kjuge, Scania.

+ J. Bosquet, 1854, Les Crust. Foss. du Terrain Crétacé du Duché de Limbourg,
p. 46, pl. iv. figs. 6-12; T. H. Withers, Jan. 1911, Geol. Mag. dec. v. vol. viii.
p. 23, figs. 3-4.

FOSSIL CIRRIPEDES. 531

extension of the valve is due, not merely to the upward growth of
the intraparietes, but to the almost equal upward and downward
growth of the valve from the umbo, the whole external surface of
the valve being ornamented. It is, therefore, probable that
species with an angularly bent carina, due to the upward growth
of the intrapavietes, which is apparently the more primitive type,
existed even earlier than in the Upper Senonian *.

Extreme interest attaches to the remarkably complete example
of S. arcuatum from the Albian (Gault) of Folkestone, Kent, here
described and figured (p. 534, text-fig. 65, 6). To find so many
valves of the capitulum in position is a remarkable circumstance,
very few specimens of Scalpellum having been discovered with
several valves associated. Those known up to the present come
from the Upper Senonian, and, with the exception of a single
incomplete capitulum of S. maximum, belong to the species S. fos-
sula. This specimen, therefore, represents the only fossil species
of Scalpellum obtained from below the Upper Senonian with any
considerable number of valves in pesition. Previously described
species of Scalpellum from rocks below the Upper Senonian have
all been founded on detached valves, and in most cases on single
valves. Neither the upper latus, rostral latus, or rostrum, are
included in these descriptions, but are represented in the present
specimen. A further example of S. arcwatum (described, p. 533)
has furnished some scales of the peduncle, which so far are not
known in any species found below the Upper Senonian. S. arcu-
atum is the only representative of Scalpellum known from the
Albian (Gault) of England.

Consequent on the discovery of the intraparietes in the carine
of S. arcuatum and S. trilineatwm, and since the carina is the
typical valve of the genus, it is here proposed to give fresh
diagnoses of those species and to describe the new material of
S. arcuatum. From the two examples of S. arcwatum it is
possible to construct a restoration of the capitulum, to which only
two valves are diagrammatically added (see text-fig. 65, 7, p. 534).

SCALPELLUM TRILINEATUM Darwin.

1851. Scalpellum trilineatum C. R. Darwin, Pal. Soc, Monogr.
Foss. Lepadidee, p. 38, pl. 1. fig. 5.

1854, Scalpellum trilineatum C. R. Darwin, Ray Soc. Monogr.
Sub-class Cirripedia, Balanide, Synopsis et Index
Systematicus, p. 633.

1854, Scalpellum trilineatum J. Morris, Cat. British Fossils,
2nd ed., p. 97.

1877. Scalpellum trilineatum H. Woodward, Brit. Mus. Cat.
Brit. Foss. Crustacea, p. 143.

* Since the above was written, two exceedingly small, angularly bent carinz
(length respectively 1°7 mm. and 1°8 mm.) with the umbo subcentral, due to the
upward growth of the intraparietes, have been obtained by Mr. F. Mockler from the
Cenomanian (Chalk Marl) near Cambridge. The specimens are incomplete and

poorly characterized, but of much importance as showing that species with an
angularly bent carina existed even so far back in tyne*as the Cenomanian.

532 MR. '. H. WITHERS ON

This species was founded on a carina and tergum from the
Cenomanian (Grey Chalk) of Dover. The tergum cannot now be
found, but the carina, which Darwin considered to be the typical
valve of the genus and can therefore be regarded as the holotype,
is in the British Museum (Natural History) registered 38461.

Diagnosis.—Carina with three prominent, rounded, longi-
tudinal ridges on its tectum—one central, and one on each side
separating the tectum from the parietes; intraparietes bent
inwards almost at right angles, the inner margins meeting a
short distance below the apex, upper part of valve solid and
projecting freely.

Description of Carina.—Carina narrow, widening very gradu-
ally from the apex, considerably bowed inwards, basal margin
obtusely angular. Tectum flatly-arched transversely, with a
central, prominent, rounded ridge extending from the apex to
the basal margin, and bounded on each side by a slightly coarser
but flatter ridge on the angle separating the tectum from the
parietes. Parietes narrow, less than half the width of the tectum,
bent almost at right angles to the tectum, slightly concave.
Intraparietes very narrow, bent inwards almost at right angles,
the inner margins meeting about one-sixth the length of the
valve from the apex; the upper part of the valve is solid, and
must have projected freely to the extent indicated by the meeting
of the intraparietes. Lines of growth plainly marked.

SCALPELLUM ARCUATUM Darwin. (Text-fig. 65.)

1851. Scalpellum arcuatwum C. R. Darwin, Pal. Soc. Monogr.
Foss. Lepadidee, p. 40, pl. i. fig. 7.

1854. Scalpellum .arcuatum C. R. Darwin, Ray Soc. Monogr.
Sub-class Cirripedia, Balanide, Synopsis et Index
Systematicus, p. 633.

1854. Scalpellum arcwatum J. Morris, Cat. British Fossils, 2nd ed.
p. 96.

1865. Scalpellum arcuatum J. W. Salter & H. Woodward
Cat. & Chart Foss. Crustacea, p. 27, pl. i. fig. 14.

1877. Scalpellum arcuatum H. Woodward, Brit. Mus. Cat. Brit.
Foss. Crustacea, p. 142.

The species S. arcuatum was founded on three detached valves,
namely, carina, scutum, and tergum, from the Gault of Folke-
stone, Kent. These valves were considered by Darwin to belong
to the same species, and the material now to be described proves
such to be the case. Darwin further stated that these valves
were in the Bowerbank Collection. This collection was acquired
in 1865 by the British Museum (Natural History), and among
the specimens are three valves, a carina (I. 13796), a scutum
(1. 13797), and tergum (I. 13798), mounted together on Bower-
bank’s original tablet, and labelled in Darwin’s handwriting
“iS. arcuatum.” The carina has been much broken, presumably
since Darwin described it, but the scutum and tergum are in good

FOSSIL CIRRIPEDES. 533

condition. These specimens are also exactly half the size of
Darwin’s figures, which are enlarged to two diameters, and in
these circumstances there seems little doubt that they are the
original valves figured by Darwin. The carina, since Darwin
considered it to be the typical valve of Scalpellum, is consequently
regarded as the holotype.

Two specimens in the British Museum (Natural History) add
considerably to our knowledge of this species.

One, registered (I, 13577), text-fig. 65,6, first appeared to
consist of a carina with fragments of the scuta and terga. Careful
clearing away of the matrix and the abundance of gum which
covered the specimen showed, however, that several of the valves
of the capitulum were preserved. These consist of the carina, the
paired scuta, the paired terga, an upper latus, a rostral latus, and
the rostrum. The left side of the capitulum is uppermost, and the
carina is so exposed that the intraparietes can readily be seen.
Portions of the inner surfaces of the right scutum and tergum are
also exposed owing to the fact that the upper valves are somewhat
displaced and broken.

The second specimen (I. 13580) has not so many valves pre-
served and those present are somewhat broken. The right side is
uppermost, and the plates shown are the carina, the pair of
terga, the upper portion of the left scutum showing its inner
surface, and.the left upper latus. This last valve showed only its
inner surface, but on removal from the matrix it was found to
be the left upper latus. This specimen is of interest since it
furnished eighteen scales of the peduncle. A supposed shell-
fragment was removed from between the two terga, and on being
cleaned was found to be a peduncle scale. On removing and
washing the remaining matrix from the same position, the
number of scales was increased to eighteen. There is little
doubt that these scales belong to S. arouatum, since it is the only
species of Scalpellum known to occur in ‘the Gault. Moreover,
they are ridged like the valves of the capitulum of that. species,
and the fact that they were found in such close association is
prima facie evidence that they belonged to the same individual, and
were washed into the position in which they were found. They are
somewhat similar m shape to the scales of the peduncle of the
Upper Senonian species S. maximum and 8. fossula as figured by
Th. Marsson *, but are easily distinguishable from them by their
longitudinal ridging.

Diagnosis.—Capitulum composed probably of fourteen valves f,
which are ornamented with numerous fine ridges radiating
from their apices; umbo of all valves apical. Carina with
tectum flatly arched transversely, parietes rectangularly inflected,

* Th. Marsson, 1880, “ Die Cirripedien und Ostracoden der weissen Schreibkreide
der Insel Riigen,” Mitth. naturwiss. Vereine Neu-Vorpommern und Riigen, xii,
pl. i. figs. 25, c, d, 3d, e.

+ It is possible that this species had a subcarina, in which case the number of
valves would be fifteen. A higher number of valves is not. likely.

MR. T. HW. WITHERS ON

Text-fig. 65.

SS

Fig. 6. Scalpellum arcuatum Darwin. Nearly complete capitulum showing the

Fig. 6a.
Vig. 6b.
Fig. 6c.
Fig. 6d.

Fig. 6 e.

left side uppermost, with the valves somewhat displaced and broken. X 2
diam, Albian, Gault: Folkestone, Kent. Brit. Mus. (Nat. Hist.), I, 13577.
c., carina, showing the inflected intraparietes; ¢.1, outer portion cf left
tergum; ¢.2, inner view of right tergum near apex, showing the ridges
evidently connected with the firm attachment of the corium; s.1, outer
portion of left scutum ; s.2, inner view of right scutum showing the pit
tor the adductor scutorum; U., upper latus; 7./,, rostral latus with upper
portion broken off; #., rostrum. :

Id. Outer view of incomplete rostral latus,

Id, Inner view of same.

Id. Outer view of rostrum. x 4 diam.

Id. Hypothetical transverse section of same.

Id, Side view of same.

Fig. 7. Restored capitulum of Sealpellum arcuatwm Darwin. This figure is based

on the nearly complete capitulum represented by fig. 6, with the addition
of a carinal latus and an infrasmedian Jatus. These two yalves have not yet
been discovered in the Gault. XX 2 diam, Albian, Gault; Folkestone,
Kent. ./., carinal latus ; 7./., infra-median latus.

Fig, 8, Sealpellum arcuatum Darwin. Two scales of the peduncle. a, outer view,

b, inner view; ¢c, outer view; d, mner view. X 8 diam. Albian, Gault;
Folkestone, Kent, Brit, Mus. (Nat. Hist.), I, 1880,

FOSSIL CIRRIPEDES. aeay3)

intraparietes bent inwards and meeting for about one-fourth the
length of the valve from the apex, upper part of valve solid and
freely projecting. Scutum with tergo-lateral angle almost in line
with the middle of the valve. Tergum subrhomboidal, a delicate
furrow extending from the apex to the basal angle. Upper latus
subtriangular, apex acutely angular, slightly bowed towards scuta,
basal margin rounded. Rostral latus acutely angular transversely,
about 24 times as long as wide. Rostrum subtriangular, with a
strong median keel extending from the apex, widening towards
the convex basal margin,

Description of valves.—Carina narrow, widening gradually from
the apex, considerably bowed inwards, basal margin obtusely
V-shaped. Tectum flatly-arched transversely, obscurely carinate,
and ornamented with numerous fine longitudinal ridges. Parietes
narrow, less than half the width of tectum, not longitudinally
ridged, bent almost at right angles to the tectum, slightly concave,
Intraparietes very narrow, bent inwards almost at right angles,
the inner margins meeting about one-fourth the length of the
valve from the apex, above which the valve is solid and must
have projected freely,

Scutum moderately convex, divided unequally by a prominent
ridge running from the apex to the basi-lateral angle, the basi-
lateral angle being slightly produced, Apex acuminate. Basal
margin sinuous, about two-thirds the length of the valve;
eccludent margin slightly convex and nearly parallel to the lateral
margin ; tergal and lateral margins of almost equal length and
forming an angle of about 145°, either margin being about half
the length of the valve. Surface of valve ornamented with fine,
closely-set, longitudinal ridges; a narrow slip along the tergal
margin is somewhat bent downwards and is devoid of ridges,

Tergum sub-rhomboidal in general outline, slightly convex,
with a delicate furrow extending from the apex to the basal
angle. Apex and basal angle acuminate, more so than is indicated
in Darwin’s original figure; scutal angle somewhat protuberant,
Carinal margin convex ; scutal margin sinuous, longer than the
ocecludent margin, which is nearly straight. Surface of valve -
ornamented with numerous fine longitudinal ridges. Inner
surface, in the region of the apex, is marked on its edges with
oblique lines of growth, these indicating the extent to which the
valve projected freely. A little below the apex, nearer to the
occludent margin, are three or four small ridges ending abruptly
about one-fourth the length of the valve from the apex. These
ridges were evidently connected with the firm attachment of the
corium, or membrane which lined the inner surfaces of the valves,
and are homologous with the series of tubercles on the inner
surfaces of the terga of S. darwinianwm Bosquet.

Upper latus subtriangular, slightly curved towards the scuta,
almost flat transversely, convex longitudinally ; umbo slightly
projecting, with a thick ledge formed beneath it, which thins out
towards the lateral angles; tergal margin slightly convex ; scutal

536 MR. T. H. WITHERS ON

margin slightly concave and about the same length as the tergal
margin, the two margins if represented by lines from the apex to
the lateral angles would enclose an angle of about 55°; basal
margin rounded, and indistinctly marked off into three lines ;
a portion of the valve on either side, parallel with the scutal and
tergal margins, is somewhat raised, and the lines of growth on
these parts are upturned sharply towards the umbo. Surface of
valve between the raised portions ornamented with several longi-
tudinal ridges.

Rostral latus.—This valve is imperfect, the upper portion being
broken off. Valve very narrow, acutely angular transversely,
about 24 times as long as wide, widening gradually from the
inner acute extremity to the rostral margin, which is abruptly
truncate; umbo apical; the inner extremity is marked by a
strong rounded keel; outer (rostral) half of valve ornamented
with fine longitudinal ridges, two fine ridges close together almost
dividing the basal margin into two equal portions. At the point
where these two ridges reach the basal margin the valve is some-
what convex, no doubt indicating the extent to which the valve
was bounded by the inframedian latus.

Rostrum sub-triangular, strongly convex transversely, bowed
inwards ; lateral margins bounded by strong ridges; basal margin
convex ; a strong median rounded keel extends from the apex,
widens considerably towards the basal margin, and is bounded
on either side by indistinct longitudinal ridges.

Peduncle scales varying in shape from semilunar to trapezoidal,
the basal margin of the former being straight, while that of the
latter is somewhat concave; immediately below the apex the
trapezoidal scales are slightly constricted, the truncated top
appearing to overhang ; scales thickest at one-third from the base,
below which, on the inner surface, they are somewhat excavated,
this indicating the extent to which the scales were covered by the
corium, the upper two-thirds no doubt overlapping the contiguous
scales. Outer surface ornamented similarly to the valves of the
capitulum with a number of longitudinal ridges, These ridges
number about seven on the larger scales, and four or five on the
smaller scales.

Measurements,—Owing to the fact that the valves in these two
specimens of S. arcwatwm are broken, it is impossible to give
their accurate measurements. Since, however, it is desirable
that we should have some idea of the relative sizes of the valves
in an individual, approximate measurements are given where the
correct measurements cannot be obtained,

Specimen J, 13577,

Carina. Length circa 21 mm,

Scutum (right valve). Length (from apex to rostral angle)
13°5 mm.; breadth 7:4 mm.

Tereum., Length circa 17 mm,; breadth 8°2 mm,

FOSSIL CIRRIPEDES. 537

Upper latus. Length 6°8 mm.; breadth cirea 7-5 mm.
Rostral latus. Length 6°38 mm.; breadth 2:6 mm.
Rostrum. Length 3-4 mm.; breadth 2:2 mm.

Specimen IJ. 13580.

Carina. Length circa 20 mm.

Tergum. Length cirea 17 mm.; breadth 7-5 mm.

Upper latus. Length 5°9 mm.; breadth cirea 4-2 mm.

Scales of peduncle. Length from 0-5 mm. to 1-2 mm.; breadth
1:3 mm. to 2 mm.

Remarks, and comparison with other Species —The restoration
of S. arcuatum (text-fig. 65, 7) is based on the nearly complete
capitulum figured on the same page, and, to complete the capitu-
lum, a carinal latus and an inframedian latus have been added.
The carinal latus figured in the restoration was found amongst a
number of detached plates of S. arcwatwm from the Cambridge
Greensand, and is longitudinally ridged as in the valves of
S. arcuatum. It possibly belongs to the same species. In any
case the only carinal latera known to the writer from the Lower
Cretaceous rocks are of the type figured, although they evidently
belong to several different species. Judging from the hiatus
between the carinal latus and rostral latus, an inframedian
latus was undoubtedly present, and was probably very like the
homologous valve in S. fossula from the Upper Senonian.

The specimen of S. arcuatwm here figured (text-fig. 65, 6) is,
up to the present, the oldest fossil Scalpellum from which one
can gain any idea of the appearance of the complete capitulum.
It comes from the Albian (Gault) of Folkestone, Kent, and the
only undoubted valves of Scalpellum older than this oceur in the
Aptian (Lower Greensand) *. These Lower Greensand forms
comprise only three species, two of which—vJS. simples Darwin
and S. accumulatum Withers—are respectively represented by
a single carina; the third, 5. comptwm Withers, is represented
by two detached terga.

Our knowledge of these early forms of Scalpellwm is therefore
not very extensive, and the fact that they are represented by
such a small number of valves, and those only of carine and
terga, serves to emphasize the importance of this fine example of
S. arcuatum (text-fie. 65, 6).

A comparison of the carina and tergum of S. arcwatwm with the
corresponding valves of S. sclidwhwm Steenstrup, as figured by
Darwin (1851, p. 42, pl. i. fig. 8), shows how closely these two
species resemble each other. They are evidently related. The

* This statement is made with full knowledge that certain valves from the
Paleeozoic and Jurassic rocks have been referred by various authors to the genus
Scalpellum. There is, however, not sufficient evidence to justify the reference of
these valves to the genus Scalpellum. Notwithstanding this, it is possible that
some of the Mesozoic Cirripedes referred to Pollicipes may eventually be shown
to be ancestral forms of Scalpellwm, but this cannot be done until more is known of
the various valves comprising the capitulum.

Proc. Zoou. Soc.—1912, No. XXXVI. 36

5388 ON FOSSIL CIRRIPEDES.

carine are easily distinguished, for whilst in S. arcuatwm the
tectum is flatly-arched transversely, and the intraparietes are
bent inwards almost at right angles, the tectum in WS. solidulwm
is strongly convex, and the intraparietes join to form a prominent
crest. Darwin considered the scutum of S. solidulum to be like
that of S. arcwatum, with the exception of the longitudinal ridges
being proportionally broader and further apart, closely resembling
those in the carina of S. solidulum.

Unfortunately the only known complete capitulum from the
Cretaceous rocks with which that of S. arcwatwm can be compared
is that of the Upper Senonian species S. fossula. In the relative
positions of the valves both species are alike, but in the structure
of the carina and scutum there are important differences. The
intraparietes of the carina of S. arcwatwm are sharply bent
inwards, the upper part of the valve is solid and must have
projected freely to some considerable extent. The carina of
S. fossula, on the contrary, projected freely but little, and the
intraparietes form a thin wall on each side of the carina. In
S. arcuatum the tergo-lateral angle of the scutum is situated
much further from the apex than in S. fossula, and im this
respect is further removed from the more advanced scuta which
have the tergo-lateral angle almost in line with the apex, above
which the valve is added to, the umbo consequently being sub-
central. The valves of S. arcuatwm are longitudinally ridged,
while those of S. fossula are smooth.

Affinities of the Species mentioned.

S. arcuatum is no doubt an ancestral form of a group of almost
exclusively deep-sea species, which Dr. P. P. C. Hoek* has
separated as a sub-genus under the name <drcoscalpellwm.
S. trilineatum, S. accumulatum, S. conptum, S. maximum var.
cylindraceum, and S. solidulwm, which appear to be related to
S. arcuatum, possibly belong to the same group, but we know too
little of these species to say much about them. SS. simplex
probably does not belong here.

The species embraced by the sub-genus <Azcoscalpellum have
no sub-carina, and it is impossible at present to say whether
S. arcuatum had a sub-carina or not. The Senonian species
S. fossula also comes nearest to the subgenus Arcoscalpellum, but
this species is said by Ed. Hébert to have a sub-carina, and
in view of this it is possible that the Albian S. arcwatwm also
may have had one. Assuming this to be the case, we have two
forms agreeing in all essential~characters with the forms of
Arcoscalpellum, except that they (possibly) have a sub-carina.
Moreover, the two species differ in the form of the carina, S. arcua-
tum having the intraparietes bent inwards at right angles and

* P. P. C. Hoek, Oct. 1907, Siboga-Expeditie, Cirripedia Pedunculata, p. 59.

+ Ed. Hébert, 1855, Mém. Soc. Géol. France, 2° sér. vol. v. p. 356, pl. xxviii. fig. 1
(S. gallicum = 8. fossula).

1D ZS), Ua, Weak, AL INW!

West,Newman ad nat. chr.

FPRATHERS FROM PHEASANTS, 3d &°.

a * “
‘i ern - ig ee EE EE

ayo yeu pe WemMsahNy “ASsoM

TSE Wel BUlsi) 1S A cl

ce
Eee

4

Ash de), Weal, Loe.

West, Newman ad nat chr

FEATHERS FROM PHEASANTS, &.

PA Selo dae Ea eynte

,

9

West,Newman ad nat. chr.

I 8

FEATHERS FROM PHEASANTS, ¢.

ON EXPERIMENTAL PHEASANT-BREEDING, 539

fo)
freely, while the carina of S. fossula has the intraparietes

forming a thin wall on each side of the carina, which is a more
advanced type. There is not as yet suflicient evidence to prove
whether S. fossula was derived from S. arcuatem or not. It is
patent, however, that from the foregoing species subsequently
arose the forms grouped as Arcoscalpellum and characterized by a
reduction in the number of plates of the capitulum, by a sup-
pression of the rostrum, and a tendency towards the reduction
of the pair of infra-median latera.

joining, the upper part of the valve beine solid and projecting
J > proj s

For help in connexion with this paper I wish to express my
indebtedness to Dr. F. A. Bather, Dr. W. T. Calman, Mr, C. P.
Chatwin, and Dr. P. P. C. Hoek.

Key to Species mentioned.

A. Carina without intraparietes, parietes not
reaching to the basal margin and bent almost
at right angles to the tectum .....................

B. Carina with intraparietes bent inwards and
joining, the upper part of the valve solid and
projecting freely.

1. Carina with basal margin rounded...............
2. Carina with tectum flatly-arched transversely,
and marked with numerous fine longi-
huaiivallinid es! c:.....5 eee eRe meee eees en.
3. Carina with tectum flatly-arched transversely,
and marked with three prominent longi-
tudinal ridges, one central, and one on
either side separating the tectum from the
parietes Se CREO sahnes cnechaada ca neeacemee nan
4, Carina with tectum strongly convex trans-
versely and marked with several longitudinal
SHG FESS pwc AAR IRE RRs ean oa nG ei ae eee
5. Carina with tectum strongly convex trans-
versely with smooth surface............ ........ S. maximum var. cylindra-

©, Carina with intraparietes forming a thin wall [eewm Darwin.
on each side of the valve.

1. Carina with tectum and parietes smooth,
dorsal surface and inner margin much

S. simplex Darwin,

S. accumulatum Withers.

S. arcuatum Darwin.

S. trilineatum Darwin.

S. solidulum Steenstrup sp.

PPKCUEWTEGL soo osnenu een oeanoo soc ahoacoconsses-hocounasea So aSamuoD IDEAL,
2. Carina with tectum bordered on each side by
a large, protuberant, flat-topped ridge ... .. S. fossula Darwin.

28. Experimental Pheasant-breeding.

By Rosr Haig THomas, F.Z.S.
[Received December 4, 1911; Read February 6, 1912. ]

(Plates LXIV.-LXVIL.*)

The experiment with which I deal in this paper was under-
taken to test the truth of the result of one previously made
(P. Z.S. 1909, pp. 884-885), in which it was shown that a male
Pheasant had transmitted to his female offspring of the second

* For explanation of the Plates see p. 545.
36%

540 MRS. R. HAIG THOMAS ON

generation the female plumage of his species, under the following

scheme of mating :—
Silver 2 X Swinhoe g

a)
F.1 2 X Swinhoe 3
ee te e

(Q) F.2 9 Sw. @) eee (3) 3 Sw. hyb.

It is to be observed that the Swinhoe male transmitted the
pure Swinhoe plumage only to one of his male offspring. As
such an isolated instance, though interesting, had small value
without the support of other evidence, the following experiment
was arranged. I chose species which have mature plumage in
five months from hatching, so as to get quicker at the results,
instead of, as in the case of gennewus, waiting eighteen months
for the adult plumage of F.2. Phasianus formosus, one of the
species selected, is found only on the Island of Formosa ; somewhat
resembling P. torquatus, it differs from it in one special feature ;
in the cock, the collar, instead of being an even white ring
forming a complete circle round the neck, is formed by a deep
Vandyke on each side with a narrow connecting-line round the
back of the neck but not connected round the throat, an incomplete
circle. The other species, P. versicolor, the Japanese Pheasant, is
too well known to require description ; it has no white collar.

Below is appended a table of the differences between the males
and females of the two species which might be expected to prove
Mendelian pairs.

P. formosus 3. P. versicolor 3.
Moult: Rapid, early. Moult: Slow, late.
Leg: Les:
Colour: Pale bluish grey. Colour: Wark, reddish grey. ©
Dimension: Thick. Dimension: Thin.
Bill: Large. Bill: Small.
Crest: Pale. Crest: Dark.
Breast : Rich brown. Breast : Metallic green.
Tail: . Long. Tail: Short.
All rectrices banded. Some laterals not banded.
Neck: White collar. Neck : White collar absent.
Bird; Large. Bird: Small.
P. formosus 2. P. versicolor 2.
Leg: Leg ;
Colour: Pale bluish grey. Colour: Reddish grey.
Dimension: Thick. Dimension: Thin.
Bill: Large. Bill: Small.
Colour: Greyish green, paler at Colour: Darker greyish green, same
gape. : all over.
Crest; Pale ground, black oval | Crest: Dark ground, irregular
spots. black patches.
Breast : Pale buff, no pattern. Breast : Dark buff, thickly patterned.
Tail; Banded pattern on pale | Tail: Banded pattern on dark
ground. ground.
Bird: Large. Bird: Small.
Kgg: Large. Hee: Small.
Calin temperament; quiet, tame. Excitable temperament: wild, untame-

able.

BXPERIMENTAL PHEASANT-BREEDING. 5Al

Scheme of mating in second experiment :—

P. formosus 2 X P. versicolor 3
|

a ee ps)

(te ae (= (eae ee
(3) F.1 3 F.19“Y” F.19“Z” X P. versicolor
SS Se

(A mais | aa
(6) F.2' 2 Ve: (2) 3 Ve. hyb.

The second experiment confirmed the results obtained in the
first.

Four skins of the F. 2 2’s are exhibited (the fifth female has
been kept to breed from), together with skins of pure Versicolor
and pure Formosan females, also the skin of the F.1 2 “Z” parent.
of F. 2. If these are all turned breast uppermost, it is seen at a
glance that Versicolor pattern and coloration are present to a
certain extent in F.1 2 ‘“ Z,” mother of the F. 2 generation, and
that the F. 2 females appear to be identical with Versicolor in size,
pattern, and coloration, except for small differences which might
be found existing between individuals of any species. Besides
plumage, these five hens had the habit and temperament of the
Versicolor, the leg-colour and eye-skin also.

The records of this experiment constantly refer to the quick
sharp movements, the wild scared appearance of the F. 2 females,
so characteristic of the habits and ways of, the untameable
Versicolor. If the pen was entered for any purpose, even when
exercising the greatest care, there was always a chance the birds
might break their necks in their terrified flights and violent
dashes against the wire netting, and in this manner one or two
were scalped to the bone.

A list of the Versicolor characters found in the F. 1 females is
interesting, for there are certain characters which can only be
classified in the living bird: Leg-colour, eye-skin, bill, moult, habit,
temperament, voice.

F.1 29 “Z” (P. formosus 9 x P. versicolor S).

Parent of Hye-skin. )
1D, 2 (IBC S< Crest. |
Ve.xVe.).: ‘Neck. |
Flank. &
Breast. ‘. S
Interscapulars. aie
Scapulars. iS
Secondaries. Ss
Back.
Size of egg. J
a, 8 Formosan.
Bill. =
Primaries. | [=
Dimension. =
Leg. ees

542 MRS. R. HAIG THOMAS ON

NINO Reon?

Parent of Hye-skin, >
inter se EK. 2. Crest. |
Neck. tee
Breast. | =
Flanks.. eS
Interscapulars. as
Scapulars. | &
Secondaries. po
Back. |
Size of ege. }
Whe cra yee: 8 Formosan.
Bill. Wee
Primaries. | &
Dimension. ( =
Tail. | a

In both F. 1 females the size of egg was transmitted by the
Versicolor male.

Sy 1 Qe Crest.
Neek:.
Flank.
Breast.
Legs.
Bull, sizeand colour.
Interscapulars,
Scapulars.
Secondaries.
Primaries.
Tail.
Back.
Dimension.
Habit of moult.
Temperament.

a

Wy

Versicolor.

ea eS

The fifth female, reserved to breed from, has a pale grey stripe
down the back of the shank of one leg.

The presence of a mosaic of pale grey and dark grey seen in
the legs of F. 1 in this experiment led to the inference that the
two parents had severally pale grey legs and dark grey legs,
which an immediate examination of the Formosan and Versicolor
species confirmed. A curious independent double segregation of
allelomorphs was observed in the autumn of 19J1 in the crests of
the F. 1 males. The centre feathers were dark—colour Versicolor ;
fully developed, rapid early moult—habit Formosan : the feathers
on the outer edge of the crest were pale—colour Formosan ;

EXPERIMENTAL PHEASAN'T-BREEDING. 545

undeveloped, many still in the quill—habit Versicolor: two
Mendelian pairs coupled in each parent yet repulsing one
another in the same area in F. 1. An examination made at the
same time of the Formosan male showed the crest fully developed,
and an inspection of the Versicolor male showed the crest
undeveloped, mostly still in the quill.

F. 1 generation is to me always the most interesting in these
artificial Pheasant crosses. For Mendelian segregation already
shows, and it is sometimes possible to select the strain of parentage
desired to reappear more strongly in the F. 2 generation. Also
F. 1 occasionally produces remarkable mosaics of sex—a sort of
sex-hybridism accompanied by sterility and extraordinary develop-
ments of plumage in the female, phenomena I will not touch upon
now. ‘To illustrate more clearly the points I wish to bring to your
notice, here is a selection of secondaries (3rds from the last primary)
extracted from the wings of the parent species and of all the
birds connected with the experiment (P]. LXTV.): the Formosan
male secondaries have a peculiar Vandyke pattern like a feather
laid on a feather, and the female secondaries of this species are
banded, both are extremely light in colour. The Versicolor male
secondaries have a mottled grey oblique banding on a very dark
grey ground; the female secondaries of this species are rich
iitonray with snide bands of darker brown; also placed in the same
frame are the secondaries of F.1 and F. 2 males and females.
After examining the parents’ secondaries, we perceive that the
Versicolor male has transmitted to his female offspring of the
F. 1 generation the female secondaries of his species, and that
conversely the Formosan female has transmitted to her male
offspring of the F. 1 generation a pattern resembling the male
secondaries of her species on one vane, though not on the other
vane of the feather. In the F.2 generation (‘Fo x Ve. x Ve.”
Pl. LXV.) the influence of the Versicolor male on his female
offspring continues in pattern and size and more or less in colour,
for in F. 2 females the secondaries seem to be Versicolor, with
slight differences of colour not unlikely to be found between
individuals of the same species; whilst the F. 2 male secondaries,
though most resembling Versicolor, are somewhat hybrid in size
and pattern, and still show slight traces of Formosan influence.

A selection of interscapulars extracted from the parents and
from the two generations of the Formosan Versicolor cross is
shown ; the females are in one frame, the males in another, and in
these the same phenomena appear. Those of the F. 1 females
seem to be Versicolor in pattern and colour, as also do the five
F. 2 females, with certain modifications that might be readily
found amongst individuals of a pure race (Pl. LXVI.). The
frame containing the male interscapulars shows amongst the F. 1
males a Formosan and also a hybrid pattern, whilst the F. 2 males
are also hybrid (Pl. LX VIT.).

The phenomenon of pattern-transference has occurred in all
my Pheasant crosses ; sometimes from the male to the female, or

544 MRS. R. HAIG THOMAS ON

conversely from the female to the male, or a pattern may be
transferred from one area in the parent species to another area
in the F. 2 offspring. J have noticed that these pattern-trans-
ferences are inclined to remain fixed and constant.

Colour-transference also takes place, and sometimes where it has
occurred seemed to inhibit the appearance of pattern.

One instance was noticed where the breasts of the males of the
two races crossed, Th. amhersti and Th. picta, differing widely in
colour, both colours were found on the breast of F. 2 Th. obscura,
the red of picta overlying the metallic green of amhersti.

These appear to be the results of the second experiment :—

(1) The male parent transmitted to his F. 1 female offspring
much of the female plumage of his species and the
dimension of the egg.

(2) The female parent transmitted to her F. 1 male offspring
much of the male plumage of her species.

(3) In the F. 2 generation, the offspring of F. 1 female x
Versicolor male, the Versicolor male seems to have trans-
mitted every character—bill, leg-colour, plumage, habit, and
temperament—of the female of his species to his F. 2
female offspring, whilst he has not transmitted every
character of the male of his species to his F. 2 male
offspring; repeating exactly the results of the original
experiment with genneus.

Are we, then, to suppose that some of the gametes of this
Versicolor male contained all the factors representing the tem-
perament and habit, the colour, pattern, and dimension of plumage,
leg, and bill, and the bulk of the female of his species and
even the factor for size of egg, with the one exception of the
factor for the sex to which these belonged ¢

To the practical experimentalist, to the non-mathematical
simple observer, the hypothesis is difficult to conceive.

These phenomena seem to be of the nature of a sex-limitation
opposed to expectation.

T have read with much interest Mr. Doncaster’s account of game-
togenesis in the Gall-fly, also his researches on sex-limitations
published in ‘Genetics,’ and am interested to know how he
would consider the above facts in relation to his theory of sex:
Male gametes § O female gametes ¢ 9 with selective fertilisation
between the male gamete O (a non-determinant of sex) and the
female gamete ¢.

T hope the material collected in these two experiments may be
thought of sutticient importance for the higher students of
Genetics to give it some attention, when probably the apparently
complicated problem will receive a simple explanation.

EXPERIMENTAL PHEASANT-BREEDING. 545d

IXPLANATION OF THE PLATES.
Prarr LXIV.
3S & & Secondaries (3rd from primary) of the two Parent species
and of F. 1 (Ko. x Ve.).
1. P. versicolor 9. Horizontally set, broad bands, bifurcated ends; ground-
colour brown.
2. P. versicolor . Mottled light bands V-shaped, apex towards rachis; ground-
colour dark grey.
3. P. formosus 2. Obliquely set narrow bands, pointed ends; ground-colour
nearly white.
4. P. formosus $. Vandyke pattern, light, mottled; ground-colour light grey.
5. F.12 “Z” (P. form.? XP. vers.g). Colour, banding, length of feather,
(Parent of F. 2, Ko.x Ve.X Ve.). versicolor; a similar mottling to that
here seen is found on the 3rd Second-
aries of a pure versicolor 2 unrelated
now living in my pheasantry.
G. F.1g“C” (P. form. 9 XP. vers. 3). Left vane Vandyke, Formosan ¢ pattern.
Right vane hybrid banding.
Colour hybrid.

Prats LXV.
6 & 2 Secondaries (8rd from primary) of F. 2 (Fo. x Ve. X Ve.).
1. F.2 ¢ “G” (F.1 Fo.XVe. 2 XP. vers. ¢). Pattern of banding nearest to
versicolor @.
Colour hybrid.

2. 3. 4. 5. 6. F.2 9 9 (F.1 Fo.xXVe. 2 XP. vers. f).
LOK Oise | ae | eH ” OG] ne | Oo Be |

Although slight differences exist between these five secondaries, yet the length

and colour of them, the horizontal setting of the bands, their breadth and bifurcated
ends, are all pure female versicolor characters.

Prats LXVI.
© posterior interscapulars of the two Parent species, of F. 1 and F. 2.
1. P. versicolor ¢.
2. P. formosus 9 (parent of F. 1).
3. F.19“Y” (parent'of inter se F.2). Pattern transference. Pattern found on
several anterior interscapulars of P. ver-
sicolor 2.
4. F.1 2 “Z” (parent of F. 2, Fo.X Ve. x Ve.).
5. F.2 2 “EK” (moult completed). versicolor posterior interscapular pattern.
6. F.2 9 “” (moult completed). Pattern transference. The pattern seen on
7%. F.2 % “H” (moult completed). these two feathers is found on some anterior
interscapulars of 2 versicolor.
8. F.2 2 “C” (aged 4 months, moult incomplete). Pattern found amongst
posterior interscapulars on Y versicolor.
9. F.2 9“B” (moult completed). Pattern found amongst anterior and posterior
interscapulars on 9 versicolor.
Note, July 2nd, 1912.—A number of posterior and anterior interscapulars extracted
from a living female versicolor in my pheasantry showed all the patterns figured
on Plate LX VI.

Prats LXVII.

6 posterior interscapulars of the two Parent species, of F. 1 and F. 2.

si i PR wersicolor 6. | . zs k P. formosus 6.

046 SIR GEORGE H. KENRICK ON

-F.1 og “A” (¥o.XVe.). Similar patterns found on posterior interscapulars of
3 versicolor.

5

6. F. 1 g “B” (Ko.XVe.). Coarser lines but same pattern as found in some
posterior interseapulars on ¢ versicolor.

7. F.1 6 “C” (Fo.XVe.). Similar pattern found amongst posterior inter-
scapulars on 3g versicolor.

8. ¥.2 g “A” (Fo.X Ve.X Ve.).

9. F.2 g “G” (Ko.XVe.X Ve). Pattern found amongst interseapulars on

& versicolor.

Note, 2nd July, 1912.—A number of posterior and anterior interseapulars were

extracted from a male versicolor now living in my pheasantry ; amongst these were

found all the various patterns pictured on Plate LXVII. There is a ditterence

between central and lateral interscapulars, the centrals have the pattern both sides,

5

the left laterals have the pattern on the left side, the right laterals have the pattern
on the right side.

27. A List of Moths of the Family Pyralidse collected by
Felix B. Pratt and Charles B. Pratt in Dutch New
Guinea in 1909-10 ; with Descriptions of new Species.
By Sir Georg H. Keynrics, F.Z.8 :

[ Received January 30, 1912: Read March 19, 1912. |

(Plate LX VIII.*)

INDEX.

Systematic :— Page
DWGIGCUUG), CHARHIALDISOS,, S\Os Ws concseossececo-cascoveascnos-ccon DEL7/
Merolicalistspans Vs. 3a3. ck ee Pee ee 547
RL, MAOSWATTASD x Men cee eee ee OLE
SWAGRIGED TOES BOs Wass sancseosscee ban enesoree-osancaves Hts}
Perisyntrocha suffusa, Sp. VW. ...... eee 54S
Ambia novagwinensts, SP. Ve... ec cee eee eee cece eee. BAD
Margarosticha plumbealis, sp. nn. .........--...04..-.. 549
Brevicella emarginata, gen. et sp. un. ................. 549
Sceliodes grisealis, Sp. . ...-......2...00ee et 000
Hintephria mioswari, 8p. i. ... 220-0000. eee eee 50
Df GPPUSCOUIS, SDs Wo Gasesoccoossgo. ban vanbnssesoo0ss s5saccen OO)
JEOIGCUD, jf CHOSGy BDo W5 Saooosscases<osanvccescnesscooesase BON
Sylepta dinawa, Sp. Ws... .2.-..-2e0s eee ese ON
Glyphodes magnificalis, Sp. M. ... 2.6... ceeeeeve eee ees 802
G. nigricincta, sp. N. 552
G. pseudocesalis, sp. 0. 552
Omphisa variegata, sp. n. 593
Noorda arfakensis, sp. 0. 503
Pilocrocis anguliferd, Sp> W. 0-225... seeeee eee ee ese 5d
Pyrausta flammealis, sp. n. 554

Variation :—

Gilypliodeswpner iene rare ee ee eee eee 5a4
Calamachrous albipunctalis 554

After some considerable experience in British New Guinea,
the two sons of Mr. A. KE. Pratt made several attempts to explore
various portions of the Dutch territory, and although disappointed
in some directions they made a very successful ascent of the Arfak
Mountains, formerly visited by d’Albertis, and spent a considerable

* Vor explanation of the Plate see p. 555.

PaaS, USy2 ledh die OVAUN.

West, Newman chr.

Horace Knight del.et lith.

PYRALIDA FROM DUTCH NEW GUINEA.

MOTHS FROM DUTCH NEW GUINEA. HAT

time in the vicinity of two lakes, which proved to be excellent
collecting-grounds for the lepidopterist.

Although for the most part the insects were the same as those
collected in British New Guinea in the south, a fair number of
different species occurred, a list of which I give herewith, together
with descriptions of new species.

The island of Mioswar, on the north-east coast, was also visited ;
but the nights were unfavourable for collecting and few moths
were taken, although it proved te be an excellent locality for
Lycenide.

The specimens were nearly all taken at light, as in the case of
those enumerated in my former list.

Sub-fam. HPrpascHitn as.
MACALLA SYRICHTUSALIS Walk.

MACALLA ARRUENSIS, sp.n. (PI. LX VIII. fig. 3.)

Head, legs, antenne, and palpi fawn-colour; abdomen the
same, with darker segments at the tuft; tarsi dark ringed with
paler.

Fore wing fawn-colour, with a dark brown basal patch out-
wardly oblique followed by a whitish irregular spot on the costa
and re: iching as far as the bottom of the cell. Following this on
the costa is a darker mark developing into a faint central shade.
Beyond this is a curved postmedial line ending in a white mark
on the costa, and at the apex there 1s a dark patch.

Hind wing pale ochreous ; fringes pale.

Exp. 32 mm.

From Arfak Mts., 4000 ft.; but there is an undescribed spe-
cimen from Arru in the British Museum.

MACALLA OLIVALIS, sp.n. (Pl. LX VIII. fig. 1.)

Head, legs, palpi, and thorax olive and white ; antenne brown ;
abdomen pale grey.

Fore wing olive-green with white markings ; three equidistant
white spots on the costa, an antemedian inte:rupted white band,
a median interrupted white line, and a postmedian distinct white
line preceded by an irregular white blotch which does not reach
the costa or the inner margin; a subterminal white line; fringe
orange and white; underside of fore wing with a number of large
silvery scales in the cell and at the base of the wing.

Hind wing grey, semi-hyaline ; fringes paler.

Exp. 29 mm.

Arfak Mts., Dutch New Guinea.

MACALLA MIOSWARI, sp.n. (PI. LX VIII. fig. 5.)

Head, thorax, antennee, and palpi reddish ochreous ; legs darker,
ringed with ochreous.
Fore wing pale ochreous with chestnut patches, one at the base

548 SIR GEORGE H. KENRICK ON

terminating in a dark antemedian line, one at the apex, and another
at the angle. Between these two is an extension of the median
pale ochreous area extending to the hind margin; an angulated
dark postmedian line; a small dark central spot and a dark mark
on the inner margin between the lines.

Hind wing grey. Fringes of both wings bright chestnut.

Exp. 27 mm.

Island of Mioswar, Dutch New Guinea.

STERICTA RUREALIS, sp.n. (PI. LX VIII. fig. 2.)

Head, antenne, and palpi pale ochreous; patagia chestnut ;
abdomen pale at the base, but with chestnut tuft; legs dark,
paler at the joints.

Fore wing ochreous, with a yellower shade along the costa and
the raised scales red. A basal patch reddish chestnut, bounded
obliquely and extending along the inner margin nearly halfway.
An apical chestnut patch and a smaller patch at the angle.

Hind wing pale ochreous, with a fine marginal line darker.
Fringes ochreous, but damiger at the apex.

Exp. 44 mm.

Artfak Mts., 4000 ft.

Sub-fam. PyRALINE.
CURENA EXTERNALIS Walk.

Sub-fam. HyDROCAMPINA,
AULACODES BRUNNEALIS Hmpsn.

AULACODES METALOXALIS Hmpsn.
NYMPHULA CHRYSEIS Meyr.
NYMPHULA POLYSTICTALIS Hmpsn.
CLUPEOSOMA POLUSALIS Walk.
PILETOCERA @GIMISALIS Walk.
PILETOCERA DIPLATYLA Walk.
PILETOCERA TORSICUTALIS Hmpsn.
NEOGENESIS FLAVOPLAGIALIS Hmpsn.

PERISYNTROCHA SUFFUSA, sp.n. (PI. LXVITI. fig. 4.)

Head, antenne, palpi, and legs pale ochreous ; thorax darker,
with paler patagia; abdomen grey, with pale band at base and
paler rings at the segments.

Fore wing hyaline, pale golden; a darker margin to the costa.
A patch of dark scales at end of cell, developing into a triangular
fuscous mark reaching to the inner margin; the apical area of
the wing is occupied iy a fuscous patch, with the exception of a
pale patch extending from the costa to the central nervure.

MOTHS FROM DUTCH NEW GUINEA. ye)

- Hind wing pale with fuscous margin, broad at apex but dis-
appearing at the angle; fringes pale.
Exp. 24 mm.

Arfak Mts., 4000 ft.
AMBIA NOVAGUINENSIS, sp.n. (PI. LX VIII. fig. 6.)

Head, legs, antenne, palpi, and thorax brown.

Fore wing whitish, with numerous brown markings ; two ante-
median lines containing a white spot near the costa: a dark spot
at end of cell, with a white triangular centre; an oblique line
running from near apex to middle of inner margin, followed by
a darker space containing five triangular white spots with dark
outer margins.

Hind wing white, with an irregular dark median line and a
dark subterminal line.

Exp. 18 mm.

This insect has veins 4 and 5 stalked in both wings.

Warmasin Lake, Arfak Mountains, 6000 ft.

MARGAROSTICHA PLUMBEALIS, sp.n. (Pl. LX VIII. fig. 9.)

Head, legs, antenne, and thorax pale chestnut ; patagia white ;
abdomen bright chestnut, with black tuft.

Fore wing bright chestnut, with silvery-white blotches: the
first is square and extends from the median nervure to the hind
margin, the second is triangular and near the apex; before the
apex itself is a narrow white streak.

Hind wing chestnut, with a broad white transverse band ;
beyond this is a lead-coloured round patch reaching the outer
margin: it is in reality black, with a number of scattered white
scales; on the outer edge of this are three black dots, each having
a centre of chestnut, and between them silvery-white dots.

Underside silvery, with the markings of the upper wings
showing through ; fringes darker.

Exp. 22 mm.

Arfak Mts., 4000 ft.

New genus under sub-fam. Hydrocampine :—

BREVICELLA (type emarginata, © ).

Palpi porrect, twice the length of head, not scaled: antenne
simple; legs not hairy; tibize with usual spies. Hind wings
with long hairs on parts of the costa.

Venation—Fore wing: 2 and 3 stalked; 4, 5, 6, 7 from cell ;
9 and 10 stalked, with 8-11 present. Hind wing: 2 and 3
stalked slightly ; 4, 5, 6 from cell; 7 stalked with 8.

BREVICELLA EMARGINATA, Side Tg (LEG WWIII ais, tek)

Body and wings dark brown ; a postmedian curved line bordered

within by seven triangular whitish dashes; at the upper end of

550 STR GEORGE H. KENRICK ON

this line is an oblique intensely black dash reaching nearly to the
costa.

Hind wing with an antemedian whitish spot and a postmedian
darker band. There is a narrow darker marginal line to both
wings, and the fringes, which are dark, are strongly emarginate.

Exp. 30 mm.

Momi-Arfak, 4000 ft.

ScELIODES GRISEALIS, sp. n. (Pl. LX VIII. fig. 7.)

Head, antenne, legs, and palpi ochreous ; thorax and abdomen
pale brown ; tuft pale.

Fore wing ochreous, powdered with black scales and with three
white angulated streaks, the first much angulated before the middle,
the second beginning in the middle of the costa and narrowing to
a point on the lower edge of cell, the third also from the costa
extending as an angulated fine line to the middle of the wing; a
subterminal ill-defined whitish band, in which is a dark sub-
terminal line followed by a dark cloud.

Hind wing whitish, with two dark dots on the disc, a dark
subterminal line followed by a dark cloud ; fringes brown.

Exp. 30 mm.

Arfak Mts., 4000 ft.

Sub-fam. PyrausTina.
XANTHOMELMHNA SCHEMATIAS Meyr.

ENTEPHRIA GLAUCIAS Meyr.

ENrEPHRIA MIoswaRI, sp. n. (Pl. LXVIII. fig. 10.)

Head, antenne, legs, and palpi silvery white. Thorax and
abdomen white, the latter with black markings on three or four
of the last segments. Fore wing white, a small soot-black spot
near the base of the cell and a larger round one at the end.
Before the apex is a costal black spot, from which springs a pale
transverse ochreous line reaching the inner margin. In some
specimens there is a dark spot at the apex, and in all the hind
margin is dark.

Hind wing white, with a small dark spot near the base and a
thin median line; hind margin and fringes darker,

Exp. 24 mm.

Island of Mioswar, Dutch New Guinea.

ENTEPHRIA GRISEALIS, sp. n. (PI. LX VIII. fig. 12.)

Head, palpi, antenne, and legs silvery white; thorax and
abdomen white, the last three segments darker above.

Fore wing white shaded with grey; two transverse ochreous
antemedian lines, the first somewhat obscure; a median line from
inner margin to vein 2, where it disappears in a broad shade; a
transverse distinct line from costa to vein 2 beyond the cell,

MOTHS FROM DUTCH NEW GUINEA. ol

Hind wing white; a transverse dark line from angle to costa
and another from costa beyond cell to vein 2.

The margins of both wings are bordered by a dark, thin, double
line enclosing a paler portion, Fringes white.

Exp. 32 mm.

Arfak Mountains, Dutch New Guinea.

REHIMENA CIssopHORA Turner.
ZAINCKENIA LOPHOCERALIS Hmpsn.
AGROTERA PYROSTICTA Hmpsn.
PaGyba BOTYDALIS Snell.

PaGYDA FuMOSA, sp.n. (Pl. LX VIII. fig. 13.)

Head, legs, antenne, and palpi straw-colour ; thorax and abdo-
men rather more orange.

Fore wing straw-colour, a short, orange, transverse line near
base; an antemedian and a postmedian line, the middle line
curved, dark orange; an oblique curved band of pale orange from
apex to hind margin and a paler marginal band.

Hind wing suffused with a smoky patch in which there are
three whitish transverse lines and a dark fine marginal line ;
fringes paler.

Exp. 22 mm.

Arfak Mts., 4000 feet.

CAPRINIA DIAPHANALIS Walk.
PHRYGANODES EREBUSALIS Hmpsn.
PHRYGANODES ANALIS Hmpsn.
PHRYGANODES USITALIS Guen.
PHRYGANODES TETRAPLAGALIS Hmpsn.
PHRYGANODES CENTRABALIS Hmpsn.
NACOLEIA PERDENTALIS Hmpsn.
Proropes MrIMIcA Swinh.

SyLepra POLYDONTA Hmpsn.
SYLEPTA LEUCODONTA Hmpsn.

SYLEPTA DINAWA, sp.n. (PI. LX VIII. fig. 11.)

Head, thorax, legs, antenne, and palpi dark brown ; abdomen
dark from with a conspicuous pale ochreous tuft and a pair of
black tufts of hair at base. Fore wing pale ochreous clouded with
dark grey ; an irregular dark antemedian line and a vague post-
median line with a branch towards the angle; the apex is broadly

552, SIR GEORGE H. KENRICK ON

margined with dark; below the costa at end of cell is a lunated
mark and there is another smaller one nearer the base.

Hind wing pale semihyaline, with traces of dark ante- and post-
median lines, the hind margin with a broad dark shade; fringes
pale.

Exp. 42 mm.

Dinawa, British New Guinea.
GLYPHODES POLYZONALIS Hmpsn.

GLYPHODES MAGNIFICALIS, sp.n. (Pl. LX VIII. fig. 17.)

Head, palpi, and antenne black; legs and underside golden.
Thorax black, with collar and patagia ochreous ; abdomen black
above.

Fore wing pale ochreous; an oblique black band beginning as a
fine line on the base of the costa reaches a point two-thirds along
the inner margin, from this there stretches a nearly vertical
black band with curved outer side to the costa. There is a sub-
terminal cloud of brown intersected by two fine metallic purple
lines, on the inner side of this are two triangular black marks
united in the middle. All the black marks are ‘shot with metallic
blue.

Hind wing pale ochreous, with an mterrupted dark postmedian
line. On the middle of the edge of the wing are three white-
centred black dots on a faint bright line.

Exp. 40 mm.

Arfak Mts., 4000 feet.

The only specimen is a female.

GLYPHODES NIGRICINCTA, sp. n. (Pl. LXVIII. fig. 16.)

Head, palpi, and antenne dark brown; legs white, the joints
dark brown on the outer side; front of thorax black; patagia
white ; abdomen grey, with a black band at base.

Fore wing white, a dark grey spot in the middle of the cell and
a second at “dhe end; the outer half of the wing very dark grey.

Hind wing white, with grey lunule and dark grey apical patch.
Fringes smoky, fading into white towards the angle of the hind
wing.

Exp. 26 mm.

Arfak Mts., 4000 feet.

GLYPHODES PSEUDOCAISALIS, sp. n. (PI. LX VIII. fig. 18.)

Differs from cesalis Wk. in the absence of hyaline patches, in
having the fringes unspotted whereas in céesalis the dark spots
give the wings a distinctly emarginate appearance, and in the
markings of the fore wing : these consist of a basal line indicated
by dots, antemedian, median, and postmedian lines, each of which
is double and the {nner lines of the median and postmedian are
connected. In cesalis these double lines are wider apart, more
irregular, and filled in with darker instead of with the ground-
colour as in pseudocesalis. Beyond the postmedian line in ceesalis

MOTHS FROM DUTCH NEW GUINEA. 553

is a dark patch with four pale lunules, which are absent in pseudo-
cesalis.

Exp. 30 mm.

Arfak Mts., Island of Mioswar, Mt. Goliath, and British New
Guinea.

It is possible that there is another allied species, but it may be
simply a larger form of cesalis.

PYGOSPILA TYRES Cram.
CROCIDOLOMIA BINOTALIS Zell.
SAMEODES POLYTHLIPTALIS Hmpsn.

OMPHISA REPETITALIS Moore.

OMPHISA VARIEGATA, sp.n. (PI. LX VIII. fig. 19.)

Head, legs, antennx, and palpi chestnut-brown ; thorax chest-
nut-brown, the patagia pale; abdomen darker, spotted with paler
on back.

Fore wing pale ochreous with brown markings.

Fore wing: basal patch brown, an antemedian and a postmedian
line enclosing a darker band which does not reach the costa, but
which surrounds a conspicuous subtriangular pale spot near the
end of the cell; beyond this is an angulated line followed by a
darker shade; costa with six lunules.

Hind wing pale, with an irregular sharply-defined median band
and a double outer dark line.

Exp. 32 mm.

Island of Mioswar, Dutch New Guinea.

NoorDA NIGRIPUNCTALIS Hmpsn.

NOoRDA ARFAKENSIS, sp.n. (Pl. LX VIII. fig. 14.)

Antenne and palpi brown; legs white, with brown tips to the
joints; head ochreous ; thorax white ; abdomen whitish, the last
segment grey, anal tuft white.

Fore wing creamy white, the costa reddish ; a dark brown. dot
at base, another at the beginning of the cell, and a third at the
end; a dark brown annulated subterminal line; the remainder
of the wing to the outer margin of a chestnut colour mottled
with darker.

Hind wing white. Fringes pale brown, shading into white at
the angle of the hind wing.

Exp. 30 mm.

Arfak Mts., 4000 feet.

DAUSARA AMETHYSTINA Butler.

Pitocrocis (a subdivision of Crocidophora) ANGULIFERA, sp. Nn.
(Pl. LXVIIT. fig. 15.)

Head, legs, antennz, and palpi straw-colour ; thorax and abdo-
men pale golden, the last few segments becoming fuscous.

Proc. Zoou. Soc.—1912, No. XX XVII. 37

554 ON MOTHS FROM DUTCH NEW GUINEA.

Fore wing pale golden, a very short basal dark line, an oblique
antemedian line, a sharply angulated median Jine, a regular
curved postmedian line (all these are dark fuscous and the last is
lost in a fuscous patch which extends from the middle of the hind
margin halfway up the hind margin).

Hind wing pale at base, with the whole of the outer portion
dark fuscous, the boundary between these is marked by a very
irregular angulated line. Fringes rather paler.

Exp. 28 mm.

Arfak Mts., 4000 feet.

APHYTOCEROS GROSALIS Meyr.
CURICTA OPPOSITALIS Walk.
PYRAUSTA OCCULTILINEA Walk.
PYRAUSTA ALENTIALIS Snell.
PYRAUSTA DEDUCTALIS Walk.

PYRAUSTA FLAMMEALIS, sp.n. (Pl. LX VIII. fig. 20.)

Male with a tuft of hair at base of fore wing below and a strong
fold of costa extending to end of cell; female with a smaller tuft
of hair; both sexes have the base of the antenne surmounted by
a tuft of hair, but this is larger in the male. Head, legs, and an-
tenne pinkish brown, also palpi; abdomen ochreous above, darker
below; thorax and patagia flame-colour.

Fore wing flame-colour, with three transverse fine dark lines: the
first ante median and oblique, the second median and angulated in
the middle, the third angulated and not reaching below the third
nervure; all the margins of the wing are darker.

Hind wing uniformly straw-colour with fringes slightly darker ;
fringes of fore wing dark. In the female the flame-colour is
replaced by Indian yellow.

Exp. 30 mm.

Arfak Mts., 6000 feet.

In addition to these species I have figured two varieties of
species already described :—

Pl. LX VIII. fig. 21 represents a form of Glyphodes pfeifferce in
which the marginal border to both wings is much deeper than any
specimen I have received from New Guinea.

Pl. LX VIII. fig. 22 represents a form of Calamachrous albi-
punctalis in which the deep maroon-purple of the wings of this
species is replaced by bright chestnut.

Both of these forms are from the island of Mioswar.

Although in some cases a good series of the species was obtained,
in others only single specimens or two examples were taken, so
that the differences in sex could not be ascertained.

From this circumstance I am inclined to think that many other

THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 555

species would turn up if the localities could be worked throughout
the year, and I believe any collector willing to face the difficulties
of a camp life would be amply rewarded by the results.

I desire to tender my sincere thanks to Sir George Hampson
for advice and assistance in regard to species of this large group
and for permission to study the excellent collection at South
Kensington.

EXPLANATION OF PLATE LXVIII.

Fig. 1. Macalla olivalis. | Fig. 13. Pagyda fumosa.

2. Stericta rurealis. 14. Noorda arfakensis.

3. Macalla arruensis. 15. Pilocrocis angulifera.

4. Perisyntrocha suffusa. 16. Glyphodes nigricincta.

5. Macalla mioswari. 17. Glyphodes magnificalis.

6. Ambia novaguinensis. 18. Glyphodes pseudocesalis.

7. Sceliodes grisealis. 19. Omphisa variegata.

8. Brevicella emarginata. 20. Pyrausta flammealis.

9. Margarosticha plumbealis. 21. Glyphodes pfeiffere (var.).
10. Hntephria mioswari. 22. Calamachrous albipunctalis
11. Sylepta dinawa. (var.).

12. Entephria grisealis.

EXHIBITIONS AND NOTICES.
April 2, 1912.

Dr. A. Smrruh Woopwarp, F.R.S., Vice-President,
in the Chair.

The Secretary read the following report on the additions that
had been made to the Society’s Menagerie during the months of
February and March, 1912.

FEBRUARY.

The registered additions to the Society’s Menagerie during the
month of February were 128 in number. Of these, 50 were
acquired by presentation, 29 by purchase, 13 were received on
deposit, 19 in exchange, and 17 were born in the Gardens.

The number of departures during the same period, by deaths
and removals, was 236.

Amongst the additions special attention may be directed to :—

1 Snow-Leopard (Felis wncia) and 1 Musk-Deer (Moschus
moschiferus), from Nepal, presented by H.M. Tur Kine on
February 29th.

2 Snow-Leopards (Melis uncia), from Kashmir, presented by
Capt. G. Douglas Oliver, F.Z.S., on February 26th.

2 Jaguars (Felis onca), born in the Menagerie on February 20th.

2 Striped Hyznas (Hyena hyena), from N. Nigeria, pre-
sented by H. P. Lobb, Esq., on February 4th.

1 Spurrell’s Dormouse (Graphiurus spurrelli), from Dunkwa,

37*

556 DR. R. W. SHUFELDT ON VIRGINIA OPOSSUMS.

Gold Coast, new to the Collection, presented by Dr. H. G.
F. Spurrell, F.Z.8., on February 12th.

1 White-capped Redstart (Chimarrhornis lewcocephalus), from
the Himalayas, presented by Alfred Hzra, Hsq., F.Z.S., on
February 29th.

Marcu.

The registered additions to the Society’s Menagerie during
the month of March were 110 in number. Of these 45 were
acquired by presentation, 32 by purchase, 21 were received on
deposit, 8 in exchange, and 4 were born in the Gardens.

The number of departures during the same period, by death
and removals, was 192.

Amongst the additions special attention may be directed to :—

1 Wallich’s Deer (Cervus wallichii) 3, new to the Collection,
1 Burrhel (Ovis burrhel) 3, 1 Tahr (Hemitragus jemlaicus) ° ,
and 6 Tibetan Mastiffs (Canis familiaris), from Nepal, presented
by H.M. the King on March 13th and 25th.

1 Alpaca (Lama pacos) 3, from Peru, received in exchange on
March 5th.

1 Racket-tailed Drongo (Dissemurus paradiseus), from India,
presented by Alfred Ezra, Esq., F.Z.S., on March 14th.

1 African Tantalus (Pseudotantalus ibis), from. Southern
Nigeria, presented by Sir Walter Egerton, K.C.M.G., F.Z.S., on
March 15th.

1 Ashy-headed Goose (Chloéphaga poliocephala) bred in
Holland, received in exchange on March 22nd.

3 Prairie-hens (7ympanuchus americanus), from North America,
received in exchange on March 21st.

Mr. R. I. Pocock, F.RBR.S., F.LS., F.Z.S., Superintendent of
the Gardens, exhibited a lantern-slide of two Polar Bear cubs
(Ursus maritimus) born in the Gardens in November 1911, and
made some remarks upon the causes of the difficulty experienced
in Zoological Gardens in rearing the offspring of this species.

Mr. C. Tate Reean, M.A., F.Z.S., 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 ground on which it lies.

Dr. R. W. SHuretpt, C.M.Z.S., 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

Pm Z S, IN!, Pil, LxXIDK.

Photos by D. Seth-Smith. Bale & Danielsson Lt? imp.

YOUING (GOANRIVAIMUN (CIRUUS TWAIN,

MR. D. SETH-SMITH ON A YOUNG CARIAMA. 557

elementary 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, etc. ‘The 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 vertebrz in the spine of
the Virginia Opossum was 7 cervical, 13 thoracic, 6 lumbar, and
26 caudal. This was probably correct for the adult animal of
this species, while in subadults, 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.

In both these specimens the dentition in use was as follows :—

- 6 1 5
07 OF pm +m ,= 42,

the third of the cheek-teeth being the molar-like predecessor of
the one premolar which changes in Marsupials. Two further
molars would have come up in later life, making the adult
formula :—

5 1
apy ae
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.

E 3 4
a pms, m4=90.

Mr. D. Sera-Smira, F.Z.8., 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 (Pl. LXIX. *) 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.

The nest was formed of sticks and small twigs on a platform of
branches that had been specially erected at about eight feet from
the ground in the Hastern Aviary. Of the two eggs laid, one
was accidentally broken by the birds, but the other hatched on
June 6th, incubation, performed chiefly by the female, having
occupied twenty-nine or thirty days. The young bird was
covered with down of a pale brown colour mottled with darker
brown on the back, that on the head being extraordinarily long
and hairlike. It was fed by the parents with small pieces of meat,

* For explanation of the Plate see p. 558.

558 MR. R. I. POCOCK ON A

mice, hard-boiled egg, and cockroaches. It left the nest at a month
old, and at five weeks old may be described as follows :—

Upper parts pale brown, darker and with a rufous shade on the
mantle, the rump and upper tail-coverts being dark brown. The
crown of the head and neck barred with dark brown; the wings
also covered with fine wavy lines of dark brown on a paler ground.

The abdomen and under tail-coverts whitish.

The bill very dark brown, the legs and feet dark grey, and the
iris very pale silvery grey.

EXPLANATION OF PLATE LXIX.

Hig. 1. Young Cariama cristata 24 days old, in nest.

Fig. 2. 5. 53 Pe 30 days old, with parent.
Figs. 3 &4.,, an Bs 2 months old.
PAPERS.

30. On a rare Stag (Cervus wallichii) from Nepal recently
presented to the Zoological Society by His Majesty
[SinggGeorve eb yee ococerstaly: Seush aliases
F.Z.8., Superintendent of the Gardens.

[Received March 19, 1912: Read April 2, 1912. ]
(Text-figures 66-71.)

INDEX.
Systematic : Page
Cervus wallichii Cuv., Western Tibet, text-figs.
CE =69 RES... hah Ee ee Ce eR BODO
» hanglu Wagn., syn. cashmeriensis Adams,
cashmeerianus Falc., Kashmir, text-fig.70 566
» 4affinis Hodgs., Choombi Valley, Lhasa, etc. 567

» macneilli Lydd., Szechuen, text-fig.71 ... 570
» kanswensis, nov., Kansu ........................ 642
Characters of these species tabulated .................. 574
Cervus albirostris, Tibet ...............cc00eceeeeeeeeess BTA

Krom the scientific standpoint the most interesting animal
in the collection from Nepal presented by H.M. the King to the
Zoological Society was a large Stag belonging to a species
(Cervus wallichii) which has never been previously exported
alive from India, and, so far as available records show, has
never been shot or preserved by any English sportsman, and is
therefore unrepresented in any of the large museums of the
world. The arrival of this specimen, moreover, has made it
possible to classify a species which for nearly a century has
been a puzzle to all systematic workers on the Deer.

RARE STAG FROM NEPAL. 559

Cervus wallichii was described by G. Cuvier (Rech. Oss. Foss.
ed. 3, iv. p. 504, 1825*; ed. 4, vi. pp. 88-89, 1835) from a
native sketch, sent to him by Duvaucel, of an animal, one of a
pair, according to Blyth, living in the menagerie at Barrackpore
and said to have come from Muktinath, north of Dwalagiri in
Nepalt. This sketch was reproduced by F. Cuvier & Geoffroy
St. Hilaire (Hist. Nat. Mamm. iv. no, 356, 1823), and modifications
of it were reproduced subsequently by Jardine £ and Hamilton
Smith §.

Text-fig. 66.

Cervus wallichii.

Photograph of Cuvier’s figure of the type-specimen.

The main characters of the animal are made clear by the
descriptions and figures of the French authors. It was described
as dark grey-brown or yellowish grey-brown with pale legs, a

* JT have not seen this edition.—R. I. P.

+ In support of this Blanford (Mamm. Brit. India, p. 538), copying Blyth, cites
Hardwicke (Tr. Linn. Soc., Zool. xiv. p. 581, 1823). Hardwicke, however, does not
there state that the stag from Muktinath he identified as ‘ C. pygargus, Pall.’ was
the type from which the sketch was taken. The coincidence of the dates, however
leaves little doubt on this point.

{ Naturalistst’s Library, iii. pl. 10, p. 161, 1835.

§ Griffith’s Animal Kingdom, iv. p. 103, pl., 1827.

560 MR. R. I. POCOCK ON A

very short white tail and a large white disk upon the crowp*.
The figure shows no less clearly that the caudal disk in size and
extension over the croup, reaching nearly to its summit, resembled
that of a Wapiti but was white instead of yellowish buff in
colour. ‘The antlers were short, only a little exceeding the head
in length, and carried a brow and a bez tine, the beam being
simply forked and the trez tine absent. The brow and bez tines
were close together at the base and diverged at about an angle of
45°, the brow tine projecting straight forwards and the bez in-
clining obliquely upwards. The beam, which receded at an obtuse
angle of about 135° from the bez, appears from the figure to
have formed a fairly regular arch, there being no indication that
either of the terminal tines was turned inwards.

For reasons given below I am quite of Hamilton Smith’s
opinion (Griffith’s An. King. iv. p. 103, 1827) that these antlers
show the stag to have been an old animal at the time the
illustration was made.

A pair of the antlers of this species was figured and described
by Blyth (J. A. 8. Bengal, x. fig. 7 of pl. facing p. 750, 1841)
and subsequently stated by this author in the same journal
(vol. xxx. p. 188, 1861) to have belonged to the animal of which
a sketch was sent to Cuvier. They were judged by Blyth
to have been produced by a stag in its third year. It will be
noticed, however, that neither antler of the pair has a trez tine ;
and since in the Red Deer this tine normally, at all events,
appears in the antlers of the third year, one would expect it to
be present in a specimen of C. wallichit of that age. It will
also be seen that in the left antler figured by Blyth, the beam is
straight and terminates in a single spike, while in the right
antler the beam is forked and much more curved. The want
of symmetry between the two in these and other particulars,
coupled with the absence of the tines mentioned, forcibly
suggests to me degeneration with age. It is, at all events, quite
clear that these antlers were not those carried by the stag at the
time the figure sent to Cuvier by Duvaucel was drawn. They
were probably a later growth and still more decadent. Even the
right antler was less curved than were those of the type, as figured ;
and although the basal juxtaposition and mutual divergence of
the brow and bez tines were much the same in the two cases, the
angle of their divergence from the beam was in both instances
much smaller. But, although these differences and resemblances
between the two sets of antlers are interesting, I do not think
they afford much clue to the affinity of C. wallichii, since these
antlers were, in my opinion, obviously degenerate.

The specimen the Society has just received is a comparatively
large Stag, though not approaching a Wapiti in size. It stands
about 4 ft. 3 ins. (just under 13 hands) at the shoulders. It
has a long face, a small sleepy looking eye, and its longish ears

* The italics are mine.—R. I. P

RARE STAG FROM NEPAL. 561

differ from those of all living Deer that I have seen in having
the upper edge markedly sinuous (emarginate) towards the tip.
The hairs on the neck and throat are longer than elsewhere
and constitute a small mane. The tail is quite short, the legs
strong, the hoofs rather broad and the metatarsal tuft a little
below the hock. The colour (in March) is remarkably pale, being
a tolerably uniform yellowish or sandy-brown all over, except on

Text-fig. 67.

Cervus wallichii.

The King’s specimen shortly after its arrival at the Gardens in March.

the forehead where the matted hair is browner, on the cheeks
and the backs of the ears which are greyer, and on the muzzle
and chin which are pale fawn with an indistinct darker patch at
the corner of the mouth. The backs of the thighs are white,
the pale portion of the rump being only as wide as in the
Hangul or Kashmir Stag, but from the level of the ischia the
white bends sharply forwards and extends nearly as far as

562 MR. R. I. POCOCK ON A

the summit of the croup, forming a very large pure white disk
without trace of median dividing dark line. Upon the croup
this disk is not emphasized by a bordering of darker hairs; but
below the level of the tail, the white on the back of the thighs
is set off by a margin of brown. This disk, so far as size is
concerned, differs from that of the Wapiti in being narrower
below, its width on each side on a level with the tip of the tail
only equalling the width of that organ, whereas in the Wapiti
the disk at this point is equal to twice the width of the tail.

It is also noticeable that the hairs adjoining the croup-disk are
quite white at the base, so that if they were elevated or deprived
of the yellowish-brown pigment of their distal portions, a far
larger white rump patch would be displayed than is normally the
ease. None of the hairs show signs of speckling apically or
elsewhere, and their concealed portion is greyish brown on the
body, though darker on the neck. Both on the body and over
the croup a very large number of them are curled forwards
apically, like those on the croup of the type specimen of C.
wallichit as described by Cuvier, but a mounted specimen of the
Kashmir Stag in the British Museum shows, to a lesser degree,
this same peculiarity, which is no doubt a sign of old hair.

The antlers, which were shed early in March on the voyage
from India, are short but massive. The right one is abnormal,
the left probably normal but, possibly, reduced in size from age
degeneracy. Both are brown with pale tips to the tines. In the
left antler the burr is short, the brow tine rising about an inch
above it. This tine is about 84 inches long, projects nearly
horizontally forwards and is bent slightly down just before the
tip. The bez tine is straight, about half an inch shorter than the
brow and diverges from it at an angle of about 45°, their basal
separation being about 1 inch. Just above the root of the bez
on the inner side there is a short snag. Beyond the bez the
beam curves gently outwards and upwards for about 73 inches to
the origin of the trez, which is about 7 inches long and lightly
curved, and lies in approximately the same plane as the bez and
brow tines. Above the trez the beam ascends, showing upon the
anterior aspect a slight inward inclination, but from the external
profile view it exhibits a decided inclination backwards with a
light upward curve, its axis lying almost at an angle of 45° to
that of the trez, and forming a very obtuse angle with the lower
part of the beam, there being a marked concavity behind the
root of the trez. It terminates above in two tines, one short and
erect and continuing the line of the beam, the other twice as
long and inclined upwards and inwards.

The antler is thus five-pointed. It resembles the antlers of
the typical specimen in the basal juxtaposition and marked
divergence of the brow and bez tines, but, apart from the
presence of the trez tine, differs from them in the more upward
curvature of the basal part of the beam, which is thus inclined at
sharper angles to the axis of the brow and bez tines. In this

RARE STAG FROM NEPAL. 563

latter particular, however, the antlers figured by Blyth are inter-
mediate between those of the example figured by Cuvier and the
one just described.

Text-fig. 68.

Cervus wallichii.

A. Anterior aspect of the shed antlers.
B. External aspect of left antler.

In the right antler the brow and bez tines resemble those of
the left antler; but above the bez the beam bifurcates into

564 MR. R. I. POCOCK ON A

anterior and posterior branches ; the former, occupying almost the
position of the trez in the normal antler, is stout, 15 mches long,
evenly curved upwards and outwards and two-tined, one tine
being short and forwardly directed, the other, which is terminal,
being long and lightly curved upwards, backwards and inwards.
This branch may be described as an attempt at a reduplication
of a normal beam. The posterior branch is thinner than the
anterior and ascends with an even curvature upwards, outwards
and then inwards, to terminate in two tines, an external which
is merely a short bud, and an internal which follows the line
of the beam without showing any marked inward inclination,
although it clearly corresponds to the inwardly inclined terminal
tine of the left antler.

Judging from their small size and asymmetry, these antlers
may be decadent; but, if so, the left antler has suffered less
than the right from that process and less than those of the
type specimen of C. wallichii described above. Still I do not
think it safe to infer that this left antler resembles that of a stag
of this species in full vigour. ‘The possibility, however, must be
borne in mind.

Of the exact geographical area inhabited by C. wallichii we
have at present no trustworthy information. The type of the
species in the Barrackpore menagerie was said by Hardwicke
(Tr. Linn. Soc. xiv. p. 581, 1823) to have been brought from
Muktinath, away to the north of Dwalagiri in Central Nepal.
The example now in the Gardens was presumably in captivity in
Nepal, since it was presented to King George by the Maharajah
of that country (see note below).

The only other Stag, known to me, which seems to belong
to this species is the one which Mr. Lydekker identified from a
photograph as C. affinis (P. Z.S. 1909, p. 599, fig. 182). This,
too, was a captive animal and was alleged to have come from
Sikhim. I judge from the size of the rump patch that this stag
belongs to CO. wallichii. My. Lydekker described the colour
as “very like that of the Hangul with a large white rump-
patch.” This seems to be correct so far as the size of the
rump-patch is concerned; but if the colour of the body was
inferred from the photograph, the inference is untrustworthy
since the camera is notoriously deceptive in accurately indicating
the differences between dark and light tones * (text-fig. 69),

* [Note added June 1st, 1912.|—While this paper was in the printers’ hands
Mr. Lydekker wrote to the ‘ Field’ (May 11th, 1912), on behalf of Col. J. Manners-
Smith, to say that the Stag represented by the photograph above discussed is the
same individual as the one presented to the King by the Maharajah of Nepal. The
history of this animal was recorded by Col. Manners-Smith in the ‘ Field, July 31st,
1909, p. 239. It came from the upper reaches of the Sanpo Valley, close to Lake
Mansarowar, where it was captured as a fawn, and was in its second year in 1909.
It is therefore in its fifth year at the present time, and is not, as I supposed, an old
animal. That the colour was the same three years ago as now may be inferred from
Col. Manners-Smith’s reference to it as “very light.” As regards the distribution
of the species, reports stated that the deer was plentiful in various places in Western
Tibet, near the source of the Brahmaputra River, the hills north of Mount Kailas,

RARE STAG FROM NEPAL. 565
Since then the only known specimens belonging, or presumably

belonging, to C. wallichii were menagerie animals, no confidence
can be placed in their alleged localities.

Text-fic. 69.

Cervus wallichii.

The photograph, forwarded to Mr. Lydekker, of the specimen shown in text-fig. 67
(p. 561).

For a variety of reasons, into whica it is needless to enter,
C. wallichit has hitherto found no abiding place in the chronicles
of the Cervide. Some authors have added its name to the
synonymy of the Kashmir Stag Cervus hanglu (= cashmeriensis),
others have supposed the Stag to be identical with Cervus affinis ;
others have left it as indeterminable. To settle, as far as may
be, its status and affinities it is necessary to examine in some
detail the characters of the two above mentioned species with
which it has been confounded, and of certain other Stags
belonging to the same group of the genus Cervus.

and other less well-known localities being mentioned; the available evidence indi-
cating, in Col. Manners-Smith’s opinion, that the species does not cross the southern
watershed of the Brahmaputra and is not found in Nepalese territory. He suggested,
therefore, that the original specimen of C. wallichii had been brought to Muktinath
from somewhere in Western Tibet round about Lake Mansarowar. It may be
added that Col. Manners-Smith, apparently unaware of the discrepancies between
the descriptions of C. walliehii and C. affinis, believed these two forms to be
identical.

566 MR. R. I. POCOCK ON A

Cervus hanglu*, better but wrongly known as cashmeerianus, is
a not uncommon Stag. Moderately good figures of it have been
published by Dr. P. L. Sclater (Tr. Zool. Soe. vii. pi. 30, 1870),
and by Mr. Lydekker (Deer of all Lands, pl. iv.), while there are
several photographs in existence of specimens that have been
exhibited in the Gardens or kept at Woburn. One recently

.

Cervus hanglu.

Specimen, living in the Gardens in March, to show the resemblance of the
caudal disk to that of C. affinis, as described by Hodgson.

published in the ‘ Field,’ Dec. 30, 1911, shows very clearly the
shape and extent of the caudal disk characteristic of the species.
The white on the inner side of the back of the thighs extends a
short distance upwards above the root of the short tail, but the

* Waener in Schreber’s Siug. iv. p. 352 (note), 1843. Various modifications of
the word cashmeerianus are also in use; but of these cashmeriensis appears to be the
oldest, since it was first introduced by Leith Adams in connection with a description
of this species (P. Z.S. 1858, p. 529).

RARE STAG FROM NEPAL. 567

yellowish or dirty-white right and left moieties of this area do
not meet in front of the base of that organ but are separated by
a median brown band, continuous with the colour of the back ;
this band may or may not run along the middle of the tail
itself. Thus the caudal disk is small, smaller in fact than in
most Deer of the Elaphine group. Nevertheless it is very
conspicuous by reason of the blackish setting formed by the
adjacent hairs of the hind quarters. So far as my experience of
two specimens goes, this caudal disk does not vary appreciably
with the season, but remains fairly constant in extent, shape and
distinctness. Nor does it alter with age, judging from the fact
that in a very old animal with worn teeth, which was recently
shot in the Gardens for senile decay, the disk was exactly like
that of a younger animal in perfect condition now on exhibition.
No doubt, however, it exhibits a certain amount of individual
variation. It will thus be seen that C. hanglw may be at once
distinguished from C. wallichii by its totally dissimilar caudal
disk *.

It is also darker in colour, being very decidedly brown but
fading to a rather paler hue on the flanks, which are lighter than
the legs, the latter being as brown as the back both externally
and internally. The coat, moreover, is markedly speckled owing
to the presence of a subapical pale annulus on the individual
hairs.

Another very noticeable distinction between all the specimens
of C. hanglu and the one example of C. wallichii that I have seen
is in the colour of the hairs about the mouth. The chin and
lower lip of C. hanglw are white and the upper lip up to the
nostrils is cream or dirty-white, so that the muzzle is rather
sharply contrasted in colour with the browner hue of the rest of
the face. Light rings round the eyes, too, are very marked.
They are not so marked in ©. wallichii and, as has been stated,
the muzzle and chin are decidedly fawn-coloured and darker than
the rest of the face at least in the winter coat.

Cervus affinis Hodgson was based upon the skull and antlers
of a Stag wrongly alleged to have come from the Sal forests in
Nepal (J. A. S. Bengal, x. pt. 2, pp. 721-724, pl., 1841). The
brow tine was long, projected forwards over the face and was
straight or had a sharply upturned tip. The bez was subequal
and subparallel to it and also had an upturned tip, the distance
between their points of origin being 24 inches. Above the bez
the beam reclined backwards and outwards for a short distance,
then bent sharply upwards below the origin of the trez and ended
in a pair of subequal terminal tines. This sharp upward bend of

* Blyth, who believed the Kashmir Stag and C. wallichii to be specifically the
same, explained away this difference by assuming that the caudal disk in the figure
of the type of C. wallichii was exaggerated (J. A. S. Bengal, xxx. p. 188, 1861).

568 MR. R. I. POCOCK ON A

the beam, obvious both in profile and front view, is absent in the
known antlers of C. wallichit.

In the J. A. S. Bengal, xix. p. 466, pl., 1851, Hodgson figured
and described another pair of antlers sent to him by Dr. Campbell
from Ding-cham, north of Sikhim. These he referred to
C. affinis; but they only serve to illustrate the variability to
which the antlers are liable.

Subsequently (J. A. S. Bengal, xx. pp. 388-394, pl. vii., 1851)
Hodgson redescribed the species “from abundant supplies of the
spoils [also sent by Dr. Campbell] ..... exhibiting both sexes
in various states of maturity ..... the skulls and leg-bones
being attached to the majority of the specimens.” He thus had
at his disposal what he described as “ unusually copious and
adequate material.” This is an important point to remember.

The description of the antlers given in this paper agrees
tolerably closely with that which he published in 1841. Certain
individual variations are pointed out, and it is stated that the
basal interval between the brow and bez tines varies from two to
over four inches, two being the usual distance.

After remarking that the specimens were in winter coat,
having been killed in February, Hodgson described the colour as
“earthy brown, more or less lutescent, the head and neck being
concolorous with the back; but the flanks are conspicuously
paled, and the belly as conspicuously darkened ..... the neck,
though paler below than above, is not very noticeably so. But
the paling of the flanks is as decidedly so as the nigrescence of
the belly ; the white and black forming a conspicuous contrast
on these parts..... The limbs are paler than the back, darker
than the flanks and they have an earthy-brown list down their
external and anterior aspect.”

Finally he said, ‘The tail is very short, and the caudal disc
remarkably small but conspicuous from strong contrast of colours.”
In another place he also spoke of ‘“ the small caudal disc,” adding
“ The little tail is white, like its disc, a dark mesial line dividing
the latter along the culmenal (sic) line of the tail.” *

This description of the small caudal disk with its median
dividing dark line extending on to the tail applies tolerably
accurately to the caudal disk of C. hanglw; and if Hodgson’s
published figure, bad though it be, of C. aginis be compared with
those of the former species published by Mr. Lydekker and
Dr. P. L. Sclater, it will be seen that the principal differences
between the two species lie in the form of the antlers and the
rather larger caudal disk with narrower median line in C. affins.
Taking Hodgson’s description as a whole, however, it amply
justifies and explains Blanford’s statement (Mamm. Brit. India,
p. 537) that the coloration of the two species is the same, and
equally discredits Mr. Lydekker’s assertion that “as regards

* The italics in this paragraph are mine,—R. I, P.

RARE STAG FROM NEPAL. 569

coloration [of afinis|] accurate information is wanting” (Game
Animals of India, p. 217, 1907).

The precise nature of the relationship, however, between the
two species is by no means so easily and certainly settled as the
literature seems to indicate.

In the British Museum there are three skins and a mounted
head referred to C. affinis. The head and one of the skins came
from Hodgson, the former being labelled “‘ North of Bhotan,” the
latter ‘‘ North India ” and bearing the date 1857. It is not cer-
tain, so far as I know, that these specimens are part of the material
that author described in 1851. Very possibly they are. If so, the
head is very much faded—which is likely enough considering that
it is exhibited in the gallery—hecause it is now a tolerably uniform
sandy fawn and not ‘earthy brown.” It is noticeable, however,
that the lips and chin are fawn-coloured as in the example of
C. wallichii in the Gardens, and not white as in C. hanglu. The
skin, on the contrary, which has not been exhibited, agrees fairly
well with Hodgson’s description, being for the most part dirty
brown and much darker both on the body and legs than is our
specimen of C. wallichii. The hair, moreover, is very coarse and is
of winter growth. The tail is cut away; but the white disk is not
nearly so distinctly divided mesially as Hodgson’s description would
lead us to believe, nor as is the case in C. hanglu. Remnants of
a brown dividing line are, however, traceable. Again, although
the disk is smaller than in C. wallichai and is set off by a margin
of darker hairs, the bases of these adjacent dark hairs are white,
and the disk could be made to approach that of C. wallichii in
size if the brown tips of these hairs were removed or deprived of
pigment. But be it remembered, the bases of the hairs adjoining
the croup-disk in our example of C. wallichii are also white, so
that the disk of the latter is actually, with the hair undisturbed,
as large as, or larger than, the disk in Hodgson’s affinis with the
hairs interfered with in the way supposed. In both specimens
the hairs are short and crisp.

The other two skins named C. affinis in the British Museum
belonged to the late Dr. Blanford and are labelled “Sikkim,
L. Mandelli” *. The coat is in a better state of preservation
than in Hodgson’s specimen above described. The general
colour is greyish brown, the hairs being pale basally, brownish
distally with a distinct subapical pale annulus, and a darker tip,
imparting a marked speckled appearance to the coat. The rump-
patch is small and white, but as in Hodgson’s specimen, and in
Hodgson’s published figure of C. affinis, it spreads on to the
posterior part of the croup in front of the base of the tail. In
one of the specimens it is very clearly defined all round by a
bordering of brown hairs unspeckled and considerably darker than
the back, but with white bases. In the other the hairs bordering

* Dr. Blanford once informed me that Mandelli’s localities are untrustworthy.

Proc, Zoou. Soc.—1912, No, XXX VIII, 38

570 MR. R. I. POCOCK ON A

the patch are only a little darker than the back and are speckled
to the edge of the disk. As m Hodgson’s skin the tails have
been cut away, and the croup-disk has a median abbreviated
irregular dark line, much more clearly defined in one skin than in
the other. This disk is of about the same size as in Hodgson’s
specimen, although the hairs of the coat generally are longer and
less close and crisped. The legs are paler, being fawn-brown down
the front and on the fetlocks, and the lips and chin are also fawn,
and not cream and white as in C. hanglu.

Reverting once more to the mounted head in the British
Museum, it may be added that the left ear shows signs of the
emargination of its upper border so evident in the living example
of C. wallichii. The right ear nevertheless has a straighter upper
rim. I suspect, however, that C. affinis has the ears shaped as
in C. wallichi.

Putting all these facts together, it appears to me that C. affinis
is nearly intermediate in coloration between C. wallichi and
(. hanglu, especially in the general tint of the body and limbs
and the size and division of the caudal disk, the most marked
character in which it resembles C. wallichii and differs from
CO. hanglw being the fawn colour of the lips and chin.

There is evidence also that both C. wallichit and C. affinis are
larger than C. hanglu and have longer faces*; but judging by
the standard of specific and subspecifie differences usually adopted
in the Cervide, it appears to me to be doubtful whether more
than subspecific importance should be granted to the differences
above described between C. wallichu, C. affinis, and C. hanglu.
It must be remembered, however, that wallichit is the oldest
name of the three.

Exact partieulars of the range of C. afinis are much wanted.
Only two districts are mentioned by Mr. Lydekker in the table
of horn-measurements in Rowland Ward’s ‘ Records of Big
Game,’ 1910 (p. 38), namely, the Tibetan Frontier and the
Choombi Valley; but the valleys of Bhotan near the Choombi
are added under the diagnosis of the species. The Stag has
also been recorded by Col. H. A. Iggulden from the Tsan-po
basin, near Lhasa (‘ Field,’ Oct. 1906, p. 736); but whether the
specimens were accurately determined or not, it is impossible,
without the evidence of skins, to say.

Another Stag belonging to this same group is the animal from
Szechuen described by Mr. Lydekker as C. cashmirianus macneillr
(P. Z.S. May 11, 1909, p. 588, pl. Ixix.). The coat is finely
speckled all over owing to the apical annulation of the hairs, as
in Mandelli’s skins of OC. affinis and as in fresh-coated examples of
C. hanglu; the prevailing colour, however, is strikingly grey,
especially on the sides and legs, but the back is rather darker,

* This difference is very noticeable between the examples of C. hanglw and
C. wallichii living side by side in the Gardens,

RARE STAG FROM NEPAL. 571

that is to say more fawn, than the sides, and the top of the head,
the face, and the nape of the neck are browner than the back.
There is no mane and the neck shows signs of coat change. The
legs are less distinctly speckled than the sides of the body and
are darker in front than behind. The belly is whitish, as in the
females of C. affinis described by Hodgson. Apart from the
prevailing greyness of the pelage, the most interesting features
about this Stag are the brownish fawn colour of the chin and lips

Text-fig. 71.

Cervus macneilli.

Photograph of the plate of the type-specimen (P. Z. S. 1909, pl. Ixix.).

and the coloration of the rump. The upper side of the tail is
black with a narrow edging of white. On each side of the tail
there is a narrow white area, which however barely surpasses
the root of that organ dorsally. This white area is bordered with
black, and there is a blackish-brown unspeckled croup-disk almost
as extensive as the white croup-disk of C. wallichii.

From the distribution of this Stag, its affinities might be
inferred to be rather with C. affinis than with C. hanglu ; and in

38*

572 MR. R. I. POCOCK ON A

spite of the lesser extent of the white on the rump, a feature in
which it more resembles C. hanglu, this conclusion is borne out
by the coloration of the chin and lips and by the shape of the
ears, which are pointed and appear to have had a sinuous upper
edge.

Moreover, as Mr. Lydekker records, Capt. McNeill declared
the Stags of this kind that he saw in Szechuen to be nearly as
large as Wapiti. Even making allowance for exaggeration in
this estimate, the animals must have been of considerable size.
CO. hanglu, however, is not’a large Stag, although strongly made
and “ cobby ” in build.

Unfortunately, there is no record of the date when the type
specimen of this Stag, a young hind standing, as mounted, forty
inches, was shot.

Another type of this group of Stags is represented by askin and
skull kindly lent to me for identification by Mr. W. F. H. Rosen-
berg, F.Z.S. The animal, an adult hind, was shot by Dr. J. A.
C. Smith on Mareh 23rd, 1911, 30 miles 8.E. of Tao-chou, Kansu,
in China, at an altitude of 11,000 feet.

The colour is a tolerably uniform earthy brown relieved by fine
close-set speckling due to a subapical pale annulus on each hair.
On the sides the main shaft of the hairs is greyer and less brown
than dorsally, and low down towards the belly the subapical
annulus is longer, so that the general tint is markedly paler.
The belly is white, but not the chest. On the neck the hairs are
longer with longer apical annulus, the shaft of the hairs being
browner along the nape than on the sides of the neck, so that
there is an ill-defined dorsal neck-stripe as in C. macneilli;
the front of the neck (throat) is paler than the sides ; the legs are
fawn-brown down the front and sides, paler behind. The forehead
is brown, the face grey-brown and closely speckled ; the lips and
chin are fawn-brown, unspeckled and without white, and the
black patch below the corner of the mouth is well marked. There
is a blackish-brown unspeckled croup-disk, as in C. macneilli,
and the hairs of this disk become gradually more and more white
towards the root of the tail*. The white on the buttocks is of
the same extent approximately as in C. macneilli, but the tail
itself is much whiter than in that animal, since it merely has a
narrow median dark stripe as in most examples at all events of
CO. hanglu. The ears are long and pointed, with apparently a
sinuous upper edge such as is seen in (. wallichit.

The following measurements in the flesh, taken by Dr. Smith,
may be of future use, although at the present time they indicate
nothing, because there are no corresponding measurements of
other deer wherewith to compare them :—

Head and body 1745 mm. (a little under 6 feet ; tail 145 mm.

* This is probably also the case in C. maeneilli, but I was unable to touch the
mounted specimen of the animal in the British Museum,

RARE STAG FROM NEPAL. 573

(under 6 inches); foot (? hock to heel) 525 mm. (21 inches) ; ear
225 mm. (9 inches). The skin is too shrunk to make an estimate
of the animal’s height of any value.

The skull and teeth show that the specimen was a full- grown,
but not old, hind. Its basal length from the notch between the
condyles to the tip of the premaxille is 132? inches (about 344 mm.);
and the length from the anterior edge of the orbit to the tip of
the premaxilla about 9 inches (= 225 mm.), and the width across
the orbits 67 inches (=156 mm.).

On comparing these dimensions with those recorded by Blantord
for C. hanglu and C. affinis, it would appear that this Kansu
Stag is somewhere about the size of the former. For example,
he quotes the length of the Hangul (probably of males) as from
7 to 7% feet and the length of a skull of this sex as just
over 15 inches. Probably the basal length of the latter would
be about 142 inches.

The omellise of two skulls of C. affinis had a basal length of 163
inches and a width across the orbits of about 74 inches. Thus,
allowing for the fact that the Kansu specimen is: an adult female,
it may be inferred that the males are about as large as those of
C. hanglu and considerably smaller than those of C. affinis.

The description given above of the colour of this Stag agrees in
a general way tolerably closely with that of the examples of
C. affinis that Hodgson had in his hands, and it is important to
remember the close correspondence in date, Hodgson’s specimens
having been killed in February and the Kansu specimen in March ;
and since deer of the Hlaphine group moult in April and May,
it is evident that the examples in question were still carrying
their winter coats. In their generally dark earthy brown color-
ation, both differ markedly from the living example of C. wallichi
in March.

The coloration of the Kansu specimen, however, is not identical
with that of C. afinis. I do not think it safe on the evidence of
one skin to trust much to the greater uniformity of the colour of
the body as shown by the absence of a distinct darkening of the
ave and “lutescence” of the sides to which Hodgson refers in

C. affinis, but the large size of the dark croup-patch and the
smaller extent of white at the base of the tail are probably more
dependable. It is in both these particulars especially that the
Kansu stag resembles the Szechuen stag C. macneilli. Since,
however, it differs from the latter in general coloration and in the
greater amount of white in the tail; from C. affinis in having no
white above the root of the tail and a larger dark area on the
croup; from C. wallichit in having no white on the croup at all,
except such as is concealed by the overlying black ends to the hairs ;
and from C. hanglu in the dark colour of the chin and upper lip,
the Kansu stag seems to deserve a name; and I propose to call
it C. KANSUENSIS.

5TA MR. R. I. POCOCK ON A

The following analytical key to the species above discussed
represents my views as to their probable affinities :—

a. Chin and lower lip white, muzzle pale fawn, markedly lighter
than the rest of the face; ears bluntly pointed and with
straight upper rim; no white on croup above root of tail... hanglu Wagn.

a’. Chin fawn or brown, muzzle brownish fawn with at most
a little white on the lip in front; ears loug and pointed
with sinuous upper edge (? in affinis).

6. White on back of thighs not spreading on to croup
above root of tail; tail dark down centre; a large
blackish-brown croup-patch reaching nearly er quite to
the summit of the croup.

ce. Prevailing colour grey, rather darker on the back and
still darker on the head; practically the whole of the

upper side of the tail black . Listeteeecssecteeernseeeeee macneills Lydd.
. Prevailing colour (March) brown; upper side of tail
with an irregular median dark stripe......................-. kansuensis, nov.

b’. White on back of thighs spreading upwards above the root
of the tail and encroaching more or less on the croup;
dark patch on croup smaller or absent.

d. White area above the tail comparatively small with a

more or less distinct median dark longitudinal stripe;

prevailing colour (February) earthy brown, paler
laterally . : affinis Hodgs.

d’. White area above tail very ‘large, ‘yeaching ‘summit of

croup and undivided by dark line; prevailing colour
(March) pale fawn-brown...............06c000-cceeeeeeeeeeees. Wallichii Cuv.

In selecting the colour of the chin and muzzle as the character
for eliminating C. hanglu from the rest of the species, I am not
unmindful of the fact that Cuvier wrote of C. wallichii “ comme
Vordinaire, le tour de ceil, celui de Ja bouche, sont plus pales, et
ily a du blanc sous la mAchoire et une tache noire sous langle
des lévres.” These words may suggest that the muzzle and
chin were coloured as in C. hanglu; but the coloured plate of
C. wallichit, from which the description was taken, does not bear
out this supposition nor does it quite justify Cuvier’s phraseology,
for the lips and chin are washed with the same yellowish tint as
that of the rest of the body.

If the caudal disk were taken as the primary basis for the
classification of the species, they would be grouped as follows :—
C. macneillit+ C. kansuensis; C. hanglu+C. affinis; C. wallichii.
I prefer on the whole to regard these forms as species rather than
subspecies because we have at present no proof that they inter-
grade, and the differences between them are perhaps greater than
those between the various races of Wapiti (Cervus canadensis)
and Red Deer (Cervus elaphus). As a group they resemble the
Wapiti and differ from the Red Deer in the shortness of their
tails.

The only other Stag which should perhaps come into the same
category, Judging from the shortness of the tail, the size of the
caudal disk, and the shape of the ears, is the Tibetan species
commonly known as C’.. albirostris which Blanford described as
C. thoroldi (P. Z. 8. 1893, p. 444). Judging from the mounted
specimen of this stag in the British Museum the coloured figures
of this species published by Blanford and by Mr. Lydekker

RARE STAG FROM NEPAL. 575

(‘Deer of all Lands,’ pl. v.) have the yellow caudal disk too
large and extending too high up the croup, and the ears, which
are really very long and pointed, with the upper edge sinuous,
represented as much too short and blunt. In the whiteness of
the chin and muzzle this species goes a stage beyond C. hanglu ;
but of course the stag differs from those discussed in this paper
by the reversal of the hairs along the spine between the croup
and withers and, so far as is known, by the absence of the bez
tine.

In the above given table no use has been made of the antlers
as distinguishing features. These vary so much with age and
from a variety of unknown causes that I am convinced too
much confidence is placed in them in most systematic treatises
on Deer*. At the same time it may be useful to bear in mind
that the recorded antlers of C. wallichit appear to differ from
average antlers of C. hanglu and C. affinis in the basal juxta-
position and marked distal divergence between the brow and bez
tines, and at least from those of C. affinis in the comparative
straightness of the beam which lacks the sharp upward curvature
of its distal half. It will be highly interesting to see if the
example of C. wallichii now living in the Gardens grows, under
the changed conditions of its existence, antlers resembling those
that it shed on its journey from India. ‘This and the colour of
its summer coat I hope to have the chance of recording later on.

As regards the stature of the Deer described above, Blanford
gave the height of C. hanglu as ranging from 48 to 52 inches.
The example of this species now in the Gardens is barely 48 inches.
C’. affinis, according to Hodgson, stands from 54 to 60 inches
-and thus rivals a Wapiti in stature. A small Canadian stag
Wapiti now in the Gardens is about 56 inches, and an example
of C. xwanthopygus 58 inches. But since Hodgson’s measurements
of C. affinis were taken from dried skins, his estimate must have
been largely guess-work and probably erred on the side of
exaggeration than otherwise. Blanford, or the authorities from
whom his information was derived, measured in all probability
freshly killed specimens of C. hanglw; and since dead animals
give higher stature-measurements than living ones, exaggeration
in connection with this species must also be allowed for.

Our living example of C. wallichit is as nearly as possible
51 inches, and is manifestly much smaller than any stag of the
Wapiti group, whether American or Asiatic, that I have seen.
It is, however, exactly the height assigned by Hardwicke to
the typical example of this species that was exhibited in the
menagerie at Barrackpore.

* T can see no reason for adopting Mr. Lydekker’s suggestion (‘ Field,’ May 11th,
1912) that the Stag he described from a pair of shed antlers picked up in Szechuen
as Oervus canadensis wardi (P.Z.S. 1910, pp. 987-989) is identical with Cervus
wallichii.

576 DR. F. E. BEDDARD ON

31. Contributions to the Anatomy and Systematic Arrange-
ment of the Cestoidea. By Frank H. Bepparp, M.A.,
D.Sc., F.R.S., F.Z.8., Prosector to the Society.

[Received February 6, 1912: Read April 2, 1912. |
(Text-figures 72-83.)

IV. On A Species or /WERMICAPSIFER FROM THE HyRAX, AND
ON THE GENERA ZSCHOKKEELLA, THYSANOTZNIA, AND

HYVRACOTANIA.
INDEX.

Systematic : Page
Inermicapsifer capensis, sp.n. .................. 598
yracotenia, SON. VW. ......... cc cee creer reese sees. 604
TELS QEYDAUSID, Sb Ws cosccdeoaes veasopcesosacosesccsces ODE!
Jel OTRUHOES GOs Wo cooadanoose0 congener oooctascéoanocn (OOS
Thysanotenia (corrections and additions) ...... 606
ZSCHOWK CELLO Rte ciecctaeeehnes acnmp ase see seee eho LOOe.

I obtained during the month of October 1911 a large number
of fair-sized unarmed tapeworms from the gut and from the gall-
bladder of an example of Procavia capensis, and I referred them
provisionally to the genus Zschokkeella of Fuhrmann* in my
Report to the Prosectorial Committee, in spite of the fact that
the type species of that genus is a parasite of the Guinea-Fow]
Numida ptilorhyncha. I now refer them partly to the more
recently instituted genus /nermicapsifer t, the aftinities of which
with Zschokkeella and my own genus Thysanotenia, I shall deal
with in the present communication. The author of the genus
Inermicapsifer—and in the paper referred to—assigns to that genus ©
with certainty or with more or less doubt eight species. I shall
compare these individually with the species I have found myself.
Dr. Janicki is certainly justified in saying that “the fact is of
interest that the genus Jnermicapsifer is parasitic in Procavia
with several nearly related species.”

From the intestine of Procavia capensis I have gathered
specimens of several distinct species, of which one was much
more abundantly represented than the others. The more
abundant species is represented in the accompanying text-figure
(text-fig. 72).

It is a long and rather slender worm, about 95 mm. in length
and of 2 mm. greatest diameter. There are fully 200 proglottids
in this example. The head is very conspicuous and half as wide
again as the part of the strobila which ensues. Anteriorly the
worm is more slender, of less diameter, than it is posteriorly ;
the increase is gradual, there being no sharp demarcation. The
body of these worms has a great tendency in specimens preserved

* Centralbl. f. Parasit. etc. Bd. xxxii. 1902.
+ Janicki, “ Die Cestoden aus Procavia,” in Schultze’s Zool. Ergebn. Forschungs-
reise in Stidafrika, Jena Denkschr. med. Ges. xvi., 1910.

NEW TAPEWORMS FROM THE HYRAX. 577

in alcohol to contract upwards or downwards, thus forming a
hollow or convex surface as the case may be. The characters
already given allow us to separate the present species from
I. hyracis*, in which the scolex is not of greater diameter than
the ensuing region of the strobila, and from J/. seté#i t, where the
total number of proglottids is very small (7. e. 30-70).

Text-fig. 72.

A nearly complete example and two fragments of Inermicapsifer capensis.
X 2.

I do not at this stage of my description attempt to differentiate
between this species and the others dealt with in Janicki’s

* Janicki, loc. cit. pl. xii. fig. 1.
+ Janicki, loc. cit. pl. xii. fig. 10.

578 DR. F, E. BEDDARD ON

monograph, since there is some little variation in the characters
of the numerous examples which I have to consider here, which
variations do not militate against the exclusion of J. hyracis and
I. settii, though they render difficult the distinguishing of my
species from some of the others referred to by Janicki. Thus
two examples measured only 68 mm. and 75 mm. respectively,
and one of them was quite 3 mm. in diameter at the widest part.
The segments were never longer than broad, except perhaps a
trifle longer in the thin anterior region of the body. They
varied somewhat in relative dimensions, as is the case* with
those of /. hyracis. Occasionally the unilateral genital pores were
very visible on slight papille near to the hinder end of segments.

The scolex is quite unarmed and conspicuous by its size, as
already mentioned. It is usually more or less globular in form,
with a slightly prominent and somewhat pointed apex, as is
shown in Janicki’s figure of /. seééiit, and which is, of course,
the rostellar region. Occasionally, however, this apical region
does not protrude, but is represented by a depression on the
surface. ‘The scolex is, moreover, sometimes flattened in its
entirety from above downwards, and presents a mushroom-like
appearance, the edges of the disc projecting round the neck,
which thus presents the appearance of the stalk of the
mushroom.

Longitudinal sections through the scolex confirm the total
absence of hooks or of any trace of a rostellum, save the slight
projection already mentioned. Nor could I detect anything
peculiar in the structure of the suckers, which, according to
Janicki and others, are remarkable for a funnel-like ingrowth
leading to the actual sucker, which thus lies at the bottom of
a depression. It is true that in the present species, as in many
tapeworms of the group Tetracotylea which I have examined,
the sucker does not lie externally on the scolex, but is covered by
an outer layer of body-wall which is only interrupted at the
orifice of the sucker. The free edge of this, when depressed
towards the interior of the sucker, is doubtless funnel-shaped
and would give rise to the appearances represented by Janicki.
Perhaps, however, there is some divergence from the normal
condition of the suckers in the members of the genus referred
to by him, which I certainly have not found in the species
with which I am at present concerned. The direction of the
suckers seems to me to vary somewhat; generally they are
lateral, but sometimes with a forward inclination.

It has been already observed that in this genus water-vessels
of unusual width extend far into the head. Such are figured by
Janicki, and referred to by Pagenstecher { (quoted by Janicki §)
as “einen mit Ringmuskeln umspinnenen Wasserbehiilter.” In.

* Janicki, loc. cit. pl. xii. figs. 2-5.

+ Loc. cit. pl. xii. fig. 10.

[ Zeitschr. f. wiss. Zool. Bd. xxx. 1878.
§ Loc. cit. p. 389.

NEW TAPEWORMS FROM THE HYRAX. 579

the present species a good deal of the head lying between the
suckers is occupied by the coils of water-tubes, which approach
very nearly to the external surface at the apex of the head.
These tubes are not in any way irregularly dilated, but their
diameter throughout is rather greater than that of the tubes in
the immediately following neck-region of the body. Janicki is
not disposed to lay great weight upon the absence or presence of
a neck in these tapeworms. In the present form the strobili-
sations begin very close to the head, which is supported by a
segment broadening towards the head, which may be regarded as
the neck. In the accompanying text-figure (text-fig. 72) will be
seen the general characters of the strobila of this worm.

The body-wall of the worm is relatively thick, but here the
cortical layer is not so definitely marked off from the medullary
parenchyma as is so often the case. The absence of a strong de-
limitation is due to the very feeble development of the transverse
musculature, which is generally hardly, or not at all, recognisable
in transverse sections. Another reason which renders the two
layers more uniform in this species than in my Thysanotcnia
gambiana and in the species of Hyracotenia described in the
present paper, is that there are stout muscular fibres in the
medullary parenchyma comparable in thickness to the longitu-
dinal muscular fibres of the cortical layer.

Water Vascular Tubes.—These tubes are, as usual, two on each
side. They lie, as a rule, almost parallel with each other, the
smaller dorsal tube having, however, an inclination to the dorsal
side which is more or less pronounced as the case may be. The
more medianly situate ventral tube is the wider, but the difference
between the two is not perhaps quite so well-marked as in
Thysanotenia gambiana. The two vessels are not united across the
proglottids by a regular series of transverse vessels, one to each
proglottid, but, as in other species of this genus and in Thysano-
tenia gambiana, are united by a network of tubes found all over
the segments. I did not follow out into any detail the precise
arrangement of the branching connecting-tubes.

The ovary, when fully mature, appears to be definitely a single
body lying anterior to and in contact with the vitelline gland.
The latter lies on a level with the group of testes of the pore side.
The ovary lies, when less fully mature, entirely to the median
side of the two water vascular tubes. Later on, it appears to
extend somewhat and is larger actually, and considerably larger
relatively, than I found it to be in Thysanotenia gambiana.
The ovary is clearly also larger in the present species than in
Inermicapsifer hyracis, according to Janicki’s figures. In the
later stages of the ovary, when it is gorged with ripe eggs and
these are beginning to spread through the proglottid, a peculiarity
of structure, perfectly plain in earlier stages, is not obvious.
One such example, representing the ovaries, is represented in
text-figs. 73 and 74. It will be there seen that the ovary really
consists of clumps of ripe and unripe eggs situated at the ends of

580 DR. F. E. BEDDARD ON

stalk-like processes of the oviduct, from which they radiate out.
There is some evidence that these are probably hollow outgrowths
of the oviduct, the main stem of which is continued on to form
the rudimentary (at any rate, short) uterus. There is also some
evidence that these outgrowths of the oviducal canal form here
and there anastomoses with each other. In the fully mature
ovary the ripe eggs are seen to be in groups surrounded bya thin
wall and with empty spaces surrounding eggs or groups of eggs.

Text-fig. 73. Text-fig. 74.

Text-fig. 73.—Ovary and vitelline gland of Inermicapsifer capensis in section.
o. Ovary. V. Vitelline gland.
The masses of ova are seen at the end of stalk-like processes.

Text-fig. 74.—Another section through the same ovary to show the rudimentary
uterus (wt.).

It seems to me that we have here a later stage of the tubes
communicating with the oviduct. From this point of view the
ovary would appear to be a compound body, in reality consisting
of many ovaries; the indications of a network among these
ducts is particularly interesting in view of the network formed by
the testicular ducts, and brings the male and female organs into
closer agreement.

NEW TAPEWORMS FROM THE HYRAX. 581

The vagina opens on the exterior behind the aperture of the
cirrus sac and immediately contiguous to it. As will be men-
tioned in describing the cirrus sac there is no common cloaca.
The direction of the tube from its opening into the receptaculum
seminis is obliquely backward, the external pore lying at the side
of the segment posteriorly. The terminal portion of the vagina,
the length of which is not far from equal to that of the cirrus sac,
is somewhat dilated and with a wider lumen ; it is plainly ciliated.
The whole of the vagina is very muscular, the muscular layer
being as thick as that of the cirrus sac. The lining epithelium
is deeply stained. The course of the vagina is very straight until
it bends somewhat at its entrance into the large receptaculum
seminis. The latter is bent round the outermost of the two longi-
tudinal water vessels. From its opposite end arises the wide
common duct, which after a short course divides into the posterior
vitelline duct and the anterior oviduct. The two latter are in
the same straight line.

A closer examination, however, of the long muscular vagina
shows that it is really divisible into two regions. These are
inequisized, the larger section being that which is nearer to the
external pore. This section has a much wider lumen than the
ensuing section, which opens into the wide and thin-walled
oviduct. The latter has a narrow thread-like lumen, but still
thick and muscular walls. It is not so long as the distal region
of the vagina. This part corresponds to the very short and
suddenly abbreviated part of the vagina in Thysanotenia
gambiana ; but is obviously very much more developed. It is
not in the same straight line as the wider distai section of the
vagina, but bends back to meet the forwardly running oviduct.
This division of the vagina appears to be very characteristic of
the present species.

At the time that I was preparing my account of the anatomy of
the two species of my genus Thysanotenia I was not acquainted
with Janicki’s memoir upon his genus /nermicapsifer, and therefore
laid no stress upon some structural features which are of
importance in comparing these forms. I have therefore carefully
re-examined my sections of Thysanotenia gambiana, and made in
addition several series of fresh sections, in order the better to
accomplish this comparison. By these means I am in a position
to make a few corrections in, and additions to, my earlier
account.

I find that account, however, to be correct in all essentials,
excepting with regard to the presence of a vesicula seminalis
in the neighbourhood of the ovary. ‘This was doubtfully
asserted, however, and I now find that the structure in question
is really a part of the dilated and thin-walled oviduct, which runs
straight, as I have said, from the thick-walled region near to the
external pore and gradually increases somewhat in width as it
approaches the ovary. I confirm my account of the variability
in the lateral extension of the ovary and vitelline gland, which

582 DR. F. E. BEDDARD ON

are sometimes restricted in their extent to the space lying
between the dorsal and ventral vessels. In incompletely mature
proglottids this restriction appeared to be (at least usually) the
case, the gonadial tissue not extending outside or inside of the two
excretory tubes. I should also emphasize the fact that the ovary
and vitelline glands are, at least for part of their extent, situated
side by side. That is to say, the vitelline gland is not wholly
behind the ovary. In young proglottids the general lie of the
female gonad seemed to me to be rather oblique dorso-ventrally,
in accordance with the fact that the duct passes to the exterior
ventrally of the dorsal excretory vessel and that the two excretory
vessels lie in a nearly parallel plane.

T have also a new fact of some little importance to add. I
have described the uterus in the sexually mature segments
as being a tubular structure running across the segments. I
have in younger specimens discovered the rudimentary uterus,
which consists of a solid chord of closely packed cells which
enclose no lumen. This is connected with the gonad tissue
lying between the two excretory tubes and passes towards, but not
far towards, the middle line of the body above the larger ventral
excretory tube, but below the sperm-duct. The rudimentary
uterus ends in a club-shaped blind extremity.

My former account of the testes of Thysanotenia gambiana
is confirmed by the examination of fresh material. I have,
however, two points to add: the first is that while the testes
are practically grouped as there stated into two groups, there
are, nevertheless, a very few testes lying between the two ventral
excretory tubes, and thus really constituting a bridge between
the two separate masses, and bringing the arrangement of the
testes more into line with that of the second species of the
genus Thysanotenia described in the paper referred to. These,
however, are indeed few; I counted but three in a single
proglottid. The general statement with regard to the testes
remains, therefore, unaffected. Moreover, each testis is rounded
or oval in form, but I could not find the distinctly pear-shaped
outline that I describe in the species of nermicapsifer dealt with
in the present paper. I should, furthermore, add to my former
account the fact that the testes are mature before the ovaries,
and that in segments where the ovary is still immature the
vagina, as well as the vas deferens, is filled with sperm. The
sperm-duct is coiled and the coil hes principally between the
nerve-chord and the dorsal excretory tube, extending to a short
distance mediad of the latter. The coils have some regularity
and lie dorso-ventrally and parallel. I have found granular
tissue accompanying the sperm-duct. Cells lay here and there
between the coils, sometimes in numbers.

The testes of the worm from the Hyrax are very distinctly
arranged in two sets in each proglottid. The great bulk of these
organs are on the side furthest away from the external pore. I did
not find a fewscattered testes uniting these two groups such as occur

NEW TAPEWORMS FROM THE HYRAX. 583

in Thysanotenia gambiana. The point of difference, however, is
not a large one, since in other species of the present genus there
is no differentiation of the testes into two masses, but they form
a continuous mass reaching from side to side of the proglottid.
In the case of the group of testes on the pore side, they lie to the
pore side of the ovary and vitelline gland. These testes were
very conspicuously pear-shaped, the duct naturally arising from
the pointed end. In the case of the testes on the pore side, at
any rate, the narrower end faced forwards in the segment. In the
case of each group the testes are closely packed together. They
are restricted to the medullary part of the parenchyma, The
efferent sperm-ducts from the testes which lie away from the
pore side form a widish trunk which breaks off and forms a slight
rete mirabile not so marked as in the species next to be described.
This opens evidently by one thick tube into the large vesicula
seminalis lying close to the ovary and to the receptaculum seminis,
The cirrus sac is not conspicuously large, and it lies close to
the lateral margin of each proglottid and thus well within the
cortical parenchyma. Its position is oblique, the end of the
sac being anterior to its aperture on to the exterior, and is
thus better shown in horizontal than transverse sections of the
proglottids. The general outline of the cirrus sac, when displayed
in such sections, is, like that of many other species (e. g.
Otiditenia eupodotidis*), flask-shaped, there being a narrower
proximal neck and a wider distal region. It lies parallel to the
vagina and very close to it, but in a different plane, so that a
single horizontal section does not display both tubes lying side
by side. Such a section, however, does show the extreme
terminal part including the external orifice of both cirrus sac and
vagina; and it will be seen that they lie accurately side by side,
the male pore being, of course, anterior to the female pore. Such
a section also shows very plainly that there is no trace of a
cloaca genitalis ; the two pores are upon the surface of the body
upon a slightly raised papilla of not very great dimensions. The
papilla which bears the two orifices is clearly of less importance
than that which characterises the two worms for which I have
recently formed the genus Thysanotenia t. Ona re-examination
of my sections of the species Thysanotenia lemuris I find that
in that species, as well as in that which forms the subject of the
present communication, there is no sinus genitalis, but that both
cirrus sac and vagina open side by side on the summit of the
papilla. The case is otherwise with Thysanotenia gambiana,
where there is a distinct though not very large sinus or cloaca
genitalis. There are also corresponding differences in the vagina
and cirrus sac of the latter, which I shall deal with after describing
the arrangements found in the Anoplocephalid from the Hyrax.
The cirrus sac has a thickish muscular coat which does not
become at all markedly thicker in the “neck” region. In

* P.Z. 8. 1912, p. 206, text-fie. 27,
+ PZ, 8. 1911, p. 1002.

584 DR. F. E. BEDDARD ON

longitudinal sections it appears that the fibres constituting
together this muscular wall run more or less along its longer
axis. It will be observed that in both of these characters the
cirrus sac of this species differs from that of a genus recently
described by myself as Otiditenia *, and, moreover, the cirrus sac
is very much larger in the latter genus. The cirrus itself is very
inconspicuous and occupies only the neck region of the sac; it is
thus necessarily very short. It is distinguishable from the
sperm-duct by reason of the fact that it is very darkly stained
with both carmine and hematoxylin. Between the cirrus and
the muscular walls of that part of the cirrus sac in which it lies
is a great accumulation of nuclei, which belong, as I presume, to
slender muscular fibres concerned with the retraction of the
cirrus.

As in other species the cirrus sac, where it swells out into the
rounded flask-like body, is filled with a delicate packing tissue
with abundant nuclei. Through this passes the sperm-duct in
two or three coils. The delicate sperm-duct takes up but little
stain and is thus very distinct from the cirrus. I found this
condition of the sperm-duct to exist in a segment posterior to
others in which the sperm-duct had undergone even further
modification. In the latter segments the sperm-duct lying within
the cirrus sac is dilated to form a vesicula seminalis. This
dilated duct is also coiled; but the two or at most three pieces
seen in an individual section completely fill the lumen of the
cirrus sac, with the exception of dividing lines filled with nuclei
belonging to the internal tissue of the cirrus sac. The fact
that an unaltered sperm-duct may lie behind one which is
converted into a vesicula seminalis is important.

It is clear that, on the whole, the cirrus sac of this worm is
more like that of the two species which I have referred to a
new genus, Thysanotenia, than it is to that of, for example,
Otiditeniat or Anoploteniat. It differs, however, in details
from the cirrus sac of both of the two species which I have
temporarily placed in the genus Thysanotenia. In the con-
cluding part of the present communication it will be necessary
to go fully into the systematic position of this worm and _ to
compare it especially with the two species of the genus Thysano-
tenia. It will be therefore convenient at the present moment
to compare the cirrus sac in these different forms. They agree
generally in the absence of a distal region, which I have termed
penis in Anoplotenia, the cirrus being rod-like up to its free
extremity and not lying at the bottom of an invaginated part of
the cloaca genitalis. Again, in all three species the general form
is the same, and the muscular layer runs along the longer diameter
of the sac and is not specially thickened at the “neck” end.
These features exhaust the general resemblances between the

* P.Z.S. 1912, p. 206, text-fig. 27.
+ P.Z.S. 1912, p. 194.
{ P. Z.S. 1911, p. 1015, text-fig. 215.

NEW TAPEWORMS FROM THE HYRAX, 585

cirrus sacs of the three tapeworms. That of Thysanotenia lemuris
differs from the other two by its larger size, the cirrus sac in the
two remaining species being of about the same size. The cirrus
sac of Thysanotenia lemuris is, indeed, fully twice the size that it
is in the two other species. In the species which forms the
subject of the present communication the cirrus is very small
and limited to the neck part of the cirrus sac, than which it is
no longer. The same characteristics apply to Thysanotenia
lemuris, only that as the cirrus sac itself is larger, the cirrus also
is larger than that of the species with which I compare it. In
both of these species the rest of the cirrus sac is filled with the
sperm-duct, which is, in both, dilated to form a vesicula seminalis;
but in Thysanotenia lemuris the dilatation fills the whole sac,
while in the other species it is coiled and thus fills the sac in a
different way. In the species Thysanotenia gambiana there is no
such conspicuous vesicula seminalis, but just after the entry of
the sperm-duct into the cirrus sac it is dilated for a short space.
On the whole, it appears that the cirrus sac of the present species
is more like that of Thysanotenia lemuris than of Th. gambiana.

One of the principal differences which this species shows from
Thysanotenia gambiana is in the character of the uterus. As I
have already mentioned in the present paper, as well as in my
memoir dealing with that species, the uterus is very plain, first
as a solid cord and then a narrow tube with an obvious lumen.
An examination of a large number of sections of the species of
Inermicapsifer described here has shown nothing exactly like the
uterus of Thysanotenia gambiana. Nor can I reconcile what I
have seen with the figures of Janicki’s illustration of the uterus
of Inermicapsifer hyracis. In the latter species the uterus is
figured * in horizontal sections through the ripe proglottid as an
irregularly shaped sac with numerous projections and outpushings
of its lumen—as, for example, in the genus Tetrabothrium +.
It is shown, however {, to commence as a sinuous tubular
structure, which appears to me to resemble very much the uterus
as I have described it in Thysanotenia gambiana. Farlier still
than this a solid cord of cells issues from the generative mass
which again would appear to be exactly comparable to what I
have seen in Thysanotenia gambiana. Janicki, however, com-
ments upon the remarkable fact that the formation of a lumen in
this cord begins, not where it would be expected to begin, at the
ovarian end, but towards the middle of the segment. It is here,
moreover, that in Thysanotenia gambiana the solid cord of cells,
which subsequently becomes the hollow uterus, widens out into a
club-shaped extremity ; but I am unable to confirm or differ from
Janicki in fixing the point at which the uterus begins to be
hollowed out to be coincident with this club-shaped extremity.
It seems, however, to be very likely.

* Loe. cit. pl. xii. fig. 13.
+ Spatlich, in Zool. Jahrb. Bd. 28 (1909).
{ Loe. cit. pl. xii. fig. 17, ut.

Proc. Zoou, Soc.—1912, No. XX XIX, 39

586 DR. F. E. BEDDARD ON

The examination of more than one series of sections has shown
me that, in the species of /nermicapsifer which forms the subject
of the present communication, the commencement of the forma-
tion of the uterus is in many respects different from that of
Inermicapsifer hyracis as described by Janicki and of Thysano-
tenia gambiana as described by myself. The earliest stages that
I have observed are perhaps best seen in horizontal sections,
though I have also observed them in transverse sections. In
horizontal sections, such as is represented in text-fig. 74, the
ovary lies in the middle of the segment from before behind and
extends some way towards the middle of the segment from right
to left. The oviduct with its numerous branches, which has been

Text-fig. 75. Text-fig. 76.

Text-fig. 75.—A portion of the medullary parenchyma of Inermicapsifer capensis,
showing centrally a condensation of tissue which is connected with the
rudimentary uterus.

Text-fig. 76.—A portion of the ovary of the same species with eggs (0.) detached
from the ovary and lying in the parenchyma, and a portion of the
network of tissue (#.) which may possibly represent the uterus.

already described, is seen to be prolonged into an extension not
distinguishable from it, which runs for a little way into the
medullary parenchyma towards the median point of the proglottid.
It is quite short, and ends more or less abruptly in a strand of
condensed parenchymal tissue, which is apt to be branched,
sending out shorter strands of a similar appearance in an oblique
direction. Some of these ceased after a short course. In neigh-
bouring regions of the proglottid (text-figs. 75 and 76) there were
patches or vather strand-like parts of the medullary parenchyma

NEW TAPEWORMS FROM THE HYRAX. 587

exactly similar to those which have been described as continuous
with the prolonged oviduct, but not always continuous with it.
They would seem to be produced independently of the generative
ducts. A network arrangement often characterised these regions
of the medullary parenchyma. ‘This condition of the uterus
characterised proglottids in the same stage of evolution as those
which in Thysanotenia gambiana possessed an uterus with a distinct
lumen. Butif weare to compare them with earlier proglottids in
Thysanotenia, then it must be remarked that in the present
species of Jnermicapsifer there is no such regular band of nuclei,
marking out the future uterus, as exists in the former worm.
The appearance is, in fact, totally different. In Jnermicapsifer
the suggestion is merely of a crowding together in a quite
irregular fashion and a local multiplication of the nuclei of the
cells of the medullary parenchyma. In Thysanotenia the nuclei
are in orderly arrangement and regular sequence, obviously
belonging to an organ which, as already pointed out, swells at
its medianly situate end into an oval region of greater diameter.
In Thysanotenia we have obviously a regular outgrowth of the
generative system; in my species of Jnermicapsifer what would
appear to be a condensation of parenchymal tissue in contact
with and continuing on a very short, and in this species hollow,
outgrowth of the generative duct. I am not clear whether the
species described by Janicki agrees most nearly with the species
described in the present paper or with Thysanotenia gambiana.
Tam disposed to think that the branched strand in which the
slightly prolonged oviduct ends in this Jnermicapsifer is not the
homologue of the solid mass of cells in Thysanotenia gambiana,
which afterwards becomes hollowed out to form the uterus
of that worm; in this case Jnermucapsifer hyracis agrees more
closely with Thysanotenia gambiana than with the species
described here.

It might possibly be held that this network, which permeates
the segment when seen in horizontal section, is merely a stage
subsequent to that figured by Janicki in a proglottid of some-
where near the same stage of maturity as that which I am now
considering in the species of /nermicapsifer studied by myself—
that the lumen had, in fact, existed and had disappeared. That
this is not the case is clear from the fact that in this stage there
were not any ova contained in the meshwork, and in fact no
ripe ova anywhere outside of the ovary. The meshwork formed
by condensation of the parenchyma, in fact, precedes the extrusion
of ova from the ovary. Still, it may represent the imperfect
remains of a retiform uterus such as is characteristic of the genus
Dipylidium or, better perhaps, in relation to the present genus,
the Anoplocephalid genus Andrya.

I have also been able to approach the question of the identity
or non-identity of the cavity. in which the eggs lie with the
cavity of the uterus from another side. Text-fig. 77 represents
portions of horizontal sections through a proglottid in which the

39*

588 DR. F. E. BEDDARD ON

eggs are leaving the ovary to be scattered through the medullary
parenchyma, though in this particular stage the whole paren-
chyma is not filled with eggs as it is later and at the end when
the whole medullary region is crammed with the paruterine
organs. As will be seen by a reference to the figures cited, the
eggs lie partly in round or oval cavities which might well be

Text-fig. 77.

Three sections through the medullary tissue in the neighbourhood of the
ovary of Inermicapsifer capensis.

In A a single ovum is shown to the left. In the centre is an oval area with
strongly marked walls, but filled with the ordinary medullary tissue.
B shows eggs (0.) lying each in a single vesicle of the medullary tissue and three
eges lying in a cavity (o.c.).
C. Three of the cavities (0.c.) containing eggs and also showing remains of the
medullary tissue which is shown unaltered in A.

thought to be uterine cavities were no further information
forthcoming. I believe, however, that these cavities are not
remnants of a uterus, as is held by Janicki to be the case with
apparently similar cavities in /nermicapsifer hyracis. A close

NEW TAPEWORMS FROM THE HYRAX. 589

examination of the cavities seems to show that they are lined by
a flattened epithelium, of which the nuclei are apparent and are
represented in the figure, and there is a slight accentuation of
the wall of the cavity, which is merely, as I think, the portion
of the parenchymatous network which abuts upon the cavity and
is not a special layer distinct from that; in this case the ap-
parent lining epithelium should be regarded merely as the nuclei
belonging to this part of the parenchymatous network. I believe
this to be the real explanation; for when we look carefully into
the cavity itself, in which eggs lie in various numbers, there is
nearly always, if not quite always, some granular detritus to be
seen and which is shown in the figures referred to. This formless
detritus (as it often, but not always, is) might, of course, be inter-
preted as simply coagulated fluid which it would not be surprising
to meet with in the interior of a uterus were this system of
cavities the remains of a uterus. But, as will be seen, this
detritus is susceptible of another explanation ; it is, as I think,
the remains of the delicate parenchymal network originally
present, as is shown in text-fig. 77 A, in these regions of the
parenchyma set apart for the development of the eggs. The
figure referred to shows plainly an area oval in section and with
a slightly accentuated wall, marking it off from the surrounding
tissue, which is filled with parenchyma network, but does not as
yet contain any eggs. If my contention is right, then it must
follow that the space containing developing eggs is at least not
always to be referred to a uterus. Nor is it in the least against
this view that it is possible to meet with these circular spaces in
which nothing is apparent but eggs—that is to say, no remains
of the originally present parenchymal network, for this may have
completely disappeared. I would furthermore point out that
the position of the eggs in the nearly empty cavity shown in
text-fig. 77 B suggests that they have only just forced themselves
into the cavity.

I have yet another argument to show reason against regarding
the cavities of the paruterine organs in which the embryos are ~
finally lodged as detached fragments of the pre-existing uterus ;
or, at any rate, to show that they cannot always be so regarded.
In text-fig. 778 and text-fig. 76 is represented an ovum lodged
in one of the meshes of the parenchymatous network of the
medullary region without any special relation to the larger egg-
containing cavities which have been looked upon as detached
fragments of a uterus. There are plenty of such examples to
be seen in sections of this age, and it is plain to me that eggs are
constantly lodged singly in the parenchymal network. I argue
this from the fact that in these cases the cavities lodging the
eggs are in every way indistinguishable in size and appear-
ance from the cavities of the parenchyma in which an egg is
not lodged. It may be, of course, that these are eggs which
got extruded from a uterine cavity and forced into the sur-
rounding parenchymatous network. But.it is equally reasonable

590. DR. F, E. BEDDARD ON

to assume that they have got directly to their situation from the
ovary (see text-fig. 76). The lax parenchymal tissue, the mesh-
work of which is filled with a substance plainly visible as granular
matter after staining with Ehrlich’s hematoxylin, but not to be
seen after staining with borax carmine, can offer little obstacle
to the immigration of eggs; so that in any case some of the
paruterine organs are without vestiges of an uterine cavity.
I believe, as a matter of fact, that all are so, and that there is
no persistent uterus in this worm.

The paruterine organs of this species resemble those of the
species which I described as Thysanotenia*. At the time when
that description was written, I believe that Janicki’s careful
account of the genus Jnermicapsifer with similar egg-capsules
had not actually appeared. J had not realised from the descrip-
tions of Zschokkeella that the organs containing the ripe eggs
were doubtless of the same structure. I had considered that
those organs probably resembled the figure given by Ransom 7,
not entirely grasping the fact that that figure was intended rather
as a diagram to distinguish between those species of Davainea
which had several eggs enclosed in one capsule and those species
in which each capsule had within it but one egg. I was thus
misled, though not through Mr. Ransom’s fault. There was, of
course, no other genus with which I could directly compare
Thysanotenia gambiana.

In defining the genus Zschokkeella, Ransom speaks of the fate
of the uterus in the following words :—‘“ Uterus early breaks
down into egg-capsules.” Earlier in his résumé, Ransom defines
the subfamily Linstowine in the same way ; he remarks that the
‘uterus breaks down into egg-capsules.” As the subfamily
Thysanosomine is defined by the presence (inter alia) of numerous
paruterine organs, I thought myself justified in placing my genus,
as I regarded it, in the latter subfamily and marked its affinities
by the use of the generic name Thysanotenia. I was indeed of
opinion that the uterus in Zschokkeella really persisted in separate
pieces, each containing so many eggs. It appeared to me, in fact,
after studying a tapeworm which I have lately described in the
‘Proceedings’ of this Society as Otiditeniat, that the fate of
the uterus in Zschokkeella might be like that of Otiditenia.
No figures are given by Fuhrmann in his account of Zschokkeella
linstowt§ which illustrate this particular point, and the only
reference to the matter is the assertion that the eggs are
surrounded by a “¢ Parenchymhiille.”

Janicki ||, however, is apparently of my earlier opinion ;
for, in distinguishing between the genus Zschokkeella (written,
as originally—Zschokkea) of Fuhrmann and his own genus

* PZ. 8. 1911, p. 1001.

+ Bull. U.S. Nat. Mus. No. 69, 1909, p. 17, fig. 8.
t P.Z.S. 1912, p. 194.

§ Centralbl. f. Bakt. u. Paras. Bd. xxxii. (1902).

|| Loe. cit. p. 893.

NEW TAPEWORMS FROM THE HYRAX. 591

Inermicapsifer, he uses the differences between the egg-capsules
in the two genera. He considers that in Zschokkeella the eggs
lie ‘“einzeln in einfache Bindegewebskapseln,” a difference which
also appeared to me to hold good. Ido not, however, feel confident
about this point, and in view of other points of likeness between
the genera am disposed to compare more nearly the paruterine
organs in the two. In my paper upon Otiditenia just referred
to I have dealt to some extent with the ‘ egg-capsules” of
Davainea, of which the various figures published are not quite
in unison. I find a justification for this in the paruterine organs
of Inermicapsifer capensis, where the appearances vary slightly
among examples which I cannot refer to different species.

To begin with, I prefer the name of paruterine organs for the
structures in question, because they seem to me to be exactly
comparable to structures so named in other tapeworms. They
are, in fact, sacs formed out of the parenchymal tissue, whether
they have or have not ultimately a connection with the uterus.
I should prefer to term the egg-containing spaces in Otiditenia
“egs-sacs” which are formed in a different way, %.¢. by a
breaking up of the uterus. When specimens of this Jnermi-
capsifer were examined alive in salt-solution, the individual
paruterine organs could easily be squeezed out of the body
by crushing it. They were then spherical or egg-shaped and
appeared to be surrounded by a thick, colourless, and almost
structureless membrane. This membrane exhibited only faint
striz in a longitudinal direction, being thus concentric in reference
to the whole body. The interior was filled with quite transparent
spherical embryos, between which were abundant cells with
granular contents.

In stained preparations (text-fig. 78) the outer layer was also
perfectly distinguishable. It was stained more lightly than the
inner by carmine and more deeply by logwood. A figure of the
mature organ in Jnermicapsifer hyracis is given by Janicki *, with
which may be compared my own figure of the same organ in the
species described here. The appearances shown by the organ in
the living condition are not borne out by preserved and stained
material. The outer layer is not fibrous but cellular, as shown
by Janicki and others. This layer is, however, as a rule, quite
distinct from the inner mass of cells immediately surrounding
the embryos. The distinction between the two partly depends,
as already mentioned, upon stains, and is not always obvious.
I do not see in my specimens of Jnermicapsifer capensis so great
a distinction between the outer and inner cells in point of size
and shape as does Dr. Janicki, which is perhaps rather remarkable
in consideration of the very different appearances they present
when living. The cells of both layers are, in fact, rounded and
nucleated, and not greatly different in size and shape. Those
of the inner layer are filled with larger spherules. They are

* Loc. cit. pl. xiv. fig. 28.

592 DR. F. E. BEDDARD ON

separated by a more deeply staining reticulum, which appears to
be similar to that of the general parenchyma. ‘This forms a some-
what thicker layer enclosing the whole organ. The reticulum of
the inner mass of cells is thicker than that of the outer layer, and
the nuclei lying at intervals along its strands are very obvious.
The reticulum also forms a distinct layer separating the outer from
the inner mass of cells. In fact, the whole organ is of an appear-
ance more like that represented by Fuhrmann* for Davainea
than that by Janicki. This is so, at any rate, as regards the
outer layer. The interstitial substance of the inner mass of cells
forming the reticulum is represented by Janicki, but as of much
greater relative extent than I have found it.

Text-fig. 78.

Paruterine organ of Inermicapsifer capensis, showing differentiation between
outer and inner coat.

The species that has just been described cannot, as I think, be
referred to any of those enumerated by Janicki. It differs from
I. hyracis in the form of the scolex, in the fact that the testes
are arranged in two separate masses in each proglottid imstead
of forming a continuous row. From J. interpositus my species
differs also in the arrangement of the testes, and in the fact that
the genital pores of J. inéerpositus are anterior in position and
open into a well-marked cloaca, and also in the fact that the
sexual products are ripe earlier in the body in J. inéerpositus.

Nor can J. seétii be confused with my species. For in the
former the body is very short and consists of not more than
70 proglottids. Moreover, the excretory vessels do not approach

* Rev. Zool. Suisse, iv. (1896).

NEW TAPEWORMS FROM THE HYRAX. 593

so nearly to the apex of the scolex, nor are they so coiled in that
region as in my species. On the other hand, the two species
agree in the posterior position of the genital pores and the
separation of the testes into two masses. There may be a
difference also in the lack of an excretory network in J. sedti,
of the existence of which Janicki is not certain.

There remain certain less-known species referred by Janicki to
the genus Jnermicapsifer with more or less certainty and of whose
characters he gives some account. Of these, “ Zawnia paronar”
cannot be identical with my species, since it possesses hooks ;
“ Tenia spatula” of von Linstow is too imperfectly described to
admit of its definite inclusion in the genus Jnermicapsifer.
It cannot, however, be identical with my species, since the cirrus
sac is apparently much larger (‘Der Cirrusbeutel nimmt 1/7
des Querdurchmessers ein ”) and the testes are scattered through
the greater part of the segment. Tania ghondhorensis of
Klaptocz is very imperfectly known, but a pit upon the scolex
shows that it is not identical with my species.

From J. criticus of Pagenstecher (which is perhaps identical
with another species described below), the present species can
be distinguished by the grouping of the testes into two masses.

I. pagenstechert of Setti differs from my species by its few
proglottids (not more than 80); otherwise it seems to present
more resemblances to my species than any other form except
I. settti by virtue of the posterior position of the genital pores.

To a species termed by Nassonov Anoplocephala hyracis Rud.
var. hepatica, and by Janicki “ Jnermicapsifer spec.?” I shall
recur in considering some worms from the gall-bladder of Procavia
capensis. In the meantime my own species may be thus defined
and named :—

Inermicapsifer capensis, sp. n.

Length about 95 mm., breadth 2mm.; number of proglottids 200.
Scolex wider than the neck. Proglottids at end of body nearly as
long as wide. Genital pores unilateral near to posterior end of -
proglottids, not borne upon a conspicuous projection. Testes in
two separate growps, one on pore side and one on opposite side of
proglottid. Vas deferens forms a network; a large vesicula
seminalis present ; cirrus sac small, filled with slightly coiled and
dilated sperm-duct. Uterus short and not persistent. Many
paruterine organs, each containing several embryos.

Hab. Procavia capensis.

On Species of the Genus Hyracotenia, gen. nov.

Along with the numerous examples of the tapeworm which I
have described here as /nernucapsifer capensis, I found in the gut
of the Hyrax two complete or nearly complete examples of a tape-
worm which has quite a different external appearance, and whose

594 DR. F, E. BEDDARD ON

internal structure is different from that of the former species.
Before examining it anatomically it was, of course, impossible to
say whether or no these worms belonged to the genus referred to,
and even now it is possible that they may be of the same species as
one or more of those which Janicki has—provisionally, at any rate,
—assigned to the genus Jnermicapsifer. They are, however,

Text-fig. 79.

Hyracotenia hyracis. X 2.

clearly not members of that genus, as the following account of
their structure will prove. Nor can I, with any confidence, refer
them to any other known genus. HenceI propose the above name,
which is indicative of their habitat. I shall enquire later as to

NEW TAPEWORMS FROM THE HYRAX. 595

the possibility of their identity with any other species known from
the Hyrax. The two worms belong, as I think, to two different
species, but are referable to the same genus without any doubt.
I shall consider the anatomy of both of them together.

The larger specimen (text-fig. 79) measured about 90 mm. in
length by a greatest diameter of 5-6 mm. The proglottids are
very short in an antero-posterior direction, but, at any rate some
way back in the body, rather thick. If there is a neck present at
all it is very short. The scolex is unarmed and distinctly marked
off from the strobila, but not much wider than the ensuing body.
The latter increases gradually in width up to the widest point,
and towards the end of the body again decreases. Of the smaller
specimen I cannot give such precise details, since, believing at first
that both specimens were of the same species, I investigated this
individual anatomically without making full notes on its external
characters. It was, however, rather shorter and of less breadth,
while the anterior narrower region of the body widened out to
the full dimensions rather sooner than in the larger specimen.

J have investigated the scolex by means of longitudinal sections
only in the smaller specimen last referred to. The four suckers
look directly upwards, their orifice being terminal in such sections.
There is nothing remarkable that I could detect about their
structure. They do not bulge to any extent from the sides of
the scolex, and these sections show that the scolex is hardly, if
indeed at all, wider than the immediately following strobila. ‘The
rudimentary rostellum is merely a hemispherical elevation lying
between the suckers. There is no terminal pit of any kind and
no hooks discoverable. The water vascular tubes extend into the
rostellum. Of the larger specimen I only examined the scolex
without destroying it. It is clear that the structure is the same
in all the points mentioned above, but I am not able to report
upon the water vascular tubes in this region.

The structure of the body-wall (text-fig. 80) also differentiates
these two species from the /nermicapsifer whose anatomy has been
described above. The principal difference lies in the much more.
marked layer of transversely running fibres which bound the
cortical layer internally and the medullary parenchyma externally.
This layer is very much the same—I think exactly the same—in
both of the two individuals of this genus which I refer later to
different species. It is composed of very delicate fibres; but the
layer, as a whole, is rendered more conspicuous by the fact that large
fibres belonging to the longitudinal layer occur between the trans-
versely running fibres. The cortical parenchyma is nearly as thick
as the medullary. The stout longitudinal fibres which run in the
former are to be found in the greatest numbers at about the
middle of the cortical layer, but they occur elsewhere. They are
not massed into large bundles, but two or three are here and there
closely associated. This massing of the longitudinal fibres is not
obvious in the larger specimen.

596 DR. F. E. BEDDARD ON

The excretory vessels consist of the usual dorsal and ventral tubes
running continuously through the strobila. Anteriorly there is
less difference in the calibre of these tubes than posteriorly. The
position in relation to each other also becomes altered. Inimmature
segments (where I have studied the excretory tubes) the rather
smaller dorsal vessel lies obliquely above the ventral vessel to the

Text-fig. 80.

Hyracotenia hyracis.

Transverse section through a proglottid.

dv. Dorsal excretory tube. Z. Row of testes. V. Vitelline gland.
v.v. Ventral excretory tube.

pore side of the same. In maturer proglottids the two vessels are
practically superposed, and the dorsal vessel is at times so minute
as to escape observation. Both of these vessels are connected
with a network of larger and smaller water vascular capillaries
which traverse the medullary region of the body. I have seen
branches of these ending in a testis, and it becomes a matter of

NEW TAPEWORMS FROM THE HYRAX. 597

great interest to enquire if this network is directly connected with
the network of vasa efferentia. But I have at present no further
facts to offer.

The ovary of this tapeworm is unquestionably single. It lies
on the pore side of each proglottid in the neighbourhood of the two
longitudinal water vessels of that side. The ovary is apparently
sometimes quite anterior to the testes, and sometimes is sur-
rounded laterally by them. The vitelline gland is posterior to the
ovary, which is thus the most anterior of the generative organs.

Text-fig. 81. Text-fig. 82.

‘ 4 Vv.

ein

VV.

Sp.d. &-S. Vg.
Text-fig. 81.—Part of a transverse section through a proglottid of Hyracotenia
hyracis, to illustrate the immature ovary.
dv. Dorsal excretory tube. o. Ovary. v.v. Ventral excretory tube.

Text-fig. 82.—Transverse section of one-half of a proglottid of the same species.

dv. Dorsal excretory tube. o. Part of ovary. r.s. Receptaculum seminis.
sp.d. Sperm-duct. ¢. Testes. v.g. Vitelline gland. v.v. Ventral excretory tube.

Its exact position with reference to the two water vascular vessels
is not always identical. It is more dorsal or more ventral, as the
case may be, and is sometimes entirely to the pore side of the
ventral tube of the excretory system, and sometimes entirely to
the median side of that tube ; sometimes it extends to both sides.
In any case it lies more or less between the dorsal and ventral
excretory tubes, The ovary isnot solid and compact, but arranged
in a series of finger-like outgrowths radiating outwards and

598 DR. F. E. BEDDARD ON

dorsally. In young ovaries, such as the one figured in text-
fig. 81, the riper eggs lie in straight lines connected by delicate
' threads with the central mass, and suggestive almost of the
pseudopodia of a Rhizopod with the eggs carried along them,
This radiating appearance of the ovary is retained until the organ
is quite mature, when it consists of a group of sacs with, for the
most part, definite walls, enclosed in which lie the ripe ova. I
refer to this appearance of the ovary in considering the uterus on
a later page. It seems possible that this condition of the ovary
is to be compared to the testes, and that there are really several
separate ovaries, which, however, are more closely adpressed than
are the testes. In any case we have instances, like Stilesia,
where the single ovary consists simply of a single mass of egg-
cells, a condition which is to be compared with one of the sub-
divisions of the ovary in the present species.

The vitelline gland in the immature segments lies exactly
opposite to the ovary, the vagina dividing, as described later, into
two ducts, which end respectively in the ovary and vitelline gland.
It is large and conspicuous in the mature proglottid. The shell-
gland is also quite conspicuous in this tapeworm.

The vagina is wider and with thicker walls for a short space
after its orifice on to the exterior. It then narrows and runs a
very straight course towards the interior of the proglottid. It
then becomes again wider, and opens gradually or abruptly into
a dilated receptaculum seminis, which lies beside the vesicula
seminalis. This region of the female duct is thin-walled. It is
on a level with the ventral water-vessel. From the median end
of the swollen receptaculum arise two tubes, one of them being
the vitelline duct and the other the duct leading to the shell-gland
and to the ovary. These two ducts are very much narrower than
the receptaculum, into which they suddenly open. In immature
proglottids the receptaculum is rather wider than the vagina, and
gradually widens towards the internally situated end, there
diverging into two horns which are respectively the vitelline
duct and the ovarian duct. These ducts are in these immature
segments of hardly less calibre than the end of the receptaculum
into which they open. These and the sperm-duct pass towards
the exterior between the dorsal and ventral water vascular tubes.

The terminal region of the vagina, 7. e. that part nearest to the
external orifice of the tube, has a lining which is very deeply
stained by logwood, and so has the narrow region which imme-
diately ensues ; the rest of the vagina is not deeply stained in this
way. I noticed in the larger of the two specimens which I report
upon in the present communication that the narrow region of the
vagina lying between the terminal part and the portion which
may be termed receptaculum was much shorter than in the other
example. I am not certain, however, whether there may not be
some variation in this matter from segment to segment, an ex-
pansion of the lumen accounting for the different appearance, It

NEW TAPEWORMS FROM THE HYRAX,. 599

is certain, moreover, that the wide internal region of the vagina,
before it divides into vitelline duct and ovarian duct, is susceptible
to variation ; for in some cases it was an abruptly formed spherical
sac, at other times merely a wider tube than the immediately pre-
ceding region of the vagina. I mention later that the end of the
receptaculum contains ripe ova in the mature proglottids. I have
never seen spermatozoa therein.

Uterus.—In both of the two examples of this species the last few
segments became somewhat shorter from side to side than those in
front, and were also longer in the antero-posterior direction. One
would naturally associate these changes in the facies of the pro-
glottids with complete maturity and the existence in those segments
ofembryos. As I find in these segments completely ripe eggs quite
detached from the ovary and associated together in small masses,
I shall assume that the anatomical structure of these proglottids
is that of complete maturity. In this case the present species
differs from those which we have been considering by the entire
absence of numerous paruterine organs like those of Jnermi-
capsifer, etc. Kven were these terminal segments not so fully
mature as I presume them to be, there would be, I should
imagine, at least some preparation for the formation of the
paruterine organs. But there is none.

The ripe eggs were massed into more or less spherical groups
surrounded by a membrane. ‘These were not to be distinguished
from the lobes of the ripe ovary, and I imagine that they were
merely the persistent ovary. In addition to these masses of eggs,
the end of the vagina, 7. e. the dilated portion which I have termed
receptaculum seminis, was found in many cases to be full or nearly
full of ripe eggs unaccompanied by any interstitial cells. This
was not only the case witha receptaculum which was swollen into
a spherical contour at its base, but also in cases where the re-
ceptaculum ended merely as a slightly wider tubular sac. I have
no reason whatever to doubt these facts, as the receptaculum is
quite easy of identification. I am therefore disposed to think
that there is no uterus as a distinct and separate structure, but
that the eggs are partly voided into the receptaculum and partly
remain im situ awaiting the loosening and perhaps disintegration
of the proglottids. It may be that one of the rounded sacs which I
regard as an ovary is really a uterus. Of this I cannot be positive,
especially in view of the very few completely mature proglottids
in both specimens. That both specimens were identical looks as
if the conditions above described are to be regarded as normal.

The éestes extend through a good part of each proglottid, and,
as seen in sagittal sections, there are five or six rows of them
laterally, though not so many in the median region. In transverse
sections they are seen to extend from edge to edge of each segment,
being nowhere interrupted save where they meet with the female
reproductive organ. This row is mostly one deep, but in places two
or even three deep. I counted from 40 to 50 or so of separate testes

600 DR. F. E. BEDDARD ON

in a single complete transverse series. There are thus altogether
two or three hundred of these gonads to each mature proglottid.
In more immature segments the testes did not appear to extend
to the pore side of the longitudinal water-vessels but to stop
short before quite reaching the median side of those vessels. In
a mature segment I found that 44 out of the 50 sections which
displayed it in its entirety were occupied by the testes, which thus
fill up most of the segment, though the proportions were not always
exactly as stated in the above instance. In proglottids from the
other example of this species 3 sections without testes were fol-
lowed by 15 sections showing testes, and these again by 5 without
testes, and thereafter 14 with testes. Itis therefore obviously the
case that the testes occupy a great deal of the segments.

It will be observed that there is no grouping of the testes into
two masses such as I have described in /nermicapsifer capensis.
They lie mainly behind the ovary and vitelline gland, and in some
proglottids the ovary lay rather more distinctly in front of the male
gonads. ‘The testes are more or less spherical or egg-shaped, and
when ripe are seen to be surrounded by a layer of spermatozoa,
which lie therefore, as I take it, in a cavity surrounding the
testis, a ceelomic cavity. I never found the testes of this tape-
worm to be pear-shaped, like those of Jnermicapsifer. Further-
more, the testes, all of them, lie dispersed in quite unaltered
parenchyma. As is very generally the case among the members
of this group, the testes were mature much more anteriorly in the
body to the ovary. It is, indeed, a striking feature of the present
species, and one in which it contrasts, for example, with the
species of /nermicapsifer that has just been described, that the
mature testes occupy so many segments of the body, while the
mature ovaries are so exceedingly limited in the number of seg-
ments in which they are found. The efferent tubules which
collect the sperm form a very definite network (see text-fig. 83),
which is copious and formed often of unequally sized vessels. A
similar network has been described in other tapeworms, for
example in Chapmannia*.

The vas deferens of this species (text-fig. 82, p. 597) is quite dif-
ferent from that of the species which we have already considered.
The reticulate efferent ducts finally find their way into a large sac,
which in the mature segments is stuffed with sperm, and which lies
in the female generative mass alongside of the receptaculum
ovorum. This large vesicula seminalis is flask-shaped, and there-
fore gradually narrows and emerges from the female generative
mass as it passes towards the genital orifice. It is impossible to
draw a hard-and-fast line between the vesicula seminalis and the
sperm-duct proper with which it is continuous, for the gradual
diminution in calibre of the entire tube forbids such a delimitation
The tube pursues a winding course, narrowing gradually and but
slightly ; it never forms an actual coil like the sperm-duct of so

* See Fuhrmann, Swedish Zool. Exped. Egypt, 1909, pt. iii,

NEW TAPEWORMS FROM THE HYRAX. 601

many .other—indeed the majority of —Tetracotylea, but is at most
once or twice bent upon itself. It becomes very narrow for a

Text-fig. 83.

Portion of medullary region of a proglottid of Hyracotenia hyracis, to illustrate
network formed by vasa efferentia (v.d.).

T. Testis. ¢r. Transverse muscular layer. v.v. Ventral excretory tube.

Proc. Zoou. Soc.—1912, No. XL. 40

602 DR. F. E. BEDDARD ON

short distance in front of its opening into the cirrus sac. The
course of the sperm-duct is roughly parallel to that of the vagina,
with which it might be sometimes confused in those cases where
the vagina has not so abrupt a transition into the receptaculum.

Along the course of the sperm-duct, which in ripe segments is
gorged with sperm almost throughout, there lie masses of what
appear to be prostatic cells, similar in the fact of their existence
to those of Znermicapsifer, but different in appearance. In the
present species these cells are of a clear, almost hyaline, ap-
pearance, which is possibly due to the state of their activity at
the time when the worm was killed. In Jnermicapsifer capensis
and in the species which I originally named Thysanotenia gam-
biana, the prostatic cells were darkly staining and granular.
Nevertheless, they appear to be equivalent structures in the two
tapeworms. In sections where the sperm-duct appears in trans-
verse section, these cells present the appearance of a winding duct
cut transversely. This appearance is due to the clear cells
clustered round the actual sperm-duct, which, as already said, is
narrow of calibre close to where it opens into the cirrus sac, and
thus not obvious in such sections. There can be no mistake,
however, in transverse sections of proglottids, where the course
of the sperm-duct is easily to be followed owing to its being filled
with sperm.

In the second and larger individual there are certain definite
differences in the form of the sperm-duct. The tube has no such
great dilatation into a vesicula seminalis, and it is very much more
coiled as it approaches the cirrus sac. It has, in fact, the large
and close coil which is so typical of tapeworms. There is certainly
nothing of the kind in the other individual. The clear cells
already spoken of form a complete layer one cell thick round
the mass of sperm in the sperm-duct, and are therefore, I take it,
simply the epithelial wall of the sperm-duct. As the sperm-duct
was in parts full of sperm, this difference cannot be owing, I
believe, to the different stages of the maturity of the proglottids
in this tapeworm as compared with those already described. It
must, I think, be a specific difference, with which also, it will be
observed, go differences in the position of the ovary and vitelline

land.
: The cirrus sac of this worm is not at all large as in the allied
forms comprised in the genera Jnermicapsifer and Zschokkeella.
It can be seen in sagittal sections to lie straight in front of the
vagina close to the external aperture, and I have not noticed any
genital cloaca. There is certainly nothing of any size, and in one
section the penis was seen to protrude on to the exterior directly
from the cirrus sac without any intermediate and common
chamber. The cirrus sac is oval in form and surrounded, as is
usual, by a strong layer of muscles. I could not see any indications
of a flask shape such as is so common in tapeworms. In the in-
terior of the sac are the usual nuclei belonging, it is to be presumed,

NEW TAPEWORMS FROM THE HYRAX. 603

to delicate muscles which retract the cirrus. The latter was
relatively wide and short and the sperm-duct within the cirrus
sac not coiled. The protruded male copulatory organ reminded
one rather of the penis of Anoplotenia* than of a cirrus, for it
was wider at the free end than just within the cirrus sac.

It is evident that this genus presents many resemblances
to the genera Zschokkeella and Inermicapsifert. It agrees with
those genera in the following assemblage of characters :—The head
is unarmed and the excretory tubules form a plexus, or at least a
coil, at the very extremity of the rostellum, as in the species which
has just been described ; the segments are narrow and the genital
pores are unilateral. The excretory tubules furthermore form a
plexus within the medullary parenchyma throughout the body.
The ovary lies to the pore side of the segments and is distinctly
not a double organ; the vagina dilates into a wide receptaculum
seminis. The cirrus sac, moreover, is small as contrasted with that
of many other tapeworms. On the other hand, there are certain
characters which argue against this placing of the worm whose
anatomy has just been described. These are as follows :—The
sperm-duct in our species is short and almost immediately dilates
into a large and long seminal vesicle, a state of affairs which is not
met with in the species of Jnermicapsifer known at present.
Finally, the network formed by the vasa efferentia is a feature
hitherto unknown in the genus Zschokkeella, though it occurs in
Inermicapsifer. Inasmuch as a reticular disposition of the vasa
efferentia is not necessarily diagnostic of a given genus as far as
we know, for it occurs in Chapmannia lapica and Hymenolepis
reticulata and not in other (at any rate in some other) species of that
genus, this fact alone would not perhaps necessitate the removal
of the present species from the genus Zschokkeella to which other
important characters would appear to assign it. But there is a
negative character which may be of very great importance. In
neither of the two individuals which I have studied was there the
least trace of the formation of the characteristic “ egg-capsules,”
which I prefer for reasons already given to call paruterine
organs.

In all the species of Zschokkeella and Inermicapsifer examined
from this point of view, the formation of these capsules began
perhaps rather far back in the body, but still a long way before
the actual termination. Now both examples of the present genus
in my possession ended posteriorly in a few segments which
were rather longer than those preceding them and at the same
time rather narrower, suggesting, in fact, the end of the body.
They were, moreover, thicker than unripe segments. If this be
not the completely mature end of the body, the worm would be
very exceptional in the deferring of the egg-reservoirs to a point
so very far behind the scolex. Besides, two specimens selected at

* See Beddard, P. Z.S. 1911, p. 1015, text-fig. 216.
+ For the generic distinctions see below, p. 607.

40*

604 DR. F. E. BEDDARD ON

random from the same host would hardly be likely to prove both
abnormal in any way. Finally, we, have in these segments
rounded sacs with ripe eggs, though it must be admitted that
these were eggs and not embryos. Some riper proglottids may,
however, be missing; but even then the commencement of the
paruterine organs would surely be visible. a

Tt is therefore, as I think, impossible to include these worms
in either of the genera with which I have just compared them.
Of the remaining Anoplocephalide (to which family I think
that these worms must be referred) there are only the genera
belonging to the subfamily Anoplocephaline. Of these Cittotenia
and Moniezia need not be considered, since their generative
apparatus is double in each segment. Of the remaining genera
none agree with the two worms under consideration in all of the
following points, viz., uterus at most inconspicuous, cirrus sac
small, genital pores unilateral, ovary to pore side of proglottids,
genital ducts pass between excretory vessels, testes posterior. 1
believe, therefore, that they must be referred to a new genus.

This new genus may be thus defined :—

Hyracotenia.

Scolex unarmed, with four unarmed suckers. Proglottids wide
and very short, a little longer at extreme end of body, but always
much wider than long. Genital pores unilateral, not borne wpon
papille. Cortical parenchyma thick, separated from medullary by
a thin layer of circular fibres. Water vascular tubes four, dorsal
and ventral, the latter larger, connected by a network of capil-
laries. Testes numerous, dorsal in position, lying behind and to
sides of ovary; vasa efferentia forming a network ; sperm-duct
wide and sinuous or coiled ; cirrus sac small; a short blunt wide
penis protrusible. Ovary near water-tubes of pore side, single, in
front of vitelline gland ; dilated receptaculum seminis and very
narrow vagina. Uterus small and sac-hke ; paruterine organs
absent.

Hab. Procavia capensis.

It is not possible for me to distinguish definitely at present
between generic and specific characters. The above embody
characters usually considered in generic definitions. The two
species may be, for the present at least, defined as follows :—

(1) Hyracotznia procavie, sp. n.

Length about 90 mm. ; greatest diameter 5-6 mm. Body attains
its greatest width about 25 mm. from anterior end. Testes very
numerous. Sperm-duct rather dilated posteriorly, much coiled
anteriorly. Ovary ventral, on outer side only or both sides of
ventral vessel ; vitelline glands dorsal. Vagina not greatly dilated
posteriorly.

NEW TAPEWORMS FROM THE HYRAX. 605

(2) Hyracotenia hyracis, sp. n.

Length about 70 mm.,; greatest diameter 4:5 mm. Body attains
its greatest width about 6 mm. from anterior end. Testes less
numerous. Sperm-duct much dilated posteriorly, sinuous but not
coiled anteriorly. Ovary more dorsal, to median side of water-
vessels ; vitelline glands ventral. Vagina usually much dilated
posteriorly.

We now come to the consideration of the question of the
possible identity of either or both of the above species with any of
those enumerated from the Hyrax by Janicki. The only species
of that series that can be considered (if, that is to say, there are
really no paruterine organs in the forms described by myself)
are Tenia (Anoplocephala®?) gondokorensis of Klaptocz*, Tenia
(Anoplocephala) spatula of v. Linstowt, and Anoplocephala hyracis,
var. hepaticaof Nassonow ¢, termed /nermicapsifer spec. ¢ by Janicki.
Of these Klaptocz’s species has a small scolex like the species
described by me, but also an apical depression (?a rudimentary
rostellum) which my species have not. Furthermore, the pro-
glottids appear to be much smaller. In the species of v. Linstow
we find too great a breadth, and the cirrus sac is too large for
comparison with my species. The shape and proportions of
Nassonow’s species are like mine, but the scolex has a conical
process. In all these species the details are insufficient.

There are thus in African animals—chiefly in Mammals (mostly
in the Hyrax (Procavia), but also in Rodents and Lemurs), but
extending to Birds (Wumida ptilorhyncha)—a group of worms
which show at least specific differences, but all of which have the
following characters in common, viz., head unarmed and no neck,
proglottids wider than long and as a rule very much so, genital
pores unilateral, cirrus sac not very large, testes numerous, ovary
not double. To these characters may possibly be added, if we
exclude the species described above as Hyracotenia spp., or are
led by further material to interpret their anatomy differently, the
formation of numerous paruterine organs—or egg-capsules as they .
have been termed by others.

These characters (excluding the fate of the uterus) seem to me
to necessitate the inclusion of this group of worms in the family
Anoplocephalidee.

The various species which agree in the foregoing characters
cannot, however, on these alone be massed into one and the same
genus without further consideration. If we subtract from the
assemblage the species which I have described as Hyracotenia
procavie, and which, as I think, must in any case be withdrawn
from the group, the reasons for uniting the rest under a single
generic name become more striking. For in this case all of the
species possess paruterine organs of the same kind, unless, indeed,

* S.B. Wien. Ak. 1906. + Jen. Zeitschr. Naturw. xxxv. 1901.
{ Arb. Zool. Lab. Univ. Warschau, 1897.

606 ON NEW TAPEWORMS FROM THE HYRAX.

Janicki be right in inferring that Zschokkeella really differs, a
point which I have already gone into above (see p. 590). The
description of a second species of Zschokkeella* from a Cerco-
pithecus does not throw any further light upon this particular
matter. If it were not for the fact that Davainea seems in
some of its species to possess paruterine organs of the same type,
the African worms referred to might well be regarded as all
congeneric, in which case, of course, Zschokkeella would have to be
the name.

Janicki appears to me to be rather hard put to it to separate
his Inermicapsifer from Zschokkeella. 'The differences are certainly
small. As already stated, he relies upon supposed differences in
the paruterine organs of which I am disposed to doubt the
existence. He also mentions the thickness of the muscular walls
in Zschokkeella as compared with /nermicapsifer, and a few other
points which seem to me to be of minor importance and not even
collectively as of generic rank. Janicki’s comparisons are based
chiefly upon his own account of Jnermicapsifer hyracis, which was
the only species investigated by him in a detailed fashion. I do
not think that a further examination of other species referred to
by Janicki will necessarily prove the identity of /nermicapsifer and
Zschokkeella throughout. I would point out that my own account
in the present paper of /. capensis shows some differences between
that species and J. hyracis. ‘These differences are mainly the
posterior position of the genital pore, the existence of a vesicula
seminalis, the complete separation of two groups of testes, and the
presence of a rete mirabile along the course of the sperm-duct.
Finally, the uterus is much more rudimentary in J. capensis than
in I. hyracis. In some of these characters it would appear that
I. setiti agrees with my species and differs from J. hyracis. A
further examination of these species may show that they agree
in other characters not referred to by Janicki in his résumé of
these forms.

I would reserve the generic name /nermicapsifer for these forms
and refer ‘“ /nermicapsifer” hyracis to Zschokkeella. 'There now
remains my genus Zhysanotenia. Of that genus I have described
two species which show many differences of structure. Thysano-
tenia gambiana is, as I now think, undoubtedly to be referred to
Zschokkeella, with which it agrees in all points, if we may assume
that the paruterine bodies are identical in the two. On the other
hand, it will be, as I think, advisable to retain the name Thysano-
tcenia for the second species of the genus (7’. lemuris), which differs
mainly in the following points :—There is no plexus of excretory
tubes and the ventral vessel is very large, the dorsal being appa-
rently absent in mature segments; the ventral vessels are connected
in each segment by the usual transverse trunks; the receptaculum
seminis is quite different from that of the other forms; the uterus

* Z, remota, see v. Linstow, Zeitschr. wiss. Zool. 1xxxu. 1905.

ON THE AUSTRALIAN LUNG-FISH. 607

is more rudimentary than in Zschokkeella. I should distinguish
the various genera thus :—

I. Excretory system forms a network in each segment. Recep-
taculum seminis long and forming end of vagina. Cirrus sac
small.

A. Genital pores median on edge of segment. Testes forming a
continuous row. No vesicula seminalis. Uterus well
developed at firss . . . . . . . . 4schokkeella.

B. Genital pores posterior on edge of segment. Testes in two
groups. Large vesicula seminalis present. Seminal ducts

form a network. Uterus never well developed.
Inermicapsifer.

II. No excretory network. Receptaculum short and globular
along the course of vagina. Cirrus sac rather large.
A. Genital pores on conspicuous papilla. Testes forming

continuous row. Uterus never well developed.
Thysanotenia.

This arrangement is naturally only tentative, since we are at
present in need of more information concerning the majority of
the species already known from the Hyrax and enumerated by
Janicki in the paper which has been so often referred to. There
are also points in the structure of the species referred to the genus
Zschokkeella which demand further investigation.

32. Additional Notes on the Living Specimens of the
Australian Lung-fish ( Ceratodus forsteri) in the Collection
of the Zoological Society of London. By Basurorp
DEan.*

[Received February 9, 1912: Read April 2, 1912.]

(Text-figures 84 & 85.)

INDEX.
Page
Development and Coloration ........................ 608
Regeneration of injured portion ..................... 611

The two specimens of the Australian Lung-fish in the Zoolo-
gical Society’s collection have been living under unchanged
conditions since 1898, 7. ¢. about fourteen years. In this time
they have been observed repeatedly by zoologists, whose interest
in these important and rare batrachian-like fishes has led them,
in several instances, to publish their notes in detail. There
is still, however, much to learn about the habits of these
fishes, and it is to be hoped that the opportunity will be
seized generally to observe the present specimens, especially

* Communicated by the SecRETARY.

608 DR, BASHFORD DEAN ON

since it is fair to assume that they are living under fairly
normal conditions. In this connection it may be mentioned
that the fishes have been subjected to no changes in their
aquarial habitat; indeed, they have remained practically undis-
turbed for over a decade.

Among the notes dealing with these Specie nS are those of
the pr esent writer, published six years ago in the ‘ Proceedings’ ct
the Zoological Society of London (1906, vol. 1. pp. 168-178), a con-
tribution which gave also data about Ceratodus summarized from
earlier literature. In this paper details were recorded regarding
the movements of the fishes, their mode of breathing, both with
gills and lungs, their manner of feeding, their nocturnal activity,
and in general their salamander-like habits.

The writer had again the opportunity of observing these
specimens during June 1911; his supplemental notes are as
follows :—

Colour.—One of the fishes, the larger one, remains notably
darker than the other. This distinction in colour, therefore,
is neither abnormal nor seasonal. Norcan there be vast adaptive
colour changes in Ceratodus, for the reaction to similar sur-
roundings would then be the same in the two fishes. Is the
darker specimen a male ?

So far as could be learned, the fishes have shown no evidence
of sexual activity. The colours have remained constant, and
there have been no signs of the brilliant tones noted by Schmeltz
(1876, J. Mus. Godeffr., vol. viii. p. 138). According to his
account, the ventral side of Ceratodus is of a deep orange-red,
and several scales on the sides are margined with red;
nothing is said, however, of the relation of these colours to the
season, From the characters of the present specimens, and by
analogy with Ama, we can safely conclude that the tones of
orange and red appear only at the time of spawning. In Ama,
aquarium-kept fishes show no bright colours, but under native
conditions the male fish develops wonderful brilliancy; the spot
at the base of the tail is conspicuous, red scale-margins appear,
and the hinder abdomen glows with tones of orange.

It was noted (1911) that the paired fins were margined with
a narrow white: band. Could this have been an indication of
a breeding colour? No coloration of this kind was seen on the
edges of the unpaired fins.

Size—At present the darker specimen measures 334 inches,
the lighter 293 inches, having grown but a very few inches during
the past seven years. They have reached, accordingly, nearly
their greatest length, Macleay leading us to infer that 3 feet is
about their maximum (Cat. Austr. Fishes, p. 284). Exception-
ally, however, a specimen may measure 45 inches, such a case
having been cited by O'Connor (1897).

Age.—The present specimens give us an idea of the age to which
Ceratodus may attain. We have in the first place data that they
have grown, broadly speaking, at the rate of three quarters of an

Tht

i ae

se

HE AUSTRALIAN LUNG-FISH. 609

inch annually for the last thirteen years. At this rate the present
fishes are over forty years old, and a fish of the record size
(45 inches) would be over sixty. But this assumes that the rate
of growth in length is approximately uniform in fishes of different
ages, In point of fact it is known that fishes in all groups grow
quickly when young, and slowly, if at all, when old: it is also
known that under favourable conditions a fish may grow with
far greater rapidity. In the case of Ceratodus the rate of growth
of the young is as follows :—

Length of specimen. Age.

inches
8 e i ‘Notes from stages
ard eet areicte eee At hatching. wales Teter
te sh he sa Seer Pe ; month. aquaria in Gayndah,
“9 2 anenth Queensland, by Mr.
Die Peeieterocras Sess eke teeth 4 . Thomas Illidge.)
dee legen een er 2 months.

TL Reet area atte Stari 21 months.

IF aera abe cele alse 6 months.

Teh i jawaasdeeueeeeae 7 months.
3 1 x

heli al eres ae eens 74 months.
en Siete te Ree 8 months.

In stages lately hatched the growth is seen to be rapid, its
rate suggesting that a specimen 12 inches in length might be
not older than a year. The rate, however, changes notably when
the young fish no longer subsists on its yolk. In fact, for a
period of about two months it actually decreases in size, a state
of affairs, however, probably abnormal and due to the lack of
proper food in the aquarium. Young an inch long are nine
weeks old; young two inches long are over seven months old.
At the end of the first year the young Ceratodus measures, we
infer, about five inches, a rate of growth which would be not
unlike that in young Ama, gar-pikes, or in a number of teleostean
fishes. If the analogy with known ganoids continues, a two-
years’ Ceratodus would measure 8-9 inches; and a specimen
25 inches in length, approximately the size of one of the present
fishes when it appeared in London, is estimated to be from eight
to ten years old; this added to the thirteen years of their
living in the Society’s aquarium, makes the total age of the
present specimens between, roundly, twenty and twenty-five
years. Older and larger specimens it is fair to credit with
great age, probably fifty years.

Breathing.—The aquatic respiration of Ceratodus varies con-
siderably according to the temperature of the water, but its range
has not been observed. In September (1904), on a cool day,
with water temperature not far from 65° Fahrenheit, the respi-
ratory movements were “slow and regular ; the opercular cavity
filled and emptied about twelve times a minute.” In late
June, when the water temperature was nearly 75°, the successive

610 DR. BASHFORD DEAN ON

movements of the gills varied from twenty-two to thirty-one
aminute. Both fish had long been quiet; if their movements
had been active there is no doubt that this rate would have been
notably exceeded.

Rhythmic movements in breathing are well shown in the
opercular membrane. In the early stage of breathing the cheek
in the subopercular region is seen to dilate slowly and strongly ;
this dilated region is then passed (rather slowly) backward, and

Text-fig. 84.

Opercular movement in breathing of Ceratodus forsteri.

The crest of the undulatory wave is indicated by the asterisk.

its enclosed water is discharged (text-fig. 84). During this
process the dilated part becomes more and more conspicuous
until the final discharge, and at that moment the free rim of the ~
gill-opening is thrown outward and drawn forward, exposing the
lighter coloured hinder border of the gill-slit. The free border up
to that time has been closely apposed to the head.

It was earlier noted that Ceratodus is a “ nostril breather.”

THE AUSTRALIAN LUNG-FISH. 611

The mouth itself showed no noticeable movement of opening
or closing ; it was indeed hardly open, the gape scarcely more than
3 inch. During the later observations, the mouth was seen
slightly to open and close; its maximum gape noted (measured
close to the glass) was 7 inch, its minimum + inch. The opening
of the mouth was here doubtless correlated with the higher
temperature of the water and the more rapid respiratory
movements. In general, however, the mouth margin was
almost motionless, the fish breathing through the nostrils.

In the matter of breathing air at the surface, Ceratodus shows
greater variability than earlier noted. On one occasion over
seventy minutes elapsed without either fish coming to the surface.

Feeding.—Little was added to the former notes. The only
detail suggests that minute food, in the form of alge together
with vegetable débris, forms a part of the normal diet. It was
observed that the fishes would ‘ nose” about in corners and suck
in this finer material. In the process little pebbles would some-
times be taken in and retained for afew moments, then rejected—
the process several times repeated, in a fashion which suggested
that the stones thus mouthed yielded food sought for by the
fish. It was noted in this connection that the stones in the
aquarium were in many cases well coloured with alge.

Text-fig. 85.

Pectoral fin of Ceratodus forsteri, showing regenerating margin.

Movements.—The writer’s preceding paper gave a number of
figures showing characteristic movements of Ceratodus. The
only addition to this series would be a variant of the fig. 11
there given; the right hand pectoral: remained in its resting
position (as in fig. 14); the left, however, was suddenly twitched
up over the back several times (as in fig. 11), but in this case
brushed close to the body, giving the observer the amusing
impression that the fish was thoughtfully rubbing its head. The
habit was curiously unlike that of a fish; it suggested rather
the movement of a tetrapod; and a very similar movement is
known in urodeles.

Regeneration.—The larger (dark coloured) specimen had suffered
an injury to the left ventral fin; a portion of it near the tip

612 MR. C.. H. O'DONOGHUE ON THE

had been lost and the margin was regenerating. Text-fig. 85 illus-
trates how far this process had extended. It will be seen that
the restitutive proliferations were most active along the free distal
margin of the fin. . Here several (four) eminences were present, each
suggesting the pointed tip of the fin; there can, however, be no
doubt as to which of these 7s the terminal one, since the skeleton
of the fin can be followed into the lowest of these lappets. The
case 1s evidently akin to one known to teratologists, for when
certain areas in injured limbs of batrachians are stimulated, there
appears polydactyly or polypody. It may therefore be worthy of
record that a similar condition occurs in the lung-fish Ceratodus.

33. The Circulatory System of the Common Grass-Snake
(Tropidonotus natriz). By CuHas. H. O’Donoeuus,
B.Sc., F.Z.8., Assistant to the Jodrell Professor of
Zoology, University College, London.

[Received April 1, 1912: Read April 23, 1912.]
(Plates LX X.-LXXII. and Text-figures 86-91.)

INDEX.
Page
Dt, Mntrodwetion::.3.. ce. cassorea he chee meee Ue eet reo GTO,
II. The Heart.
A Developmient yy: omc cac. cation ee sek esa ete OLS)
Be Adu baloranlca ha scucs aes aan tee ok Uae Pee ane 7
III. The Arterial System.
Ao Developmierit 4-0: epetawee ite sai acco ete Veale GAO,
Bs ACUIRG AHO ene, dks Stat Ov an Mee Aue AGO
(@)PAntertommViesselsiaee eee Oo
(5) Posterior Vessels .......00 0. cee cecceeceeeeecee eee eeeeee 623
IV. The Venous System.
AL Development) .ise.eaia toe, a aeRO ee COU EMuE NG DG
Bec Adult Mornay, ne. pamet tee: ee alee SES ()
(@) A\TM@TIO? WORE 255 connas aoc apeocecse coo noo cos nonocoone 630
(6) Posterior Vessels .....0....00.c0cccceeeeceeeeeeeeecee ees 681
V. The Vessels of the Head.
tA SATUCTIES) ta reex shee: sav er statin adenh ee ak ten, AUPE, Ole GSS
IBS Meimisiiy oh. ctasac eas tackndn sarge belts dee ope Saoeenti ma I Me GAD
ME. sihistiof References te ate eG
WET Explanaivon) of) iE latesip acres trae ann en eee nent

I. InrRoDUCTION.

Our knowledge of the circulatory system in snakes is far from
exhaustive ; indeed, we have only a complete account of the
vessels in the Python by Hopkinson and Pancoat (25), and a
later and a more full one by Jaquart (26), and in Pelophilus
madagascariensis by Gadow*. Although Zropidonotus natrix is

* This is incorporated in the account given by Hoffmann (23).

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CIRCULATORY SYSTEM OF THE GRASS SNAKE.

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CIRCULATORY SYSTEM OF THE GRASS SNAKE.

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CIRCULATORY SYSTEM OF THE GRASS SNAKE.

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 613

the snake most commonly dissected in the laboratories of this
country, no description of its vascular system has appeared’ since
the anatomical account of the blood-vessels in snakes written by
Schlemm (35) in 1827, which was based largely on Coluber
(i. e. Tropidonotus) natrix and Trigonocephalus mutus. This
account by Schlemm, although excellent in many respects, is by
no means complete, and, owing to the overlooking of the cerebral
carotid artery, the remaining arteries. of the head are misinter-
preted. A great deal of work has been done, however, on
different parts of the circulatory system of this animal by various
authors. Weare indebted to Rathke for a valuable account of
its development (30) and also of the arteries of the head and
neck (31); the last is the best general account of these vessels in
snakes that has been written as yet. Hochstetter has dealt with
the development of the posterior veins (20) and of the blood-
vessels in general (22), Grosser and Brezina (19) with the
development of the veins in the head and neck, and Bruner (12)
with the veins and sinuses in the head of the adult.

In addition to these works bearing directly on 7. natrix,
Beddard (1-6) has added considerably to our knowledge of the
blood-vessels of other snakes, and the intracranial circulation has
been dealt with in the vertebrate series in general by De Vriese
(14) and Hofmann (24) and in reptiles by Dendy (18).

The blood-vessels of the Grass-Snake were investigated by
means of the dissection of a number of injected specimens. The
injection fluid used for the main vessels was that recommended
by Kingsley (28), 2. e.,

Corn starch and 2 per cent. chloral hydrate (each) 400 vols.
95 per cent. alcohol 100 vols. and Colour and glycerine (equal
parts) 100 vols.

For the finer vessels a gelatine mixture advised by Tandler (36)
was used, 2. é.,

5 gms. of gelatine in 100 c.c. of distilled water coloured with
Berlin blue or carmine.

5-6 gms. of potassium iodide added slowly while warming
gently.

These are two very good mixtures, as the first will keep almost
indefinitely, and the second, with the addition of a few crystals
of thymol, will keep in a stoppered bottle for months, and, in
addition to being useable when almost cold, will withstand acids.
By mixing a quantity of the gelatine mass with about one-third
or less of its volume of the solid residue that settles to the
bottom of the starch mixture, an extremely useful general in-
jection mass is obtained. The latter mass, which flows very
readily if only slightly warmed, and sets firmly and fairly quickly
in 70 per cent. aleohol or in 4-5 per cent. formalin, was the one
most frequently employed in making the preparations for this
investigation, ;

614 MR. C. H. O DONOGHUE ON THE

For the sake of clearness in description, the account of the
blood-vessels of the head is not included in the general description
of the vascular system of the whole animal, but is dealt with
separately later. A brief account of the development of the
heart and of the arterial and venous systems has been introduced
in order to throw some light on the condition that obtains in the
adult.

Two or three features of general interest in connection with
the elongation of the body and the loss of limbs in the Ophidia
are clearly brought out in dealing with the vascular system of the
Grass-Snake. The first is the marked asymmetry of the viscera
and their blood-supply ; not only are the organs of the right side
anterior to those of the left, but they are also considerably larger.
Thus the right ovary, supra-renal body, and the kidney are in
front of and larger than the corresponding organs on the left,
and, as is well known, in the case of the lungs the left one is
entirely suppressed.

Secondly, the tendency to form longittdinal systems of vessels,
common to all Ophidia, as Beddard (1) pointed out, is well
marked. The various arteries supplying the intestine and the
fat-bodies are in each instance indirectly connected into one long
system. The ovarian artery forms a longitudinal trunk along the
corresponding supra-renal body. Among the veins also we find
that the hepatic portal vein runs from one end of the intestine
to the other, and that each oviduct possesses a sinus running
beside it for the greater part of its length. This oviducal sinus
is very conspicuous in 7’, natria, although it does not appear to
have been described previously in other snakes. The liver, too,
is greatly elongated, and the post-caval vein and the hepatic
portal vein pass along its opposite faces from one end to the
other.

Lastly, the blood-vessels of the adult, with the exception of a
small pair of veins in the cloacal region, which may represent the
pelvic veins of Lacertilia, give no indication of their derivation
from those of a limb-bearing ancestor.

In conclusion I should like to express my sincere thanks to
Professor J. P. Hill of this College for the kindly assistance and
advice he has given me throughout the work.

II. Tue Heart. (Pl. LXX.)

(A) Development.

The development of the heart has been very fully dealt with
by Rathke (30), and as this account differs but slightly from that
of Lacerta given by Greil (18) and Hochstetter (20), it does not
appear necessary to give more than a brief outline here.

The primitive heart is in the form of a simple tube stretching
in an antero-posterior direction in the region of the gill-slits.
Its posterior end is formed by the union of the two omphalo-

CIRCULATORY SYSTEM OF THE GRASS-SNAKE, 615

mesenteric veins, and the anterior is continued as the short
common stem (truncus arteriosus) of the first pair of branchial
arches. It soon bends towards the right, and as the two ends
remain in approximately the same position, while the tube itself
grows longer, it is forced to take on a curved form. In this
twisted condition three portions can be distinguished: first, a
posterior part running from the union of the omphalo-mesenteric
veins, close to which now open the paired ductus cuvieri,
ventrally and towards the left; secondly, a median part situated
ventrally and running obliquely from left to right; and thirdly,
an anterior part running from the right to the median line, where
it bends sharply dorsally before passing over into the truncus
arteriosus, from which, by this time, two pairs of branchial arches
are given off.

Grooves appear on the posterior part of the tube which indicate
the divisions between sinus venosus and atrium and between
atrium and ventricle. Into the sinus venosus now open the
paired umbilical veins. The middle part, afterwards to become
the ventricle, becomes dilated ventrally, and as the anterior part
also dilates, the two parts are separated by a deep furrow.
According to Hochstetter (20) and Langer (29) this anterior
part is homologous with the bulbus cordis of the Batrachia.

The ventricle expands still further and moves caudally, so that
the atrium, which has also become dilated, comes to le close to
the bulbus cordis. The further dilatation of the atrium takes
place cranially and towards the left, causing it to take up a
position on the left side of the bulbus. The constriction between
the atrium and ventricle, corresponding to the auricular canal,
becomes mere marked as these two structures swell out. At this
stage, too, the sinus venosus is sharply constricted off from the
atrium and the truncus arteriosus gives off the six pairs of
branchialarches. Rathke erroneously described only five branchial
arches, but this will be referred to again later.

After this the ventricle gradually assumes its adult shape
The base of the bulbus cordis, originally joining the ventricle on
the left, moves into an almost mid-ventral position. Spirally
twisted grooves appear between the branchial arches, now reduced
to three in number, and extend downwards over the bulbus.
The atrium now gives off another dilatation, but this time to the
right, and consequently the truncus arteriosus and bulbus cordis
lie in a deep groove between the outgrowths of the atrium.
These two dilatations are the definitive auricles, and already
the sinus venosus, which lies in the atrio-ventricular sulcus on the
dorsal side of the heart, opens into the one on the right.

The groove between the bulbus and the ventricle gradually
disappears, and ultimately the proximal part of the bulbus
becomes incorporated with the latter, while its distal portion
becomes assimilated to the truncus arteriosus. The spiral
grooves finally extend over the whole of the so-formed truncus,
and they indicate its internal division into three parts by the

616 MR, C. H. O'DONOGHUE ON THE

backward growth of two septa. One, the septum aortico-
pulmonale, arises from the edge of the pulmonary artery, and as
it grows it divides the truncus cavity into two tubes, an aortic and
a pulmonary. The other, the septum aorticum, arises between
the two aortic arches, and so subdivides the aortic cavity of
the truncus into two, a right and a left. Of the three tubes
formed in this way, one lies to the left, ventrally, and leads to the
pulmonary arch ; another lies to the right, ventrally, and leads
to the left aortic arch ; and the third lies dorsally and leads to
the right aortic arch. ~ .

Text-fig. 86.

Ventral view. of the heart and adjoining vessels.

Az. Azygos vein. C.A. Left common carotid artery. L.A. Left auricle.
L.C.A. Left coronary artery. L.J. Left common jugular vein. L.S. Left systemic
arch. P.A. Pulmonary artery. P.C. Primary carotid. P.V. Pulmonary vein.
P.V.C. Post-caval vein. R.A. Right auricle. R.C.A. Right coronary artery.
R.J. Right common jugular vein. R.S. Right systemic arch. Th.A. Thyroid
artery. V. Ventricle. V.A. Vertebral artery.

Lastly, the sinus venosus also assumes its definitive form.
After the disappearance of the umbilical and omphalo-mesenteric
veins, it has opening into it, on the right, the right ductus

Cuvieri and the post-caval, while on the left it has the left ductus -

Cuvieri. Thus we have practically the adult condition of the

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 617

heart, which, however, in the course of the further development
moves caudally, and ultimately comes to lie a long way from its
primitive position.

Text-fig. 87.
Ro
CA
(ai V. A.
ThA.
RC. Ras
Ng
LA face
LPC ae
rt PA.
V1.
(ei
(CAVE
be
Pv RV.C.

Dorsal view of the heart and adjoining vessels.

C.V. Coronary vein. U.P.C. Left pre-caval vein (left common jugular).
R.P.C. Right pre-caval vem (right common jugular). §.V. Major part of sinus
venosus. §.V.1. Minor part of sus venosus. Other letters as in text-fig. 86.

(B) Adult Form,

Tropidonotus in common with all the reptiles, except the
Crocodilia, possesses a three-chambered heart. This is situated a
considerable distance behind the head and slightly towards the
right. It is enclosed in a pericardium in which it lies freely,
not being attached to it by a gubernaculum cordis as is the case
in the heart of the Lacertilia and Crocodilia, Beddard (2) has
pointed out that although a gubernaculum cordis is generally
absent in snakes, it is not completely so, as a homologous structure

oceurs in some species*, The pericardium on the right side lies

* EH. g. Coronella getula, Coelopeltis monspessulana, and Ophiophagus bungarus,
Beddard (loc. cit.).

Proc. Zoou. Soc.—1912, No. X LI. 4]

618 MR. C. H. O DONOGHUE ON THE

against the body-wall, while on the left side it is separated from
it by the intervention of the cesophagus. In conjunction with
the elongated form of the body, we find that the heart also is
long and narrow.

The Sinws Venosus is situated on the dorsal surface of the
heart, and appears as a saccular structure divided into two parts
and formed by the swollen extremities of the common jugular
veins and the post-caval veins (text-fig. 87). It is hardly dis-
tinguishable externally from the right auricle, although internally
the two cavities are separated by the bicuspid sinu-auricular valve.
The right common jugular vein from the anterior end of the body
and the post-caval from the posterior end join together to form
the major part of the sinus venosus, which lies on the right of
the dorsal surface of the heart. The left common jugular vein
runs down along the outer edge of the left auricle and then across
in the groove between the left auricle and ventricle (text-fig - 87).
Tts mouth opens into the smaller part of the sinus venosus *
which is partially separated from the major part by a valvular
septum. The efficacy of this septum is seen when injecting, for
while the right common jugular and post-caval veins may be easily
injectel from the major part of the sinus, it is almost impossible
to inject the left common jugular from it.

The thin-walled Awricles (text-fig. 86) are unequal in size, the
right, of an elongated oval form, being much larger than the left,
which is shorter and more rectangular. Into the right auricle
opens the sinus venosus and into the left the single pulmonary
vein (text-fig. 87). The opening of the pulmonary vein is not
guarded by a valve as Fritsch (16) pointed out, but it seems
highly probable, as Sabatier (33) suggested, that during systole
a fold of the auricle in this region “functions as a valve and so
prevents regurgitation. The auricles are completely separated
by an imperforate inter-auricular septum which is continued
caudally so as to divide the auriculo-ventricular aperture into two.
The internal surfaces of the auricles possess a network of raised
muscular ridges, the musculi pectinatz.

The Ventricle is somewhat oval in shape, but very asymmetrical.
The posterior end forms a bluntly conical apex, and the base,
although more or less transverse on the right side, is produced
anteriorly on the left side into a conical process, so that the left
side of the ventricle is nearly as long again as the right. It is
extremely thick-walled, and its cavity contains a large number of
muscular trabeculee, some of which interlace in such a way as to
form an oblique, incomplete ventricular septum. ‘This partial
septum keeps the aerated blood brought in by the left auricle
more or less completely separated from the non-aerated blood
from the right auricle. Two valves, a right and a left, similar in

arrangement to those in Lacertilia, guard the auriculo-ventricular
apertures.

* They do not open separately into the auricle as stated in Rolleston (32).

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 619

The Bulbus cordis, as has been pointed out above, is not to be
found as a separate structure in the adult, and so the three aortic
arches arise directly from the ventricle. The base of each of
these is guarded by two semilunar valves, which Langer (29) has
shown to be homologous with the distal row of valves in the
amphibian heart.

III. Tar Arteria System. (Pl. LXX.)
(A) Development.

The development of the aortic arches in Z’ropidonotus is very
similar to that of other Reptilia*. It was first described by
Rathke (30), whose general account has been confirmed since by
Van Bemmelen (7 & 8) except in one particular. Rathke

Text-fig. 88.

CEO TTT
i

LN : Apa

ea

DA:

Diagram of embryonic arterial arches. It represents the condition after the
disappearance of arches 1, 2, and 5, and shows also the division of the truncus
arteriosus into three arterial roots.

C.A. Common carotid. I).A. Dorsal aorta. E.C. External carotid. I.C. Internal
carotid. -L.S.A. Left systemic arch. P.A. Pulmonary artery. P.Ar. Pulmonary
arch. R.S.A. Right systemic arch.

Adapted from Hochstetter (22).

describes the development of only five visceral arches on each
side, which he numbered from the anterior end 1-5. Van Bem-
melen, however, showed that there was another arch, which has
however, a somewhat transient existence, between arches 4 and 5
of Rathke, so that the latter’s fifth arch is in reality the sixth of
the series, and thus the snake is brought into line with other
Amniota.
* For a general account of this see Hochstetter cai iS

620 MR. C. H. O'DONOGHUE ON THE

These arches soon become reduced to three on each side, viz.
3, 4, and 6, by the disappearance of arches 1, 2,and 5. Of the
remaining arches, 3 is the carotid, 4 the systemic, and 6 the
pulmonary. By the separation of the truncus arteriosus into
three tubes the two carotids and the right systemic have a common
opening into the ventricle; the left systemic opens separately,
and the two pulmonaries open by a common vessel (text-fig. 88).
The most remarkable change in the development is the enormous
lengthening of the carotids, brought about partly by the elonga-
tion of the neck but largely by the caudal shifting of the heart.
Thus it happens that in the adult condition the 3rd arch is far
removed from the 4th and 6th arches.

Text-fig. 89.

Diagram to show changes in embryonic arterial arches. It shows the change
from the condition in text-fig. 88 to the definitive state. The filled-in portions
represent the vessels left in the adult, and those indicated by dotted lines the
vessels that disappear in the course of development.

A.B. Basilar artery. C.C. Left common carotid. P.C. Primary earotid.
Th.A. Thyroid. Other letters as in text-fig. 88.
Adapted from Hochstetter (22).

The Carotid Arch goes through considerable changes in the
course of its development. The two common carotids arise from
the systemic by one root, the primary carotid (carotis primaria

CIRCULATORY SYSTEM OF THE GRASS-SNAKE, 621

of Rathke), which remains short as in most snakes*. After the
common carotids have split into internal and external carotids,
each internal vessel gives off a branch which enters the neural
canal with the first spinal nerve and joins the basilar artery on
the ventral side of the nerve-cord. These branches quickly
widen out and, forming as they do an anastomosis between the
two internal carotids, make it possible for the right common
carotid to degenerate. This it does completely + from the point
where it divides into internal and external branches down to
close to its union with the left common carotid, but the last part
of it remains and is to be found in the adult as a small artery
supplying the Thyroid gland (text-fig. 89).

The Systemic Arch undergoes very little change during
development.

The Pulmonary Arch degenerates almost completely on the
left side. In conjunction with the suppression of the left lung
in T’ropidonotus, we find that the pulmonary branch of the 6th
arch is only developed on the right side. In the adult only one
pulmonary artery is to be found.

(B) Adult Form.

(a) Anterior Vessels.

The Left Aorta bends dorsally around the csophagus and
trachea and then posteriorly to unite with the right aorta in the
mid-dorsal line. During this course it gives off two very small
branches to the cesophagus, but none whatever to the parietes.

The Right Aorta takes a corresponding course on the other side,
during which it gives off the following branches :—-

J. The Left and Right Coronary Arteries arise behind the two
semi-lunar valves which guard the base of the aorta. The right
coronary artery runs in the groove between the auricle and
ventricle, and is the chief supply of the dorsal surface of the
heart. The left passes around the base of the pulmonary artery
and spreads out over the ventral side of the heart.

* Tn some snakes it is absent altogether, so that the common carotids come off
separately from the systemic arch, e.g. Boa.

+ It is interesting to note, however, that in some variations of 7. natrix this
does not occur, and so the two common carotids persist in the adult. The first
specimen of 7’. natrix that I examined was in this condition, although in all other
respects it appeared a perfectly normal adult male. The two common carotids, left
and right, sprang from a common stem, the primary carotid, and were about equal
in calibre. On the left side the carotid pursued a normal course. The abnormal
right carotid passed ventral to the cesophagus, just behind the thyroid gland, to
which it sent a small branch, over to the right side of the neck. From here up to
the posterior end of the skull it followed a similar course to its fellow on the left.
Unfortunately the vessels of this specimen were not injected, so that the relation of
the persistent right carotid to the basilar artery could not be ascertained. However,
this apparent anomaly in the arterial system is quite readily understood in the light
of the developmental history of these vessels. Only one other example of this
peculiar abnormality seems to have been described before, and that by Van Bemmelen
(Q), but in this case the right carotid was only a fine tube.

622 MR. C. H. O'DONOGHUE ON THE

II. The Primary Carotid (Carotis primaria, Rathke) is a short
trunk which quickly divides into two unequal branches :—

i. The Thyroid Artery is the smaller of the two, and, in addition
to supplying the thyroid gland, it sends a twig to the
right thymus glands. This is the sole remnant of the
right common carotid.

ii. The Lefé Common Carotid (Arteria carotis communis,
Rathke, Arteria cephalica, Schlemm) runs along the left
side of the esophagus and trachea, to which it sends three
or four slender branches, until it reaches the posterior
region of the head. Here it divides into internal and
external branches, and supplies the whole of both sides of
the head. (The distribution of this vessel in the head will
be dealt with later.)

IM. The Vertebral Artery (Arteria vertebralis, Rathke, Arteria
collaris, Schlemm) arises from the anterior dorsal part of the right
carotid arch and runs forward, a little to the right of the vertebral
column about half-way to the head. It gives from three to
seven branches to the parietes and one or more to the cesophagus
before disappearing into the vertebral musculature in the mid-
dorsal line.

IV. Five Parietal Arteries are then given off. The first three
are very slender and close together, while the remaining two are
of the same size as the regular parietal arteries.

V. One or two small Wsophageal Arteries run to the cesophagus.
After this the two systemic arches unite to form a single
vessel, the dorsal aorta.

The Left Ductus Botalli is not completely closed up in the
course of development, and its proximal portion is to be found in
the adult animal as a cul-de-sac running cranially from a point
low down on the root of the right carotid arch*. This remnant
varies somewhat in size in different individuals, and although
it is always more or less short, is of nearly the same calibre as
the right carotid. It is completely hidden by the left auricle, but
is readily seen if that body be carefully removed. From its
somewhat bluntly conical end comes off a thin strand of tissue
which runs forward into the left systemic arch at the pomt where
it bends over to run backwards. This represents the closed part
of the left pulmonary arch, and is therefore the left ligamentum
Botalli, such as has been described by Brenner (11) and Hoch-
stetter (21). I find myself in agreement with the former author
also when he states that he was unable to find a right igamentum
Botalli in Tropidonotus natria.

According to Brandt (10), quoted also by Hoffman (23), there

* Such a saccular appendage appears to be present generally in those snakes with
one lung suppressed, and has been recorded by Hochstetter (21) in Tropidonotus
natrix, T. tesselatus, Coluber esculapii, Coronella levis, Vipera berus, and
Cerastes vipera.

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 623

is present in the Grass-Snake a solid strand of tissue joining the
primary carotid to the transverse part of the left aortic arch and
called by him the ligamentum caroticum. ‘This, he states, in
exceptional cases may remain open and may then be described as
a ductus caroticum, and is a vestigeal structure somewhat similar
to the ductus Botalli. In the hearts I have examined no trace of
this vessel or cord could be found, and, indeed, no such connection
exists in the course of embryonic development, as a glance at text-
figs. 88 and 89 will show. A connection between the carotid and
left aortic arches is present in the embryo, but in the adult it
would run from the dorsal part of the left systemic arch along the
whole length of the neck up to the point of origin of the internal
carotid (text-fig. 89). This, however, does not fit in with Brandt’s
description of the ligamentum caroticum.

(>) Posterior Vessels.

The Light Pulmonary Artery (Arteria pulmonalis, Schlemm)
arises separately from the ventricle and leaves the heart the most
dorsal of the three arterial roots. It runs backwards alongside
the cesophagus almost parallel with the right systemic over which,
however, it passes ventrally, and then runs dorsal of the post-
caval vein to the anterior end of the lung. As it passes along
the right border of that organ it gradually gets smaller and
smaller until it disappears as a distinct vessel at the level of the
posterior end of the liver, although the lung is continued on for
some distance.

In correlation with the suppression of the left lung no left
pulmonary artery is found at any time.

The Right and Left Aortic Arches unite posterior to the heart to
form the dorsal aorta.

The Dorsal Aorta runs in the body-cavity just ventral to the
vertebral column, back to the level of the cloaca. Just posterior
to this it leaves the body-cavity and enters the hemal canal, and
in this is continued along the tail as the Caudal Artery. During
its course through the ccelom it gives off a number of branches.

The Parietal Arteries form a numerous and more or less regular
series of branches going to the body-wall, of which there are
about twelve up to the point of origin of the superior mesenteric
artery. These arteries enter the body-wall in the mid-dorsal line,
a characteristic of most colubrine snakes, and do not split into
two before so doing, as in the pythonine snakes (cf. Beddard,
4 and 1).

The @sophageal and Hepatic Arteries.—In front of the liver
the dorsal aorta gives off two or three slender branches to the
cesophagus. After these come a series of common trunks, about
fifteen in number, which divide into two branches, one going to
the liver and one to the esophagus or posteriorly to the stomach.
The last of this series is considerably larger than the others and
has more branches, some of which go to the anterior end of the
stomach.

624 MR. C. H. O DONOGHUE ON THE

The following vessels then come off from the dorsal aorta in
order :—

1. The Lzeno-gastric Artery is the first of these. Its gastric
branch is the main artery supplying the stomach, and it also sends
a branch to the spleen, and yet a third, the cystic artery, to the
gall-bladder. No branch of it goes to the pancreas, nor does
the superior mesenteric artery send twigs to the spleen and gall-
bladder as Beddard (1) has described in T’ropidonotus fasciatus.

2. The Superior Mesenteric Artery is the largest vessel arising
from the dorsal aorta. Soon after its origin at about the level
of the pancreas it divides into two branches ; a smaller one, the
duodenal, running anteriorly supplies the part of the intestine
immediately after the pylorus and also the pancreas; a much
larger one running posteriorly supplies the many coils of the
intestine as far back as the posterior end of the right ovary.
Small branches from it also supply the anterior part of the fat-
body, I have been unable to find any branch of this artery
running to the right ovary such as Beddard (1) recorded in
Tropidonotus fasciatus.

3. The Right Ovarian Artery, a moderate-sized vessel, runs to
the right supra-renal body and, dividing into anterior and _ pos-
terior branches, forms a longitudinal vessel along it. From this
longitudinal trunk are given off :—

(a) Six small ovarian arteries of equal size.

(6) Three somewhat larger oviducal arteries. One of these
arises from the anterior end of the longitudinal vessel
and supplies the fimbriated opening of the oviduct, and
the other two arise from the posterior end.

(c) Three fat-body arteries—an anterior, a small median, and
a posterior.

(dz) A number of fine twigs to the supra-renal body.

The posterior of the three fat-body arteries in some cases has
an independent origin from the aorta.

4. The Mirst Inferior Mesenteric Artery supplies the coils of the
intestine just posterior to the right ovary. Its point of origin
varies, however, in different individuals and in the two sexes, as
will be pointed out below.

5. The Anterior Right Renal Artery conveys blood to the
anterior half of the right kidney, and also gives off a branch to
the right oviduct.

6. The Left Ovarian Artery is distributed in a very similar way
to the right, and from the longitudinal trunk it forms along the
supra-renal body come off :—

(a) Five equisized ovarian arteries.

(6) Three oviducal arteries. The anterior again supplies the
oviducal funnel.

(c) Two fat-body arteries.

(d) A number of fine twigs to the supra-renal body.

As on the right side the posterior of the fat-body arteries may
arise separately from the aorta.

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 625

7. The Anterior Left Renal Artery divides soon after its origin
into two branches, one going to the fat-body and the other to
the kidney. This latter branch supplies the anterior half of the
kidney and sends a twig to the oviduct.

8. The Second Inferior Mesenteric Artery arises about the level
of the anterior end of the left kidney, and supplies the intestine
in the region of the median part of the right kidney.

9. The Median Right Renal Artery, in addition to taking blood
to the posterior median part of the kidney, sends a branch to the
right oviduct.

10. The Posterior Right Renal Artery feeds the posterior end of
the kidney and gives off two branches to the oviduct.

ll. The Third Inferior Mesenteric Artery, arising near the level
of the posterior end of the right kidney, supplies the intestine in
the region of the posterior part of the left kidney.

12. The Median Left Renal Artery is distributed very similarly
to the corresponding vessel on the right.

13. The Fourth Inferior Mesenteric Artery, arising at the level
of the posterior end of the left kidney, supplies the last part of
the intestine.

14. The Posterior Left Renal Artery resembles its fellow on the
right, save that it does not send a branch to the oviduct.

15. The Posterior Oviducal Arteries are a pair of arteries
running to the posterior ends of the oviducts. In addition to
this there may be one or two small twigs going to the rectum.

16. The Rectal Artery is a small vessel supplying the last part
of the rectum.

An Epigastric Artery is present, and it appears to be similar to
that described by Beddard (1) in Ophiophagus bungaris, that is,
it runs along the body-wall in the mid-ventral line in close con-
nection with the epigastric vein, and is fed by branches from
the carotid anteriorly and the fat-body posteriorly. Its precise
relations are somewhat difficult to make out, for it is a vessel too
small for individual injection, and in order to get a satisfactory
injection of the arterial system it is necessary to open the snake.
from end to end, dissect away the skin, and free the gut to some
extent. This, however, necessitates cutting either the epigastric
artery itself or some of its smail tributaries.

The arrangement of the vessels in the male snake is very
similar to that just described for the female. All the arteries
anterior to and just posterior to the heart are precisely the same,
and it is not until the region of the urino-genital organs is
reached that we find any difference.

Each testis receives one spermatic artery as is general in snakes,
which first runs to the supra-renal body, whence it sends branches
to the testis and also to the vas deferens. These two spermatic
arteries are each followed by another vessel that runs to the pos-
terior part of the supra-renal body and also supplies the vas
deferens for a considerable distance, that on the right side also
sends a branch to the fat-body.

626 MR. C. H. O DONOGHUE ON THE

The kidneys each possess an anterior and a median renal artery
as before, but instead of one posterior trunk there are at least
two, but most often three *. These vessels also send small twigs
to their respective vasa deferentia, and the left anterior renal
supplies the fat-body with a large artery.

The supply to the alimentary canal is also slightly different.
The first of the inferior mesenteric arteries arises posterior to the
anterior left renal artery, whereas in the female it is anterior to
the anterior right renal +. This is followed by a series of about
four smaller inferior mesenteric arteries. The various mesenteric
arteries are connected by their small branches and so form more
or less of a longitudinal system along the gut.

The origin of the arteries supplying the fat-bodies in both sexes
is fairly similar and is somewhat interesting. The anterior part
is supplied by branches from the superior mesenteric artery, the
next portion by vessels from the right genital artery, and the
posterior end is fed by branches from the left genital and left
renal supply. In addition to which the posterior branch from the
genital artery on each side may in some cases arise independently
from the dorsal aorta. All these branches are joined one to
the other by small twigs into a longitudinal system running the
whole length of the fat-body, but there does not appear to be one
unbroken artery, an arteria epiploica, traversing the whole length

of the fat-body.

IV. Tue Venous System. (Pl). LXX.)
(A) Development.

The general course of the development of the venous system
in 7ropidonotus is similar to that of other reptiles (vide Hoch-
stetter, 22), but it has some points peculiar to itself (vide Rathke,
30, und Hochstetter, 20).

The first veins to appear are the two omphalo-mesenterics, of
which the right is somewhat stouter than the left (the reverse is
the case in Lacerta), and they open into the sinus venosus. Soon
after their appearance the anterior and posterior cardinal veins
arise on each side, and their common stem unites with the
umbilical vein on each side to form the ductus Cuvieri, which
becomes associated with the omphalo-mesenteric veins at the

* The number of renal arteries varies in different species, being only one in Python
spilotes and eight in Coronella catenifer, Beddard (1).

+ This differs from the account of 7. fasciatus in Beddard (1), where “it springs
from the aorta in both sexes close to the second (i.e. left) gonad artery, in front of it
in the male, behind it in the female.” Some variation is to be found, however, in the
position of this artery in different individuals, for in the females of 7. natrix that I
have examined, although it has generally been in front of the right anterior renal and
left ovarian arteries, it may be behind these vessels. Inthe male it is usually behind
the left anterior renal artery, but it may be in front of it. It has not occurred
anterior to the right anterior renal artery in any male snake that I have examined.
Again, we find considerable variation in the number of these gut-arteries in various
snakes. Lachesis gramineus has only one, while in the genus Coluber there may be
ten or eleven (Beddard, Joc. cit.).

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 627

point where they open into the sinus venosus. An anastomosis
between the two omphalo-mesenteric veins forms on the dorsal
side of the gut just posterior to the pancreas rudiment, and the
portion of the left vein between this point and the sinus venosus
disappears. A similar anastomosis between the two veins now
forms on the ventral side of the gut, and thus a complete ring is
formed. In a short time, however, the right half of this ring
disappears, leaving a single vein which runs in a spiral manner
round the gut. While these latter changes are taking place, the
middle part of the right omphalo-mesenteric vein between the
sinus venosus and the dorsal anastomosis spreads out and forms a
venous network in the liver. The portion of the right omphalo-
mesenteric vein in front of the hepatic network persists as the
hepatic vein, and the part behind it always remains as the anterior
end of the hepatic portal vein (text-fig. 90).

Text-fig. 90.

Diagram of posterior veins in the embryo. It shows the change from an early to
a late embryonic condition. The shaded vessels being the first to disappear.

A.C, Alimentary canal. D.C. Ductus Cuvieri. H.N. Hepatic network. 1.0.
Left omphalo-mesenteric (soon disappears). L.U. Left umbilical. O. Omphalo-
mesenteric. P.C. Post-caval. R.O. Right omphalo-mesenteric (disappears later
than L.O.). R.U. Right umbilical. S.V. Sinus venosus.

Adapted from Hochstetter (20).

The posterior cardinal vein originates at the caudal end of the
mesonephros, and runs forward along the dorso-lateral edge of
that body. After leaving the kidney, however, it breaks into
several branches, which soon reunite and then it runs into the
ductus Cuvieri. When the caudal vein develops it divides into
two branches at its anterior end, each of which runs to the
extremity of the corresponding cardinal vein (not along the mesial
wall of the kidney as in Lacerta). The post-caval vein springs
from the union of the right umbilical and omphalo-mesenteric

628 MR. C. H. 0 DONOGHUE ON THE

veins and runs backwards in the mesentery to the kidneys.
Between them it breaks up into two branches, which run pos-
teriorly along their mesial borders. These branches meet the
kidneys at a point some distance from their front end, and there
each receives a branch from the anterior part of the kidneys.
Subsequently the parts of the posterior cardinal veins in front of
each mesonephros disappear, and so all the blood from the caudal
veins has to pass through those organs.

Still later the adult kidney arises caudally to the mesonephros,
and then the end part of each posterior cardinal vein comes to lie
on its ventral and external border, while a continuation of one of
the posterior mesonephric branches of the post-caval vein extends
along the inner border of the permanent kidney.

The left umbilical vein loses its opening into the ductus Cuvier,
and all its blood is taken to the liver network. The right
umbilical vein, on the other hand, retains its original opening
into the ductus for some time (text- -fig. 90). In the subsequent
changes the post-caval vein increases in size, and the part of the
omplialo- -mesenteric vein joining it breaks up into a venous net-
work with the caudal extension of the liver. As the two umbilical
veins are joined by an anastomosis at the navel, it is possible for
the right to disappear, which it does, leaving the left, which,
however, disappears soon after birth*. Ultimately the omphalo-
mesenteric vein, lying on the dorsal side of the liver, can only
communicate with the sinus venosus vid the hepatic network and
the post-caval vein, which is on the ventral side of the liver.

The anterior cardinal veins originate in a similar way to those
of Selachians and Amphibians, but the parts of these veins in the
head are completely replaced in an interesting way (vide Grosser

-and Brezina, 19). The original cardinal vein runs backwards
from the infraorbital and the anterior cerebral veins ventrally to
the cranial nerves into the ductus Cuvieri. Three venous rings
are now formed in succession; the first around the root of the
facial nerve and the auditory vesicle, the second around the root
of the glossopharyngeal nerve, and the third around the vagus
root. Their median portions afterwards disappear, and the external
ones unite to form one lateral trunk. In the meantime two new
vessels come to open into the anterior cardinal: one, the median
cerebral vein, coming from the cerebellum, opens just posterior to
the trigeminal nerve; and the other, the posterior cerebral vein,
coming from the medulla, opens posterior to the vagus, leaving
the skull by the foramen magnum.

Still later the lateral trunk opens anteriorly into the infra-
orbital vein by means of an extension by the side of the second
and third branches of the trigeminal nerve, and posteriorly past
the hypoglossus into the anterior cardinal opposite to the pos-
terior cerebral vein. In this way is formed a new, complete

* Generally the umbilical vein disappears in the adult snake, but remains of it
persist in some species, e. g. Boa constrictor, B. diviniloqua, Python regius, Hunectes
murinus, Corallus cookii (Beddard, 4 & 5).

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 629

lateral trunk, the lateral cephalic vein (V. capitis lateralis, Grosser
and Brezina). At the same time the three cerebral veins become
connected by a median longitudinal vessel (text-fig. 91). Further,
two new anastomoses arise from the median cerebral vein, one
goes to the anterior cerebral vein and the other, the secondary
median cerebral vein, leaving the skull with the trigeminus, goes
to the lateral cephalic vein. This becomes the main vein leading
from the anterior part of the brain.

Gf

Lule,

Diagram of veins in the region of the head in the embryo. It shows the original -
veins indicated by the shaded portions and the definitive vessels indicated in
black.

A.C.V. Anterior cerebral. A.V. Auditory vesicle. A.V.C. Anterior cardinal.
T.0.V. Infra-orbital. J.V. Right common jugular. L.C.V. Lateral cephalic.
M.C.V. Median cerebral. P.C.V. Posterior cerebral. P.V. Prosencephalic.
S.M.C.V. Secondary median cerebral. II, V, VII, 1X, X, XII, Primordia of cranial

nerves.

Adapted from Grosser and Brezina (19).

The adult condition is reached by the regression of the dorsal
part of the anterior cerebral vein, the complete disappearance of
the portion of the anterior cardinal vein in the head, leaving its
cervical part, however, as the common jugular vein, and the
formation of a vein bringing back blood from the upper and
lower jaws.

630 MR. C. H. O DONOGHUE ON THE

(B) Adult Form.

(a) Anterior Vessels.

The whole of the blood from the head is brought back to the
heart by the two common jugular veins. The distribution of the
veins in the head and their union to form the common jugulars
will be dealt with later.

The Left Common Jugular Vein * (V. jugularis sinister, Schlemm)
runs from the anterior end of the neck parallel with and close to
the left common carotid artery to the heart. It passes ventral to
the left systemic arch to the dorsal side of the left auricle, and it
runs along the dorso-lateral edge of this structure to the auriculo-
ventricular sulcus. Here it bends sharply to the right and runs
into a part of the sinus venosus somewhat sharply marked off
from the remainder. It receives :—

i. In its anterior part a few small tributaries from the anterior
end of the csophagus.

i. The Coronary Veins (text-fig. 87), a series of small vessels
which join it in its course along the auriculo-ventricular
sulcus.

The Right Common Jugular Vein* (V. jugularis dextra,
Schlemm) is similar in disposition to the left, but at the level of
the anterior end of the right auricle it receives a large tributary,
the azygos vein. After this it runs straight on and unites with
the post-caval vein to form the major part of the sinus venosus.

I. The Azygos Vein (V. azygos, Schlemm) is a short trunk
leading into the right common jugular vein from the union of the
anterior and posterior azygos veins.

A. The Anterior Azygos Vein (V. azygos anterior, Schlemm) lies
in front of the heart between the esophagus and the
backbone, and extends from the beginning of the neck
back to the heart. It collects blood from the dorsal body-
wall along this region by means of a fairly regular series
of intercostal veins, all of which come off to the right of
the vertebral column. Just before it unites with the
posterior azygos vein it receives a branch from the
cesophagus.

(a) The Esophageal Vein is formed at the level of the front
end of the thyroid gland by the union of two branches.
One of these branches comes from the anterior dorsal
part of the cesophagus, and the other from the posterior
dorsal part. Their common stem runs almost straight
to the right, ventral to the vertebral artery and dorsal
to the right common jugular vein, and opens into the
anterior azygos vein.

* Tt is to be noted that the proximal parts of these two veins are homologous

with the precaval veins of Lacertilia. As, however, there is no representative of the

subclavian veins to mark the beginning of the precaval portion, it is convenient to
apply the one name to the whole vessel.

CIRCULATORY SYSTEM OF THE GRASS-SNAKE, 631

B, The Posterior Azygos Vein (V. Azygos posterior, Schlemm)
is a much shorter vessel than the anterior, and also than
the homologous vein in Lacerta. It originates at about
the level of the middle of the ventricle, and runs forward
to meet the anterior azygos just anterior to the right
auricle. In its course it receives three or four intercostal
veins, all of which come from the body-wall to the right of
the vertebral column.

(6) Posterior Vessels.

The Right Pulmonary Vein (V. pulmonalis, Schlemm) first
becomes noticeable as a definite vessel on the left side of the lung
at the level of the posterior end of the liver. It runs forward
along that organ to its anterior end, receiving numerous branches
and, leaving the lung, it runs parallel to the post-caval vein and
ventral to the right systemic arch to open into the right auricle.

A left pulmonary vein never exists at any time.

The Caudal Vein (V. caudalis, Schlemm) arises far back in the
tail and runs forward in the hemal canal, together with the caudal
artery. It leaves this canal and divides into two branches, the
renal portal veins, a short distance before reaching the cloaca.

The Renal Portal Vein (V. venalis advehens, Schlemm) on each
side runs forward from the bifurcation of the caudal vein over
the cloaca and along the outer side of each kidney. It receives
soon after its origin a lateral tributary, which from its position
and distribution appears to correspond to a pelvic vein *. If this
be the case, it is interesting as being the only indication in the
circulatory system of the derivation of the snake from a limb-
bearing ancestry. Each renal portal vein receives several small
tributaries from the dorsal body-wall, the cloaca, and the lateral
part of the tail. It then passes over the cloaca and lies between
the oviduct which is on its outer side and the ureter which is on
its Inner side. Shortly after passing the cloaca each vein gives
off a well-marked branch, the right being anterior to the left,
which passes dorsally around the ureter and then runs ventrally
to unite with its fellow on the dorsal wall of the gut. The vein
so formed, the “‘ Veine mésentérique postérieure” of Jourdain
(27), runs forward along the gut and is continued as the hepatic
portal vein. In its course from the cloaca to the kidney each
renal portal vein receives a number of small tributaries from both
ureter and oviduct, one of which, the Posterior Oviducal Vein, is
well marked, and also one or more from the dorsal parietes. On
reaching the kidney it runs along its lateral border closely accom-
panying the ureter, and, gradually getting smaller, disappears at
the anterior end of that organ. It is not continued anterior to
the kidney as in some snakes, e. g. the Boide and also in Zamenis
gemonensis (Beddard, 4 & 2), a feature recalling the condition in
the Lacertilia. During its course along the posterior part of the
kidney this vein receives a well-marked tributary from the
oviduct.

* A similar pair of veins is to be found in Hunectes murinus (Beddard, 2).

632 MR. GC. H. O DONOGHUE ON THE

The hight Efferent Renal Vein (V. venalis revehens, Schlemm)
originates along the inner margin of the kidney at its posterior
end, runs to the anterior end, and after a short independent
course in front of the kidney unites with its fellow of the opposite
side to give rise to the post-caval vein.

The Left Hfferent Renal Vein also has its origin along the inner
margin of the kidney. But, unlike the right, it receives branches
from the supra-renal body, the ovary, and the left oviduct in its
coursé from the kidney to the point where it unites with its
fellow. Thus :—

1. The Left Ovarian Veins are numerous small tributaries
flowing into the left efferent renal veins.

u. The Left Supra-Renal Veins are a large number of small
branches running from the supra-renal body into the
efferent renal vein, to which it is closely attached. They
return the blood brought to that body by the supra-renal
portal veins.

il. The Left Oviducal Vein is a large vein leaving the oviduct
at the level of the fimbriated funnel and flowing into the
efferent renal vein just in front of the supra-renal body.
It returns blood from the oviducal sinus.

A. The Left Oviducal Sinus is a wide thin-walled vessel
which runs from the extreme anterior end of the ovi-
ducal funnel backwards to about the level of the middle
of the corresponding kidney. It is very conspicuous in
injected specimens and was present in all the female
examples of 7’. natrix that I have examined, although it
does not appear to have been recorded in any other
Ophidian.

The Post-Caval Vein (V. cava posterior, Schlemm) is formed,
as has been stated above, by the union of the two efferent renal
veins. It passes forward, dorsal to the gut, freely in the mesen-
tery to the posterior extremity of the liver, and then along in a
groove in the ventral surface of that organ to its anterior end.
After leaving the liver it runs almost straight forward, ventral to
the right systemic arch and unites with the right pre-caval to
form the larger division of the sinus venosus. Soon after its
origin it receives veins from the right ovary, the supra-renal
body, and the oviduct. Thus :—

i. The Right Ovarian Veins are similar to those of the opposite

side.

i. The Right Supra-Renal Veins also resemble those on the
left and return the blood gathered by the supra-renal
portal veins.

il. The Right Oviducal Vein is like its fellow on the left and
comes from an oviducal sinus.

A. The Right Oviducal Sinus extends along the oviduct from
the anterior end of the funnel to about the level of the
middle of the right kidney.

CIRCULATORY SYSLEM OF THE GRASS-SNAKE. 633

The Supra-renal Portal System.—Hach supra-renal body has
a portal supply, an arrangement of vessels which is universally
present in snakes according to Beddard (1), who, however, attributes
the discovery of this system to Gratiolet in 1853 (17), whereas it
had been described seven years previously by Ecker (15). The
supply consists of two, but in more rare cases of three, intercostal
veins which arise from the corresponding side of the dorsal body-
wall and also of a vessel from the oviducal sinus in the female,
all of these pour their blood into the supra-renal network, The
last of these vessels is rather small and runs at the level of the
middle of the ovary on each side,

The Hepatic Portal Vein (Vena porte, Schlemm) arises by two
roots, from the renal portal veins, which unite to form a single
vessel on the dorsal wall ef the rectum in a way that has already
been described *. It passes forward through the gut mesentery,
receiving on its way numerous branches from the many coils of
the small intestine and also from the fat-body. Towards its
anterior end a tributary joins it bringing blood from the front
part of the intestine, the pancreas, and the spleen. This is shortly
followed by another vein coming from the gall-bladder. In the
same region it is Joined by the anterior abdominal vein which
runs down from the fat-body, From this point instead of being
on the ventral side of the post-caval vein it passes dorsally to the
left of this vessel and enters a furrow on the dorsal side of the
liver. Between the entrance of the vein from the pancreas and
the spleen and the posterior end of the liver, 7. ¢. just anterior to
the superior mesenteric artery, the hepatic portal vein receives
three large intercostal veins from the parietes of the right side.
The vessel runs in the groove of the liver right to its anterior end,
gradually diminishing in calibre, and it receives in this part of its
course a more or less regular series of intercostal veins arising to
the left of the vertebral column and a greater number of small
veins from the stomach and esophagus. A very similar condition
obtains in 7’, fasciatus (Beddard, 1).

I. The Anterior Abdominal Vein T, corresponding to the simi-
larly-named vein in Lacertilia, is a single small vein arising at

* According to Schlemm (35) the hepatic portal vein has only one root, and that
arises from the right renal portal vein. This statement is also made in Hoffmann
(23), but it should be noted that this author quotes nearly the whole of Schlemm’s
account of the venous system almost verbatim without indicating in any way that
he is so doimg. My own investigations confirm those of Jourdain (2'7) and Hoch-
stetter (20), who describe a double root for this vein in Tvopidonotus, one part
avising from each renal portal vein. ‘The latter author makes a similar statement
with regard to Coluber esculapii. Beddard (4 & 3) has also recorded the same
arrangement in Coluber corais, Zamenis gemonensis, and Ancistrodon piscivorus.
According to Jaquart (26) there are a number of anastomoses between the hepatic
portal vein and the right renal portal vein in Python. In Gadow’s account of
Pelophilus madagascariensis, quoted hy Hoffmann (28), it is stated that there is no
connection between the hepatic portai and renal portal veins.

+ This vein is especially interesting, as it is subject to considerable variation among
the Ophidia. In Lizards, as is well known, it arises by two roots from the renal
portal veins, and a similar condition is to be found in some snakes, viz., Hryx jaculus,
Ei. johni, Python sebe, and Boa diviniloqua, Beddard (2,3, & 4). In other snakes

Proc. Zoot. Soc.—1912, No. XLII. A2

634 MR. C. I. O DONOGHUE ON THE

the posterior end of the fat-body and running forward in it
to the level of the spleen, where it passes dorsally and opens into
the hepatic portal vein. It is only connected in an indirect way
by small anastomosing branches with the renal portal veins.
Along its course it receives little twigs from the epigastric vein.

The Lpigastric Vein lies beside the epigastric artery in the mid-
ventral line of the abdominal wall. In the region of the liver it
gives off five or six small branches, all of which enter directly
into the left side of that body and are not connected with the
hepatic portal vein. Behind the liver the epigastric vein is
connected by a number of small venules with the anterior
abdominal vein. As Beddard (4) has pointed out, this is one
of the most constant veins in Snakes, and is single save in
Lioheterodon madagascariensis, where it is alternately single and
double.

The veins in the male are, like the arteries, on the whole very
similar to those in the female. Those in front of the heart are
precisely similar in both sexes.

The caudal vein bifurcates to form the renal portal veins which,
at the level of the cloaca, receive the paired pelvic veins, and in
addition, in the male a vein from each corpus cavernosum. The
renal portals give rise to the two branches which unite above the
gut to form the beginning of the hepatic portal vein and then
pass forwards to the kidneys between the vasa deferentia and the
ureters. On the kidney they receive no specially marked tribu-
tary from the vas deferens to correspond with the one from the
oviduct in the female.

Each testis gives off one spermatic vein, whereas in the female
there are a number of small ovarian veins, just in front of the
corresponding supra-renal body, that of the left side opening into
the left efferent renal vein, and that of the right into the post-
caval vein.

There is no vessel in the male to correspond with the oviducal
sinus, and consequently no branch from it to the supra-renal
body. The supra-renal portal supply consists of two intercostal
veins, one at each end of that body, which arise from the corre-
sponding side of the vertebral column.

Some variation is to be met with in’ the position of the union
of the two efferent renal veins with regard to the kidneys. The
junction may be as much as an inch in front of the right kidney,
cr, on the other hand, this kidney may overlap the point of union,
in which case several small veins bring back the blood from the

it has only a single origin from the left renal portal vein, viz., Hryx conicus, Hunectes
murinus, and H. noteus, Beddard (8). Lastly, it may have no direct connection
with the renal portal veins, but only indirect ones by means of anastomosing twigs,
viz., Zamenis gemonensis, Beddard (2), Coluber escula, and Tropidonotus natria,
Hochstetter (20). This last observation I have been able to confirm. Further, the
anterior abdominal vein may be partly double throughout, as in Boa constrictor,
B. diviniloqua, Eryx jaculus, Beddard (4), and Python sebe, Jaquart (26), or
single as in Zamenis gemonensis, Causus rhombeatus, Eryx johni, Beddard (2), and
Tropidonotus natrix.

CIRCULATORY SYSTEM OF TIE GRASS-SNAKE. 635

anterior end of the kidney into the right efferent renal vein. On
the whole this distance appears to be greater in the female than
in the male.

The remaining vessels of the male, the hepatic portal factors,
the anterior abdominal vein, and the epigastric vein correspond
in all respects to those of the female.

V. Tse Vessens or tHe Heap. (Pls. LX XI. & LXXII.)
(A) Arteries.

It has been pointed out above that the whole of the blood
is brought to the head by the left common carotid, the right
common carotid having disappeared early in the course of
development. To compensate for this absence of an artery on
the right ride we find developed three arterial anastomoses
between the two sides of the head. The first lies beneath the
medulla oblongata and joins the two internal carotids ; the second
is situated beneath the fore-brain just in front of the optic
chiasma and unites the anterior cerebral and facial carotids ; and
the third is behind the symphysis of the lower jaw and joins the
two external carotids.

The Left Huternal Carotid (Carotis externa, Rathke; Arteria
inframaxillaris, Schlemm) arises from the common carotid internal
to the articulation of the lower jaw and the quadrate bone. It
runs forward between the floor of the pharynx and the broad
mylohyoideus * muscle, first inwards towards the toneue sheath
and then outwards to the inner side of the mandible, being
accompanied throughout the greater part of its course by its
corresponding vein, the glossopharyngeal nerve, and the cutaneous
branch of the hypoglossal nerve. At the anterior end of the
lower jaw, just a short distance behind the symphysis, the left
external carotid anastomoses with its fellow by a well-marked
vessel.

The Right Huternal Carotid is similar to the left, save that the
common carotid from which it originated has disappeared and is
represented only by a small branch vessel, It receives its blood-
supply partly from the anastomosis just mentioned, and partly
from the anastomosis between the internal carotids.

The distribution of the arteries in the dorsal part of the head
is the same on both sides, so that the ene description will apply
equally well to either side.

The Internal Carotid (Carotis interna, Rathke; Art. cephalica
and Art. carotis communis, Schlemm) starts from the origin of
the external carotid and bends in a sharp curve dorsally round
the angle of the lower jaw on the inner side of the vagus and
hypoglossal nerves. It then passes forward under the columella
and along the inner side of the quadrate to a point behind the
orbit and above the posterior pterygo-spheneidalis muscle, where

* The nomenclature of the muscles is that adopted by Hoffmann (28).
4Q%

=

636 MR. CG. If. O DONOGHUE ON THE

it divides into two branches, the cerebral carotid and facial carotid
arteries,
During this course it gives off the following branches :-—

I. One well marked and sometimes also a smaller artery that
supply the cesophagus.

II. A branch that supplies a part of the cervico-mandibularis
muscle, the lateral and ventral walls of the pharynx and skin in
this region.

TI. A branch (Ramus pterygoideus, Schlemm) which supplies
the transverso-maxillo-pterygo-mandibularis, and the cervico-
mandibularis muscles and the skin near them.

IV. The First Spinal Artery (Ramus spinalis, Rathke; Art.
nervi spinalis I, Hofmann) which arises at the same level or
slightly in front of the preceding. The vessel passes through the
atlanto-occipital membrane between the axis and the basi-occipital
bone and, running into the loop of the basilar artery beneath the
medulla oblongata, forms an anastomosis with its fellow of the
opposite side, a relationship first described by Schlemm (35), who

ealled this vessel the Truncus anonymus. By means of this
anastomosis the blood can pass from the left to the right internal
carotid. Shortly before piercing the atlanto- occipital membrane
this artery gives off two branches :—

A. A small branch (Ramus pterygoideus, Schlemm) to the
Pterygo-sphenoidalis muscles.

B. A fairly large Occipital artery (Art. occipitalis, Schlemm),
which however soon divides into two vessels, a small and a
large.

The smaller is the occipital branch (Ramus occipitalis, Schlemm),
supplying the muscles and skin of the occipital region.

The larger is the Cervical Artery (Art. cervicalis, Schlemm), a
long and fairly well-marked vessel. It runs backwards
near the hypapophyses of the anterior vertebrae, covered
by the rectus capitus anticus muscles to which it sends
small branches, to the fourteenth vertebra. Here it joins
with a small branch from the bifurcated extremity of the
vertebral artery. During its course it gives off a series of
branches that he beside the spinal nerves, and each of
these sends branches not only to the skin and muscles,
but also a small one, a spinal artery, that enters the
neural canal through the intervertebral foramen.

V. The Maxillary Artery (Art. maxillaris, or dentalis inferior,
Rathke ; Art. alveolaris inferior, Schlemm) arises about midway
between the spinal artery and the point just behind the orbit
where the internal carotid divides. It runs outwards, accompany-

ing the inferior maxillary branch of the trigeminal nerve between

the second and third parts of the parietali-quadrato-mandibularis
muscle, to which it gives branches. After giving off several twigs
also to the transverso-maxillo-pterygo-mandibularis muscle, it
passes on through the posterior maxillary foramen into the lower

CIRCULATORY SYSTEM OF THLE GRASS-SNAKE. 677

jaw. It soon gives off a branch to the inferior Jabial gland *, and
then runs on in the lower jaw to its anterior end. Here it comes
out through the anterior maxillary foramen as the Mentalis Artery
(Art. mentalis, Rathke) and supplies the anterior part of the
inferior labial gland and the skin in the region of the chin.

Just anterior to this branch the internal carotid splits into two
branches, the cerebral carotid and the facial carotid arteries.

The Cerebral Carotid (Art. carotis cerebralis, Rathke and Hof-
mann) is a vessel of moderate calibre running downwards just
behind the infra-maxillary branch of the trigeminal nerve.
It then turns inwards under the skull and enters the cranial
cavity through a foramen in the basisphenoid. This vessel was
overlooked by Schlemm. In its course it gives off a small artery
just outside the skull going to the posterior pterygo-sphenoidalis
muscle. Inside the skull near the hinder end of the pituitary
fossa 1t splits into three fairly equal vessels, the posterior, median,
and anterior cerebral carotids.

I. The posterior branch (Ramus caudalis, Hofmann) runs back-
wards round the outside of the pituitary fossa, giving off branches
to the cerebellum and, about half-way to the foramen magnum,
unites with the similar vessel of the other side to form the median
basilar artery.

The Basilar Artery (Art. basilaris, Rathke and Hofmann)
passes backwards until just before the foramen magnum, where
it splits up into a loop in the form of an isosceles triangle. Into
the corners of the base of this loop open the right and left first
spinal arteries, and thus it forms the anastomosis between these
two vessels. Before it divides to form the loop, the basilar artery
gives off on either side a well-marked internal auditory artery
(Art. auditiva interna, Rathke and Hofmann) that enters the
ear with the auditory nerve, and also a series of smaller branches,
some of which supply the medulla, and some run on to the small
choroid plexus of the fourth ventricle.

Il. The median branch fT runs outwards behind the cerebral
hemisphere, to which it gives some twigs, and a short distance .
from its origin divides into two.

A. The Anterior Choroid Artery (Art. choroides anterior,
Hofmann) is the anterior branch. It passes around the
hemispheres, supplying them with smail twigs, to the
dorsal side of the brain near the pineal stalk, where it
breaks up in the choroid plexus of the third and lateral
ventricles and anastomoses with a branch from the olfactory
artery.

B. The posterior division passes behind the optic lobes and
spreads itself out over their dorsal surface.

* The nomenclature of these glands in the head of the snake is that given by West
(37 & 38).

+ In some examples this median branch does not arise at the point where the
cerebral carotid splits into anterior and posterior branches, but a little way down the
latter ; consequently it appears as a branch of the posterior cerebral carotid and is
described as such by Hofmann (24).

638 MR. C. H. O DONOGHUE ON THE

III. The anterior branch (Ramus cranialis, Hofmann) runs
forward along the side of the pituitary fossa and unites with the
similar vessel from the other side under the bases of the optic
nerves immediately in front of the chiasma. By the union of
the two anterior and the two posterior branches of the cerebral
carotids a complete arterial ring, the circle of Willis (circulus
arteriosus cerebralis, Hofmann) is formed around the pituitary
fossa. During its course this anterior branch gives off :—

A. A Median Cerebral Artery (Art. cerebri media, Hofmann),
which is a large vessel running round to the dorsal side
of the brain and supplying the anterior end of the hemi-
spheres and the olfactory lobes.

B. An Ophthalmic Artery (Art. ophthalmica, Hofmann), which
is given off immediately before the two anterior branches
of the cerebral carotids unite to form the circle of Willis.
This passes out of the skull with the optic nerve, and
immediately on reaching the orbit anastomoses with a
branch of the facial carotid.

The Olfactory Artery * (Art. olfactoria, Schlemm) arises from
the mid-point of the anastomosis of the two anterior branches of
the cerebral carotids and runs forward ih the skull in the furrow
hetween the two olfactory lobes, to which it sends branches. At
the anterior end of the lobes it gives off two symmetrical branches.
Kach of these again divides into two (Aa. ethmoidales, Rathke ;
Aa. bulbi olfactorii mediales, Hofmann), which pass out of the
skull with the olfactory nerve to ramify over the olfactory
membrane, The main trunk then recurves dorsally and runs back
in the fissure between the two hemispheres, to which it sends
numerous small branches, and near the pineal body anastomoses
on each side with a branch of the anterior division of the median
branch of the cerebral artery.

Tre Inferior Spinal Artery (Art. spinalis inferior, Rathke ;
Tractus spinalis ventralis, Hofmann) is situated just below the
ventral fissure of the spinal cord, and runs in a fairly straight line
caudally from the anastomosis between the right and left first
spinal arteries at the posterior end of the basilar artery. On its
course it gives off branches to the spinal cord, some of which pass
around to the dorsal side and others enter the ventral fissure, and
it also receives the paired spinal arteries which come in through
the vertebral column at the points of exit of the spinal nerves.
In some places where a pair of such arteries enter it, the inferior
spinal artery splits into a diamond-shaped loop.

The facial Carotid (Carotis facialis, Rathke) takes a fairly
straight course forward through the temporal fossa close to the
infra-maxillary branch of the trigeminal nerve to the orbit.
Here it passes under the post-frontal bone into the orbit and

* This artery 1s double for the greater part of its length in Python molurus,
Beddard (6).

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 639

immediately divides into two fairly equisized terminal branches.
On its way it gives off :—

I. A branch (Ramus glandule maxillee superioris posterior,
Schlemm) which arises a short distance behind the orbit and runs
outwards to the posterior part of the parotid gland, @. e., the gland
corresponding to the poison-gland of other snakes. It ramifies
in the gland, and one of its branches anastomoses with a branch
of the artery supplying the anterior part of the gland. During
its course it gives off a palatine branch and branches to several
muscles.

II. Several small branches and one fairly well-marked one that
supply the Harderian gland.

III. Two or three slender branches to the skin overlying the
skull and one or two to the under side of the skull.

One terminal branch (Carotis facialis, Schlemm) of the facial
carotid passes along the back of the orbit downwards to its floor,
where it runs forward (Ramus palatinus anterior, Schlemm),
accompanying the infra-orbital branch of the trigeminal nerve.
It goes on forward out of the orbit, close to the outer wall of the
internal nar es, and finally spreads out over the skin and muscles.
at the front end of the snout. During its course it gives off :—

I. A well-marked vessel (Ramus glandule maxillaris superioris
anterior, Schlemm) that accompanies the supra-maxillary branch
of the trigeminal nerve outwards in the posterior wall of the
orbit and along the upper jaw. It supplies the teeth in the pos-
terior part of the upper jaw, the superior labial gland, the
anterior part of the parotid gland, within which one of its small
twigs anastomoses with a similar twig from the artery supplying
the posterior part of this gland, and finally it gives off small
vessels to the skin in this region.

II. A posterior palatine branch (Ramus palatinus recurrens,
Schlemm) that goes to the posterior palatine teeth.

III. Small branches to the anterior palatine teeth.
TV. Branches to the teeth in the anterior end of the jaw.
V. Branches to the nasal gland.

The other slightly stouter terminal branch of the facial carotid
artery (Art. carotis cerebralis, Schlemm) runs in an irregular arch
along the upper, inner, and hinder sides of the orbit to the optic
foramen, where it anastomoses with the ophthalmic artery. Ton
this way the blood from the left facial carotid can pass vid the
ophthalmic artery into the circle of Willis, and thence to the
encephalic arteries and also over into the right facial carotid. In
its course this artery sends out the following branches :——

I. Several well-marked twigs to the part of the Harderian

gland within the orbit.
II. Branches to the muscles of the eye (Aa. musculares oculi,

Rathke).
III. Two short ciliary branches (Aa. ciliares postice breves,

Rathke).

640 MR. C. I. O DONOGHUE ON THE

IV. Two longer ciliary branches (Aa. ciliares postice longe,
Rathke), one of which runs forward on the inner side of the eye
and the other outward on its hinder side.

V. A Retinal Artery (Art. centralis retine, Rathke) which
enters the eyeball with the optic nerve and spreads out over the
retina.

(B) Veins.

The veins of the head of Zropidonotus natrix have recently
been described in great detail by Bruner (12), who also describes
a muscular mechanism in the head whereby the blood-pressure in
its veins and sinuses may be considerably increased. It is not
possible by ordinary dissection to make out all the small vessels
given by that author, and, as my own results agree closely with
his, it will only be necessary, for the sake of completeness, to give
a brief description of the cephalic veins.

The Mandibular Vein (V. inframaxillaris, Schlemm ; V. mandi-
bularis, Br runer’) arises from a small sinus at the anterior end of
the lower jaw and runs backwards close to and on the outer side of
the external carotid artery. On its way it receives veins from the
trachea, tongue-sheath, muscles of the floor of the mouth, and
the pharynx. It runs into the maxillary vein immediately before
the latter joins with the lateral cephalic vein to form the common
jugular vein. The bases of the mandibular, maxillary, and lateral
cephalic veins and the anterior end of the common jugular vein
are surrounded by constrictor muscles whose morphology and
function are described by Bruner (loc. cit.).

The Mawillary Vein (V. palatina, Schlemm; V. maxillaris,
Bruner) also commences in a small sinus which is situated just
behind the premaxilla. This sinus has a double anastomosis with
the similar one on the other side. From this point it runs back-
wards beneath the nasal cavity along the floor of the orbit and
then above the palate to join the mandibular vein at the angle of
the lower jaw. During its course it receives :—

I. The Rostral Vein (V. rostralis, Bruner), which enters at the
level of the anterior anastomosis and brings the blood from a
venous network at the front end of the snout. It also receives
nasal veins (V. nasales externe, dorsalis, and ventralis, Bruner)
from the nasal gland.

Il. The Subnasal Vein (Sinus subnasalis, Bruner), which enters
at the level of the posterior anastomosis. It drains a somewhat
complex system of subnasal sinuses, which anastomose with one
another at the posterior end of the nasal cavity and which receive
also the palato-pterygoid vein.

The Palato-pterygoid Vein (Sinus palato-pterygoideus, Bruner)
runs from near the anterior to near the posterior end of
the skull on the inner side of the palatine and pterygoid
bones, and flows into thé anastomosis between the sub-
nasal sinuses.

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 641

III. A vein that joins it at the anterior end of the orbit and
forms an anastomosis between it and the orbital sinus.

IV. An Inferior Palpebral Vein (V. palpebralis inferior, Bruner)
that also enters at the front end of the orbit and runs backwards
in the lower eyelid to its posterior end, where it enters the orbital
sinus together with the superior palpebral vein.

V. Several small veins from the orbital sinus just posterior to
the orbit.

VI. An Oblique Palatine Vein (V. palatina obliqua, Bruner)
that runs obliquely forward beneath the skull at the level of the
hypophysis to join the similar vessel of the opposite side and form
a median palatine sinus which runs forward for a short distance.
Before reaching the middle line it gives off a palato-cerebral
vein.

The Palato-cerebral Vein (V. palato-cerebrales, Bruner) runs
dorsally around the skull and enters the median cerebral
vein soon after this leaves the cranium.

The Orbital Sinus (Sinus orbitalis, Bruner) is a fairly large well-
defined sinus occupying the inner and hinder parts of the orbit.
At the outer end of the hinder part it is prolonged outside the
orbit beneath the Harderian gland. It receives the following
branches :—

I. A small vein at its anterior end which comes from the nasal
gland.

Il. The vein joining it to the maxillary vein which also enters
at the anterior end.

IIl. The Superior Palpebral Vein (V. palpebralis superior,
Bruner), which arises at the anterior end of the sinus and runs
backwards in the upper eyelid, re-entering the sinus at its
posterior end and receiving just before it does so the inferior
palpebral vein.

IV. The Secondary Anterior Cerebral Vein (Sekundare Verbind-
ung der v. cerebralis media mit der v. cerebralis anterior, Grosser
und Brezina) which runs from the posterior internal corner of the
orbital sinus backwards inside the skull into the median cerebral
vein just as the latter is leaving the cranial cavity. The anterior
segment of this vessel is formed by a part of the original anterior
cerebral vein.

The Lateral Cephalic Vein (V. capitis lateralis, Grosser und
Brezina; V. jugularis interna, Bruner) arises from the posterior
prolongation of the orbital sinus and runs inwards and backwards
to the side of the internal carotid artery. It passes backward
closely accompanying this artery to the posterior end of the head,
where it bends round to the ventral side and unites with the
maxillary and mandibular veins to form the common jugular vein.
On its course it receives :—

J. A vein from the Harderian gland.
II. The Median Cerebral Vein (V. cerebralis media, Bruner).—
This runs from the longitudinal cerebral vein on the dorsal side

642 MR. C. H. O'DONOGHUE ON THE

of the brain, outwards, and around the posterior face of the optic
lobes to the ventral side of the brain. Here it goes forward and
leaves the skull by the foramen for the trigeminal nerve. Out-
side the skull it bends sharply backwards and joins the lateral
cephalic vein as the latter reaches the internal carotid artery.
The last part of this vessel outside the skull is a secondary con-
nection (V. cerebralis media secundaria, Grosser and Brezina)
developed between the median cerebral vein, which originally
opened into the internal jugular, and the lateral cephalic vein.
During its course it receives :—

A. The Dorsal Cephalic Vein (V. capitis dorsalis, Bruner), which
arises from its dorsal side within the. skull and passes
outward through a special foramen. It runs baekwards
between the pro-otic and squamosal bones, receiving one
or two cutaneous veins, and then bends laterally and enters
the lateral cephalic vein.

B. The Secondary Anterior Cerebral Vein, which runs on the
floor of the cranium and joins it to the orbital sinus (vide
supra).

C. The Palato-cerebral Vein, which connects it with the oblique
palatine vein (vide supra). According to Bruner there
is also an external secondary anastomosis with the anterior
cerebral vein, as well as the internal one described above.
I have been unable to find this yein by dissection.

IIT. A large vein from the parotid gland and the muscles of
the head, which closely accompanies the maxillary artery and
enters the lateral cephalic vein close to the place where the
maxillary artery leaves the internal carotid.

IV. A Dorsal Cephalic Vein (vide supra), which joins it to the
median cerebral vein.

V. The Posterior Cerebral Vein (V.cerebralis posterior, Bruner),
which runs from the end of the longitudinal cerebral vein a little
behind the posterior end of the optic lobes obliquely outwards
over the medulla oblongata and leaves the skull by the foramen
magnum. Just before leaving the cranium it gives off a Spinal
vein Which runs caudally on the ventral side of the spinal cord,
where it unites with the similar vessel from the other side.

VI. One well-marked and several smaller veins from the
muscles of the posterior end of the skull.

VIL. A Cervical vein which returns blood from the muscles of
the neck.

The Longitudinal Cerebral Vein (V. longitudinis cerebri, Bruner)
is a vessel running backwards along the mid-dorsal aspect of the
brain from between the olfactory lobes. At the posterior end of
the optic lobes it gives off the median cerebral veins, and a short
distance further back divides to form the posterior cerebral
veins,

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 643

VI. List or REFERENCES.

(1) Bepparp, F. E.—Contributions to our Knowledge of the
Circulatory System in the Ophidia. Proce. Zool. Soc.
1904, vol. i. p. 331.

(2) Bepparp, F. E.—Notes upon the Anatomy of Certain Snakes
of the Family Boide. Proc. Zool. Soc. 1904, vol. ii. p. 107.

(3) Bepparp, F. E.—Contributions to the Anatomy of the
Ophidia. Proce. Zool. Soc. 1906, p. 12.

(4) Bepparp, F. E.—Contributions to the Knowledge of the
Vascular and Respiratory Systems in the Ophidia, and to
the Anatomy of the Genera Boa and Corallus. Proc.
Zool. Soc. 1906, p. 499.

(5) Bepparp, F. E.—A Comparison of the Neotropical Species
of Corallus, C. cookti, with C. madagascariensis ; and on
some Points in the Anatomy of Corallus caninus. Proc.
Zool. Soc. 1908, p. 135.

(6) Brepparp, F, E—A Contribution to the Knowledge of the
Encephalic Arterial System in Sauropsida. Proc. Zool.
Soc. 1905, vol. i. p. 59.

(7) Bemmeten, J. F. van.— Entwikkelung en metamorphose der
kieuw of veseralspalten en der aortenbogen bij Embryonen
van Tropidonetus natria en Lacerta muralis. Kon. Akad.
v. Wet. Amsterdam, Afd. Natuurk., 1885.

(8) Demmeten, J. F. van.—Die Visceraltaschen und Aortenbogen
bei Reptilien und Vogeln. Zool. Anz., 1886.

(9) Bemmeren, J. F. van.—Beitrage zur Kenntnis der Halsgegend
bei Reptilien. 1. Anatomischer Teil. Mededeelingen tot
de Dierkunde. Tijdschrift van het Genootschap Natura
artis magistra te Amsterdam, 1887,

(10) Branpr, F.—Ueber einen eigenthumlichen, spater meist
obliterivenden Ductus Caroticus der gemeinen Kreuzotter
(Pelias berus). Mélange biologique, Bd. v., 1865.

(11) Brenner, A.—Uber das Verhiltnisse des N. laryngeus
inferior vagi zur eigen Aortenvarietiiten des Menschen
und zu dem Aortensystem der durch Lungen athmenden
Wirbelthiere tiberhaupt. Archiv fur Anat. und Physiol.,
Anat.-Abt., 1883.

(12) Bruner, H. L.—On the Cephalic Veins and Sinuses of
Reptiles, with a description of a mechanism for raising
the venous blood pressure in the head. American Jour.
of Anat., 1907.

(13) Denpy, A.—The Intracranial Vascular System of Sphenodon.
iPlnui, Abrenais., IO

(14) Ds Vrresz, B.—Sur la Signification morphologique des
Arteres cérébrales. Archives de Biologie, xxi., 1905.

(15) Ecxer, A.—Der feinere Bau der Nebennieren beim Menschen
und den vier Wirbelthierklassen. Braunschweig, 1846.

(16) Frirscu, G.—Z4ur vergleichenden Anatomie der Amphibien
herzen. Archiv fiir Anat. und Physiol., 1869.

644 MR. C. I. O DONOGHUE ON THE

(17) Gratioter, P.—Note sur Je systéme veineux des Reptiles.
L'Institut xxi., 1853.

(18) Grett, A.—Beitriige zur vergleichenden Anatomie und
Entwicklungsgeschichte des Herzen und des Truncus
arteriosus der Wirbelthiere. Morph. Jahr., Bd. 31, 1903.

(19) Grosser, O., und Brezina, E.—Ueber die Entwicklung der
Venen des Kopfes und Halzes bei Reptilien. Morph.
Jahr., Bd. 23, 1895.

(20) Hocusrerrer, F.—Beitriige zur Kntwicklungsgeschichte der
Amnioten. 2. Reptilien (Zropidonotus, Lacerta). Morph.
Jahr., Bd. 19, 1893.

(21) Hocusrerrer, F.—Ueber Varietaiten der Aortenbogen,
Aorten wurzeln und der von ihnen entspringenden Arterien
bei Reptilien. Morph. Jahr., Bd. 29, 1902.

(22) Hocusrerrer, F.—Die Entwicklung des Blutgefiisssystem.
In Hertwig’s Handbuch der Entwicklungslehre der Wir-
beltiere, 1901-06.

(23) Horrmann, CO. K.—Schlangen und Entwicklungsgeschichte
der Schlangen. SBronn’s Thier-Reich, 1890.

(24) Hormann, M.—Zur vergleichenden Anatomie der Gehirn-
und Riickenmarksarterien der Vertebraten. Zeitschrift
fiir Morph., Bd. 11., 1900.

~ (25) Hopkinson, J. P., and Pancoat, J.—On the Visceral Anatomy
of the Python (Cuvier) described by Daudin as the Boa
reticulata. Trans. American Phil. Soc., 1837.

(26) Jaquart, H.—Meémoire sur les Organes de la Circulation
chez le serpent Python. Ann. des Sci. Nat., Tome iv.,
1855.

(27) Jourpain, 8.—Recherches sur la vein Porte Rénale. Ann.
des Sci. Nat., 1859.

(28) Kinestey.—Guides for Vertebrate Dissection. New York,
USOT.

(29) Lancer, A.—Uber die Entwicklungsgeschichte des Bulbus
eordis bet Amphibien und Reptilien. Morph. Jahr.,
Bd. 21, 1894.

(30) Raruxe, H.—Entwicklungsgeschichte der Natter. Kénigs-
berg, 1839.

(31) Raruxe, H.— Ueber die Carotiden der Schlangen.
Denkschr. Wien. Akad. Wiss., Math.-nat. K1., Bd. xi., 1856.

(32) Ronieston, G., and Jackson, W. H.—Forms of Animal
Life, 2nd Ed., p. 70. Oxford, 1888.

(33) Sasatier, A.—Ktudes sur le cceur et la circulation centrale
dans la série des Vertébrés. Ann. des Sci. Nat., Tome 18,
1873.

(34) Sapatrer, A.—Observations sur les transformations du
Systeme aortique dans la série des Vertébrés. Ann. des
Sci. Nat., Tome 19, 1874.

(35) Scutemm, F.—Anatomischer Beschreibung des Blutgefiiss-
system der Schlangen. ‘Treviranus’s Zeitschrift fiir

Physiologie, Bd. 3, 1827.

CIRCULATORY SYSTEM OF TILE GRASS-SNAKE.

645

(36) 'Tanpier, J.—Mikroskopische Tnjectionen mit kaltflussiger

Gelatin.

OU
(37) West, G. S.—On the Buccal Glands and Teeth of certain

Poisonous Snakes.

Zeitsch. f. wiss. Mikros. v. f. Mikros. Tech.,

Proe. Zool. Soc. 1895, p. 812.

(38) West, G. S.—On two little-known Opisthoglyphous Snakes.
Jour. Linn. Soe., Zool. 1896.

VII. EXPLANATION OF PLATES LXX—-LXXII,
Lettering.
A.A.L. Left Aortic Arch. G.A, Gastric Artery,
A.A.R. Right Aortic Arch. Gall. Bl. Gall-bladder.
A.A.V. Anterior Abdomimal Vein. H.A. Hepatic Artery.
A.C.A. Anterior Choroid Artery. Har. Gl. Harderian Gland,
A.V. Azygos Vein. H.G.A. Harderian Gland Artery,
A.V.A. Anterior Azygos Vein. H.P.V. Hepatic Portal Vein.
A.V.P. Posterior Azygos Vein. H.P.V.’ The posterior part of the
B.A. Basilar Artery. Hepatic Portal Vein, the
B.A.L. Loop of Basilar Artery. “Veine mésentérique
C.A. Left Common Carotid postérieure” of Jourdain
Artery. (27).
Ca. A. Caudal Artery. H.P.V.O. Origin of the Hepatic Portal
C,.C.A. Cerebral Carotid Artery. Vein from the Renal Portal
C.C.Ant. Anterior branch of the Vein,
Cerebral Carotid Artery. J.A.A. Internal Auditory Artery.
C.C.M. Median branch of the 1.C.A. Internal Carotid Artery.
Cerebral Carotid Artery. I.C.M. Muscular branch of In-
C.C.P. Posterior branch of the ternal Carotid Artery.
Cerebral Carotid Artery. 1,C.O. Esophageal branch of In-
Cep. V.D. Dorsal Cephalic Vein. ternal Carotid Artery.
Cep. V.L. Longitudmal Cephalic I.M.A. First Inferior Mesenteric
Vein. Artery.
Cer. A. Cervical Artery. I.M,A.2. Second Inferior Mesenteric
Cer. V. Cervical Vein. Artery.
Cer. Hem. Cerebral Hemispheres. I.M.A.3. Third Inferior Mesenteri¢
C.W. Circle of Willis. Artery,
C.J.L. Left Common Jugular I.M.A.4. Fourth Inferior Mesenteric
Vein. Artery.
C.J.R. Right Common Jugular Inf. Infundibulum,
Vein. I.P.V. Inferior Palpebral Vein.
C.V. Caudal Vein. I.S.A. Inferior Spinal Artery.
C.V.L. Longitudinal Cerebral Kid. L. Wett Kidney.
Vein. Kid. R. Right Kidney.
C.V.M. Median Cerebra! Vein. L.A. Left Auricle.
C.V.P, Posterior Cerebral Vein. L.C.A. Long Ciliary Artery.
C.V.S.A. Secondary Anterior Cere- L.G.A. Lieno-gastric Artery.
bral Vein. L.R.A. Anterior Left Renal
D.A. Dorsal Aorta. Artery.
E.C.A. Anastomosis of the two Ex- L.R.M. Median Left Renal Artery,
ternal Carotid Arteries, L.R.P. Posterior Lett Renal
E.C.L. Left External Carotid Artery.
Artery. M.A. Maxillary Artery.
E.C.R. Right External Carotid M.C.A. Median Cerebral Artery.
Artery, M.C.P. Posterior branch of the
E.M.A. Eye Muscle Artery. Median branch of Cere-
H.R.L. Lett Efferent Renal Vein. bral Carotid Artery,
E.R.R. Right Efferent Renal Vein. Med. Medulla oblongata,
Ext. Nar. External Nares. M.V. Mandibular Vein.
F.A. Fat-body Artery. Mx. Maxillary Vein,
.C.A. Facial Carotid Artery. Mx.A. The double anastomosis of
F.C.P. Palatine branch of the the Maxillary Vein.
Facial Carotid Artery, O.A,L. Left Ovarian Artery,

646 ON THE CIRCULATORY SYSTEM OF THE GRASS-SNAKE.
O.A.R. Right Ovarian Artery. Pel. V. Vein probably corres-
Oc. A. Occipital Artery. ponding to the Pelvic
Od. A. Oviducal Artery. Vein ot Lacertilia.
Od. A.P. Posterior Oviducal Artery. Pit. Fos. Pituitary Fossa.
Od.S.L. Lett Oviducal Sinus. P1.C.V. Palato-cerebral Vein.
Od. S.R. Right Ovidueal Sinus. Pl. P.V. Palato-pterygoid Vein.
Od. V.L. Lett Oviducal Vein. Pul. A. Pulmonary Artery.
Od. V.L.P. Left Posterior Oviducal Pul. V. Pulmonary Vein.
Vein. P.V. Parietal Vein.
Od. V.R. Right Oviducal Vein. R.A. Right Auricle.
Od. V.R.R. Right Posterior Oviducal Ret. A. Retinal Artery.
Vein. R.P.L. Left Renal Portal Vein.
Oe. A. Hsophageal Artery. R.P.R. Right Renal Portal Vein.
Oe. V. @sophageal Vein. R.R.A. Anterior Right Renal
Ol. A. Olfactory Artery. Artery.
OLf. lo. Olfactory Lobe. R.R.M. Median Right Renal
Op. A. Ophthalmic Artery. Artery.
Opt.lo. Optic Lobe. R.R.P. Posterior Right Renal
Opt. Ner. Optic Nerve. Artery.
O.P.V. Oblique Palatine Vein. Rt. A. Rectal Artery.
Orb. Orbit. R.V. Rostral Vein.
O.S. Orbital Sinus. S.C.A. Short Ciliary Artery.
O.S.A. Anastomosis between the S.M.A. Superior Mesenteric
Orbital Sinus and the Artery. :
Maxillary Vein. S.N.A. Anastomosis of the Sub-
Ov. A. Smaller Ovarian Artery. nasal Sinuses.
Ovar. L. Lett Ovary. S.N.V. Sub-nasal Vein.
Ovar. R. Right Ovary. Sp. A. First Spinal Artery.
P.A. Parietal Artery. Sp. V. Spinal Vein.
Pan. Pancreas. Spin. Cor. Spinal Cord.
Par. Parotid Gland. Spl. Spleen.
Par. A. Anastomosis between a S.P.L. Lett Supra-renal Portal
branch of the Anterior Vein.
and a branch of the 8.P.R. Right Supra-renal Portal
Posterior Parotid Gland Vein.
Arteries. S.R.L. Left Supra-renal Vein.
Par. A.A. Anterior Parotid Gland S.R.R. Right Supra-renal Vein.
Artery. Sup. Lab. Gl. Superior Labial Gland.
Par. P.A. Posterior Parotid Gland Sup. Ren. L. Lett Supra-renal body.
and Superior Maxillary | Sup. Rev. R. Right Supra-renal body.
Artery. Th. A. Thyroid Artery.
Par. V. Vein from Parotid Gland Thy. Gl. Thyroid Gland.
and also from neigh- Ton. Mus. Tongue Muscle.
bouring muscles. Ton. Sh. Tongue Sheath.
P.C. Primary Carotid Artery. V. Ventricle.
P.C.V. Post-Caval Vein. V.A. Vertebral Artery.

Prats LXX.

Fig. 1. Diagram of the Heart and Blood-vesseis in the anterior part of Tropidonotus
natrix. The two main arteries and veins on the ventral side of-the head
are indicated, but their precise distribution is dealt with on Plate LX XII.
Between the lines X.X. and Y,Y.a portion of the circulatory system is
omitted, as the relations of the Hepatic and Pulmonary Vessels and the
Dorsal Aorta are similar throughout the whole length of the liver. The
part omitted is about the same length as the distance from the thyroid
gland to the line X.X. For the purposes of diagram the liver is repre-
sented as pulled out to the right of the animal. ‘he positions of the Liver
and Lungs are indicated by simple outline and those of the Gall-bladder,
Spleen, and Pancreas by shaded areas.

Fig. 2. Diagram of the posterior Blood-vessels in a female Tropidonotus natrix.
The positions of the kidneys and ovaries are indicated by simple outline
and those of the supra-renal bodies by shaded areas. For the purposes of
diagram the intestine is represented as pulled out to the right of the
animal, the fat-bodies as external to the oviducts, and the oviducts as
external to the ureters. Owing to the fact that the anterior end of the
Fat-body is supplied from theright ovarian artery, while a little further

ON THE COURTSHIP OF TITE REDSHANK. 647

back it is fed from the left ovarian artery, if 1s necessary to divide the
Fat-body between the two gonads. Thus it comes that the anterior part
of the Fat-body is represented on the right side of the animal and the
posterior part on the left. As the Anterior Abdominal Vein runs in the
Fat-body it is also divided and is so represented in the diagram, in which
its two ends are joined by a dotted line.

Prate LXXI.
Diagrams of the Blood-vessels in the head of Tropidonotus natria.

Fig. 3. Diagram of the Cranial Arteries seen on the ventral surface of the brain.
The brain is represented as removed from the skull leaving behind,
however, the Pituitary Body.

Fig. 4. Diagram of the arteries of the head seen from the dorsal side. The brain is
removed, but some of the cranial arteriesare left behind. The positions of
the glands of the head are indicated by shaded areas.

Prate LXXII.

Vig. 5. Diagram of the vessels on the ventral side of the lower jaw after the removal
of the superficial muscles.

Vig. 6. Diagram of the Veins and Sinuses in the head seen from the dorsal side.
The deeper vessels are represented in lighter shading and with a dotted
outline. The position of the brain is indicated in simple outline, while tha
positions of the glands are indicated by shaded areas, For the purposes of
diagrem the Superior Palpebral Vein is omitted. It runs from the upper
anterior part of the Orbital Sinus above the Maxillary Vein and joins the
Inferior Palpebral Vem as the latter enters the Orbital Sinus. In dis-
secting out the vessels of the orbit it is almost always removed with the
upper eyelid.

34. A First Account of the Courtship of the Redshank
(Totanus calidris Li.). By Juntan 8. Huxuey, Lecturer
of Balliol College, Oxford.

[Received February 2, 1912: Read April 23, 1912.]

InpEx.

Page
TeAnitroductionn |, 208s aan ee eee ee eee aN GIT
De TO Cali tye’ y:.unageee es meeRee coat e eee ee ee eee SE GAS
Sa ihe Countshipsproperntesn sspears eee ee eee eee ee mG

4. Other habits of the Pairmg-Season ;
(@)\ Whe Wove= fic litle same earn eete a ashi neae cael Goll
(6) The Combats of the Males ...........0..0...0..0..-.. 652
(c) Calling from a conspicuous perch —............... 632
5: WISCUSSIONE » cdassccemcees sian cee cee cE ae OOS,

1. IytRropucttIon.

While staying last spring in a lonely corner of North Wales
it was my good fortune to come across a number of rare and
interesting birds. But great as was the pleasure of seeing, for
the first time, such comparatively uncommon species as the Grey
Plover and Black-tailed Godwit, it was far surpassed by that of
being able to study, under the most favourable conditions, the
natural behaviour and home life of some of the commoner shore-
birds. Of these I was particularly fortunate with the Redshank,

648 MR. J. S. HUXLEY ON THE

and was able to see the whole of the courtship and pairing.
When I say the whole, I do not mean that I saw every detail,
nor that every detail 1 saw is clear to me, But I mean that
IT know what is the general course of events, and can interpret
the birds’ behaviour more or less consistently.

On returning to civilization and libraries, to my surprise I
could find very little on the subject: the observations recorded
were either fragmentary or inaccurate. It was not until most
of this paper was written that I discovered a fairly complete
account by Selous *. This had remained undiscovered owing to
the absence of any reference to Redshanks in the title of the
paper or in the index of the volume.

I have thought it worth while to publish my observations,
however, since they differ in several points from those of Selous.
Meanwhile, I fully realize their incompleteness, and recognize
that they cannot as yet be properly used in any general discussion
of the theory of sexual selection. I hope to continue my own
observations when opportunity offers, but venture to publish
this general outline at once as a stimulus to other bird-watchers
and naturalists.

2, Locatity, ete,

Before passing on to the birds’ actions I must first just
mention the theatre where I saw them played. This was part of
a small estuary in the northern half of Cardigan Bay: an arm
runs out on one side at right angles to the river, thus giving
during high spring tides a land-locked sheet of water nearly a
mile long and half a mile wide; during neaps, even high tide
failed to cover it. Numbers of Redshanks and other birds fre-
quented this expanse, and especially its head or most landward
end, where they were close to a thick bed of reeds and tussocks ;
the mud here was scarcely ever covered by the tide, though kept
always moist by a little stream.

At one side of the head was a low ridge of grass-covered
dunes, about five feet high and thirty or forty | feet long, with level
ground behind them. Thus, by crawling over the flat, I could
get up to the dunes into an excellent position for viewing the
whole top of the bay ; every bird was easily seen against the wet
mud. So I kept watch with the naked eye until some disturb-
ance or unusual behaviour attracted attention ; then, being armed
with a telescope magnifying 30 diameters (for the loan of which
I have to thank my brother, Mr. N. T. Huxley), I focussed this
on the spot, and could see the minutest details of attitude and
behaviour in the nearer birds, and even in those on the far side
of the bay could quite well interpret what I saw. I made a
number of notes on the spot, and usually within twenty-four
hours embodied what I had seen the day before in a letter to
an ornithological friend.

* WH. Selous, “ Observations tending to throw light on the Question of Sexual
Selection in birds,” etc., Part 1: Zoologist (4) x., 1906, pp. 201-219.

COURTSHIP OF THE REDSHANK 649

3. THe CouRTSHIP PROPER.

I will begin with an account of the typical course of the
courtship and pairing, such as I have seen repeated, with but
slight variations, a considerable number of times.

Among the forty or fifty birds that usually would be quietly

feeding on the flats, walking or running in short starts from
mouthful to mouthful, a disturbance would every now and then
be visible—two birds running, one pursuing the other. 'These
two are cock and hen. A cock takes a fancy to one of the
hens, leaves his feeding, and starts running towards her. She at
once runs away from him, and there ensues a regular game of
follow-my-leader. The hen never goes far in a straight line;
she usually runs in a series of curves, often doubling sharply
back, and sometimes describing a complete circle or even a figure
of eight. Where she goes the cock goes after her, following
exactly, but some yards behind. The couple would be ridieulous
enough in their devious course, with heads somewhat down and
quick-moving legs—rather the action of a fast-trotting horse—
but the attitude of the cock adds to the effect: his eyes being set
on the sides of his head, his neck has to be stretched stiff out and
markedly sideways (atan angle of at least twenty degrees with the
line of his body), in order to keep the hen in sight. In addition
he spreads his pure white tail, so that you see half the fan of it on
either side of the tips of the folded wings; but whether the hen
so far ahead can see anything of this I do not know.
_ This pursuit goes on often for quite a long time, the birds
covering maybe a quarter of a mile. The hen usually flies away,
leaving the cock disconsolate, but sometimes she will consent to
inaugurate the second stage of the courtship, in which she is
able to inspect the suitor more closely. This she does by suddenly
coming to a dead stop. The cock then perhaps runs a yard or
two further, but soon he too stops, and begins his part of the
second stage of the courtship.

The first stage was almost mere pursuit: the second is pure
display. He first unfolds his wings and raises them right above
his back, so as to expose their conspicuous under-surface of pure
white somewhat clouded or barred with grey. Then, fluttering
them tremulously, but keeping them raised all the time, he
advances very, very slowly towards the hen, lifting his feet high
in the air, and often putting them down scarcely in advance of
where they were before. To the human eye the whole action
seems the expression of eager excitement tempered by uncer-
tainty, and that, presumably, is what the bird is actually feeling.
Meanwhile, as he steps on, he stretches his neck a little forward,
opens his mouth, and gives utterance to a single continuous note,
which is changed into a long roll or rattle by the quick vibration
of the lower mandible. The sound is quite like that of a Night-
jar, but higher, and without any of the little breaks in the pitch
of the note. So he advances closer and closer, the hen usually

Proc. Zoot, Soc.—1912, No. XLITI. 43

650 MR. J. 8S. HUXLEY ON THE

remaining motionless. Again at any time during this stage she
may reject his suit by flying off, but if she is going to accept him,
she simply stays still, often without moving a muscle the whole
time.

As the cock gets closer, he gets more and more excited, vibrates
his wings more and more rapidly, at length so fast that almost
his whole weight is supported by them, though he still continues
to execute the high-stepping movements with his feet. At last,
when just behind the hen, he abandons the ground, and flutters up
on to her back, on which he half alights. The period while he
is thus on her back is the third and last stage of the courtship:
it is very short, and is of course in a sense nothing more than
getting into the proper position for the actual pairing. But it
should still be called part of the courtship, for even now the cock
is not assured of his desire. Sometimes the hen, suddenly repug-
nant, or annoyed by the series of shriller, less continuous cries
which the cock is now uttering, gives a violent jerk or sideways
twist, and shakes him forcibly off on to the ground, herself
running or flying away.

Occasionally, however, she apparently is satisfied ; she spreads
her tail diagonally, and the cock, with a quick and wonderfully
graceful motion, half supported all the time by his fluttering
wings, accomplishes the act of pairing. Then the hen gives
the same violent twist that I have just mentioned, he gets shaken
off, and they both begin quietly feeding, often side by side, but
now no longer taking the least interest in each other.

That the course of true love should run so smooth was, how-
ever, quite the exception. Though I did not keep a record of
the number of unsuccessful cocks that came under my observa-
tion, there must have been at the lowest estimate fifty of them,
while but three times did I see the courtship consummated.
Thus in something well over 90 per cent. of the cases I saw,
pairing did not take place; and this was always due to the rejec-
tion of the male by the female, as the cock, once he had started a
pursuit, never of his own accord abandoned it. Thus, though the
hen does not actively select her mate from among a bevy of
competing cocks, yet, like the modern European woman, she has
the power of saying yes or no to each individual male who
may choose (here literally, there metaphorically) to run after
her.

The hen may reject her suitor at any time during the whole
proceeding. On the one hand, I have seen her break off the
courtship after a few seconds, while on the other I have seen her
stop still after running, stand and watch the cock’s display
and gradual approach, even let him fly up on to her back, and
only then with a sudden jerk throw him off and take to flight.
Between these extremes there were, of course, all intermediates.
During the first or pursuit stage, rejection was accomplished by
the hen simply taking wing and flying off some fifty or a hundred

COURTSHIP OF THE REDSHANK. 651

yards. The cock sometimes (though not usually) flew after her,
settled, and began his running once more; but then she would
very soon fly off again, and I never saw a cock persevere after a
second such repulse. The first stage may be very short, as I
have said, owing to her rejecting her suitor at once; and it may
be short for the reverse reason, the hen stopping almost at once
to let the cock come up for inspection. Usually, however, it is of
considerable length, and in well over half the number of cases 16
ends in the rejection of the cock and the cutting short of the
courtship before the second stage is reached.

Of the minority who survived thus far, a still larger percentage
were rejected before getting to the third stage. This was most
often accomplished by the hen simply flying off, leaving her
suitor to fold his wings and pretend nothing had happened.
Sometimes, however, if more undecided, she would behave in a
curiously human way. As the cock got close she would as it
were lean away from him, and at last, giving a little quiver all
over, break into a short quick run of about four or five steps,
like a frightened horse shying across the road. Then she stood
stillagain, and, when the cock advanced again, very likely repeated
the action. Of the five or six hens I saw act thus, all as a matter
of fact at length flew off; but I have no doubt that occasionally
they make up their minds in the opposite way.

4. OTHER HABITS OF THE PAIRING-SEASON.

There are other habits of the pairing-season which call for
special remark, JI do not understand their relations to the court-
ship proper, and shall merely mention them here. They are
introduced partly for the sake of completeness and partly to show
what a number of unsolved problems still exist concerning the
habits of common birds.

(a) The Love-flight.—This is a well-known habit. A Redshank
(presumably a cock) rises up into the air and there flies in a
series of switchbacks. I will quote from my own notes on a
particular bird: ‘‘ Just before the bottom of each switchback he
gave very quick wing-flaps, almost fluttering, one would call it;
this made him start up again. He went on fluttering or flapping
till he was about half-way up, and for the rest of the up-stroke
of the switchback he soared up with the impetus he had gained.
His wings now were set back and down, his neck and head
thrown up in a beautiful proud attitude, his tail spread out.
Then he turned the angle of his wings and glided down, still in
the same attitude.” While flying thus, he gives vent to what
one may well call a song—a series of pure sweet single notes,
never uttered on other occasions. The flight may be quite short,
or may go on and on for several minutes. It is usually, I think,
gone through by single birds, but I have fairly often seen it done

43*

652, MR. J. S. HUXLEY ON THE

by one of two or one of three birds. Jf so, however, it was
hardly ever repeated more than a very few times. I have several
times seen it take place when I have frightened a bird up from
feeding.

The meaning of this habit is hard to iBecawen, and its relation
(if any) to the courtship proper is equally obscure. It is per-
formed, I believe, only in the spring, and would certainly seem
to be of the same nature as the dr umming of the Snipe or the
short soaring flight of the Wood-Pigeon.

(b) The Combats of the Males. These have been well described
by Selous (oc. cit.). I agree with his opinion that the combats
of many birds are now at least merely formal. This was well
shown in the Redshank; the birds scarcely ever touched each
other, and often did not even seem fierce. It was mostly a mere
running up and down of two birds facing each other, often with
a very formal-looking character about it.. Further observation
alone can reveal its significance.

(c) Calling from a conspicuous perch.—This is mentioned by
various writers, and is a very noticeable custom. A single bird
will settle on a gate- post, railing, notice-board, or other prominent
perch, and will sit there, moving its head from side to side, and
uttering a single note many times repeated. Usually after every
few notes there comes a short pause. Long pauses arerare. One
bird continued calling thus from the same perch for 45 minutes,
and was still going on when I had to stop watching. I have at
present no idea as to the sex of the calling bird, or as to the
purpose of the whole proceeding.

5. DIscussion.

I will confine myself in this paper to a short discussion of the
courtship proper, and will begin by comparing my observations
with those of Selous (loc. cit.); all other descriptions can here be
left out of account, owing to thei vague and fragmentary
character.

Selous, who made his observations in Holland, seems, to start
with, to have seen a greater proportion of successful courtships
than I did. His description (often hard to follow, as there is no
arrangement—only notes in chronological order) differs in the
following chief points from mine :—

1. He seems to have seen only two examples of the first stage
of the complete courtship (pp. 212, 213), which is curious, since
the follow-my-leader evolutions of my birds were most con-
spicuous.

His first stage is described as follows :—“ The male, ap-
proaching the female, ran about her twice or thrice, in so man
half-cireles, fanning his tail as he did so, and inclining his body
towards her.”.... That is to say, the hen was stationary,

COURTSHIP OF THE REDSHANK. 653

instead of leading the cock a long chase, as in the Welsh
birds.

3. Sometimes directly after pairing there was a curious set
run or antic of the couple, e.g. (p. 206) :—‘‘'T'wo Redshanks,
after pairing, run, in an excited and curious-looking manner,
over the sand, following one another.” Another time (p. 204),
(in perhaps slightly different circumstances) :—“ the birds walked,
for a little, about and very near each other, fanning out their
tails, whilst bending them inwards, so that, had their legs been
short, they would have swept the ground at intervals, as does
that of the courting pigeon.”

4. He sometimes saw a male which had been definitely rejected
begin courting the same female again after a short time, once or
twice successfully ; I never sawthishappen. (This is presumably
an error of omission on my part.)

These are the chief differences. His observations on the
second and third stages and on the pairing itself are very similar
to mine.

The differences may be ascribed either to real differences of
behaviour of Votanus calidris in different parts of its range, or
to errors or omissions of observation. Further watching alone
will reveal the truth.

Further watching too must elucidate the following points, all
of which ought to be known before a full discussion of the facts
and of their bearing on the theory of sexual selection can take
place :—

1. How often does each bird go through the act of pairing 2

2. Is pairing promiscuous, or do birds pair for the season, or

for life ?

3. What is the relation between pairing and nest-building,

and between pairing and each act of oviposition ?

4, When does pairing begin in the spring, and for how long

is it continued 2

5. What is the relation of the love-flight, the combats, and

the calling from a perch, to the courtship proper ?

6. Does the female, who possesses all the structures used by

the male in his display, ever use them herself for ‘showing
off,” or for any other purpose (recognition signals, etc.) ?

With regard to 1, it is interesting to note that when the air
was calm the Nightjar-like note described above (p. 649) could
be heard at every hour of the night and day; on some nights
there were one or more birds giving utterance to it practically
all the time that I was listening. Now I am prepared to assert
that this call is only used during the second stage of the court-
ship, so that the number of courtships which advanced as far
as the second stage must have been very great (yet less than the
number of those which never reached the second stage at all;

cf. p. 651.)

654 MR. J. 8S. HUXLEY ON THE

The number of Redshanks near the head of the estuary (where
alone they really congregated) was never very large. I should
put fifty as an outside limit, and from observations on this and
other species (¢. g. Oyster-catchers), I believe that they are the
same individual birds day after day. Therefore, even allowing
that only a fraction of the courtships which reached the second
stage were consummated, yet the number of acts of pairing must
have been many times greater than the number of birds; pro-
bably, therefore, each bird pairs several times a day.

Tt is thus likely, as Selous says, that the performance of the
courtship will vary very much in different circumstances ; there
may be satiety or eagerness in either sex, as well as timidity and
shyness in the female, at different times. Selous adduees the
Pheasant to prove that the display of the male may be gradually
much abbreviated and scamped after the first few pairmgs.

As regards 4, I have very meagre evidence. I can only say
that pairing had begun before the 8th of April, and was still
going on very vigorously when I left Wales on the 18th. All
the other questions I must leave for the present unanswered.

Selous’ and my observations, however, are in themselves enough
to establish one important point, namely, that the actions of the
birds which lead up to each single act of pairing are explicable
only on the Daywinian theory of Sexual Selection, or on some
modification of that theory.

On the one hand, there is a very marked display by the male: the
fanning of the tail in the first two stages, the lifting and flutter-
ing of the wings, and the high-stepping with the legs in the
second stage, are all obviously calculated to show off to the best
advantage certain conspicuous markings which are usually con-
cealed, while the rattling note of the second stage is to my mind
equally to be considered as an excitant.

On the other hand, there is an equally marked power of choice
shown by the female; it is perfectly clear that if a female Red-
shank does not want to pair with any given male, he has no
possible means of forcing her to do so. He can only persuade
her, or rather attempt to persuade her (by means of his display),
and if she is very unfavourable to his suit, she can even prevent
him from doing this, by flying off directly he begins.

Thus, though the male in this particular species has the
initiative, the final decision must rest with the female.

The chief postulate of Darwinian Sexual Selection therefore
holds good in the case of the Redshank:—The females have a
power of choice, and the cocks have to go through a display
before pairing. Moreover, the one certainly seems to stand in a
causal relation to the other.

It is interesting to note, however, that although Sexual
Selection is at work, yet it has not produced any appreciable

difference between the sexes. This can:be explained in one of
three ways.

COURTSHIP OF THE REDSHANK, 655

(1) The markings used in display (ved legs, white tail, and
whitish under surface of wings) were acquired previously by the
species for some distinct purpose (e.g. as recognition or oblitera-
tion markings) or possibly accidentally, and then were turned to
account by the cock as the ‘‘ physical basis ” of his display. The
hen would of course possess the markings too; she differs from
the male in not having secondarily acquired the instinct to display
them.

(2) The markings were acquired by the cock first or primarily
as secondary sexual characters, to form the basis of his display,
but were either at once or later found to be of use in other ways:
they would then tend to be transferred to the hen as well, either
from their first beginnings, or subsequently to their definitive
development in the cock, by the operation of Natural Selection.

(3) The markings were acquired for purposes of display, while
in other respects they are neither harmful nor the reverse; the
instinct to use them for display, however, depends on a physio-
logical stimulus only present in the male sex. Then they would
tend to be transferred to the female sex, for we generally find
that the two sexes resemble each other unless there is some
definite reason for their differing. It appears to be both more
primitive and easier for hereditary characters to be transmitted
equally to both sexes.

It is at present very hard to decide between these possibilities.
Of the three, the last appears the least probable. What does
emerge clearly, however, is that in considering the facts of Sexual
Selection, as so often elsewhere, we must be careful not to isolate
structure from function. When we speak of secondary sexual
characters, we usually think of structures only. In reality the real
character is the structure plus the instinct to use the structure,
for it is the use of the structure which alone has any significance
for the species: it is that which constitutes a unity, 1t is that
which has been really acquired by the species. For purposes of
convenience we separate it into two components—structure and
function; but in any question of its origin and history we must
always be careful to think of it asa whole. To take a concrete
example: if it were proved (as is probable) that the female Red-
shank never used her white tail, etc. for purposes of display, we
should be justified in saying that the Redshank showed secondary
sexual characters—these characters being the various actions of
display found in the male, and in the male alone.

This point of view will perhaps help to make more intelligible
the various cases which have been described where the sexes are
alike in plumage, but the male alone goes through a display.

656 MRS. E. W. SEXTON ON

35. Some Brackish-water Amphipcda from the mouths of the
Weser and the Elbe, and from the Baltic*. By H. W.
Sexton, Marine Biological Laboratory, Plymouth f.

[Received March 1, 1912: Read April 23, 1912. ]

(Plates LX XIII. & LXXIV.£)

INDEX.
Page
Variation ........... : Paar toss . 656
Systematic: New Species ander Ganmaruss, .. 657

While engaged in determining the Copepoda of the harbour at
Bremerhaven, Herr W. Klie met with certain Gammaridea, some
of which were not easily referable to known forms. Through the
intervention of a friend these were submitted to me for examina-
tion, and the present paper contains the result, with notes on
some other collections of brackish-water Gammarus. There
proved to be three species in Herr Klie’s collection, one of
them (Leptocheirus pilosus Zaddach) already known, the other
two apparently new, which I described. But before the descrip-
tion was published I received a paper from Dr. Vanhoffen, which
contained the description and figures of one of these species
under the name of Corophiwm lacustre. I have therefore in-
cluded here only a few notes on this species. Dr. Vanhoffen
kindly allowed me to examine his specimens of Amphipoda from
the Frische Haff, and it is interesting to note that the three
species, Leptocheirus pilosus Zadd., Corophiwm lacustre Vanhoften,
and Gammarus zaddachi, sp. n., are present in both his and Herr
Klie’s collections. All three appear to flourish equally well in
absolutely fresh water as in the brackish water of river estuaries
and harbour basins.

The classification of the Amphipoda is rendered exceedingly
difficult by the changes or modifications resulting from each
successive monlt. We know practically nothing yet of the
factors influencing the development of any given species, except
those of growth amd sew. The modifications caused by these two
alone are responsible for an excessive multiplication of synonyms;
in some cases, /assa for instance, almost every moult has been
given a different specific and sometimes a different generic
name. by different observers. But in the species of Gammarus

* (Since this paper was communicated to the Society, under the title “Some
Amphipoda from Bremerhaven,” the author received some large collections of the
Gammarus described, which showed remarkable differences between those living in
salt and in fresh water. These facts have been included in the text, and the title
has been slightly modified. |—Ep. P. Z. S.

ut Communicated by the Rev. T. R. R. Sresprne, M.A., F.R.S., F.Z.S.

¢ For explanation of the Plates see p. 665.

PZ.S.1912, Pl. LAXMI.

Hoth sc, et imp.

E.W.Sexton del,
AMPHIPODA FROM BREMERHAVEN.

\

Io) iS SILO IL, ILO,

f

==

SSS
SSS

—

E,W.Sexton del,

Huth sc.et imp.
AMPHIPODA FROM BREMERHAVEN,

SOME BRACKISH-WATER AMPHIPOD2. 657.

described below, certain marked modifications occur which cannot
be referred to either of these influences. They would appear
to be caused by the animal’s environment, according as it lives
in fresh water or in salt. The difference in appearance between
a typical adult freshwater specimen and a typical marine or
brackish-water one is so extreme as to suggest their belonging
to distinct species, but structurally they are identical, and
intermediate forms are very common (see below, “ Moorflether
Concave”), apparently varying with the degree of salinity, though
on this point I cannot as yet speak definitely. Experiments
have been instituted at the Laboratory here with an allied species
to try and determine the question of the effect of salinity on
the animal and the length of time for such effect to become
evident.

GAMMARUS ZADDACHI, sp. n. (Pls. LXXIII. & LXXIV.
figs. 1-12.)

=1844. Gammarus locusta Fabr. ?, Zaddach, Syn. Crust. Pruss.
p. 4.

=1878. Gammarus locusta Fabr., Zaddach, Die Meeres Fauna
an der preussischen Kiiste, pp. 26-32.

= 1886. Gammarus pulex Kraepelin, Die Fauna der Hamburger
Wasserleitung. Abhand. Geb. d. Naturw. Verein in Hamburg,
Istobsap-ey dalle

=1907. Gammarus pulex Volk, Mitteil. biol. Elbeuntersuch-
ung. Naturh. Museums in Hamburg. Verhandl. Naturw. Ver.
Hamburg.

=1911. Gammarus locusta L., Vanhoffen, Beitriige z. Kennt.
d. Brackwasserfauna im Frischen Haff. Sitzung. d. Gesellsch.
naturl. Freunde, Beriin, 1911, no. 9.

It is with reluctance that I have felt myself obliged to institute
a new species in the already overcrowded and confused genus
Gammarus. The species of Gammarus are difficult to separate
except by the secondary sexual characters of the adult males;
the immature of all the species are practically indistinguishable .
from each other, and even the females are not easy to differen-
tiate. The species now under discussion has been frequently
confounded with others, the freshwater form with G. pulex, and
the marine and brackish-water form with G. locusta and G. due-
benit; but it can be distinguished from them by a glance at the
antennee, both of which are characterised by clusters of long out-
standing hairs, and by the form of the 4th sideplate and the
3rd uropod.

The brackish-water specimens are characterised by their slender-
ness and transparency, and the tenuity of their epidermis, while
the freshwater ones are broad, very robustly built, the epidermis
thick, strong and opaque, the basal joints of the hinder pereo-
pods narrower especially in the old males, and with a much
denser supply of. the long fine hairs developed on the antenne,
the peropods, particularly the hinder ones, the pleon, the 3rd

658 MRS. E. W. SEXTON ON

uropods, and the telson ; in some cases these long hairs appear to
replace the spines of the brackish-water form, e.g. those inset on
the inferior margins of the 2nd and 3rd pleon epimera and the
posterior margins of the 4th and 5th persopods.

The finest brackish-water specimens I have seen come from
the Konigsberg Collection, taken at Rauschen and Zoppot, the
largest freshwater ones from the Hamburg “ Wasserleitung.”
These are larger than any of the others examined by me, probably
owing to the more protected life they lead in the underground
pipes, and the rich and constant supply of food.

The collections I have examined are as follows, with the names
of those to whom I am greatly indebted for the opportunity of so
doing :— .

1. Konigsberg Museum: Dr. Lithe. Four tubes containing
Zaddach’s specimens and others from Danzig Bucht, Zoppot,
Koliebk, Redlan, and Rauschen, in all about 125. Most of the
specimens are of the typical brackish-water form, but in one
sample from Danzig in which a specimen of the freshwater
G. pulex was also found, many of the animals were ‘intermediate ”
forms, more solidly built and with the long fine hairs developed
in great numbers.

2. Frisches Haff: Dr. Vanhoften. Two tubes labelled ‘‘ Gam-
marus locusta L. Frisches Haff bei Pillau Ostpreussen. 20.ix. 1911.
Vanhoffen. 5.” One tube contained 16 specimens, 2—5:°5 mm. in
length. It is interesting to note a few young Leptocheirus
pilosus with the young G. zaddachi; these two species, with
Corophium volutator and C'. lacustre, seem to flourish equally well
in fresh or brackish water and are frequently recorded together.
There were 8 specimens of G. zaddachi in the second tube, 3°5—
10 mm. long, all of the brackish-water form ; the largest, however,
approaching the freshwater form, strongly built and opaque.

3. Mouth of the Oder: Dr. Vanhoffen. One tube, “Gammarus
locusta L. Dievenow Odermundung Pommern 1889. Hilgen-
dorf. 5.” 26 specimens, 1 young male G. locusta and 25 G. zad-
dachi exactly like those described from Pillau.

4, Mouth of the Elbe, from Schulau to Gauert, together with a
collection from the Hamburg Wasserleitung; Dr. Steinhaus. This
is a most interesting series taken at regular stations in the Elbe
(see Volk, with map of the district), containing in all about 500
specimens. Commencing with the stations nearest the mouth of
the river, where the water has nearly the full salinity of the
North Sea, the details are as follows :—

““Suder-Elbe siidl. Blankanese. Grund. 11.vi.00.” 11 speci-
mens 4—10 mm.

“Hafen siidl. Nienstedten. Grund. 2.vi.00.” 385 specimens
1-5-10 mm.

‘““ Wafen sudl. Nienstedten. 11.vi.00.” 3 specimens, largest a
female 11 mm.

‘““Linkes Hlb-Ufer siidl. Nienstedten. 2.ix.00.” 3 specimens,
largest a female 8 mm.

SOME BRACKISH-WATER AMPHIPODA. 659

“ Altona. Ponton. 9.vii.00.” 107 specimens, all rather small,
averaging 8-9 mm. in length; the eyes in these have reddish-
brown pigment.

“ Altona. Duc d’Alben. 1.x.00.” 1 specimen, a female.

All these belong indubitably to the brackish-water type. The
very young animals differ slightly from the adult, the eyes are
round, the peduncles of the antenne are about equal in length,
the pleon clusters are represented each by a single spine, and the
rami of the third uropod are much more unequal than in the
adult. Some of the specimens, especially those from Blankanese,
are more setose than the others.

“Hamburg.” The “ Fleet” referred to is one of the canals
which run through the inner town. These canals are mostly
dependent on the tide, with mud-banks exposed at low tide.
‘“« Grasbrokhafen” and “ Indiahafen ” are blind alleys as it were
opening out of the Harbour basin ; they are deep and broad, the
bottom consisting of clay, sand and mud, with organic detritus
deposits providing food for a rich animal life.

“ Bleet 1. Grund. 25.vi.00.” 1 specimen, a male 11 mm.

* Fleet 1. Krotzprobe. 23.vii.00.” 1 specimen, a male, with
exceptionally large eyes.

‘“¢ Grasbrookhafen. Grund. 19.xi.00.” 2 specimens 19-20 mm.

“Grasbrookhafen. Due d’Alben. 20.vi.00.” 18 specimens
mostly small, the largest 9-10 mm.

“ Tndiahafen. Ponton. Oberfliiche. 22.v.00.” 77 specimens, all
very young.

The specimens from the “ Fleet” samples show intermediate
characters, forming the link between the two extremes of the
species. They are more solid in appearance than the brackish-
water form, but the chitin is not as thick as in the freshwater
animal. The spines and long hairs vary greatly in number, and
the narrowing of the basal joints of the hinder pereopods also
appears variable. Of the “ Grasbrookhafen ” specimens the two
from the bottom are typical freshwater ones; the eighteen spe-
cimens from the ‘“ Duc d’Alben” (piles driven in and used for
making ships fast) are all delicate in appearance, the largest
showing intermediate characters. Some of the small ones, 3-4 mm.
long, are extremely slender and compressed, almost thread-like,
and with very few hairs. The “ Indiahafen” specimens are all
very small, the brackish-water form.

“ Alte Dove Elbe and Moorflether Concave.” Volk says of
these that, taken together, they form a quiet bay with the bio-
logical characters of a backwater (“ Altwasser”), but influenced
to some degree by the tide. These samples are the most interest-
ing of the series ; in some both the fresh- and the brackish-water
forms occur together, with the ‘intermediate ” form.

‘“‘Moorflether Concave. Grund I. 23.vii.00.” 1 specimen, a
female 9 mm., with the delicate appearance of the brackish-water
form, but with dense clusters of long hairs on antenne, per:zeo-
pods, and telson.

660 i MRS. E. W. SEXTON ON

‘“‘Moorflether Concave. Oberfliiche II. .23.vii.00.” 11 speci-
mens; the largest, males, measured 10-11 mm.,; all the brackish-
water form.

‘“‘Moorflether Concave. 23.vii1.00.” 129 specimens; largest
15mm. Freshwater, brackish, and “intermediate” forms, and
one specimen of the exceedingly narrow, thread-like form described
from the Grasbrookhafen sample.

‘“‘ Moorflether Concave. Ponton. 23.vii.00.” 5 specimens, the
largest, a male of 14 mm., shows the freshwater characters ; the
others are of the brackish-water type.

‘“‘Moorflether Concave. 24.i1x.00.” 37 specimens, brackish-
water form, largest 12 mm.

The three following samples are all unmistakable brackish-
water form. The “ Prielen” are little channels cut in the banks
of the river; the current is less strong, and the growth of vege-

tation and animal life richer than in the Elbe.

“Dove Elbe. Oberfl. 2.vii.00.” 6 specimens, largest about
8 mm.

“JT. P.r. O. 7.vi.00” (= Rechtsseitiger Elbpriel am Spaden-
land, Oberfliiche). 6 specimens, largest 10 mm.

«T, P.r. O. 2.vii.00.” 14 specimens, largest 8 mm.

The remaining samples are all fine examples of the typical
freshwater form.

‘“‘Kaltenhofe. ‘ Wasserprobe aus Filter No. 16 entnommen
am 6. Dec. 1894.’” 3 specimens, the largest 22 mm.

‘‘ Hamburger Wasserleitung.” 5 specimens, 18-20 mm. long.

“Hamburger Wasserleitung. Samuelson.” 71 specimens.
Dr. Kraepelin has given a very interesting account of the condi-
tions of animal life in the underground “ Wasserleitung ” of Ham-
burg, conditions much more favourable than in the Elbe, owing
to the greater abundance of food, the protection from many
enemies, and the lesser temperature variations. He mentions
this species under the name of G. pulea as second only in numbers
to Asellus aquaticus among the Edriophthalma met with in the
series of samples taken.

5. Bremerhaven: Herr Kiie. One tube “ Alter Hafen. x.11.”
10 specimens, the largest a female 9 mm., from which several of
the figures are drawn. This sample contains one of the thread-
like form described above. . These Bremerhaven specimens, which
come from salter water than the preceding, have scarcely any of
the long fine hairs developed, which so strikingly characterise the
freshwater animals.

6. Irish Lakes: British Museum (coll. by Major Trevelyan).

“Lough Nadarragh.” 4 large specimens, male and female,
about 18 mm, 1n length. Freshwater form, exactly like those
from the Hamburg Wasserleitung.

‘* Lough Keenaghan,” 2 specimens, male and female G. pulea.

‘f Lough Keenaghan.” 1 specimen, a female, freshwater form,
G', zaddachi.

SOME BRACKISH-WATER AMPHIPODA. 661

“Tough Awaddy and Tullynabour.” 4 specimens, three males
and one female, largest 14-15 mm. Freshwater form.

* Lough Erne.” 4 specimens; two G’. zaddachi, male and female,
freshwater form; and two G. pulew.

The first to observe the species now under discussion was
Zaddach, after whom I have accordingly named it.

In 1844 he described it under the name of “ Gammarus locusta
Fabr. 2,” but pointed out the characters in which it differed from
Milne-Edwards’s deseription, adding that if the latter were correct,
his (Zaddach’s) species must be regarded as new. In his later
work he gives a more detailed description and figures of the species,
again, however, emphasizing the differences in the antenne and
the 3rd uropods. Zaddach’s specimens are still preserved at
Konigsberg Museum. The G. locusta, described by Chevreux
(Bull. Soc. Zool. v. xvii. p. 141) as inhabiting the Loire and
the mouths of rivers of Corsica and Provence, is very probably
the species here described.

The principal points of difference between this newly-established
species and G. locusta lie in the antenne, in the proportions of the
peduncle-joints of antenna I, and in the sete, in the gnathopoda,
in the 4th side-plate, in the armature of the pleon, and in the
3rd uropod.

It differs from G. duebenii, specimens of which were kindly
sent to me by Professor Sars for comparison, in the following
points:—The upper antenna of G'. duwebenii is only sparsely fur-
nished with hairs, and has none of the regular graduated clusters
so characteristic of G'. zaddachi ; the hands of gnaths. 1 and 2 are
smaller, and practically subequal in size, in G. zaddachi the hand
of gnath. 2 is decidedly larger than that of gnath. 1 ; in the adult
G. duebenit the basal joints of pereeopods 3, 4, and 5 are all
expanded, with the hind corners free, while in G. zaddachi these
joints are hardly expanded at all in the male, and only the 3rd
has the hind corner free ;.in the largest G'. duebenti the 1st and
2nd uropods and the telson reach to the level of the peduncle of
uropod 3, and the outer ramus of urop. 3 is twice the length of
the outer ramus of urop. | (in the smaller specimens the propor-
tions are as figured by Sars, Crust. Norway, vol. i. ee 177); in
G. zaddachi urop. | is considerably longer than urop. 2, and the
outer ramus of urop. 3 only half as long again as that of urop. 1 ;
the telson in G. dwebenii is shorter, broader, and more spinose.

From G. pulex it is distinguished at once by the shape of the
eye, small and rounded in G. puwlex, large and reniform in
G. zaddachi; the antenne, gnathopods, 4th sideplate, pleon
armature, and the 3rd uropods also differ.

The Bremerhaven and Frisches Haff Specimens are all young,
the largest, a female figured on Pl. LX XITII., measuring 9 mm.
The largest male in the K dnigsberg collection measures 18°5 mm. ;
the Remalies are much smaller and broader, 12-13 mm., but none

662 MRS. E. W. SEXTON ON

of them are ovigerous. The largest Hamburg specimen measures
22 mm.

The body is slender and compressed, the brackish-water speci-
mens much more delicate in appearance than the freshwater
ones.

Head (fig. 1) about equal in length to the first two perzon-seg-
ments, in G. locusta it is distinctly shorter ; lateral lobes obliquely
truncate, upper angle usually subacute in large specimens, and
obtusely rounded in smaller ones, rounded below, sinus small.

Side-plates smaller than in G’. locusta, about as deep as the corre-
sponding segments, 1—4 serrated at the anterior angle. The 4th,
which forms one of the distinguishing characters of the species,
is as broad as deep, with the posterior expansion short and round-
ing into the inferior margin. In G‘. locusta the posterior expansion
is very deep, with the hind margin straight and much more
serrated.

Pleon-segments 1-3 with the postero-lateral corners acutely
produced, with only two or three setules inset on the hind
margin. Segments 4—6 each with three groups of spines. The
number of spines is very variable: generally speaking, the median
group has two spines, and the lateral groups three each set fan-
wise, and accompanied asa rule by long fine hairs, more numerous
in the freshwater animals; but many of the larger animals have
three spines in the median group of the 4th segment, and two
in the 5th and 6th, and four spines in the lateral groups of
the 4th and 5th segments, and three in those of the 6th. The
number varies, however, even in animals of the same size.
‘The spines are longer than in G. locusta, and inset at a rather
different level ; the dorsal groups are only slightly raised, whereas
in G. locusta they are elevated and prominent.

Eyes large, reniform, very dark; outer row of ommatidia
colourless,

Antenna 1 not quite half as long as the body, much more
setose in the male than in the female. The peduncle is longer
than in any other known species of Gammarus, nearly equalling
the peduncle of ant. 2 in length in the adult, and quite equal to
it in the young animal. It is furnished inferiorly with the out-
standing clusters of stiff setee characteristic of the species, some
of the sete in each cluster extending far beyond the rest, and
graduating in length to the distal end of each joint. The
Ist joint is longer than the 2nd, but not as long as the 2nd
and 3rd taken together. The primary flagellum is 18-jointed
in the largest Bremerhaven specimen, with 5 joints in the
accessory flagellum; the largest male from Rauschen had 33
jomts in the primary, and 8 in the accessory, the smaller
animals averaged 23-27 in the one, and 4-6 in the other.
Zaddach gives the range as from 25-35 in the primary, and 5-9
in the accessory. EKach jointin the primary flagellum from about
the 5th carries a small stalked sensory filament in addition to the
small sete, and a long seta on alternate joints. In G. locusta the

SOME BRACKISH-WATER AMPHIPODA. 663.

peduncle is noticeably shorter than that of ant. 2, extending only
to the distal end of the 4th joint of the latter, instead of to half
the length of the 5th, as in our species; both the peduncle and
flagella are very sparsely provided with sete, and the accessory
flagellum is much longer, 13—14-jointed in the male, with about
A7 joints in the primary.

Antenna 2 shorter; 4th joint of the peduncle slightly shorter
than the 5th, both furnished inferiorly with the characteristic
clusters of sete. The flagellum is about as long as the 4th and
5th joints of the peduncle combined, 11-jointed in the largest
Bremerhaven specimen, 15-19-jointed in the large males from
Rauschen ; in both sexes calceole occur on the proximal joints
and clusters of setz similar to those of the peduncle.

In the male of G. locusta, and to a less degree in the female
also, both the peduncle and flagellum are clothed with dense tufts
of long fine delicate sete. The almost glabrous first antenna
and the exceedingly hairy second antenna of (. locusta are sufti-
cient to distinguish the species from G. zaddachi at a glance.

Gnathopoda.—Vhe difference between the gnathopods of the
males of the two species will be better seen by referring to
Pl. LX XIII. figs. 2-5. In G. zaddachi (fig. 2) they are much
broader in proportion to their length, with the palm less oblique
and the palmar margin crenulate. In G. locusta (fig. 3) the
palmar margin is sinuous, the clusters of sete on the hind margin
are much denser, the sets, as in ant. 2, are very fine and long,
and the palmar spines are of different structure.

In the female (figs. 4&5) the second gnathopod only carries a
spine midway on the palm; the finger is much more pointed than
in the male.

Perwopoda.—The proportions differ from G. locusta, the 4th
joint being much shorter in: proportion and broader, and the 6th
joint, except in pereeopod 1, always exceeds it in length. The
2nd joint in pereopod 3 has the lower hind corner produced, sub-
acute; this joint in pereopods 4 and 5 is narrowed distally, the
hind corners not free, but with one or two strong spines inset at
the angle (see fig. 10). In most of the large males the basal joints
are long, very narrow, and scarcely at all expanded posteriorly; in
the female and immature specimens they are always shorter and
more expanded than in the adult male. Generally speaking the
posterior margins of the basal joints and the inferior margins of
the 2nd and 3rd pleon segments are beset with small spines in
the brackish-water form, and with long setiform spines or setz
in the freshwater form, but this again is not an invariable rule,
some specimens having spines on the hinder perzeopods and setz
on the pleon epimera or vice versd. ‘The hinder pereeopods in the
freshwater animals are covered with long fine hairs set in clusters
with the spines ; these hairs are not much developed in the animals
from salt water, but, on the other hand, they are provided with
more spines.

The wropoda are more slender than in G. locusta. In uropod 3

664 MRS; E. W: SEXTON ON

(fig. 11) the inner ramus is about three-quarters the length of
the Ist joint of the outer ramus; in G. locusta it reaches to the
distal end of the 1st joint, and both rami are more richly furnished
with spines and plumose sete.

Cororuium LAcustTRE Vanhoffen, 1911. (Pl. LXXIV. figs. 13-
ds))

1911. Beitr. z. Kennt. d. Brackwasserfauna im Frischen Haff.

The largest male measured 3° mm.; the largest female
5°) mm.

Body broad, hardly at all compressed. Side-plate 1 with three
very long plumose sete at the anterior angle.

Pleon-segments 4-6 coalesced, strongly resembling C. acutwm
Chev.

Head. The rostrum in the male is much produced, apex sub-
acute; not so prominent in the female, apex obtuse. Lateral
lobes rounded, produced, but not to the level of the rostrum.

Eyes large for the genus, irregularly round, pigment black.

Antenna 1—The peduncle in the male is clothed with long
hairs; Ist joint nearly as long as the 2nd and 3rd combined ; in
the female (Pl. LX XIV. fig. 13) the first joint is longer than the
2nd and 3rd taken together ; in both, it is produced below apically
and tipped with one short stout spine; 3rd joint much shorter
than the 2nd. ‘The flagellum is shorter than the peduncle, about
9-jointed in the male and 7-jointed im the female; terminal
joint rudimentary, the last three joints each carry a cluster of
sensory filaments.

Antenna 2 very stout; in the male the large 4th joint of the

peduncle carries two apical teeth, the outer one much the larger,
recurved. The 5th joint and the flagellum are furnished in-
feriorly with clusters of very long. fine hairs. The flagellum
consists of two joints, the small terminal one with two strong
spines. In the female (fig. 14) the 3rd joint of the peduncle is
produced apically into a small process tipped with a spine; the
Ath joint has a similar process midway on the inferior margin,
and a long apical tooth with a spine inset at the tip: 5th joint
almost as long as the 4th. The flagellum as in the male, but
showing one joint-division more; one spine on the terminal
joint.
Gnathopod 1 (figs. 15 & 16).—5th joint longer than the 6th,
tapering distally ; 6th as wide at the palm as at the base, but
with hind margin lightly curved; front margin with transverse
rows of sete; the palm convex, serrulate, fringed with six small
spines; the finger curved, acute, inner margin serrulate, with
three or four setules inset subapically.

Gnathopod 2 (fig. 17) much as in C. triaenonyx Stebbing. The
2nd joint is expanded, more so in the male than in the female ;
6th joint long, tapering distally. The finger is tridentate ; teeth
upturned, the 3rd the largest, with small stiff sete inset between,

SOME BRACKISH-WATER AMPHIPODA. 665

Pereopods 1 and 2 stoutly built; the posterior margin of the
2nd joint and both margins of the 4th are fringed with long fine
sete in the male, but carry only a few sete at intervals in the
female. The finger has the glandular aperture opening at
the tip.

The hinder percopods are alike in both sexes. The 5th is
long and slender, about twice the length of the 3rd. The fingers
of pereeopods 3 and 4 are strongly recurved, that of perzeopod 5
falciform. The posterior margin of the 3rd is furnished with a
few simple sete, that of the 4th with plumose sets, while both
margins of the 5th are densely fringed with long and _ short
plumose sete ; the 6th joint of this pereeopod carries two or three
clusters of remarkably long setz posteriorly.

The incubatory lamelle are long, narrow, and fringed with
very long hairs, that of perzeopod 3 almost equals the perzeopod
in length.

Telscn the same in both sexes, about twice as broad as long,
the apex obtusely truncate; a pair of mobile setz inset on each
side near the base.

EXPLANATION OF THE PLATES.
PratE LXXIII.

Fig. 1. Head of Gammarus zaddachi, sp. ., young female, 9 mm. Bremerhaven.
The clusters of hairs are much denser in the males and in the older

females. x 42.

2. Hands of gnathopods 1 and 2, G. zaddachi, large male, 17 mm. Rauschen.

x 20.

3. Hands of 5 » G. locusta, large male, 17 mm. Plymouth

Sound. x 20.
4. Gnathopod1. Young female. Bremerhaven. G. zaddachi. X 42.
5. 29 2. ” bP) 39 bb] x 42.
6. Perzeopod 1. 35 > 5 5 Xx 42.
Uo 3? 2. 9 ”? ” bb) x 42.
8. 23 3. bb) bb] bb] ae) x 42.

PravTE LXXIV.

Fic. 9. Pereopod4. Youngfemale,9mm. Bremerhaven. G.zaddachi, sp.n. X 42.
x 42

”? y ” ” ” ” 23 2? 2
11. Uropod 3. 6 $3 5 - ss Gaze
12. Diagrammatic dorsal view of pleon segments 4-6 and telson, showing the
arrangement of the spine clusters; in the males and older females, par-
ticularly in the freshwater form, these clusters are always accompanied by

long fine hairs. Young female, 9 mm. x 42.
13. Antennal. Female. Corophiwm lacustre Vanhoffen. Bremerhaven. X 42.
14. yD) 2. yy) oy) 29 ot) oP) x 42.
15. Gnathopod 1. Male. - A i Br x 58.
16. Finger and palm, gnathopod 1. Male. Corophiwm lacustre Vanhoften.
Bremerhaven. x 145.

17. Gnathopod 2. Male. Corophium lacustre Vanhéften. Bremerhaven. X 58.

Proc. Zoou. Soc.—1912, No. XLIV. 44

666 MR. C. TATE REGAN ON

36. Descriptions of new Fishes of the Family Loricariidee

in the British Museum Collection. By C. Tare Rrean,
Males IM ZetSs

[Received March 5, 1912; Read April 23, 1912. |

(Plates LXXV.-LXXVII.*)

INDEX.

Systematic: Page
Plecostomus hond@, SP. Nei... cecsev ene crv eer seneereserse 666
Chetostomus palmeri, SP. De... .eeceeeeeeeeeecees avon 667
(OM UGPRATHATIS, (No 116 Bonecbooneen oon pcosonconeposceuascopcescxe (OLSV/
ONparicispinis, SWeillseece ene reer eee seer eee OO
Xenocara heterorhynchus, Sp. V..........0.e0eeeee-s ss 668
XG nuléisprnis, Spo We ene aecsceeseeeenaesaciecsee esas) OOS
Otocinclus maculipinnis, Sp. UN. ......... 02.02 serene 668
Oxyloricaria tamane, SP. Me 2.0.2 ..0.. 0.0 ee eee eee eee 669
O} UIGIEOREs BMs Wo o2e.00000000 cocanocan oHaaonooosaoseacencce HAY)
ANRACS GHARETTUBS SVs Worocooscenssoecs cossds ona sccceccsoanacee (HN)

During the last three years several Loricariid fishes have been
added to the British Museum Collection, including examples of
the following ten species new to science.

1. PiEcostomus HONDA, sp.n. (PI. LX XVI. fig. 3.)

Depth of body 43 in the length, length of head 3, Depth of
head 12 in its length, breadth 17, snout 13, eye 7, interorbital
width 21. Length of mandibular ramus 3 in interorbital width ;
jaws with about 12 teeth on each side. Snout ovate; supra-
orbital margins slightly raised ; supraoccipital convex, without
distinct ridge; temporal plates not or but feebly keeled. Scutes
spinulose, weakly carinate, 27 or 28 in a longitudinal series,
7 between dorsal and adipose fin, 13 between anal and caudal ;
supraoccipital entirely bordered posteriorly by a single scute ;
lower surface of head and abdomen with a few scattered granules
(probably covered with granuiar scales in the adult). Dorsal 17;
first ray nearly as long as head, last 3 as long; base more than
distance from tip of spine of adipose fin. AnalI 4. Pectoral
spine extending to anterior 7 of ventrals. Caudal emarginate,
the middle rays 3 as long as the outer ones, which are longer
than the head. Caudal peduncle 3 times as long as deep.
Reddish brown, with traces of darker spots on head and anterior
part of body ; dorsal red, with three series of dark spots between
the rays; caudal with alternate red and dusky bars; lower fins
reddish, with dark spots.

Two specimens, 80 mm. in total length, from Honda, Colombia
(300-400 ft.), presented in 1909 by Sir Bryan Leighton.

* For explanation of the Plates see p. 670.

PZ. S. LOI PL oxy.

A.H, Searle, del et lith.

C. PALMERL.

C. PAUCISPINIS. 3.

a.

CHAETOSTOMUS LEPTURUS.

1.

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NEW SOUTH-AMERICAN FISHES. 667

2. CH#TOSTOMUS PALMERI, sp.n. (Pl. LXXYV. fig. 3.)

Nearest to C. fischeri Steind. Depth of body 53 in the length,
length of head 34. Head a little longer than broad and nearly
twice as long as ase. Diameter of eye 64 to 8 in length of head,
interorbital width 3 to 33, length of snout 12 to 12. Length of
mandibular ramus some to interorbital wat. Interopereulum
with 2 or 3 spines. 24 scutes in a longitudinal series, 6 or 7
between dorsal and adipose fin, 11 between anal and “caudal.
Dorsal I 8; first ray 2 to ? the length of head, last 3 as long;
length of base equal to distance from posterior part of spine of
adipose fin. Anal I5. Pectoral spine just reaching base of
ventral. Caudal peduncle 23 times as long as deep. Coloration
uniform ; dorsal and caudal with or without series of spots on
the rays.

Two specimens, 75 and 95 mm. in total length, from the Rio
Tamana, Rio San Juan, Choco, $.W. Colombia, collected by
Mr. G. Palmer.

3. CHETOSTOMUS LEPTURUS, sp. n. (Pl. LXXYV. fig. 1.)

Depth of body 53 to 6 in the length, length of head 33 to 33. -
Head nearly as ‘amon as long and almost twice as long as Alea.
L touuetes of eye 8 to 94 in length of head, interorbital width
34 to 33, length of snout 12 to 12 rf Length of mandibular ramus
equal a interorbital width. Tinteropereulam with 5 to 8 spines.
25 scutes in a longitudinal series, 6 or 7 between dorsal and
adipose fin, 11 or 12 between anal and caudal. Dorsal I 8; first
ray nearly as long as head, last 2 to 4 as long; length of base
equal to or slightly more than its distance ‘trom adipose fin.
Anal I 5. Caudal obliquely emarginate. Caudal peduncle 33 to
4 times as long as deep. Coloration as in C. marginatus Regan,
from which hic species differs especially in the form of “hs
dorsal fin and the more slender tail.

Three specimens, 135 to 215 mm. in total length, from the Rio
Tamana, Rio San Juan, Choco, 8.W. Cslonmi, collected by
Mr. G. Palmer.

4, CHATOSTOMUS PAUCISPINIS, sp. n. (PI. LXXYV. fig. 2.)

_ Depth of body 5 in the length, length of head 3}. Head as
broad as long and 14 as long as deep. Diameter of eye 8 in
length of Neude interorbital width 3, length of snout 12. Inter-
operculum with 2 spines. 24 scutes in a longitudinal series,
6 between dorsal and adipose fin, 10 between anal and caudal.
Dorsal I 9; first ray a little less than 3 length of head, last 2 as
long ; length of base equal to distance from tip of spine of adipose
fin. Anal I 5. Pectoral spine extending to base of ventral.
Caudal very slightly emarginate. Caudal Tpedunele 27 times as
long as deep. Back with dark cross-bars; dorsal fin with series
of spots on the rays; tips of all the fins reddish.
A single specimen, 90 mm. in total length, from Tado, Rio
San Juan, Choco, Colombia, collected by Mr, G. Palmer.
44*

668 MR. CG. TATE REGAN ON

5. XENOCARA HETERORHYNCHUS, sp.n. (Pl. LXXVI. fig. 2.)

Depth of body 6 to 63 in the length, length of head 22 to 23.
Head longer than broad and more than twice as long as deep.
Diameter of eye 7 or 8 in length of head, length of snout 13 to
2, interorbital width 3. Length of mandibular ramus 1! to 13
in interorbital width. Snout with tentacles. Interoperculum
with about 10 spines, the longest about + the length of head.
23 or 24 scutes in a longitudinal series, 7 or 8 between dorsal
and adipose fin, 11 or 12 between anal and caudal. Dorsal I 7;
first ray 3 to 2 the length of head, last, when laid back, separated
by 3 keeled scutes from adipose fin ; base of dorsal equal to or
less than its distance from adipose fin. Anal 1I3. Pectoral spine
extending to base of ventral. Caudal obliquely truncate. Caudal
peduncle 3 times as long as deep. Fins with series of dark spots.

Two specimens (¢), 58 and 80 mm. in total length, from
Uruhuasi, Peru (4000 ft.), collected by Messrs. H. & C.
Watkins.

This species is related to X. bufoniwm, differing somewhat in
proportions and also in the much greater width of the naked
margin of the snout, into which the bony plates extend as a narrow
median prominence nearly reaching the apex of the snout.

6. XENOCARA MULTISPINIS, sp. n. (Pl. LX XVI. fig. 1.)

Depth of body about 5 in the length, length of head 27. Head
longer than broad and twice as long as deep. Diameter of eye
8 to 94 in length of head, interorbital width 22 to 24, length of
snout 1¢to 2. Length of mandibular ramus 2 in the interorbital
width. Snout with many long tentacles(¢). Interoperculum
with 20 to 25 spines, the longest 4 the length of head. 24 scutes
in a longitudinal series, 6 or 7 between dorsal and adipose fin,
10 or 11 between anal and caudal. Dorsal I 7; first ray about
3 length of head, last about = as long; base equal to distance
from posterior part of spine of adipose fin. Anal I 4. Pectoral
spine extending to anterior 4 or middle of ventral. Caudal
obliquely truncate. Caudal peduncle 24 to 22 times as long
as deep. Traces of rather large pale spots on head; dorsal and
lower fins with series of dark spots; caudal dusky.

Three specimens (3), 90 to 120 mm. in total length, from the
Humboldt and Novo Rivers, Sta. Catherina, S.E. Brazil, collected
by Mr. W. Ehrhardt.

This species is near to X. stigmaticum Kigenm., and except for
the larger size of the spots on the head might as justly be referred
to that species as those that I described as X. stigmaticum from
the Mogy-guassu River. Dr. R. von Ihering has written to me
that this locality is not 250, but only 25 miles inland of Santos.

7. OTOCINCLUS MACULIPINNIS, sp.n. (PI. LX XVII. fig. 3.)

Closely related to O. nigricauda and O. perforatus. Depth of
body about 53 in the length, length of head 23. Diameter of eye

NEW SOUTH-AMERICAN FISHES. 669

about 7 in length of head, interorbital width 2%, length of snout
2 to 21. Occipital region evenly convex, without crests. Scutes
spinulose, not carinate, 22 to 24 in a longitudinal series. Lower
surface of head with a naked area in front of clavicles ; abdomen
with 4 or 5 irregular series of plates between the lateral series.
Dorsal I 7; origin a little behind base of pelvics, but not much
nearer to end of snout than to base of caudal. AnalI5. Pectoral
spine extending beyond middle of ventral. Caudal emarginate.
Caudal peduncle 3 times as long as deep. Fins with series of
dark spots.

Four specimens, 30 to 35 mm. in total length, from the La
Plata, presented in 1909 by Herr J. Paul Arnold.

8. OXYLORICARIA TAMAN#, sp.n. (PI. LX XVII. figs. 1, la, 1b.)

Head 53 in the length. Breadth of head 14 (d) or 14 (2) in
its length, diameter of eye 7 to 8, interorbital width 31, length of
snout 13. Snout without distinct rostrum; supraorbital edges
not or scarcely raised; sides of the snout, in the male, with
bristles. 32 or 33 scutes in a longitudinal series (14-15+18) ;
lateral keels weak, obsolete anteriorly, united posteriorly. <A
naked area surrounding the lips ; abdomen posteriorly with 3 series
of plates between the lateral series; an anal plate bordered
anteriorly by 3, these 3 again by 5. Dorsal elevated, its height
+ the length of the fish; pectoral spine extending to anterior 4
of ventral; caudal emarginate, the outer rays produced. Breadth
of body at level of last anal ray 42 to 5 in the distance from
the caudal. Anterior half of dorsal blackish ; caudal blackish at
the base; pectoral blackish, pelvics more or less dusky, anal
spotted anteriorly.

Two examples (¢, 2), 225 and 200 mm. in total length, from
the Rio Tamana, Rio San Juan, Choco, 8.W. Colombia (200 ft.),
collected by Mr. G. Palmer.

9. OXYLORICARIA LEIGHTONI, sp.n. (Pl. LX XVII. fig. 2.)

Head 44 in the length. Breadth of head 12 to 14 in its length,
diameter of eye 8, interorbital width 34 to 33, length of snout 14
to 2. Snout without distinct rostrum; supraorbital edges not or
scarcely raised. 30 scutes in a longitudinal series (15+ 15); lateral
keels weak, united posteriorly. A naked area surrounding the
lips ; abdominal plates posteriorly forming 3 or 4, anteriorly 6 or 7,
irregular series between the lateral series; an anal plate bordered
anteriorly by 3. Height of dorsal equal to length of head ; pectoral
spine extending to anterior 7 of ventral; caudal emarginate, the
outer rays produced. Breadth of body at level of last anal ray
1 the distance from caudal. Fins with blackish spots; anterior
part of dorsal blackish at base and near tip.

Two specimens, 55 and 65 mm. in total length, from
Honda, Colombia (300-400 ft.), presented in 1909 by Sir Bryan
Leighton.

670 ON NEW SOUTH-AMERICAN FISHES.

10. ARGES CIRRATUS, sp. n.

Length of head 32 in the length of the fish. Interocular width
rather more than distance from eye to posterior nostril, rather
less than 4 the length of head. Nasal flap produced into a barbel
which is as long as “the diameter of the eye; maxillary barbel not
quite reaching” gill-opening; teeth of outer series of preemaxil-
laries unicuspid, acute; mandibulary teeth bicuspid. First dorsal
ray 2 the length of head ; pectoral extending to base of ventral ;

ventrals inserted below anterior part of dorsal, nearly reaching
anal. Spine of adipose fin well developed, moval connected’
by a membrane with the caudal peduncle, inserted at a distance
from caudal equal to about 4 the length of the middle rays al
that fin. Distance from tip “Of snout to origin of dorsal fin 2
in the length of the fish, from base of last anal ray to caudal se
A few large dark spots on body; a blackish bar across middle of
caudal fine

A single specimen, 24 mm. in total length, from the Rio
Condoto, Rio San Juan, S.W. Colombia, collected by Mr. G.

Palmer.

EXPLANATION OF THE PLATES.

Prate LXXV.

Fig. 1. Chetostomus lepturus.
Fig. 2. 5 paucispinis.
Fig. 3. 5 palmeri.

PrateE LXXVI.

Fig. 1. Xenocara multispinis.
Fig. 2. 3 heterorhynchus.
Fig. 3. Plecostomus honde.

Pirate LXXVII.

Fig. 1. Oxyloricaria tamane (x4).
Figs. 1a, 16. Heads of g & 9.

Fig. 2. O. leightoni.

Fig. 3. Otocinclus maculipinnis (X2).

ON EPIDERMAL SHEATHS FROM BILL OF THE KING PENGUIN. 6/71

EXHIBITIONS AND NOTICES.

April 23, 1912.

Dr. 8. F. Harmer, F.R.S8., Vice-President,
in the Chair.

The Secrerary 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 Secrerary exhibited a number of photographs of an
Hlephant Kraal in Siam which had been presented to the
Society by the Rt. Hon. Sir Cecil Clementi Smith, P.C.,
G.C.M.G.

May 7, 1912.

Prof. EK. A. Mincuin, F.R.S., Vice-President,
in the Chair.

Mr. R. I. Pocock, F.R.S8., F.LS., F.Z.S., 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. Seru-Smiru, 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 (Aptenodytes
pennanti) living in the Society’s Gardens. Mr. W. EK. de Winton
had observed the shedding of this epidermal sheath in a bird living
in the Gardens in 1898 (P. Z.S. 1898, p. 900); but although the
present specimen had been carefully watched during two suc-
cessive moults in March and October 1911 (P. Z.8. 1912, p. 60),
no sign of this process was observed. 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 the orange-coloured patches became loose and finally

672 MR. G. A. BOULENGER ON

fell off; the pieces somewhat resembled the wing-cases of a large
beetle, being semi-transparent and of a clear orange-colour.

Dr. Francis WArD, 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 in a 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.

PAPERS.

37. On a Collection of Fishes made by Mr. A. Blayney
Percival in British Hast Africa to the Hast of Lake
Baringo. By G. A. BounenceEr, F.R.S., F.Z.8.*

[Received April 2, 1912: Read May 7, 1912.]

(Plates LX XVIII.-LXXX.+)

INDEX.

Systematic : Page
IFAD FHARCORTM| Gy HO D> soocoaceoabsondoncsbsvoowcdessoeoss OE
Barbus argyrot@nid, Sp. Ne .......eiceecec eee eeeeeeee eee O74
BEMUNUSASPS Meee. ae eee ee Ree eee OTA
Amphilius oxyrhinus, Spo. ..ceceee ceeeeeeceeeee 675
DBUGHOIG FDARCODGH, SDs Wa csaveroanc0e00000000000 090808000 676

The collection here reported upon, presented to the British
Museum by Mr. Blayney Percival, is of special importance as
coming from a district the fishes of which had not been collected
before, from a watershed of its own without any communication
with sea. The river called EKusso Nyiro, or Guaso Nyiro, quite
distinct from the Southern Guaso Nyiro in German Hast Africa,
takes its source to the east of Lake Baringo, separated from it
and from Lake Rudolf by hill-ranges 1000 to 1500 feet higher
(4000 to 5000 feet above sea-level) and is lost in the Lorian

* Published by permission of the Trustees of the British Museum.
+ For explanation of the Plates see p. 676.

Fe A, MEAs IP, LOC WANIL ,

J.Green del.et lith.

LAN IE © PIB IRCIVAILIL,

ad

12), Axe) NOB, JED. OID

J.Green del.et lith.

ILBARBUS ARGYROTANITA. 25 1S), MOMS,
Sp yA PIU TEI IRC INN).

SOUNDNSCSLAS CG) {SQUAW

“UAT 1eT>p us229 'P

POG Nel “Glee (Sve wal

Siar)
Pee es

-

FISHES FROM BRITISH EAST AFRICA. 673

Swamp. The Eusso Mara is a tributary of the Eusso Nyiro.
The Saya is a separate river, between the Anagata Borita and the
Eusso Nyiro, flowing north and lost in a small swamp some
50 miles north of Kusso Nyiro. Unless otherwise stated, the
fishes here listed were obtained in the Eusso Nyiro, below the
falls.

The fishes of this basin have much in common with those of the
Webi Shebeli and Juba, and nothing with those of Lake Rudolf.
Curiously, the small Barbus, B. nairobiensis and B. percivali,
previously discovered by Mr. Percival in the Nairobi (Athi River
system), on the east side of Kilimanjaro, were found again in
abundance in the Eusso Nyiro.

1. Mormyroprs DEetictosus Leach.
2. MormyRus KANNUME Forsk.

3. LABEO PERCIVALI, sp.n. (PI. LX XVIII.)

Body strongly compressed, its depth 33 to 32 times in total
length. Head 4 to 43 times in total length, 14 to 1} times as
long as broad; snout rounded, 3 length of head; eye perfectly
lateral, 25 (young) to 32 times in length of head; interorbital
width 4, or a little over 4, length of head; lips with small papille
forming transverse plice ; lower lip with a fringe of large papillee;
rostral flap moderately large, with entire or indistinctly denticulate
edge; a small barbel in the corner of the mouth. Dorsal III
10-11, equally distant from end of snout and from root of caudal,
or slightly nearer the former ; upper edge concave ; last simple ray
in adult males much produced, twice, or nearly twice, as long as
head, not or but little longer than head in females and young.
Anal III 5, reaching or nearly reaching root of caudal. Pectoral
considerably longer than head in males, reaching base of ventral,
the first ray of which falls below fourth or fifth branched ray of
dorsal, shorter in females. Caudal deeply emarginate, with long,
pointed lobes. Caudal peduncle 13 to 14 times as long as deep. —

Seales 38-42 —- 5 between Jateral line and root of ventral,

18 round caudal peduncle. Silvery, back brownish grey ; vertical
fins greyish.

Numerous specimens, measuring 55 to 190 mm.

Allied to Z. neumanni Blgr. Well distinguished by the larger
eye and the shape of the dorsal in the males.

4, LABEO CYLINDRICUS Peters.

Also from the Eusso Mara and the Saya, and the Lorian
Swamp.

5. DiscoGNATHUS DEMBEENSIS Rupp.
Also from the Eusso Mara and the Saya.

674 MR. G. A. BOULENGER ON

6. Barpus ERLANGER! Bley.

Also from the hot springs at Chandler Falls, upper Kusso Nyiro,
and from the Saya.

7. BARBUS NATROBIENSIS Bler.
From the Eusso Nyiro above the falls,

8. BARBUS PERCIVALI Bler.

From the above locality, and also from the Saya.
The largest specimen measures 105 nm.

9. BARBUS ARGYROTHNIA, sp.n. (PI. LX XIX. fig. 1.)

Depth of body 2? to 3 times in total length, length of head 34
to 3? times. Snout rounded, shorter than the eye, which is 25 to
3 times in length of head and equals or nearly equals interorbital
width; mouth small, subinferior; lips feebly developed; two
barbels on each side, anterior about % diameter of eye, posterior
as long as or slightly longer than eye. Dorsal III 8, equally
distant from posterior border or centre of eye and from base of
caudal, border feebly concave; last simple ray enlarged, bony,
strongly serrated behind. Anal III 5, not reaching caudal.
Pectoral a little shorter than head, not reaching ventral ; base of
latter below origin of dorsal. Caudal peduncle a little longer
than deep. Scales radiately striated, 30-31 = 3 between lateral

line and ventral, 14-16 round caudal peduncle. Pale sand colour,
with a silvery lateral band which is sometimes edged above with
a streak of black pigment; fins white, often tinged with pale
orange at the base.

Total length 53 mm.

Numerous specimens from the Eusso Nyiro below the falls.
BL. argyrotenia stands nearest to B. zanzibaricus Peters.

10. BarBus mimus, sp.n. (Pl. LX XIX. fig. 2.)

Depth of body 24 to 22 times in total length, length of head
33 to 4 times. Snout rounded, much shorter than the eye, which
is 25 to 3 times in length of head and equals interorbital width ;
mouth small, subinferior; lips feebly developed; two barbels on
each side, anterior about # diameter of eye, posterior a little
longer than eye Dorsal III 8, equally distant from posterior
border or centre of eye and from base of caudal, border feebly
concave; last simple ray not enlarged, not serrated, as long as or
a little longer than head. Anal III 5, not reaching caudal.
Pectoral as long as head, not quite reaching ventral; base of
latter below origin of dorsal. Caudal peduncle as long as deep or
a little longer than deep. Scales radiately striated, 25-26 = 2-24

2
between lateral line and ventral, 12 round caudal peduncle.
Pale sand-colour, with a silvery lateral band, which is sometimes

FISHES FROM BRITISH EAST AFRICA. 675

edged above with a streak of black pigment; fins white and
transparent, or pale orange at the base.

Total length 55 mm.

Numerous specimens from the Eusso Nyiro below the falls.

Closely allied to B. neglectus Blgy. from the Nile, and strikingly
resembling the preceding species in general appearance, although
differing widely in the feeble, non-serrated last simple dorsal ray
and in the much lower number of scales.

11. Ciarias LAzERA Lacep.
Kusso Nyiro, near the Lorian Swamp.

12. EurrRoprus DEPRESSIROSTRIS Peters.

13. Bacrus vrostiema Vincig.
Eusso Nyiro, above the Chandler Falls.

14. CLAROTES LATICEPS Riipp.
15, AMPHILIUS GRANDIS Bler.

16. AMPHILIUS OXYRHINUS, sp.n. (Pl. LX XX.)

Depth of body 6 times in total length, length of head 3? times.
Head much depressed, slightly longer than broad ; snout pointed,
3 length of head; eye very small, 11 times in length of head,
25 times in interorbital width ; posterior nostril midway between
eye and end of snout; maxillary barbel $ length of head, reaching
root of pectoral; outer mandibular barbel 2 length of head, inner
nearly 2. Gill-rakers rather long, 7 on lower part of anterior
arch. Dorsal I 6, much nearer end of snout than root of caudal.
Adipose dorsal low, 34 times as long as deep, twice as long as
rayed dorsal, its length $ its distance from latter. Anal III 5.
Pectoral a little more than 3 length of head. Veutral a little
shorter than pectoral, far behind base of dorsal. Caudal feebly
emarginate. Caudal peduncle 13 times as long as deep. Reddish
brown above, spotted with blackish, dirty white beneath.

Total length 195 mm.

Distinguished from 4. grandis by the pointed snout, the longer
caudal peduncle, and fewer anal rays.

A single specimen from the Eusso Mara, a swift mountain-
stream.

17. SYNODONTIS SCHALL BI. Schn.

18. SYNODONTIS GELEDENSIS Gthr.
From the Eusso Nyiro, close to the Lorian Swamp.

19. Tinapia niztorica L.
From the Eusso Nyiro, above and below the falls.

676 ON FISHES FROM BRITISH EAST AFRICA.

20. TILAPIA PERCIVALI, sp.n. (Pl. LX XIX. fig. 3.)

Depth of body equal to length of head, 23 times in total length.
Head about twice as long as broad; snout rounded, with siciglo
or slightly convex upper profile, a little broader than long, ? post-
ocular part of head ; eye 33 (young) to 4 times in length of head,
greater than preorbital depth ; mouth large, ? width of head,
extending to below anterior border of eye; lips very strongly
developed ; teeth moderately slender, in 3 or 4 series, 40 to 50
in outer series of upper jaw; 2 series of scales on the cheek, width
of scaly part rather less than diameter of eye. Giull-rakers short,
13 or 14 on lower part of anterior arch. Dorsal XIII-XV 11;
spines increasing in length to the last, which measures 2 length of
head ; longest soft rays § length of head. Anal ITI (exceptionally
IV) Ga 10; . third spine 3 length of head. Pectoral about 3 length of
head, not ‘veaching origin of anal. Ventral reaching vont or anal.
Caudal rounded, subtruncate. Caudal peduncle deeper than long.
Scales cycloid, 30-31 aS; lateral lines are Dark brown, with
black vertical bars on the body, or entirely black.

Total length 75 mm.

Hot springs at Chandler Falls, Northern Eusso Nyiro, 2500 ft.
Caught in water about 100° Fahr.

Near allies of this new T7%lapia are J’. alcalica Hilgend. from
alkaline water in volcanic holes near the Nguruman Salt Lake,
into which the Southern Guaso Nyiro flows, on the boundary of
British and German East Africa, and 7’. grahami Bley. from Lake
Magadé, a perfectly isolated hot soda lake in the Rift Valley,
British East Africa, not far from the Southern Guaso Nyiro.
The former has, like 7’. percivali, D. XIII 11, but the scales on
the cheek are in two series and the caudal is truncate; the latter
has D. XI 11-12, 3 series of scales on the cheek, and the caudal
peduncle is deeper than long.

EXPLANATION OF THE PLATES.

PuatTEe LXXVIII.
Labeo percivali, p. 673. 2 nat. size.

PratE LXXIX.

1. Barbus argyrotenia, p. 674.
2. 4 mimus, p. 674.
3. Tilapia percivali, p. 676.

Vig.

Puate LXXX.,
Amphilius oxyrhinus, p. 675.

ON NEW CESTODES FROM THE TASMANIAN DEVIL. 677

38. Contributions to the Anatomy and Systematic Arrange-
ment of the Cestoidea. By Frank H. Bepparp, M.A.,
D.Se., F.R.S., F.Z.8., Prosector to the Society.

[Received April 2, 1912: Read May 7, 1912.]
(Text-figures 92-101.)

V. On A NEw Genus (Dasywrotenia) FROM THE TASMANIAN
Devit (Dasyurus ursinus), THE TYPE OF A NEW FAMILy.

INDEX.
Systematic : Page
Dasyurotenia, GEM. We 0.2.0... .0. 02 cee ces eeecee neces ss 694
Ds HOOUSEAY SPH Is | scecesheeretesnstosececeestiasssmaccasaces OOS
Résumé of anatomical points...............0ccceeeeseee 695

The Cestoidea hitherto recorded from Marsupials appear to
belong to the following genera only, viz. T’riplotenia, Moniezia,
Lertiella, Linstowia, Anoplotenia, Oochoristica, and Cittotenia.
To these must now be added an eighth genus, of which I have
recently examined examples from the Tasmanian Devil, in whose
small intestine they occurred.

Text-fig. 92.

Dasyurotenia robusta, nat. size.

Two views of the same two specimens, to show their attachment to the gut.

The left-hand figure represents the interior of the gut, the right-hand
figure the outside.

The accompanying drawings (text-fig. 92) show two of the
three specimens which I obtained in situ. Each worm is firml
imbedded at the head end in the wall of the gut. The two

678 DR. F. E. BEDDARD ON

drawings show the inner surface and the outer surface of the
intestine. The body of the worm is rather flat though at the
same time fairly thick, and is curved im a sickle-like fashion.
The length of the larger of the two individuals is about 17 inches,
and the greatest breadth does not exceed 5mm. There is nota
very marked difference in diameter at the neck end of the body,
and the worm thus presents a very solid and strong appearance,
which is correlated with the very firm way in which it is 1m-
bedded in the gut. The narrowness of the segments and the
lack of overlap of successive segments contributes further to the
stout appearance of the worm.

Text- fig. oO 93 .

Head of scolex of Dasywrotenia robusta, shown by opening up the cyst on the gut.

The last two or three segments of the body are rather narrower
from side to side and are curved, thus tending to encircle each
other successively. This state of affairs-i 1S shown i in the drawings
referred to. Anteriorly the strobila is seen to perforate the wall
of the intestine, which forms a fold round it, like a collar not
fitting very closely. The strobila at this point disappears from
view when the intestine is examined from its internal surface.

—

NEW CESTODES FROM THE TASMANIAN DEVIL. 679

When the piece of gut to which the worms are attached is turned
over they reappear on the outer surface (as is shown in text-
fig. 92) in the form of a cyst. This cyst forms a simple hemi-
spherical bulging upon the wall of the gut consisting apparently
of the peritoneum. When this peritoneal wall is cut through a

Text-fic. 94.

Transverse section through scolex of D. robusta at about the middle.

The central medulla shows the problematical vesicles referred to in the text in
the middle of its extent and the coils of the water-tubes at the ends.

r. Irregular furrows upon scolex. ¢. Tube formed by these.
A second tube of the same nature lies above and to the right of that lettered.

cavity is opened up in which lies the large scolex. This is
illustrated in text-fig. 93. The scolex, it will be observed, does
not completely fill the cavity in which it lies.

680 DR. F. E. BEDDARD ON

The large size of the scolex of Dasyurotenia contrasts with the
relatively small or even minute scolex of other Tetracotylea, and
thus recalls that of the Tetraphyllidea and others of the ‘‘ lower”
groups of Tapeworms. So also do certain other characters of
the scolex in this worm. ‘The greatest diameter of the scolex is
3°5 mm., and it is of much the same length, so that its dimen-
sions are but little less than the diameter and thickness of the
body, and are greater than those of the neck-region which imme-
diately ensues. Broadly speaking, the scolex is of a globular form
and has a soft, easily indentable, irregular surface marked by faint
ridges and depressions, and not at all like the vertex of a typical
member of the Tetracotylea. Two flat plate-like areas are to be
seen when the scolex is examined in this way with a pocket-lens.
These are, as I believe, the expanded suckers which I have seen
in a contracted condition in another specimen whose scolex was
studied by means of transverse sections.

Transverse sections of the scolex (text-fig. 94, p. 679) are
approximately circular in outline, in the middle part at any rate.
This particular scolex, which I studied by the section method,
was apparently more retracted than the one represented in text-
fig. 93. For at first the sections (text-fig. 95) showed two semi-
circular masses closely applied along the straight margin which
is obviously the expression of an apical groove in the scolex.
It was lower in the series than this that the contour’ of the
sections became circular. At the body end of the scolex the
latter slightly overlaps the neck for the whole of its circum-
ference. But although the sections through the scolex are on the
whole circular in form, the peripheral layer is not at all uniform
and shows numerous processes and grooves occurring everywhere,
which is an expression of the irregular grooving and ridging of
the scolex apparent when that region is examined with a hand-
lens. I could detect no symmetrical arrangement in these pro-
jections and grooves, which, however, may collectively represent
‘“‘bothria” meandering over the surface of the scolex.

It is furthermore the fact that these grooves are converted in
places into tubes which run along the interior of the peripheral
layer of the scolex in a longitudinal direction, though not for a
long distance. These short tubes end blindly. Their existence,
together with the grooves and ridges of which they are a further
development, must facilitate the adhesion of the scolex to the
surrounding walls of the cyst. While the more or less irregular
grooves and furrows upon the surface of the scolex might be put
down to irregular contraction of its outer layer, this can hardly
be the case—one would suppose—with these invaginated tubes.
In any case the arrangement of the suckers does not conform to
the arrangement usually met with among the Tetracotylea (to
which group this worm would be expected to belong). The scolex,
however, possesses four suckers which are of very small size when
compared with the diameter and circumference of the scolex.

NEW CESTODES FROM THE TASMANIAN DEVIL. 681

The greatest diameter of one of the suckers is not more than
one-tenth the diameter of the scolex at its widest part.

Text-fig. 95.

Section through apex of scolex to illustrate relative size of one of two
smaller suckers (s.).

g. Median groove formed by retraction of sucker.

On either side of the apical groove already mentioned was a
single cup-shaped sucker bearing hooks. I assume, therefore,
that these two suckers are really anterior in the fully expanded
condition of the scolex, and that they are the two disk-like bodies

Proc. Zoou. Soc.—1912, No. XLV. 45

682 DR. F. E. BEDDARD ON

described above in the scolex, which was examined entire and by
the aid of a lens only. These two suckers were not exactly
opposite on the opposite sides of the groove. They represent
all that I can find in the seolex comparable to a rostellum. I
believe, however, that they do not represent the rostellum of the

Tetracotylea.

Text-fig. 96.

Two consecutive sections through one of two larger suckers bearing hooks.

A. Muscular layer of sucker. B. Cellular layer in which lie hooks (#.).

In these sections (text-fig. 96) the sucker appears cup-shaped,
the orifice naturally opening into the median transverse groove

NEW CESTODES FROM THE TASMANIAN DEVIL. 683

upon the scolex, The structure of the sucker is not like that of
the typical Tetracotylean sucker. In the latter there is one thick
layer of typically arranged muscle-fibres (into the nature of which
arrangement it is not my purpose to enter—it is well known)
which is quite distinct from the surrounding tissues of the scolex,
but never covered by them on its free surface, which faces towards
the aperture. In the present species the structure of this sucker
is illustrated in the accompanying text-figure (text-fig. 96), which
is drawn from one sucker only which appears to me to be rather
larger than its fellow, the two being thus mutually asymmetrical.
Examining the sections from below upwards, the sucker is seen
at first to be apparently like that of the Tetracotylea generally.
It consists of a strongly marked layer of muscle-fibres ete. lying
in a cavity of the body parenchyma. This is shown in the
figure. As we approach more nearly to the rostellar region,
another layer in which the hooks are implanted pushes itself
in front of the sucker proper, as is also shown in the text-
figure referred to. This layer is continuous with and similar
in structure to the general parenchyma of the head. The
true (?) sucker thus gets as it were buried beneath a layer
of tissue bearing hooks. So far the structure of this sucker
will be plain from the sections figured. In the corresponding
sucker of the opposite side of the body the hooks are obvious
and quite similar, and imbedded in a perfectly corresponding
tissue. But I could find no trace whatever of the muscular
structure independent of these hooks.

In addition to these two suckers the scolex of Dasyurotenia
possesses two others, thus making the normal four. The latter
are roughly opposite to each other, but do not—obviously,
at least —alternate with the others. These two suckers
are opposite to each other and lie on the outside of the
scolex, and are about on a level with the hooked suckers
already described. It is quite possible that when the scolex
is fully expanded and not warped through unequal contrac-
tion with alcohol, the four suckers might alternate with more
regularity than is apparent in my preparations. In any case,
these suckers are quite different from those already described.
They are smaller than, at any rate, the larger of the more apically
placed suckers; and there is no trace whatever of any hooks
associated with them. Hach appears (see text-fig. 95) in section
to have the form of a flat disk. Their structure seems to be
quite like that of suckers generally among Tapeworms. What
is very apparent about these suckers is their very small size com-
pared with the wide periphery of the scolex. I could not detect
them at all on an examination of the scolex with a lens (see
text-fig. 93, p. 678).

We have next to consider the histological structure of the
scolex, which presents certain peculiarities. The anterior region
of the scolex has no distinction into medullary and cortical layers.
The commencement of such a differentiation is marked by the

45*

084 DR. F. E. BEDDARD ON

depth of the invagination of the anterior end of the scolex referred
to above. This part of the scolex presents a very lax appearance
in sections, which is doubtless to be correlated with the irregular
outline of the scolex. Through this lax tissue stray Im every
direction muscle-fibres which no doubt effect the retraction of this
anterior part of the scolex. ‘The main mass of tissue in which the
muscular fibres run has the usual amorphous ground-substance of
the Cestoid body with many nuclei; but I have not made a
particular histological study of these tissues.

Text-fig. 97.

Part of transverse section through neck-region of Dasyurotenia robusta.

C. Cortical layer outside of longitudinal muscles (/.m.).
me. The thin medullary layer.

Further back in the scolex the medullary region appears as
quite distinct from the cortical. In the latter the muscular
fibres gradually collect into a massive bundle of longitudinally
running fibres which immediately surround the medulla and
occupy a great deal of the cortex. Within the scolex there is no
subdivision of this muscular mass into bundles such as I shall

NEW CESTODES FROM THE TASMANIAN DEVIL. 685

describe in the strobila and even in the neck-region of the same.
In the medullary region at either end is to be seen the water
vascular tube, which is here rather coiled, a tube appearing five
or six times in a single cross section. In the middle of the
medullary area are certain remarkable structures, which are
represented in text-fig. 94 (p. 679). These are apparently
hollow spheres excavated in the parenchyma but with very
definite walls. These vesicles contain lumps of an amorphous
matter into the nature of which I have not enquired. There
‘is no connection that I could find between these vesicles and
the water vascular system. Iam quite uncertain as to their
nature.

The neck-region of the worm after it has issued from the cyst
which contains the scolex is represented in transverse section in
text-fig. 97. It will be observed at once that the medullary
parenchyma is very greatly reduced. It forms a thin layer very
much narrower than the extremely thick cortical layer. Nor
could I discern in it the typical retiform appearance of the
Cestoid medullary parenchyma with spherical masses of granular
matter lying between the meshes of the network. The whole of
the available space, save between the individual testes at the two
sides of each proglottid, is occupied by the rudimentary female
organs, which together with the testes seem to occur in the very
first segments. The whole space thus occupied by the generative
masses and by what remains of the medullary parenchyma is not
one half of the diameter of the cortex. It has been already
pointed out that at the posterior end of the scolex, a little before
it merges with the neck, the medulla is surrounded by a very
thick layer of muscles composed of large fibres. In this layer no
marked arrangement of the fibres into bundles could be made
out. But in the next few sections, which we here speak of as
the neck, these fibres are very definitely disposed in a series of
bundles (text-fig. 98).

These bundles are disposed in four to six layers, and at the
two lateral extremities of each segment they are rather more
numerous than medianly. The bundles consist of a large number
of individual fibres, often as many as sixty or so. ‘They are sepa-
rated from each other by a dense nucleated tissue. Later in the
body we have the same four to six rows of bundles of fibres ;
but in these latter segments each row is separated from those
which lie above and below it by a delicate layer of transverse
fibres. These fibres are not apparent in the neck-region. I
imagine that this enormously powerful muscular system is corre-
lated with the distance to which the scolex is enabled to force
itself into the wall of the intestine of its host. These rows of
bundles of longitudinal fibres are reminiscent of what is charac-
teristic of the family Acoleidz, as is to be seen, for example, in
the genus Proterogynotenia,* where they have been figured by

* Abh. Senck. Ges. 1911, p. 260, fig. 14.

686 DR. F. E. BEDDARD ON

Fubrmann. Outside of these layers of fibres the cortex presents
no features which differentiate it in any marked way from other
allied forms.

Text-fig. 98,

(pairs iy Se
EOD. lage
Section through the muscular layer of the cortex, taken at a point further
back than that represented in text-fig. 97.

im. Bundles of longitudinal fibres. ¢.m. Transverse fibres.

The water vascular system of this tapeworm is in more than
one respect remarkable. In transverse sections through the
ripe proglottids only two vessels are as a rule visible, one on
each side of the body. The most careful examination often
failed to reveal the presence of another, even minute, tube. Nor
did longitudinal sections show any trace of this second vessel.
It is not, however, really absent in this worm, as I found it
to be in Thysanotenia lemuris, for it is present as a very minute
tube in some segments. ‘The vessels which are obviously present

NEW CESTODES FROM THE TASMANIAN DEVIL. 687

are of very large size, and it appeared to me that that on the pore
side of the segment was a little the larger of the two. This, then,
is the first important point about these vessels, 7.¢. that there is
a very large ventral tube on each side of the body and a very
minute dorsal tube. In following out a series of transverse
sections, it is seen that the lumen of the large ventral water-
vessels is occasionally occluded by a delicate diaphragm-like
sheet of membrane which is abundantly nucleated. There is no
question of a narrowing of the calibre of the tube, but of an
actual membrane which extends partly across it here and there.

In sagittal sections the existence of these membranes stretch-
ing partly across the lumen of the water vascular tube is quite
obvious. They occur, moreover, on both sides of the body, that is
to say in the case of both ventral tubes. The reason for emphasizing
this fact will be apparent later. In the longitudinal sagittel
sections referred to it will be seen that there are several of these
membranes which stretch a good way across the water-vessel,
and though two membranes arising from different sides of the
vessel do not actually meet, the edge of each stretches beyond the
edge of the other, so that the tube would appear, when viewed in
the direction of its length, to be entirely occluded. It is note-
worthy that these diaphragms, so to speak, arise indifferently
from both sides. The exact arrangement will be plain from the
annexed drawing (text-fig. 99, p. 688). I have noticed that in some
of the posterior proglottids the lumen is actually occluded once
in each proglottid. The two ends’of two oppositely projecting
membranes are connected by a continuous though very thin
membrane which connects the thicker extremities of the lateral
projections. This median part appears to be nowhere deficient,
and the water vascular system is thus divided up in these regions
of the body into a series of chambers.

I presume that these numerous membranes stretched across the
large vessels correspond to what has been figured in other tape-
worms as valves. I have emphasized the fact that they occur on
both sides of the body, because they carry on the pore side the.
genital ducts which actually perforate their substance and lie in
their thickness. This perforation is limited to the larger of these
valve-like structures which arise from the outer side, and it has
been produced I imagine by the extension round the ducts of the
water-vessel, which is of much greater diameter in the posterior
than in the most anterior segments. In the case of these latter,
as already mentioned, the generative ducts pass to one side of the
water-tube.

Another important feature in the water vascular system of this
tapeworm is the total absence of transverse vessels uniting the
longitudinal trunks in such segments. This state of affairs is
not unknown among other Cestoidea—it occurs, for example, in
Hymenolepis acuta *—but it is not common. Nor is this lack of

* vy, Janicki, Zool. Anz. Bd. xxvii. 1904, p. 776.

688 DR. F. E. BEDDARD ON

transverse vessels compensated by any network of excretory tubes
pervading the medulla, such as is met with in the genera /nermi-
capsifer and Zschokkeella *.

Text-fic. 99.

Sagittal section to illustrate ventral water vascular tube.

V. Projecting valves. v.d. Vas deferens. v.a. Vagina.
T. Testes.

* Beddard, P. Z.S. 1912, p. 596, and literature quoted there.

NEW GESTODES FROM THE TASMANIAN DEVIL. 689

The Generative organs of this tapeworm begin to be recog-
nisable very early in the body, only a segment or two behind the
head. But it is a long way back before the ovaries are ripe. As
in the vast majority of Cestoidea, the testes ripen earlier than the
ovaries. This being the case, the testes are recognisable earlier
as distinct bodies, and only cease to be so clear in the more pos-
terior segments, where the uterus is gorged with eggs. A remark-
able point of interest in the generative organs is the fact that the
duct leading to the exterior, or to be more exact the formative
mass of cells which will be both vagina and cirrus sac, is seen to
alternate in position in relation to the single water vascular tube
and the nerve-cord. In all segments the actual opening is on the
sane side of the body, but the generative duct passes towards it
either between the water-tube and the nerve-cord or outside of
both ; in the latter case only to one side—there is no alternation
between a dorsal and a ventral position. We shall see, when the
cirrus sac and vagina come to be described later, that there is
also variation in the exact relationship in position between these
two.

The ovaries are large and consist of two wings, which are
symmetrical or very nearly so, the middle point between them
being the middle line of the body, where are situated the shell-
gland etc. The ovaries are posterior in the segment, and behind
them lie the vitelline glands. These latter are of much the same
shape as the ovaries, and in rather immature segments differ only
from them by their rather darker staining with hematoxylin.
They also form two wings symmetrical with their middle point,
and are in contact with the ovaries in front. The two glands are
composed of many lobes, which reach as far as the testes at the
sides; altogether an ovary occupies fully half of the segment, and
rather more when it is fully mature. Immediately in front of it
are the sperm-duct and vagina. At the sides are to be found the
testes, which are also dorsal to it.

The vagina offers no very remarkable character. It has at first
a contracted lumen, which widens out for a considerable space,
and then contracts again before it suddenly opens into the rather
pear-shaped receptaculum seminis. The course of the vagina is
quite straight between its two ends and oblique in direction.
The narrow part of the vagina which opens into the receptaculum
is of some length, but shorter than the wider part. The vagina
shows quite the same characters in the most mature proglottids.
The receptaculum seminis is full of spermatozoa, and very
frequently contained ova at its wider end, close to where the
oviduct opens into it. The receptaculum and the vagina lie
anteriorly in the segment in front of the ovary, but behind the
uterus.

The wterus of this worm is persistent and found as a large
cavity extending right across the segment in the most mature
proglottids that I have examined. It begins as a small rounded
cavity lying in the front part of each segment.

690 DR. F. E. BEDDARD ON

The testes of this worm are chiefly massed at the two sides of
the proglottid, but these two masses are connected by a string
of testes which pass dorsally along the proglottid. They therefore
nearly surround the ovary and the female organs generally, so far
partly resembling Cyclorchis. In transverse sections the testes
appear circular in section; but sagittal sections such as that
represented in text-fig. 99 (p. 688) show that the form of the testes
is that of a flat plate, for in those sections they appear more linear
in shape. ‘These sections also show quite plainly that each testis

Text-fig. 100.

A portion of the coil of the sperm-duct (sp.) gorged with sperm.

ce. Interstitial prostatic cells.

lies in a space, unless, indeed, the appearances produced are due
to shrinkage through reagents. In any case, however, the testes
of these worms frequently present the appearance of being sur-
rounded by the spermatozoa which they produce, thus showing
that a chink exists or can be formed for their reception when
pressed out of the testis. The testes are very numerous and
quite crowded together,—so much so that the delimitations
between successive segments so far as concerns these organs

NEW CESTODES FROM THE TASMANIAN DEVIL. 691

are not at all visible in sagittal sections; they appear as a
continuous mass.

The sperm-duct runs straight for a short way after it has left
the cirrus sac. It forms a copious coil (text-fig. 100) occupying the
middle of the body and lying partly dorsally to the receptaculum
seminis and rather nearer to the pore side of each proglottid. The
rest of the coil—that part which is nearest to the cirrus sac—is

Text-fig. 101.

Sagittal section showing in consecutive segments the varying relations of
cirrus sac (the larger tube) and vagina.

to the pore side of the receptaculum, and therefore ventral in
position. In fact, regarded as one coil, this region appears oblique
in direction in transverse sections of the proglottids. Between the
individual loops of the sperm-duct are cells which quite fill up
the interstices, and are thus numerous in proportion to the width

692 DR. F. E, BEDDARD ON

of those interstices. The cells are rather clear, with well-stained
nuclei. They correspond exactly, as it seems to me, with the
prostatic cells of Jnermicapsifer and Zschokkeella, dealt with in
these genera by v. Janicki* and myself T, and which appear to
occur also elsewhere. In immature segments the cells bulk more
largely than the coils of the sperm-duct. But the reverse is the
case in the mature proglottid.

The genital ducts open into a common cloaca genitalis, which in
its turn opens on to the exterior. The cloaca genitalis is of some
depth, but it is not borne upon any process of the body. Into it
open, close together, the vagina and the cirrus sac, whose mutual
relations I have investigated by means of sagittal sections (text-
fig. 101) through portions of the strobila. There is a con-
siderable variation in these relations. Although the vaginal pore
is apparently never in front of the male pore, it is not always
directly behind the male pore. ‘The commencing vagina lies
obliquely behind the commencing cirrus sac—the direction of the
obliquity being now dorsal now ventral in this segment and in that.
There is, in fact, an irregular alternation from segment to seg-
ment ; sometimes the two tubes are not merely oblique, but actually
dorsal or ventral to each other as the case may be, lying there-
fore side by side in sagittal sections. This recalls to mind the
alternation that occurs in certain (but not all) species of the
genus Moniezia, where the vagina may be dorsal or ventral to the
clirus sac. But in this latter genus the alternation is of the
right and left set of generative organs of a single segment.

The cirrus sac of this tapeworm is large and has the very common
flask-shape. The neck-region has very thick circular muscular
walls, forming a sheath which thins out over the more distended
region of the sac. In less mature segments (in which, however,
the testes are fully developed, though with no mature sperma-
tozoa) the cirrus sac is elongated, gradually diminishing in breadth
towards the external pore; there is no marked division into neck
and flask. It is very long and extends towards the middle line
of the body, a little beyond the water vascular tube of its side of
the body, or at least reaches the internal side of that tuke. The
cirrus runs straight from end to end of the cirrus sac and
anteriorly presents a moniliform appearance, which is due to suc-
cessive dilatations of the lumen of the cirrus. This region of the
cirrus is both preceded and succeeded by a perfectly straight
section of that tube with very narrow lumen and thick walls.

In later segments the cirrus sac acquires the flask-shape already
referred to. Coincidently with this is an actual shortening of the
length of the entire sac and a coiling of the cirrus within it.
The cirrus sac in these and in subsequent segments hardly reaches
beyond the outer edge of the water vascular tube. It seems clear
therefore that the shortening is due to an actual contraction of

* Denkschr. Ges. Jena, xvi. 1910.
+ P. ZS. 1912, p. 602.

NEW CESTODES FROM THE TASMANIAN DEVIL. 693

the length caused by a bulging of the walls of the cirrus sac due
in its turn to the rapid growth and consequent coiling of the
cirrus. When the cirrus sac is in this fully formed condition, the
cirrus itself is differentiated more thoroughly into those regions
less markedly indicated in earlier stages. The sperm-duct enters
the cirrus sac at the apex and its lumen contracts to a fine line
for a short space near to its entry. This is particularly obvious
in the last few segments of the body, where the sperm-duct has
become much dilated before its entry into the cirrus sac, and thus
offers a greater contrast to this exceedingly narrow region.

In these more mature cirrus sacs the flask-shape has been
acquired, as already mentioned. But the neck of the flask is
much longer than the body part. The latter is so fully occupied
by the coils of the cirrus itself that there is but little of the inter-
stitial packing tissue to be seen. At its entry into the cirrus sac
and for a considerable time thereafter the duct is thick-walled,
with a very narrow lumen, and much coiled. This region of the
cirrus is succeeded by a not very long but coiled tract, which is
much wider and has thinner walls. The lning membrane bears
numerous spinelets. Finally, the distal region of the cirrus is
again thick-walled and with a narrow lumen: it opens into the
genital cloaca without any alteration of character. In the most
posterior segments of the body the greater part of the cirrus sac
is filled with sperm, the posterior region alone showing a group of
coils of the cirrus. Whether the anterior part of the cirrus has
become ruptured, as it appeared, or has been simply enormously
expanded and its walls reduced to extreme tenuity by the enclosed
sperm, I am unable to say.

The systematic position of this tapeworm is difficult to fix with
any confidence. ‘Ihe generative system, and, indeed, the internal
anatomy generally, presents no differences of importance from
many ‘l'etracotylea; and there are, indeed, no reasons so far why
the worm should not be placed in the Anoplocephalide, which
family, as has been pointed out, contains nearly all the Marsupial
tapeworms. On the other hand, the very much developed layers
of longitudinal muscles in the body-wall suggest the family ~
Acoleidze. The difficulty, however, of accurately placing the worm
lies in the peculiarities of the scolex. here is no doubt that it
contrasts very considerably with the general form of the scolex
among the ‘Vetracotylea in a number of points, of which the
principal ones are:—(1) its large size, both relatively to the body
and actually ; (2) the presence of numerous grooves which cannot
be, at any rate, entirely artefact, as they are converted here and
there into tubes running win the thickness of the head;
(3) the relatively minute size of the four suckers and the fact that
two of them and two only are furnished with hooks*.

* Furthermore, these hooks are distinctly hollow at their broader end, “like a
Ruminant’s horn,” as Shipley (Willey’s Zool. Res., Entozoa, 1900) notes of C ‘allio-
bothriwm, one of the Tetraphyllidea.

694 DR. F. E, BEDDARD ON

These characteristics are collectively different from anything
met with among the Tetracotylea that is known to me. They
are not, however, inconsistent with the conditions known to occur
among the Tetraphyllidea, if we may admit the grooves upon the
scolex to represent the bothria of such tapeworms. The suckers
would then correspond with the small accessory suckers so
frequently possessed by these latter worms, and their small size
relatively to the scolex would be thus intelligible. The apparently
numerous bothria not reducible (by me) to symmetry is suggestive
of a type like Phyllobothrium* slightly modified, or perhaps
Peltidocotyle t. We cannot, however, place Dasywrotenia among
these Tetraphyllidea on account of the Tetracotylean character of
its yolk-gland. But with reference to this gland it may be borne
in mind that it is in structure much more diffuse than is usual
with the generally solid vitelline gland of the Tetracotylea.

The genus may be thus defined :—

Dasyurotenia, gen. nov.

Stoutly bwilt worms with large scolex bearing four small suckers,
of which the inner two bear hooks. No rostellwm, but anterior end
of scolex, including hooked suckers, retractile. Segments very
short. Inner layer of (longitudinal) muscles very thick, consisting
of four to six rows of bundles of fibres. Ventral excretory tubes
large, with numerous valves not communicating with each other in
the strobila. Dorsal vessels minute, not always visible. Genital
pores unilateral. Testes numerous, chiefly lateral, anterior, and
dorsal. Vas deferens with a large coil in middle of segment sur-
rounded by prostatic cells ; cirrus sac large, cirrus with spinelets.
Ovaries with two wings, median ~ and posterior and ventral in
position, in front of vitelline gland, which is also symmetrical.
Shell-gland median, dorsal. Lreceptaculum seminis present, nearly
median, ventral. Uterus sac-like, persistent, fills nearly whole of
ripe proglottid. Eggs thin-shelled.

Hab. Marsupials.

The species I term “robusta” on account of its very stout build.
It is, however, quite impossible for me to venture upon an
enumeration of the peculiar specific characteristics for the present.

This genus and species cannot, as I think, be identified with
any other form that has been described from an Australian
Marsupial. From the present genus we only know a species
described by myself § a little time since as dnoplotenia dasyuri.
Nor can I identify it with any of the genera enumerated at the
beginning of this paper in other Marsupials. In fact, Bertiella is

* Bronn’s ‘ Thierreich,’ Bd. iv. Cestoiden, pl. xli. fig. 10.

+ Ibid. pl. xlii. fig. 1.

~ Middle line of female apparatus only slightly displaced towards pore side.
§ P.Z.S. 1911, p. 1003.

NEW CESTODES FROM THE TASMANIAN DEVIL. 695

the only one of these genera to which it bears any likeness in the
reproductive system, and from this genus the characters of the
scolex at once distinguish Dasyurotenia. Indeed, its inclusion
among the Tetracotylea (=Tzenioidea) is not, to my mind, an
obvious certainty. In any case the hooked suckers exclude it
from the family Anoplocephalide, to which nearly all the
Marsupial tapeworms belong up to the present.

The most salient points of anatomical interest in this worm
appear to me to be the following :—

(1) The immense size, relatively speaking, of the scolex and the
small size in comparison with it of the suckers. The fact that
two suckers are armed with hooks while the other two suckers
are not so armed.

(2) The great thickness of the longitudinal muscles, which
consist of at least four layers of bundles each containing very
many individual fibres of considerable stoutness.

(3) The existence for the most part of only a single water
vascular tube on each side of the body, which is, moreover, in the
posterior segments completely divided up into a series of com-
partments, one to each segment, and whose lumen is also here and
more anteriorly divided by delicate septa jutting into its cavity.
Furthermore, by the fact that these tubes are not connected
in successive segments by transverse vessels, as is so nearly
universally the case.

(4) As a remarkable structural feature, which is at present
mysterious in nature, may be mentioned the isolated cavities in
the medullary region of the head which have no connection with
the water vascular tubes.

(5) An anatomical feature of some importance is the very
variable relation to each other in position of the extremities of
the male and female ducts, which is correlated with an orifice
upon one side of the body only. An alternation in the position
of the external pore may, we know, be accompanied with difference
in the relative position of the ducts as, for example, in the double
series of genital tubes of Moniezia.

(6) In view of the very considerable peculiarities of structure
briefly indicated in the foregoing résumé, it may be worth men-
tioning, as a remarkable fact, that the generative organs do not
show any marked features of interest as compared with those of
other tapeworms.

696 MR. R. E. TURNER ON

39. Studies in the Fossorial Wasps of the Family Scoliidze,
Subfamilies Elidinee and Anthoboseinee. By RowLanp
EK. Turner, F.Z.S., F.H.S.

[Received March 29, 1912: Read May 7, 1912.]

(Plates LXXXI-LXXXIIL*)

INDEX.
Geographical Zoology : ‘ Page
Anthoboscas DistributlonsOie ee eteeteeeteeneeeete eee emo

Systematic :

Anthobosca occipitalis SP. Ve ...cieceeeeeseeteeeeeeeeeeeeeee V4
IST ROPSOMIGINUG FHEIG 15 coc uno coo syoe00 60D oAnOF baDnodeapcooCaseas OIL
ss quadraticeps, SP. Ve vccevcieeereeresreeeeee 698
* HARICEOS Ss 15 soaotacanosnosnesnndesccdenoso | ONY)
TUS mag on As py ashes. ve Rapes eee tectees seo cae eceeee eee RISO
spin SELLOWL, ISP Mla, nde sk et nenetes o sa deccesteasns shieustaleessins sommes
59 (GEES) GHABO, SIM WE sooace cdocaocdgane eccoaeAcacosensess DE!
5 Dy CYDICUDETIDOS, Go Wo esoecnaos so sod oosccpcess00020 707
es Plt 06-0 A0P8Ip S100 Necernes bapesosaneenonbperpamenecccis).e (0)
3 Fy) ANCELLA SPAM ee ah sea eeahaanettaceaneke cose ALO.
43 fy CORE TODOS SOs 1) paoceoncocarcsconscescscce 1
is a) SpaldatadspauMey | stucsorsusensccccesonaatecsrsh er MLL
WIG EUDG WiRACHIROs Fo 5 choosen 05 900000 990300 99590900 506 700
45 combusta Sm. tO Hlis.......:-.cesccceoreseeeee sees TRA
g9 COMSELECELVENETIS, SP. Deo... sce ceeeseveeereereeees ero!
» sauakinensis Grib. to Anthobosca ...... ........ 740
Sod ISLOGULAS SDS Ds, | anon: tascmcetare setqseckine tee. ae ceo OOO,
Do EMD RALTCE AIS MMM et on cee esate ee suisse oe ame
Odontothynnus Cam. f to Anthobosed.................065 724
Plesia continua Cam. 6 of Myzina abdominalis Guér. 703
» erythronota Cam. to Anthobosca .................. 738
» leucospilaCam. ,, 5s) Wg HAR ee 739
» melanaria Cam. ,, a Meee erscoeicthey aie)

The following notes on the Elidine and Anthoboscinze will, it is
hoped, facilitate the study of these neglected groups, the latter of
which especially has been very little understood by many authors,
who have touched on it merely as describers of new species,
Saussure alone having seriously studied the group. ‘The best
work on the Elidine has been done by the same author, but the
material at his disposal was very limited.

I am indebted to Dr. Brauns, of Willowmore, 8. Africa, for
valuable assistance with many carefully collected specimens. The
material available is still insufficient for a revision of the species
of Myzine, as to which much confusion still exists.

The species which I have not seen are marked with an asterisk.

* Wor explanation of the Plates see pp. 753-754.

IP ZS. WB Ae al IOI,

West, Newman chr.

Horace Knight del.et lith.

FOSSORIAL WASPS.

PBS, UGA, IPI, USOT.

ZO

West,Newman lith.
WING NEURATION OF FOSSORIAL WASPS.

Catherine A.M.Pearce del.

Miibears:*¥

12S AZ, PIL IL OO.

Catherine A.M.Pearce del. West,Newman lith.
HRXOSKELETAL STRUCTURES OF FOSSORIAL WASPS.

STUDIES IN THE FOSSORIAL WASPS. 697

Family SCOLIIDA.
Subfamily Hxurin 2.
BRAUNSOMERIA, gen. nov.

2. Apterous; mandibles acute at the apex, with a rather in-
distinct tooth on the inner margin near the apex; antenne twelve-
jointed, the first joint of the flagellum very small and almost con-
cealed by the apex of the scape. Head almost rectangular, the
posterior angles slightly rounded ; eyes oval, touching the base of
the mandibles, rather small, separated by a distance at least as
great as their gwn length from the posterior angles of the head ;
ocelli absent, their pasition indicated by large punctures. Thorax
much narrower than the head ; pronotum rather longer than its
greatest breadth; mesonotum very short, almost covered by the
pronotum, the tegule more or less developed ; scutellum narrower
than the pronotum, broader than long; median segment nearly as
long as the pronotum, flattened on the dorsal surface, broadened
from the base to the apex. Sides of the head and thorax and base
of the abdomen thinly covered with long hairs, Abdomen longer
than the head and thorax combined, shining, the apical segment
long and more or less acute at the apex, stricture between the first
and second ventral segments well developed. Intermediate coxze
rather widely separated, posterior coxe contiguous, intermediate
and posterior tibize spinose, tarsal ungues simple.

3. Winged; stigma rather large, situated at about three-fifths
from the base of the wing; radial cell shorter than the stigma ;
three cubital cells, the second and third small, not reaching the
apex of the radial cell, each receiving a recurrent nervure; cubital
and discoidal nervures not continued beyond the cells. Medial
cell of the hind wing not emitting veins from the apex. Antenne
in the typical species long and slender, thirteen-jointed, the first
joint of the flagellum almost concealed in the apex of the scape,
the antenne much longer than the abdomen ; antennal tubercles
well developed. Head strongly convex; ocelli present. First
abdominai segment with a short petiole, the segment, including
the petiole, a little longer than the second segment, suddenly
widened at the apex of the petiole. Apical segment with a re-
curved spine, the apical emargination of the dorsal segment
shallow. Eyes entire, not emarginate.

The characters given here for the male will doubtless be found
not to apply to all species of the genus; but the important
characters in the neuration separating the males from dyzine
are the larger stigma, the blunter apex of the radial cell, the fact
that the cubital and discoidal nervures are not continued beyond
the cells as in Myzine, and that no veins are emitted from the
median cell of the hind wing, there being two veins in J/yzine.

Tvpe of the genus, Lrawnsomeria quadraticeps.

Proc. Zoou. Soc.—1912, No. XLVI. 46

698 MR. R. E. TURNER ON

BRAUNSOMERIA QUADRATICEPS, sp. n. (Pl. LXXXTI. figs. 9, 10;
PU OXOXGT I fies fe)

2. Rufo-ferruginea ; mandibulis apice, vertice, capite lateribus ;
segmentis abdominalibus tribus basalibus nigris ; femoribus tibtisque
Juscis, calcariis albidis.

Long. 8 mm.

@. Head broader than long, fully half as broad again as the
pronotum ; mandibles with a blunt tooth on the inner margin
near the apex, another near the middle of the imner margin,
acute, and another smaller nearer the base. The whole insect
shining, with a few scattered punctures. Pronotum longer than
broad, a little longer than the median segment. Dorsal abdo-
minal segments broadly depressed at the apex, the basal portion
of the segments produced into a slightly raised rounded mark

op each side; apical segment very narrowly rounded at the
extremity.

3. Niger; mandibulis, clypeo macula mediana nigra, scapo
subtus apice, tuberculis antennalibus, macula parva frontali, mar-
gine interiore oculorum, linea undulata verticali, pronoto macula
utrinque antice et fascia lata postice, mesonoto macula quadrata,
scutello fascia lata, postscutello, mesopleuris fascia, segmento dor-
sali primo fascia apicali, ceteris fascia apicali macula nigra
utringue, segmentisque ventralibus 2-6 fascia bisinuata apicali
pallide flavis ; alis hyalinis, venis testaceis.

Long. 6 mm.

3g. Antenne slender, longer than the abdomen, the inter-
antennal tubercles prominent. Clypeus very short and broad,
very shallowly emarginate at the apex. Head very strongly
convex, cheeks as broad as the eyes. Head and thorax coarsely
but not very closely punctured, median segment finely and closely
punctured-rugulose; abdomen shining, very sparsely and shallowly
punctured. Pronotum shorter than the mesonotum, narrowed
anteriorly, the anterior margin straight, posterior margin very
feebly arched. Median segment steeply sloped posteriorly, not
truncate. Petiole of the basal abdominal segment occupying less
than half the length of the segment, the remainder of the seement
slightly inflated ; “lhe first segment, including the petiole, only a
little longer than the second. The segments not constricted ;
hypopygium forming a long recurved spine; apical dorsal sesment
convex, shallowly emareinate at the apex. Stigma large, “twice
as long on the costa as broad, nearly twice as long as the radial
cell, which is broadly rounded at the apex. Three cubital cells
on the right side, two on the left; on the right the second
abscissa of the radius is very short, the third about equal to the
first and second combined, but shorter than the second transverse
cubital nervure, second recurrent nervure received close to the
apex of the third cubital cell.

Hab. Willowmore, Cape Colony ; January (Dr. Brauns).
The female is the type.

STUDIES IN THE FOSSORIAL WASPS. 699

T have little doubt that links will be discovered connecting both
sexes of this genus with M/yzine through the short-winged Pseudo-
meria section in the females, and through species with somewhat
more extended neuration in the males. But the apterous con-
dition of the female and the differences of neuration pointed out
in the description of the male seem to me to be sufficient reason
for founding a new genus. The female shows a strong resem-
blance to female Thynnide of the genus Mirone, also to the
Bethylid genus Apenesia. J have not been able to examine the
mouth-parts, but it is likely that they would show atrophy of
some parts. The entire eyes of the male are also noticeable as
contrasted with the shallowly emarginate eyes of Myzine.

BRAUNSOMERIA ATRICEPS, Sp, 1.

2. Nigra; mandibulis basi, clypeo, antennis, therace, seymento
mediano, pedibus pygidioque apice ferrugineis,

Long. 5 mm.

2. Mandibles acute, with a very small ill-defined tooth on the
inner margin near the apex. Head rectangular, a little broader
than long, very slightly convex, shining, with a few scattered
punctures. Thorax shining, with a few scattered punctures on
the pronotum, the median segment more closely punctured.
Pronotum longer than broad, shghtly narrowed anteriorly,
narrower than the head by about one-third; mesonotum very
short, the tegule rather better defined than in quadraticeps ;
scutellum rounded posteriorly, broader than long. Median seg-
ment a little shorter than the pronotum, slightly broadened from
the base, a little broader than long and obliquely sloped poster-
iorly ; sides of the thorax and median segment sparsely clothed
with long yellowish hairs, Abdomen shining, finely aciculate,
the basal segment truncate anteriorly, with a short petiole not
more than half as long as the posterior coxe ; the third segment
the broadest; a semicircular small raised mark on each side of
dorsal segments 2-5; sixth segment smooth, pointed at the apex.
The constriction between the two basal ventral segments is well
marked. The eyes are smaller than in quadraticeps and are
separated from the posterior angle of the head by a distance
equal to about three times their own length.

Hab. Algoa Bay, Cape Colony; November (Pr, Brauns).

Myzine (?) stigma, sp,n. (Pl. LXXXI. fig, J1; Pl, LX XXII.
fig. 13.)

3. Niger; mandibulis basi, pronoto anguste postice, tegulis,
tibtis subtus tarsisque basi pallide flavis; alis hyalinis, venis
perlucidis, stigmate maximo, pallide flavescenti; oculis haud
emarginatis, antennis abdomime brevioribus ; cellula radiali oblite-
rata, cellula cubitali secundo pene obliterata.

Long. 7 mm.

¢. Clypeus very short, transverse. Antennze about as long as
46*

7CO MR. R. E. TURNER ON

the thorax and median segment combined, not very slender, of
about even thickness throughout, inserted a little nearer to the
eyes than to each other. Eyes not emarginate, their inner
margins parallel; posterior ocelli a little nearer to each other
than to the eyes. Head convex, closely punctured, with a
frontal suleus reaching to the ocellus; antennal tubercies not
developed. Thorax rather sparsely punctured. Pronotum rather
short, as broad as the head, the anterior margin straight, the
posterior margin widely and feebly arched. Scutellum large, a
little shorter than the mesonotum. Median segment short, almost

smooth, with a median sulcus, truncate posteriorly. Abdomen —
subsessile, the basal segment broad, not constricted at the apex
on the dorsal surface, deeply divided from the second on the
ventral surface, all the segments sparsely punctured and shining ;
the apical segment rather deeply triangularly incised for the
reception of the Jong aculeus of the hypopygium; the whole
abdomen about equal in length to the head, thorax, and median
segment combined. Stigma very large, about twice as Jong as
the greatest breadth ; only one cubital cell and one recurrent
nervure, which is received on the cubitus just beyond the angle
of the cubital cell, the cubitus continued just beyond the point of
reception of the recurrent nervure, the radial cell and all neura-
tion beyond the stigma obliterated. Median and submedian cells
of the hind wing present, but no neuration beyond them, the
median cell not extending very far beyond the submedian.

ITab. Willowmore, Cape Colony (Dr. Brauns).

This very distinct species will probably prove to be generically
distinct from MMyzine. It approaches most nearly to MW. swalec
Turn. and J. brawnst Turn., but differs in the reduced neuration,
the entire eyes, the more robust and subsessile abdomen, and the
much broader stigma. But until the female is known I prefer to
leave it provisionally in Myzine. From Braunsomeria, to which
the neuration approaches more nearly than to M/yzine, it may be
distinguished by the much more robust build, the much shorter
and stouter antenne, and the deeper emargination of the apical

segment.

MyzinE BRAUNSI, sp. n. (Pl. LXXXI. fig. 14; Pl. LX XXII.
fig. 14.)

3. Niger; mandibulis basi, pronoto fascia angusta postice,
segmentis dorsalibus 2-6 macula transversa mediali apice, mac-
laque curvata laterali utrinque, tegulis, tarsis articulo apicali
excepto, tibis anterioribus omnino, imtermedus posterioribusque
basi pallide flavis; alis hyalinis, venis perlucidis, stigmate pallide
testaceo.

Variat segmentis dorsalibus macula mediali obliterata.

Long. 5-8 mm.

3. Clypeus much broader than long, closely punctured and
shallowly emarginate at the apex. Antenne gradually thickened

STUDIES IN THE FOSSORIAL WASPS. 701

towards the apex, the terminal joint twice as thick as the fourth ;
the antennz about as long as the head, thorax,and median seement
combined. Eyes slightly emarginate on the inner margin ;
posterior ocelli nearly as far from: each other as from the eyes.
Head and thorax closely and not very finely punctured ; pronotum
narrower than the head, shorter than the mesonotum, narrowed
anteriorly, the anterior margin straight, the posterior margin
widely but not strongly arched. Median segment transversely
rugose, with a longitudinal depression in the middle, truncate
posteriorly, the surface of the truncation coarsely transversely
striated. Abdomen narrower than the thorax and nearly half as
long again as the head, thorax, and median segment combined,
finely and sparsely punctured, the segments scarcely constricted
at the base; petiole of the basal seement very short, the segment
abruptly broadened and slightly olan, nearly as Jong as the
second segment without including the petiole. Apical seement
convex, the incision at the apex subtriangular, not quite as deep
as its apical breadth. ‘Tarsal ungues simple. Cubital and
discoidal nervures not continued beyond the cells, stigma not
rounded on the inner margin, three times as long as the greatest
breadth ; radial cell acute at the apex, produced far beyond the
third cubital cell, second and third abscissee of the radius nearly
equal in length, second recurrent nerv ure received just before the
middle of the third cubital cell. Median cell of the hind wing not
emitting any veins from the apex.

Hab. Willowmore. Cape Colony; January to March (Dr. Brauns).

This is allied to AZ. swalei Turn., but differs in the shorter and
stouter basal abdominal segment, in the sculpture of the median
segment, and in the translucent nervures of the wings. Both
differ from typical Myzine by not having the cubital andi discoidal
nervures continued beyond the cells vad) i the absence of the two
veins emitted from the apex of the median cell of the hind wings.

MYZINE CONSTRICTIVENTRIS, sp. n. (Pl LUXXXII. fig. 15;
Pl. LX XXIII. fig. 12.)

3. Niger, dense albo-pilosus ; mandibulis basi, pronoto margine
posteriore, tegulis bast, segmentis dorsalibus 2-6 macula mediana
apicali et macula transverse tr inque, tibiis basi tarsisque pallide
flavis ; alis hyalinis, venis brunneis.

Long. 10-12 mm.

3g. Clypeus broad and short, shallowly emarginate at the apex.
Hyes distinctly convergent towards the elypeus, the inner margin
not emarginate, almost straight. Antenne inserted nearer to
each other than to the eyes, almost as long as the abdomen,
moderately stout and of almost even fhiclnees throughout.
Posterior ocelli a little nearer to the eyes than to each ‘oth er,
Head and thorax closely and strongly punctured; pronotum
short, narrower than the head, the anterior margin straight, the
posterior margin very feebly arched. Median seoment coarsely

702 MR. R. E. TURNER ON

rugose, almost vertically truncate posteriorly. Abdomen much
longer than the head and thorax combined, not very slender,
sparsely punctured, the segments rather strongly constricted at
the base, subsessile, the first segment no longer than the second.
Sixth dorsal segment more coarsely punctured, not convex, the
lateral margins raised, the apical emargination shallow, much
broader at the apex than deep. Radial cell broad, nearly twice
as long on the costa as the greatest breadth ; second abscissa of the
radius distinctly longer than the third, second recurrent nervure
received close to the middle of the third cubital cell.

Hab. Willowmore, Cape Colony; October to January (Dr.
Brawne).

This species is easily distinguished by the absence of any
emargination of the eyes, the shallow emargination of the apical
dorsal segment, the strongly constricted abdominal segments, and
the broad radial cell. The cubital and discoidal nervures are
continued very little beyond the cells, and the two nervures
emitted from the apex of sls median cell of the hind wing are
very short.

MyYZINE UMBRATICA, Sp. i.
Nigra, mandibulis pygidioque fusco-ferrugineis ; segmentis
dorsalibus 2-3 macula laterali utrinque alba; alis fusco-violaceis.
Long. 10 mm.

@. Shining and almost smooth, coarsely but not very closely
punctured round the base of the antenne; a few scattered
punctures on the vertex, pronotum, mesonotum, and scutellum,
pro- and mesopleurz strongly but not very closely punctured ;
median segment closely and rather finely punctured at the base,
almost smooth in the middle and at the apex, a few obscure striz
at the posterior angles; abdomen almost smooth, with a few
small punctures on the apical portion of the segments. Long
black pubescence on the sides of the thorax; calcaria whitish.
Hyes rather narrowly ovate, cheeks as broad as the eyes; ocelli
small, the posterior pair about as far from each other as from the
eyes. Antennze smooth and shining, thescape beneath punctured
and clothed with long hairs. Head subrectangular, half as broad
again as long, much broader than the thorax. Pronotum about
twice as broad as long, the posterior margin almost straight.
Mesonotum only half as long as the pronotum and a little shorter
than the scutellum, the parapsidal furrows very distinct. - Apical
segment of the abdomen convex, long, and pointed. Wings of
moderate length, reaching to the fifth abdominal segment, the
stigma situated about halfway between the base and apex;
neuration similar to that of rufifrons Fabr.

Hab. Fourteen Streams, Cape Colony; January (Dr, Brawns).

There is only a very obscure median suleus on the median
segment: The slit in the fore wing extends from the termination

of “the cubitus just beyond the third cubital cell to the margin of
the wing.

STUDIES IN THE FOSSORIAL WASPS. 703

LXXXT. Qe

Guér.

MyZINE ABDOMINALIS (Pale

Pl. LXXXIITI. fig. 5.)

Meria abdominalis Guér, Rev. de Zool. iii. p. 365 (1839), 2.
Plesia continua Cam. Rec. Albany Mus. i. p. 299 (1905), 3

Hab. Willowmore; Burghersdorp, Cape Colony.

Taken in copuld by Dr. Brauns.

fig.

variable, the head being sometimes ferruginous.

10.
11.

13.

. Sixth dorsal segment punctured ;

. Lateral margins of the srareekenn segment acute ;

Key to the Ethiopian Species of Elis (Mesa).

females.

. Basal joint of Beene tarsi with a row of pa

beneath ak
Basal joint of posteri ior tarsi unarmed or with a ‘scopa

only beneath .......

- Radial cell distinctly ‘separated from the costa for

Black, two

more than half the length. apical

abdominal seements red
Radial cell separated from the costa at the apex

only
Abdomen ferruginous, the basal segment only black.

Abdomen black

. Thorax and abdomen entirely black

Thorax more or less red, the pronotum shorter than
the scutellum

calearia of the

hind tibie black
Sixth dorsal segment striate ;

calearia whitisn

. Vertex red, much more sparsely punctured than the

front, mesonotum and scutellum black. Spur of
the Se tibia not strongly bent near the
base ........

Vertex black, as s closely ‘punctured : as the “front,
mesonotum and scutellum red. Spur of the
posterior tibiz strongly bent near the base .........

. Abdomen wholly bright ferruginous ........

Abdomen black, the “two apical segments sometimes
ferr uginous red
. head,

thorax, legs, and abdomen black :
Lateral margins of the median segment not acute be

. Median segment twice as broad as the Jength in the

middle, the lateral carine sharply defined. An-
tenne fusco-ferruginous at the pa black at
apex ee

Median segment ‘about three times as "broad | as the
length in the middle, the lateral carme not
sharply defined. Antenne wholly orange

Head, thorax, and abdomen black

Head, thorax, and abdomen more or less red

Legs black. Punctures of front and pronotum coarse
and confluent longitudinally .........

Legs ferruginous. Punctures of front and pronotum
not coarse and well separated ..

. Apical or two apical segments of abdomen ferru-

ginous red, head and thorax black

Abdomen wholly black, head and sometimes thorax
more or less red et

Two apical segments of the abdomen red ; “fore v wings
fuscous except at the apex; sixth dorsal seement
punctured :

Apical segment of abdomen. only red ;

“wings hyaline ;
sixth dorsal segment striate NED

ts

FE. alicie Turn.

3.

*E. abdominalis Guér.

4.
5.

6,

*H. peringueyi Sauss.
*H. hottentota Sauss.

. adelogamia Turn.

. auriflua Turn.
. torrida Sm.

10.

. Sdussuret Turn.

. xanthocera Gerst.
11.
12%,
. nnotata Turn.
erythropoda Turn.
13.

14.

Hf. apicipennis Turn.

LE. pyxidata Turn.

The colour in the female is

704 MR. R. E. TURNER ON

14. Wings subhyaline; head, pronotum, mesonotum, and

scutellum red; vertex punctured ..................... Hi. heterogamia Sauss.
WATERS THOSCOMOEEROWE, Voaaochnooodsebonao sen vay 000 soa bog 000 15.
15. Vertex punctured ; pronotum closely age legs
blacked . E.hova Turn.
Vertex smooth; pronotum “sparsely ‘punctured ; ‘lees 7"
LOG haa tasjaas sehen casiseasinecn use scum tte memes eee :
NS, Jae] mack, moore NECK cosecococpacos onconsnonannAuoosannn E. ruficeps ruficeps Sm.
Bivoraxsmorelow less red Ase ss-cee ae eee eee Peete LE. ruficeps atopogamia
[Sauss.
Males.
1. Apical dorsal segment not incised at the apex ........ 2.
Apicil dorsal segment more or less incised at the
2) OLS iE PEERS o drs det Si eee Gan cen tmstad cone sue oc 4.
2. Third dorsal segment of the abdomen measured
from the transverse basal furrow distinetly shorter
than its basal width. Abdomen wholly without
Jellow markings; asso emeacw eyes nee en eee EF. incerta Turn.
Third dorsal segment measured from the transverse
basal furrow distinctly longer than its basal
width. Abdomen with very small ey
markings... 3.
3. Apical dorsal sezment flattened, ‘with raised mar gins
and a median car ina; pronotum entirely black .:. E asmarensis Turn.
Apical dorsal segment convex; the margins not raised,
without a distinct median carina ; pronotum with
a yellow band on the posterior margin .:,............ E. anetalla Turn.
4. Abdomen entirely black, sometimes with blue sheen . 5.
Abdomen banded with yellow Bese aee a acta ne CR Coc een egene 6.

5. Wings more or less shaded with fuscous or viola-
ceous ; Incisio# of the apical segment halfas deep _
as its apical width. Length 15- “48 mam: wi. ves ruficeps Sm.
Wings clear hyaline; incision of apical segment
much less tian half as deep as its ata width.
Length 13 mnt. . HH, nodosa Guér.
6. Apical “dorsal segment flattened with eal | margins ;
head large, the oheeay more than half as broad as
the eyes ..:.... .. FE, capitata Sm.
Apical dorsal samen convex, with a median. carina ;
head small, the cheeks much less than half as

brdad as the NESEY Sctoniaee deta a Uae uae GOR couloeaanebnads Ue
7. Not very sleider ; the apical slowed sacanery with an
incision nearly as deep as broad at the apex ie... Li. $poliaté Turn.

Very slender; incision df, the apical dorsal segment ‘
nét more than half as déep as its Apical breadth... #. lowgiventris Turn.

Exis (Musa) aticra, sp.n. (Pl. LXCXXI, fig. 12; Pl. LXX XII.
fig. 8; Pl. LX XXIII. fig. 9.)

Q. Nigra; segmentis abdominalibus qwinto sectoque rufo~
Jerruginers ; pedibus albo=pilosus, caleartis albidis; alis nigro-
ewruleis.

Long. 18 mm.

@. Head and thorax coarsely punctured-rugose, the punctures
on the front finer than on the vertex; clypeus very broadly
rounded at the apex, with two or three indistinct teeth on the
margin ; scape shining and sparsely punctured, clothed beneath
with long whitish hairs; the nine apical joints of the flagellum
opaque. Front thinly clothed with white hairs, the interantennal
prominence well developed and feebly bilobed. Inner margin of
the eyes slightly sinuate; posterior ocelli about twice as far from

STUDIES IN THE FOSSORIAL WASPS. 705

the eyes as from each other. Pronotum nearly as broad as the
head, widely emarginate anteriorly ; scutellum with a large
triangular rugose area from the base to the narrowly truncated
apex, the sides smooth and opaque. Postscutellum and median
segment smooth and opaque, a few large punctures on the middle
of the postscutellum ; median segment raised towards the median
line, with a narrow margined median groove, the posterior trunca-
tion of the segment coarsely but shallowly punctured, the sides of
the segment closely obliquely striated. Abdomen shining, finely
and very sparsely punctured, the punctures larger on the ventral
than on the dorsal surface, and rather closer at the apex of the
segments than at the base; the apical dorsal segment broadly
rounded, and longitudinally punctured-striate. Second abscissa
of the radius a little shorter than the third; first recurrent
nervure received beyond the middle of the second cubital cell,
second at two-thirds from the base of the third cubital cell.
Radial cell detached from the costa for about half its length.

Hab. British East Africa, Makindu, 3300 ft. (S. A. Weave);
April 5-7, 1911 (4. & R. C.).

This fine species does not seem to be very nearly allied to any
other, being well distinguished from the rujficeps group by the
very coarse sculpture of the head and thorax. The basal joint of
the posterior tarsi is furnished with a closely-set comb of small
spines beneath.

The description of Cosila donaldsont Fox corresponds rather
closely to this species, but the clypeus is tridentate on the apical
margin, not rounded, with indistinct teeth as in the present
species, and I think Fox was too careful a worker to have con=
fused the genera.

Eis (Mesa) AURIFLUA, sp. n.

2. Nigra; mandibulis basi, pronoto, mesonoto sculelloque rufo-
JSerrugineis ; alis infuscatis.

Long. 12 mm.

2. Head and pronotum closely and rather strongly punctured ;
mesonotum and scutellum much more sparsely punctured, smooth
in the middle} pleur coarsely punctured; median segment finely
and closely punctured, the median groove narrow and shallow,
margined by low caine, the posterior truncation more shallowly
punctured. Abdomen rather closely and not very finely punctured,
the apical segment closely longitudinally striated. Sides of the
median segment obliquely striated. Clypeus transverse at the
apex. Eyes vety feebly and widely emarginate on the inner
margin; posterior ocelli further from the eyes than from each
other. Head more than half as broad again as long, broader than
the thorax. Pronotm short, not as long in the middle as the
scutellum, the anterior margin straight, the posterior margin
widely arched. Pubescence sparse and whitish, calearia whitish,
Radial cell only separated from the costa at the apex, which is
subtruncate; first abscissa of the radius as long as the second,

706 MR. R. E. TURNER ON

but shorter than the third; recurrent nervures received close to
the middle of the second and third cubital cells.

Hab. Johannesburg, Transvaal (Kobrow). Received from
Dr. Brauns.

Nearly allied to adelogamia Turn. and diapherogamia Sauss.
From the former it may be distinguished by the entirely black
head, and by the closer and more even puncturation of the head
and pronotum, the vertex being almost smooth in adelogamia;
from diapherogamia it may be distinguished by the much shorter
pronotum, the stronger and closer punctuation, and the colour of
the head. #. hova has the pronotum longer than in the present
species or adelogamia.

The comb on the underside of the basal joint of the posterior
tarsi is present, but the teeth are few. ‘The upper spur of the
hind tibie is strongly bent near the base.

Exis (Mesa) AaDELoGAMiIA Turn.

Plesia (Mesa) adelogamia Turn. Ann. & Mag. Nat. Hist. (8) i.
p. 503 (1908), @.

Hab. Maseru, Basutoland; Lichtenburg, Transvaal.

This is nearly allied to #. auriflua, but differs as noticed in the
key.

Evrs (Mesa) ruricers Sm. (PI. LXXXII. figs. 9, 10;
Pl. LXXXITI. figs. 2, 4, 10, 15, 16.)

Myzine ruficeps Sm. Cat. Hym. B. M. iii. p. 75 (1855), ° ;
Turner, Ann. & Mag. Nat. Hist. (8) i. p. 503, ¢ ; Turner, Ann.
& Mag. Nat. (8) vii. p. 304 (1911), 5 @.

Eis (Musa) RUFIcEPS, subsp. ATOPOGAMIA Sauss.

Plesia( Mesa) atopogamia Sauss.in Grandidier, Hist. Madagascar,
xx. p. 244 (1892), ©.

Plesia (Mesa) diapherogamia Sauss. ; Distant, Naturalist in the
Transvaal, p. 225 (1892), 2.

Plesia (Mesa) disjuncta Turn. Ann. & Mag. Nat. Hist. (8) i.
p- 902 (1908), 3.

Lilis (Mesa) ruficeps, subsp. atopogamia and diapherogamia Turn.
Ann. & Mag. Nat. Hist. (8) vii. p. 304 (1911), 3 2.

Hab. Zanzibar; Nyasaland; Transvaal.

The colour-differences between atopogamia and diapherogamia
are not constant, though in the former the mesonotum is usually
red and in the latter black. The wings of the male are darker in
specimens from Nyasaland than in those from the Transvaal.
A single male received from Harar, in 8. Abyssinia, has the wings
entirely hyaline, slightly iridescent.

Ents (Mrsa) HerpRocamsA Sauss.

Plesia (Mesa) heterogamia Sauss. in Grandidier, Hist. Mada-
gascar, xx. p. 244 (1892), 2.

Hab. Delagoa Bay ; Manica; South Nyasaland.

STUDIES IN THE FOSSORIAL WASPS. 707

Euis (Mzsa) nova Turn. :
Pilesia (Mesa) hova Turn. Ann. & Mag. Nat. Hist. (8) i. p. 504
(1908), 2.

Hab. Madagascar.
Probably the female of nodosa Guér.

*Hiiis (MESA) ABDOMINALIS Gueéer.

Plesia abdominalis Guér. Rev. Zool. i. p. 57 (1838), 2.

Plesia (Mesa) abdominalis Sauss. in Grandidier, Hist. Madagas-
car, xx. p. 244 (1892), ©.

Hab. South Africa.

Eis (Musa) APICIPENNIS, sp. 0.

2. Nigra; segmentis abdominalibus quarto apice, quinto sextoque
rufo-ferrugineis ; mandibulis bast antennisque ferrugineis, cal-
cariis albidis ; alis fuscis, apice anguste hyalinis.

Long. 10 mm.

@. Clypeus finely punctured, broadly rounded at the apex.
Head sparsely and rather finely punctured, shining, almost
smooth round the anterior ocellus; interantennal prominence
bilobed, divided by a longitudinal suleus which does not reach the
anterior ocellus. Scape shining, very sparsely punctured; flagel-
lum opaque, the two basal joints shining. Posterior ocelli nearly
as far from each other as from the eyes. Pronotum and pleuree
closely but not coarsely punctured; mesonotum and scutellum °
more sparsely punctured; median segment opaque, finely
punctured, the punctures more or less confluent longitudinally,
the median groove shallow and not distinctly margined. Abdo-
men shining, finely and rather closely punctured, with a very
short petiole, the first segment broadly truncate at the base; the
sixth dorsal segment punctured, rounded at the apex. The sides
of the median segment are closely obliquely striated. Third
abscissa of the radius at least half as long again as the second 5
first recurrent nervure received just beyond the middle of the
second cubital cell, second at two-thirds from the base of the
third cubital cell. Radial cell not distinctly separated from the
costa, narrowly rounded at the apex.

Hab. British East Africa, Makindu, 3300 ft. (S. A. Weave);
April 5—/, L911 (4. 2. Re: C.).

Nearly allied to #. pyxidata Turn. from N.E. Rhodesia, but
differs in the broader shape of the third cubital cell, the position
of the second recurrent nervure, the colour of the scape, of the
wings, and of the fourth and fifth abdominal segments, in the
finer and sparser puncturation and in the sculpture of the
pygidium. The wings are clear hyaline beyond the radial and
cubital cells. Basal joint of the posterior tarsi with a scopa of
white hairs beneath ; spur of the posterior tibia bent near the
base.

708 MR. R. E. TURNER ON

Eis (Mesa) rorripA Sm.
Myzine torrida Sm. Desc. new spec. Hymen. p. 178 (1879), 9.

@. Nigra; mandibulis basi, scapo apice abdomineque toto
ferrugeners ; tegulis lestacets ; tibiis tarsisque fusco-ferrugineis ;
alis hyalinis, venis nigris.

Long. 12 mm.

. Clypeus short, transverse at the apex, subcarinate in the
middle, finely punctured at the base, smooth at the apex, Front,

ronotum, and mesopleure closely and strongly punctured, ver tex
a little more sparsely punctured; mesonotum and scutellum
coarsely but sparsely punctured; pronotum as long as the
scutellum. Median segment twice as broad as the length in the
middle, not margined, almost smooth, but not shining ; the median
groove narrow but well defined, with the margins of the groove
raised. Abdomen shining, with a few scattered punctures, the
apical dorsal segment finely longitudinally striated. Basal joint
of the posterior tarsi unarmed beneath, with a scopa of very fine
hairs. Radial cell detached from the costa at the apex, third
abscissa of the radius longer than the second, but a little shorter
than the first, second recurrent nervure received just beyond two-
thirds from the base of the third cubital cell.

Hab. Gambia (ex coll. Shuckard).

*Hiiis (MEsA) PERINGUEYI Sauss.

Plesia (Mesa) peringueyi Sauss. in Grandidier, Hist. Madagas-
car, xx. p. 245 (1892),

“ Areola radialis apice minute truncata. Secunda v. recurr. in
ipso medio margine tertiz ar. cubitalis exserta. Metatarsus
posticus subtus scopa spinarum brevium instructus. Majuscula,
nigra, nitida, cinereo-hirta. Caput et thorax cribrosa. Caput
validum, quam pronoto paulo latius. Metathorax leviusculus,
superne tenuiter punctatus, subtiliter carinatus (carina a latere
visa subbituberculata) utrinque pago polito; ejus facies postica
plana, subrugulata ; metapleura strigata. Hpipygium elongato-
trigonale, punctatum. Spinee tib. post. nigre, acute. Metatarsus
posticus subtus pectinatus, pilis vel spinis albidis intermixtis,
spinisque nonnullis dilatatis. Ale bruneo-nebulose, venis fuscis ;
2° ar. cubit. intus valde acuta; 2° vena recurrens transvevsalis,
cum v. discoidali angulum fere rectum efficiens.”

*Kiis (MmsA) HOTrENTOTA Sauss.

Plesia (Mesa) hottentota Sauss. in Grandidier, Hist. Madagascar,
xx. p. 245 (1892), ©.

“* Areola radialis apice minute truncata. Secunda v. recurr. in
ipso medio margine tertiz ar, cubitalis exserta. Metatarsus
posticus subtus scopa spinarum brevium instructus. Minor,
nigra, cinereo-hirta. Antenne imo apice flavo. Caput et thorax
sat tenuiter cribrosa. Caput quam thorax vix latius. Meta-
thorax lviusculus, subtilissime punctato-rugulatus, superne

STUDIES IN THE FOSSORIAL WASPS. 709

obsolete roundato-carinatus ; facie postica punctulata, supra
distincte angulata, obtusangula, fere rectangula. Epipygium
trigonale, striolatum, margine levi. Spine tib. post. albescentes,
squamose. Alz subhyaline venis bruneis et fuscis ; parte apicali
nebuiosa, 2* ar. cubit. intus breviter acute producta ; 2° vena
recurrens arcuata, obliqua, In alis posticis v, discoidalis longius
ultra venulam transverso-discoidalem furcata, Long 14 mill. ;
al, 10 mill. (Africa meridionalis),”

*Hiuis (Mesa) CAPENsIS Lep.

Tiphia capensis Lep. in Hist, Nat. Insect., Hym. iii. p, 554
(1845), ©.

“Caput nigrum, supra nigro, subtus rufo subvillosum. An-
tenne nigre, articulo primo nigro hirto. Thorax niger rufo
subhirtus. Abdomen nigrum, subnudum. Pedes nigri, rufopallido
subvillosi, femoribus duobus posticis angulatis compressisque.
Ale rufo- fuscee, nervuris costiaque vufo-fuscis ; squama, nigra,
Cellula radialis clausa. Femina,” ;

‘““ Long. 7 lignes.

“ Cap ‘de Bonne Esperance. Musée de M. Serville.”

The figure shows that this species is an His.

Enis (Mesa) innorata Turn.

Plesia (Mesa) innotata Turn. Ann. & Mag. Nat. Hist. (8) i,
p- 506 (1908), @.

Hab. Loangwa River, N.E. Rhodesia; 8. Nyasaland.

Exits (MesA) SAUSSUREI Turn.

Plesia (Mesa) saussureti Turn, Trans. Ent. Soc. London, 1910,
p- 394, °.

Hab. Madagascar.

This is near #. xanthocera, but the median segment is longer in
proportion and much more ‘distinctly marg gined; the colour of
the antenne is also different.

Exits (MzsA) XANTHOCERA Gerst,

Myzine «xanthocera Gerst. Arch, f. Naturg, xxxvii. p. 353 (1870),
2; v.d. Decken, Reise in Ost-Afrika, Gliedethiere, p. 339, pl. 14,
ime, (USS), Qe

Plesia (Mesa) xanthocera Sauss, in Grandidier, Hist. Madagas-
car, xx. p. 245 (1892),

Hab. Howick, Natal; Zoutpansberg, Transvaal; Mozambique ;
Harar, Abyssinia.

Eis (Mesa) ERYTHROPODA Turn,

Plesia (Mesa) erythropoda Turn. Ann. & Mag, Nat. Hist, (8) i.
p- 505 (1908), @.

Hab. Lake Ngami.
The scopa beneath the basal joint of the hind tarsus is rather

710 MR. R. E. TURNER ON

coarse at the base, but I do not think that any spines are
present.

Eis (Mzsa) INCERTA, sp. n.

3. Niger, cano-pilosus ; clypeo macula mediana minuta, tabiis
anticis supra pallide flavis ;. tegulis pedibusque fuscis; alis
hyalinis, venis nigris ; pygidio haud inciso.

Long. 13 mm.

¢. Front coarsely reticulated, the vertex punctured ; thorax
closely but not coarsely punctured, median segment punctured-
rugose, abdomen finely and closely punctured. Pronotum shorter
than the scutellum, the anterior margins straight, the angles
subacute. First abdominal segment petiolate, the narrow petiole
scarcely more than half as long as the dilated apical portion of
the segment, which is constricted at the apex; second segment
twice as broad at the apex as at the base, all the segments slightly
constricted at the base. Apical dorsal segment sparsely punctured,
without an incision, - somewhat convex, subcarinate in the middle,
pointed at the apex, the lateral margins raised near the apex.
Third abscissa of the radius a little longer than the second, which
is fully twice as long as the fourth. First transverse cubital
nervure oblique, sharply bent close to the cubitus, second
recurrent nervure received just before one-third from the base of
the third cubital cell, curved outwards below the middle, the ends
slightly curved inwards.

Hab. Howick, Natal (J. P. Cregoe); Cape Colony.

This is very near the description of Plesia carbonaria Cam., but
in that species the seventh dorsal segment is said to be shortly
incised at the apex, and the fourth abscissa of the radius is almost
as long as the second or third.

Euis (Mesa) capiraTa Sm.
Myzine capitata Sm. Cat, Hym. B.M, ii. p. 74 (1855), 3.

3. Niger, albido-pilosus ; clypeo, mandibulis, linea obliqua
uiringue supra antennas, pronoto fascia angusta postice, tegulis
basi, segmento dorsali primo fascia apical emarginata, segmentis
dorsalibus et ventralibus 2-6 fascia bisinuata angusta flavis ;
pedibus flavis nigro-variegatis ; alis hyalinis, venis testacets.

Long. 14-17 mm.

$. Head large, much broader than the thorax, the cheeks more
than half as broad as the eyes; antenne stout, the apical joint
truncate at the apex. Head rugose, thorax closely punctured,
median segment punctured-rugose; abdomen shining, glossed
with blue, very finely and closely punctured. Pronotum shorter
than the scutellum, slightly narrowed anteriorly, the anterior
margin straight, the angles not prominent, posterior margin
widely arched. First abdominal segment petiolate, the petiole
only about half as long as the rather strongly swollen apical

STUDIES IN THE FOSSORIAL WASPS. 711

portion, the apex slightly constricted. Apical dorsal segment
flattened, subcarinate in the middle, the lateral margins raised
and nearly parallel, the apical incision subtriangular, not quite as
deep as the apical width. Second and third abscisse of the radius
nearly equal in length ; second recurrent nervure strongly curved
outwards below the middle, joining the cubitus just before one-
fourth from the base of the third cubital cell.

Hab. Johannesburg, Transvaal; Kroonstad.

The type is much damaged and without wings; the details of
neuration are taken from a more recent specimen in which the
cubitus of the hind wing on the right side is almost interstitial
with the transverse median nervure, but is further removed
towards the base on the left side.

Huts (MEsA) SPOLIATA, sp. n.

3. Niger; mandibulis, clypeo macula parva nigra utrinque,
linea utrinque supra antennas, pronoto angulis anticis et fascia
postice, tegulis, segmento dorsali primo fascia angusta apicali,
segmentis dorsalibus et ventralibus 2-6 fascia bisinuata angusta,
coxis apice, femoribus posticis supra, tibiis anticis et intermediis
supra tarsisque subtus pallide flavis; pedibus rufo-testaceis ; alis
hyalinis, venis nigris, stigmate rufo-testaceo.

Long. 13 mm.

3. Clypeus short and broad, rather narrowly produced in the
middle and very feebly emarginate at the apex. Eyes widely
emarginate ; cheeks very much narrower than the eyes ; posterior
ocelli more than half as far again from the eyes as from each
other. Antennal tubercles well developed ; antennz longer than
the head, thorax, and median segment combined, of even thick-
ness throughout. Head punctured-rugose, much wider than the
thorax; the whole thorax finely and closely punctured, with
sparse white pubescence ; pronotum a little longer in the middle
than the scutellum, narrowed anteriorly, the anterior margin
straight, posterior margin strongly arched. Median segment
rounded posteriorly, not truncate, very closely but not coarsely
punctured. Abdomen slender, slightly shining, very finely and
sparsely punctured, petiolate; the petiole occupying the basal
third of the first segment, the apical two-thirds elongate pyriform,
the whole segment half as long again as the second, which is
gradually broadened from the base, a little longer than the apical
width ; third segment broader than long. Apical dorsal segment
slightly convex, subcarinate longitudinally in the middle, shining,
with a few large punctures, the apical emargination almost as
deep as its breadth at the apex. Radial cell pointed; second
abscissa of the radius shorter than the third, but much longer
than the first; second recurrent nervure received at one-third
from the base of the third cubital cell.

Hab. Algoa Bay, Cape Colony; March (Dr. Brauns).

Nearly allied to #, capitata Sm., but in that species the head

712 MR. R. E. TURNER ON

is larger, the cheeks much broader, and the first abdominal
segment a little shorter and more swollen towards the apex.
The apical dorsal segment in capitata is flat, not convex, and
has the sides distinctly raised into marginal carine.

Enis (Mzsa) LONGIVENTRIS, sp. n. (PI. LXXXI. fig. 13;
Pl. LXX XIII. fig. 11.)

3. Niger; mandibulis, clypeo, linea obliqua utrinque supra
antennas, pronoto fascia angusta postice, segmento dorsali primo
Jascia angusta apicali, segmentis dorsalibus et ventralibus 2-6 fascia
angusta apical bisinuata, coxis subtus. tibtis tarsisque anterioribus
supra, intermediisque subtus albido-flavis; alis hyalinis, venis
negris.

Long. 10-13 mm.

3d. Very slender. Head rugose ; thorax strongly and closely
punctured, median segment punctured-rugose, Antenne rather
slender, as long as the head, thorax, and median segment com-
bined, of even thickness throughout. Pronatum longer than the
scutellum, the sides almost parallel, the anterior margin straight,
with acute angles, the posterior margin widely arched. Sides of
the head and thorax rather thicky clothed with long white
pubescence. Median segment longer than broad, rounded pos-
terlorly. Abdomen slender, shining, very finely punctured ; the
basal segment as long as the secand and third combined, petiolate,
the narrow petiole nearly as long as the slightly swollen apical
portion ; second segment broadened from the base, about equal
in length to the third, the segments scarcely constricted at the
base; the apical dorsal segment convex, subcarinate in the
middle, sparsely punctured, the punctures large, the apical
incision shallow, not as deep as its breadth at the apex. Wings
reaching to the apex of the fourth dorsal segment; second and
third abscisse of the radius usually about equal in length, the
fourth distinetly shorter ; second recurrent nervure received just
before one-third from the base of the third cubital cell, curved
outwards in the middle.

Hab. Willowmore, Cape Colony (Dr. Brauns).

This corresponds rather closely with the description of Plesia
meisa Cam,, but in that description the incision of the apical
dorsal segment is said to be twice longer than wide, and some of
the details of neuration are not quite the same. But the latter
character is of little importance in the genus, especially as to the
length of the second and third abscisse of the radius. Many
variations occur which are not of specific value.

Enis (Mesa) AsMARENSIS Turn,

Plesia asmarensis Turn, Ann. & Mag. Nat, Hist. (8) iii, p: 481
(1909), 3. | .

Tkab. Krythrea,

he

STUDIES IN THE FOSSORIAL WASPS. 713

Enis (Musa) AMETALLA Turn.

Elis (Mesa) ametalla Turn. Ann. & Mag. Nat. Hist. (8) vil.
joes 0259 GAS) ecg

Hab. 8. Nyasaland.

Almost certainly the male of innotata,

Enis (Mesa) Noposa Guer.

Myzine nodosa Guér, Dict, pitt. hist, nat. v. p. 584 (1837), 3;
Sauss. in Grandidier Hist. Madagascar, xx. p. 240 (1892), ¢@.

Hab. Madagascar.

* His (Mzsa) CLAVATA Sauss,

Myzine clavate Sauss, in Grandidier, Hist, Madagascar, xx.
p. 242 (1892), ¢

Hab. Transvaal.

*Exis (Mesa) CARBONARIA Cam.

Plesia carbonaria Cam, Rec, Albany Mus. i, p, 317 (1905), ¢.
fab. Dunbrody, Oape Colony,

*Hiiis (Musa) RETICULATA Cam.

Plesia reticulata Cam, Rec, Albany Mus, i. p, 300 (1905), ¢
Hab. Brak Kloof, Cape Colony,

* Huis (MESA) RUFO-FEMORATA Cam,

Plesia rufo-femorata Cam, Rec, Albany Mus, i.p. 298 (1905), 3
Hab. O’okiep, Cape Colony,

* Huis (Mesa) mncisa Cam,

Plesia incisa Cam. Rec. Albany Mus. 1, p. 320 (1905), 3
Hab. Dunbrody, Cape Colony,

Key to the Oriental Species of Klis (Mesa).

Females.
1. Sixth dorsal segment longitudinally striated ............ 2.
Sixth dorsal segment punctured... ...............00..2. 2000s 8.
2. Abdomen more or less ferruginous Be
Abdomen wholly black : 4.
3. Clypeus with a median carina ; ; two ‘apical ‘abdominal
segments and the apex of the fourth black............ H. dimidiata Guér.
Clypeus without a distinct carina; apical abdominal
segment only black ............ ...... H. mandalensis Magy.
4. Median segment distinctly margined posteriorly | ae HL. fuscipennis Sm.
Median segment not ee margined pees ee :
5. Wings FRED US: oe tira ete tela wry 2A 6.
Wines subhyaline... uy ney Cee ao neo ee eae We
6. Pronotum longitudinally rugose ; “second abscissa of
the radius as long as the third. Length 16 mm.... 2. mandibularis Sm.

Pronotum coarsely punctured ; second abscissa of the
radius much shorter than the third. Length 10mm. E. petiolata Sm.

Proc. Zoou. Soc.—1912, No. XLVII. AZ

714 MR. R. E.. TURNER ON

ae

8.
Ss)

10.

11.
12.
13. Mesonotum closely punctured

14.

Propleure sparsely punctured
Propleurwe tinely striated.. by eee
Abdomen not marked with ‘yellow : ea eas
Abdominal segments with yellow apical hanidsn, .aee
Head black ; abdomen black or black and ferrugimous.
Head red; abdomen steel-blue ................0222..0e een
Abdomen mostly ferruginous.. Sergcumsens
Abdomen entirely black, or with ‘the apice al segment
only ferruginous ..... Stemess
Four basal segments of the abdomen ferruginous ae als
Four apical segments of the abdomen ferruginous ......
Apical segment of the abdomen ferruginous cig:
Apical seement of the abdomen black PELE nama te wee ats

Mesonotum shining, almost impunctate ..
Lees ferruginous; head coarsely and closely punctured.
Legs black ; head sparsely punctured

Fi, claripennis Bingh.
HH. ustulata Tourn.
9.

*E. picticollis Moy.

10.
FL. tricolor Sm.
11.

FH. bengalensis Cam.
E.. apimacula Cam.
13.
14,

... *H. fedtschenkoi Sauss.
. *H. dubia Mor.

HH. rothneyi Cam.
EL. opacifrons Turn.

Myzine anthracina yecordel by Bingham as Indian is
Australian and belongs to the genus Anthobosca; and Myzine

c=)

combusta also recorded as Indian is undoubtedly West Indian,
and a true His.

1.

~

Males.

First dorsal segment elongate, age more or less
nodose at the apex ; Ra. cen aac eae

First dorsal segment subsessile, “not nodose at the
apex, broader “at the apex than long..

Seventh dorsal segment distinctly incised at the apex
abdomen wholly black.. pay

Seventh dorsal segment not incised at the apex pepedaace

. Legs black; seventh dorsal segment smooth and

SINS AU ERE Bem ee oan ee mene ae
Lees fusco - ferruginous; seventh dorsal segment
Gommselhy FOUMCUMINEC! 2.4200 002 aso cov pd coo cao cca boo eee bod san 005

4. Abdomen wholly black, without yellow markings,
sometimes glossed with blue .. PanaS atte acy Ease
Abdomen more or less marked with yellow. Peeebemacedtcd:
5. Median segment without a distinct sulcus; pygidial

area clearly detined, with marginal caringe reaching
nearly to the base of the sesment . Ree ae.

Median segment with a distinct median. “sulcus;
pygidial area less clearly defined, the marginal
carine lower and only on the apical third of the
segment .

. First abdominal seoment very little, if at all, longer

than the hind femur and trochanter combined ee
First abdominal segment much longer than the hind
femur and trochanter combined ..................0.. 008

. Head punctured-rugose ; ee without a Bee

band .
Head punctured ; pronotum, with a “yellow band on the
posterior margin ......

. Wings hyaline ; “pronotum ‘with | a yellow band c on : the

posterior margin, closely punctured ..
Wings pale fusco- hy aline ; pronotum without a | yellow
band, rather sparsely punctured .

. Narrow. petiole of first abdominal sezment scarcely

more than half as long as the sw ollen apical portion
ot the segment .

Narrow petiole of first abdominal ‘segment ‘nearly a as
long as the swollen apical portion of the segment...

Myzine pallida Sm. and Myzine orientalis Sm., placed by Bing-

iy

ae

E. dimidiaticornis
[Bingh.
3.
A,
E. burmanica Bingh.
HH. leta Bingh.
5.
6.

EL. dimidiata Guér.

HH. mandibularis Sm.
To
8.

HH. nurset Turn.

HEH. mandalensis Magy.
9.

FH. extensa Turn.

EH. petiolata Sm.
LA. claripennis Bingh.

t=)

ham and Smith in J/yzine, together with many species of is,

STUDIES IN THE FOSSORIAL WASPS. 715

belong to the genus Zswara Westw., which also belongs to the
Elidine.

Huis (Mesa) Dimrpiara Guer.
Myzine dimidiata Guér. Dict. pitt. hist. nat. v. p. 584 (1837),

Methoca orientalis Sm. Cat. Hym. B. M. iii. p. 66 (1855),
(nec Smith, 1875).

Myzine madraspatana Sm. Cat. Hym. B. M. iii. p. 72 (1855), @.

Myzine violaceipennis Cam. Mem. Manchester Lit. & Phil. Soe.
sdhina jon All (Cusihs)), 2

Hab. The whole of India, except the North-West.

@. Head and thorax coarsely rugosely punctured, smooth
round the anterior ocellus and at the base of the mesonotum ;
median segment opaque, with a narrow median sulcus in which is
a low carina, the sides of the sulcus raised and forming low
carine, the segment not margined at the base of the posterior
truncation. Radial cell distinctly separated from the costa for
more than half its length, third abscissa of the radius very
distinctly longer than the second. Sixth dorsal segment finely
longitudinally striated ; basal joint of hind tarsi with a scopa of
short hairs beneath.

Black ; second, third, and the base of the fourth abdominal
segments ferruginous. Oalcaria whitish ; wings fusco-violaceous.

3. Head and thorax closely punctured; median segment
rugose; basal third of first abdominal segment forming a narrow
petiole, apical two-thirds swollen and slightly constricted at the
apex, the whole segment more than half as long again as the
second segment. Abdomen shining, very finely punctured ;
pygidial area coarsely punctured, fully twice as long as broad,
the sides raised and forming carine, a well-defined median carina,
the apex narrowly truncate, not emarginate.

Black ; the abdomen slightly glossed with blue; base of the
mandibles, fore tibie im front, fore tarsi in front, base of the
tegule and the hind margin of the pronotum narrowly and
* obscurely pale yellow. Wings hyaline at the base, fusco-hyaline
beyond the basal nervure.

Length 17 mm.

The association of the sexes was suggested by me in 1908, and
positive proof was published by Mr. Lefroy a year later, a pair
having been taken im coitu by Mr. Dutt at Pusa.

Eis (MssA) MANDIBULARIS Sm.

Methoca mandibularis Sm. Trans, Ent, Soc, London, p, 301
(i869). or

Plesia (Mesa) mandibularis Turn. Ann. & Mag, Nat. Hist, (8)
1. p. 509 (1908), 3

Plesia (Mesa) purpureipennis Turn. Ann. & Mag, Nat. Hist.
(8) 1. p. 508 (1908), ©.

This will almost certainly prove to be the Chinese subspecies

47T*

716 MR. R. E. TURNER ON

of ZL. dimidiata Guér., from which the female differs in the
entirely black abdomen and the greater length of the second
abscissa of the radius, which is nearly as long as the third. ‘The
male differs from dimidiata in having the second abscissa of the
vadius very distinctly longer than the third, in the paler colour
of the apical portion of the wings, in the presence of a distinct
longitudinal sulcus on the median segment, and in the somewhat
shorter and broader form of the first abdominal segment. The
pygidial area of the male is also wider and less distinctly margined
than in dumidiata.

ITab. Shanghai.

Kis (Musa) BENGALENSIS Cam.

Myzine bengalensis Cam. Mem. Manchester Lit. & Phil. Soe.
slits joy MUL (CiSBke),

The four basal abdominal segments are ferruginous ; the wings
violaceous, the base of the hind wings hyaline. Pronotum
sparsely but coarsely punctured.

Length 15 mm.

This is quite distinct from dimidiata, and seems, as Cameron
suggests, to be more nearly related to mandalensis.

Eis (Masa) MANDALENSIS Magr.

Plesia mandalensis Magy. Ann. Mus. Civ. Gen. (2) xii. p. 257
(1892), @.

@. The five basal abdominal segments are ferruginous.
Calearia and spines of the hind tibiz whitish. Wings hyaline,
dark fusco-hyaline beyond the basal nervure of the fore wing
to the apex, the apex of the hind wing fusco-hyaline. Pronotum
closely and somewhat coarsely punctured; sixth dorsal segment
finely longitudinally striated.

$. Head and thorax closely and rather finely punctured ;
median segment punctured-rugose ; abdomen shining, micro-
scopically punctured. First abdominal segment scarcely longer
than the hind femur and trochanter combined, the narrow petiole
a little more than half as long as the moderately swollen apical
portion of the segment; second segment longer than the third
by about one-quarter. Seventh dorsal segment pointed, not
incised ; pygidial area well defined, very narrow, coarsely
punctured and with a well-marked median carina, Third
abscissa of the radius distinetly longer than the second.

Black; mandibles at the base, clypeus, apex of the inter-
antennal prominence, posterior margin of the pronotum, tegule,
an apical band on dorsal segments 1—6, strongly bisimuate on
segments 2-6 and on ventral segments 2-6, tarsi, fore and
intermediate tibiz in front, and the base of the hind tibiz yellow.
Wings hyaline, nervures fuscous.

Length, 2 10 mm., ¢ 10-11 mm.

Hab. Mandalay, Burma.

Taken in copula by Colonel Bingham.

|
|

~~

STUDIES IN THE FOSSORIAL WASPS. Tle

Huis (Mesa) roruneyi Cam.

Myzine rothneyi Cam, Ann. & Mag. Nat. Hist. (7) x. p. 88
(1902), ©.

2. This fine species is easily distinguished, being black with
ferruginous legs; the wings fusco-hyaline, flushed with purple.
The sixth dorsal segment is coarsely punctured, the punctures
tending to become confluent longitudinally towards the apex.
Head and thorax very coarsely punctured, rugose on the
pronotum.

Hab. Khasi Hills, Assam.

Eis (MzsA) FUSCIPENNIS Sm.

Myzine fuscipennis Sm. Cat. Hym. B. M. ii. p. 72 (1855), 2 ;
Bingh. Fauna Brit. India, Hymen. i. p. 67 (1897), 2.

@. This species may be distinguished from the female of
petiolata Sm., which it closely resembles, by the sharply margined
median segment, the posterior slope of which is abrupt and steep,
not gradual as in petiolata. The colour is black; the calearia
whitish, mandibles fusco-ferruginous, wings fuscous. ‘The spines
on the outer margin of the hind tibiz are black in fuscipennis,
whitish in petiolata.

Length 12 mm.

Hab. India.

The type is unique in the British Museum collection and was
obtained from Shuckard in exchange, so that the locality is
uncertain. Bingham’s description is taken from the type, but
he evidently confused the species with petiolata, a specimen of
which is labelled fusctpenmis by him in the British Museum
collection. I have not seen the specimens he records from
Burma, but consider it very doubtful if they belong to this
species.

Exts (Mesa) PerioLata Sm.

Myzine petiolata Sm. Cat. Hym. B. M. iii. p. 72 (1855), 3.

Myzine ceylonica Cam, Ann. & Mag. Nat. Hist. (7) v. p. 18
(S00) oe

Plesia (Mesa) petiolata Turn. Ann. & Mag. Nat. Hist. (8) i.
p. 512 (1908); 3) O-

®. Differs from fuscipennis Sm. in the absence of a distinct
margin separating the dorsal surface of the median segment
from the surface of the posterior slope and in the white spines of
the hind tibie. From claripennis Bingh. it differs in the fuscous
colour of the wings, the somewhat coarser puncturation, and the
lesser length of the second abscissa of the radius, which is only
about half as long as the third in petiolata and almost or quite as
long as the third in claripennis. In fuscipennis the second
abscissa of the radius is shorter than in peticlata, being distinctly
less than half as long as the third.

18 MR. R. E. TURNER ON

3S. This is very near the male of claripennis, but may be
distinguished by the less elongate petiole, the narrow basal
portion of which is scarcely more than half as long as the
moderately swollen apical portion of the segment.

Length, ¢ 2, 10 mm.

Hab. Bengal, Bombay, and Ceylon.

Eis (Musa) CLARIPENNIS Bingh.

Myzine claripennis Bingh. Fauna Brit. India, Hymen. i. p. 68-
(EIN), 2 -

Myzine hortata Nurse, Journ. Bombay Nat. Hist. Soc. xiv.
joo toll (IO2), Qe

@. Differs from petiolata as noticed under that species.
Bingham’s description of the species is inaccurate as to the
median segment, and this has misled Nurse. The segment is not
smooth and shining, and the longitudinal impression, though
not very long, cannot be said to be triangular.

3. The male, which has not been previously described, closely
resembles petiolata Sm. Head and thorax closely punctured,
most coarsely on the front, median segment rugose; abdomen
very slender, shining, minutely punctured. First abdominal
segment more than half as long again as the hind femur, the
narrow petiole very little shorter than the feebly swollen apical

ortion ; second segment very narrow at the base, nearly half as
éng again as the third segment. Seventh dorsal segment not
imciced) at the apex, pointed, convex and without a distinct
pygidial area.

Black ; mandibles, clypeus, the apex of the interantennal
prominence, posterior margin of the pronotum, an apical narrow
band on dorsal segments 1-6, strongly bisinuate on segments
9-6, base of the tecule, fore tarsi, anterior and intermediate
tibize in front, base of hind tibize and base of intermediate and
hind tarsi pale yellow. Wings hyaline, nervures fuscous.

Length, ¢ 8mm., 9 10 mm.

Hab. Burma, Ceylon, Bengal, and Deesa.

The male has the two basal abdominal segments more slender
than in petiolata, the first with the apical portion less swollen,
the basal narrow: petiole longer in proportion ; the second
narrower at the base.

Eurs (Mesa) USTULATA Turn.

Plesia (Mesa) ustulata Turn, Ann. & Nag. Nat. Hist. (8) i.
p- 510 (1908), 2.

Q. This is nearest to claripénnis Bingh., but is a larger and
more robust species, somewhat more closely punctured, with the
wings distinctly darker and the punctures oh the abdomen
larger. ,

Hab. Yunzalin Valley, Tenasserim.

a

Se

STUDIES IN THE FOSSORIAL WASPS. 719

Eis (Mzsa) opactrrons Turn. -

- Plesia (Mesa) opacifrons Turn. Ann. & Mag. Nat. Hist. (8)
1. p. 509 (1908), 9.

2. This black species may be distinguished by the very sparse
puncturation of the head, pronotum, and sixth dorsal segment.
The wings are pale fusco-hyaline. It is a larger and more robust
species than petiolata.

Hab. Salwen Valley, Tenasserim.

*His (Mesa) pubra Mor.
» Plesia dubia Mor. Hor. Soc. Ent. Ross. xxiv. p, 627 (1890), @.

This species is black, with the apical abdominal segment,
tarsi, hind tibize, mandibles, and antenne beneath ferruginous,
the mesonotum shining and almost impunctate, the sixth dorsal
segment finely and closely punctured.

Length 113 mm.

Hab. Turkestan.

Presta (Musa) APIMACULA Cam.

Myzine apimacula Cam. Journ. Bombay Nat. Hist. Soe. xiv.
p. 272 (1902), ©.

2. This is allied to dubia Mor., but differs in having the four
apical abdominal segments ferruginous, the wings paler, and the
puncturation somewhat different.

Hab. Deesa, N.W. India.

The male described as belonging to this species by Colonel
Nurse is Pecilotiphia albomaculata Cam., but has three cubital
cells instead of two as in the type of the species, which is
evidently an aberration from the usual neuration. The male
has the form of a Myzine, not of a Mesa, but it is by no means
improbable that Nurse is correct in the association of the sexes,
though he informs me that he does not recollect his reasons for
placing them together. The apical dorsal segment is deeply
incised as in Myzine. Cameron’s action in forming a new genus
on a specimen with abnormal neuration is quite unjustified and
due to ignorance of the variable character of neuration in the
Scoliidze, but I am at present doubtful if Pecilotiphia should be
treated as a synonym of Myzine or of Klis (Mesa).

*Huis (MmEsA) FEDTSCHENKOI Sauss.

Plesia jfedischenkot Saussure in Fedtschenko: Turkestan,
Scoliide, p. 29 (1880), 2 (Plesia tartara on plate).

Plesia tartara Sauss. 1. c. pl. 11. fig. 12, 2.

This seems to differ from dubia Mor. in the coarser and closer
puncturation of the mesonotum. The name tartara is used on
the plate through a fault in the editing, and has unfortunately
been recorded in Dalla Torre’s Catalogue as a distinct species.

720 MR. R. E. TURNER ON

*Hirs (Mesa) prorrconiis Mor.

Plesia picticollis Mor. Hor, Soc. Ent. Ross. xxiv. p. 624 (1890), 2 .

This appears to be a very distinct species, strongly marked
with yellow on the head, thorax, and abdomen. The colour would
suggest an Anthobosca allied to A. arabica Turn., but the structure
of the apical abdominal segment and the indistinct striation of
the metapleure render it unlikely that it belongs to that genus.

Eis (Mesa) TRIcoLOR Sm. (PI. LXXXT, fig. 16; Pl. LX XXII.
eres)

Myzine tricolor 8m. Journ. Linn. Soc., Zool. ii. p: 91 (1858), 2.

This fine species is easily distinguished by the large size
(19 mm.), the bright red head, and steel-blue abdomen. The
wings are fusco-hyaline, flushed with purple, almost hyaline at
the base. The head is large, quadrate, sparsely punctured,
smooth on the vertex; pronotum coarsely punctured, median
segment subconcave on the posterior slope; sixth dorsal segment
strongly punctured. Second abscissa of the radius nearly as
long as the first and third combined. :
Hab. Borneo (typical) ; Dibrughar, Assam; W. India (7. R.
Bell).

mh only specimen I have seen from Assam has the head
distinctly longer than broad, longer behind the eyes than in the
typical form, and the scape and three basal joints of the flagellum
are red; the median segment is not at all concave on the posterior
slope. The differences will probably prove to be subspecific.

Mr. Bell informed me that he bred this species from the larva of
a longicorn beetle.

Euis (Mesa) ? pimrpraticornis Bingh. (Pl. LXX XI. fig. 15.)

Myzine dimidiaticornis Bingh. Journ, Linn. Soc., Zool. xxv.
p. 423 (1896), 3 ; Turn. Ann. & Mag. Nat. Hist. (8) i. p. 501
(1908), 3.

¢. Antennte stout, a little longer than the thorax and median
segment combined. Front rugose, vertex sparsely punctured.
Pronotum transversely rugulose, much longer than the meso-
hotum; the sides almost parallel. Head slightly narrowed and
produced from behind the eyes. Thorax and median segment
rugose, the mediah segment distinctly margined posteriorly and
vertically truncate. First abdominal segment vertically truncate
anteriorly, with a distinct transverse carina above the base of
the truncation, attached to the abdomen by a very short petiole,
nearly as broad as the second segment, the sides parallel.
Apical dorsal segment not incised at the apex. Without a pygidial
area. Second abscissa of the radius as long as the first and third
combined.

Black; scape and four basal joints of the flagellum, clypeus,
and apex of the interantennal prominence dull ferruginous ;
abdomen glossed with steely blue. Wings hyaline; fusco-

STUDIES IN THE FOSSORIAL WASPS. (Pal

violaceous from the basal nervure of the fore wing to the apex
and at the apex of the hind wing.

Length 13 mm.

Hab. Kumaon, N.W. India.

Although this resembles Myzine rather than His in the form
of the first abdominal segment and the antenne, I consider that
it will probably be found to be the male of #. tricolor Sm. The
proportions of the cubital cells, the colour of the abdomen and
antenne, and the colour of the wings are all very similar in the
two forms and unlike any other species, either of Myzine or Elis.
I consider it probable that the group of Hiis containing dubia,
fedtschenkoi, and apimacula will also be found to have males
showing the facies of Mysine rather than of Hlis. These cases
render it very difficult to reach absolute certainty in distinguishing
the males of Hlis from IMyzine, though all males with a long
petiole may be assigned to Eis.

I think JJethoca rugosa Cam. (Mem. Manchester Soc. 1896)
from Ceylon will prove to be a local form of this species, and also
an individual aberration as far as the loss of one of the transverse
cubital nervures is concerned. It is certainly not a JMJethoca.

Eis (Mzsa) nurse Turn.

Plesia nursei Turn. Ann, & Mag. Nat. Hist. (8) iii. p. 480
(ESOS) Sie

The first abdominal segment is no longer than the hind femur
and trochanter combined, the narrowed petiole being rather less
than half as long as the strongly dilated apical portion of the
segment. In general appearance there is a strong resemblance
to petiolata, but there is no yellow band on the pronotum and
sixth abdominal segment. It is also a larger species, and the
shorter petiole is a good distinguishing character, in which the

resemblance is nearer to mandalensis than to petioluta.
Hab. Simla.

Ents (Masa) EXTENSA Turn.

Plesia (Mesa) extensa Turn, Ann. & Mag. Nat. Hist. (8) i.
p. 511 (1908), 3.

The first abdominal segment is much longer than the hind
femur and trochanter combined ; the antenne are longer than the
head, thorax, and median segment combined. ‘The wings are
strongly suffused with yellowish brown, and the yellow bands on
the abdomen are reduced to short transverse lines on each side of
dorsal segments 2-5.

Length 12 mm.

Hab. Upper Burma.

Eis (Mzsa) tara Bingh.

Myzine leta Bingh. Faun. Brit. India, Hymen. i. p. 70
(1897), ¢.

dé. Black; the legs fusco-ferruginous; wings hyaline. First

722, MR. R. E. TURNER ON

abdominal segment a little longer than the hind femur and
trochanter combined, the narrow petiole about half as long as
the dilated apical portion ; seventh dorsal segment distinctly
incised at the apex, the incision nearly as deep as its apical
width ; second segment about half as long again as the third.
Second abscissa of the radius distinetly longer than the third.
Pygidial area clearly defined and strongly “punctured, Hind
tibia more distinctly serrate than in other species of the genus.

Length 9-11 mm.

Hab. Moulmein, Tenasserim.

Exis (Mesa) surmaAnica Bing.

Myzine burmanica Bing. Faun. Brit. India, Hymen. i. p. 70
(USB), és

3. This differs from deta in the black colour of the legs; the
pygidial area is flatter, less distinctly punctured, and not distinctly
margined, and the third abscissa of the radius is distinctly longer
than the second. Whether these differences are of specific value
or not it is not easy to say from a single specimen of each, but I
am inclined to regard them as merely variations of one species.

fab. Amherst, Tenasserim.

ELIS SELLOWI, sp. n.

@. Nigra ; postscutello linea transversa, segmento dorsali primo
macula magna utringue, segmentisque dorsalibus 2-5 fascia basali
angusta utrinque flavis ; femoribus, tiblis tarsisque rufo-testaceis ;
mandibulis fusco-ferrugineis ; alis fusco-hyalinis, costa obscura,
venis fuscis.

Long. 11-13 mm.

2. Clypeus strongly punctured, broadly rounded at the apex,
with a longitudinal carina from the base almost reaching the
apex. Front strongly and closely punctured, vertex and ocellar
region sparsely and less deeply punctured, cheeks smooth, an
arched impressed mark above the posterior ocelli; scape strongly
punctured. Thorax deeply and closely punctured, especially on
the pronotum and mesopleure, the disc of the mesonotum and
the scutellum rather sparsely punctured; propleure obliquely
striated. Median segment closely punctured in the middle,
opaque and almost smooth near the anterior angles; the posterior
slope almost vertical, concave in the middle and shagreened, with
short transverse strize on the sides, the sides of the segment
strongly striated. Abdomen shining, with a few scattered
punctures; the apical segment closely longitudinally striated,
with a few large punctures between the striz, very broadly
rounded at the apex. The three abscisse of the radius almost
equal in length; first recurrent nervure received just beyond the
middle of the second cubital cell, second just beyond one-quarter
from the base of the third cubital cell, The ventral abdominal

STUDIES IN THE FOSSORIAL WASPS. 723

segments are very sparsely punctured, with a row of piliferous
punctures near the apex ; the pubescence is whitish.
Hab. San Juan del Rey, Brazil (Sellow). Two specimens.
Type in the Berlin Museum.

KLIS MAJOR, sp. Nn.

2. Nigra; fascia angusta transversa supra antennas, margine
interiore oculorum anguste, linea pone oculos, postscutello, segmentis
dorsalibus primo secundoque macula magna obliqua utrinque,
segmentisque 3-5 fascia lata sub-basali flavis ; mandibulis, tibus
tarsisque fuscis ; alis flavo-hyalinis, venis testaceis.

Long. 24 mm.

@. Clypeus coarsely punctured, broadly rounded at the apex,
raised and longitudinally carinate in the middle. Front coarsely
punctured, the prominence between the antenne almost trans-
verse at the apex, vertex and ocellar region a little more sparsely
punctured, cheeks very sparsely and more finely punctured ;
posterior ocelli nearly twice as far from the eyes as from each
other. Thorax closely and coarsely punctured, especially on the
pronotum and mesopleurz ; propleure finely obliquely striated.
Median segment smooth, opaque, rugosely punctured in the
middle, with a few irregular transverse strize at the apex and
also on the sides of the posterior truncation ; the sides of the
segment striated, with a small yellow band near the apical angle.
Aibdomen shining, with a few scattered punctures, the apical
segment closely longitudinally striated and broadly rounded at
the apex. Ventral segments smooth, more or less punctured at
the apex. First and third abscisse of the radius about
equal in length, the second nearly half as long again. First
recurrent nervure received beyond the middle, sometimes at
two-thirds from the base of the second cubital cell, second
received between the middle and one-quarter from the base of
the third cubital cell. Calcaria whitish.

Hab. Central Brazil (Sellow).

Described from three specimens in the Berlin Museum.

The probable male of this species has the usual yellow markings :
the mandibles, clypeus, a small spot on each side above the base
of the antenna, the margins of .the pronotum, interrupted in the
middle on the anterior margin, the tegule, a spot on the meso-
notum, one on the centre of the scutellum, a large spot on the
mesopleure below the anterior wings and another before the
intermediate cox, the postscutellum, a lateral longitudinal band
on each side of the posterior slope of the median segment, apical
bands on dorsal segments 1-6, straight and narrow on segments
4-6, broader and shallowly emarginate on segments 2-3 and very
deeply emarginate on the first segment, the tibie, tarsi, and the
greater part of the femora yellow. No yellow spots on the apical
segment. Wings hyaline, nervures black. HEmargination of the
apical segment broader at the apex than deep, the dorsal surface

724 MR. R. BE. TURNER ON

of the segment flattened and distinctly margined lateraily,
shallowly longitudinally depressed in the middle and sparsely
punctured ; the basal segment distinctly less than twice as long
as the breadth at the apex. Clypeus broadly rounded or sub-
truncate at the apex, not emarginate. Second recurrent nervure
received at the base of the third cubital cell, almost interstitial
with the second transverse cubital nervure; third abscissa of the
radius a little longer than the second. Pronotum slightly
emarginate anteriorly.

Length 19 mm.

The details of neuration in the Elidine are not reliable for
specific differences, often showing slight differences on the
opposite sides of the same insect. The really important
distinctions in the males of His are to be found in the structure
of the apical and basal segments of the abdomen.

Exis compusta Sm. (Pl. LX XXII. fig. 12.)

Myzine combusta Sm. Descr. new spec. Hymen. p. 179 (1879),
2; Bingh. Faun. Brit. Ind., Hymen. i. p. 67 (1897), @.

The type is from Shuckard’s collection. I do not know why
Smith gave the locality as India or Africa, nor why Colonel
Bingham included it in the Fauna of India without any note of
doubt. The structure is that of the true American section of
Elis, and I look on it as a mere colour-variety, or possibly a local
race confined to one island, of #. ephippiwm Fabr., a West Indian
species which I have recorded from St. Thomas, St. John, Antigua,
and Porto Rico.

Subfamily ANTHOBOSCIN &.

Genus AnrHoposoa Guér.

Anthobosca Guér. Voy. ‘ Coquille,’ Zool. 11. p. 214 (1839).

Cosila Guér. Voy. ‘ Coquille,’ Zool. 11. p. 249 (1839).

Dimorphoptera Sm. Trans. Ent. Soc. London, p. 238 (1868).

Myzine Burm. Stett. ent. Zeit. xxxvil. p. 168 (1876) (nec
Latreille).

Odontothynnus Cam. Rec. Albany Mus. i. p. 161 (1904).

Austrotiphia Cockerell, Bull. Mus. Comp. Anat. Harvard, p. 49
(1906).

ae Cockerell, Bull. Mus. Comp. Anat. Harvard, p. 49
1906).

Anthobosca Turn. Proc. Linn. Soc. N.S.W. xxxii. p. 514
(1907).

There has been so much confusion in connection with this
interesting genus that I think it may be useful to give a short
account of the species, which may serve as a basis for a future
monograph. Owing to the great differences between the sexes,
they were for a long time placed in different families, most of the

STUDIES IN THE FOSSORIAL WASPS. 725

males being placed in <Anéhobosca in the Thynnide, and the
females in Cosila among the Scolide. Guérin used both
names in the same work, but although he placed Cosila
chilensis correctly, with the Scoliide, he failed to see the
relationship to Anthobosca australasice which he had described
in an earlier portion of the same work and classed with the
Thynnide. Smith in 1868 placed a single male in his
genus Dimorphoptera, which is a synonym of Cosila, but both
before and after placed other males in Anthobosca among the
Thynnide. He also described both sexes of A. albomaciulata Sm,
which were taken coupled by Bates, as I/yzine, but this work was
published after his death and without his revision. Burmeister
also in 1876 correctly associated the sexes, but placed his species
in Myzine. Although three or four males had been correctly
associated with the females, they do not appear to have been
connected in any way with the males described in the genus
Anthobosca till my revision of the Australian species of the
genus appeared in 1907. ‘The best work on the genus was done
by Saussure in 1892; but he treated the males as unknown,
except in the case of chilensis, and did not connect Cosila with
Anthobosca. In Dalla Torre’s great Catalogue, published in
1897, there is much confusion in regard to this genus, species
appearing under the genera Zhyninus, Myzine, and Cosila. There
is much confusion over the genus in the recent papers of Cameron
on South African Hymenoptera. Ashmead treats the group as a
family, Cosilide, not as a subfamily of the Scoliide, which I
consider the more natural course. But he places Anthobosca in
the Thynnide and Dimorphoptera in his family Myzinide. He
includes in his family Cosilide several genera of doubtful
affinities, of which, in my opinion, Vursew should be treated as
an aberrant genus of the Sphecoidea, whilst Maurillus belongs to
the Pompilide. The position of Sierolomorpha and Dicrogeniwm
seems to me very doubtful, but I have not seen specimens. I am
compelled to look on Axthobosca as the only genus which can
be placed in the subiamily Anthoboscine with any certainty,
Ashmead states that the intermediate coxze in his Cosilide are
contiguous or nearly so; but this is quite incorrect as to the
females, and even in the males the separation is quite distinct.
In my key to the species 1 have included several species which I
have not seen, one or two of which may possibly not belong to
the genus.

The females are distinguished from other Scoliide by the
absence of a deep groove between the two basal ventral segments
of the abdomen. The hind coxe are separated as in Tiphiu,
not contiguous as in “lis; but the intermediate coxe are less
widely separated than in either of those groups, though very little
less so than in Hlis. The males are distinguished from all other
Scolide by the unarmed rounded hypopygium—in this character
approaching most nearly to the Thynnide of the genus Hirone.

726 MR. R. E. TURNER ON

It is quite possible that the genus will have to be subdivided
owing to differences of structure, especially in the tarsal ungues,
which in most of the Australian species have a blunt lobe at the
base, but in the African, South American, and a few Australian
species are bifid. The neuration 1s so variable, even in the same
species, that I do not consider that genera should be founded on
the shape of the radial cell alone, on which character the sub-
genera Colobosila Sichel and Callosila Saussure have been based.
In two Australian species the cell is acute at the apex, in others
blunt, and African species may similarly be divided into two
eroups. But until more species are known in both sexes any
subdivision would be rash and unnecessary.

The geographical distribution of the genus is interesting, being
almost entirely Southern, including Australia (where the species
are most numerous), South America as far north as the Amazon,
Madagascar and South Africa spreading up the East African
coast to Suakin and crossing to Aden. lam aware that many
regard similar cases of distribution as a proof of southern origin,
and explain them by former northern extensions of the Southern
continent connecting at different times with southern extensions
of the land in the Southern Hemisphere. But taking into
consideration the enormous depth of the Southern oceans I
cannot look on this explanation as satisfactory, and think that it
is more reasonable to look on this and other similar cases of
distribution as instances of the survival in the south of genera
which in former times had a much more extensive range. It
has been pointed out by Darwin that the struggle for existence
is more severe on large land-areas than on smaller ones, owing to
the more complex conditions of life from the larger number of
existing species which are able to come into competition. Now
the land-areas in the south are very much smaller than in the
north, so that it is reasouable to suppose that many genera may
have been able to survive in the south with little or no modi-
fication, which have been exterminated by the more severe
struggle for existence in the north. In this case we should
expect to find fossil remains of such genera, or at all events of
nearly related forms, in the north; and in a great number
of cases such fossils have been found. In the present genus I
look on Cockerell’s Geotiphia found fossil in Colorado as absolutely
congeneric with existing South-American species ; and the plant
genus Araucaria and many others, which are now southern, had
in early geological times a wide range in the Northern Hemi-
sphere. It may, of course, be argued that such genera originally
had their home in the south and at one time extended their
range northward, but in that case it may be conceded that
they may have reached the different portions of the Southern
Hemisphere by way of the north and not from a southern
eontinent. I must own to an objection to calling up continents
from the extreme ocean depths ‘‘ nisi dignus vindice nodus.”

5.

10.

16.

: Second npcersen of the radius no longer

. Pronotum and scutellum strongly punctured

. Front punctured; legs wholly black ;

. Abdomen black ;

STUDIES IN THE FOSSORIAL WASPS. fi

bo
~I

Key to the Species of Anthobosca.

Females.

> SSSOES shropen ne Olal WVOP Cl co napocaseu cuecoanosceacnoace sae

South-American species

. Australian species ........

African and Arabian species

. Tarsal ungues with a lobe at the base ‘either younded

or pointed... is eee
Tarsal ungues w ithout a basal lobe, deeply. bifid

. Radial cell pointed at the apex

Radial cell blunt at the apex .....

Pubescence black, calcaria black, “abdomen “wholly
lac kamen sere

Pubescence whitish, ‘calcaria white, abdomen. usually
with a yellow spot on each side of the third dorsal
segment ......

Wings short, the length ‘of the costa ‘of the fore v wing

not exceeding two. and a half times the breadth of
the mesonotum. Pronotum, median segment, and
three basal abdominal segments rufo-testaceous ...
Wings longer, the length of the costa of the fore
wing three times as great as the breadth of the
mesonotum. ‘Thorax and abdomen black

. Antenne orange; stigma about three times as long

on the costa as broad AGe ps coAS aE HSE MOR BAERS
Antenne black or fuscous; stigma scarcely more
than twice as long on the costa as broad ............

. Legs bright ferruginous

Legs black, sometimes fusco- ferruginous beneath —

than the
third ; first recurrent nervure “ieuannall just before
the middle of the second cubital cell

Second abscissa of the radius longer than the third;
first recurrent nervure received at one-quarter from
the base of the second cubital cell

Wings fuscous; four basal abdominal segments with
Onllied Oie WOM SEMAN WNT ocean nes ononcn oso accesses

Wings hyaline or subhyaline; abdominal segments
without cilixe of scale-like hans See ae

Pronotum and scutellum Seanad and very oe
punctured

. Third cubital cell nearly a as long on 1 the radius : as on

the cubitus or longer; first recurrent nervure
received slightly beyond the middle of the second
cubital cell .

Third cubital cell much shorter on the radius than
on the cubitus: first recurrent nervure received
well before the middle of the second cubital cell ...

leneth 11 mm.

Front smooth and shining; legs fusco-ferrugimous
beneath ; length 7 mm..

second and third cubital cells each
recelying a recurrent nervure

Abdominal segments 2-4 very broadly rufo-testaceous
at the base; second cubital cell receiving both
TELOWIB EOIN, TVET ABU ROS ooo ong oacene dan opansoasosovenouobseowenne

. Sixth dorsal segment strongly striated at the base;

radial cell detached from costa for half its length .

Sixth dorsal segment punctured: radial cell detached
from costa at extreme apex only .

Radial cell subacute at apex; abdomen not marked
with yellowish white..

Radial cell broadly rounded or 1 narrowly truncate at
apex; abdomen marked with yellowish white ......

A, australis Sichel.

A.

signata Sm.

A, fastuosa Sin.

Me

A, flavicornis Sauss.

A.

&.
9.
10.

. cognata Sm.

strandi Turn,
11.
12.

. albopilosa Turn.

. argenteocincta Grib,

. anthracina Sm,

13.

. wnicolor Sm.

. levifrons Sm.

15.

. clypeata Sm,
. nubilipennis Turn.

. occipitalis Turn,

17,

728

MR. R, E. TURNER ON

17. Apical segment at least of abdomen red; wings
fusco-v iolaceous .. 18.
Abdomen wholly black ; ; wings 's flavo- or fusco- hy aline. 19.
18. Apical abdominal segment only red; thorax not
coarsely punctured . *A, melanaria Cam.

Two apical abdominal sements ‘red; “thorax very
coarsely sculptured

Ele

donaldsoni Fox.

19. Wings flavo-hyaline; third transverse cubital nervure
received close to the apex of the radial cell ......... A. insularis Sm.
Wings fusco-hyaline; fourth abscissa of the radius
as long as the first.. alate A. natalica Turn.
20. Radial cell narrow ly tr uncate at apex. “Length 5mm. 4. minima Turn.
Radial cell broadly rounded at apex. Length 9mm.
(Yen esTOV RS Meera ee Raper ddpocuuoods ccc Cop‘oosnbadon ebooden050d 21.
21. Head and thorax black with pale yellow markings ... 22.
Head and thorax without yellow markings, thorax
black or more or less ved. ...,......00ss0ceeseeeeeeeeeeeeee A. erythronota Cam.
22. Abdomen ferruginOUS............eecceeeereeeeeeeesereeeesseeee A. Sauakinensis Magy.
Abdomen black) s.cccoscneeseotacemencuasslasieccccre seein A. arabica Turn.
93. Abdomen black without markings ............00.ceseee eee 24.
Abdomen marked with yellow or white.................. 25.
24, Antenne orange; head and thorax closely punctured. A. chilensis Guér.

Antenne black; head and thorax very sparsely
punctured

A.

carbonaria Burm.

25. Wings flavo-hyaline; abdomen with four or five
interrupted yellow bands .......- 26.
Wings fusco-hyaline ; abdomen with one or two
yellowish lateral spots age eed pes cenceeictenemns ene 27.
26. Legs ferruginous A. antennata Sm.

Lee $ inflewle

. Vertex smooth and ‘shining ; ‘third dorsal. segment

only with lateral spots ..
Vertex sparsely punctured ; ‘second and third dorsal
segments with whitish lateral spots ..................

1. ANTHOBOSCA AUSTRALIS Sichel.

Pl, LX XXIII, figs. 1, 3, 6, 14.)
Cosila australis Sichel, Sauss. et Sichel, Cat. spec. gen. Scolia,

p-

261 (1864), 2.

. erythropyga Burm.

A. bipunctata Perty.

A. albomaculata Sm.

(Pl. LXXXII. figs. 1, 2

Dimorphoptera scoliiformis Sm. Trans. Ent. Soc. London, p. 238
(1868), °.
Cosila (Callosta) australis Sauss. in Grandidier, Hist. Mada-
gasear, xx. p. 232 (1892), 2.
Dimorphoptera nigripennis Sm. Trans. Ent. Soc. London, p. 239
(1868), 3.

2

The radial cell is pointed and detached from the costa at

the apex, receiving the second transverse cubital nervure close to
the middle, the second abscissa of the radius almost as long as the
third. First recurrent nervure received close to the middle of
the second cubital cell, second at one-third from the base of the
third cubital cell. Third cubital cell fully half as long again on
the cubital as on the radial nervure. Basal lobe of tarsal ungues
bluntly rounded. Basal joint of hind tarsi with a scopa beneath,
in which is a row of longer black spines. Sparsely punctured,
shining; pronotum and scutellum more closely punctured ;
pygidium densely clothed with long black pubescence.

Black, with black pubescence. Wings fusco-violaceous.

Length 23 mm.

STUDIES IN THE FOSSORIAL WASPS. 729

3. Second abscissa of the radius about half as long as the
third. Basal lobe of tarsal ungues bluntly rounded. Apical
joint of the flagellum about half as long again as the penultimate
and slightly curved ; the eighth, ninth, tenth, and eleventh joints
subtuberculate in the middle beneath. Posterior tibie very
feebly spined on the outer margin. Pronotum short, scarcely
narrowed anteriorly. First abdominal segment very little longer
than the second, broader at the apex than long. Head and
thorax closely and not very minutely punctured.

Black, with cinereous pubescence, calcaria black. Wings
fusco-violaceous.

Length 17 mm.

Hab. EK. Coast of Australia between Sydney and Brisbane.

The association of the sexes is not certain. The structure of
the male antenne is very remarkable.

2. ANTHOBOSCA SIGNATA Sim.

Myzine signata Sm. Cat. Hym. B. M. iii. p. 75 (1855), 9.

Cosila biguttata Sichel, Sauss. et Sichel Cat. spec. gen. Scolia,
p- 262 (1864), 9.

Dimorphoptera signata Sm. Trans. Ent. Soc. London, p. 238
(1868), ©.

Cosila (Callosila) signata Sauss. in Grandidier, Hist. Madagascar,
Oe Oh a vad (ksh) y 2.

Var. Dimorphoptera morosa Sm. Trans. Ent. Soc. London, p. 239
(S68) o=

9. Radial cell pomted and detached from the costa at the
apex, second recurrent nervure received at about one-quarter
from the base of the second cubital cell. Other details of
neuration as in australis Sich. Basal joint of hind tarsi with a
comb of thickly set spines beneath. Basal lobe of tarsal ungues
bluntly rounded. Shining and sparsely punctured, the front and
pronotum more closely punctured; sixth -dorsal segment of
abdomen rugosely punctured. |

Black, with whitish pubescence, the calcaria white. Wings
fusco-violaceous. Third dorsal segment with an orange spot on
each side.

Length 22 mm.

Hab. %. Australia, from Sydney to Cairns.

I look on morosa as a small variety in which the orange spots
on the abdomen are wanting. The second abscissa of the radius
is distinctly shorter than the third in the type of morosa, owing to
the sharp curve toward the base in the second transverse cubital
nervure, but I do not think that this distinction is of specific
importance.

3. ANTHOBOSCA FASTUOSA Sm. (Pl. LXXXT. fig. 3.)

Dimorphoptera fastuosa Sm. Trans. Ent. Soc. London, p. 240

(1868), 2.
Myzine fastuosa D, T. Cat. Hym. viii. p. 123 (1897).

Proc. Zoot. Soc.—1912, No. XLVIII. 48

730 MR. R. E. TURNER ON

2. Wings short, the length of the costa of the fore wing not
exceeding two and a half times the breadth of the mesonotum.
Radial cell broad, the apex detached from the costa and narrowly
rounded. Third abscissa of the radius nearly half as long again
as the second, and more than twice as long as the first; second
transverse cubital nervure received at about two-fifths from the
base of the radial cell. Second cubital cell not very strongly
produced towards the base on the cubitus, receiving the first
recurrent nervure near the middle, second recurrent nervure
received just beyond one-quarter from the base of the third
cubital cell. Basal joint of hind tarsi finely pectinate beneath ;
basal joint of fore tarsi with six spines above and about eight
shorter ones beneath; basal lobe of tarsal ungues bluntly rounded.
Sparsely punctured, more closely on the front, pronotum, and
abdomen ; the apical abdominal segments more strongly punctured
than the three basal segments.

Rufo-testaceous; head, mesothorax, and three apical abdominal
segments black; mandibles at the base and scape fusco-ferruginous;
pubescence rufo-testaceous. Wings hyaline, tinged with yellow ;
nervures testaceous, the stigma black.

Length 17 mm.

Hab. Champion Bay, W. Australia.

The process beneath the hind femora near the apex is much less
developed in this species than in most Australian species, though
the femora are by no means convex beneath as in most of the
South-American species and in A. nudilipennis.

4, ANTHOBOSCA FLAVICORNIS Sauss.

Cosila (Callosila) flavicornis Sauss. in Grandidier, Hist. Mada-
gascar, xx. p. 233 (1892), ©.

Q@. Radial cell with a slight angle at the apex ; second abscissa
of the radius shorter than the first, the third longer than the
first and second combined, third cubital cell of about equal
length on the radial and cubital nervures. Second cubital cell
much produced towards the base on the cubitus; first recurrent
nervure received about the middle of the second cubital cell,
second just before the middle of the third cubital cell. Basal
joint of hind tarsi with a scopa beneath, in which are a few
spines longer than the hairs; basal joint of fore tarsi with six
stout spines above and a row of very fine spines beneath; basal
lobe of tarsal ungues rounded. The lobe beneath the apical
portion of the hind femora has the lower margin nearly straight,
only very slightly rounded, and is strongly developed. Shining
and rather sparsely punctured, the anterior half of the pronotum
and the median segment closely and finely punctured and sub-
opaque.

Black, with white pubescence; the flagellum orange; calcaria
white, the spines of the tibize and tarsi reddish brown. Wings
fusco-hyaline, nervures fuscous.

STUDIES IN THE FOSSORIAL WASPS. 731

Length 11-14 mm.

Hab. Victoria.

There is also a specimen in the National Collection from
Tasmania and another from Cairns, Queensland. In the former
the radial cell is rounded at the apex and the tibiz and tarsi are
reddish; in the latter the radial cell is almost truncate at the
apex, though narrowly, and the second abscissa of the radius is
as long as the first. I do not think that small differences of
neuration in this genus will prove to be of specific value.

5. ANTHOBOSCA COGNATA Sm.

Dimorphoptera cognata Sm. Descr. new spec. Hymen. p. 188
(CSS) 5 ae

@. Radial cell rather broadly rounded at the apex; the four
abscissee of the radius of about equal length; the third cubital
cell much longer on the cubitus than on the radius. Recurrent
nervures received close to the middle of the second and third
cubital cells; the second cubital cell strongly produced toward
the base on the cubitus. Basal joint of hind tarsi with a scopa
beneath in which are five or six longer spines; basal joint of fore
tarsi with six long and rather slender spines above and a closely
set row of very small spines beneath ;_ basal lobe of tarsal ungues
less bluntly rounded than in flavicornis. The lobe beneath the
apical half of the hind femora is very broadly rounded. Closely
and finely punctured, more finely and closely on the abdomen
and median segment than on the head and thorax, some larger
punctures on the disc of the mesonotum.

Black, with sparse whitish pubescence; mandibles at the base
and legs, except the cox, ferruginous.

Length 11 mm.

Hab. Swan River, Western Australia.

6. ANTHOBOSCA STRANDI Turn.

Anthobosca strandi Turn. Proc. Zool. Soc. London, p. 306
(1910), @.

@. Radial cell broadly rounded at the apex; second abscissa
of the radius twice as long as the first and distinctly longer than
the third; first recurrent nervure received at one-quarter from
the base of the second cubital cell, second at one-fifth from the
base of the third cubital cell. Apical lobe beneath the hind
femora broadly rounded and strongly developed. Pronotum,
median segment, and abdomen very finely and closely punctured,
front coarsely and closely, mesonotum and scutellum less coarsely
and rather sparsely punctured.

Black; the pubescence pale fulvous on the head and pronotum,
grey on the sides of the abdomen, fusco-ferruginous on the apical
dorsal segment; apex of the scape and flagellum beneath fusco-
ferruginous, a dull ferruginous spot on each side close to the
summit of the eyes; mandibles at the base, tegule, tibie, tarsi,

48*

732 MR. R. E. TURNER ON

femora at the apex, and the whole of the hind femora ferruginous.
Wings subhyaline, nervures fuscous, stigma ferruginous.

Length 13 mm.

Hab. &. Australia; probably from Victoria.

I have not a specimen of this species before me at the time of
writing.

7. ANTHOBOSCA ALBOPILOSA Turn.

Anthobosca albopilosa Turn, Proc. Linn. Soc. N.S.W. xxxii.
p. 520 (1907), @.

Q. Radial cell slightly angular at the apex; third abscissa of
the radius the longest, but shorter than the first and second
combined, the first scarcely as long as the second. First re-
current nervure received near the middle of the second cubital
cell, second just beyond one-quarter from the base of the third
cubital cell. Second cubital cell less strongly produced towards
the base than in flavicornis. Basal joint of hind tarsi with a
scopa beneath, in which are a few longer spines; basal joint of
fore tarsi with six or seven well-developed spines below and the
usual six above; basal lobe of tarsal ungues rounded; lobe
beneath the apical half of the hind femora broadly rounded and
less strongly developed than in flavicornis. Deeply and rather
closely punctured, the abdomen more shallowly punctured.

Black; the pubescence on the head and sides of the thorax
white, on the abdomen and dorsal surface of the thorax black ;
calearia black; four basal dorsal segments of abdomen and
ventral segments 2—4 with apical ciliz of scale-like white hairs.
Wings fuscous, with faint violet reflections.

Length 14 mm.

Hab. Queensland.

8. ANTHOBOSCA ARGENTEOCINCTA Grib.

Oosila argenteocincta Gribodo, Ann. Mus. Civ. Stor. Nat. Genova,
Xvill. p. 261 (1883), @.

@. Radial cell with a slight angle at the apex; second abscissa
of the radius longer than the first, the third equal to the first
and second combined, the third cubital cell a little longer on the
cubitus than on the radius. First recurrent nervure received
just beyond the middle of the second cubital cell, second at one-
third from the base of the third cubital cell; the second cubital
cell moderately produced towards the base on the cubitus. Basal
joint of hind tarsi with a close row of small spines beneath ; basal
joint of fore tarsi with seven spines above, and a close-set row of
slender spines beneath; basal lobe of tarsal ungues bluntly
rounded; lobe beneath the basal half of the posterior femora
broadly rounded. Shining, finely and very sparsely punctured ;
the median segment subopaque and very finely and closely
punctured on the sides; abdomen shallowly but much more
coarsely punctured.

STUDIES IN THE FOSSORIAL WASPS. 733

Black, with sparse grey pubescence; flagellum beneath fuscous ;
tarsal ungues ferruginous ; calcaria white. Wings light fuscous,
shaded with bronze; nervures black.

Length 13 mm.

Hab. Adelaide, 8. Australia.

I think I have identified this species correctly. Gribodo
mentions a tubercle at the base of the first ventral segment,
which does not seem to be any more developed in the specimen
described above than in other species.

9, ANTHOBOSCA ANTHRACINA Sm. (PI. LX XXII. fig. 3.)

Myzine anthracina Sm, Cat. Hym. B. M. iii. p. 71 (1855), 9.
Myzine sabulosa Sm. Cat. Hym. B. M. iii. p. 76 (1855), 2.
Dimorphoptera anthracina Sm. Trans. Ent. Soc. London, p. 238
1868).
sR subulosa Sm. Trans. Ent. Soc. London, p. 238
1868).
Ausrotitphia kirbyt Cockerell, Bull. Mus. Comp. Anat. Harvard,
p. 49 (1906), 2.

2. Radial cell very bluntly rounded at the apex; second
abscissa of the radius shorter than the first, the third abscissa
half as long again as the first and second combined. First
recurrent nervure received just beyond the middle of the second
cubital cell, which is more sharply produced towards the base
than in wnicolor Sm. or levifrons Sm.; second recurrent nervure
received at two-fifths from the base of the third cubital cell,
which is longer on the radius than on the cubitus. Basal joint
of hind tarsi finely pectinate beneath ; basal joint of fore tarsi
with six spines above and eight finer beneath ; basal lobe of tarsal
ungues bluntly rounded. Finely and rather sparsely punctured,
pronotum more coarsely punctured, median segment very finely
and closely punctured with a smooth line in the middle.

Black with whitish pubescence; spines of -the tibie and tarsi
white, those of the anterior pair slightly reddish. Wings
hyaline, shaded with fuscous towards the apex, nervures pale
testaceous.

Length 14 mm.

Hab, 5.K. Australia ; Adelaide to Sydney.

The description is taken from the type of anthracina. The
type of sabulosa has the first and second abscisse of the radius
about equal in length, combined equal to the third; the third
cubital cell no longer on the radius than on the cubitus; the
nervures are fuscous. But excepting in these points I cannot
detect any difference, and do not consider that small differences
of neuration can be held to be of specific importance, considering
how much variation occurs in other groups of Scoliide in such -
detail. The type of Austretiphia kirbyi is similar to sabulosa.

734 MR. R. E. TURNER ON

10. ANTHOBOSCA UNICOLOR Sm.

Myzine unicolor Sm. Cat. Hym. B. M. iii. p. 75 (1855), 2.

Dimorphoptera unicolor Sm. Trans. Ent. Soc. London, p. 238
(1868), °.

Cosila (Callosila) minuta Sauss. in Grandidier, Hist. Madagascar,
ke Ps 290)( 1892), Oo".

@. Radial cell rounded at the apex; second abscissa of the
radius as long as the first, the two combined a little longer than
the third; first recurrent nervure received a little before the
middle of the second cubital cell, second before one-third from
the base of the third cubital cell, which is much longer on the
cubitus than on the radius; second cubital cell not very strongly
produced towards the base. Basal joint of hind tarsi with a few
small spines beneath ; basal lobe of tarsal ungues rounded; lobe
on the apical third of the hind femora beneath not very strongly
prominent. Shining and sparsely punctured, the front closely
and strongly punctured; median segment subopaque, very finely
and closely punctured.

Black, with white pubescence; mandibles and scape beneath
fuscous; calcaria white. Wings hyaline, faintly tinted with
fuscous.

Length 10 mm.

Hab. Eastern Australia; Tasmania to Cairns.

11. ANTHOBOSCA LEVIFRONS Sm,

Dimorphoptera levifrons Sm. Descr. new spec. Hymen. p. 188,
(er), 2.

Myzine levifrons D. T. Cat. Hymen. viii. p. 124 (1897).

2. Radial cell broadly rounded at the apex; second abscissa of
the radius shorter than the first, the two combined longer than the
third ; first recurrent nervure received at two-fifths from the base
of the second cubital cell, second just before the middle of the third.
cubital cell, which is much longer on the cubitus than on the radius.
Basal joint of hind tarsi with a row of fine hairs beneath, in which
are two or three longer spines; the lobe beneath the apical third
of the hind femora broadly rounded. Basal lobe of tarsal ungues
small and blunt. Shining, head and thorax very sparsely punc-
tured, the front smooth; median segment and abdomen closely

-and minutely punctured.

Black, with sparse white pubescence; mandibles, flagellum
beneath, and legs beneath fusco-ferruginous. Wings hyaline,
tinged with fuscous ; nervures fusco-ferruginous.

Length 7 mm.

fab. Adelaide, 8. Australia.

12. ANTHOBOSCA OCCIPITALIS, sp. n.

2. Nigra, vertice macula utrinque supra oculos fusco-sanguines ;
unguiculis bifidis ; alis subhyalinis, venis fusco-ferrugineis.
Long. 14 mm

TUDIES IN THE FOSSORIAL WASPS. (5x

2. Radial cell rounded at theapex; third abscissa of the radius
shorter than the second ; third cubital cell as long on the radius as
on the costa; first recurrent nervure received just beyond one-
third from the base of the second cubital cell, second at one-fifth
from the base of the third cubital cell; second cubital cell not
very strongly produced towards the base. Tarsal ungues bifid, |
without a basal lobe; basal joint of hind tarsi with rather long
hairs beneath ; lobe beneath the apical half of the hind femora
broadly rounded. Closely punctured; rather strongly on the
head, finely on the thorax, coarsely on the scutellum, rather
sparsely and shallowly on the abdomen; sixth dorsal segment
rather broadly truncate at the apex.

Black, with grey pubescence, a fusco-sanguineous spot on the
vertex on each side close to the summit of the eyes; sixth dorsal
segment fuscous at the apex, calcaria whitish.

Hab. S. Australia; probably from Adelaide.

This species is remarkable for the very long second abscissa of
the radius, and may be easily distinguished from the majority of
Australian species by the bifid tarsal ungues.

13. ANTHOBOSCA NUBILIPENNIS Turn.
Anthobosea wubilipennis Turn. Proc. Zool. Soc. London, p. 307

(SLO) Soe

Q@. Radial cell detached from the costa for fully half its length,
slightly angular at the apex; second abscissa of the radius longer
than the first and about equal in length to the third; first
recurrent nervure received beyond two-thirds from the base
of the second cubital cell, second before one-quarter from the
base of the third cubital cell, which is nearly twice as long

on the cubitus as on the radius. Tarsal ungues bifid, without a
basal lobe ; hind femora convex beneath, the lobe not apical but
almost medial; basal joint of hind tarsi with three or four short
spines beneath. Deeply but not very closely punctured ; meso-
notum and scutellum very sparsely punctured; sixth dorsal
abdominal segment coarsely striated at the base, rounded at the
apex.

“ines with white pubescence; calcaria black. Wings fusco-
hyaline, nervures fuscous; the stigma small. Median segment
abruptly truncate.

Length 16 mm.

Hab. Perth district, S.W. Australia.

A very distinct species in the characters of the hind legs,
neuration, and median segment.

14. ANTHOBOSCA CLYPEATA Sm. (P]. LUX X XI. fig.2; Pl. LX XXII.
Hi pedals AGO.GUUL, tee 7.)

Dimorphoptera clypeata Sm. Trans. Ent. Soc. London, p. 239
(1868), @.

@. Radial cell rounded at the apex, detached from the costa

I 2

13 MR. R. E. TURNER ON

at the extreme apex only; second abscissa of the radius twice as
long as the first and about equal in length to the third; second
cubital ceil receiving both recurrent nervures, the first at one-
third from the base, the second just before the apex; third
cubital cell as long on the radins as on the cubitus. Tarsal
ungues bifid; lobe beneath the apical half of the hind femora
strongly rounded ; basal jot of hind tarsi with a row of short
hairs beneath, in which are a few longer spines, Closely punc-
tured; the thorax very closely and finely punctured, with larger
and sparser punctures intermixed ; abdomen shining and sparsely
punctured.

Black, with whitish pubescence; sixth dorsal segment densely
clothed with long golden hairs; clypeus exeept at the apex and
the margins of the eyes broadly interrupted on the summit,
yellow ; basal two-thirds of dorsal segments 2-4 and of ventral
segments 2-3 emarginate in the middle posteriorly, rufo-
testaceous. Wings subhyaline, nervures fusco-ferruginous.

Length 22 mm.

Hab. Swan River, W. Australia.

An obscure scar runs from the base of the stigma to close to
the base of the first transverse cubital nervure. I have little
doubt that the male of this species will prove to be A. crassicornis
Sm., which corresponds with it in the position of the recurrent
nervures, the bifid tarsal ungues, and also to some extent in the
colour of the clypeus and abdomen.

*15, ANTHOBOSCA FASCICULATA Sichel.

Cosila (Colobosila) fasciculata Sichel, Sauss. et Sichel, Cat. spec.
gen. Scolia, p. 263 (1864), 2.

Hab. Australia.

I fail to recognise this species with any certainty, but do not
consider that the subgenus Colobosila can stand, the truncation
of the apex of the radial cell being insufficient as a subgeneric
character unless supported by others.

*16, ANTHOBOSCA INORNATA Sauss.
Cosila inornata Saussure in Grandidier, Hist. Madagascar, xx.

p, 233 (1892), 2.

J do not recognise this species as a synonym of any known to
me, though it seems to be related to A. anthracina, differing in
the position of the first recurrent nervure. Considering how
unreliable the details of neuration are in this genus, it is quite
possible that it is only a variety of anthracina.

*|7, ANTHOBOSCA MELANARIA Cam.

Plesia melanaria Cam. Rec. Albany Mus. i. 5, p. 297 (1905), 2.

@. From the description I have no doubt that this is an
Anthobosca, the characters ‘‘ apex of radial cell sharply pointed,”
and ‘‘pygidium piceous red, its base fringed with long, bright
rufous hair,” and “ metapleurze smooth,” agreeing much better

STUDIES IN THE FOSSORIAL WASPS. 137

with Anthobosca than Plesia. . Although the two genera belong
to different subfamilies of the Scoliide, the females are likely to
be confused by a beginner. Cameron places male Scoliidz of the
subfamily Anthoboscine in the Thynnide, and females of the same
group with the Scoliide of the subfamily Elidine.

*18. ANTHOBOSCA DONALDSONI Fox.

Cosila donaldsoni Fox, Proc. Acad. Philadelphia, p. 549
(LSYO), Qs

@. Third transverse cubital nervure received close to the apex
of the radial cell. Clypeus tridentate on the anterior margin.
Head and thorax very coarsely punctured, pronotum and scu-
tellum scabrous; median segment finely striato-punctate, the
sides obliquely striated ; abdomen strongly but sparsely punc-
tured, sixth dorsal segment striato-punctate; tarsal ungues
cleft.

Black, with greyish pubescence ; two apical abdominal segments
red; calearia whitish. Wings black, strongly violaceous.

Length 18 mm.

Hab. Somaliland.

I have not seen this species, but the description corresponds
almost exactly with Hlis aliciw described in this paper, though
the clypeus in that species is not distinctly tridentate. But I do
not believe that Fox would have confused the genera.

19. AnrHoBoscA INSULARIS Sm. (Pl. LXXXII. fig. 6;
Pl. LX XXIII. fig. 8.)

Myzine insularis Sm. Descr. new spec. Hymen. p. 178
(879) 2.

Cosila insularis Saussure in Grandidier, Hist. Madagascar, xx.
pale (S92) OR

@. Radial cell subacute at the apex, receiving the third trans-
verse cubital nervure very near the apex, in some specimens
almost at the apex; second abscissa of the radius longer than the
first, the two combined not quite as long as the third; first
recurrent nervure received at two-fifths from the base of the
second cubital cell, second at one-quarter from the base of
the third eubital cell, which is nearly or quite as long on the
radius as on the cubitus. Tarsal ungues bifid; hind femora
very broadly rounded beneath on the apical two-thirds, basal
joint of hind tarsi with three long spines beneath. Shining and
sparsely punctured, closely on the front and closely and very
finely on the median segment.

Black, with black pubescence; calcaria brown. Wings flavo-
hyaline, nervures ferruginous.

Length 23-29 mm.

Hab. Madagascar.

The type has a rufous spot in the middle of the first dorsal
segment, and obscure reddish shading on the head and thorax.

738 MR. R. E. TURNER ON

20. AnrHOBOSCA NATALICA Turn.
Anthobosca natalica Turn. Trans. Ent. Soc. London, p. 85

(1908), @.

@. Radial cell narrowly rounded at the apex, subacute; third
abscissa of the radius twice as long as the second, the fourth
shorter than the first ; first recurrent nervure received just beyond
the middle of the second cubital cell, second before the middle of the
third cubital cell, which is as long on the radius as on the cubitus.
Tarsal ungues bifid; basal joint of hind tarsi with a scopa beneath.
Shining and sparsely punctured, more closely on the front and
pronotum; median segment very finely and closely punctured ;
abdomen finely but more sparsely punctured.

Black ; pubescence on the apical dorsal segment brown and
long ; tegule and extreme apical margin of the abdominal segments
testaceous brown; calearia whitish ; fore tarsi fusco-ferruginous.
Wings fusco-hyaline, nervures fusco-ferruginous.

Length 12-17 mm.

Hab. Malvern, Natal.

The neuration given is as in the type, but in another specimen
the second abscissa of the radius is much more than half as long
as the third, and both recurrent nervures are received nearer the
base of the cells.

21. ANTHOBOSCA ERYTHRONOTA Cam. (PI. LXXXI. figs. 5, 6;
Pl. LXXXII. fig. 4.)

Plesia erythronota Cam. Rec. Albany Mus. i. 5, p. 320 (1905).

2. Nigra; prothorace, mesothorace, tibiis tarsisque rufo-ferru-
giners ; segmentis abdominalibus 2-4 macula laterali alba utrinque ;
alis fusco-hyalinis, venis nigris.

3. Niger, gracilis; mandibulis basi, clypeo, oculis margine
interiore, pronoto late postice, tegulis, mesonoto macula, mesopleuris
macula, scutello macula mediana, postscutello macula mediana
transversa, segmentisque abdominalibus 2-4 macula magna trans-
versa utrinque albido-flavis ; pedibus nigris albo-variegatis ; alis
hyalinis, venis nigris.

Variat 2 prothorace mesothoraceque nigris.

Long., 2 10-11 mm., 5 8-11 mm.

@. Clypeus shining in the middle and somewhat flattened,
narrowly truncate at the apex. Head convex, broader than long,
not much broader than the thorax; rather sparsely punctured,
the space round the base of the antenne closely punctured and
clothed with fulvous hairs. Eyes touching the base of the
mandibles, the line of the interior margin slightly undulating ;
posterior ocelli much nearer to each other than to the eyes.
Thorax rather sparsely punctured ; pronotum slightly emarginate
anteriorly, the posterior margin widely but not strongly arched :
the projection of the mesosternum between the intermediate cox
very deeply bilobed. Median segment very finely and closely

STUDIES IN THE FOSSORIAL WASPS, 739

punctured, the posterior slope very steep. Abdomen rather
closely punctured; the apical margins of the segments rather
broadly depressed and smooth, more broadly in the middle than
at the sides, the apical segment testaceous and thickly covered
with fulvous bristles. Radial cell bluntly rounded at the apex,
not detached from the costa; second abscissa of the radius a
little longer than the first, but scarcely more than half as long as
the third, the third cubital cell longer on the radius than on the
cubitus, second recurrent nervure received at about one-quarter
from the base of the third cubital cell. Ungues cleft.

3. Clypeus slightly convex, rather ‘narrowly truncate at the
apex and armed with a row of three very minute teeth. Antenne
no longer than the thorax and median segment combined, tapering
slightly towards the apex. Eyes convergent towards the clypeus,
the inner margin almost straight. Head and thorax finely and
closely punctured ; pronotum strongly narrowed anteriorly, the
anterior margin slightly emarginate, posterior margin strongly
arched. Abdomen shagreened, very slender, slightly tapering
to the extremities, the basal segment distinctly longer than
the second. Hind tibiz finely serrate, all the tarsal ungues
cleft. Second abscissa of the radius half as long again as the first
and only a little more than half as long as the third, second
recurrent nervure received at about one-quarter from the base of
the third cubital cell.

Hab. Willowmore, Cape Colony; November to January
(Dr. Brauns).

The male comes very near Cameron’s genus Odontothynnus,
which I have elsewhere treated with some doubt as a synonym of
Anthobosca; but Cameron states that the posterior tarsal ungues
in his genus are simple. Even if he is correct as to this character
it would not be sufficient to justify the formation of a genus on
one sex only, and his remarks show that he has no knowledge of
the genus Anthobosca. I suspect that the present species may
prove to be identical with Plesia leucospila Cam., with the
description of which it agrees fairly well, but the mesopleurze
are rather strongly though not very closely punctured, not almost
smooth as in Cameron’s description. In the broadly rounded
apex of the radial cell this species differs from A. natalica
Turn., in which the cell is subacute, but agrees with 4. arabica
Turn.

*99, ANTHOBOSCA LEUCOSPILA Cam.

Plesia leucospila Cam. Rec. Albany Mus. i. 5, p. 319
(UOD), Oe

@. It is almost certain that Cameron has misplaced this
species. It is possibly identical with A. erythronota Cam., which
has a similar colour variety, in which case the name lewcospila
should be used for the species. But Cameron states that the
pleure are almost smooth, whereas in erythronota the mesopleure

740 MR. R. E. TURNER ON

are strongly though not closely punctured. But this is possibly
an error in Cameron’s description.

*23. ANTHOBOSCA SAUAKINENSIS Magr.

Myzine sawakinensis Magr. Ann. Mus. Civ. Genova, xxi. p. 560
(1884), @.

I think it probable that 4. arabica Turn. is only a colour variety
of this species, with the abdomen black instead of ferruginous.
The difference in size between the two forms is considerable, but
both size and colour vary much in this genus. But as I have not
seen sauakinensis I think it better to keep the two separate at
present.

94, ANTHOBOSCA ARABICA Turn.

Anthobosca arabica Turn. Trans. Ent. Soc. London, p. 397
(CUO) Se

@. Radial cell broadly rounded at the apex ; second abscissa
of the radius longer than the first, the two combined equal in
length to the third, first recurrent nervure received a little before
the middlJe of the second cubital cell, second before one-third from
the base of the third cubital cell, which is as long on the radius
as on the cubitus. Tarsal ungues bifid; the lobe beneath the hind
femora occupying nearly the whole length of the joint and scarcely
rounded ; basal joint of fore tarsi with a comb of nine rather short
spines on the outer margin and a row of short fine spines on the
inner margin, the outer angle strongly produced and almost
reaching the apex of the second joint. Shining, finely and sparsely
punctured, more closely on the front and pronotum.

Black, with grey pubescence, a narrow band on the inner margin
of the eyes, continued and arched on the vertex, a spot on the
front, hind margin, and anterior angles of the pronotum, a curved
band on the scutellum, a median spot and the posterior angles on
the median segment, and a transverse band on each side on dorsal
segments 1-4 pale yellow; mandibles, tegule, tarsi, and pygidium
testaceous brown.

Length 9 mm.

Hab. Aden district.

25. ANTHOBOSCA MINIMA Turn.

Anthobosca minuma Turn. Trans. Ent. Soc. London, p. 398
(@Si0) oe

@. Radial cell narrowly truncate at the apex; second abscissa
of the radius more than twice as long as the first, but much shorter
than the third. 'Tarsal ungues bifid. Shining, sparsely and finely
punctured ; apical dorsal segment strongly punctured and covered
with stiff fulvous hairs.

Black ; mandibles, antenne, and legs testaceous brown ;

STUDIES IN THE FOSSORIAL WASPS. 741

abdomen dark reddish brown; yellow marks as in arabica, but
the yellow band on each side of the first dorsal segment is
absent.

Length 5 mm.

Hab. Mombasa.

26. ANTHOBOSCA CHILENSIS Gueér.

Cosila chilensis Guér. Voy. ‘Coquille,’ Zool. ii. p. 249 (1839), 2;
Spinola, in Gay’s Hist. Fis. Chile, Zool. vi. p. 312 (1851), g 2.

Myzine flavicornis Sm. Descr. new spec. Hymen. p. 183
(AESiAS) Oe

2. Radial cell rounded at the apex; second abscissa of the
radius twice as long as the first, but distinctly shorter than the
third; first recurrent nervure received close to the middle of
the second cubital cell, second at about one-fifth from the base
of the third cubital cell. Tarsal ungues bifid; lobe beneath the
apical third of the hind femora scarcely rounded, basal joint of
hind tarsi with a thinly-set row of very short spines beneath.
Finely and rather closely punctured, median segment finely
rugulose ; abdomen shining, very finely and sparsely punctured.
Basal joint of fore tarsi not strongly produced at the outer apical
angle.

Black, with long black pubescence ; calcaria black; flagellum
bright orange. Wings fusco-violaceous.

3. Third abscissa of the radius twice as long as the second in
some specimens, shorter than the second in others ; first recurrent
nervure received by the second cubital cell at a distance from the
base slightly less than the length of the first transverse cubital
nervure, second either interstitial with the second transverse
cubital nervure or received a little before the apex of the second
eubital cell. Apical joint of antenne no longer than the
penultimate. Basal abdominal segment nearly twice as broad
at the apex as long. Very finely and closely punctured, minutely
on the abdomen.

Colours as in the female.

Length, 2 22 mm., ¢ 16 mm.

Hab. Chile.

27. ANTHOBOSCA CARBONARIA Burm.
Myzine carbonaria Burm. Stett. ent. Zeit. xxxvii. p- 168

(1876), @.

Anthobosca carbonaria Turn. Trans. Ent. Soc. London, p. 83
(1908), 2.

Cosila carbonaria Bréthes, Ann. Mus. Buen. Aires, xx. p. 256
(1910), 2.

Q@. Radial cell narrowly rounded at the apex; second abscissa
of the radius longer than the first and third combined; first
recurrent nervure received at about two-fifths from the base of

742 MR. R. E. TURNER ON

the second cubital cell, second before one-third from the base
of the third cubital cell, which is nearly twice as long on the
cubitus as on the radius. Basal joint of hind tarsi with a row of
fine spines beneath; basal joint of fore tarsi with seven spines
above and a row of short spines beneath; tarsal ungues cleft ;
apical half of hind femora broadly rounded beneath ; tarsal ungues
cleft. Shining, finely and sparsely punctured; front and pro-
notum more closely and coarsely punctured; median segment
subopaque, very finely and closely punctured, the posterior slope
rather indistinctly transversely striated. Pygidium densely clothed
with fusco-ferruginous pubescence. Basal joint of fore tarsi not
very strongly produced at outer angle.

Black, with grey pubescence; calcaria pale brownish. Wings
rather light fuscous, fusco-hyaline at the apex.

Length 20 mm.

Hab. Nova Fribourg, 8. Brazil.

28. ANTHOBOSCA BIPUNCTATA Perty.

Tiphia bipunctata Perty, Delect. anim. artic. Brasil. p. 139
(1833), 2.

Myzine bipunctata Sm. Cat. Hym. B. M. ii. p. 76 (1855).

Anthobosca bipunctata Turn. Trans. Ent. Soc. London, p. 83
(1908).

@. Radial cell narrowly rounded at the apex; first and fourth
abscisse of the radius about equal in length, second and third also
nearly equal to each other and twice as long as the first; the first
recurrent nervure received just beyond two-fifths from the base of
the second cubital cell, second at the middle of the third cubital
cell, which is shorter on the radius than on the cubitus. Tarsal
ungues bifid; lobe beneath the posterior femora extending over
the apicai two-thirds of their length and scarcely rounded. Shining
and very sparsely punctured, the front smooth, apical dorsal seg-
ment covered with long fuscous hair. Basal joint of fore tarsi
strongly produced at the outer apical angle.

Black ; a yellow spot on each side of the third dorsal segment.
Wings fusco-hyaline, nervures fuscous; calearia whitish.

Length 16 mm.

Hab. Minas Geraes, Brazil.

Allied to carbonaria Burm., but has the lobe beneath the hind
femora longer, the basal joint of the fore tarsi is more produced,
and the puncturation is sparser. A. bipustulata Turn. is probably
the male of this species.

29. ANTHOBOSCA ALBOMACULATA Sin. (PI. LXXXTI. figs. 7, 8.)
Myzine albomaculata Sm. Descr. new spec. Hymen. p. 181

Gus) "2 Se
Myzine lecointet Diicke, Rev. entom. p. 146 (1907), 9.

@. Radial cell narrowly rounded at the apex; second abscissa

STUDIES IN THE FOSSORIAL WASPS, 743

of the radius a little longer than the first, the third longer than
the first and second combined ; first recurrent nervure received
just beyond two-fifths from the base of the second cubital cell,
second close to the middle of the third cubital cell. Tarsal ungues
bifid ; lobe beneath the basal half of the hind femora not very
prominent and very feebly rounded; basal joint of hind tarsi
with a row of short hairs beneath; basal joint of fore tarsi
strongly produced at the outer apical angle. Shining and sparsely
punctured ; median segment opaque, closely and minutely punc-
tured; apical dorsal segment coarsely punctured and_ thickly
clothed with long fuscous hairs.

Black ; a spot on each side near the anterior angles of the pro-
notum, a spot on the mesonotum, a spot at the base of the median
segment and one at each of the posterior angles, and a spot on each
side of the second and third dorsal segments yellowish white ;
calearia whitish. Wings pale fusco-hyaline, nervures fuscous.

3. Second abscissa of radius twice as long as first, third about
half as long again as second; first recurrent nervure received a
little beyond the middle of the second cubital cell, second at two-
fifths from the base of the third cubital cell. Tarsal ungues bifid ;
hind tibie serrate on the outer margin. Basal abdominal segment
nearly half as long again as the breadth at the apex. Finely and
rather closely punctured, abdomen finely shagreened.

Black ; mandibles, clypeus, scape beneath, a spot on each side
of the pronotum, a spot on the mesonotum, a spot on the
scuteilum, one on the postscutellum, another at the apex of
the median segment, the base of the tibiz, and the tarsi pale
yellow. Wings hyaline iridescent, nervures nearly black.

Length, 2 12 mm., ¢ 9 mm.

Hab. Amazon, from Para to Ega.

30. ANTHOBOSCA ANTENNATA Sm.

Anthobosca antennata Sm. Descr. new spec. Hymen. p. 174
(USS) Sc

Cosila jheringi Saussure in Grandidier, Hist. Madagascar, xx.
p. 234 (1892), 2.

Thynnus antennatus D. T. Cat. Hymen. viii. p. 101 (1897), 3.

@. Radial cell very narrowly rounded at the apex; third
abscissa of radius half as long again as the second, which is
nearly twice as long as the first, the fourth shorter than the first ;
first recurrent nervure received a little before the middle of the
second cubital cell, second at one-fifth from the base of the third
cubital cell. Tarsal ungues bifid ; lobe beneath the hind femora
commencing near the base and extending to the apex, broadly
rounded ; basal joint of hind tarsi with a row of very short fine
spines beneath ; basal joint of fore tarsi strongly produced at the
outer apical angle. Shining, very finely and rather sparsely
punctured ; median segment subopaque, very closely and minutely

744 MR. R. E. TURNER ON

punctured ; apical dorsal segment densely clothed with long fulvous
hairs.

Black ; mandibles at the base, antenne, and legs ferruginous ;
an interrupted band on the posterior margin of the pronotum, a
spot on the mesonotum, one at the base of the median segment
and another at each of the posterior angles, and an interrupted
band on the four basal dorsal segments, more broadly interrupted
on the second than on the other segments, yellow. Wings very
pale flavo-hyaline, nervures ferruginous.

3. Third abscissa of the radius half as long again as the second,
which is about equal to the fourth and twice as long as the first ;
first recurrent nervure received at or a little beyond the middle
of the second cubital cell, second at one-third from the base of the
third cubital cell. Tarsal ungues bifid; hind tibie serrate on the
outer margin. Antenne short and stout, tapering to the apex;
pronotum rounded anteriorly ; first abdominal segment a little
longer than the breadth at the apex, the apical dorsal margin of
the segment broadly rounded. Finely and closely punctured ;
abdomen finely shagreened.

Black; mandibles, clypeus, scape beneath, posterior margin of
the pronotum broadly, a spot on the mesonotum, one on the
scutellum, another on the postscutellum, and the greater part of
the tibiz and tarsi yellow. Wings hyaline, nervures fuscous.

Length, 2 13-16 mm., ¢ 12-14 mm.

Hab. 8. Brazil; Rio Grande do Sul.

As I have noticed before, the antennz in the typeof antennata
are a little shorter and stouter than in the males sent with jhering?.
This difference may possibly prove to be specific, but I do not think
it is.

*31. ANTHOBOSCA ERYTHROPYGA Burm.

Myzine erythropyga Burm. Stett. ent. Zeit. xxxvil. p. 169
(USO, Dei

Anthobosca erythropyga Turn. Trans. Ent. Soc. London, p. 83
1908).
eb erythropyga Bréthes, Ann. Mus. Buen. Aires, xx. p. 256
(1910).

I have not seen this species. Schrottky suggests that it is
identical with A. jheringi Sauss., but Burmeister’s description
gives the legs of the male as red, of the female black, and the five
basal abdominal segments in both sexes with lateral yellow spots,
whereas in jheringi the four basal segments are banded with yellow
and the legs red in the female, and the male has the abdomen
without yellow marks and the legs yellow and black. I do not
think there is sufficient ground for considering the two to be
identical, especially as Schrottky had not seen specimens of
jheringt.

tee

1.
2.
3.

10.

11.

13.

14.

STUDIES IN THE FOSSORIAL WASPS.

. ANTHOBOSCA APICALIS Sichel.

745

ae ae. Sichel, Sauss. et Sichel, Cat. spec. gen. eSeolia,

p. 262 (1864), 2.

my have not seen this species, which may possibly be identical
with carbonaria Burm., but the dorsal segments of that species
are not reddish laterally.

Males.
Species from the Old’) World :.).....000050. 0.0. cs ei eee
Species from South America............ 0.0.60 eee cee cee eee eee
Australian species 000.000. 0. ccc seeseceencen sce eee eed eee cee ceeces

African species .... selaecas

Tarsal ungues with ¢ a blunt: ‘lobe ‘at ‘the base ; second
and third cubital cells each receiving a recurrent
nervure

Tarsal ungues bitid, “without a : basal lobe ; 5 second cubital
cell receiving both recurrent nervures

. Highth and three following joints of the flagellum sub-

tuberculate beneath, apical jomt longer than penulti-
mate and slightly curved. Wings fusco-violaceous...

None of the joints: of the flagellum subtuberculate
beneath, apical joint no longer than penultimate.
Wings hyaline or subhyaline ....

. First abdominal segment distinctly Jonger ‘than the

second; fully half as long again as its breadth at the
apex .

First abdominal. segment not distinctly longer. than the
second ; scarcely, if at all, longer than its apical
breadth

. Thorax and abdomen v wholly black .

Pronotum at least marked with yellow .. She taal A bene
Legs ferruginous

Legs black

5 Median segment transversely rugulose, very “short and

broad ; ‘anteri ior tibiee black beneath .....................
Median segment finely punctured, not very short or
broad ; anterior tibize ferruginous beneath

. Legs ferruginous..

Legs black, more or less marked with yellow... ria

Postscutellum yellow, legs marked with yellow ; first
transverse cubital nervure as long as the second or
longer and strongly bent near the eubitus os...

Postscutellum and legs without yellow markings ;
transverse cubital nervure much shorter than the
second, and not bent near the cubitus..

Five basal abdominal segments with yellowish lateral
spots; cubitus of hind wing originating just before
the transverse median nervure, almost interstitial

Five basal abdominal segments immaculate; cubitus
of hind wing originating very distinctly before the
transverse median nervure

. Seventh dorsal segment with yellowish 1 lateral ‘spots ;

postscutellum yellowish SPE ee mens rafeoaecucsecicesteieke
Seventh dorsal segment and postscutellum immaculate .
Clypeus black ; mesonotum without a yellow spot ; hind
tibize feebly spinose .......
Clypeus yellow ; mesonotum ‘with a “yellow ‘spot ; “hind
tibize feebly Sertater circcscmseheeseeeacd eet eune seatones
Apical margin of clypeus not toothed .. neduie oat
Apical margin of clypeus with two small teeth —

Proc. Zoou., Soc.—1912, No. XLIX.

4,
A, crassicornis Sm.
A. australis Sichel
(=nigripennis Sm.).

6.
A. australasie Guér. |

6.
Uc
9.
A, varipes Sm..'
8.
A. ethiops Sm.
A. nigra Sm.

10.
11.

A. torresensis Turn. |
A, gilesit Turn.
A. moderata Turn.

12.

13.
A. frenchi Tarn.

A. longipalpa Turn.
A. lagardet Turn.

15.
16.

49

746 MR. R. E. TURNER ON

15. Legs wholly ferruginous; mesonotum with a yellow

‘Spot ; tegulee black.. veresterneeesee A, Errans Sm.
Legs ferruginous marked with “yellow ; “mesonotum
= ‘immaculate ; tegulze ae _iSguitushacnenie oeeoaaaen te meee A. flavopicta Turn.
17.

16. Wings hyaline ; Be

Apical half of wings ; with a distinct fusco-violaceous
tinge .... sucies tenons senractignestcatdes deeen dagazaste mes OCdenbaton Camu

17. Abdomen immaculate ....c.cc0ccccccsececsecsssseen A, lacteipennis Cam.

Dorsal abdominal segments 2-4 with large eben

lateral spots 3 Sino A, erythronota Cam.

18. Antenne orange ; wings fusco-violaceous seseseceessssseeee A. Chilensis Guér.
Antennze black; wings hyaline ..........................000 19.

19. Abdomen spotted with ok sohoodssuscasepennbapodddocooson “ele LR AOROVOLIAG LENT.
Abdomen immaculate dc byatahe Noten aunte doe Smamnareerc Matera Meee 20.

20. Legs ferruginous _... neadeaedeansannarnndagsnaonpas 4) (oopanalianng, Iwona.
Legs black, marked with ‘yellow ales sta ech Sh aes 21.

21. Leneth 13mm. Yellow band on pronotum entire ...... A. antennata Sm.

Length 10mm. Yellow band on pronotum interrupted. 4. albomaculata Sm.

33, ANTHOBOSCA AUSTRALASLE Guér. (Pl. LX XXIII. fig. 13.)

Anthobosca australasie Guérin ; Duperry, Voy. ‘ Coquille,’ Zool.
li, p. 237 (1839), ¢.

Anthobosca crabroniformis Sm, Cat. Hym. B. M. vi. p. 59
(1859), 3

Thynaus cathreinit D. T. Cat. Hym. viii. p. 103 (1897), ¢

¢. Pronotum much narrowed anteriorly, nearly as long as the
scutellum. First dorsal abdominal segment slender, more than
half as long again as the apical breadth, much longer than the
second segment. Seventh dorsal segment very broadly rounded
at the apex. Hind tibiz feebly spined. Third abscissa of the
radius longer than the first and second combined ; first recurrent
nervure received close to the middle of the second cubital cell,
second at one-quarter from the base of the third cubital cell ;
first transverse cubital nervure curved, not sharply bent at the
base. Cubitus of the hind wing originating before the transverse
median nervure, separated from it by a distance exceeding half
the length of that nervure.

Black ; mandibles, clypeus, margins of the eyes interrupted on
the summit, pronotum very broadly posteriorly, tegule, a
quadrate spot on the mesonotum, a spot on the mesopleure, the
middle of the scutellum and postscutellum, a curved transverse
band at the apex of the median segment, an interrupted trans-
verse band on dorsal segments 2-4, sometimes also a spot on each
side of the first and fifth segments, a spot on the anterior coxze
and a line on the anterior tibie yellow; legs ferruginous. Wings
hyaline, tinged with yellow ; nervures ferruginous.

Length 16 mm.

Hab. New South Wales, Queensland, as far north as Cairns.

The female is unknown, but the range of the species is about
the same as that of signata Sm.

STUDIES IN THE FOSSORIAL WASPS. TAT

34. ANTHOBOSCA GILESI Turn.

Anthobosca gilesi Turn. Proc. Zool. Soc. London (1910),
p. 308, 3.

g. Antenne shorter than the thorax and median segment
combined ; pronotum much narrowed anteriorly, shorter than
the scutellum. First dorsal segment as broad at the apex as
long, a little shorter than the second segment. Seventh dorsal
segment subtruncate at the apex. Neuration similar to that of
A. australasic, but the position of the first recurrent is variable—
it is received beyond the middle of the second cubital cell in the
type, but at the middle in another specimen.

Black; the base of the mandibles, part of the clypeus, the
pronotum very broadly posteriorly, and the tegule yellow ;
legs ferruginous. Wings hyaline, nervures fuscous. Calcaria
white.

Length 11-14 mm.

Hab. Perth, West Australia.

35. ANTHOBOSCA TORRESENSIS Turn.

Anthobosca torresensis Turn. Proc. Linn. Soc. N.S.W. xxxil.
p- 518 (1907), 3s.

3. Pronotum narrowed anteriorly, nearly as long as the
scutellum. First dorsal segment as broad at the apex as long,
no longer than the second segment; seventh dorsal segment
narrowly truncate at the apex. Second abscissa of the radius
shorter than the first, the two combined much shorter than the
third. First transverse cubital nervure longer than the second,
very sharply bent near the cubitus; first recurrent nervure
received close to the middle of the second cubital cell, second
beyond one-quarter from the base of the third cubital cell.

Black ; legs ferruginous; mandibles at the base, scape beneath,
pronotum very broadly posteriorly, postscutellum, and a line on
the anterior tibie yellow. Wings hyaline, nervures fuscous.
Calearia whitish.

The hind tibiz are almost smooth, the spines nearly obsolete.
Neuration of hind wing as in A. australasic.

Length 7-8 mm.

Hab. Cape York, Queensland.

36. ANTHOBOSCA VARIPES Sin.

Anthobosca varipes Sm. Cat. Hym. B. M. vit. p. 59 (1859), ¢.

Thynnus fischeri D. T. Cat. Hym. viii. p. 106 (1897), ¢.

3g. Pronotum shorter than the scutellum, not very strongly
narrowed anteriorly ; scutellum with a delicate carina from the
base to the apex. First dorsal segment nearly as broad at the
apex as long, very little longer than the second segment ;
seventh dorsal segment broadly subtruncate at the apex. Hind
tibie very feebly serrate. First transverse cubital nervure

748 , MR. R. E. TURNER ON

oblique, not sharply bent near the cubitus, longer than the
second. Second abscissa of the radius about. twice as long as
the first, the two combined shorter than the third. First
recurrent nervure received distinctly beyond the middle of the
second cubital cell, second beyond one-third from the base of
the third cubital cell. Hind wing as in A. australasiw. Apical
joint of maxillary palpi as in A. australasic, not filiform and
scarcely longer than the penultimate joint.

Black ; the legs ferruginous. Wings hyaline, nervures
fuscous.

Length 12-15 mm.

Hab. Adelaide, S. Australia; Ararat and Ringwood, Victoria.

37. ANTHOBOSCA NIGRA Sin.

Anthobosca nigra Sm. Cat. Hym. B. M. vii. p. 59 (1859), 3.

Thynnus reischit D, T. Cat. Hymen. viii. p. 114 (1897), 3

¢. Clypeus truncate at the apex; antennz nearly as long as
the thorax and median segment combined. Pronotum short,
scarcely more than half as long as the scutellum, very slightly
narrowed anteriorly. First dorsal segment longer than the apical
breadth ; seventh dorsal segment broadly rounded at the apex.
Hind tibie rather feebly serrate ; hind femora without a lobe at
the apex beneath. Second abscissa of the radius longer than the
first, the two combined much shorter than the third; first
transverse cubital nervure oblique, sharply bent close to the
cubitus; first recurrent nervure received at the middle of the
second cubital cell, second at two-fifths from the base of the third
cubital cell. The distance between the cubitus of the hind wing
and the transverse median nervure is greater than half the length
of that nervure.

Black ; the fore tibie ferruginous within; calcaria whitish.
Wings hyaline, nervures fuscous.

Length 9-12 mm.

Hab. Victoria and South Australia.
_. Differs from varipes in the absence of a carina on the scutellitm,
in the colour of the legs, and the. more distinet serration of the
hind tibie.

38. ANTHOBOSCA ZTHIOPS Sm.
Anthobosca wthiops Sm. Descr. new spec. Hymen. p. 175

(1879); o-
Thynnus stolzii D. T. Catal. Hymen. viii. p. 116 (1897), 6

g. Antenne stouter than in other Australian species ;»pro-
notum very slightly narrowed anteriorly, shorter than the
seutellum. Head and thorax very closely and finely punctured,
median segment transversely rugulose, short and not as strongly
convex as in other species. First dorsal segment very little
longer than the apical breadth ; seventh dorsal segment narrowly

|
|
;

SRE a or ee

STUDIES IN THE FOSSORIAL WASPS, 749

truncate at the apex. Hind tibie spinose ; hind femora without
a rounded lobe at the apex beneath. Second abscissa of the
radius equal to the first, the two combined shorter than the third ;
first transverse cubital nervure oblique, sharply bent close to the
cubitus; first recurrent nervure received close to the middle of
the second cubital cell, second just before one-third from the
apex of the third cubital cell. Cubitus of the hind wing
separated from the transverse median nervure by a distance equal
to one-third of the length of that nervure.

Black; the inner margin of the eyes narrowly whitish; calcaria
whitish ; wings hyaline, tinged with fuscous, nervures fuscous.

Length 14 mm.

Hab. Champion Bay, W. Australia.

39. ANTHOBOSCA MODERATA Turn. (Pl. LX XXTI. fig. 4.)

Anthobosca moderata Turn, Ann. & Mag. Nat. Hist. (8) ii.
p- 482 (1909), 3.

3d. A minute tubercle in the middle of the clypeus; pronotum
not much narrowed anteriorly, not more than half as long as the
scutellum ; first dorsal segment a little narrower at the apex than
long, a little longer than the second segment. Hind tibie very
distinctly spinose ; hind femora with a very distinct rounded lobe
at the apex beneath. Second abscissa of the radius longer than
the first, the two combined about equal to the third; first
recurrent nervure received at the middle of the second cubital
cell, second beyond one-third from the base of the third cubital
cell. Cubitus of the hind wing originating just before the
transverse median nervure.

Black; mandibles at the base, clypeus, posterior margin of
pronotum, tegule, a spot on the mesonotum, postscutellum, a
spot on each side at the apex of the median segment, a small
lateral spot on each side of dorsal segments 1—5, a large spot on
each side of the seventh segment, the base of the tibize and
the base of the tarsi whitish yellow; the apex of the seventh
dorsal segment testaceous. Wings hyaline, tinged with fuscous ;
nervures fuscous.

Length 12 mm.

Hab. Townsville, Queensland.

40. ANTHOBOSCA FRENCHI Turn.

Anthobosca frenchi Turn. Proc. Linn. Soc. N.S.W. xxxil.
p- 518 (1907), 3.

3. Clypeus truncate at the apex; pronotum much shorter
than the scutellum, very slightly narrowed anteriorly. First
dorsal segment no longer than the apical breadth ; seventh dorsal
segment broadly rounded at the apex. Hind tibize very feebly
serrate; hind femora without a lobe at the apex beneath.
Second abscissa of the radius longer than the first, the two

750 MR. R. bE. TURNER ON

combined as long as the third; first transverse cubital nervure
oblique, strongly bent near the cubitus, longer than the second ;
first recurrent nervure received at the middle of the second
cubital cell, second at one-third from the apex of the third cubital
cell ; the distance between the cubitus of the hind wing and the
transverse median nervure equal to less than half the length of
that nervure.

Black ; base of the mandibles, a small spot round the base of
each antenna, the hind margin of the pronotum very narrowly,
tegule, anterior tibize in front, and the base of the hind tibiz pale
yellow ; apex of the mandibles and fore tarsi fusco-ferruginous.
Wings hyaline, nervures dark fuscous.

Length 7-8 mm.

Hab. Victoria.

41. ANTHOBOSCA LAGARDE! Turn.

Anthobosca lagardei Turn. Trans. Ent. Soc. London (1908),
[Oa chO5, Cie

$. Differs from A. frenchi in the yellow clypeus, yellow spot
on the mesonotum, postscutellum, apex of median segment, and
on each side of the seventh dorsal segment ; in the rounded lobe
at the apex of the hind femora beneath, which is well developed
in lagardet, and in the yellow basal joints of the tarsi.

Length 8 mm.

Hub. Sydney, New South Wales.

42. ANTHOBOSCA LONGIPALPA Turn.

Anthobosca longipalpa Turn. Proc. Linn. Soc. N.S.W. xxxii.
jos Dil CUM), ere

3. Clypeus very feebly emarginate at the apex; the apical
joint of the maxillary palpi filiform, longer than the penultimate
joint, the three apical joints much more slender than the basal
ones. Pronotum short, very slightly narrowed anteriorly. First
dorsal segment a little longer than the apical breadth ; seventh
dorsal segment narrowly truncate at the apex. Hind tibiz feebly
spinose; the lobe at the apex of the hind femora beneath not
distinct. Second abscissa of the radius nearly twice as long
as the first, the two combined longer than the third. First
transverse cubital nervure curved, much longer than the second,
and rather sharply bent near the cubitus; recurrent nervures
received a little before the middle of the second and third cubital
cells. Cubitus of the hind wing originating before the transverse
median nervure, the distance between it and that nervure equal
to about one-third of the length of the transverse median
nervure.

Black; posterior margin of the pronotum very narrowly, a
transverse line on the postscutellum, a spot on each side of the
seventh dorsal segment, and the base of the hind tibie whitish

STUDIES IN THE FOSSORTAL WASPS, 751

yellow ; tibiz fusco-ferruginous; calcaria white. Wings hyaline,
nervures fuscous,

Length 12 mm.

Hab. Cairns, Queensland.

43. ANTHOBOSCA CRASSICORNIS Sm. (Pl. LX XXI. fig. 1.)

Tachypterus crassicornis Sm. Cat. Hym. B. M. vi. p. 64
(1859), 3.

Trachypterus crassicornis D. T. Cat. Hymen. vii. p. 120
(USIMGs

Anthobosca crassicornis Turn. Proc. Linn. Soc. N.S.W. xxxii.
p. 519 (1907), 3.

3. Clypeus produced in the middle and rather narrowly trun-
cate at the apex. Antenne slightly longer than the thorax and
median segment combined. Head and thorax very closely and
not very finely punctured. First abdominal segment scarcely
longer than the second ; the apex of the dorsal segment rounded
and a little, but not very distinctly, narrower than the first
segment is long. Posterior tibize very distinctly serrate, tarsal
ungues bifid. Second abscissa of the radius longer than the
third; both recurrent nervures received by the second cubital
cell, the first at two-fifths from the base, the second at about
nine-tenths or more from the base. Third cubital cell as long on
the radius as on the cubitus.

Black, with long grey pubescence; mandibles, clypeus, lower
portion of the inner margin of the eyes, anterior coxe, trochanters
and femora beneath, and a line beneath the intermediate femora
yellow ; second and third abdominal segments and the base of
the fourth, tibie, tarsi, hind femora, and intermediate femora
above ferruginous. Wings hyaline, nervures fusco-ferruginous,
stigma fuscous.

Length 13 mm.

Hab. Austyalia.

A variety has the ferruginous colour much obscured. As
mentioned under that species, I believe this to be the male of
A. clypeata Sm , owing to the similarity of the structure of the
tarsal ungues, the position of the recurrent nervures, the length
of the second abscissa of the radius, and some similarity of
colour.

44, ANTHOBOSCA ERRANS Sm.

Anthobosea errans Sm. Descr. new spec. Hymen. p. 174
(1879), 3.

3. Clypeus broadly truncate at the apex, not toothed; an-
tenne short and stout. Pronotum shorter than the scutellum,
distinctly narrowed anteriorly ; scutellum and median segment
obscurely longitudinally carinated in the middle. First dorsal
segment longer than the apical breadth; seventh dorsal segment

752 MR. R. E. TURNER ON

narrowly truncate at the apex. Hind tibiz serrate ; hind femora
with a rounded lobe at the apex beneath. Second abscissa of the
radius nearly twice as long as the first, the two combined shorter
than the third; first transverse cubital nervure oblique, strongly
bent close to the cubitus, longer than the second ; the distance
between the cubitus of the hind wing and the transverse median
nervure equal to about one-third of the length of that nervure.
Tarsal ungues bifid.

Black ; base of the mandibles, clypeus, a spot on each side of
the pronotum, a spot on the mesonotum, one on the scutellum
and a transverse line on the postscutellum yellow; legs ferru-
ginous. Wings hyaline, nervures fusco-ferruginous.

Length 13 mm.

Hab. Zaluland.

45, ANTHOBOSCGA FLAVOPICTA Turn.

Anthobosca flavopicta Turn. Trans. Ent. Soc. London, p. 399
(1910), ¢ pl. L. fig. 4.

3. Clypeus subtruncate at the apex, not toothed. Posterior
tibiz serrate, tarsal ungues bifid.

Black; clypeus with a black spot in the middle, inner margin
of the eyes narrowly, a small spot behind the eyes near the
summit, a broad band on the posterior margin of the pronotum,
tegule, a large spot on the scutellum, one on the postscutellum,
the apex of the fore femora, and the anterior and intermediate
tibie and tarsi above pale yellow; tibie and tarsi ferruginous
brown. Wings hyaline, nervures black.

Length 11 mm.

Hab. Zanzibar.

46. ANTHOBOSCA BIDENTATA Cam.

Odontothynnus bidentatus Cam. Rec. Albany Mus. i. p. 162
(1904), 3.

3. Apex of clypeus bidentate. Hind tibie serrate. First
transverse cubital nervure rounded. Wings hyaline, the apical
half fusco-violaceous.

Black ; lower half of inner orbits of the eyes, face, clypeus,
base of mandibles, base and outer side of fore and intermediate
tibiz, calcaria, tarsi, base of hind tibie, and the posterior margin
of the pronotum broadly yellow.

Length 11-12 mm.

Hab. Grahamstown, 8. Africa.

47. ANTHOBOSCA LACTEIPENNIS Cam.

Odontothynnus lacteipennis Cam. Rec. Albany Mus. i. p. 162
(1904), 3.
3g. Apex of clypeus bidentate. Hind tibize serrate. First

STUDIES IN THE FOSSORIAL WASPS. (5S

transverse cubital nervure oblique. Wings hyaline, nervures
white.

Colours as in bidentata.

Length 10 mm.

Hab. Grahamstown, 8. Africa.

I have seen neither bidentata nor lacteipennis. Cameron’s
statement that the ungues of the hind tarsi are simple, not bifid,
needs confirmation.

48. ANTHOBOSCA BIPUSTULATA Turn.

Anthobosca bipustulata Turn. Ann. & Mag. Nat. Hist. (8) iv.
p. 343 (1909), 3.

do. Slender; antenne stout, tapering towards the apex;
pronotum scarcely more than half as long as the scutellum,
moderately narrowed anteriorly. First dorsal segment nearly half
as long again as the apical breadth; seventh dorsal segment
rather narrowly rounded at the apex. ‘Tarsal ungues bifid ;
hind tibiz serrate; hind femora without a distinct lobe at the
apex beneath. Apical joint of maxillary palpi filiform. Second
abscissa of the radius twice as long as the first, the two combined
nearly as long as the third; first transverse cubital nervure
oblique, sharply bent near the cubitus, longer than the second ;
the distance between the cubitus of the hind wing and the
transverse median nervure equal to half the length of that
nervure.

Black: mandibles, clypeus, scape beneath, a spot on each side
on the posterior margin of the pronotum, the base of the tegule,
a small spot on the mesonotum, one on the scutellum, another on
the postscutellum, the two latter spots sometimes absent, and the
anterior coxze beneath yellow; legs ferruginous. Wings hyaline,
nervures fuscous.

Length 8-12 mm.

Hab. Barbacena, Minas Geraes.

A variety has the legs black, marked with yellow.

EXPLANATION OF THE PLATES.

Prats LXXXI.

Fig. Hig
1. Anthobosca crassicornis Sm. 6. 10. Braunsomeria quadraticeps Turn.
2. Anthobosca clypeata Sm. &. ©
3. Anthobosca fastuosa Sm. @. 11. Myzine stigma Turn. ¢.
4, Anthobosca moderata Turn. 6. 12. Elis (Mesa) alicie Turn. 2.
5. Anthobosca erythronota Cam. 6. 13. Elis (Mesa) longiventris Turn. 6.
6. Anthobosca erythronota Cam. 2. 14. Myzine braunsi Turn. 3.
4. Anthobosca albomaculata Sm. 6. 15. Hlis (Mesa) dimidiaticornis
8. Anthobosca albomaculata Sm. &. Bingh. ¢.
9. Braunsomeria quadraticeps Turn. | 16. lis (Mesa) tricolor Sm. &,
3

Proc. Zoot. Soc.—1912, No. L. 50

st
or
&

MR. ABEL CHAPMAN ON

Prare LXXXII.
Neuration of Wings.

Fig. Fig.

1. Anthobosca australis Sichel. 9. 9. Elis (Mesa) ruficeps Sm. 9.

2. Anthobosca australis Sichel. g. 10. Hlis (Mesa) ruficeps Sm. ¢.

3. Anthobosca anthracina Sm. 9. 11. Hlis (Mesa) tricolor Sm. ¢.

4, Anthobosca erythronota Cam. ?. 12. Elis combusta Sm. 2.

5. Anthobosca elypeata smn, @ 13. Myzine stigma Turn. g.

6. Anthobosea insularis Sm. §. 14. Myzine brawnsi Turn. 6.

7. Braunsomeria quadraticeps Turn. | 15. Myzine constrictiventris Turn. 3.

3. 16. Myzine abdominalis Guér. 2.
8. Elis (Mesa) alicie Turn. ¢.

Prats LXXXIIt.
Exoskeletal Structures.

Vig. 1. Anthobosca australis Sichel. 9. Suture between two basal abdominal
segments.
2. Elis (Mesa) ruficeps Sm. §. Suture between two basal abdominal
segments.
3. Anthobosca australis Sichel. 6. Apical ventral segment.
4. Elis (Mesa) ruficepsSm. g. Apical dorsal segment.
5. Myzine abdominalis Guér. ¢. 3 5 33
6. Anthobosca australis Sichel. 9. 'Tarsal unguis.
7. Anthobosca clypeata Sm. &. 3 5
8. Anthobosca insularis Sm. 9. Intermediate and hind coxee.
9. Hlis (Mesa) alicie Turn. 2. Basal jot of hind tarsus.
10. Elis (Mesa) ruficeps Sm. &. o a a
11. Elis (Mesa) longiventris Turn. 3. Two basal segments of abdomen.
12. Myzine constrictiventris Turn. g.
13. Anthobosca australasie Guér. 3.
14, Anthobosca australis Sichel. g. RS
15. Hlis (Mesa) ruficepsSm. 6. us s =
16. Elis re ruficeps Sm. 2. Intermediate and hind coxe.

40. Notes on the Spanish Ibex.
By ABEL CHapMAN, F.Z.S.

[Received April 13, 1912: Read May 7, 1919. ]

INDEX.
Page

Distal buts ones ee boo: wag EES

In Professor Angel Cabrera’s most informative paper on Capra
pyrenaica,* two small points occur to me as worthy of brief
comment. In discussing the present distribution of the Spanish
Wild-Goat in some six isolated colonies, Dr. Cabrera takes
exception to the remark in ‘ Unexplored Spain > (by Walter Buck,
U.M.Z.S., and myself) that they had been so isolated ‘‘during ages.”
Well, the term used in our former book (‘ Wild Spain’) was
“during centuries,” and that is certainly more definitive and
probably more accurate. Dr. Cabrera, however, goes on to state
that there exist “strong reasons for believing that in the past

* P.Z,S. 1911, p. 963.

THE SPANISH IBEX, 15D.

|inferentially as late as the middle of the seventeenth century—
say 250 years ago], Ibex inhabited every suitable point of almost
every mountain-ridge in Spain.” ‘The only reason actually
adduced, however, is the prevalence of place-names based upon,
or compounded with, the Spanish word Cabra = goat. Such
names, lt is true, are ubiquitous; but it would never have
occurred to me that those names necessarily refer to the wild
goat. Spain is a land of goats, and many localities bearing names
such as Sierra de las Cabras, Cabrales, Cebrero, and so on, are not
at all adapted to the nature and requirements of the wild Ibex.
I would suggest that, in many cases, the names merely indicate
the existence of suitable local pasturage for domestic goats, which
are herded everywhere.

Again, Dr. Cabrera translates the Spanish name of the Ibex,
Cabra montés, as equivalent to ‘“‘ Mountain-Goat.” Now it would
be nothing less than presumptuous for me, a foreigner with but a
limited colloquial knowledge of the Spanish tongue, to question
his rendering. Ido not doso. I accept that as the pure classic
Castilian of Madrid. But I do venture to say that, in wilder
Spain, the term monte, with its derivative adjective montés
(pronounced montéss), possesses quite a different signification.
Monte may occasionally, and in combination, be used to indicate
a hill or mountain ; but in its ordinary provincial sense, it signifies
scrub or brushwood, Thus the wild-cat, which is equally common
on lowland or sierra, is known as Gato montés = Scrub-Cat: on
the low-lying plains of Andalucia or Estremadura, the expression
feses montéses includes all the scrub-haunting animals—such as
deer, wild-boar, lynx, ete.

Now, viewed in this light, it has always appeared to me utterly
inexplicable and incongruous to apply the name Cabra montés, ox
Scrub-Goat, to the Ibex of the higher ranges, such as the Sierras
de Grédos and Nevada, where the Ibex live exclusively amidst
rock-regions far above the topmost levels of scrub, But such
incongruity would disappear if Dy. Cabrera’s assumption were
correct, that the Ibex, up toa couple of centuries ago, occupied the
whole vast area shown in the map at p. 965, suprd. Was such the
distribution, that name would become appropriate enough, since
an immense proportion of the dotted area consists, not of high
mountains at all, but of low serub-clad hills, Such country
might appear, to preconceived ideas, totally unsuitable for Ibex :
but we have the fact before us (as fully explained in our books on
Spain) that in several of the lower Mediterranean sierras (some
of which are bush-clad to the summits) the local Ibex do to-day
take kindly to a bush-haunting habit. Indeed, in such situations,
it is obvious, they have no other option.

This latter point tends to support Dr. Cabrera’s assumption,
and equally, of course, undermines our own.

In our two books we had pointed out that the Ibex of the two
extreme ends of Spain (7. e., those of the Pyrenees and those of

156 ON THE SPANISH IBEX.

the Mediterranean sierras) most nearly assimilated to each other
in their more flattened and laterally compressed horns*. It is
gratifying to find that our rough field-observations are now
corroborated by Dr. Cabrera’s skilled investigation and careful
cross-sections. But again, it appears anomalous—assuming that
complete isolation only commenced some two centuries ago—that
the central group (now honoured with subspecific rank as C. p.
victori@) should have developed the greater difference.

The females and young males of the Spanish Ibex are devoid
of the dark dorsal stripe, as is correctly shown in the plate in
‘ Unexplored Spain,’ at p. 140. They are of a dun-brown, uni-
colorous in coat as the Spanish Red-Deer ; but Dr. Cabrera is quite
justified in criticizing the second plate (op. cit. p. 967), at p. 216.
That slip should be debited, in the first instance, to the artist,
Mr. E. Caldwell, but the fault is wholly mine, since I should have
detected the mistake and had it corrected before passing the
drawing for reproduction.

If permissible to express an opinion on the three beautiful
plates given by Dr. Cabrera, I would say that in life the Spanish
Ibex is rounder and bulkier in the barrel than can ever be
gathered from museum specimens, since skins shrink.

Im conclusion, may I express a fervent hope (since interest in
her vanishing Ibex has been aroused in Spain) that further
protection may be extended to the few surviving colonies 2
Within my own time, Ibex have been exterminated in several of
their earlier haunts. To-day they are at their last gasp in the
Pyrenees and in the Gerez (Portugal). Fortunately, in Grédos,
Morena, and Bermeja, their future has been assured—though
only at the eleventh hour. Can our Spanish friends not see to
safeguarding the much-menaced remnant that yet survives on the
main chain of Nevada ?

* Unfortunately, in ‘ Unexplored Spain,’ in an effort to be concise, and to avoid
repetition, we omitted the word “laterally”; but a reference to ‘Wild Spain,’
p. 129, makes our meaning clear.

Papers (continued).

Page

28. Hxperimental Pheasant-breeding. By Ross Haic Tuomas, F.Z.8. (Pls. LXIV.-LXVII.) 539

30.

31.

3a.

30.

86.

37.

38.

39.

40.

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. By
Sir Georer H. Kenricn, F.Z.S. (Pl. LXVIIL.) 22... eccese sec e cece cece sceeee

On a rare Stag (Cervus wallichii) from Nepal recently presented to the Zoological
Society by His Majesty King George. By R. I. Pococn, F.R.S., F.LS., F.Z.S.,
Superintendent of the Gardens. (Text-figs. 66-71.) .........0-00 leslie sVarmoleferetans

Contributions to the Anatomy and Systematic Arrangement of the Cestoidea.—IV. On
a Species of Znermicapsifer from the Hyrax, and on the Genera Zschokkheella, Thy-
sanotenia, and Hyracotenia, By Frank HE. Bepparp, M.A., D.Sc., F.R.S., F.Z.S.,
Prosector to the Society. (Text-figs. 72-83) .o0......-eee..+0> Miao Gente Asana 3e

. Additional Notes on the Living Specimens of the Australian Lung-fish (Ceratodus

forsteri) in the Collection of the Zoological Society of London. By Basurorp Daay.
(Text-figs. 84 & 85.) seeceeeeeeesese sore eco ee se 20 88 e@eoean se 2a 22202888828 25 oO e@seee

The Circulatory System of the Common Grass-Snake (Tropidonotus natrix). By
Cuas. H. C’Donocuuz, B.Sc., F.Z.S., Assistant to the Jodrell Professor of Zoology,
University College, London. (Pls. LXX.—LXXII. and Text-figs. 86-91.) ...,-...- .

. A First Account of the Courtship of the Redshank (Totanus calidris), By Juuian 8.

Huxtny, Lecturer of Balliol College, Oxford ........c.ccsceersee Sears ataieletaket ers

Some Brackish-water Amphipoda from the mouths of the Weser and the Elbe, and from
the Baltic. By E. W. Sexton, Marine Biological Laboratory, Plymouth. (Pls. LXXITT.
Ge DRO.) 2a a sae pa aoOGhe De UO MO BOROOr Sbarro ons SPS ale cche orale cioetey everehe ate.

Descriptions of new Fishes of the Family Loricariide in the British Museum
Collection. By C. Tarm Ruean, M.A., F.Z.S. (Pls. LXXV.-LXXVII.) ..........

On a Collection of Fishes made by Mr. A. Blayney Percival in British Hast Africa to
the East of Lake Baringo. By G. A. Bounenerr, F.B.S., F.Z.S. (Pls. LXXVIIT—
HIANONIK 2) oc aie odie eT ea nS eet BS a MSG eI) CIE oh GU AG eo erent ana

Contributions to the Anatomy and Systematic Arrangement of the Cestoidea.——V. On

‘a new Genus (Dasywrotenia) from the Tasmanian Devil (Dasyurus wrsinus), the type

of a new Family. By Frank EH. Bupparp, M.A., D.Sc., F.R.S., F.Z.S., Prosector to
iiesocicty.) (Vext-tigss Q2-UOL yin aeiste «yee ciate oe ESSA aR NAA) ie Mae :

Studies in the Fossorial Wasps of the Family Scoliide, Subfamilies EHlidinz and
Anthoboscinez, By Rowzanp E. Turner, F.Z.8., F.E.S. (Pls. LXXXI-LXXXIIT.)

Notes on the Spanish Ibex. By Apen Cuarman, F.Z.S. ........--e0ee sees startin eta :

046

558

576

612

656

666

(er)
=~]
Lo

696

jos

LIST OF PLATES.

1912, Part III. (pp. 505-576). /
Plate . Pags
LXI.
LXII. Alcyonaria from Singapore ...+....+-..6 ye eeecees -.. 505
LXIII.
LXIV. }
bes. \Meathers from Pheasantsis? sels. « «=/e10 +. e\= telein'e vielen ema
LXVI.
_XVIL.)
LXVIII. Pyralids: from Dutch New Guinea ....+.....s2.-2c..02 946
XIX. | Young Cariama cristata 6. . 0.2 enews Seance lacs 557.
LXX.
vxxt| Circulatory System of the Grass-Snake ...........-.... 612
LXXII.
LXXIiIl.

Pexiy } Amphipoda from Bremerhaven’, : .cic- sis sce: ts esi eae e aoe

LXXV. 1. Chetostomus lepturus. 2. C. paucispinis. 3. C. palmeri. \

LXXVI, 1. Xenocara multispinis. 2. X. heterorhynchus. 3. Pleco- |
stomus hond@...... BORE Hee choad Tojeta sien ioe eine + 666

LXXVIL 1. Onyloricaria tamane. 2. O. leightoni. 3. Otocinclus |

TURCULUBIRUIVES sa a sual uvelata cielo) cel > aie mieten aki =/ol tae wheat

LXXVIII. Labeo percivali ....... Hime omic oats So bo Coe CaueacocnC \
LXXIX. 1. Barbus argyrotenia. 2. B. mimus. 3. Tilapia
(OGRE) Weaaonoo en caGnKT be Gdodwoodan A0 Ddoaaags a6 ; 672
LXXX. Amphilius oxyrhinus ......++ wee ceeeeee Sais eee kines )
LXXXI. Fossorial Wasps ......... sdododonGode apse
LXXXII. Wing Neuration of ekeorial eee EO OO CHO OG OO OOS dc 696
LXXXIII. Exoskeletal structures of Fossorial Wasps ....-+.......-
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 March.

ge eelpenes, June.
er IE het ied September.
seal Waris December,

‘ Proceedings,’ 1912, Part II. (pp. 241-504), were published on
June 22nd, 1912,

L

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

OOLOGICAL SOCIETY
OF LONDON.

1912.

PARTI.

_ CONTAINING Paces 757 to 913, witH 14 Puates

AND 25 TEXT-FIGURES.

fs JAW 16 1918

DECEMBER 191 y

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

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

[Price Twelve Shillings. |

2 ey Bae be I
Riengl Museve>

LIST OF CONTENTS.

1912, Parr IV. (pp. 757-913).

EXHIBITIONS AND NOTICES,
The Secretary. Report on the Additions to the Society’s Menagerie during the month
Ole Sepia ertetons a cieie's oie is BOUodoogOdSHDaID DOT AGA OF siale(eJofaisie,steeiercha eietel mente tate

Mr. A. Buayney Prrcrvat, F.Z.S8. Exhibition of photographs and lantern-slides of Game
Animals from British Hast Adria i i.crekes - iter « etiete cistela aie ateimetetetietie lee etree

Page

806

806

Mr. D. Seru-Sirtn, F.Z.8. Exhibition of two living specimens of the Lory, Calliptilus —

SOUULGHUUS, LOMO I) eie\lare nie tabeteronetetetel tayreteiatedeetteye) = hers efietatfer<tekeesfeieis ies fete Mege delay tel stele (ed ete

Mr. G. A. Boutenerr, F.RS.,F.Z.8. Notice of a paper entitled “Second Contribution to
our Knowledge of the Varieties of the Wall-Lizard” ........++..0+--- +e sees eee

Mr. E. G.-Boviencrr, F.Z.8. Exhibition of a cocoon of the African Lung-fish
CEMA CHAOGHIGOD, cea ib 98500905 0.090.050.0000 40b508 caso as odo DaDOsOS Neve areicteke

Mr. E. G. B. Mrapze-Wanpo and oruers. Discussion on the Preservation of the Native
Fauna of Great dbritaim #syatelsis« ace siscs tte stare tun ensuc eave cisvcss or wuava ceri ot omeiane aire eee eto

The Secretary. Report on the Additions to the Society's Menagerie during the months
of: May to September, Mn sete cs sirens cis w egenal ye eieiststs siete aeienein > = eter te latent eee ee

Mr. R. Lypuxxer, F.R.S., F.Z.8. Note stating that “ Gazella hayi” = Gazella fuscifrons .

The Rey. T. R. R. Strppine, M.A., F.B.S., F.Z.S. Notice of a memoir on the Crustacea

Tsopoda collected by the ‘ Porcupine’ Expedition in 1869-1870 ..........-05...00

Sir Epuunp G. Loprr, Bt., V.P.Z.S. Demonstration of the capacity of the electric lantern
presented by him to the Society ..............+.- sodeqcoosos Fatelates aiare oot rates

Mrs. Ross Hara Tuomas, F.Z.S8. Exhibition of the Eggs of Phasianus formosanus,
P. versicolor, and their F. 1 and F. 2 offspring. (Text-fig. 126.) ........-..eeee0es

PAPERS.

41, The Local Races of Burchell’s Zebra. By Major J. Srzvenson-Hamitton, C.M.Z.S.,
Game Warden of the Transvaal. (Text-figs. 102-106.) ..... sence ee ee teense eeeees

42. On Dipteropeltis, a New Genus of the Crustacean Order Branchiura. By W. T.
Carman, DiSe.RVZS. AB) XeXeXaVic) se einro ye oars ole cote ne pete nie ence erent teee rectors

43. On two new Larval Trematodes from the Striped Snake (Tropidonotus ordinatus
sirtalis). By Wriu1am Nicom, M.A., D.Sc.. M.D., F.Z.8., Lister Institute of
Preventive Medicine, London. (Text-fiz. 107.) ......-.seceeceeceweees Saat

806

807

807

280%

908

911

912 |

913

912

757

Contents continued on page 3 of Wrapper.

oT

ee Se ee ee ee ee

ee

ee se

THE ZOOLOGICAL SOCIETY OF LONDGN.

Tars Society was founded in 1826 by Sir Sramrorp Rarruuzs,
Mr. J. Sasine, Mr. N. A. Vigors, and other eminent Naturalists,
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/

Patron.
HIS MAJESTY THE KING,

COUNCIL.
HIS GRACH THH DUKE OF BEDFORD, K.G., President.
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Ricnarp H. Burne, Ese., M.A.
‘ ‘ P. Caatmers Mircnent, Ksa.,

pee oe ae ues AEA Dieee iE Deen

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AtrreD Henrace Cocks, Hse., J
M.A. W. R. Ogitvin-Grant, Ese.
F, G. Dawrrry Drewitr, Kse., || AvBerr Pam, Esa.
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Antony H. Wrinerietp, Ese.

2

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their names in the book at the entrance gate, and may introduce
Two Companions daily.

The Wire or Hussanp of a Fetitow can exercise these privileges
in the absence of the Fellow.

For the year 1913, and until further notice, Frtnows will
receive 40 undated Green Cards, available on any Sunday or week-
day up to the end of February of the year following the year of
issue, and 20 White Cards available on any week-day up to the
same date. Twenty of the Green Cards may be exchanged for a
book containing two Orders for each Sunday in the year, and the
twenty White Cards may be exchanged for a similar book of
Saturday Orders, Special children’s tickets will no longer be issued,
but the Green and White Cards will be perforated, and each half
will be valid for a Child under twelve years of age.

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

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

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

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

4

Any Fetxow who intends to be absent from the United Kingdom
duririg the space of at least one year, may, upon giving to the
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“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 Frttow, 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.,
December, 1912.
MEETINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON
FOR

SCIENTIFIC BUSINESS,

1913.
TursDAY, FEBRUARY ...... 4 and 18
Es IM ASRORLS "thad2 Uae 4, 8
iy ANI Poin 8 deo Cr py 22
mee NPA nb. lara cat nese g Ores 20
se eh Sel RUINGES ova yer wea US 3
ee OCnoBERiiy sonic 28
» NGVEMBER.... dE vee 259

The Chair will be taken at half-past Eight o'clock in the Evening
precisely.

ZOOLOGICAL SOCIETY, OF LONDON,

LIST OF PUBLICATIONS.

THE 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, indexed 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. 1s 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 80s.
(which includes cost of delivery), payable on the Ist. of July
in each year; but this privilege is forfeited unless the
subscription be paid before the Ist. of December following.

The following is a complete list of the publications of the
Society already issued.

TRANSACTIONS * OF THE ZOOLOGICAL SOCIETY OF LONDON.

4to. 19 vols. and Index. que Precworie

Vol. Ticontaiming 59 Plates)... (iess—3)) an aio mlommOn i. 4 elon Oi
3 TB a aL he 6 (1830-41) an. pe OmOm 5 6 6t
» TE. + 63, vee (1842-49) F. MomOmea ae AN EL Os
eps Tae ee a een arpa . (1851-62). ae OE Odor
AON 67 CISE2-66) ce em OREO)
at oo i D2) ess ‘ine (866-62) Feeble oman 15 0 0
ee CLs OE O).. 1S 12 6
A AVOUT ISS ey 18 82s navies QISR2 20ers eons 20
Ci ee EOE) eA Oca. 18 2 ©
aol Mee. Oasis 9D ee aCe) soe5 IO Or, rey 7
moles, WOlkks ILS, 9 dino oo bode 06 (icss—78) conn U7 Oo; 010 0
Vol. XI., containing 97 Plates.. (1880-85) .... 912 0.... 1216 O
Paes. Mee Giver eso (sls 00) goes od) Sle One 7 4 0
ae PANU, 59 OD” a lan CLSOIEO aie, tr OSes SI
Fy AEN 5 pen iee eo ain (USCE oy e one! (Oo Ge Or 2 7 OER
Es NOV 33 ieee oo (evel 901) oy Om On Oue 714 0
Cm VE; Pech pa 50 UICC) Ge, SS One 7 4 0
7 EONEEL., yo AL, 1 ee A908 S1 906) orb DES RG See marae ene
py ON VINE pce 40! Hap Mae (LOOT NOMI) ey x4 kOe een ame)
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. OG alerreh iy (O2MSp IAG) oo (ies IGUZ)) 5 OLS © - 140
OR oe 9 (2s MUL.) Coyne WON) 30 SO). 301 F0

PROCEEDINGS OF THE COMMITTEE OF SCIENCE AND

PROCEEDINGS OF THE ZOOLOGICAL SOCIETY OF LONDON.
(First Series.)

Part

”
”
”
”
”

CORRESPONDENCE OF THE ZOOLOGICAL
LONDON. 8vo. 2 vols. (Letterpress only).

Part I. 1830-81.

”

II. 1832.

1 vol. Syo.

33

e@oeee®eeeace

ee eeeee tore

8vo. 15 vols. (Letterpress only) and Index.

Price to Price to the

Fellows. ublie.

TI. 1833. 1 vol. 8vo. 4s. 6d. .. 6s.

TI TEM a Gs 54 Ce
Wan, JES, = yy) LG ay Ge
Thy, ICR we OGL ow Gs.

V. 1837. 4s. 6d. .. Gs.

VI. 1888. 5 4s. 6d. .. 6s.
POVTGISSO. = ) 486d sweet
MANA SAO) 12,0 Weds, Goss Get

8vo. 13 vols. aa Index.

Letterpress only.

Price to

‘ Fellows.
Part XVI. 1848. 1 vol. 8vo. 4s. 6d.
5s XVII. 1849, s As. 6d.

bs XVIII. 1850. 5 As. 6d.
a XIX. 1851. 3 4s. 6d.

. XX. 1852. 5 4s. 6d.

5 XXI, 1858. - 4s. 6d.

= XXII. 1854. is 4s, 6d.

» AXIIT. 1855. , 4s, 6d.

> XXIV. 1856. $s As. 6d.

9 XXV. 1857. As. 6d.

5 XX VI. 1858. 3 4s, 6d.

» AXVII. 1859. 3 4s. 6d.
XXVIII. 1860. 5 As. Gd.
Index 1848-1860. ds. 6d.

* In consequence of a re-arrangement of the stock of the ‘ ‘Transactions,’ the Society is
now able to offer for sale, at the reduced price of £80, sets of Vols. v.-xvi. inclusive, and
separate papers, of which a list can be supplied, at about one-fourth their published price.

Price to
Fellows.

As,

6d.

SOCIETY OF

Price to the
Public.

6s.F
6s.

Price to Price to the

Fellows. Public.
Part TX. 1841.1 vol. 8vo. 4s. 6d. .. 6s.F
5 X. 1842. 5 4s. 6d. .. 6s.
9 XI. 18438. 2 4s. 6d. .. 6s.
pt OSU lKeel e 4s. Gd. .. 6s.
» XIII. 1845. 5 4s. 6d. .. 6s.
y ALY. 1846. “5 4s. Gd. .. 68.+
XV. 1847. us 4s, 6d. .. 6s.f
Index 1830- 1847. 95 4s. 6d. .. 638.
(Second Series.)
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Price to the Price to Price to the
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GSA orc tart £1 0) 88 Sl eos
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Gs. i

t Out of print.

PROCEEDINGS OF THE SCIENTIFIC MEETINGS OF THE
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BRS GUURON Sicha cetersrute Cek CORR ONO ICRC 9s ee Sick ree oes. 9d. .... 458.7
US Zh os aie terare eee ronan nics. s 9s Lge er 5 Oa Oh cose 4s
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NS owe Micha ace stolees eee ere other Qs AC AAES Aen 6 36s 48s
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SOS RL Soins ete sree Bachar cramtieh rari erete ras en aici vera a temmin tes 36s . 48s
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* No perfect copies in stock. t Out of print.

PROCEEDINGS or THE GENERAL MEETINGS ror SCIENTIFIC
BUSINESS or tuzr ZOOLOGICAL SOCIETY OF LONDON.
Svo. 24 vols. and Index.

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MOL evolen gle matan tte wn Gea ataio seg ave 4. cualrece S-ciotesei etoae ema areas ISsraceo as:
ie op dlls i eae eas ie era eMC PCR Cow M5 GteualG oH cio 1SS.5. 8 onan se
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‘5 in beeen nom coe nomad Gans 6.0'.0.6 0b gd vat TBS, ie ince Mass
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LISTS OF THE ANIMALS IN THE SOCIETY’S GARDENS.

List of the Vertebrated Animals now or lately Living in the Gardens
of the Zoological Society of London. (Mighth Edition.) 8vo.
1883. Cloth, 4s. 6d.

List of the Vertebrated Animals now or lately Living in the Gardens
of the Zoological Society of London. (Ninth Edition.) 8vo.
1896. Cloth, 6s.

Catalogue of the Library of the Zoological Society of London
(Fifth Edition.) 8yvo. 1902. Cloth, 6s.

THE OFFICIAL ILLUSTRATED GARDEN GUIDE—10th Edition
—with (1) a Railway and Street Map, showing a direct
route to the “Zoo” from all parts of London and the Suburbs;
(2) a Plan of the Grounds, showing at a glance the location of
the animals ; (3) a short description of some of the principal
animals in the Collection (mow containing over 3000 spe-
cimens), together with 50 Photographic Illustrations and
Index. Price 6d. in Stiff Paper Cover, postage 14d., or in
Green Cloth Cover price 1s. 2d. post free.

P. CHALMERS MITCHELL,
Secretary.

Regent's Park, London, N.W.,,
December, 1912.

These publications may be obtained at the Socrery’s Orvrcn,
at Messrs, Longmans’ (Paternoster Row, E.C.), or through any
bookseller.

‘ZOOLOGICAL SOCIETY OF LONDON.

THE ZOOLOGICAL RECORD.

(ee object of the ZootogicaL Rucorp is to give, by means of an

annual Volume, complete lists of the Works and Publications
relating to Zoology in all its branches that have appeared during
the year preceding the issue of the Volume, together with full
information as to the points they deal with, arranged in such a
manner as to serve as an Index to the literature of Zoology in all
parts of the globe, and thus to form a repertory that will retain its
value for the Student in future years.

The ‘ Zoological Record’ having been amalgamated with the
International Catalogue of Scientific Literature, Zoology, Volumes
from 43 onwards can now be obtained only from Messrs. Harrison
& Sons, except when purchasing complete sets from the Zoological
Society.

Under the scheme of amalgamation, Fellows of the Society, and
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following.

The Society is able to supply complete sets of the Record on the
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Vols. 1 to 37, price £14 10s. net.

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Vol. 43 and onwards at 40s. each.

The prices for separate volumes are as follows :—

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The price of the ‘Zoological Record,’ Vol. 43 and subsequent volumes,
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InpEx Zootocicus. An alphabetical list of names of genera
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‘Zoological Record,’ 1880-1900; together with other names not
included in the ‘ Nomenclator Zoologicus’ of S. H. Scudder. Com-
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Record.’ London, 1902. Price to Fellows, 18s.; price to the
public, 20s., or if sold with a set of the ‘ Zoological Record,’ 10s.

Inpex Zoouoeicus, No. II. An alphabetical list of names of
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the ‘ Zoological Record,’ Vols. 38-47 inclusive (1901-1910), and
the Zoology volumes of the ‘ International Catalogue of Scientific
Literature, Annual Issues 1-10. Compiled (for the Zoological
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edited by Davip Sarr, M.A., F.R.S., Editor of the ‘Zoological
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Divisions of Vols. 39 to 42 of the ‘ Zoological Record’ can be
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of Messrs. Harrison & Sons, 46 St. Martin’s Lane, W.C.

ize Bo,

SEPARATE DIVISIONS OF THE ZOOLOGICAL RECORD.

Divisions of the ‘Zoological Record,’ Vols. 39-42, containing
the literature of the years 1902-1905, may be obtained separately
as follows :—

Ss
List of abbreviations of journals, etc. .. .. 2

Special Records, viz. :—
I. General Subjects ..
II. Mammalia
III. Aves nae tear SN ae
IV. Reptilia and Batrachia. .
V. Pisces
VI. Tunicata
VII. Mollusca
VIII. Brachiopoda ..
IX. Bryozoa
X. Crustacea
XI. Arachnida
XII. Myriopoda
XII. Insecta
XIV. Echinoderma
XY. Vermes ..
XVI. Coelenterata ..
XVII. Spongiz
XVIII. Protozoa

eBpnmrRereerepwpohwt ob wb

js
bo

LS) fo I= BS CH
SoS Ooageage oq eq Oooo q & © @® &

Index of new names of genera and subgenera.

INS

Divisions from Vol. 43 onwards are now supplied by Messrs.
Harrison & Sons, 46 St. Martin’s Lane, London, W.C.

P. CHALMERS MITCHELL,
Secretary. 5
; Recents Parx, Lonpon, N.W. ;

December, 1912.

ON BURCHELL’S ZEBRA. 757

41. The Local Races of Burchell’s Zebra. By Major J.
StTEVENSON-Hamiuton, C.M.Z.8., Game Warden of the
Transvaal.

[Received April 1, 1912: Read May 21, 1912.]

(Text-figures 102-106.)

INDEX.
Page
VeariatlOneen te ceeeeeeeee ee See cde neces senna TOT

Burchell’s Zebra (Lquus burchelli) south of the Zambesi is
usually divided by naturalists into several local races—wahlbergi,
transvaalensis, chapmanni, selousi. ‘These derive their subspecific
distinctions from certain assumed differences in their leg and body
striping as observed in museum specimens. Thus :—

“Tn wahlbergi the body stripes meet the ventral stripe
inferiorly, while the legs are more or less fully striped ; the shadow
stripes on the hind quarters are strongly developed and not much
narrower than the main stripes, which are narrower than the
intervening spaces, while the fetlocks and pasterns are devoid
of stripes and spots. A female zebra from the Transvaal
differing from the typical wahlbergi by the extension of the
shadow stripes to the neck, has been named LZ, 6. transvaalensis.
In £. b. chapmanni the shadow stripes have become faint and
narrow, the legs are marked to the hoofs, but the stripes on the
lower portions tend to break up into spots, and the inferior part
of the pasterns is not wholly black. ‘This race inhabits the
country between Damaraland and Matabeleland. The last repre-
sentative of the species with distinct shadow stripes is the
Mashonaland bonte quagga (. b. selousi), which differs from the
last in that the striping of the legs is complete right down to
the hoofs, the pasterns being striped on both sides, and their
lower portion, owing to the fusion of several stripes, wholly
black. The sides of the tail are also striped.” (Lydekker.)

Long observation of the herds of Burchell’s Zebra, running
in the Transvaal Game Reserves, has convinced me that all or
nearly all the above distinctions are found among them, frequently
in the same herd. I have often discussed the matter with
Dr. Gunning, of the Transvaal Museum, and have with him
examined skins and live specimens obtained from the Western
and Northern Transvaal. There is no doubt that any of these
might, from their markings, have come from the Eastern part of
the Province. Among the herds found in the latter occurs a
very great variety of striping. Some animals show heavy, wide,
and deeply tinted shadow stripes, while others display only the
slightest indications of them. Some are strongly ringed down
to the fetlocks, while others have no signs at all of any markings

Proc. Zoou. Soc.—1912, No. LI. 51

758 MAJOR J. STEVENSON-HAMILTON ON

below the knees and hocks. In some cases the body stripes are
continued right round the barrel to the ventral stripe, in others

Text-fig. 102.

Burchell’s Zebra six years old, from south of the Sabi River,
? bE) y
Eastern Transvaal.

they stop far short of it. The striping is usually carried right
round the buttocks, but in I should say 20 per cent. at least of the

BURCHELL’S ZEBRA, 759

animals in each herd this is not the case. In fact there appears
to be no special distinctive marking common to any locality or

Text-fig. 103.

Burchell’s Zebra, 3, ten years old, from south of the Sabi River,
Eastern Transvaal.

| 51*

760 MAJOR J. STEVENSON-HAMILTON ON

even herd. ‘The shadow stripes on the neck, said to be dis-
tinctive of Z. 6. transvaalensis, have been occasionally, but very

rarely, noticed in individuals, and I cannot conceive their presence

to be more than an individual eccentricity, occasionally perhaps
transmitted to offspring, but by no means constant.

Text-fig. 104.

Burchell’s Zebra, 9, eight years old, from south of the Sabi River,
Kastern Transvaal.

During the year 1911 I was asked to obtain a couple of skins
for the Transvaal Museum, and selected two stallions from the
same herd at a point a few miles south of the Sabi River (roughly
25 degrees 8. lat. by 31 H. long.). I chose them merely as being
typical of the darker and lighter forms of striping respectively,
neither of them being in any way exceptional. Some animals.
present displayed deeper striping than the specimen shown in
text-fig. 102, while others showed more white than that in text-
fig. 103. These figures are from photographs of the skins. Text-

BURCHELL’S ZEBRA. 761

fig. 102 is that of a stallion six years old, and text-fig. 103 is of
one about ten years of age. A little later I obtained the skins of
a mare and her foal killed at a spot less than ten miles from
where I had shot the two stallions. The photographs of these are
reproduced as text-figs. 104 and 105.

Text-fig. 105.

Burchell’s Zebra, , foal, trom south of the Sabi River, Eastern Transvaal.

In the first stallion (text-fig. 102) both the ordinary and the
shadow stripings are exceptionally strong, and the former is con-
tinued down and nearly all round the legs to the pasterns, which
are quite black. The body stripes meet the ventral line through-
out. The dorsal stripe is wide and very strongly marked. The
neck stripes are exceptionally deep, and set very close together.

In the second stallion (text-fig. 103) the shadow striping is not
nearly so well defined and the black stripes also are narrower
The body stripes do not extend all round the barrel, and there is
vractically no connection between them and the not very distinct

762 ON BURCHELL’S ZEBRA.

ventral stripe. The dorsal stripe is a very narrow black line, and
-there is a distinct gridiron pattern on the back. The legs are
very slightly striped ipalow. the knees and hocks, and there are only
a few black hairs on the pasterns.

In the case of the mare (text-fig. 104), while there is a well-
marked ventral and a wide dorsal stripe, and complete union
between the former and the barrel stripes, there is an entire
absence of any black markings from all the pasterns, and from
the fore legs below the knees.

Text-fig. 106.

Burchell’s Zebra, ¢ ; the same animal as that shown in text-fig. 103,
photographed just after death.

In the foal again (text-fig. 105), while the legs all bear faint
indications only of stripes, the pasterns are all quite black, and
there is some indication of a gridiron pattern on the back.

In none of the specimens is there the smallest indication of
any shadow striping on the neck. It will be noticed also that in
the younger stallion and the mare (text-figs. 102 & 104) the
frontal stripes on the face run straight from forehead to nose,
while in the older stallion (text-fig. 103) (and to some extent in

the foal (text-fig. 105), though it does not show up well in the
figure) the fr onal stripes cena to form concentric rings.

LXXXIV.

IP, 2 So WZ, Pi.

Bale & Danielsson, L*? imp.

IDO WMIDIODIDIO WIS, IBIS INI) ©).

ard, del.

G.M. Wood

ON A NEW CRUSYACEAN GENUS. 763

It would be interesting to know whether this absolute want of
uniformity found among Burchell’s Zebras in the Transvaal is
peculiar to this country, or whether it is the case throughout
Africa south of the Zambesi. Obviously it would be very
dangerous to attempt any classification from isolated museum
specimens obtained from the Tiansvaal at any rate.

Gunning proposes, I understand, to deal more fully in
the forthcoming‘ Annals’ of the Transvaal Museum with a matter
which I here only lightly touch upon.

On Dipteropeltis *,a New Genus of the Crustacean
Order Branchiura. By W. T. Catmay, D.8e., F.Z.8.f

[Received April 13, 1912: Read May 21, 1912.]

(Plate LX XXIV.2)

INDEX.

Page
SHSTPEKCLDUNE) OW” AHO OINOM OE ocboncag-oosecesecpnsendsnopnere Zee)
JBhelnolkorsny (ER ISHINSION)) © ssyaonnovosnonnseacansasssavansdonse | Cae}
Geographical Zoology ............ ee ee OS
Systematic: Dipter yaa! aap. tek re Mba eu e 2 763

The specimens described in this paper were presented to the
British Museum twenty years ago by Spencer Moore, Esq., by
whom they were collected in Southern Brazil. They were found
among a number of specimens of Dolops longicauda Heller bearing
the label “ Parasite on the gills and body of the fish known
all up the river as ‘Dorado’ from its golden colour. From
Corumba and neighbourhood, Matto Grosso.”

DIPTEROPELTIS HIRUNDOS. (PI. LX XXIV. figs. 1, 2.)
Calman, Abstract P. Z.S. 1912, p. 34 (May 28th).

Description of female :—The most striking feature of the species
is the form of the carapace, the lateral lobes of which are drawn
out into narrow lanceolate wings directed backwards and ex-
tending far beyond the tips of the long abdominal lobes. The
wings are fleshy in texture, traversed by numerous branching
blood-channels, but without spines or other armature on the
under surface.

The head is small, defined on each side from the carapace wings

* The complete account of the new form described in this paper appears here,
but since the name and a preliminary diagnosis were published in the ‘Abstract,’ it
1s distinguished by being underlined —Eprror.

t Published by permission of the Trustees of the British Museum.

{ For explanation of the Plate see p. 766.

§ For definition of the genus see p. 766.

764 DR. W. T. CALMAN ON A

by a shallow antero-lateral sinus from which two converging
grooves run in on the dorsal surface. These grooves unite into
one on each side, but the resulting groove dies out before reaching
the middle line. A pair of inter-ocular chitinous rods are well-
marked, closely approximated in front, diverging posteriorly, and
united by an indistinct articulation with a nearly parallel posterior
pair. The paired eyes are very small and close together near the
front margin. The unpaired eye could not be detected.

The free thoracic region is not distinctly segmented ; it is of
equal width throughout and more than three times as long as
wide.

The abdominal lobes are very long, narrowly lanceolate, and
cleft nearly to the base. No trace of furcal rami can be found.

The antennules and antenne (fig. 3) are very minute and nearly
hidden from view under the in-turned front margin of the head.
The antennules, in particular, are very easily overlooked; they
are not divided into segments but have a blunt lobe at the base
carrying a short apical tooth. The antenne are about the same
length, and consist of a stout basal part and a subglobular terminal
part separated from it by a constriction.

The mouth-parts form a prominent cone (fig. 4). The two large
and well-separated lobes of the upper lip are unarmed. The lower
lip bears a pair of slender conical papille (“maxille ” of Claus)
which project slightly from the opening of the mouth. The
mandibles are crescentic in form, with the concave edge very
finely serrated and the convex edge bearing a few teeth near the
tip.

ie front of the oral cone and continuous with it at the base is
a papilla, directed forward, having at its tip the opening of a duct.
This, no doubt, represents the sheath of the preoral ‘‘sting” of
Argulus, but no trace of the spine or sting itself could be detected
in the dissection of two specimens. At the base of this papilla
and of the mouth-cone are groups of very large cells, presumably
glandular, with deeply-staining nuclei.

The suckers (first maxillee, second maxille, or first maxillipeds
of various authors) are placed close together in front of the
mouth-cone, generally concealing altogether the preoral papilla.
In place of the usual radial supports, the whole of the membranous
border of the sucker is covered with discoidal scales (fig. 5).

The maxillipeds (fig. 6) (maxillee of some authors) are very short,
very stout at the base, and tapering rapidly to the tip. They are
each composed of five segments, of which the second and third
have numerous pectinate or branched spines on the ventral and
anterior surfaces. The terminal segment bears two minute claws
and a process which lies alongside of them.

The successive pairs of legs (figs. 7-10) are set wide apart from
each other along the sides of the thoracic region. ‘The rami are
in all cases shorter than the protopodite and carry only a few
short setee ; there is no flagellum on any of the legs. In the first

NEW CRUSTACEAN GENUS. 765

pair the endopod may be a good deal shorter than the exopod, or,
as in the specimen figured, nearly equal to it; both rami are un-
segmented. In the second pair also the rami are unsegmented,
and the endopod is from about one-half to two-thirds as long as
the exopod. In the third and fourth pairs the rami are subequal
and the endopod is divided into two subequal segments. The
basal lobe of the protopodite in the fourth pair is tongue-shaped,
with a slight protuberance at the base of its distal edge.

Dimensions of Holotype in millimetres.

Length of body to tip of abdominal lobes............ 20-0
Total length to tip of carapace wings ............... 26-0
Breadithyotg mea dire. .cttc en etech A. eackerette see brane I 2°
Isength: ofvthowacie VesiOme Wet shcnbk «st sstees «tek <a 8-0
Breadthwon bloracte reer. s20).22 tc a eos eee eee os 2°5
Breadthyotswanesaitebasetinl 00... isch. Hh. doe pt Alee 1:8
Greatest breadth of wing, at about 8 mm. from base 4:8
encthrotabdomunalblobes iin: 6) -s0-h.cal. bss. ack 68
Greatest breadth of abdominal lobes.................. 13
Length of antennules and antenne, about ......... 0-13
eng bhiohtonal(comes: teases sel). Jas ts dest eee ea 0:5
Greatest diameter of sucker ...................00e000 0 lal
enetingotesecomdglesy 5.2. scale thie nc scsi seen aslods 1:8
WDiamletersOneves pa DOM . 2 52c:dacd-cace sous Sashes ones. « 0-1
Distanceyapartromeyes, about... 2.5... -c2e-s2 e+ 65m: 0-45

Locality.—Corumba, on the Paraguay River, Matto Grosso,
Brazil. Four female specimens taken (in company with Dolops
dongicauda) on the fish known as “ Dorado” (probably a species
of Salminus).

Holotype.—¥emale, No, 92.10.24.2 in British Museum Register
of Crustacea.

A ffinities.—In having the so-called maxille or first maxillipeds
modified into suckers this species differs from the genus Dolops,
and in possessing antennules and a preoral papilla it differs from
Chonopeltis. From all the species hitherto referred to Argulus it
differs in (1) the remarkable form of the lateral wings of the
carapace ; (2) the length of the abdominal lobes and the absence
of furcal rami, in which characters it resembles some species of
Dolops ; (3) the entire absence of conspicuous spines or hooks on
the under side of the body and appendages, in which it resembles
Chonopeltis ; (4) the vestigial condition of antennules and
antenne ; (5) the absence of a spine or sting on the preoral
papilla; (6) the absence of the usual radial supports on the disc
of the suckers. Some of these characters, especially Nos. 1, 3,
and 6, are possibly not of great systematic importance, but
together they seem to show that the new form is less closely
related to any of the species included in the genus Argulus than

een ata

766 ON A NEW CRUSTACEAN GENUS.

these are to one another. I have proposed, therefore, to establish
for it a new genus, which may be defined as follows :—

DIPTEROPELTIS.
Calman, Abstract P. Z.S. 1912, p. 34 (May 28th).

Argulidee with the so-called maxille modified as suckers ; with
the preoral papilla present, but without a spine ; with antennules
and antenne very minute, imperfectly segmented ; without large
spines or hooks on under side of carapace, body, or appendages ;
without furcal rami on the abdominal lobes; with the lateral
wings of the carapace greatly elongated.

Genotype.—D. hirundo Calman.

It is to be noted that the single species of Chonopeltis,
C. inermis Thiele *, is known only from a single specimen (from
Lake Nyasa). It is not at all impossible, therefore, that the
antennules, if they are as small as in Dipteropeltis, may have
escaped observation, and it is even possible, though less probable,
that an unarmed preoral papilla may be present. Should this
prove to be the case it is doubtful whether, in spite of the great
difference in the form of the carapace, the separation of Diptero-
peltis from Chonopeltis could be maintained.

EXPLANATION OF PLATE LXXXIV.

(Lhe magnifications given are only approximate.)

Fig. 1. Dipteropeltis hirundo. Female (holotype), from the dorsal side. x 3.

2. Anterior part of body of same specimen from ventral side. The suckers.
have been drawn apart and slightly backwards. In the preserved
specimens they meet in the middle line and entirely conceal the preoral
papilla. x 6.

3. Antennule and antenna. x 450.

4. Tip of oral cone from antero-ventral aspect. x 100.

5. Part of membranous margin of one of the suckers. x 160.

6. Maxilliped. x 20.

7. Leg of first pair. x 20.

8. Leg of second pair. 20.

9. Leg of third pair. x 20.

10. Leg of fourth pair. x 20.

Figures 3-10 are drawn from one of the paratypes.

* Thiele, Zool. Anz. xxiii. p. 47 (1900); Mitth. Zool. Mus. Berlin, 1. p. 44,
figs. 110-116 (1904).

ON NEW LARVAL TREMATODES. 767

43. On two new Larval Trematodes from the Striped Snake
(Tropidonotus ordinatus sirtalis). By Wiii1aAm NIcott,
D.Se., F.Z.S., Lister Institute of Preventive Medicine,

London.
[Received April 9, 1912: Read May 21, 1912.]

(Text-figure 107.)

INDEX.
Page
Ethology: Two larval Trematodes in the mesenteric fat of
Striped Snake (Tropidonotus ordinatus sirtalis). Adult

form probably in a bird . 767
Geographical Zoology : North “America ; “Striped ‘Snake
infected with two larval Trematodes ........ 767

Systematic: Cercaria ordinata, sp. n., and Diplostomum
igi sp.n., from mcs in the mesentery of the Striped
Snake .. NOPE Ey ers eA cee MER Pn ne may OS

The occurrence of encysted larval parasites in snakes is evidence,
if such were wanting, that some snakes are eaten by other
animals. What is more important, the character of the parasites
may indicate what variety of animal is in the habit of eating the
snake in question. Conversely, the presence of any particular
species of adult parasite in an animal is almost always a sure
proot that such animal eats the snake in which the larval stage
is found. It is unfortunately in many cases a matter of difficulty
to diagnose the systematic characters of a larval parasite. In a
number of cases, however, it is possible to assign it to a definite
genus, rarely to a particular species.

The two cases to be dealt with here present a certain amount
of difficulty. The first larva is evidently a Distomate Trematode,
but beyond that it is impossible to go; the second larva is just
as obviously a Holostomid, and almost certainly belongs to the
genus Hemistomum. Such a diagnosis does not lead very far,
but it at least enables one to say that in all probability the
Striped Snake (Zropidonotus ordinatus, var. sirtalis) is eaten by
some bird, for adult Holostomata are known to occur only in
birds. This will possibly be confirmed by direct observation.

Both forms were met with together in each of three Striped
Snakes from North America, which died in the Society’s Gardens
on the 5th and 20th December, 1910, and on 10th March, 1911.
They occurred in enormous numbers in the mesenteric fat along
the whole length of the intestine. Hach was enclosed in a small
spherical or ovoid cyst with unusually thin and soft walls.
Unlike what is generally found in a Trematode cyst, the wall
gave the impression of being a thin membrane instead of the
more usual tough chitinous investment. On this account, not
only could the larve be extracted from the cysts without dif-.
ficulty, but when placed in water they escaped readily of their
own accord. When a piece of the cyst-infested mesentery was.

768 DR, WILLIAM NICOLL ON

suspended in water, a continuous shower of larve was observed to
fall to the bottom of the vessel.

The first form, which I name CERCARIA ORDINATA, Sp. n. (text-
fig. 107 a), was much more numerous than the other. It isa
typical tailless encysted cercaria, about °5 mm, (‘4-55 mm.) in
length and :2-:25 mm. in greatest breadth. In shape it is ovoid
and flattened dorso-ventrally. The entire surface of the body is
covered with minute regularly-arranged spines. The oral sucker
is almost terminal and has a diameter of ‘07 mm. (:06--08 mm.).

Text-fig. 107.

Se eS
SS ee

A. Cercaria ordinata, sp.n. Ventral view, x 150.

B. Diplostomum sirtale, sp.n. Ventral view, X 150.

The length is usually slightly greater than the breadth, and the
sucker has a somewhat characteristic funnel-shaped appearance.
The globular ventral sucker is situated rather in front of the
centre of the body, and has a diameter of 083 mm. (:(075—097mm.).
Its distance from the anterior end of the body is on an average
22 mm. (17-25 mm.). On the dorsal lip of the oral sucker are
the two symmetrical apertures of the cystogenous ducts. The
cystogenous glands are conspicuous structures, and consist of four

=r. 2

NEW LARVAL TREMATODES. 769

large cells situated in a transverse row immediately in front of
the ventral sucker. There are two pairs,a right and a left. The
ducts from each pair unite almost at once, and the united ducts
then pass forward in an irregular course; but just before they
reach the oral sucker each makes a characteristic twist, following
which there is a gradual increase in calibre until near the termi-
nation, when they contract slightly again. The cystogenous
glands have an irregular rectangular outline, and measure
"048x037 mm. In direct contact with the oral sucker is the
small muscular pharynx measuring °024 mm., which is continued
by a short, somewhat dilated cesophagus of the same length as
the pharynx. The intestinal bifurcation is about midway between
the two suckers. ‘The diverticula are simple, somewhat wide
tubes, which terminate not far behind the ventral sucker (7. e. a.
third of the distance from the sucker to the posterior end of the
body). The excretory vesicle is V-shaped. A common trunk is
practically absent, and the limbs extend forward to near the
terminations of the intestinal diverticula. The excretory tubules
are very fine; the main tube on each side extends forward to
near the oral sucker, where it turns back. The excretory aperture
appears to be slightly dorsal.

No trace of other organs could be made out, so that no accurate
idea can be obtained of the systematic position of this larva.
The peculiar configuration of the excretory vesicle may, in con-
junction with the shape of the alimentary canal, eventually lead
to the identification of its adult form, but at present, so far as I
am aware, there is no known adult Distome to which this larva
can be ascribed.

The second form, which I name DrpLosromum siRTALs, sp. n.
(text-fig. 107 B), differs markedly from the first. It is about the
same size, and occurs in somewhat similar cysts, but its shape and
colour are entirely different. The body of Cercaria ordinata is
light and transparent, whereas that of Diplostomum sirtale is
dark and almost opaque. The opacity is due to the presence of
innumerable small granules distributed throughout the whole
body. The shape is that of a typical Diplostomum larva, being
scoop-like with a short handle. The shape is due to the rolling
over of the postero-lateral margins of the body. In life, however,
these margins are capable of more or less eversion, so that on
occasion the body may appear almost flat. The dimensions of
this larva are -48—55 x -28-"32 mm., the short stumpy tail being
°06 mm. long.

The oral sucker measures (039 mm. in diameter; the ventral
sucker ‘042-045 mm. The latter is situated a little in front
of the middle of the body, :25--28 mm. from the anterior end.
Midway between it and the posterior end of the body occurs
the characteristic Holostomid fixing disc, which appears as
a transparent disc about the same size as the ventral sucker.
Of the internal organs only the alimentary canal and excretory
vesicle were visible. The former comprises a pharynx contiguous

770 SIR CHARLES ELIOT ON A RARE NUDIBRANCH.

with the oral sucker and measuring :024x:020 mm. This is
followed by an cesophagus about twice as long as the pharynx.
The intestinal bifurcation takes place rather nearer the ventral
sucker than the oral sucker, and the simple diverticula extend
a little beyond the posterior border of the fixing dise. The
excretory vesicle consists of a wedge-shaped sac, which extends
forward as far as the ends of the intestinal diverticula.

This is in all probability the larval stage of some species of
Hemistomum, parasitic in a bird.

44, A Note on the rare British Nudibranch Hancochia
cudactylota Gosse. By Sir Cuartes Eniot, K.C.M.G.,
C.B., F.Z.S.

[Received April 18, 1912: Read May 21, 1912. |
(Plate LXXXV.)

See Gosse, On Hancockia eudactylota, Ann. Mag. Nat. Hist. ser.
4, xx. 1877, pp. 316-318; Gamble, On two rare British Nudi-
branchs, Lomanotus genet and Hancockia eudactylota, ib. ser. 6, 1x.
1892, pp. 378-885; Trinchese, Ricerche anatom. sul genere
Govia ( = Hancockia), Mem. della R. Accad. delle Sei. dell’
Istituto di Bologna, ser. 5, vii. pp. 183-191, 1886; Eliot, Sup-
plement to Alder and Hancock’s Monograph of the British
Nudibranchiate Mollusca, Ray Society, 1910, pp. 17, 72,
118-120, 163.

No coloured figure of this rare British Nudibranch has yet
been published so far as Iam aware. I endeavoured to include
one in my Supplement to Alder & Hancock’s Monograph, but
no specimen of the animal could be obtained before the time fixed
for publication. Shortly afterwards a single individual was
captured at Plymouth and drawn by a local artist under the
supervision of the naturalists who were working in the Laboratory
of the Marine Biological Association. I have not seen the animal
alive, but these drawings agree with what is known of its struc-
ture, and I have no doubt that they faithfully reproduce its
appearance.

For an account of the genus and species see the references to
my Supplement given above.

EXPLANATION OF PLATE LXXXYV.

Hancockia eudactylota.

Dorsal view of animal.
Ventral view of animal.
Side view of head.

A rhinophore.

Fig.

SI OUR go No

Different views of lateral processes.

ONS a ee

a ee

PLoS, TOG, Pil, LX XX XW.

London Stereoscopic Co. imp.

HANCOCKIA EUDACTYLOTA.

ash WON, Jl, WCW,

London Stereoseopiec Co imp.

GIRAFEA GCAMELOFARDALIS ST HORNITCGRO En.

ON THE NORTH RHODESIAN GIRAFFE, CHL

45, The North Rhodesian Giraffe.
By R. LypuxkKer, F.R.S., F.Z.S.*

[Received April 25, 1912: Read June 4, 1912.]

(Plate LXX XVI. +)

INDEX.

IR heoyoKeevey, Caieninr> Olt’ \ sosdsdodnbasduaabaseoedd sanders vos ccd eeslves 771
Giraffa camelopardalis thornicrofti, Description of ... 771

On the 20th of October, 1910, the British Museum (Nat. Hist.)
received from Mr. H. Thornicroft, Native Commissioner, Petauke,
North-east Rhodesia, the skin, skull, and limb-bones of an adult
male Giraffe shot by himself in that district. Mr. Thornicroft
had previously called on me in London, and expressed his willing-
ness to shoot and present to the Museum a Giraffe from the
single herd in this part of Rhodesia, if the necessary permit could
be obtained. This was in due course procured, and was followed,
after an interval, by the arrival of the above-mentioned skin and
bones.

The skin was soon afterwads set up by Rowland Ward Ltd.,
and the mounted specimen placed in the big case at the head of
the staircase leading to the Hast Corridor of the Museum, along-
side the male and female of the East African Girajfa camelo-
pardalis rothschildi. As it is mounted with the neck bent, it is
difficult to ascertain the exact height, but I estimate this at close
on 18 feet, or possibly rather more.

When the specimen was installed in its case, it became essential
that it should receive a distinctive name; and I accordingly
communicated the following preliminary note to ‘ Nature’ + :—

“This Giraffe is characterised by the low and conical frontal
horn, the grey colour and scattered spotting of the sides of the
face, the chestnut-brown forehead, deepening into black on the
tips of the horns, the absence of a distinctly stellate pattern in
the neck and body spots, which are light brown on a yellowish
fawn ground, and the uniformly tawny colour of the lower portion
of the limbs, This Giraffe, which I propose to call Giraffa
camelopardalis thornicrofti, appears to be related to the Kiliman-
jaro G. c. tippelskirchi, but differs by the more compact frontal
horn, the brown, in place of grey, forehead, and the uniformly
fawn lower part of the legs, the latter being whitish in adult
bulls [of tippelskirchi|, but fawn and spotted in cows and young
bulls.”

The last statement rests on the authority of Messrs. M. de

* Published by permission of the Trustees of the British Museum.
+ For explanation of the Plate see p. 773.
~ Vol. Ixxxvii. p. 484 (1911).

172 ON THE NORTH RHODESIAN GIRAFFE.

Rothschild and H. Neuville*, who state that in the East African
Giraffe which they describe as rothschildi, but which—despite the
locality whence it is stated to come—is certainly tippelskirchi,
these age and sex differences are observable. I have, however,
doubts whether they hold good in all cases; and it is still possible,
in spite of what I have previously written, that there may be one
form (schillingsi) in which the shanks of adult bulls are white and
another (tippelskirchi) in which they are fawn and spotted 7, and
further, that these two types may intergrade.

That the nearest relative of the Rhodesian Giraffe is G. c. tip-
pelskirchi, may be considered certain. Of the latter I have had
for comparison the mounted head and neck of an adult male, a
mounted immature female, and the mounted head and neck of
a calf, as well as,a coloured plate in Messrs. de Rothschild and
Neuville’s memoir f.

Elaborating to a certain extent the foregoing brief diagnosis,
attention may be directed to the fact that tippelskirchi and
thornicrofti agree (and thereby differ markedly from rothschildi)
in having the triangular space between the eye and the nostril
devoid of spots. In the adult male of tippelskirchi, however, the
zround-colour of the whole head is dirty greyish white, whereas
in thornicrofti the forehead is chestnut or umber-brown, deepening
into black at the tips of the horns, which are grey in the
Kilimanjaro race.

In the Rhodesian Giraffe the spots on the region behind the
eye and the side of the lower jaw are very faintly marked, and
blackish grey in colour; whereas in the Kilimanjaro bull they
are larger, more numerous, and chocolate-brown in colour, being
deeper in tint than the neck-spots (this feature being also shown
in the immature female and the calf).

In thornicrofti the spots on the neck are burnt-umber in colour
and markedly elongated in form, with their terminal ends jagged. .
There are about eight of them in the longitudinal row which
starts immediately in advance of the point of the shoulder. In
tippelskirchi they are more numercus (ten or eleven in what
appears to be the corresponding row), less elongated, and much
more irregular in shape.

Compared with the young cow teppelskirchi, the spots on the
body of thornicrofti are less numerous, more especially on the
hind-quarters, while many of them are more deeply incised on one
side, although they are less jagged in general contour. The
spotting on the inner side of the thighs and of the upper part
of the fore-legs is also much less pronounced. In the original
description (which was drawn up when the specimen was in the
basement of the Museum) it is stated that the shanks of the
legs are uniformly fawn, but, as a matter of fact, they are

* Ann. Sci. Nat., Zool. ser. 9, vol. xiii. pp. 124, 129 (1911).
+ See Proc. Zool. Soc. 1904, vol. i. p. 219.
f Op. cit. pl. ii. fig. 1, lettered G. ¢. rothschildi.

ON ANTLER-GROWTH IN THE CERVIDZ. CUS

rufous-fawn with very faint traces of spotting nearly down to
the fetlocks; while from the latter to the hoofs they are dirty
greyish white. ‘

The foregoing evidence clearly establishes the right of the
North Rhodesian Giraffe to rank as a distinct local race; and if
it be true that the one herd is completely isolated, there is
probably no intergradation with the Kilimanjaro race.

EXPLANATION OF PLATE LXXXVI.
Adult bull of Giraffa camelopardaiis thornicrofti.

46. On Antler-Growth in the Cervide, with special reference
to Elaphurus and Odocoileus (Doreelaphus). By
R. I. Pococn, F.R.S., F.L.S., F.Z.8., Superintendent
of the Gardens and Curator of Mammals.
[Received and Read June 4, 1912.]

(Text-figures 108-112.)

InDEX.
Page
ClassmiGanmonne Gi ne Olnyiles oogrccssecte nda nconne oan ononseavsaraene 773
Mode of growth of Antlers in typical Old World Deer ...... 775
Interpretation of the Antlers of Hlaphurus davidianus ...... UTe
Interpretation of the Antlers of Odocoileus sp. incert. ...... 780

Date of Antler-change in American and Old World Deer ... 783 (Note)

Introduction.

Most, if not all, the attempts that have hitherto been made to
understand the antlers of Deer and arrive at correct conclusions
regarding the homology of the tines have been based upon com-
parisons between the fully formed antlers of different species.
This, in my opinion, is the reason why there has been failure in
some cases to detect homologies which study of the growth of
individual antlers reveals.

The importance of this question depends upon the circumstance
that twenty years ago Mr. Gordon Cameron * proposed a classi-
fication of the Cervide, based upon the antlers, as a substitute
for the classification, founded upon the skeletal structure of the
fore feet, which Sir Victor Brooke had suggested 7.

To make clear the purpose of the present paper, it is necessary
to summarise briefly the rival classifications put forward by these
two authors. Sir Victor Brooke divided the Cervide into two

* «The Field,’ 1892, pp. 265, 703, 741, 860.
+ P.Z.S. 1878, pp. 883-928.

Proc. Zoou. Soc.—1912, No. LIT.

or
i)

14 MR. R. I. POCOCK ON

sections. The first, which he called Telemetacarpi (Telemeta-
carpalia) because the distal ends of the lateral metacarpals
persist, comprises the Roe (Capreolus), the Chinese Water-Deer
(Hydropotes), the Reindeer (Rangifer), the Elk (Alce), and all the
exclusively American deer with the single exception of the typical
Wapiti (Cervus canadensis); the second, called Plesiometacarpi
(Plesiometacarpalia) because the proximal ends of the lateral
metacarpals are usually present, whereas their distal ends are
suppressed, comprises all the deer of the Old World, except
the four genera mentioned above, but none of those of the
New World apart from the Wapiti. Amongst the Old World
forms the most important species for’ the moment figuring
amongst the Plesiometacarpalia is Pére David’s Chinese Deer
(Hlaphurus davidianus).

Mr. Cameron’s classification was widely different. Dismissing
as unimportant the character relied upon by Brooke, he divided
the Cervide into three sections: one for the Reindeer with
antlers in both sexes, the second for the Elk with laterally
extended antlers, the third for the remaining species with antlers
restricted to the male and erect or suberect. This third section,
which alone concerns us now, was subdivided into two categories
of species, one comprising those in which the antlers consist, as
in the typical Old World deer and the Wapiti, of a ‘“ brow-tine”
and a “beam,” to use Gordon Cameron’s terminology, and the
other those in which the antler has, as he thinks, no brow-tine
but consists of a ‘forked beam,” as in all typical American deer
(except the Wapiti) and in the Roe and Pere David’s Deer
amongst the Old-World species.

Now with regard to the affinities of the species composing
Cameron’s third division, there is only one point in which there
is complete divergence between him and Brooke. This concerns
Pére David’s Deer, a species classified by Brooke with the Red
Deer, Sambar, and other Hlaphine stags, and by Cameron with
the American forms allied to the Virginian and Mule Deer, the
correct name of which seems to be Odocoileus *.

So far as I can see, the only @ priori objection to be raised
against Mr. Cameron’s system, if we accept his premises, is that.
it is based upon a secondary sexual character. But although it.
cannot be justifiably consigned to oblivion on that account, it may
be doubted if it would ever have come into sufficient prominence
for serious discussion had it not been for the unqualified acceptance
accorded it by Mr. Lydekker. However that may be, it is clear
that if Mr. Cameron’s assumption that there is a fundamental
difference in structure between the antlers of the groups of deer
mentioned above is wrong, his classification, based on that claim,
goes by the board.

In the following pages J shall endeavour to show that his
classification is untenable, because a study of the seasonal growth

* Dorcelaphus and Cariacus are better known but superseded terms.

ANTLER-GROWTH IN THE CERVIDA. 775

of an antler of Pére David's deer and of an American deer, allied
to the Virginian, proves that the homologue of the brow-tine of
the Elaphine stage is present in both—a conclusion which is by
no means evident from an examination of the fully-formed
antlers.

Antler-Growth in typical Old-World Deer.

In the Zoological Gardens I have repeatedly watched, year
after year, the growth of the antlers of deer belonging to the
Elaphine, Sikine, and Rusine ty es without finding any variation

Text-fig. 108. /

b
---2
Wr MW! . SS
C 3 wn
BD

Early growth-stages of Antlers of some Old-World Deer.

A & B. Successive stages observed in Cervus hanglu. C. Cervus canadensis.
D, Rusa aristotelis.

a, the anterior branch or “ brow-tine” ; p, the posterior branch or “ beam ”
b, the rudiment of the bez-tine arising from the posterior branch.

of moment in the method of their development. The antler
starts as an undivided bud. This bud then shows signs of

division into two buds, an anterior and a posterior. These buds
52*

776 MR. R. I. POCOCK ON

gradually, and with nearly equal rapidity, increase in length, the
anterior growing forwards and the posterior backwards. In the
Sambar (Rusa, text-fig. 108, D) and some other species they show
a marked inclination upwards; so that at one stage the antler may
be likened somewhat to a short-stalked Y, and at this or even at a
later stage in deer like the Sambar (/twsa) and others which have
no “ bez ”-tine, the antler may be indifferently described as an
‘‘unbranched beam with a brow-tine” or as a ‘ forked beam” or
as a biramous antler. The anterior and posterior branches some-
times, as in Cervus eldi, grow at approximately the same speed
until the anterior has almost attained its limit; but usually the
growth of the posterior tine is from the first more rapid. However
that may be, the equivalence of the two branches in the early
stages is plain enough ; but afterwards this becomes less and less
evident as the posterior branch continues to lengthen and
develops its accessory tines.

Text-fig. 109.

p
P a B a
DEIN yn ny

aa
HN ay

pine E
D

Five stages (4 to EZ) in the growth of an antler of Rucervus duvaucelli.

a, anterior branch or “brow-tine” ; p, posterior branch or “ beam.”

These facts are shown in the annexed figure (text-fig. 109),
representing five stages in the growth of an antler of a specimen of
the Swamp Deer or Barasingha (Rucervus duvaucelli). These were

ohare,

ANTLER-GROWTH IN THE CERVIDA, ah

sketched on May 13, 16, 22, June 6 and 12. Similar stages may
be observed in other typical deer of the Old World*. In the
Elaphine stags, however, which normally grow a “ bez”-tine,
the biramous stage is early complicated by the appearance of the
bud of this tine. Now this tine has been regarded as a dupli-
cation of the brow-tine; and in Max Weber’st diagram showing
suggested homologies of the tines in certain deer the brow- and
bez-tines are tinted alike, suggesting his adoption of this view.
Nevertheless I believe it to be quite incorrect, for in all cases
where I have watched its origin, the bud of the “bez ”-tine
arises, not from the brow-tine at all, but from the “beam.” It
is, in fact, the basal or proximal tine of the posterior branch of
the antler. This is illustrated in text-fig. 108, A—C, showing the
early stages of the growing antler of the Hangul (Cervus hanglu)
and of the Wapiti (Cervus canadensis).

Antler-Growth i Pere David's Deer (Elaphurus davidianus).

There is no stag whose systematic position has troubled
zoologists so much as Hlaphurus. On the one side are those,
like Dr. Gray, Mr. Cameron, and Mr. Lydekker, who, relying
upon the structure of the antlers of the adult, placed the genus
with the American deer. On the other side are those, like
Sir Victor Brooke, Flower, Max Weber, and others, who, adopting
the skeleton of the foot as a basis, classified it with the typical
Old- World species.

The antlers of this stag have often been figured and described,
and a good idea of their form in the adult may be gathered from
text-fig. 110, C, and text-fig. 111, 7. They typically consist of a
comparatively long basal portion from which two branches arise :
one long, slender, simple or divided, projects backwards parallel,
or nearly so, with the animal’s back; the other stout, erect, or
curved slightly forwards, terminates in a pair of strong tines.

At first sight, these antlers appear to have-no trace of a brow-
tine. This was evidently Sir Victor Brooke’s opinion, and it was
adopted by Mr. Cameron and Mr. Lydekker, who, on the strength
of this belief, boldly claimed that this stag belonged to the same
group as the American deer, also held to have no brow-tine,
despite the resemblances in other respects pointed out by Brooke
between Hlaphurus and the typical Cervide of the Old World.
Prof. Garrod was more cautious, and frankly gave up the attempt
to interpret the antlers of Hlaphurus when he remarked that they
‘‘are at present quite beyond my comprehension.”

This, then, was the state of the case when my researches on
the specialised cutaneous glands of Ruminants { showed that the

* Mr. J. G. Millais (‘Mammals of Great Britain and Ireland,’ iii. plate facing
p. 140, 1906) has published a series of figures of antler-growth in the Fallow Deer
(Dama) illustrating precisely the same phenomenon.

+ Die Saug. p. 667, 1904.

t P. Z.S. 1910, p. 840.

778 MR. R. I, POCOCK ON

absence of interdigital glands on the feet and the smoothness of the
integument between the hoofs in Hlaphurus corroborated Sir Victor
Brooke’s views as to the relationship between this animal and sucn
Old World deer as Rusa, Rucervus, and Cervus, and weakened to

a corresponding degree the claim for affinity between it and the

Text-fig. 110.

Diagram of the Antlers of four genera of Cervide, to illustrate the homologies
established in this paper.

A. Cervus. B. Rusa. C. Elaphurus. D.. Odocoileus.

@, anterior and p, posterior branch. In 4 and B the anterior branch is called the
“brow-time” and the posterior branch the “beam.” B is somewhat inter-
mediate between A and C. They differ collectively from D in having the
anterior branch well developed. In D it is small and concealed behind the
highly developed posterior branch.

Telemetacarpal species, in all the members of which examined by
me, belonging to the genera Mazama, Odocoileus (Dorcelaphus),
Capreolus, Rangifer, and Alee, the skin between the hoofs is

ANTLER-GROWTH IN THE CERVIDA. 779

thickly hairy and, in all but Alce, a large pouch-like inter digital
gland is present at least in the hind foot.

Text-fig. 111.

A

WN aes <td
G F

Tv
H

Nine stages (4 to Z) in the growth of an Antler of Elaphur us davidianus, showing
that the branches marked a and p correspond precisely in origin with the
brow-tine and the beam of other genera of Old World Deer. Compare D and H
with fig. 108, D.

(From sketches made at Woburn and kindly supplied by ord Tavistock.)

780 MR. R. I. POCOCK ON

Thinking, for these reasons, that there must be some flaw in
the claim that the antlers of Hlaphurus differ fundamentally from
those of, say, Rusa, I suggested the following homologies :—In
Rusa the antlers have a short base, a short undivided anterior
branch or brow-tine, and a large divided posterior branch or
beam ; in Hlaphurus they have a longer base, a very large divided
and more erect anterior branch or brow-tine, and a correspondingly
reduced, comparatively slender, divided or undivided posterior
branch or beam.

This, however, was a mere guess, which I was unable to.
substantiate by any evidence of much value. Believing, how-
ever, that the growth of the antlers in Hlaphurus would finally
settle the question one way or the other, I asked Lord Tavistock
if he would kindly observe the process for me on one of the stags.
at Woburn. This he was good enough to do, and sent me in
addition the series of sketches reproduced in text-figure 111.

These sketches show, in my opinion, that my guess was, as I
expected, correct. In the first three stages the antler is little
more than an excrescence dividing into an anterior and a posterior
bud. In the fourth stage the base is beginning to lengthen, the
anterior bud to grow upwards, and the posterior bud nearly
straight backwards. This process continues during the succeeding
stages, the anterior branch gradually taking the lead in size and
importance, and becoming divided distally into two tines. I can
see no escape from the conclusion that the anterior and posterior
buds of the very young antler in this stag are the homologues of
the corresponding buds in the young antler of the Barasingha
(Rucervus duvaucelli) shown in text-fig. 109. That being so,
it is clear that the anterior branch of the antler of Hlaphurus is
homologous with the “ brow-tine” and the posterior branch with
the ‘beam ” of the antler in the Red Deer, Sambar, Barasingha,
and other deer characteristic of the Old World. The differences.
between them are mainly a matter of size and direction of growth ;
that is to say, they are differences of degree and not of kind*.

Antler-Growth in a Species of Odocoileus.

Writing of the antlers of the typical American Deer, Mr.
Lydekker said +:—‘ A large amount of misconception has arisen
with regard to the structure of the antlers of this group. In
1872 the late Dr. Gray rightly termed the single upright prong
arising from the inner side of the lower part of the beam of the
antlers of the Virginian Deer the ‘subbasal snag’; but this snag

* The subdivision of the anterior branch of the antler in Hlaphurus is, of course,
no argument against it being the homologue of the “ brow-tine,”’ for the Jatter not
infrequently, though abnormally, produces an additional snag in Elaphine and allied
eroups of deer. In some species indeed, as in the Irish Elk and Cervus eldi, it is.
commonly and normally provided with supplementary processes.

+ ‘ Deer of All Lands,’ p. 246.

ANTLER-GROWTH IN THE CERVIDA. 781

Sir Victor Brooke incorrectly identified with the brow-tine of
the typical Old World deer. This error has been pointed out by
Mr. A. Gordon Cameron, [who stated that] these characteristic
tines have nothing in common with the true brows of Old World
types, and rise vertically from the inner side of the beam between
the coronet and the main furcation, usually converging at the
apex. They are subject, in common with the antlers that produce.
them, to all kinds of eccentricities; are frequently forked or sub-
palmate.”

Mr. Lydekker writes as if Mr. Cameron’s dictwm settled the
question at issue; but it does not appear to me that much weight.
can be attached to the reasons adduced by the latter for his
dogmatic denial of the truth of Sir Victor Brooke’s interpretation
of what Gray called the ‘“subbasal snag” in the Virginian deer.
Except that the tine in question is situated on the inner side of
the antler, there is no great difference between it and the brow-
tine of the Old World stags, which is highly variable in direction,
as a comparison between the antlers of, e. g., Cervus affinis and
Rusa aristotelis will show. Not less does it vary in size and
structure even in nearly allied forms, as is testified by Dama
dama, where it is large, by Dama mesopotanvica, where it is
sometimes almost suppressed, and by the Ivish Elk, believed to
be a Damine stag, where it may be palmated and branched.

The question to be settled, then, is this :— Does the position of
this tine on the inner side of the antler in the Virginian deer
preclude its being the homologue of the brow-tine situated on the
front of the antler in the Old World deer? Study of the growth
of the antler justifies, in my opinion, a negative answer to this.
question and shows that Sir Victor Brooke’s opinion was correct.

Karly last year the Society received from the northern part of
South America a male specimen of Odocoilews, which I cannot
determine accurately. It is smaller and browner than a Venezuela
specimen identified as O. savannarum, but is otherwise very like
it. Its antlers are short, with the beam curved forwards in the
upper portion and ending in two tines, an anterior and a posterior 3.
while on the inner side, near the base, arises the so-called ‘“ sub-
basal snag.”

The growth of these antlers was very instructive. They started
as a simple exerescence, which soon began to divide into an anterior
and a posterior bud, the only difference between the antlers at
this stage and those of a typical Old World deer being that the
anterior bud was slightly internal and projected a little inwards
as well as forwards. Nevertheless the two buds were perfectly
visible in profile view. The appearance of the antler at this
stage is shown in text-fig. 112, 4, takenon May 12th. Four more-
stages of the growth are represented in the following figures, b—/,
taken respectively on May 22nd, May 30th, June 6th, and
June 17th, which show very markedly the gradual assumption of
an apparently more internal position by the anterior branch, its.

782 MR. R. I. POCOCK ON

Text-fig. 112.

Wy
\ SN

\S
\
\y
DY)

A

> Pe

—_!ll,

Five stages (4 to #) in the growth of an Antler of an American Deer (Odocoileus
sp. incert.), showing that the “subbasal snag” (a) and the “forked beam ”
(p) were respectively the homologues of the ‘‘ brow-tine ” and the “beam ” of
the typical Old World Deer. Compare B with fig. 108, D.

ANTLER-GROWTH IN THE CERVIDA. 783

point of attachment to the posterior branch being completely
concealed from the external aspect in’ the last three stages *.

In view of these facts, I do not think it can be doubted that
the anterior bud which develops into the ‘“subbasal snag” in
Odocoileus is the homologue of the anterior bud which forms
the brow-tine in Cervus. In that case the “ subbasal snag ” and
the ‘‘brow-tine” are homologous structures passing under different
names, and to state that Odocoilews has no brow-tine is merely
playing with terminology.

If this interpretation of the structure of the antlers in Hlaphurus
and in the species of Odocoilews above referred to be, as I believe,
correct, it shows that these two genera are widely divergent in the
very point upon which relationship between them has been claimed
to exist, and that the likeness, such as it is, between the antlers
of Hlaphurus and of the Mule Deer (O. hemionus), for instance,
which has the so-called forked antlers without a brow-tine or with
the merest vestige of it, is purely a question of parallelism in
development ; that is to say, it has been brought about by growth
and modification of fundamentally different parts of the antler.
In the Mule Deer the anterior branch or brow-tine is to all
intents and purposes suppressed, practically the whole antler
being composed of the posterior branch or “‘ beam,”’ which is
highly developed and heavily tined. In “laphurus, on the
contrary, the principal part of the antler is composed of the
anterior branch or ‘“ brow-tine,” which attains a large size and is
divided into two prongs, while the posterior branch or beam
remains comparatively small and slender and projects straight
backwards as a long often undivided prong.

* In connection with the date of antler-change in this Stag, attention may be
directed to its approximate coincidence with that of the typical elaphine deer of the
Old World; that is to say, the antlers were in the velvet during the summer months
and functional during the autumn and winter. ‘Khey were shed in the early spring
and at the time of writing (July 3) the new antlers are nearly full-sized though stall
in the velvet, exactly as in our Wapiti, Red Deer, Japanese Deer, and other Old
World species. The same is true of a specimen ‘ot Odocoileus americanus. On
the other hand an example of Mazama bricenii which shed in April 1908, and again
in April 1909, did not repeat the process till May 1911. He then carried a pair of
antlers for 25 months; and those that started to grow in May 1911 are still on his
head. Thus Dr. Scharff (Distribution and Origin of Life in America,’ p. 111) is
mistaken in saying that the antler-change in American deer takes place at a quite
different time of year from that of Old World deer. It is well known too that the
time of antler-change at all events in some tropical Old World deer is highly variable
within specific limits. For instance, one example of CO. dwvaucelli in the Gardens
regularly carries his antlers till about the end of May, while another of the same
species has antlers at least half their full size at that time.

’

ie
784 MISS HELEN PIXELL ON

47. Polycheeta from the Pacific Coast of North America.—
Part I. Serputipm, with a Revised Table of Classi-
fication of the Genus Spirorbis. By Hentmn L. M.
PIxELL, B.Sc., F.Z.8., Demonstrator of Zoology and
Reid Fellow, Bedford College, University of London.

[Received May 6, 1912: Read June 4, 1912. ]
(Plates LXXXVIL-LXXXIX.*)

Systematic: InpDEx. Page
ZY TOMASO GODS, Dy Ws egacocnvcaccracauerave ooosbecceess Msi
JEROME FNCU IG, NOs 1s aghcaonanccadeovee wasaeectos pease Usts)
Classitication\ot Spiro7bisee.. seen en oe
JETRO COSMO) SURI My oo agecocgrs avosovecesvonsavecca BE
Spirorbis ambilateralis, sp. N...............00-....... 798
Sb PUCCMIOSUIS; De WS aga coonshtsy so ssousnbonbosacccebacpacesoo TOO
SGOCUCUSy S105: Ue ice, comsddane aan waddle daus boa dom cpeeseetosocboc: me OO)

General Characteristics of the Family Serpulide.
1. Tube calcareous, nearly always attached to rocks or other
substratum for some part of its length.
2. Generally one or more branchiew on dorsal side terminated
by an operculum.
3. Thorax, generally provided with a thoracic membrane,

.
{

representing the fused cirri and having 3-9 (usually 7) ;
segments. i
4. Gland shields in thorax only. i

Genus Serputa Linné (12) 1767, Philippi (21) 1844.
Generic characteristics fT :—
1. Collar sete of bayonet-shape, with spines at base of blade.
2. Operculum funnel-shaped, with numerous radii ending in ;
serrations on margin. '
3. Uncini with only a few large teeth.

1. SeRPULA CoLUMBIANA Johnson (9), 1901. (Pl. LXXXVII.
fig. 1.)
Serpula splendens Bush (8), 1905.
Serpula columbiana Moore (19), 1909.
Specific characteristics :—
1. Anterior abdominal sete with flaring fringed ends, short
and deeply embedded, posteriorly replaced by small fascicles.
of very long stiff spines.

1 ge ee sit

* For explanation of the Plates see p. 805.

+ An attempt is here made to summarise briefly the generic and _ specific
characteristics in every case. Such a procedure has not previously been adopted, so
far as I know, and it will, no doubt, in some cases be necessary at some future time
to modify such characteristics, but in the present confused state of our systematic
knowledge of the Serpulids, this seems to be a course likely to eliminate some of the
difficulties.

IY So WSN 2 LO.

yy

yh IssI IDI,

29>:
Za » mn,

29999
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London Stereoscopic Co. imp.

NORTH AMERICAN SERPULID£.

= 2 S 1S tn iecoev nin,

London Stereoscopic Co. imp.

NORTE AMERICAN | SERP UES 7.

1 Loot). LNA el hp OOD:

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London Stereoscopic Co. imp.

NORTH AMERICAN SERPULEID A.

POLYCH#HTA FROM NORTH AMERICA. 785

2. Uncini generally 6 or 7 teeth, the anterior one being the
largest (figs. 202 & 203 Jonnson, 9).

3. Very large size (fig. 1) with numerous abdominal segments,
30 to 54 branchize and from 80 to 160 serrations on edge
of operculum.

Numerous specimens from Departure Bay, Dodds Narrows,

and two from Puget Sound (Prof. Kincaid’s collection).

Isolated tubes are to be found attached to the undersides of
stones on rocky shores near extreme low-water mark, and small
colonies may be found above this level in rock-pools.

In other places attached to rocks near low-water level the
masses of large white calcareous tubes are very striking—they
are thick and often finely ridged, the lower parts being much
intertwined, the free distal ends often overgrown with Polyzoa,
Spirorbis, ete.

The brilliant red colour on the branchial crown may involve
the whole of the gills and operculum, or these may be colourless
except for the tips, or they may be barred and mottled in a
large variety of ways. The remainder of the body is generally
yellowish.

As regards collar, thoracic membrane, and operculum with its
tubercles, this species agrees very closely with S. vermicularis,
so fully described by St. -Joseph (24. pp. 328-335). Johnson (9)
presumably has made it into a new species on account of its
very much greater size with corresponding greater number of
abdominal seements, branchie and ade eins on operculum,
together with small differences in the sete and uncini. He
cannot, I think, have examined many specimens, for he states
that the functional operculum is on the right side (9. p. 432).
The position of this in the genus Serpula may be right or left as
shown by Zeleny (27. p. 34), but out of 50 specimens that I
have examined 28 had it on the left, 21 on the right, and the
remaining specimen had one on each side. Consequently, when
he says that there are about 100 serrations on the edge of the
operculum, I cannot think that Bush (8) is justified in recording
this as a distinction between this species and S. splendens with
127 to 150 serrations.

Moore (19) gives 140 for the one specimen of S. columbiana in
which he counted the serrations. In my specimens they vary
from about 80 to 160—the number apparently increasing with
age.

Another distinction given by Bush is that there are in
S. columbiana “but 250 abdominal segments in a length of
55 mm.,” whereas Johnson says ‘250 or more” (9. p. 432), and
she gives 313 as the number in a specimen of her so-called
S. splendens, of which she does not state the size, and 190 ina
specimen 35 mm. long. These figures speak for themselves I
think as creating nothing but confusion. In 15 of my specimens
the average number of abdominal segments was 236, including 79
in a specimen 10 mm. long, 142 in a specimen 4] mm. long, and
310 in one 81 mm. long.

786 MISS HELEN PIXELL ON

The third point of difference given by Bush is that S. colwm-
biana has more numerous branchie—54 in each lobe as given
by Johnson—than her S. splendens, for which she records 45 to
50 pairs.

In the specimen of the former species counted by Moore (19)
there were only 38. One small specimen of mine had only 18,
but the general number was from 30 to 50 on each side, one or
two large specimens having as many as 54—the number given
by Johnson. These facts show clearly I think that S. splendens
is a quite unnecessary species and can be included in S. columbiana,
which in turn, as pointed out by Johnson (9. p. 433), may be
identical with WS. jukesti Baird, for which however no satisfactory
description was given.

Numerous large free Selenidia in the trophozoite stage were
found in the alimentary canal of nearly every specimen of
S. columbiana examined.

Genus Crucicrra Benedict (1), 1886.

Generic characteristics :—

1. Collar sete and uncini similar to those of Serpula.

2. Operculum with comparatively few radii forming a scalloped
margin to the funnel and with conspicuous basal pro-

' cesses.

2. CruCcIGERA zYGopHORA Johnson. (Pl. LXXXVII, fig. 2.)

Serpula zygophora Johnson (9), 1901.
Crucigera zygophora Bush (8), 1905.

Specific characteristics :—
1. Branchize about 30 pairs with long filamentous ends to

rachises.
2. Operculum thick, shallow, with about 30 radii aad 3 rounded

processes at its base ; attached by a long pedicle.
One specimen from Puget Sound was 45 mm. long. A smaller

incomplete specimen came from Victoria (fig. 2). Another
specimen was only 7 mm. long and had a much thinner operculum,

but seemed otherwise Similan

3. CRUCIGERA IRREGULARIS Bush (8), 1905. (Pl. LXXXVII.
fig. 3.)

Specific characteristics :—

1. Branchize much coiled and with comparatively short fila-
mentous ends to rachises.

2. Operculum irregular, apex of funnel displaced ventrally and
the posterior eral lager walls deeper and rolled over to
some extent, Not more than two basal processes which
may, however, be bi-lobed and attached by long stout
pedicle (fig. 3).

About 12 specimens from the Channel outside Departure Bay

POLYCHZTA FROM NORTH AMERICA. 787

and one from Dodds Narrows. Depth 15 to 25 fathoms. Tubes
generally solitary, attached to stones, shells, old wood, ete. Only
one specimen is recorded by Bush from Juneau. Tubes much
coiled, with flaring ends and one or two other conspicuous ridges
at intervals indicating the flaring end of a younger tube. Young
tubes seem to develop with a centimetre or so attached more
or less straight along the substratum, then to coil indifferently
to right or left, and only ata much later stage, if at all, to ascend
and form the flaring end,

General colour pale orange, the branchie and operculum
variously mottled and barred with red. Pinne sometimes
golden; ova greenish, Length varies from about 14 mm. to
50 mm. (fig. 3).

Largest diameter of operculum 1:3 to 4:5 mm.; the latter had
32 radii forming a thick scalloped edge. The pedicle was bifid
and contracted at the top just before joining the basal processes
of the operculum (fig. 3). Sete as figured by Bush (8); in the
posterior region of the abdomen the ordinary sete are replaced
by small fascicles of long slender spines.

Genus Apomatus Philippi (21), 1844.

Generic characteristics :—

1. Opereulum globular, terminating a gill retaining its pinne.

2. Some thoracic sete bladed sickles (sete of Apomatus)
(fig. 4c).

3. Terminal dorsal gland present.

4, APOMATUS TIMSII, sp.n. (Pl. LXXXVII. figs. 4 a-4f.)
Specific characteristics :—

1. Collar setz simple tapered blades (fig. 4 6).
2. Branchiz about 40 pairs with pinne nearly to the ends of
rachises.

3. Uncinigerous tori begin on third setigerous segment.

4, Uncini with numerous small teeth, the posterior one larger
with terminal enlargement (fig. 4 f

Abdominal setz more or less sickle-shaped with some long
filiform ones in the last segments.

Or

Eleven specimens from the Channel outside Departure Bay in
about 20 fathoms. This is the first time apparently that the
genus has been recorded from the Pacific Coast of America.

The tubes are solitary, adherent, and sinuous—one was attached
along its whole length to a portion of the Hexactinellid sponge
Aphrocallistes whiteavesianus.

When full-grown about 80 mm. with 150 abdominal segments
(fig. 4a).

The branchial rachises are much coiled and almost colourless,
with pairs of red spots up their outer surfaces and only short.
filiform extremities—the pinne appear green due to the contained
blood. The thorax is more or less orange or red, the thickened

788 MISS HELEN PIXELL ON

ridges bearing the tori being especially deep in colour. <Ab-
domen pinkish; there is a short anterior asetigerous region, and
posteriorly nearly covering the last centimetre or so of dorsal
surface is a chalky white strap-shaped raised glandular area. This
is the terminal dorsal gland mentioned by St.-Joseph (24) as
possibly serving in the construction of the tube. It tapers
anteriorly and ends abruptly posteriorly just above the vertical
slit-like anus.* Along nearly the whole of this glandular region
the ordinary abdominal sete are replaced by fascicles of about five
slender capillary setze without blades and nearly equal in length
to the width of the abdomen at this place. The collar is entire
ventrally and has a deep incision on each side—the lateral lobes
being continuous with the wide thoracic membrane.

The functional operculum is large and transparent and carried
by the second branchia from the dorsal side. The pedicle is
generally coiled twice like the other gills; except in very young
specimens there is a small club-shaped one terminating the
branchia on the opposite side.

All the collar sete and most of the other thoracic setz have
simple narrow blades; amongst them in the posterior thoracic
fascicles are to be found a few of the typical bladed sickles
(fig. 4c). The abdominal sete are more or less sickle-shaped,
though they easily become folded (figs. 4d, e) or straightened
out.

Several of the specimens examined were much smaller and
obviously young forms, with 70, 83, 101, 114 abdominal segments
and fewer branchiz than in adult specimens.

Genus Proruta Risso (23), 1826.
Generic characteristics :—
1. No operculum.

2. Collar sete simple tapered blades (fig. 5 6).
3. Terminal dorsal gland present.

5, PROTULA PACIFICA, sp.n. (Pl. LXXXVII. figs. 5 a—5 f.)
Specific characteristics :—
1. Ventral lobe of collar notched.
2. Abdominal setz somewhat sickle-shaped in the anterior
region, narrow terminal bladed ones posteriorly.
3. Uncinigerous tori extend from segment 3 to the end of the
abdomen.
4, Uncini with numerous small teeth, the posterior one long
with a bulbous extremity (fig. 5 /).
5. Thoracic sete simple blades with sometimes a few sete of
Apomatus posteriorly.
* Tn section the gland is seen to consist of epithelial cells crowded with spherical
granules which stain easily with iron hematoxylin but not with Delafield’s. The

gland is apparently unchanged by the presence of acid in the preservatives and
therefore does not appear to be calcareous.

.

POLYCH&TA FROM NORTH AMERICA, 789

Three specimens from Fairway Channel outside Departure Bay
in about 30 fathoms; one from Puget Sound (Prof. Kincaid’s
collection). No tube.

The specimens were colourless except for the branchie, which
appear greenish due to the contained blood. Down the outer
side of each branchial rachis was a line of opaque white spots.

The branchial crown (10-15 mm. high) easily falls off, leaving a
scar; the remainder of the body is 38 to 60 mm. long, rather more
than one-third of which is thorax. This is broad and flattened
with nearly parallel sides (fig. 5a) and the usual 7 setigerous
segments. The thoracic membrane is very wide, with an entire
margin which can extend beyond the sete but is generally
considerably crumpled. The collar is notched in the median
ventral line and has a deep fissure on each side. The inward
coil of the stout base of the branchial crown is seen on the right
side of the specimen in fig. 5a. It takes 14 to 2 turns inwards
and upwards, the short inner gills reaching about the same
height as the outer long ones. An interbranchial membrane
connects the lower third of the gills, of which there are about
60 pairs having small closely placed pinne almost to their
extremities. At the junction of the branchie with the stout
basal membrane which carries them there is a slight ridge
visible externally, and at this place on the inner side arises
the oral membrane, which is continuous across the median line
and up each of the spirals. On the dorsal side of the mouth is
another shorter membranous lip.

Uncini begin on the third setigerous segment. They have the
characteristic shape (fig. 5). The first two millimetres or so of
the abdomen is achztous, nearly round, and of smaller diameter
than the wide dorso-ventrally depressed part which follows.
The ventral surface has deep segmental grooves showing 83-110
segments, and a wide fzcal groove which turns to the right on
reaching the thorax. Laterally the short tori are raised on
distinct parapodial processes which extend to the posterior end
of the body. The last 30 or so segments on the dorsal surface
are covered with the calcareous-looking gland which anteriorly
tapers off to two points, suggesting a paired origin.

In section the gland is seen to occupy nearly the whole
thickness of the dorsal body-wall, the longitudinal muscles being
pushed towards the sides. Nearly all the cells are crowded with
the spherical granules or globules which stain easily with iron
hematoxylin, and are apparently similar to those in the glandular
cells so frequently found in the epidermis.

This hind region of the abdomen has very long sete which
extend 2 mm. or more on each side but are easily broken. At
first sight they appear to be simple spines very slightly bent at
the extremities, but, with high magnification, a narrow striated
wing may be made out (fig. 5e), shorter sete found with these
are more distinctly winged (fig. 5d). The other abdominal
segments have ventral fascicles containing about 13 shiort, stout,

Proc, Zoou, Soc.—1912, No. LITT. 53

790 MISS HELEN PIXELL ON

somewhat sickle-shaped setz (fig. 5c). These setz were, un-
fortunately, not examined before being preserved, and as
St.-Joseph (24. p. 338) points out, a lengthened immersion in
alcohol tends to reduce the curve of the sickle.

In general structure this species resembles P. capensis McIntosh
(13) fairly closely, but differs in the shape of the set# and
uncini: in the latter respect and in some other points it differs,
too, from P. diomedece Benedict (1). There are many character-
istics distinguishing it from P. superba Moore (19) and other
Pacific species that have been described. P. atypha Bush (8)
might possibly be a young specimen of this same species.

Genus Currrnopoma Levinsen (10), 1883.

Generic characteristics :—

. No thoracic membrane.

. Collar sete with fin-like expansion at base of blade.

Some of the other thoracic setz are sickle-shaped.
Abdominal sete geniculate.

. Uncini with 9 or 10 fine teeth, the anterior one being
_ larger and blunter than the others.

6. Operculum with horny plate.

Our co bo

6. CHITINOPOMA GREENLANDICA. (PJ. LX XXVIII. figs. 6 a—6 ¢.)

Serpula triqueter Fabricius (7), 1780.

Hydroides norvegica var. grénlandica Mérch (20), 1863.
Hydroides (¢) grénlandica Malmgren (14), 1867.
Chitinopoma fabricit Levinsen (10), 1883.

Specific characteristics :—
1. Bodies elongated, somewhat cylindrical.

2. About 6 pairs of branchiz with ends free from pinne.
3. Operculum enclosing central stalked vesicle.

Numerous specimens from Departure Bay and neighbourhood,
one incomplete one from Victoria. In thick sinuous tubes
adherent to shells, stones, etc., and having a very conspicuous
dorsal keel generally ending in‘a spine overhanging the aperture.
One tube was U-shaped with the two ends close together. The
largest specimen was about 12 mm. long (fig. 6a). The whole
animal was practically colourless, the pedicle of the operculum
and the branchiz sometimes having faint transverse bands, and
the contents of the alimentary canal were in some specimens
dark rea.

The branchiz varied very much in number (from 6 to 8 in each
lobe) and some were frequently found in a rudimentary state:
one specimen had 7 in the right, and only 3 fully-developed
functional ones in the left. The operculum was in every case
on the left dorsal side, and I have found no trace of a secondary
one on the other side.

POLYCHAHTA FROM NORTH AMERICA. 791

The full-grown branchise had about 17 pairs of ciliated pinne,
which stopped short some distance from the top of the rachis
leaving a fairly long filamentary end. In sections the rachises
are seen to be strengthened externally by a thick layer of chitin,
which also forms a protecting layer round the pedicle (fig. 6 ¢),
and is continuous over the operculum with the thick horny plate
on its top. This plate was often covered with sand, and many
specimens had infusorians, ete. attached to it. The central
coelomic space in the pedicle is lined with peritoneum provided
with very large conspicuous nuclei. There is a small vessel
running along its whole length and enlarging in the opercular
cup into a spherical vesicle (fig. 6¢). This is filled with a finely
granular precipitate, and from its wall and general appearance
seems to correspond with the branchial blood-vessels of the
ordinary gill rachises, though I do not see that it can have any
respiratory function. It is apparently suspended in a fine
reticulate connective-tissue which easily shrinks away from the
epithelial cells ; Levinsen, in his original description (10. p. 203),
suggested that it might be a new operculum forming in the old
one.

The collar is very wide—the entire ventral lobe being generally
reflexed—the latero-dorsal lobes are continued down the dorsal
side to between the second and third thoracic sete, where they end
abruptly, giving the appearance of a short thoracic membrane.
This was a constant characteristic in both small and large
specimens. I have not seen it referred to before, but do not
think it necessary on that account to separate this as another
species.

With the large collar sete (fig. 6 5) are a few shorter curved
forms with very narrow blades.

The abdominal sete agree with those described by St.-Joseph
(24) for the genus (fig. 6c).

The abdomen is long and slightly dorso-ventrally flattened,
with 25-40 segments. The tori contain about 17 uncini, but
there are only a few sete to each segment. The dorsal longi-
tudinal muscles are greatly developed——there is a small fascicle
of ventral ones on each side of the wide fecal groove. The large
ventral nerve-cords are separated from one another. The
epithelium consists of low columnar cells with numerous gland-
cells containing the usual spherical masses which stain easily
with iron hematoxylin.

This species seems to agree very closely with that described
by Bush (3) as Hyalopomatopsis occidentalis. I cannot under-
stand why this form with no thoracic membrane and “ trumpet-
shaped” abdominal sete is put in a genus, of which St.-Joseph
writes (24. p. 264): “Il m’a fallu créer un genre nouveau
Hyalopomatopsis pour le Hyalopomatus langerhansi Eh. et le
H. marenzelleri Lang., qui par la présence d’une membrane
thoracique ... ne pouvaient rentrer, comme l’avait du reste
prévu Langerhans, dans le genre Hyalopomatus tel que Vavait

FH Ow

JO”

792 MISS HELEN PIXELL ON

défini von Marenzeller.” St.-Joseph also gives as another
characteristic of his genus Hyalopomatopsis, the presence of
capillary sete in all the abdominal segments.

Genus Sprrorsis Daudin (6), 1800.
(Pls. LX XXVIII, LXX XIX. figs. 7-16.)

Generic characteristics and Schemes of classification :—

1. Caleareous tubes coiled in a dextral or sinistral spiral.
The method of coiling and the markings on the tube have
been used by Bush (3) in drawing up Table I., but the
coiling of the tubes is variable (cf. figs. 8 a and 6), being
determined to a great extent by the nature of the sub-
stratum, and as Caullery & Mesnil (5) have already
pointed out there is no constancy in either coiling or
markings.

2, Branchiz are constant in different species, the operculum
with terminal calcareous plate, always occurring as the
second on the concave side (7. e. the right in dextral forms
and left in sinistral; since the animal lies with its back
towards the substratum).

3. Thoracic segments generally 3—the first having only dorsal
setee—the two following have on each side an uncinigerous
torus as well. In the subgenera given the prefix Para-
by Caullery & Mesnil an extra torus (and in Sp. can-
cellatus a fascicle of dorsal sete also) is developed on
the concave side of the animal; this condition has been
described in the following Table of Classification (p. 794)
as 35 setigerous segments.

In Sp. ambilateralis, sp. n., there are four complete setigerous
segments, although the fourth on the convex side is very reduced.
This specimen therefore approaches the hypothetical Prospirorbis,
described by Caullery & Mesnil (5. p. 233), who point out
that the genus Spirorbis has been evolved from other Serpulide,
which have the characteristic greater number of thoracic segments,
by a gradual reduction. I have therefore placed this species in a
new sub-genus Protoleospira.

4. Abdominal segments 8-40,

Between the thorax and the abdomen is a more or less long
asetigerous region—often crowded with ova. The spermatozoa
develop in the posterior setigerous segments.

5. The thoracic sete are distinctive and the differences are of
use as specific characteristics. Table II. given by Bush

(3. p. 261) is drawn up with regard to these and the
direction of the coiling alone.

As a rule, the first thoracic segment has some slender capillary
sete forming the inferior part of the fascicle (fig. 146)—the

POLYCHZTA FROM NORTH AMERICA. 793

superior ones, referred to as collar sete, are distinctive, they may
have simple blades (fig. 13) or there may be a distinct fin-like
expansion at the base of the blade (figs. 7, 10a, 11a). In the
case of sinistral forms Caullery & Mesnil referred the former
to their sub-genus Romanchella, putting only those with a distinct
fin in the sub-genus Leospira. In the species S. verruca and
S. evolutus Bush and S. medius, sp. n., however, the set are
intermediate between these two types, having blades which are
faintly notched (fig. 14a), showing an indication of a superior
blade and inferior fin. Consequently I propose to do away with
Caullery & Mesnil’s sub-genus Romanchella altogether—taking
Leospira to include all sinistral forms with three setigerous
segments: it thus comes into line with the other three sub-genera
proposed by Caullery & Mesnil (cf. Table of Classification).

The second thoracic segment has only ordinary bladed sete,
differing very little from one species to another.

The third thoracic segment has some ordinary bladed sete,
but generally also some bladed sickles as found so generally in the
genus Apomatus (fig. 14 e).

In many Pacific species there are present instead of the
ordinary bladed sickles a peculiar shorter form which appears
almost fringed at the extremity (fig. 10).

6. Abdominal ventral setz generally geniculate.

7. Uncini similar in thorax and abdomen—plates with the free
edge provided with fairly numerous fine teeth, the anterior
one being jarger than the others.

The following Table of Classification has been adapted from
that given by Caullery & Mesnil in their excellent paper on
Spirorbis (5) to include such of the Pacific forms as have so far
been studied. These authors pointed out that such a modification
would possibly be necessary.

Unfortunately the majority of the new species described by
Bush (2, 3, 4) from California, Alaska, and Japan cannot be
included owing to absence of information as to the number of
thoracic segments. Two or three of the species are established
on details as to the tubes alone.

Sub-genus PaRaDEXIosPiRA Caullery & Mesnil (5), 1897
(modified).
Characteristics :—
1. Tube dextral.
2. Thorax with 34 setigerous segments.

7. SprrorBis virrevs Fabr., 1780. (Pl. LX XXVIII. fig. 7.)

Serpula vitrea Fabricius (7), 1780.

Spirorbis vitrews Morch (20), 1863; Malmgren (14), 1867 ;
Levinsen (10), 1883; Caullery & Mesnil (5) 1897;
Moore (17), 1902; Bush (8), 1905.

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Specific characteristics :—

1. Last thoracic segment on the right has no dorsal sete.

2. Collar sete with fin-like expansions at base of deeply serrated
blades.

3. Some seta of third segment bladed sickles.

4, Embryos incubated in the tube.

This species was quite common on stones and rocks from the
Departure Bay region and from Victoria.

The tubes vary a good deal but are always translucent, with
the whorls piled on one another. The whole tube measures
about 2°5 mm. in diameter and the aperture about 1 mm. across.
There are various markings on the exterior. Sometimes a
ridge along the median line ends in a sharp projection above
the aperture—in the grooves on either side of this there may
or may not be scalariform markings. Young specimens have
shells as figured for this species by Levinsen (10. fig. 11), older
ones were ridged more like Sp. cancellatus (10. fig. 18). The living
animals were bright pink in colour. Gills 7, each with six pairs
of long opposite pinne, opercular plate a shallow funnel. About
20 abdominal segments.

Sub-genus Dextospira (Caullery & Mesnil, 1897).
Characteristics :—

1. Tubes dextral.
2. Thorax with 3 setigerous segments.

8. Sprroreis sprrintuM Linné, 1767. (Pl LUXXXVIII.
figs, 8a—8c.)
Serpula spirillum Linné (12), 1767; Fabricius (7), 1780.
Spirorbis spirillum Malmgren (14), 1867; Levinsen (10),
1883; Caullery & Mesnil (5), 1897; Moore (17), 1902;
Bush (3), 1905.
Spirorbis lucidus Montagu (16), 1803; Mérch (20), 1863 ;
Malmgren (14), 1867.
Circeis armoricana Saint-Joseph (24), 1894.
Spirorbis borealis Fewkes (8), 1885.
Specific characteristics :—
1. Collar sete geniculate (fig. 8 ¢).
2. Operculum without brood-pouch.

3. Concave plate of operculum has a slight projection (talon)
on under side.

Two varieties of this species were fairly common in the Departure
Bay region and at Victoria, growing on calcareous polyzoa,
seaweeds, etc. . The discoid form grows only on smooth surfaces,
Laminarian thalli being often extensively covered with specimens.
These were very flat, regularly coiled tubes—the spiral with
14 to 3 coils measuring 5 to 2mm. in diameter (fig. 8a). The

POLYCH ETA FROM NORTH AMERICA, 197

ascending variety in some localities is much more common than
the discoid one, and it grows toa much larger size (fig. 8 6), often
attaining a height of 5 mm. It was generally found with its
lower coils overgrown with an orange Bryozoan growing on
Chetopterus tubes.

The living animal had a reddish colour, witha colourless trans-
parent operculum. A string of pink ova extended along the tube
beyond the posterior end of the body in some specimens. There
were 3 thoracic and from 12 to 20 abdominal segments.

The collar sete attain a length of °27 mm., much larger than
those figured by Caullery & Mesnil (5. fig. 46), they resemble
more closely in shape those given for S. armoricanus (5. fig. 5 6).

The differences between these two species were pointed out
by Caullery & Mesnil (5. p. 199) to be quite unimportant. All
that they could summarise were that Sp. armoricanus (Circeis
armoricanus St.-Joseph (24. p. 350)) was slightly larger, had more
abdominal segments (16-20) and a reduced talon to the operculum.
They record having seen intermediate forms themselves, and the
above observations as to variation in size, the number of abdom-
inal segments, and the collar setz consequently confirm their
opinion that Sp. armoricanus should be considered as a variety

only of Sp. spirillum.

9. SprrorBIs PUSILLOIDES Bush (3). (Pl. LX XXVIII. figs. 9a,

. ; 9 b.
S. pusillus Caullery & Mesnil (5), 1897. )
Mera pusilla St.-Joseph (24), 1894.

Non S. pusillus Rathke (22 a), 1836.

Specific characteristics :—

1. Collar setze of a more or less geniculate form (fig. 9 @).
2. Embryos incubated in operculum.

3. Sete of 3rd thoracic segment sickle-shaped.

4. Hepatic pigment reddish brown.

On stones from Taylor’s Bay, Gabriole Island. This animal
agrees in all important points with the full description by
St.-Joseph (24. p. 351) for Mera pusilla. The collar sete are a
little more distinctly angulated at the base of the blade (fig. 9 a),
but they have not the typical geniculate form described by
Caullery & Mesnil (5. p. 202). They are much shorter than
the sete of the second segment and are decidedly hooked. The
extent to which the sete are hooked has been pointed out by
St.-Joseph to be reduced by preservation (24. p. 338), so that
this does not seem to be a very important point.

The opercular brood-pouch is somewhat cylindrical and bounded
proximally and distally by caleareous plates, the talon is reduced
to a small quadrangular projection.

The abdomen has eight segments, with large, more or less
sickle-shaped sete (fig. 9b) and an asetigerous anal segment.

798 MISS HELEN PIXELL ON

Sub-genus PRoTOLMOSPIRA, Nov.

1. Tube sinistral.
2. Four complete setigerous segments to thorax.

10. SpPrRoRBIS AMBILATERALIS, sp. Nn. (Ely lu Xexexc yeaa
figs. 10 a-10e.)

Specific characteristics :—

1, Collar seta very large, conspicuously serrated blades with
fin-like expansion at base (fig. 10 a).

2. Operculum without brood-pouch.

3. Talon of operculum with large hook-like process (fig. 10 6).

4. Some sete of 3rd segment have fringed ends (fig. 10).

5. Some setze of 4th segment ordinary bladed sickles.

6. Abdominal sete brush-like (fig. 10).

7. Dorso-lateral brood-pouch.

Several specimens on the inner sides of shells of Balanus
nubilus from Dodds Narrows, 15-25 fathoms. ‘Tubes forming
translucent sinistral spirals measuring 3 to 4mm. across. ‘The
surface is distinctly corrugated outside, highly polished inside ;
the aperture measures 1 mm. in diameter.

Branchie 12—6 on the right, and the operculum with 5
others on the left. The pinne extend upwards and reach the
same height as the rachises. The opercular plate (fig. 106)
is very like that of S. cornu-arietis Marion & Bobretzky (15).
From close to the point of origin of the pedicle arises a wide tube
which passes across the dorsal surface beneath the left lateral
lobe of the collar, and enlarges into the thin-walled brood-sae
which lies along the dorso-lateral surface on the right side. This
structure has never, so far as [ am aware, been described before:
it is figured and more fully described for S. racemosus, in which
species it was first seen.

Collar wide and entire ; the collar sets rather more than | mm.
in length (fig. 10a). Sete of 2nd segment very numerous, all
with simple blades; these appear again in the 3rd fascicle, and
with them some shorter bladed setze with their extremities more
or less curved and fringed (fig. 10¢). The sets of the 4th thoracic
segment are few in number; on the convex side of one specimen
there were three bladed sickles (fig. 10d) and three with plain
blades. The uncigerous tori on the 4th segment are also smaller
than on the two preceding, especially on the right (convex) side.
The thoracic uncini are about 90 ~ long and have 20-25 teeth.
Those on the abdomen are only 25 y in length. About 50 of
the latter make up each of the tori which are very distinct on
the 18-20 abdominal segments: they begin quite close to the
posterior end of the thorax—the anterior asetigerous region of
the abdomen being very short.

The collar sete in this form seem to have been specially

POLYCH ETA FROM NORTH AMERICA. 799

developed, for according to Caullery & Mesnil (5) the possession
of blade and fin is not a primitive characteristic, and they are
also very large; possibly they are specially developed in correla-
tion with its habitat in rapids where the tide runs from 7—9 knots
an hour. The presence of the specially developed brood-pouch
might perhaps receive a similar explanation.

Sub-genus PARALAOSPIRA.

1. Tubes sinistral.
2. Thorax with 33 setigerous segments.

11. SprrorBis RACEMOSUS, sp. n. (Pl. LX X XIX. fig. 11.)

Specific characteristics :—

1. Collar sete with fin-like expansion at base of very coarsely
serrated blade (fig. 11 a).

2. Operculum with massive talon having lateral wings and a
hook on ventral surface (fig. 11 6).

3. Thin-walled dorsal or lateral brood-pouch attached to the
thorax at the base of the operculum (fig. 11d).

4. Some setz of 3rd fascicle bladed sickles (fig. 11 ¢).

A few large specimens from San Juan Island on barnacle-
shells overgrown with calcareous sponge; smaller ones from
Channel outside Departure Bay. Depth in both places 15 to
25 fathoms.

The largest specimen measures 5 mm. across the spiral; the
tube is corrugated and generally rather loosely coiled, leaving a
central hole.

Branchie 13—6 functional ones on each side; rachises have
short tapered extremities without pinne; operculum not unlike
that of S. ambilateralis, but the talon rather more massive and
its outer concavity almost hemispherical and filled with débris.

The collar sete are very large (fig. lla); the 2nd fascicle
consists of plain-bladed sete, and the 3rd contain some fringed
sickles as well. There is a distinct third uncinigerous torus on
the left side (fig. 11d); the uncini have the usual shape.

There are 21 well-marked abdominal segments, each with three
or four brush-like sete and along asetigerous anal segment. The
brood-pouch with its distinct wide stalk seems to be peculiar to
this and the last described species of Spirorbis: it does not appar-
ently replace a gill, for the caleareous operculum is developed as
usual on the 2nd to the left. The sac has a very thin wall, the
- large ova causing marked protuberances on its surface, the whole
somewhat resembling a bunch of grapes (fig. 11d).

Sub-genus Lospira.

1. Tube sinistral.
2. Thorax with 3 setigerous segments,

800 MISS HELEN PIXELL ON

12. SprrorBis AsPERATUS Bush (8), 1905; Sitka. (Pl. LXXXIX,
fig. 12.)

Specific characteristics :—

1. Collar sete with simple blades.

2. Operculum without brood-pouch.

3. Calcareous plate of operculum thin, with large lateral wings
on its talon.

4, Some sete of 3rd fascicle short fringed sickles (fig. 12),
others with long narrow blades.

On an old Serpulid tube inhabited by a hermit crab. One
specimen only. The opaque sinistral tube shows conspicuous
growth-lines, and the whorls are somewhat piled on one another.

There are 12 or 13 reddish gills nearly hidden by the high
collar, above which the operculum projects for some distance.
The alimentary canal was greenish and the ova along the dorsal
surface salmon-pink. The abdomen consisted of about 20
segments.

13. Sprrorprs vatipus Verrill (26), 1874. (Pl. LX X XIX. fig. 13.)
S. verruca Levinsen (10), 1883, figs.+Caullery & Mesnil
(5), 1897 ; Moore (17), 1902.
S. validus Bush (8), 1905.

Specific characteristics :—
1. All thoracie sete with long, finely serrated, narrow blades
(fig. 13).

2. Operculum with brood-pouch.

3. Opercular plate (Bush (3), pl. xliv. figs. 11-14).

4. Branchie 13.

On old shells of Balanus nubilus from Dodds Narrows, 15 to
28 fathoms.

The tubes were smooth, sinistral, and opaque, measuring 3 mm.
in diameter. The operculum was colourless and transparent,
with a large sac-like brood-pouch which was, however, empty in
specimens collected on September Ist. The gills and thorax were
of a bright orange-red colour, the rachises being thick with a double
row of small pinne extending inwards at right angles. Abdomen
with 25 segments having a few sete, some being somewhat
geniculate, others small hooks (Bush, pl. xxxvu. figs. 5 & 6).

14, SprrorBis MEDIUS, sp.n. (PI. LXXXIX. figs. 14 a-14e.)
Specific characteristics :—
1. Some collar setze with shallow posterior notch (fig. 14 a).
2. Operculum without brood-pouch.
3. Caleareous plate of operculum very large and of character-
istic shape (figs. 14c & d).
4, Some setee of 3rd segment serrated bladed sickles (fig. 14 e).

From Channel just outside Departure Bay. Large flat tubes

POLYCHATA FROM NORTH AMERICA, 801

thick and opaque, slightly roughened, but without definite growth-
lines. A slight median ridge and sometimes one on either side ;
the aperture has, however, an entire margin and measures 2 mm.
ACLOSS.

The animal is of a uniform brick-red colour, and its total
length is 455 mm; there are 14 branchiw—7 being joined at
their bases and situated on the right side; the other 6 func-
tional ones are joined with the operculum on the left. Hach
rachis has a thin membranous projecting flap which wraps round
the outside of the next, and thus gives rise to a series of imbri-
eating semilunar membranes just inside the base of the collar.
The total height of the gills is 1 mm. There is a very wide
thoracic membrane on the right side which almost envelops the
whole animal. The calcareous plate of the operculum is 1°5 mm.
along its long axis; the outer side is concave and generally
covered with sand; the talon projects obliquely inwards and has
large wing-like expansions at the sides (figs. 14c & d). The
collar setze (fig. 14 a) appear to be in an intermediate stage between
plain bladed form and that with a distinct fin-like expansion at
the base of the blade—in one specimen I could find none of these,
only forms with the ordinary blades. In the 2nd segment there
were numerous setz of the normal kind, and the 8rd fascicle was
made up of bladed sickles (fig. 14¢) and ordinary bladed sete.
The two tori in the thorax consisted of uncini of the ordinary
shape with about 20 teeth. In the abdomen were counted 20 to
25 segments, and the setz were of the ordinary geniculate type.

15. SPIRORBIS LANGERHANSI Caullery & Mesnil (5), 1897, and
Bush (3), 1905, from Panama. (Pl. LXXXIX. fig. 15.)

Specific characteristics :—

1. Collar set# with fin-lke expansion at base of coarsely
serrated blade (fig. 15).

2. Embryos incubated in operculum.

3. Operculum of characteristic shape (5. fig. 22).

On tubes of Serpula columbiana from Departure Bay. Tubes
small, marked with lines and ridges.

The structure of the animals agrees with the description given
by Cauliery & Mesnil (5. p. 217). They are 1-2 mm. long,
the abdomen being broad, with a long asetigerous region followed
by 9 short setigerous segments.

16. SprrorBis MORcHI Levinsen (10), 1883 (Greenland); Caullery
& Mesnil (5), 1897; Bush (8), 1905. (Pl. LXXXIX.
figs. 16 a-16 d.)

Specific characteristics :—

1. Collar sete about 12, with fin-like expansion at base of
coarsely serrated blade (fig. 16 a).

2. Embryos incubated in operculum (fig. 16 6),

3. Branchie 8.

802 MISS HELEN PIXELL ON

Numerous specimens on tubes of Serpula columbiana from
Departure Bay. The opaque tubes are pressed together with
their ends standing erect, their bases being overgrown with an
encrusting sponge ; the surfaces are free from regular markings, the
apertures being circular and measuring about ] mm. in diameter.

The animals vary in colour, some being almost colourless, but
were generally some shade of red or brown.

The sete of the 2nd and 3rd thoracic segments have long, very
narrow, delicately serrated blades (fig. 16c). A few of the 3rd
are shorter, with small posterior blades; these probably represent
straightened sickles (fig. 16d).

There is a deep collar and a wide thoracic membrane on the
right side only—this traverses the ventral surface obliquely
towards the end of the thorax and covers over several segments
of the abdomen. ‘The gill rachises are thick and have long
pinne, the seven normal respiratory ones are rather taller than
the one bearing the operculum. The latter is protected at
its extremity by a convex calcareous cap which extends nearly to
the base of the brood-pouch on the ventral side. The top is
quite opaque and slightly bilobed. Fifteen or more large reddish
ova were contained. Abdominal segments 20-29 with normal
geniculate sete.

17. SPIRoRBIS VARIABILIS Bush (3), 1905; Sitka.

Specific characteristics :—

1. Collar sete coarsely serrated blades with posterior fins.
2. Operculum without brood-pouch.

3. Talon of operculum without projections.

On stones and tubes of Serpula columbiana from Departure
Bay and neighbourhood. Thick tubes about 1 mm. across with
the outer whorl spreading over the others to some extent ; surface
markings were not very distinct.

General colour brick-red, with cerise ova along the dorsal
surface.

Branchiew 8 (including operculum), are nearly hidden by the
high collar. The thoracic membrane on the right side is very much
developed. The opercular plate had a marked concavity on its
outer side, and a small almost central talon projecting inwards.
The sets agree with those described by Bush (3). The abdominal
segments varied a great deal in number, from 15 to 28.

18. SprroRBIS QUADRANGULARIS Stimpson (25), 1853.

Spirorbis quadrangularis Morch (20), 1863, and Bush (8),
1905.

Spirorbis fabricii Malmgren (14), 1867.

Spirorbis carinatus Levinsen (10), 1883 + (11), 1886 +
Montagu (16), 1803.

Spirorbis afinis Levinsen (10), 1883.

Spirorbis granulatus Moore (17), 1902 + (?) Caullery &
Mesnil (5), 1897,

POLYCH ETA FROM NORTH AMERICA. 803

Specific characteristics :—

1. Collar sete finely striated blades with basal fin.

2. Operculum with large brood-pouch.

3. Convex caleareous cap with long cylindrical projection
(figs. 14, 15, pl. xliii., Bush (3) 1905).

Victoria ; two specimens only obtained.

Tubes measure 2 and 3 mm. across respectively ; each has a
very conspicuous median ridge and two lateral ones, the outer
one being at the top of the perpendicular outer wall. The stria-
tions on the blades of the collar sete could hardly be distinguished
at all. Some sets in the third fascicle were serrated, bladed
sickles. The collar and thoracic membrane were well developed
on the right side, and there was a long asetigerous region
following the thorax. There were 12 abdominal segments with
geniculate sete.

Most of the specimens described in this paper were obtained
during a stay at the Marine Biological Station, Departure Bay,
Vancouver Island, in the summer of 1911. Mr. F. A. Potts
kindly collected those from Puget Sound and some others: I
should like to take this opportunity of thanking him, also the
Rev. G. W. Taylor, F.R.S8.C., Curator of the Marine Laboratory,
Departure Bay, for his kindness and help with regard to dredging
arrangements, and Dr. Marett Tims for his advice and assistance
throughout.

REFERENCES.

1. Benepict, J. E.—Description of ten species and one new
genus of Annelids from the dredgings of the U.S. Fish
Commission Steamer ‘ Albatross.’ Proc. U.S. Nat. Museum,
ix. p. 547, pls. xx-xxv. Washington, 1886.

2. Busu, K. J.—On Spirorbis from Japan (see under Moore
(18)).

3. Busu, K. J.—Sabellide and Serpulide. Harriman Alaska
Expedition, vol. xu. 1905.

4, Busy, K. J.—Description of new Serpulids from Bermuda,
with notes on known forms from adjacent regions. Proc.
Acad. Nat. Sci. Philadelphia, vol. lxii. 1910.

5. Cauntery, M., et Musniz, F.—Etudes sur la morphologie
comparée et la phylogénie des espéces chez les Spirorbis.
Bull. Sci. France et Belgique, xxx. pp. 185-233. Paris,
1897.

6. Dauprin, F. M.—Recueil de mémoires et de notes sur les
espéces inédites ou peu connues de Mellusques, Vers et:
Zoophytes, Paris, 1800, p. 38. Nouveau genre de Ver a
tube calcaire, voisin des Serpules et des Dentales. Bull.
Sci. Soc. Philom., Paris, ii. p. 145, 1800.

7. Fasrictus, O.— Fauna Greenlandica, Copenhagen and
Leipzig, 1780,

804

10.

late

12.
13.
14.
15.
16.
17.
18.
19.

20.

21.

22.

MISS NWELEN PIXELL ON

. Fewxes, J. W. On the Larval form of Spirorbis borealis.

Am. Naturalist, xix. p. 247. Philadelphia, 1885.

. Jounson, H. P.—Polycheta of the Puget Sound Region.

Proc. Boston Soc. Nat. Hist. xxix. pp. 381-437. 1901.

Levinsen, G. M. R.—Systematisk-geografisk Oversigt over
de Nordiske Annulata, Gephyrea, Chetognathi, og Balano-
glossi. Vidensk. Medd. Natur. 4, 1882-3, pp. 92-350.
Copenhagen, 1883.

Levinsen, G. M. R.—Kara-Havets Ledorme (Annulata).
Dijmphna-Togtets Zoologisk-botaniske Udbytte, pp. 289—
303. Copenhagen, 1886.

Linné, C. von.—Systema Nature, xir. 1767.

McIntosu, W. C.—Report on the Polycheta. ‘ Challenger’
Reports, xii. London, 1885.

Matmeren, A. J.—Annulata Polycheta. Gfvers. Vet.-Akad.
Forh. pp. 1-127. Stockholm, 1867.

Marton, A. F., et Bosprerzky, N.—Annélides du golfe de
Marseille. Ann. Sci. nat. (6) 11. pp. 1-106. Paris, 1875.

Montacu, G.—Testacea Britannica. 1803.

Moors, J. P.—Descriptions of some new Polynoide, with a
list of other Polycheta from North Greenland Waters.
Proc. Acad. Nat. Sc. Philadelphia, liv. 1902.

Moors, J. P.—Sabellide and Serpulide from Japan, with an
appendix on Spirorbis by K. J. Bush. Proc. Acad. Nat.
Se. Philadelphia, lvi. p. 157. 1904.

Moors, J. P.—Polychetous Annelids from Monterey Bay
and San Diego, California. Proc. Acad. Nat. Sc. Philad.
Ixi. p. 230. 11909.

Morcu, A. L.—Revisio critica Serpulidarum et Bidrag til
Rorormenes Naturhistorie. Naturhist. Tidsskrift, 1.
pp. 347-470. Copenhagen, 1863.

Puriiprr, A.—On the genus Serpula, enumeration of Medi-
terranean species. Ann. Mag. Nat. Hist. xiv. p. 153.
London, 1844.

QuATREFAGES, De.— Histoire naturelle des Annelés, Tome il.
p. 488. (Suites 4 Buffon, 1865.)

22a. Rarake, H.—Zur Fauna der Krym. Mém. Acad. Imp.

23.
24.
25.

26.

27.

Sci. St. Pétersbourg, iii. pp. 407-429. 1836.

Risso, A.—Histoire naturelle des principales productions
de Europe méridionale, iv. Paris, 1826.

Sarnt-JoserH, Baron de.—Les Annélides polychétes des cdtes
de Dinard. Ann. Sci. nat.(7) xvii. pp. 1-395. Paris, 1894.

Stimpson, W.—Synopsis of the Marine Invertebrata of Grand
Manan. Smithsonian Contributions to Knowledge, vi.
pp. 29-36. Washington, 1553.

Verritt, A. E.—Explorations of Caseo Bay by the U.S. Fish
Commission in 1873. Proc. American Assoc. Ady. Sci.
pp- 340-395. 1874.

ZeveNy, C.—Experiments on the control of asymmetry in
the development of the Serpulid Hydroides dianthus.
Journ. Morph. vol. xxii, No. 4. Philadelphia, 1911.

es a

POLYCHATA FROM NORTH AMERICA. 805

EXPLANATION OF THE PLATES.
Prats LXXXVII.

Fig. 1. Serpula columbiana from left side, X 3.

Fig. 2. Crucigera zygophora, anterior end from right side, X 2.

Fig. 3. Crucigera irreguluris, whole animal from right side, X 2.

Fig. 4. Apomatus timsii. 4a, whole animal from the right side, X 3, d.g., dorsal
terminal gland; 46, collar seta, X 300; 4:c, bladed sickle-shaped seta
from thorax, X 300; 4d, crumpled anterior abdominal seta, X 300;
4 e, short posterior abdominal, seta X 300; 4.f, thoracic uncinus, X 300.

Fig. 5. Protula pacifica. 65a, ventral view, X $; 56, thoracic seta, X 300;
5c, anterior abdominal seta, embedded up to the dotted line, X 200;
5d, short posterior abdominal seta, X 300; 5e, tip of long posterior
abdominal seta, X 300; 5, abdominal uncinus, X 300.

Pratr LXXXVIII.

Fig. 6. Chitinopoma greenlandica. 6a, dorso-lateral view, X 2; 66, collar seta,
x 460; 6c, abdominal seta, X 460; 6d, transverse section through
pedicle of operculum, v.s. stem of vesicle, X 55; 6e, transverse section
through operculum showing vesicle (v) suspended in a loose connective
tissue, X 48.

Fig. 7. Spirorbis vitreus, collar seta, X 220.

Fig. 8. Spirorbis spirillum. 8a, tube of discoid variety, X 48; 8b, tube of
ascending variety, X 10; 8c, collar seta, X 200.

Fig. 9. Spirorbis pusilloides. 9a, collar seta, X 460; 9b, large abdominal seta,
x 460.

Fig. 10. Spirorbis ambilateralis. 10a, collar seta, X 220; 106, calcareous plate
of operculum, lateral view, X 48; 10c, seta from 3rd thoracic segment,
showing fringed sickle-shaped extremity; 10d, ordinary bladed sickle,
from 4th segment (twisted end); 10¢, brush-like abdominal seta. c-e,
X< 220.

Prats LXXXIX.

Fig. 11. Spirorbis racemosus, sp. u. 11a, collar seta, X 220; 11 6, calcareous plate
of operculum, X 48; 1c, seta from 3rd thoracic segment, X 460;
11d, dorsal view of whole animal showing stalkéd brood-pouch, x 12.

Fig. 12. Spirorbis asperatus, seta from 3rd thoracic fascicle, X 460.

Fie. 13. Spirorbis validus, collar seta, X 460.

Fig. 14. Spirorbis medius, sp. nu. 14a, collar seta showing shallow posterior

Fig. 15
Fig. 16

notch, X 460. 144, ordinary capillary seta from Ist fascicle, X 460.
14c¢, calcareous plate of operculum, dorsal view, showing talon and
lateral wings, X 16. 14d, latero-ventral view of opercular plate,
showing flat talon with lateral wing-like projections, X 16. 14, ser-
rated bladed sickle, x 460.

. Spirorbis langerhansi, collar seta, X 460.

. Spirorbis mérchi. 16a, collar seta, X 220. 166, operculum containing
ova, lateral view, X 48. 16c, narrow seta of 2nd and 3rd fascicles,
< 220. 16d, seta from 3rd fascicle, X 220.

Proc. Zoou. Soc.—1912, No. LIV. 54

806 MR. D. SETH-SMITH ON A RARE LORY.

EXHIBITIONS AND NOTICES.
May 21, 1912.

Sir Epmunp G. Loprr, Bt., Vice-President,
in the Chair.

The Secrerary read the following report on the additions
made to the Society’s Menagerie during the month of April
1912 :—

The number of registered additions to the Society’s Menagerie
during the month of April last was 291. Of these 165 were
acquired by presentation, 61 by purchase, 17 were received on
deposit, 8 in exchange, and 40 were born in the Gardens.

The number of departures during the same period, by deaths
and removals, was 189.

Amongst the additions special attention may be directed to :—

1 Riippell’s Guereza (Colobus abyssinicus occidentalis) 3, from
the Southern Cameroons, deposited on April 29th.

1 Black Jaguar (Felis onca) 9, from Para, purchased on
April 12th.

1 Capped Langur (Semnopithecus pileatus), 2 Klands (Tauro-
tragus oryx), 1 Common Camel (Camelus dromedarius), and
1 Ursine Tree-Kangaroo (Dendrolagus ursinus), born in the
Menagerie.

A collection of birds from Columbia, presented by W. K.
Pomeroy, Esq., F.Z.S., on April 6th, containing amongst others,
2 Purple Jays (Cyanocoraa affinis), new to the Collection ; 1 Harpy
Eagle (Thrasaétus harpyia); 2 Severe Macaws (Ara severa); and
2 Golden Hangnests (Icterus xanthornus).

2 Limmergeiers (Gypaétus barbatus), from Russian Turkestan,
purchased on April 19th.

1 large Reticulated Python (Python reticulatus), from the East
Indies, presented by Rowland Ward, Esq., F.Z.8., on April 19th.

Mr. A. Buayney Percivat, F.Z.S., 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. Sern-Surrn, F.Z.8., 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

ON THE PRESERVATION OF OUR NATIVE FAUNA. 807

mongoose. The exhibitor referred to a recent note on the species
by Dr. Bahr in ‘ The Ibis’ for April 1912, p. 293.

Mr. G. A. Boutenesr, F.R.S., F.Z.S., 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-Kastern 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.

June 4, 1912.

K. G. B. Meapr-Watpo, Esq., Vice-President,
in the Chair.

Mr. E. G. Boutenerr, Curator of Reptiles, exhibited a clay-
ball containing a cocoon of the African Lung-fish (Protopterus
annectens) presented to the Society by Capt. J. A. M. Vipan, F.Z.S.
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. Meape-Watpo, Vice-President of the Society,
introduced 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.
The 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.

54%

808 DR. HANS GADOW ON OVARIES

PAPERS.

48. The One-sided Reduction of the Ovaries and Ovidacts in
the Amniota, with Remarks on Mammalian Evolution.

By Hans Gapow, M.A., Ph.Ds Hass Eas:
[Received May 31, 1912: Read June 4, 1912.]

INDEX.

Anatomical Structure and Development............... 803

Sanguine morphologists reckon that it takes about ten years
for their discoveries to find their way into a text-book. It
takes a generation to evadicate erroneous statements, especially
generalisations, out of such books, since most of them are
repeated from others without consultation of the immense host
of original papers. And it is apparently hopeless to expect the
enthusiast or amateur to appreciate the difference between a
generalising text-book of comparative anatomy and a zootomical
account. 1t means progress for a branch of science if we can
inscribe upon its statute-book a few lines of true generalisation,
which, if there be no hedging, require no longer any concrete
examples to be mentioned. If only partial generalisations are
possible, of course the exceptions are to be recorded and every
new case is welcome, until their accumulation in turn permits of
being summarised. Then let there be drawn a line, and let the
discoverer of further cases keep his peace unless he has something
new to say.

The condition of the bird’s ovaries and oviducts is a case in
point. The main facts have by now become ancient history and
general knowledge to the zoologist, so ancient that the original
workers have been forgotten, as much as the name of the
originator of the term morphology.

That the ovaries and ducts of birds are one-sided was probably
known since time immemorial. Perrault * described and figured
them in the Ostrich without further comment. In the year 1810
Wolf mentioned that he had usually found two ovaries in the
Sparrow-hawk, a fact duly incorporated by Tiedemann 7 in his
excellent work, which reveals him as a zoologist far ahead of his
time. Next, Spangenberg? figured the right ovary ina Duck.
Barkow § described the occurrence of right-sided rudiments of
the female generative apparatus in various other birds. Emmert ||
observed equally large right and left ovaries in the Sparrow-

* PpRRAvULT: Mémoires pour servir 4 histoire naturelle. Amsterdam, 1736.

+ TrmpEMANN: Anatomie und Naturgeschichte der Vogel. 1810.

{ SpANGENBERG: Disquisitiones circa partes genitales feemineas Avium. Got-
tinge, 1813, 4to.

§ Barkow: Von der Kloake verschiedener Vogel. Meckel’s Archiv f. Anat. u.
Phys., 1829.

|| Emuert: Reil u. Authenrict’s Archiv.

AND OVIDUCTS IN THE AMNIOTA. 809

hawk and some other Accipitres. When Nitzsch made his
anatomical contributions to Naumann’s ‘ Naturgeschichte’ he
knew already of quite a number of cases of double ovaries.
R. Wagner* added to them in his Lehrbuch, and still more in his
Beitriige. He described the cases of double ovaries accompanied
by double ducts; he knew that vestigial ducts are much rarer
than ovaries, and was well aware of the fact that even in Accipitres
with double ovaries there may be no traces of a second duct, ete.
Stannius f added further cases in his Lehrbuch. It was, how-
ever, mainly owing to Wagner's lists that Dumeéril could give a
general summary in Cuvier’s posthumous edition of his ‘ Anatomie
comparée,’ published in 1836. Owen also felt justified in
summarising without referring any more to special cases. On
p. 247 of his work he says:—‘‘'The symmetry [of the ovaria] is
soon disturbed by concentration of development in the left
ovarium, the right one remaining stationary, and ultimately, in
most birds, disappearing.” On p. 249: *‘ Subsequently the left
oviduct alone proceeds to grow; the right is stationary, or
shrivels ; occasionally it may be discovered as a rudiment in the
mature bird, but usually all trace of it has disappeared.”

Still further instances of abnormalities have been described
and even figured without adding to or upsetting the generalisa-
tions quoted above. In most of the general text-books and
others, which have occasion to mention these organs, the
quotation of certain birds has become a regular stock in trade,
always the same, copied from one author by the next, who in
turn is quoted as the original authority by the more popular
writer.

So far as I know, I was the first to give a possible explanation
of the one-sided reduction, and I translate here what L wrote in
Bronn’s ‘ Thierreich,’ p. 842, published in 1890. “In all birds
only the left ovary is completely formed and functional; the
right is present in most cases, and may even produce unripe eggs,
but these degenerate later and seem never to become free. In
correlation herewith the right oviduct undergoes early reduction ;
at most it persists towards the cloaca as a ligamentous strand,
may be even as a tube which opens into the cloaca. This one-
sided development of ovary and oviduct may be referable to
saving of space. Two completely developed, hard-shelled eggs
would scarcely have room in the belly, and we may add that even
in the left oviduct two complete eggs seem to be very exceptional,
if such ever occur. In Reptiles, however, both ovaries and ducts
are equally developed.”

This idea could not have been expressed more guardedly than
by the following sentence in the ‘ Dictionary of Birds,’ p. 783:
“This one-sided suppression of the organs may possibly be

* Wagener; (1) Lehrbuch der vergleichenden Anatomie, 1834.

(2) Beitrage zur Anatomie der Végel. Abhandl. Minch. Akad.
Wiss. i1. 1837, pp. 271-283.
- + Sranntus’s Lehrbuch der vergleichenden Anatomie. 1846.
{ OwEn: Anatomy of Vertebrates, vol. 11. 1866.

810 DR. HANS GADOW ON OVARIES

referable to the inconvenience that might be caused were each
oviduct to contain an egg ready to be deposited.” Mr. Gunn *
takes a reporter's liberty by substituting for ‘ inconvenience ”
various gruesome calamities, as fracture of the egg-shell, rupture
of the oviduct, and even peritonitis ! ! After thus having tried to
throw ridicule upon my generally accepted notion, he assures us
that such evil sequences need not be assumed at all because
of “the frequency with which Falconide are found with paired
ovaries, which are obviously functional.” Then he proceeds to
distinguish between several theoretical possibilities besides the
only actual condition, namely, that in which the functional organs
are those of the left side.

Not a single case is known of a completely developed right
duct, whilst the left is vestigial. Further, I withdraw the state-
ment made in the ‘ Dictionary of Birds,’ that ‘but with rare
exceptions only that [ovary | on the left side becomes functional.”
‘“‘ Functional,’ I regret to say, was there used in a loose way,
since the right ovary not unfrequently forms rather large eggs.
But strictly it should be called functional only if any of those
eggs ever became ripe, 7. e. burst from the ovary. We know
that even relatively large ovarial eggs, even those of the left
side, can undergo complete reduction ft.

Mr. Gunn, however, taking for granted that growing eggs in
both ovaries mean that both are functional, and that, although
two ovaries may not be a necessity, they must be better than one
(a principle which has produced the double-barrelled gun), has to
face the question whether one oviduct can serve two ovaries.
We are told that “there is not much evidence for or against this
supposition,” and that “‘ there seems no physical objection to the
open end of the tube swinging across the mid-line of the spine,
and grasping the right ovum of the opposite ovary with nearly
the same facility as the ovum of its own side.” One physical
objection to this amazing trick-performance may be the gut with
its loops and mesenteries, and it is at least doubtful whether the
agile tube (the infundibulum of which is most carefully anchored
opposite its own ovary) can overcome these obstacles, in spite of
the best-intentioned regulating nerve-stimulus.

Let us enquire further into the meaning of the one-sided
reduction of the female bird’s reproductive organs.

Gegenbaur { favours the size of the egg, the complete egg with
albumen and shell, as the primary cause :—‘“ Bei den Vogeln
gelangt nur das linke Ovar zu seiner volligen Ausbildung dh. nur

* T. BE. Gunn.—“On the Ovaries in certain British Birds,” P. Z. S. 1912, p. 63.
Mr. Gunn and the Rev. F. C. R. Jourdain credit the late Prof. Newton with the
authorship of the article “ Reproductive Organs ” in the ‘Dictionary of Birds,’ and
they suggest as his “German source” of original information T'aschenberg’s Intro-
duction to Naumann’s ‘ Naturgeschichte,’ re-edited in 1905, eleven years “after the
‘Dictionary of Birds.’ ; d

T Cf. A. yon Brunn: “Die Rickbildung nicht ausgestossener Hierstockseier bei
Vogeln.” Beitrage zur Anatomie und Embryologie, als Festgabe ftir Jakob Henle.
Bonn, 1882.

+ Vergleichende Anatomie der Wirbelthiere, i 1. 1901, p. 503.

AND OVIDUCILS IN THE AMNIOTA. 811

in ihm kommen Hier zur Reife, und das rechte erhilt sich nur bei
einzelnen (manchen Accipitres, Schwimmvégeln und einzelnen Gat-
tungen verschiedener Abtheilungen) fort, indess es bei den iibrigen
verktimmert. Dies steht in Zusammenhang mit der Ausbildung
nur eines (des linken) Oviductes und mit dem bedeutenderen
Volum der Hier, wodurch jeweils nur einem einzigen ein lingerer
Aufenthalt in der engen Beckenhohle gestattet ist. Hs ist also
hier die kiickbildung einer Hadlfte des gesammten weiblichen
Geschlechtsapparates von der Ausbildung des Hivolums abhingig,
und dadurch an einen héheren Zustand geknipft, dass das mit
reichlichem Eiweiss und Dotter ausgestatte Ei das sich in ihm
entwickelnde Junge zu einer bedeutenderen Ausbildung gelangen
lisst.” The correlations mentioned in these cumbrously involved
sentences are valid enough, but they do not carry the question
any further than where I had left it.

Since the ovary is the prime organ and the duct merely
auxiliary, it might be assumed @ priori that the primary cause
of the reduction was the cessation of the production of ripe
eggs on one side, whereupon degeneration of the corresponding
duct would follow, as there would be no longer any work for it.
Illicit, undesirable traffic is stopped best by cutting off the
supply, in the present case by the stoppage of ripening eggs at
the source. But, as we have seen, this does not agree with the
facts, considering the frequent activity of right ovaries, whilst
right ducts are much rarer. Further, we know that even com-
paratively far advanced ovarial eggs can be resorbed. Lastly,
we should have every right to expect birds with right, and others
with left functional organs. It cannot well be assumed that the
one-sided reduction is an inheritance from reptilian ancestors, of
which unfortunately we know nothing. Of recent reptiles only
Crocodiles and Tortoises can be studied for the sake of analogy.
These produce for one clutch a considerable number of hard-
shelled eggs which pass through both ducts. The eggs are well
protected, and there is ample room for them in the broad body
of Chelonians, and there is likewise space and safety in the long
belly of a Crocodile.

Presumably therefore the cause of the asymmetry should lie in
the typical organisation of the bird. In proportion to its body
the eggs are enormous, especially in some of the nidifugous
groups which represent the lower conditions. They could not
well produce the whole clutch at once; and they incubate
their eggs, not merely because they require several days,
even weeks, to produce the full number, but because as warm-
blooded creatures they have reached a higher state of reproductive
organisation. There is no room within the pelvis for more than
one complete hard-shelled egg, leaving aside the inconvenience
of a right and a left egg which, for argument’s sake, might be
overcome, as is actually the case with reptiles. The available
space in the bird’s belly is limited ; the longitudinal distance is
relatively much shorter than in the majority of reptiles which
are devoid of a sacrum, and the peculiar pelvis of birds is as

812 . DR. HANS GADOW ON OVARIES

broad as is compatible with the upright walking and with the
flying organisation. Most likely the broadness and the absence
of symphyses have been produced in adaptation to the eggs, but
even the distention of the belly downwards must be limited in
the bird, which is essentially and primarily a flying creature.

Well, then, let us take it that it is advantageous that one of
the ducts and the activity of the corresponding ovary should be
suppressed. Instances of asymmetry, brought about by sup-
pression of one of originally paired organs, are common enough
in the Vertebrata, and they can in most cases be fairly explained
by mechanical factors. To refer them to mere accident, to a
toss up, which then becomes established, is too shallow a mode,
although not unprecedented in morpholog ry. ‘The tadpoles of
some Anura have paired “spiracles,” others a median, the majority
a left hole. The reduction of one of the lungs of Snakes and
snake-shaped Lizards is of course directly correlated with the
shape of the body, and it appears almost optional whether the
right or the left lung should be affected, since both cases have
become established in the various groups.

If we apply the principle of elimination of all those unfortunate
hen-birds which happened to produce eggs in either side, whilst
only those birds propagate the race which happen to have only
one side in working order, this would not explain the universal
right-sided suppression, which accor ding to Gegenbaur is a
weighty argument for the monophyletic origin of the class*. If
we assumed this as a proof of their monophylism, we should
logically arrive not only at the imaginary pair of “ Urvégel” but
also at the Eve of hens, which in her case would relegate the
establishment of the asymmetry to a toss up. During the
presumably long period of dawning bird-life such a one-sided
incipient suppression must have taken place over and over again
before it was firmly established. Inheritance, if not swamped
by panmixis, might have established asymmetry, but once more
we are groping in the dark for a cause which favours the left
side. It must be a factor which is very ancient and yet does
not interfere with the symmetry of the male organs, neither the
testes nor the vasa deferentia. Since the ovaries are strictly
homologous, or rather homogeneous, with the testes, this may be
taken as another hint that the ovaries are not the parts primarily
atfected. But the male ducts are not the same as the female ducts,
therefore the latter are indicated. No factor causing the
asymmetry can be derived from the vascular system, nor even
from the vestiges of the renal portal system, by the suppression
of which birds and mammals differ from their common ancestors,
the reptiles. To refer the enlarged left ovary and duct to the

* The suppression of one lung in Snakes, etc. stands on a different footing. It
may be due to an accident or sport,as much as right- and left-clawed crabs, or
right- and left-twisted shells. The remaining lung enlarges and shifts its position
s0 as to occupy most of the space originally intended for both.—H. &.

—. -. 2

AND OVIDUCTS IN THE AMNIOTA. 813

stronger arterial supply would of course mean mistaking effect
for cause *.

A sufficient cause, however, may be the fact that a full ovi-
duct is less lable to disturb the other intestines in the left half
of the body-cavity than in the right. The primary intestinal
loops are so arranged or packed, that their bases begin on the
right whilst their apices extend towards or into the left side of
the abdomen. This is especially the case with those loops which
starting to the right of the stomach (itself mostly shoved to
the left) fill the space between stomach and vent with their
distal halves. It needs no further comment that it is the
free or apical end, and not the base of a loop where the
mesenteric vessels enter, which is displaced easiest and which
will easiest resume its original position. But this packing
from right to left is not an adaptation to, and is not produced
by the preponderance of the left oviduct. It can be traced
to a much more primitive condition, namely to the fact that
the bird’s embryo comes to rest with its left side upon the yolk,
with its curved back towards the blunt pole. In all probability
this is a truly cenogenetic feature, essentially ontogenetic ; one
of those numerous phenomena which, like the gills of tadpoles,
the allantois and placenta, are originally incidental to embryonic
life, although they may by correlated after-effects profoundly
influence even the adult organism. Obvious results of this left-
sided position of the embryo are the increasing preponderance of
the left vitelline vein ; the yolk-stalk causes the first loop of the
midgut; the stomach itself sinks in, turning the pylorus to the
right, upon which side the duodenal loop descends, and further
secondary loops of the midgut follow suit. If there are large
ceca, they likewise make their way towards the right and back
of the stomach. The allantoic bag, containing fluid only, rises
and comes to lie upon the embryo, 7. e. upon its right side.

Consequently there is asymmetry introduced at an early date,
which affects the viscera, notably the gut, and introduces a bias
in their mutual behaviour within the belly. During the growth
of the embryo, by shrinking of the yolk room becomes available
for extension of the gut towards the left side. The permanent
organs will soon—speaking from the point of evolution—establish
an equilibrium, whilst it is clear that any occasional or contingent
requirement of space, or disturbance, can be met with easiest in
the left half. Such a disturbance is caused by the periodic
growth and passage of the eggs which brook no delay. The
slightest bias will turn the scales, and now we may apply the
censorship of natural selection to its fullest extent. Left eggs

* The suppression of the right oviduct has had an effect upon the male copulatory
organ. Where such is present it is asymmetrical, although unpaired, and stowed
away in a left-sided recess of the cloaca. The act invariably takes place from the
left side, and the same applies to those birds which are now devoid of such an intro-
mittent organ.

814 DR. HANS GADOW ON OVARIES

and a left duct will be the least liable to set up complications.
Two canals may be good, but one improved way is better, and if
the traffic goes in one direction only, the other duct falls into
abeyance. If goods are still produced at the terminus of the _
obsolete line, they will deteriorate, but this does not matter if
the output of the opposite factory is equal to the demand.

So far so good, and the enquiry need not be carried further
back, if it were not for the Monotremes. Although these archaic
creatures show no appreciable difference in the size of their
paived ovaries and ducts, only those of the left side are functional.
According to Semon, Ornithorhynchus invariably produces two
eggs, always in the left side; Hchidna lays only one egg, also
left-sided. The right ovary forms numerous large eggs which
never ripen, and the respective duct and uterus are swollen and
much vacuolised during theseason. In short, Monotremes behave
exactly like certain abnormal birds, e.g. the famous Sparrow-hawk,
by the retention of an ancestral feature which is now normally
lost. Since the reduction in the Monotremes has made so little
progress, it looks as if it were but of comparatively recent date,
but at the same time so ancient as not to have interfered with
the inheritance of the full symmetry by the Meta- and Hutheria.
The Monotremes are no longer quite primitive, not even in these
organs. Their eggs have lost much of the yolk; they continue
to grow in bulk within the uterus after they have received their
keratine shell. Indeed, we cannot well imagine that, compared
with oviparous reptiles and birds, the very small egg of the
Monotremes, and the imperfect, almost larval condition of
the new-born represent truly ancestral conditions, unless—
and this is well worth further enquiry—we are prepared to
assume that in all Vertebrata the viviparous condition was
primary to one in which the feetus is surrounded by a shell and
then hatched outside the mother. If this should be the case, we
should further have to distinguish between primordial viviparity
(of which recent examples are unlikely) and secondary, pseudo-
primitive viviparity, the numerous instances of which have been,
and are still being, acquired independently: many Sharks and
Teleosts; many Urodela, even one or two of the Anura, and
many scattered cases among the reptiles, as some Chameleons
and Lacertide, Iguanide and Anguide, all the Scincide, all the
thoroughly aquatic snakes, the Viperide, and here and there
some other terrestrial kinds. But to return to the Monotremes.
Can their incipient, or perhaps arrested, asymmetry be referred to
the same embryonic conditions as those which prevail in Birds 2
The bulk of the egg is formed by the yolk, the yolk-stalk might
be strong enough to cause a disturbance, the allantois protrudes
towards the right, and the left vitelline vein preponderates.
How far, and if at all, the viscera are affected by these conditions,
remains unknown. For our purpose it is significant that there
is incipient asymmetry (functional although scarcely structural),
and that this should be restricted to the only recent Mammals

AND OVIDUCTS IN THE AMNIOTA. 815

which still possess comparatively large-yolked eggs. However,
the Monotreme embryo does not turn upon its left side, it merely
sinks into the cavity of the emptied yolk-sac, forming a
proamnion ; the bird, owing to its enormous yolk, turns over
and ultimately comes to he on its curved back; the reptilian
embryo must also sink in, but it does not turn. This turning
over, so marked a feature in the bird, may be correlated with
the conditions of incubation. Everybody knows that the
chalaze keep the blastoderm ‘on the top,” i.e. nearest the
source of warmth, against the hen’s body *. This does not apply
to reptiles which deposit their eggs in the ground, nor to the one
or two eggs in the moist pouch of the Monotremes; lastly, to the
embryo of viviparous and ovoviviparous creatures ‘“ orientation ”
towards the source of heat is not only unnecessary, it would
also be impossible because the mother changes so much her own
position by moving about.

Since no trace of functional asymmetry of ovaries and ducts
appears in Meta- and Kutheria, and as that of the Monotremes
cannot well be a reptilian inheritance (because their asymmetry
is usually marked by an enlargement of the right gonads and
ducts, e.g. Snakes and Crocodiles, both sides however being
functional), the asymmetry of the Monotremes must be due to a
departure within the Prototheria, but so slight as not to have
caused any irreparable morphological reduction of either ovaries
or ducts by the time that the Prototheria entered upon the next
higher or Metatherian stage, excepting of course the Monotremes.
If Prototheria ever laid eggs much larger than they are now, the
asymmetry may have been greater and be referable to the same
primary causes as those suggested for birds, but since the recent
Monotremes seem to be actually in the process of reducing them
now, and moreover to the last possible number, the left-sidedness
seems to be a case of mere coincidence with birds.

Some simplification of the completely double female apparatus
of the Vertebrates was bound to come; it was a matter of time,
and success depended upon the grade or height of the general
organisation of those who attempted it. Any agreement
between birds and mammals versus reptiles cannot be anything
more than convergent resemblances, at best cases of Isotely.
The classes of both Birds and Mammals have gone beyond the level
of the Reptilian organisation and they represent highest termini ;
but, although the class of Birds is by far the most specialised and
in various respects has reached seemingly unsurpassable perfec-
tion, the class of Mammals is morphologically the highest, in
spite of its still comprising such lowly, undecided types as the

* The usual statement that the hen turns the eggs over from time to time in
order to ensure the equal warming of the whole egg, now the upper and then the
lower half, implies nonsense. What the sitting bird does, is to rearrange the
position of the eggs with reference to each other, to give those now lying peripherally
an equal chance of best position near the centre. With a small clutch this is not
necessary, but with a dozen eggs the frequent rearrangement is very noticeable, at
least with thoughtful sitters.

816 DR. HANS GADOW ON OVARIES

Monotremes (¢f. loss of the nucleus of red blood corpuscles,
structure of the atrio-ventricular valves, the alveolar lungs, the
abolishment of the cloaca, the cranio-dental articulation of
the underjaw, etc.).

The Elasmobranchs show various instances of precocious in-
ventions, ahead of the times, foredoomed to failure or further
improvement, because their owners are after all but low fishes.
In some the ovary is single, unpaired, but it lies in the middle.
Some Acanthopteri1 have succeeded in producing a median duct
out of their otherwise paired ovarial sacs. Birds, as we have
seen, have suppressed one side; a clumsy mode of procedure
because it has a lopsided result and implies the reduction, by
neglect, of one of the precious gonads. It is only the higher-
eraded organisation of the Mammals which has succeeded in
simplifying the apparatus in the morphologically neatest way,

namely, by the partial fusion of the two ducts into one passage,
not only unpaired, but median, whilst the upper ends and the
ovaries remain intact and functional.

Monotremes, Marsupials, and Placentals form an unbroken,
progressive, therefore most probably monophyletic series. The
reduction in question could not be brought about until the
reptilian plan of hard-shelled eggs had given way to internal
gestation. The differences between ovlparous, OVvOviviparous,
implacentally and placentally viviparous, are questions of degree
only. ‘There is for instance no difference, to be expressed in a
few words, between the ovoviviparous fruit of a Viper and the
newborn feetus of a Kangaroo, except that the newborn reptile is
complete and must shift for itself. The point is that the young
bursts its egg, and other membranes, with the act of birth.
Whilst no newborn reptile requires maternal care, most birds do,
and all mammals are absolutely dependent on their mothers for
nourishment.

Within the Class of Birds every stage from almost reptilian
to practically mammalian conditions is represented. The
typical nidifugous birds are hatched with still a considerable
amount of yolk slipped into the belly, sufficient for the little ones
to hold out for days without food until they are bodily and
mentally strong enough to feed themselves; in many cases they
have to be shown by the parents how to do it. Next come those
which are hatched in a more or less helpless condition and must
be fed by the parents with food either in its natural state or
already semidigested. Lastly, those which are nursed with a
milky secretion of the crop. The higher Altrices or Nidicole are
born with but small remnants of yolk left, the digestive organs
having been hurried on at the expense of the others. The most
significant point, however, is that through the crop-secretion
the Pigeons have established a parallel with the Mammals in so
far as the young are fed actually with parental matter, in which
proliferation and fatty degeneration of epithelial cells plays a
great part. The analogy can be carried still further, since by

AND OVIDUCTS IN THE AMNIOTA. 817

Pigeons and some of the lower mammals this “milk” is squeezed
or injected into the young.

The Mammalian evolution has probably gone through many
stages and vicissitudes, difficult to enumerate, because there were
many factors and not all the organs changed at the same time.

1. We stajt with a hypothetical stage of Sauro-mammalia.
The hard-shelled egg, when laid, contains an already far advanced
embryo, therefore ovoviviparous. This egg was not so much
incubated for warmth as covered for protection by the mother.
The young although hatched in a reptilian condition was pro-
tected by the mother.

2. Reduction of the size of the egg. Gradual preponderance
of extra-uterine nursing over uterine gestation, made possible
and introduced by the protecting insulation, in such a way that
an abdominal incubation area was developed, owing to the
reaction of the mother’s abdominal surface by the sat-upon,
adpressed, covered egg; this resulted in hypertrophic condition
of the cutaneous blood-vessels, hence of the glands, and correlated
reduction of hairs. Incidental change of absorption of moisture
through the porous shell, enhanced by loss of its calcareous
portion— When this suppression had been well established, by
progressive inheritance, the now quite porous parchment ege had
the same chances of absorbing fluid whilst still m the oviduct.
At the same time it stands to reason that the chances of external
or brood-pouch nourishing may become prevalent. The shorten-
ing of the life-period within the egg implies the birth of an
unripe foetus. Foetal life must be taken as ending with the
bursting of the egg, no matter whether this act coincides with
the moment of parturition, or whether it happens some time
after the egg has been “laid.” In either case it coincides with
the cessation of any further possibility of function of yolk-sac
and allantois.

We must further assume that the actual length of time re-
quired for the production of a young animal is the same in
equal-sized creatures from the beginning of segmentation until
it is independent. Unless we assume this, the argumentation
would become too complicated.

Let us say that it took 50 days from impregnation until the
Sauro-mammalian youngster was ready to face the Permian
world. This means:

50 days of ovoviviparous, internal uterine life ...........................(Sauro-mammal),

40 days uterine and 10 days incubation-life within the laid ege......... (Hypotheria).
BO » LON sp and 10 days nursing in pouch

(Prototheria).

20 ;; » 5? ,, 2 25 5 3 (Monotremes).

8 2” oy) 0 ,, oy) 42 ” o) (Opossum),

The suppression of the incubation-life marks the Metatherian
stage just as sharply as the introduction of incubation of a
foetus marked the early Mammal stage. The adaptation to the

818 DR. HANS GADOW ON OVARIES

new invention of pouch-life and nursing caused the development
of an entirely new category of features—of features which were
not required either by the foetus or the adult, therefore larval,
é. g. &® suctorial apparatus with its far-reaching incidental in-
fluences upon the future adult structures.

In these respects viviparous reptiles, Hypo- and Prototheria
culminating in Monotremes, and Metatheria culminating in
Marsupials, represent a continuous progressive series, with a
logical terminus characterised by the enormous preponderance
of extra- over intra-uterine development. Compared with these
terminal Marsupials the Eutheria seem to be totally different,
provided we take as their type those which are born complete,
in this respect lke the hypothetical Sauro-mammals, the whole
of the ‘50 days” being intra-uterine. And yet the Eutheria have
with certainty passed through the same Metatherian stage as
have the Marsupials, and this Metatherian stage comprised,
besides others, the following features*: Truly viviparous ; allan-
toic placenta; marsupium; diphyodont teeth, the same two
middle series of a total of prelacteal, lacteal, permanent and post-
permanent sets; nipples; semi-cloaca; absence of a corpus
callosum.

To arrange any one of these features into successive morpho-
logical stages is comparatively easy, but it does not follow that
these represent exactly the phylogeny of the groups, because of
the complicated correlations with other organs which by no
means keep step with each other, neither in the same species nor
in the greater groups. Some are precocious, even hypertelic,
while others lag behind.

Just as to the large egg of the truly oviparous Sauropsids
albumen (more watery but less fatty yolk) is added, before it
receives its calcareous shell, so in the Monotremes fluid is added
to the contents of the egg, but with the remarkable difference
that fluid matter is taken into the yolk-sac itself by osmosis
from the uterine walls, after the keratine shell has already
been developed. This process is correlated with an un-
doubted previously acquired reduction of the amount of ovarial
yolk, and is as much a secondary process as the loss of calcareous
matter in the parchment-like “‘ keratine” shell.

As Semon has shown, the whole shell-enclosed egg multiplies
its size during its passage through the oviduct. This mode of
growth finds a curious parallel analogy in various Lacertilia, the
parchment-shelled eggs of which grow considerably after they
have been deposited.

Whilst the Sauropsidan allantois comes to surround the whole
yolk-sac and also nearly the whole of the albumen, so as to
spread over most of the inner surface of the egg, the Monotreme

* Of. R. Semon: “ Monotremen u. Marsupialier,” in Zoolog. Forschung. Australien,
ii. 1894-97; further, J.T. Wilson and J. P. Hill’s papers in Q. J. M.S. 1897, 1898,
1900.

AND OYIDUCTS IN THE AMNIOTA. 819

allantois and yolk-sac balance each other. ‘The shell-enclosed egg
is for a short time transferred into the marsupium (cf. Semon,
Kchidna ; not into a bursa as was imagined by Klaatsch, Gegen-
baur, and others), which secretes fluid (the “‘ nutritive sweat” in
Gegenbaur’s unfortunate diction), and this can be taken up by
the embryo through the porous shell. This may be the reason
why the shell is soon cast off.

In Marsupials the shell is at first still present, but soon absorbed
within the oviduct. The egg-membranes, etc., of the embryo
establish no structural communication with the uterus. The
ovarial yolk is much reduced. But the yolk-sac becomes en-
larged, as in the Monotremes, still occupies a great portion
of the inner egg-surface, and has established intimate contact
with the serosa. The allantois, being independent of the expansion
of the ccelom, which results in the driving away of the yolk-sac
vessels from the somatopleure, establishes a villous placenta.
Such must have been the condition of the Metatheria.

In Perameles the allantois still reaches the surface, where it
is very vascular, and fused with the serosa, a truly respiratory
arrangement. The placenta being lost in most other Marsupials,
the allantois reverts to its primary function of urinary receptacle,
although apparently late during the feetal life.

The Metatherian stage may therefore be characterised as one
in which the posterior of the two bags, the allantois, has super-
seded the previous attempts of placentation by the yolk-sac.
The new placenta was perhaps not advanced enough to prevent
the foetus from being born soon after the limited amount of yolk
was used up. Certainly it did not pass beyond the non-deciduous
stage, and it never reached the extent of even the lowest recent
Eutherian placenta. Yet one effect of this incipient organ must
have been to render the feetus less independent than that of a
viviparous reptile. It had therefore still to be transferred into
the marsupium, there to be kept moist and suckled in as
premature a condition as the Monotreme.

Two opposite tendencies are inherent to this stage. One
palingenetic, to give birth when the yolk is used up; the other
cenogenetic, to prolong the retention of the fetus because of the
compensatory, respiratory, etc. advantages incidental with a
placenta. Obviously the Metatherian stage was a_ half-way
house at a parting of the ways to further improvements, leading
to Marsupials* and to Placentals.

* We have here an instance of the well-known fact that Group-names based upon
single anatomical features are mostly unsuitable for classificatory purposes. ‘The
taxonomic value of these characters may be good enough, but they are not diagnostic.
If such names were used merely as labels without much intrinsic meaning, well and
good, but even the best of us cannot, on occasion, resist taking their face-value for
full value. There are Mammalia Implacentalia with a placenta, and we now know
that the young Echidna does not he in a bursa. Odontornithes are a valueless,
heterogeneous assembly, and overcontideuce in “ Ratite” was responsible for
having branded Hesperornis as a “swimming Ostrich.” ‘“ Mammalia” is for-
tunately an excellent term, although invented before Monotremes were known.

820 DR. HANS GADOW ON OVARIES

To judge from analogy the new organ should continue straight
on, leading in this case through a diffuse and cotyledonary to a
solid, and from a non-deciduous to a deciduous placenta, ter-
minating in the birth of the young in an utterly helpless con-
dition. “As a rule the non-deciduous and diffuse placenta i is found
in those mammals which at birth require least maternal care.
The latter implies greater mental capacity. But we also know
from analogy that a recent invention—although it may be
capabie of much perfection—is not always kept up if the old
string to the bow is still capable of further adaptation. Where
both alternatives are good, workable, and capable of being
improved, it may take a long time to settle which is after all the
better of the two.

What has induced some Metatheria to neglect the further
development of their placenta and to intensify the marsupial
alternative? The morphological and physiological momentum

was surely on the side of improving placentation. There
must have been some external, environmental influence, and
it was Dollo’s brilliant suggestion that arboreal life was the
underlying cause. His further explanation is less acceptable,
that the climbing habits have caused premature births, which
have become habitual, and that the hapless foetus had therefore
to be nursed and located in a marsupium, the bursa, or rather
multiple burs, beimg no longer sufficient.

I think the new conditions to be faced by intense arboreal
life have acted somewhat differently, especially if, as we assume
now, all the Metatheria were already possessed of a marsupium*.

I venture to suggest that those Metatheria which were driven
to arboreal life, had to solve the question what to do with
their young. They had to carry them, and being already pos-
sessed of the pouch, this was not only retained but intensified
by prolonged use. That the facilities offered by such a pouch
are great, is shown by the still existing numerous and much
diversified Marsupials, and by the fact that some at least, e. g.
Opossum, have reduced the uterine gestation almost to the
conceivably lowest limit. Whilst these arborealised Metatheria
could not afford to leave their young behind, it was different
with those other Metatheria which during the Cretaceous epoch
somewhere “‘in the larger and more effective workshops of the
North” (to apply here one of Darwin’s happiest expressions) went

* Klaatsch’s view of the correlation between Bursee and Marsupium had to be
considerably modified by Semou’s discovery that the Monotreme does not lie in a
bursa but in an already typical, though transient, marsupium. Moreover, there are
unmistakable vestiges of marsupial - muscles, other than compressores Mamm2,
in various Placentals. Lastly, each teat with its areola does represent a bursa
(Klaatsch), but if each bursa had been a brodd-pouch instead of merely the result of
the attachment of young, the possession of a series of burs from near the vulva
up to the armpit would imply an impossible condition. In fact, bursz or nipples
can be formed independent of a marsupium and in places whereto no such organ
can have extended.

AND OVIDUCTS IN THE AMNIOTA, 821

in for that further mental development which is intimately
correlated with the origin of the corpus callosum. Competition is
most effective if backed by brain-power. Where these dawning.
Kutheria spread the other Metatheria had to go to the wall, or
were forced to get out of reach—into the trees. Those which,
already arborealised, found their way into countries like Australia,
where the Eutheria could not follow in time, found there
nothing to prevent them from becoming again terrestrial, as
Dollo has shown so conclusively. The chance of reviving the
degenerated placenta was, however, gone, and the Marsupials are
now, in spite of the retention of various low characters, the most
specialised subclass of Mammals.

The Edentates present a striking analogous case : being with-
out exception arboreal or fossorial, they ‘also seem to have got
out of the way of mentally higher, dangerous Kutheria. They
have survived, whilst those that went in for heavy bulk and
armour were not a great success. At least the American Eden-
tates seem to have come into contact with the dangerous
Kutheria long after they themselves had become typical Eutheria,
so that the former possession of a marsupium could avail them
nothing.

A few words about the Eutheria. They could afford to part
with their young for a time and to suckle them intermittently,
an advantage which needs no further comment than that it
would be impossible without higher mental faculties.

Great adaptive changes have been wrought; every chance has
been tried, now here, then there. Nearly all the Non-deciduata
are at birth precocious, able to look after themselves, no matter
whether the placenta is diffuse, cotyledonary or zonary even,
namely the aquatic Cetacea and Sirenia, and the Artio- and
Perissodactyles, the exception being the Lemurs. The Deciduata
are at birth helpless: Rodents, Insectivores, Carnivores, Chiro-
ptera, Tarsius and Simiz, except the Elephants.

The egg-laying Monotremes, the implacental Marsupials, and
the Mammals with the most advanced placenta, are born in a
condition as helpless as the highest nidicolous birds, whilst the
avowedly low non-deciduous and diffuse placenta of the Mare i
sufficient to produce a young which is as precocious and in-
dependent as a lizard.

Can these facts be brought into line? What is here primitive
and what is a roundabout return to apparent but not really
ancestral conditions 2

Proc. Zoot, Soc.—1912, No. LV. 5d

822 DR. F. E. BEDDARD ON A

49, Contributions to the Anatomy and Systematic Arrange-
ment of the Cestoidea. By Frank E. Bepparp, M.A.,
D.Sc., F.R.S., F.Z.S., Prosector to the Society.

[Received April 30, 1912: Read June 4, 1912.]
(Text-figures 113-121.)
VI. On aw AsexuAL TAPEWORM FROM THE RopENt, Fiber zibethicus,

showing a new form of Asexual Propagation, and on the
supposed Sexual Form.

INDEX.
Page
ST AAELELA DUD Sw Pe eRendrdond: cnowedederonondeeeeaadde coosaopacenesed™ |ce!13)
Systematic :
Urocystidium gemmiparum, gen. et sp. n....... 840
General Résumé and Systematic Position ............ 849
Anatomical Summary ...........ccee cece eeeeeeeeeeee ss 850

I received in February of this year two complete tapeworms
which were found in the hepatic duct of a Musquash (fiber
zibethicus) which died in the Society’s Gardens. The two worms
lay together in a part of the hepatic duct just before it receives
the bile-duct, which was much dilated by the parasites. Although
these two worms were very different in appearance, I believe them
to be respectively the sexual and asexual form of the same species.
T shall give reasons for this conclusion in the course of the
following description of the two individuals.

(1) THe AsexuaL Form.

The general appearance of this very remarkable worm is
shown in text-fig. 113 (p. 824), which illustrates its most
remarkable peculiarity, viz., the possession of two series of
what appear to be buds at one end of the body. It will be seen
in the course of my description of this “ Cysticercoid” that it is
without any doubt to be regarded as an asexual form. It does
not, however, follow that the mature worm found with it is a
further stage in its development. On the contrary, indeed; for
as a mere matter of guesswork, the assumption would be the other
way. Still, I believe that I shall be able to prove that the two
worms are stages of the same species, in which event we have the
very remarkable fact of both the sexual and the asexual form
coexisting in the same host, and, moreover, in a situation where
one would expect to meet with sexual forms only. The Hymeno-
lepis* of the mouse is a partly, but not an entirely, parallel
instance.

At first it seems likely, from an inspection of the figure

* HH, murina.

NEW ASEXUAL TAPEWORM, 823

annexed, that the wider end of the worm is the scolex and that
the narrower end is the posterior extremity where the proglottids
are being shed. The dilated extremity suggests a scolex not
altogether unlike that of the genus Dasywrotenia, which I have
lately described to this Society *, and which is characterised by
an unusually swelled scolex. Furthermore, there is a slight
diminution in the diameter of the worm towards the narrower
end, which again conforms to this view, which, as a matter of fact,
I at first held myself. It appears, however, that the reverse in
fact is the case, and that the narrower end is the scolex end.
The opposite extremity therefore is, as I suppose, to be compared
to the persisting bladder of the bladder-worm stage of the
Cestode. To the naked eye, and even on examination with a
lens, the scolex end presents every appearance of being the proli-
ferating end of the body, for. the last segments appear to be
slightly incurved, as 1s so commonly the case at that end. It was
not until I studied this region by means of a series of horizontal
sections that I was able to discover its true relations. And even
now certain important details are wanting.

T could find, indeed, no armed scolex, nor any trace of suckers ;
if the worm is a member of the Pseudophyllidia and has there-
fore only bothria, these may easily have escaped attention in such
sections, which would not be suitable for their display. I cut the
sections, in fact, under the impression that I was dealing with the
posterior end of the body, and without making a sufficiently ex-
haustive survey of the external characters. The main arguments,
therefore, which lead me to the conclusion that this is really the
scolex end are firstly the mode of imbrication of the proglottids,
and secondly the presence of large pigment granules, a condition
which would hardly be expected at the posterior end of the body,
but which is not uncommon among tapeworms in the scolex.
As to the imbrication of the proglottids it seems to me to be
necessary to regard a segment which overlaps the next one as
being anterior to it in point of origin, and therefore lying to the
scolex side of it. Judged by this conclusion, the narrower end of
the body of this very remarkable tapeworm is the scolex end.

There is no evidence that a scolex has been lost. On the
contrary, the body ends here in a slight median elevation, which
is quite unlike the termination of the body were this the region
of the detachment of proglottids. This little elevation, however,
bears no particular likeness to a scolex, and there are certainly no
suckers or hooks to be seen anywhere. Nor is there any neck, or
break of any kind, between this region and the first obvious pro-
glottid. In this latter, moreover, the lateral and transverse tubes
of the water vascular system are as large as in the more posterior
segments, and do not end in a coil such as is so frequent in the
anterior part of this systemin other worms. It may be, of course,
that the scolex is in this genus a transitory affair, as it has been

* P.Z.S. 1912, p. 677.
55*

824 DR. F. E. BEDDARD ON A

believed to be (though perhaps hardly now) in certain other tape-
worms. Or, on the contrary, we may have here a feebly developed
head like that of Zigula. But the great breadth of the body of
the present worm is not perhaps in favour of such a supposition.
Had there been a very narrow anterior neck, some suggestion of
this kind might have been put forward. Anyhow, the general
characters of the anterior region of the body of this Cestode are
as described above. We shall recur to their more minute anatomy

Text-fig. 113.

The upper figure represents the entire asexual form of Urocystidium gemmiparwn
enlarged by about one-third. The lower figure is the posterior end of the same
individual more magnified. For explanation see text.

later. From this point to the extreme posterior end are very
many proglottids. They are very short and wide, and do not
appear to differ very appreciably in length in different regions of
the body. The greatest diameter is something like 4mm. This
is at and towards the posterior end of the body. Quite anteriorly,
the diameter is not more than 25mm. The length of the entire
worm is about 80 mm.; the small posterior bladder is 3 mm. long

NEW ASEXUAL TAPEWORM. 825

by 2°5-3 mm. wide. The proliferating end of the worm ceases
abruptly with the commencement of the bladder. This region,
which bears a great many lateral buds, shown in the text-figure
referred to, measures 17 mm. The region of the body which
bears these presumed buds (we shall consider their nature later)
does not differ much from the preceding section. It looks,
perhaps, a little more transparent, but it is divided up exactly in
the same way into proglottids, which are of about the same
dimensions as those elsewhere in the body. The buds form a row
on each side of the body and are quite lateral in position ; they
- are not at all arranged in order of growth, though representing
very many stages. That is to say, the older buds alternate with
younger and older forms. Nor does every proglottid bear a bud
or pair of buds (one on each side). Between any two buds there
was often a variable number of proglottids without any trace of
budding at all. This, indeed, is necessary; for the large size of
the buds would prevent their proper growth upon immediately
adjacent proglottids. I counted altogether 17 buds upon one
side and 18 upon the other. But I may have omitted one or two
in each case; for it is a little difficult to fix the actual first
appearance of a bud. They begin, in fact, as an only just recog-
nisable rounded swelling of the edge of a proglottid. Sometimes
the swelling includes two proglottids. There is no question
whatever of the continuity of these buds (as I regard them) with
the parent stock; they are most plainly outgrowths therefrom.
In later stages the minute round bosses at the edges of the
segments swell into spherical, largely transparent, vesicles, of
which, as the text-figure (text-fig. 113) shows, the size varies—
possibly according toage. Later still, these bladders show a young
worm growing out from their distal extremity, which is usually
of considerably less diameter than the bladder which is attached
to the parent stock. It exhibits numerous wrinkles which I do
not definitely regard as the delimitations of proglottids. The
longest of these presumed young worms was about 4 mm. in
length; the bladders reached a length of 2 mm. The exact
number of these more developed buds is faithfully reproduced in
text-fig. 113.

§ Structure of the Parent Stock.

I have investigated the anatomy of the worm by transverse,
longitudinal, and sagittal sections. From an inspection of the
former (see text-fig. 114, p. 826) the depressed form of the body
-was obvious, the diameter of a section being about seven times
its depth.

At the thinner anterior end of the body, not far from the actual
extremity, transverse sections showed that the medullary region
of the proglottid was not more than two-thirds of the diameter of
the cortical region. ‘The two were plainly marked off from each
other by delicate transverse fibres forming a very thin layer, and

826 DR. F, E. BEDDARD ON A

yet quite unmistakable. The same proportions and the same
structure were visible as plainly in the wider posterior region of

Text-fig. 114.

Part of a transverse section through a proglottid of the asexual form of
Urocystidium gemmiparum.

A, spaces in middle of proglottid with darkly staining walls referred to in text.
lm. Longitudinal muscular layer of cortex, within which is seen delicate
transverse layer of muscles. .s. Nerve-cord. .d. Dorsal excretory tube,
».v. Ventral excretory tube.

the body. The medullary substance is quite well stained by hema-
toxylin and has the usual homogeneous appearance. ‘There is,

NEW ASEXUAL TAPEWORM. 827

moreover, in the present worm the usual meshwork arrangement
of fibrils round spherical masses of the homogeneous ground-
substance such as occurs in so many tapeworms, and is often a
very coarse and thus more obvious network. The homogeneous
ground-substance was, however, traversed by two sets of fibres,
which are, I presume, muscular fibres. These are absolutely at
right angles to each other and not at all closely set in either case.
The dorso-ventrally running fibres are nearly always wavy in
their course, the transverse fibres are quite straight—a matter of
different states of contraction, as I imagine.

The cortical layer is easily divisible into two regions, which are
of about equal diameter. The innermost of these is that occupied
by the bundles of longitudinal muscular fibres. The bundles of
muscular fibres are two or sometimes three deep, and each bundle
contains a large number of fibres, which, however, are not very
closely pressed together. In addition to these bundles of rather
slender fibres there lie on either side, between the nerve-cord and
the lateral margin of the proglottid, two bundles of considerably
stouter fibres which run continuously from segment to segment
and are closely associated with a longitudinally running cavity.
These bundles appear on occasions to lie actually within the cavity.
This tubular cavity is not at all like the tubes of the water
vascular system, and it lies within a very lax tissue. Iam unable
at present to suggest its nature.

The accompanying text-figure (text-fig. 115, p. 828) illustrates a
section through the hinder part of the worm not very far in front
of the budding region. ‘This region shows very plainly a system
of irregular spaces in various parts of the body belonging to the
water vascular complex. It may be remarked first of all that the
dorso-ventral diameter is greater here than in the more anterior
part of the body. The structure of the vertical and medullary
regions is the same, but the lower central region of the medulla
actually forms a cavity (text-fig. 114 A) which I do not think to
be artifact and which may be a forward extension of the cavity
of the terminal bladder, though I have not proved the fact, from
an unwillingness to sacrifice the specimen altogether. The in-
terior lining of the space was, however, so very strongly stained
as compared with the surrounding tissues that it appears to me to
be practically certain that the staining fluid gained access through
the two cut ends of the piece of worm, which was stained en bloc.
This, of course, argues a continuous central cavity.

It was quite plain that this cavity was quite distinct from that
of the transverse water vascular trunk. The peculiar central
cavities occupying nearly the middle of a segment here and there,
referred to later (see p. 830) in describing sections made from
earlier segments in the body, are present here and are again not
to be confused with the central lacuna. Their thick walls show
no trace of an opening into it. In addition to the main trunks
of the water vascular system, which are disposed here as they are
elsewhere in the body, there are scattered irregular spaces which

828 DR. F. E. BEDDARD ON A

seem to belong to the same system. These consist of larger but
very short stems with finer branches. I have not traced any con-
nection between these tubes and the main stems of the excretory
system; but I can, nevertheless, hardly doubt that it exists.

Text-fig. 115.

EV.

= a es

A more highly magnified section through excretory vessels and adjacent
structures of the asexual form of Urocystidium gemmiparum.

d.v. Dorsal vessels surrounded by circular muscle-fibre. m. Layer of transverse
muscle-fibres separating the cortex (to the right) from the medulla (to the
left). 2. One of several tubes forming the excretory network and lying, as is
shown, in the cortex as well as in the medulla. ¢.v. Transverse commissural
vessel forking to join the ventral vessel (v.v.), which is also bifurcate to receive
the branches.

These ramifications lie in the cortical layer as well as in the
medullary (text-fig. 115, 7). In one case I observed a narrow
duct leading from a wider space to the exterior—at any rate, most

NEW ASEXUAL TAPEWORM. 829

undoubtedly to the cuticle, and in other cases it appeared to me
to be highly probable that the ramifications of these tubes do
open on to the exterior here and there; they at least invade the
outermost layer of the cortical tissues.

The water vascular tubes are large in this species, and not far
from being equisized. There are, as is the rule, two upon each
side of the body. As is so often the case among the Cestodes, the
two tubes, instead of being respectively dorsal and ventral and
superposed, lie side by side. As is also quite usual among these
worms, the two tubes, dorsal and ventral, differ somewhat in
structure. I consider that the slightly smaller and more muscular
of the two tubes is the dorsal vessel, from the analogy of other
forms; in the present species, therefore, the dorsal vessel is
situated internally to the ventral. They are, however, quite in
the same straight line for the most part and also in the same
straight line with the nerve-cord. But, as a matter of fact, the
more muscular dorsal tube lies in a corkscrew fashion, which is
very plain when the proglottids are viewed in longitudinal and
sagittal sections, and thus that tube in cross-sections appears to
lie now rather dorsal of, and at times rather ventrally of, the
ventral and more external vessel. Elsewhere they are more
directly in the same straight line. It is a remarkable fact that
both of these vessels seem to be quite as large or very nearly so
in the most anterior segments of the body. There is no great
increase in calibre posteriorly.

The general relations of these two vessels are shown in text-
fig. 114. ext-fig. 115 is a more enlarged representation of the
two tubes and the adjacent parts taken from a section of the
posterior region of the body. ‘Their detailed structure is there
illustrated. In his account of the Cestoidea in Bronn’s ‘ Thier-
reichs,’ Prof. Braun remarks—‘ Ob in der Wand der Sammel-
rohren auch noch Muskelfaseren vorkommen, ist noch strittig.”
The figure just referred to is, as I think, quite conclusive as to the
presence of a particularly thick layer of circular fibres constituting
the greater part of the wall of what I regard as the dorsal vessel.
This muscular layer appears to wax and wane in thickness from
place to place. But it is always thick and thus very apparent.
The actual tube is lined with a chitinous cuticle which is also often
very thick and everywhere to be seen. Outside of the layer of
circular muscles are abundant nuclei, massed round the tube. I
did not observe any nuclei among the circular muscular fibres,
which are, indeed, very closely contiguous. There appear to be
no intrinsic longitudinal muscles to assist in the widening of the
tube again after contraction by the circular muscles; but there is
a mechanism which takes the place of such fibres to serve in their
stead.

In the neighbourhood of the dorsal excretory vessel the dorso-
ventral muscles form groups here and there of rather thicker
and more closely adpressed fibres, which, as it were, tie to or
suspend the dorsal vessel from the cortical layer ; it seems certain

830 : DR. F. E. BEDDARD ON A

that the contraction of these would dilate the lumen of the dorsal
excretory tube. These fibres, which exist both dorsally and
ventrally, have a somewhat fan-like arrangement—that is to say,
they converge upon the vessel from above and from below. It is
these various muscular layers which cause so great a variation in
the lumen of the excretory tube from place to place.

The ventral vessel differs entirely from the dorsal by the entire
absence of any muscular layers and by the possession of only a
very thin lining cuticle. It follows, therefore, that the variations
in the calibre of this tube are not so great, and when they do
occur seem to be due to a squeezing by the surrounding tissues.
The ventral vessel also runs a much straighter course than the
dorsal without any marked corkscrew-like windings exhibited by
the latter. There is a transverse vessel in each segment which
has a rather peculiar relation to the ventral longitudinal tube
into which it opens on either side of the body. A little way before
reaching the dorsal tube the transverse vessel splits into two
halves, which embrace the dorsal vessel and open separately into
the ventral vessel. This is partly shown in text-fig. 115.

We have now to deal with a series of sac-like bodies forming
closed cavities of very problematical nature, which lie in the
middle of many segments. There is by no means one of these
sacs to each segment, and thus they are very far from consti-
tuting a tube running without intermission through the body.
I compare them, however, later in this paper to a tube found in
the young buds (see p. 833). But, as a matter of fact, one of
these sacs may actually extend through two proglottids, so that
there is a hint of a formerly continuous structure such as occurs
in the buds. Asa rule, however, they appear to be limited to a
single proglottid, of which they occupy sometimes the exact centre,
at other times pushed rather to one side. They lie close to the
transverse water vascular vessel. J cannot find that these sacs
have any connection with any other spaces. They appear to be
perfectly isolated. The first of the series occurs very early in
the body—if not in the very first segment. These sacs have a
lining cuticle surrounded by a single layer of epithelial cells of
which the borders between the individual cells are not clear.
The nuclei, however, are very obvious and large. There are no
muscle-fibres at all apparent encircling the sacs. They often
appear crumpled in transverse sections, and occasionally seem
to be Y-shaped. They are impossible to miss in a series of
sections.

The only remaining characteristic of this worm upon which I
am able to report is the structure of the ‘“‘ head” end of the body,
which has been already briefly described above. In the place of
a scolex there is a small projecting process, no larger than the
rostellar process of many unarmed tapeworms. ‘This arises from
the middle of what may be termed, for the present, the first seg-
ment of the body, which is distinguished from those that follow
by its rather less breadth and greater length. It would seem to

NEW ASEXUAL TAPEWORM. 831

be really composed of two segments, if we consider the condition
of the water vascular system therein. The two main longitudinal
vessels of this system extend well forwards into this compound
segment, and the outer ventral vessels are connected by two
transverse vessels, thus indicating, as I suggest, the double
nature of this apparently single segment. Towards the centre
of the specialised anterior segment is a large group of deeply-
pigmented oval bodies, which appear to be special pigment-cells.
They are as large, or nearly so, as the calcareous corpuscles which
are very abundant in this worm. In the middle of the concave
anterior border of these two fused segments is quite a small
conical projection, than which nothing else at all comparable to
a scolex exists. There is no sign of any breakage at the summit
of this which might suggest that a scolex had been accidentally
detached.

It is a somewhat remarkable fact that these conditions (or, at
any rate, something very near to them) occur in the genus
Dioicocestus; for, as will be pointed out later, the genus with
which I am concerned in the present paper shows some likeness
to the Acoleide and even to the genus Dioicocestus, which is of
that family. In Dioicocestus acotylus, Fuhrmann * has described
and figured the practical absence of suckers and the very rudi-
mentary condition of the rostellum—a state of affairs which is
not found in the other species of this genus. The figure referred
to also shows—as I have described in my species—the absence
of any marked diminution in calibre of the body at the head end.
But it must be remembered that the Diotcocestus was a fully-
developed sexual worm.

§ Structure of the Buds.

It now remains to consider the minute anatomy of these out-
growths and their nature. As to the latter query, there are, as
it appears to me, only two alternatives. These outgrowths must
be either “tentacles” of the nature of the processes known in
Hymenolepis villosus, or young worms budded off from the
parent stock. There is very little, if indeed anything, to be
said in favour of the former view. It is true that a large out-
growth of the body to form a tentacle might have very much the
same structure as the body, even to possessing branches of the
excretory system and lateral nerve-cords. But there would
hardly be a practical identity of structure such as I shall point
out in detail later. Furthermore, the growth of processes of the
body might be expected to be more regular than are these out-
growths (cf. text-fig. 113), and above all a gradual freeing of
themselves from firm contact with the body, as shown in the
figure, is precisely what we should expect with budding offspring,
which, as it appears to me, is the obvious and only way to in-
terpret these appendages of the fully grown worm.

* Zool. Jahrb., Abth. f. Syst. Bd. xx. 1904, p. 131.

832 DR. F, E. BEDDARD ON A

I selected for investigation, by means of transverse sections,
the most adult-looking of the young budded-off worms, in which,
moreover, segmentation was obvious. I have already referred to
this individual in my account of the budding process and need
not redescribe its external characters here. The wider end
attached to the body of the parent forms a bladder with rather
thick walls. ‘This I take, therefore, to be the posterior end of
the young worm. The opposite end is very much thinner (as 1s,
of course, usual among tapeworms) and represents the scolex end
of the animal. I could observe no trace of hooks anywhere on
this scolex, nor suckers of any kind. This, however, is not sur-
prising in view of what I have already described in considering
the conditions observable in the parent stock. At about the
middle of the worm a transverse section shows the appearances
represented in text-fig. 116. The cortical layer is thick—about
the same diameter, or in places rather more than the diameter,
of the medullary region. The inedullary region is plainly marked
off from the cortical by transverse muscular fibres of delicate
constitution.

There are several parallel fibres in this layer, which is altogether
of some width, the individual fibres being widely separated. The
parenchyma of the medullary region is as usual, and contains
many nucleated cells, generally triangular in form, connected
with processes at the angles. These cells get to be particularly
numerous near the water vascular tubes, which will be referred
to later. There are also heaps of nuclei visible in some sections,
which I take to be the rudiments of the sexual organs. To one
side of the body the medullary region seems at first sight to be
invaded, as it were, by the cortical region, thus producing an
asymmetry in the transver se section, which i 18 quite apparent in
the accompanying text-figure. This asymmetry is associated with
the presence of a tubular organ (text-fig. 116, «), which will be
described in detail on a subsequent page. ‘This occupies a good
deal of space, and round it are large bundles of stout muscular
fibres, quite similar to those which occur in the cortical layer.
It is very easy, however, to observe that the transverse layer of
muscles bounding the medullary parenchyma is continued on
either side of this differentiated mass of tissue and ends laterally
at a point asymmetrical with its ending on the other side of the
body. I therefore regard this region as belonging to the medullary
part of the body. The cortical region is distinguished from the
medullary (other than that section of the medullary region just

5
referred to) by the bundles of stout longitudinally running fibres.

These bundles form a layer Sounve Eavielr outside of the transverse
fibres; but there are much smaller bundles and isolated longi-
tudinal fibres extending some way towards the exterior of the
body.

In the medullary region lie the water vascular tubes and the
nervous system. All of these lie in the same straight line with

each other and with the tubular organ already referred to, and

NEW ASEXUAL TAPEWORM. 833

this line is the longer diameter of the cross-section ; the tubes
ete. are thus laterally disposed with regard to each other. It is
necessary to emphasise what may appear to be a statement of the
obviously true, since in other regions of the body the mutual
relationships of these several organs and systems are a little
changed,

Text-fig. 116.

Transverse section through middle of strobila of oldest bud of the asexual
form of Urocystidiwm gemmiparum.

lv. Eixeretory tubes. 2. Nerve-cord. «x. Tube of doubtful significance
described in text.

In all other tapeworms known to me the water vascular tubes
of the two sides of the body are symmetrical with each other in
point of position within the medulla and distance from the lateral
cortex on either side, They are also generally correspondent in
size, though not always regularly so throughout the whole body,
In the present species the water vascular tubes are asymmetrical in
both these particulars. The tube of the one side is considerably

834 DR. F. E. BEDDARD ON A

larger than that of the other and is relatively asymmetrically
placed. ‘This latter fact is doubtless due to the bundles of longi-
_ tudinal fibres surrounding the large lateral tubular organ. This
structure has, as it were, pushed the nerve-cord and the water
vascular tube of its side towards the centre of the body. Indeed,
the vessel in question lies very nearly in the actual centre of the
section and quite in this position shortly before and during its
connection by the transverse vessel with the tube of the opposite
side of the body. These transverse vessels are at regular intervals
and quite easy to see as in the adult worm. I did not observe
any other branches of the water vascular tubes than these.

These sections from the middle of the body of the young worm
also show very plainly the remarkable constitution of the nervous
system in this Cestode. One of the two lateral cords (text-
fig. 116, 2) is very obvious indeed, lying between the water vascular
tube of that side and the tubular organ of doubtful meaning.
It cannot possibly be missed in these sections. But on the opposite
side of the body I could find no trace whatever of any nerve-cord
even of smaller size than the one represented in the text-figure.
I feel confident that a failure to observe a cord in this situation,
were it present, would be difficult. The medullary tissue is so
clearly differentiated by the stain (iron hematoxylin*) that
the nerve-cord, if of any size at all comparable to that of the
opposite side of the body, would stand out quite prominently.
This state of affairs is undoubtedly very anomalous and hard of
explanation. For we shall see later that the two nerve-cords are
both quite recognisable towards the end of the body.

Finally, this transverse section shows to one side a large tube
cut across, to which reference has been made incidentally once or
twice already. It is situated in the centre of a group of bundles
of longitudinal fibres, which bundles are like those in the cortical
layer of the body in every way, including their rather sparse
scattering round the centrally placed tube. The bundles in the
cortical layer are not densely pressed together as they are, for
example, in Dasyurotenia robustat. ‘The tube itself is of con-
siderable size and is lined by a thick layer of chitin, which is
stained precisely like the chitin which covers the body of the
worm externally. Outside of this is a single layer of stout longi-
tudinal muscle-fibres, and then a deeply staining layer of elongated
cells, interspersed among which are small bundles of stout longi-
tudinal fibres. The structure of the tube is in fact almost exactly
that of the outer layer of the body, only differing, indeed, by the
intercalation of the groups of stout longitudinal fibres among
the cells.

We shall now consider the course and structure of this tubular
organ throughout the body of the young worm. The identity of
structure with the outer layer of the body which this tube
shows in the middle region of the body, is rendered intelligible

* I am indebted to Dr. Plimmer, F.R.S., for the use of this reagent.
+ See Beddard, P. Z.S. 1912, p. 684, text-fig. 97, lm.

NEW ASEXUAL TAPEWORM. 835

by the fact that in two places, one behind the other, it communi-
cates with the exterior by a narrow duct (text-fig. 117), One of

Text-fig. 117.

Two consecutive sections from another region of the same bud as that which is

represented in text-figure 116, to illustrate the opening of the tube a on tothe
exterior.

In the upper figure the orifice (0.) on to the exterior is shown and the
commencement of the invagination leading towards the tube x.

Tn the lower figure the outgrowth of the tube a to meet the invagination is seen.

mn, Nerve-cord. 7.v. Water vascular tubes,

836 DR. F. E. BEDDARD ON A

these openings is in the bladder region and the anterior one a
very little way in front of this. Both orifices are dorsal or ventral*
in position—at any rate not lateral,—but they do not coincide
exactly. In view of the position of these orifices it is clear that
the tube of communication is of some length. In addition to
these two openings on to the external surface, the tube gives off
two short cecal processes, one of which nearly but not quite
opens into the bladder.

Anteriorly the tube gradually comes to an end and does not
end by opening on to the exterior, It diminishes in calibre for
the space of a few sections and then simply ends. At the bladder
end of the body—the end which is attached to the parent stock—
the tube gets to be more and more shoved to the side as the
bladder increases in size. In consequence also of this the tube
becomes flattened from side to side, at the anterior part of the
bladder. Followed backwards this tube can be recognised by its
thick chitinous Jining, already referred to, and can be seen
thereby to be quite distinct from the excretory tube which has
come to lie below it in consequence also of the development of
the bladder region, The next that occurs is that the tube divides
into two coincidently with the formation of two or three septa,
partly dividing up the cavity of that part of the bladder which
lies adjacent. The two tubes lie one dorsally to the other, and
thus both of them in the same straight line with the excretory
tube. The middle tube of the two after a very short course
appears to open into one of the chambers of the bladder, and is
thus a diverticulum like the one mentioned above. Furthermore,
I am not absolutely certain that it actually opens into the
bladder ; but it ends at least in close contact.

I feel, indeed, almost inclined to assert that this tube does
open into the cavity of the bladder, since the main tube, continu-
ing a little way further back, gets very narrow and undoubtedly
ends by opening into the bladder. It is to be noted that these
orifices do not involve a mere continuity of lumen. The character
of the lining membranes changes at the point of meeting. It
is further to be noted that it is not the main cavity of the
bladder into which the tube opens, but into a portion of it,
separated off by septa. This problematical tube varies in parts
in its structure and in its relations to the nerve-cord. Anteriorly
the layer of cells surrounding it are not so conspicuous as
elsewhere, and thus the longitudinal muscular fibres come into
greater prominence; the calibre of the tube also is not the same
all through the body. It has been mentioned that in a section
from about the middle of the body, that the tube lies outside
of the nerve-cord and quite laterally to it. In other parts of the
body, particularly anteriorly, the tube lies above (or, perhaps it
may prove necessary to say, below 7) the nerve-cord.

We have, therefore, in these young worms still attached to the

* T cannot differentiate between the dorsal and ventral surfaces.
+ See footnote above.

NEW ASEXUAL TAPEWORM. 837

parent, though (as the narrow stallx shows) just ready for detach-
ment, a tube of the structure of the outer layer of the body
which traverses in a straight line nearly the whole of the body,
which opens into the bladder behind and has two segmentally-
arranged orifices of communication with the external world.
There is no obvious answer to the question as to its nature. In
buds within the cyst of Hchinococcus* a cavity is formed which
is not an ingrowth of the cavity of the whole cyst, but which
is alleged to be continuous up to the scolex. This, however,
has been denied and the scolex asserted to be entirely solid. But
even supposing a comparison is possible on the grounds that
the scolex of Hcehinococcus is hollow, the short tubes leading to
the exterior in the worm of which the present paper contains
an account would seem to invalidate it. And, moreover, the
structure of this problematical tube is not that of the bladder
with which it communicates. It seems, indeed, to be a special
structure and not a space connected in any way (or, rather, any
direct way) with an evagination of the scolex.

For the greater part of the body I could only find a nerve-cord
on one side, where it was quite as obvious as in the “‘adult ” worm.
In the anterior region, however, the nerve-cord of the opposite
side was visible for a short space. It was, however, not half the
size of its fellow, though of precisely the same structure and
equally unmistakable as a nerve-cord. Both cords lie in the
medullary region just at its bordering on the corticallayer. I could
find no enlargement of the nature of a brain. Quite anteriorly, as
well as posteriorly in the bladder region, I could not recognise a
nerve-cord. It is noteworthy that the asymmetry of the two
water vascular tubes in size is exactly paralleled by that of the
two nerve-cords.

These are the facts which I have ascertained in the anatomy of
the most mature of the growing buds. It now remains to compare
the structure of the young worm with that of the parent stock of
which it is a bud.

§ Comparison of Bud with Parent Worm.

In the general form of the body and in the possession of a
relatively small bladder they agree; also the rudimentary scolex
is a marked feature of both young and old.

Tt is rather remarkable to find asymmetry in the young
worm, but this of course may be its normal method of growth.
Certainly I noticed no asymmetry in the parent form. What
is perhaps important as a difference is the presence of only two
water vascular tubes in the young and four of these in the
older worm. The additional water vascular tubes may, however,
make their appearance later. Furthermore, for the same
reason, too much stress cannot be laid upon the more copious

* Bronn’s Thierreichs, iv. Abth. 1B, p. 1549.
Proc. Zoou, Soc.—1912, No. LVI. 56

838 DR. F. E. BEDDARD ON A

bundles of longitudinal muscular fibres which are to be noted in
the large worm. I believe also that the continuous large tube in
the immature worm is represented by the cavities contained in
many of the segments of the adult worm, though the minute
structure, it must be confessed, differs in the two cases. Further-
more, the series of cavities in the adult worm lie very nearly in
the middle of the body, indeed quite in the middle of the body in
some segments; the tube in the young worm is, on the other hand,
as distinctly to one and the same side. But it must be borne in
mind that the position does vary in the young worm, and a further
flattening of its body might easily cause a greater similarity in
the position of the tube to that observable in the more fully
mature worm,

§ Comparison with other Forms and Systematic
Position of the Parasite.

Apart from the question of proliferation by budding, to which
we shall return later, this Tapeworm presents a certain number
of undoubted resemblances to Cysticercus fasciolaris (of rats and
mice, etc.), which becomes Tenia crassicollis in the Cat. Both
these forms agree (I have compared the species which forms the
subject of the present paper with examples of the Cysticercus from
the common rat) in possessing a long segmented body and a small
bladder posteriorly. But in Cysticercus fasciolaris the bladder is
smaller and the body shorter than in my species, while the hooked
and suckered anterior end renders any confusion or detailed
comparison impossible. Nevertheless, the two forms have in
common the small bladder and long strobila. But while the one
occupies the position in the body of its host of asexual worm, 7. e.
in a diverticulum of the alimentary tract, the other is found, as
are Cysticerci, encysted, and in the liver of its intermediate host.
Nor is it by any means certain that the species from the Mus-
quash is a member of the Teenioidea (Cyclophyllidea) at all. It
may well be a member of one of the lower groups of Cestodes.
We shall consider the arguments for and against the placing of
the worm among the Teenioidea.

It must be admitted at once that there are no absolutely con-
clusive arguments which point definitely one way or the other.
This, of course, may be explained on the assumption that we are
dealing here with quite a new typeof Cestode. In the meantime,
the structure of the worm so far as it can be read does not favour
such an assumption, though it does not, for the matter of that,
appear to be necessarily contradictory of this possible view. As to
the other alternatives, the absence of a marked scolex with suckers
is greatly against the reference of the worm to the Tenias, but
the character of the terminal bladder, on the other hand, is on the
whole in favour of such a placing. This latter fact is obviously
against the supposition that the worm is a Plerocercoid of any
kind; but the lack of a definite “head” is as clearly in favour of

NEW ASEXUAL TAPEWORM. 839

this supposition. There, as it appears to me, the question must be
left—in regrettable uncertainty.

The process of asexual reproduction by budding which this
worm shows is not entirely unknown among the Cestoidea, but
there are some not unimportant differences from what has been
recorded in other forms. There are two principal comparisons to
be made.

The first of these is with Cysticercus longicollis, the bladder-
worm of Zeenia crassiceps. It is of further interest from the
point of view of the present comparison to note that this Cysticercus
is also found in a Rodent, Arvicola arvalis. Besides the earlier
investigators, Prof. M. Braun* has studied this form and em-
bodied his results in brief in Bronn’s ‘ Thierreichs’ +. The buds
from the Cysticerci do not apparently {contain a prolongation of
the bladder-cavity, ‘“‘sondern durchweg als solide Wucherungen
der peripheren Schicht der Wand entstehen.” This is an obvious
point of similarity with my species, where, of course, the buds
cannot be continuous with the cavity of the bladder—that is to
say, in both cases the buds are solid outgrowths. Furthermore,
before being separated off from the parent Cysticercus the attach-
ment of the bud dwindles to a narrow stalk precisely as I have
described above. The buds, however, in the case of Cysticercus
longicollis seem to be limited to the bladder and always to the
hinder end of that; whereas in my species this is exactly the
region where no buds are formed.

I believe, however, that a nearer approximation to the condition
observable in the species which I describe in the present paper is
offered by a worm recently described by Tjima { with some detail.
This is a Plerocercus or Plerocercoid found parasitic in a human
being in Japan in cysts in the skin. It appears to give off actual
buds, which are, however, more or less irregularly arranged and
present nothing of the comparative symmetry manifested in the
species from Miber zibethicus. ‘The individuals differ in the degree
of their budding, some giving off a large series of slender pro-
cesses. These buds are, as in my species, actual outgrowths of the
parent stock and not, for instance, connected with the excretory
organs or any other definite part of the body. They occur
as much at the head end as elsewhere and there differ from the
Tapeworm of Wiber zibethicus. ‘There is, however, a resemblance,
in that in both cases the more mature buds are not in front of
or behind the less mature: there is a complete irregularity in
their order of succession.

The similarity between the two cases of budding cannot, how-
ever, be carried into any detail, since it is obvious that the two
worms are not nearly allied. The Plerocercoid described by

* See Centralbl. f. Bakt. u. Par. xx. 1896, p. 580; Zool. Anz. 1896, No. 514; ibid.
1897, No. 521. These papers are not illustrated.
+ Bd. iv. Abth. 1B, p. 1529.
£ “ On a new Cestode Larva parasitic in Man,” Journ. Coll. Sci. Japan, xx. 1905.
For a reference to this paper I am indebted to Dr. W. Nicoll.
56*

840 DR. F. E. BEDDARD ON A

Tjima is regarded by him—and, as I think, rightly—as a young
Bothriocephalid whose larval stages are also known from similar
positions in other mammals than man. These larve have no
bladder, and thus the fact that in the Plerocercoid, as in my
worm, the buds are produced from the general body surface loses
its principal significance. It is, | think, more important to note
that the worm from Viber zibethicus differs from Cysticercus
longicollis, in that the budding is not upon the bladder end of the
worm. It is remarkable, however, that both in my worm and in
the Plerocercoid there is no recognisable scolex ; but at the same
time there is in the supposed young Bothriocephalid no proper
segmentation of the body as yet visible. It is thus particularly
remarkable that my species is very fully segmented throughout.
I think, therefore, that the case of budding which I bring forward
in the present paper may be regarded as in some respects a new
form of asexual reproduction which is partly paralleled in a few
other forms of Tapeworms.

Inasmuch as the propagation of this worm is different in detail
from anything that is known, and as the worm itself does not
entirely conform with any known species or genus, it is desirable,
as I think, for mere convenience sake to give it a name and to
define as far as may be the characters of the Cestode. Had the
worm proved definitely a Tetracotylean*, I should have con-
tented myself with referring it temporarily to the genus “ Tenia,”
a name which is generally given to forms whose generic affinities
are uncertain. As, however, this cannot be done with any
certainty, I venture to give a new generic name which is non-
committal as to its position in the series, and I term it
accordingly

Urocystidium gemmiparum, gen. et sp. n.

Incompletely mature worm with small bladder and very numerous
proglottids. Scolea feebly developed and without hooks. Tubes of
water vascular system lateral to cach other, the dorsal with thick
muscular coat and inside of ventral, the ventrals connected by a trans-
verse vessel which forks and surrounds dorsal vessel ; a network present
in cortex as well as in medullary region. Longitudinal musele-
layer in cortex of at least two rows of bundles with numerous fibres
in each. Buds formed at hinder end of body on both sides, thus
forming two rows, of which the bladders are attached to the parent
stock ; the scolex is not inverted, is without hooks, and rudimentary.

Hab. Musquash (Fiber zibethicus), in liver-duct.

(2) THE Sexvat Form.

As has been already mentioned, the hepatic duct contained
besides the ‘“ cysticercoid” just described a sexual worm which I

__* Lthink it necessary to maintain an attitude of reserve with respect to the
identity of the asexual and sexual forms.

—

NEW ASEXUAL TAPEWORM. 841

believe to be the mature form of the same Tapeworm. The size of
the two worms was about the same. The sexual form measured
86 mm. in length and the greatest diameter, which occurred not

Text-fig. 118.

The sexual form of Urocystidiwm gemmiparum (P).

The left-hand figure represents the entire worm magnified about twice; the right-

hand figure represents the scolex with a double crown of hooks more highly
magnified.

far behind the head, was6 mm, The appearance of the worm
(text-fig. 118) was quite similar to that of the immature form. It

842, DR. F. E. BEDDARD ON A

was posteriorly of a rather delicate appearance, rather translucent,
and the proglottids were very short and enormously wide in
proportion. There was no increase in length of the proglottids
towards the end of the body, and their proportions were as in the
asexual worm. It would hardly, of course, be sufficient to insist
upon an identity upon these grounds alone; but, taken in con-
junction with other facts which will be dealt with, the similarity
in outward appearance is very striking. ‘The “tail” end of the
body appeared to be a little excavated, as is common in tapeworms,
the penultimate segments slightly embracing the last segment,
which was not quite so wide. I can believe that the worm was
proliferating at this end.

The scolex was not quite so wide as the ensuing strobila, and no
appreciable neck separated the two. The scolex is proportionately
large, as is indicated in the accompanying text-figure (text-fig.
118), and well armed anteriorly with two rows of hooks alter-
nating in position. The anterior circle of hooks consisted of 16
separate hooks, which were about twice the size of those of the
succeeding circle, whose number I did not count, but which were
presumably the same, as they were implanted between the larger
hooks. The usual four suckers are present; they show no un-
usual features and are unarmed ; their cavity looks forwards.

I have investigated the internal structure by means of trans-
verse and sagittal sections. The cortical layer does not differ
greatly from the medullary layer in thickness, and the general
appearance of the sections is very like that of sections of the
immature worm (see text-fig. 119). The cortex, for example, is
identical or nearly so. The same bundles of longitudinal fibres
occur and are very much of the same thickness. They are also
separated from the medulla by transversely running fibres. I
have described and figured in the asexual form the bundle or
bundles of rather stouter longitudinal fibres running outside of
the nerve-cord on either side and associated with a cavity dubiously
related to the excretory system. I find the same arrangement in
the sexual worm.

The excretory system does not needa very long description,
since it agrees in its main peculiarities with that of the supposed
asexual form. There are, in fact, the same two lateral vessels on
either side lying parallel to each other. They are, moreover,
roughly equisized, and the innermost of the two has very thick
muscular walls, the fibres being circular in their disposition. I
should add that nuclei interspersed among these fibres were very
obvious. In addition to these two longitudinal trunks each pro-
glottid possesses a transverse vessel which has the same remarkable
mode of union with the ventral excretory tube that I have figured
(see text-fig. 115) and described (see p. 830) in the presumed
Cysticercoid stage, and which I need not redescribe here as the
structure seems to be identical. There is, however, one important
difference which the sexual form shows from the Cysticercoid ; and
that is the absence in the former of the peripheral water vascular

NEW ASEXUAL TAPEWORM, 843

network of the latter. I could find no trace of this in the sexual
worm, although it was easy to see in the supposed Cysticercoid of
the same. Tt seems to me possible (perhaps necessary) to explain
this difference by assuming in the Cysticercoid a retention and
gradual metamorphosis of the bladder into the strobila, together
with its excretory system, which latter ultimately disappears on
the assumption of sexual characters. There are, of course, other
Cestodes in which the bladder is not cast off before the acquiring
of sexual characters.

Text-fig. 119.

ee

se

wen
<
ae
s

BRE
eo

ee
fs

Bes

Part of a transverse section through a proglottid of the sexual worm.

e.m. Circular muscles. d.v. Dorsal vessel of the water vascular system. N. The
three laterally running nerve-cords. ¢.v. Transverse water vascular trunk,
v.v. Ventral trunk. X. Large longitudinal muscular fibres referred to in the
text as frequently running within a space.

In the neighbourhood of the water vascular trunks delicate muscular fibres are
shown running chiefly in a dorso-ventral direction, which may be associated with
the dilation and contraction of the water vascular tubes.

The nerve-cord shows no trace of the asymmetry which I have
described above in the very young worms. There is nothing un-
usual about the position or structure of the main trunks. There

844 DR. F, E. BEDDARD ON A

is, however, on either side of the body and running parallel with
the main nerve-cord a second and a third nerve-cord (see text-
fig. 119, N) which lie above and below the former and at some
distance from it. These accessory nerve-cords appear to be
exactly like the main nerve-cord in structure and are situated
atthe boundary of the medullary and cortical regions, a trifle
nearer to the middle of the segment. In sagittal sections the
supplementary nerve-cord is very obvious and is seen to be con-
nected with the main trunk by numerous (7 or 8) transverse
cords in each segment, which produce a ladder-like appearance.
There is an obvious likeness here to Bertiella, in which genus the
nerve-cord also consists of three separate strands, and to Dioico-
cestus, where the cords are wider apart. ‘They are, however, much
closer together in Bertiella than in the present worm. I have
not observed this arrangement of three nerve-cords in the supposed
immature form already described. But it is to be observed that
the main cord is there almost as wide as the medulla, thus leaving
no room for the accessory cords which may be split off later.

§ Sexual Organs.

In the middle region of the body and possibly for some way in
front (1 have not examined sections just in front of the mid-
region) the proglottids are full of ripe eggs. I have examined
these proglottids in sagittal as well as in transverse sections. The
former bring out the important fact that there is apparently no
internal boundary-line between the proglottids ; for the eggs form
continuous masses stretching without intermission from segment
to segment. In this region the most careful search failed to show
any other genital organs than these exceedingly numerous ripe
ova. It is, of course, not uncommon in fully mature segments
of tapeworms, such as these in the present form, to find nothing
but ova in the medulla; but in those forms there are at least
considerable though often altered remains of the genital ducts
and terminal apparatus. In the present species there is no
vestige of cirrus-sac or vagina and not the least trace that I
could discover of an external pore. Nor do I think that it would
have been easy to have missed these structures were they present,
particularly in a series of sagittal sections. J am, in fact, con-
vinced that they are absent.

The eggs of this tapeworm do not lie in any space or spaces
that can be strictly called a uterus. There are, however, spaces
which suggest the remains of the uterus. These are not arranged
in strict metamerism; but in a certain number of proglottids,
and not by any means confined to a single proglottid, are large
cavities such as is represented in the accompanying text-figure
(text-fig. 120). These cavities are more or less circular and lhe
in the medullary region of the body. As a matter of fact,
they are not at all full of eggs. Indeed, they are often quite
empty; only sometimes are eggs to be seen lying within them

es

NEW ASEXUAL TAPEWORM. 845

Text-fig. 120.

Two sections through portions of proglottids of the sexual worm.

The upper figure is a longitudinal section in the more anterior part of the body,
showing the smaller ova lying among and to the inside of the longitudinal
muscles (J.); these smaller ova lie in what is probably generative tissue, not
clearly indicated in the drawing.

¢. Transverse muscle fibres. o. Mature ova scattered through medulla and
also at the opposite side of the figure in the cortex.

The lower figure represents a transverse section through a more posteriorly situated
segment.

c. A cavity which may possibly represent a uterus; two spaces are shown in the
cortex which may or may not belong to the same category. 7. Longitudinal
muscles, o. Ripe ova showing a nucleus. ¢. Transverse muscles.

846 DR. F. E. BEDDARD ON A

and adherent to their walls. I doubt, therefore, whether these
Spaces are the remains of the uterus, or whether they are not
rather cavities which have arisen in the medullary region perhaps
by stretching of the walls of the body and consequent laceration.
The eggs, in fact, lie scattered through the parenchyma, some-
times singly and more often in masses of various sizes, as is
shown in the figure referred to. The eggs are not limited
to the medullary region. It is clearly to be observed that
they extend into the cortex (text-figs. 120, 121) a good way

Text-fig. 121.

sue
38 ! :
See gata *

ohh. elk

Pay sept oN
en

+ RA WRNBG, Saks
PO Sgiseem cal mp CS
ae
Met Sheet Leary
Seer AE ALTE

Part of a transverse section through a proglottid of the sexually mature worm in
which the ripe ova (0.) have been very deeply stained and are seen to be
scattered through the cortex as well as the medulla.

1. Longitudinal muscles. ¢. Transverse muscles.

towards the external layer of the body-wall. They stop short,
however, some way below it. It is quite certain that whether
or no there may be remains of the uterus, some of the eggs lie
scattered within the parenchyma, In the cortical parenchyma
they are to be seen between the muscle-bundles and closely packed
in masses. ‘The appearance is, indeed, not at all unlike that
which I have recently described in the tapeworm Anoplotenia
dasyuri * in those parts where the cavities of the uterus were
not so conspicuous; for in the latter worm there are uterine

* P.Z.S. 1911, p. 1012, text-fig. 213.

NEW ASEXUAL TAPEWORM. 847

cavities lodging many, perhaps most, of the eggs. The present
species is, as it were, a stage beyond that exhibited by Anoplo-
tenia dasyuri. As is well known, the imbedding of the eggs in
the parenchyma is not a novelty; for Oochoristica is a genus
which is largely characterised by this very feature, though in
this case the eggs are imbedded singly and there are not the
heaps of eggs seen in the present species and perhaps in Ano-
plotenia. In Anoplotenia, however, the masses of eggs are
usually in cavities of the uterus. In both Oochoristica and Ano-
ploteenia, however, the scattered eggs are limited to the medullary
region. But the peculiarity of the species with which I am
dealing in the present paper is also known in other forms. Clerc *
has figured eggs in the cortical region in a species of Dilepis.

I have some reason to think that the scattered eggs lie in
the interstices of the network forming the ground-tissue of the
worm; but in any case eggs were also seen lying in larger
circular cavities. I believe that these cavities (shown in text-
fig. 120) are not the remains of a uterus. They are, as it
appears to me, to be directly compared with similar spaces to be
seen in mature (but less mature than in the present instance)
segments of Jnermicapsifer, where 1 have described and figured
them?. In this latter case I was able to bring forward some
positive evidence to show that the spaces in question were not,
and could not be, the isolated series of chambers left by a
vanishing uterus. In the worm with which I am dealing in the
present communication I am unable to furnish any of those
proofs of the nature of the cavities in question. I can merely
point out their general similarity to those of /nermicapsifer.

Although there were no traces of gonads like those of other
tapeworms in this region of the body, I have succeeded in dis-
covering what I regard as undoubtedly the generative tissue.
The medulla was packed with ripe ova, which, as already said,
stray into the cortical layers lying between the bundles of
muscles. But at the line of junction of the cortex and medulla
and lying in the latter is a continuous mass of tissue with
interspersed nuclei, in and just outside of which eggs in various
stages of development are to be seen. This is, as I think,
undoubtedly to be regarded as a continuous layer of gonadial
tissue. I cannot see any other obvious explanation of it. It is
clearly totally unlike the ground-tissue of the tapeworm’s body
and forms a solid mass with darkly staining nuclei. Its associa-
tion with developing ova seems to be final in the matter of
argument. It is, however, possibly the case that this gonadial
tissue is wholly or in part the origin of spermatozoa also; for
testes like those of other tapeworms were not found in the
present species. I have, however, no evidence of the presence of
spermatozoa. The tissue itself is—save where developing ova

* Rev. Zool. Suisse, t. xi. 1908, pl. xi. figs. 75, 76.

+ P. Z.S. 1912, p. 588, text-fig. 77. For the opposite view, see Janicki, Denkschr.
Med.-Nat. Ges. Jena, xvi. (1911) p. 381, pl. xiv. figs. 25-27.

848 DR. F. E. BEDDARD ON A

are present—naturally indistinctive. The only way of proving
its testicular nature is the discovery of spermatozoa, An _
inability to find these is not of itself conclusive proof of the
absence of testicular tissue. But an argument of weight in this
direction is to be derived from a study of the ripe eggs. These
are rather small as compared with some species and possess a
very thick hyaline shell, which is with difficulty penetrated by
staining reagents. Thus in many of my sections the eggs seem
to be simply oval structureless bodies. In cases, however, where
the staining has been more successful the true ovum within the
shell becomes obvious. It consists of a single cell with a large
nucleus. I say a single cell because there are not vitelline cells
enclosed within the shell, anything in the nature of a vitelline
body being completely absent. The absence of a vitelline gland
is rare among tapeworms, but is known in the genus called
on that very account Avitellina*. In no case did I find this
ovum in course of division—and I have examined a large
number of individual ova in many segments. I infer, therefore,
that no spermatozoa are formed in this individual worm and
furthermore that there is no entry of spermatozoa from another
individual—a fact which is also supported by the absence of a
vagina. But it must be remembered that the host may not
have contained in the bile-ducts another sperm-producing indi-
vidual. It may be that fertilisation occurs outside the body.
But this is clearly a mere suggestion for the present, though
not repugnant to such evidence as there is.

There being no trace of the actual uterus in the middle
region of the body, I naturally sought for these bodies elsewhere
and made a series of sections from the anterior region of the
body. I selected the rather thicker region which immediately
follows the head and examined a piece cut from about 10 mm.
behind the anterior extremity. Here, as I imagined, the:im-
mature uterus might be found or at any rate some trace of its
existence. I could, however, find in this part no essential
differences from the posterior region of the body which has just
been described. ‘The body was in the same way packed in places
with the ripe ova, which were present everywhere in a less crowded
state; indeed, they seemed to me to occur nearer to the external
surface here than posteriorly. I found the eggs only just beneath
the external cellular layer of the body-wall. The only conclusion
that I can come to is that this worm possesses no uterus, or
that it exists for a very transitory period only, and also that very
possibly the sexes are separated as in the genus Diotcocestus.
The specimen described here being a female, this conclusion
is obviously based upon negative evidence only and is thus less
valuable. It is, however, quite clear to my mind that the genera-
tive products develop simultaneously in the proglottids and that

* See Gough, Quart. Journ. Mier. Sci. vol. lvi. pt. 2, 1911, for an account of this
genus.
b=)

NEW ASEXUAL TAPEWORM. 849

therefore there is no growth of the worm when it has arrived
at sexual maturity. This conclusion, which fits the facts that I
am able to set forth, agrees also, it may be pointed out, with the
very worm-like cysticercoid stage. There is, so to speak, not
much necessity for this immature worm to grow in length before
assuming the sexual condition. Far otherwise is it with the
typical Cysticerci, which are provided with but short strobila
as compared with the mature forms of their species. There is
at the very least a relation between the two series of facts which
is worth noting. It seems to me to be furthermore likely that
at no period are there sexual ducts developed—at any rate, no
female ducts. This latter circumstance, if true, is not new; for
the absence of a female orifice has been asserted in more than
one genus of worms belonging to more than one family.
Aporina among the Anoplocephalidze and many genera among
the Acoleide are instances to the point. It is mainly, indeed,
the plain absence of a cirrus-sac which leads me to believe that
this worm possesses a dicecious habit so rare in this group.

§ General Résumé and Systematic Position.

It is, I think, obvious from the foregoing account of the
sexual form of this worm that it is the representative of a new
genus which differs in a good many particulars from any known
form. It will be convenient to give a short résumé of the
essential characters of this worm, for which the name already
given to what I regard as the asexual form may be retained.
My reasons for retaining the name of the asexual form are
firstly that I have been able to give a more complete account
of it than of the presumed sexual form of the same Cestode,
and secondly that I cannot fully define the sexual form, concerning
whose identity with the asexual form, moreover, some doubt may
be considered to remain.

In any case the following are the principal characters of the
sexual worm :—Length 86 mm., greatest breadth 6 mm. Head
with two rows of hooks 16 in each row, the hooks of the anterior
row twice the size of those of the second row. Suckers normal
and unarmed. No neck; strobila consisting of numerous pro-
glottids very short and not appreciably longer at posterior end of
worm.

Body flat, thicker anteriorly. Cortex about the same diameter
as medulla. Two layers of bundles of longitudinal muscular
fibres in cortex. Water vascular tubes two on each side lying
side by side; dorsal (?) tube with thick muscular walls. A trans-
verse vessel in each proglottid forking round dorsal vessel to join
ventral at two points. No water vascular network present.
Nervous system consisting of a larger lateral trunk and two
smaller trunks, one dorsal and one ventral, connected by many
cords in each proglottid to main trunk. The sexes are apparently
separate or the worm is protogynous or protandrous throughout.

850 ON A NEW ASEXUAL TAPEWORM.

Female organs consist of a layer of gonadial tissue lying in the
medulla at its junction with the cortical layer from which ova are
shed into the body, which they permeate even to the outer layers of
the cortex. There is no trace of segmentally arranged gonads or of
a uterus, nor are the eggs surrounded by any kind of ‘ capsule.”
There is no vagina or female passage of any kind; the vitelline
glands are totally absent. The ripe eggs are surrounded by a
thick hyaline shell and none were observed to be dividing and
no embryos were discovered in them. ‘Thus it is possible that
fertilisation occurs outside of the body.

These characteristics do not entirely fall in with those of any
other family of tapeworms. They obviously point to an affinity
with the Acoleide, but do not definitely necessitate the inclusion
of this remarkable worm within that family.

§ Anatomical Summary.

It may be convenient to extract from the foregoing account
of this new tapeworm the more remarkable anatomical facts
which I have been able to make out.

(1) The absence of a marked scolex in the Cysticercus of a tape-
worm which cannot be placed among the Bothriocephalids
and of which therefore it would be expected that the scolex
would be very prominent. And correlated with this the
necessary assumption that, as in Bothriocephalids, the
scolex only develops part passu with the growth in maturity
of the worm.

(2) The enormous size of the strobila as compared with the
bladder—a rare condition among the Tetracotylea, but
paralleled in Cysticercus fasciolaris.

(3) The very thick layer of muscle surrounding the dorsal
vessel of the excretory system and the bifurcation of the
transverse vessels round the dorsal vessel to open into the
ventral water vascular tube.

(4) The total absence of generative ducts or (presuming that
the species is dicecious) of the female tubes, which is, so
far as I am aware, a unique anatomical character.

(5) The total absence of a uterus or of any trace thereof.

(6) The diffuse and non-metameric character of the ovaries,
which are not sharply differentiated into relatively small
bodies of a definite shape.

(7) The enormous quantity of eggs produced and their exist-
ence in quite anterior as well as posterior segments; the
eges are, moreover, found quite as abundantly in the
cortical layer as in the medulla. The conditions observable
in this part of the generative system are simply an
exaggeration of what is to be met with in other genera,
where the eggs come to be ultimately scattered through
the medullary parenchyma.

ON TWO NEW TREMATODES. 851

50. On Two new Trematode Parasites from the Indian Cobra.
By Wiuuiam Nico, M.A., D.Sc., M.D., F.Z.S., Lister

Institute of Preventive Medicine, London.
[Received May 28, 1912: Read June 4, 1912. |

(Text-figure 122.)

INDEX.
Page
Ethology: Two new Trematode parasites in the gall-bladder
and ureters of the Indian Cobra (Naja tripudians) ...... 851
Geographical Zoology : India; Cobra, two new flukes from
the gall-bladder and ureters .....................cccceseseeoeene SDL
Systematic :
Xenopharyne (gen. n., Fam. Dicrocceliide) solus, sp. n.,
from gall-bladder of Indian Cobra ....................00.... 861
Styphlodora naje, sp. n., from ureters of Indian Cobra,
closely resembles S. horrida Leidy ........................... 854

In an Indian Cobra (Waja tripudians) which died in the
Zoological Society’s Gardens on 11th October, 1911, a few speci-
mens of two interesting new Trematode parasites were found.
Of the first of these a single specimen was found in the gall-
bladder; of the second, four somewhat macerated specimens were
met with in the ureters. Four different species of Nematode
parasites were also present in the same animal, so that altogether
it was infected with six different species of parasitic worms.

The first species belongs to the family Dicrocceliid and is of
particular interest from the fact that members of this family
are not commonly found in Reptiles. Hitherto the only typicai
representative in Reptiles is that described by de Faria (1910).
The present species shows most of the characteristic features of
the family, but at the same time it exhibits several divergences of
such importance as to warrant its being regarded as the type of a
new genus. For that genus I propose the name Xenopharynx.

XENOPHARYNX SOLUS, gen. et sp.n. (Text-fig. 122, B.)

Only a single specimen was found in the gall-bladder. It
measures 4°55 mm. in length and 1°68 mm. in greatest breadth,
which is about the middle of the body. The outline is almost
elliptical and the body is fairly flat. There are no cuticular
spines.

The oral sucker has a diameter of -42 mm., but its length is
only 30 mm. The ventral sucker is circular, with a diameter of
‘41 mm. It is situated 1:13 mm. from the anterior end. The
neck is therefore almost exactly one-fourth of the body-length.

The pharynx is contiguous with the oral sucker and measures
‘22x24 mm. It possesses the curious shape shown in text-fig.
122, B. At first sight this shape was thought to be the result of

852 DR. WILLIAM NICOLL ON

unequal contraction of the pharyngeal walls or perhaps due to the
drawing of the cesophagus into the lumen of the pharynx, but
neither of these possibilities seemed to accord with experience.
The condition appears to be brought about by the thinning of the

Text-fig. 122.

M. Rhodes del.
A. Styphlodora naje, sp.n. Ventral view. x 40. C., cirrus;
Ov., ovary; R.S., receptaculum seminis.

B. Xenopharynex solus, gen. et sp.n. Ventral view. x 25.
Ov., ovary.

posterior part of the pharyngeal wall, which anteriorly is of the
usual thickness. In section the appearance is somewhat horse-
shoe-shaped. It is unfortunate that no second specimen was

TWO NEW TREMATODES. 853

available for comparison. Following the pharynx, and aboyt as
long as it, is a wide cesophagus, which bifuycates about midway
between the pharynx and the ventral sucker, The diverticula are
fairly uniform, but become somewhat sinugus towards their ter-
mination, which is ‘45 mm, from the end of the body.

The excretory vesicle is Y-shaped, with a long median stem
reaching the middle of the body and two short limbs, the left of
which extends as far forward as the left testis, Owing to the
fact that sections could not be made, it was impossible to determine
whether Odhner’s view with regard to the shape of the excretory
vesicle in the Dicrocceliidze holds good in this case. To all
appearance, however, the paired limbs are part of the vesicle,
Their walls stand out as distinctly, and are abont the same
thickness, as those of the main stem,

The testes lie not far behind the ventral sucker, They are two
transversely oval bodies, separated from each other by a distance
equal to their diameter, They are situated obliquely, the right
being about half a diameter in advance of the left, and being
separated from the ventral sucker by a similar distance. They
measure ‘31 mm, in transverse diameter, The cirrus-pouch is of
small size and lies entirely in front of the yentral sucker, It
measures *29 x :13 mm., and contains a small convoluted vesicula
seminalis, a comparatively long pars prostatica, and a short narrow
ductus ejaculatorius, The genital aperture is median, just over
the intestinal bifureation.

The ovary lies a considerable distance behind the testes, about
midway between them and the end of the body. It is on the
right side overlying the right intestinal diverticulum, Its outline
is transversely oval and it is much Jarger than the testes,
measuring 31 x°47 mm. There appears to be no receptaculum
seminis, or if present it is obscured by the uterus, The yolk-
glands are situated chiefly in the neck, the sides of which they
almost completely fill. They extend on each side from the oral
sucker to well behind the ventral sucker, but they are more ex-
tensive on the left than on the right. On the right they stop at
the level of the bifurcation of the excretory vesicle, although
there are a coupie of small follicles near the ovary. On the left
they extend slightly beyond the level of the ovary, In the neck
the follicles spread well in towards the mid-line of the body, but
behind the ventyal sucker they lie entirely to the outer side of the
intestinal diverticula. The transverse yolk-ducts cross the body
about the level of the testes, and the shell-gland les just behind
the right testis,

The uterus is moderately voluminous and is thrown inta
irregular narrow convolutions, It does not extend much behind
the ovary, but in front it overlaps the intestinal diverticula, and
winding between the testes it passes forward on the left side of
the ventral sucker and makes a few short turns in front of the
sucker before passing into the short vagina. The eggs are small
and numerous, measuring *036—-039 x 018-019 mm.

Proc. Zoou. Soc.—1912, No. LVITI. 57

854 DR. WILLIAM NICOLL ON

The new species differs from all other Dicrocceliidee in the dis-
tribution of the yolk-glands and in the shape of the pharynx.
Another important feature is the distance separating the ovary
from the shell-gland complex. From the genus Dicroceliwm it is
further distinguished by the position of the testes and the ovary
and by the extent of the uterus. With Platynosomwm it is more
closely allied, yet the differences between it and this genus are
greater than those separating Dicroceliwm from Platynosomum or
Kurytrema. It is rather curious that this new species does not
show any particularly close relationship to “ Dicrocelinum” infidum
de Faria from the snake, Hwnectes marina. The latter is closely
allied to the avian genus Platynosomum and should be included
in it, unless de Faria’s doubtful observation in regard to the
excretory vesicle proves to be correct.

SryPHLODORA NAJH, sp.n. (Text-fig. 122, A.)

Four somewhat macerated specimens of this species were found
in the ureters of the Cobra. The species is a typical member of
the genus Styphlodora and presents a very great resemblance to
S. serrata Looss and to S. horrida Leidy. The features separating
it from these two species are so slight that I have some hesitation
in regarding it as a distinct species. They are, however, quite as
distinctive as those separating the above two species from each
other. The difficulty is rather increased by the fact that the
present specimens, although mature, are possibly not fully grown.

The length is 2°0-2°4 inm., the greatest breadth -42—55 mm.,
which occurs a little behind the ventral sucker. The breadth of
the whole postacetabular region is fairly uniform, and there is
only a very slight attenuation in the neck. There isa considerable
amount of dorso-ventral flattening. Im each of the specimens
cuticular spines were entirely absent, but it is practically certain
that they have fallen off, and it would be unreasonable to suppose
that this isan unarmed form. On that account I have had spines
depicted in the drawing.

The oral sucker is subterminal and has a diameter of *22--25 mm.
It is rounded and rather shallow. ‘The ventral sucker is slightly
transverse and measures *24x°25 mm. It is only very little
larger than the ora: sucker, and it is situated at a distance of
-67—84 mm. from the anterior end, 7. e. about one-third of the
body-length. There is a distinct prepharynx followed by a large
pharynx measuring about "13 x-14 mm. The esophagus is about
the same length, and the bifurcation takes place midway between
the suckers. The intestinal diverticula are fairly straight and of
considerable width. They are longer than in the other species of
Styphlodora, reaching to within 18-29 mm. of the posterior end.
They are very slightly unequal in length.

The main excretory vesicle was entirely invisible, but it was
apparent that it gave off numerous lateral branches, which,
anastomosing freely, gave the body a honeycomb-like appearance.

The testes lie obliquely behind one another, the left being in

eeu

TWO NEW TREMATODES. 855

front. They are separated by the uterus, which passes between
them, but does not overlap them to any great extent. Their
outline is irregular, but they appear to be roughly triangular or
trilobate. The anterior testis lies almost exactly midway between
the two ends of the body, but its position in relation to the ovary
varies somewhat, and it may be slightly nearer than is shown in
the figure. The posterior testis is about -27 mm. behind the
anterior. They touch the intestinal diverticula or overlap them
to a very small extent. Their dimensions are, on an average,
-21x-17 mm. and *22x'19 mm. Thecirrus-pouch is of moderate
length and uniform width, measuring 31 x-11 mm. It extends
to about the posterior border of the ventral sucker, and contains
a convoluted vesicula, a short pars prostatica, and a fairly long
ductus ejaculatorius. In each of the specimens the cirrus was
exserted and was longer than the pouch itself. The genital
aperture is situated in the middle line, almost immediately in
front of the ventral sucker,

The small round ovary lies just behind the ventral sucker, a
little to the right side, and measures "13 mm, in diameter,
Immediately behind it lies a somewhat smaller receptaculum
seminis. ‘he yolk-glands lie on each side at the level of the
ovary. They are, for the most part, external to the intestinal
diverticula, but they also overlap them dorsally. They consist on
each side of about a dozen fairly large follicles, which extend from
the posterior border of the ventral sucker to the anterior border
of the left testis. The transverse yolk-ducts cross the posterior
border of the ovary.

The uterus is poorly developed. It passes back between the
testes to near the posterior end of the body and returns along
much the same path, terminating ina short vagina. It is only
slightly convoluted and contains a comparatively small number
of eggs. It is confined within the space bounded by the intestinal
diverticula, and it does not form any convolutions behind the ends
of the intestine, as is the case in the other species of Styphlodora.
The ova are rather elongated, oval, possessing a large operculum
and, in a few cases, a small knob at the anopereular pole. They
measure *038—"048 mm. x ‘019-024 mm., the average size being
043 x :021 mm.

The features which appear to distinguish this form as a definite
species are the length of the intestinal diverticula and the
restricted extent of the uterus. It is not impossible that both
these features may be less marked in a fully-grown specimen,
In addition the yolk-glands are slightly less extensive than they
are in the other species of the genus. From S. serrata it is
further distinguished by the relatively larger size of the suckers,
Both S. condita and S. horrida also possess relatively smaller
suckers, and in these species the distance between the suckers is
considerably less. The only other species of Styphlodora, namely
S. similis Sonsino and S. bascaniensis Goldberger, present marked

features of difference from the present species.
DT*

856 DR. WILLIAM NICOLL ON
REFERENCES.

1. Ds Faria, Gomez. 1910.—Contribution towards the Classifi-
eation of Brazilian Entozoa. IJ. Dicrocelium infidum,
n. sp., Parasite of the Gall-bladder of Hunectes marina L.
Memorias do Instituto Oswaldo Cruz, i1. (1) pp. 22-28, pl. 2.

2. De Farta, Gomnz. 1911.—Beitriige zur Systematik der bra-
zilianischen Helminthen. IV. Styphlodora condita, n. sp.
Memorias do Instituto Oswaldo Cruz, ii. (1) pp. 40-45, pl. 1.

3. Gonppercer, J. 1911—A New ‘Trematode (Styphlodora
bascaniensis) with a blind Laurer’s Canal. Proc. U.S. Nat.
Mus. xl, pp. 233-239.

4. Leipy, J. 1850.—Descriptions of Two Species of Distoma, with
the partial History of one of them. Journ. Acad. Nat. Sc.
Philadelphia, ser. 2, vol. i. pp, 301-310, pl. 43.

5. Looss, A. 1899.—Weitere Beitrige zur Kenntniss der
Trematoden-Fauna Aegyptens. Zoolog. Jahrbiicher, Abt. f.
Syst, xii. pp. 707-708, pl. 26, fig. 28.

6. Opuner, T. 1910.—Nordostafrikanische Trematoden. Results
Swedish Zool. Exped. to Egypt and White Nile, 1901, p. 53,
note 2, fig, v.

5]. Statistical Note on the Worm Parasites collected from
the Animals dying in the Zoological Gardens, from
December 1910 till April 1912. By Wriu1am Nicont,
M.A., D.Se., M.D., F.Z.S., Lister Institute of Preventive
Medicine, London.

[ Received May 30, 1912: Read June 4, 1912, |

INDEX.
Page
Ethology: Animals dying in the Zoological
Society’s Gardens, numbers infected with
[OAV GHOUAOAS cooconses noes ssecsodeessbeonacunndenns eld
Geographical Zoology : Importance of ascertaining
distribution of parasitic worms .................. 857

At the scientific meeting of the Society held on May 21st last,
{ referred to the exeellent work which is being done by the
Prosectorial department in acquiring information concerning the
parasites which infect the animals living in the Gardens. This
is particularly noteworthy in regard to the worm parasites, about
which many valuable facts have been obtained, Following my
remarks at the meeting, Dr. Beddard suggested that I might be
able to supply some general information regarding the animals
which had been sent to me for examination, and acting on this
suggestion I venture to offer the following communication.

The scheme, which owes its initiation to the Secretary of the

LL OO tint

WORM PARASITES. 857

Society, was put into action in November 1910, and the practical
working has been evolved as follows : A general examination of the
viscera of all the animals killed or dying in the Gardens is made
by the Pathologist, after which certain selected examples are
forwarded to me for further examination. The desirability of this
latter procedure is shown by the fact that a very large proportion of
Entozoa are too small to be detected in the course of an ordinary
routine examination; and although the detection of such forms
does not involve any very special skill, yet it is a tedious process
and one which necessitates some experience. In justification of
this extra trouble it may be remarked that the comparatively
few animals which have been submitted to this additional
examination have yielded almost as large a number of parasitic
worms as all the other animals put together. This fact, I
venture to believe, is a plea for a still more extended and
exhaustive system of examination, dealing with all the animals
which die in the Gardens. This, naturally, would involve some.
additional expenditure, which the Society is probably not at
present in a position to incur, but it would certainly yield a very
rich return.

In addition to supplying information with regard to the
diseases and habits of the animals living in the Gardens, as was
the original intention of the scheme, it also provides oppor-
tunities of studying the parasitic fauna of the various countries
from which the animals come. This is a very important con-
sideration, for it is an unfortunate fact that in most of the large
faunistic expeditions which have hitherto been undertaken
practically no attempt has been made to deal with the internal
parasites. As important exceptions to this may be mentioned
the German Expedition to Spitzbergen in 1898, and the Swedish
Expedition to Egypt and the White Nile in 1901. By both of
these expeditions a large number of parasitic forms has been
collected, and the results have been published in a series of very
valuable monographs. Dr. Leiper has, at these meetings, on
more than one occasion called attention to this regrettable
neglect of the parasitic fauna, and has urged on collectors the
great service they would render by making even a cursory
attempt to collect parasitic worms.

During the past seventeen months a total of 198 animals have
been submitted to me for examination, the great majority of
which have been birds and reptiles. Of these, 87 were found to
be infected with worm parasites of one kind or another, 7.e.,
about 44 per cent. In most cases the infection was single, only
one species of parasite occurring ; but in an Indian Cobra as many
as six different species were found. As is generally the case,
Nematode infections were much more numerous than any others,
there being 60 cases. Trematodes were found in 28 cases, and
Cestodes in 15. The Trematodes belong to about 20 distinct:
species, a large proportion of which are new, and several of
which are new generic types,

858 ON WORM PARASITES.

From the reports of Dr. Beddard and Dr. Leiper I gather that
during the same period 41 additional cases of infection with tape-
worms, and 53 of infection with Nematodes have been observed.
To these must be added a very considerable number of infections
with larval Filariz observed by Mr. Plimmer. These figures,
excluding the larval Filarie, give a total of about 180, of which
the infection with Trematodes, Cestodes, and Nematodes is
respectively 28, 56, and 113. This appears to show that
Trematode parasites are much rarer than other forms, but that
is certainly not the case. It must be remembered that only a
comparatively small number of the animals passing through the
Prosectorium were adequately searched for Trematodes. The
extent to which Trematodes and the smaller intestinal Nematodes
escape notice except when thoroughly searched for may be
gathered from the following table.

Trematodes. Cestodes. Nematodes.

Total number of cases of infection 28 56 is
Number of cases found only on

further examination ............ 20 15 60
Rercenta ce mean earn 5 eee err 71 26 53

It may be added that the bulk of the Cestoda which were
not noted on first examination were small immature forms or
fragments of no value. It may therefore be safely affirmed that
only a negligible fraction of the mature tapeworms actually
escape detection in the Prosectorium. The matter, however, is
different with Trematodes ; and in view of the fact that only about
one-tenth of the total number of animals dying in the Gardens
have been submitted to a thorough examination, it must appear
that the number of Trematodes actually occurring is considerably
greater. Even allowing that they only occur in 5 per cent. of
cases, Which is taking an extremely low estimate, we ought to
have had instead of 28 records a total of well over 100. In the
same way the number of small intestinal Nematodes ought to
be much increased.

These remarks, however, are not intended as a criticism of the
efforts of the prosectorial staff. Parasitic worms are, after all,
only a very small part of the work of this department of the
Society, and as I have mentioned in the beginning of this com-
munication, considerable praise is due for the results actually
obtained. My chief intention is to show that parasitic worms
are undoubtedly an important concern and that their importance
justifies the attention which is being paid to them. Their claim
to notice had until recently been somewhat neglected by the
Society, and it seems necessary that someone should offer a plea
for their vindication. ‘There can be no doubt that the attention
which the Society, thanks to the efforts of Dr. Chalmers Mitchell,
is devoting to these forms will do much to benefit Zoology as a
whole.

ON NEW FOSSIL REPTILES. 859

On some new Fossil Reptiles from the Permian and
Triassic Beds of South Africa. By R. Broom, D.Sc.,
C.M.Z.8.

[Received May 13, 1912 : Read June 4, 1912. |
(Plates XC.—XCIIL.*)

INDEX.

Systematic : Page
Taurops macrodon, gen. et sp. n. ...............2..... 859
Galeops whaitsi, gen. et sp. n. ch R860,
Scymnognathus whaitsi, gen. ah SD Te aseeccararogona OIL
AMlurosaurus striatidens, sp. nM. ..............002...-.. 863
Pristerognathus platyrhinus, sp. n................06608. 863
Alopecorhinus parvidens, gen. et sp. n. ............... 864
Ictidognathus hemburyi, sp. n. ........................ 865
Endothiodon whaitsi, sp. N. .......2..0..e20ee seen. 866
E. platyceps, sp. n.. F Sees ca eae SOT
Prodicynodon enn ETS, | SPaeMlerec een ease eae OO
Dicynodon to include Oudenodon ..................:.. 868
Pristerodon to include Opisthoctenodon............... 868
Dicynodon laticeps, Sp. 0.............cceeeeereeeeeeeeeeees 868
1D), JOSIMIAXGOOS, El We, sonccoenoovmpedeodecnsudeopennondenbeoe (tos)
D. lutriceps, sp. n. Auchan seein leiiusisisdeaas cones OO.
Emydops minor, gen. wa Salley wesenecediste sce slemeaenian ef SUEL
Ictidopsis elegans, gen. et Sp. 1. ......:e:eeeeeeceeeeeeee 872
Nythosaurus browni, sp. n. He Been ccrimcl ae She
Endothiodon subdivided : see » Addanduna Peet MOTO
Emydochampsa, gen. 0..............:cseseceseeeereereeeess 875

The following series of new reptiles forms an important addition
to our knowledge of the Karroo faunas. The majority have
been found by the Rev. J. H. Whaits, of Beaufort West. Of
the others one was found by Mr. Alfred Brown, of Aliwal North,
one by Mr. H. J. Hembury, and a few by myself.

Suborder DINOCEPHALITIA.

TAUROPS MACRODON, gen. et sp.n. (PI. XC. fig. 1.)

The snout of this large Dinocephalian was obtained at Bos-
manshoek at the foot of the Komsberg. Though so little has
been obtained we have enough to give the distinguishing characters
of this new type. In size it almost equals Zapinocephalus
atherstonei, but the snout though narrower is considerably deeper,
and it differs in the great degree of development of the teeth,
especially of the incisors. Where the snout is broken across,
about the transverse plane of the back of the anterior nares, the
width is 190 mm. and the height about 100 mm. ‘The anterior
nares are unusually small and situated about 90 mm. from the front
of the snout. The distance between the two nostrils is about
45 mm. The condition of the bone renders it almost impossible
to make out the sutures, the only one that is distinct being that
between the two premaxillaries.

* For explanation of the Plates see p. 875.

860 DR. R. BROOM ON

The teeth are for the most part badly preserved, but as they
are broken off at different levels and as a large number of
replacing teeth are present, the structure can be made out without
much difticulty. There are 14 teeth preserved, of which the
first five are large and long. From the 6th backwards the teeth
steadily decrease in size and the crowns become quite short.
Possibly there are two or three teeth lost behind the 14th, but
it seems improbable that there are many. The anterior teeth
are of the usual Dinocephalian type. They are, except where the
root is being absorbed by a replacing tooth, of great length, the
whole root and crown being about 90 mm. The anterior cusp is
long and narrow, and on section is semicircular. At its base it
measures about 9 mm. across and gradually narrows towards the
point. None of the anterior upper teeth shows the whole anterior
cusp, but it probably measures about 20 mm. in length if not
more. The posterior basal cusp is about 15 mm. in width and
has a slightly concave surface. The posterior teeth have short
cusps. The 12th tooth has a crown only 12 mm. in height.

The teeth in the lower jaw are apparently closely similar in
type to those in the upper. Most of the functional teeth are
badly weathered away, but the weathering shows that at the base
of each is a well-developed replacing tooth similar in type to the
functional one. It seems probable that the long anterior cusps
of the front teeth interdigitated and that the evinding took place
by the meeting of broad posterior cusps. The edges ‘of the long
anterior cusps “would thus form an admirable cutting apparatus,
and the internal cusps would take the part of molars. In the
Dinocephalians almost the whole dental apparatus is placed in the
front of the snout, and the absence of grinding molars such as aré
seen in pe contemporaneous or slightly earlier American herbi-
vores, é6.g. Diadectes, is explained by the crushing or grinding
function Taine been taken up by the peculiarly specialiséd
incisors.

Suborder DROMASAURIA.

GALEoPS WiAirst, gen. et sp.n. (PI. XCI. fig. 6.)

This new génus is founded on the antetior half of the skeleton
of a Dromasaurian. The skull is Grushed but fairly well preserved,
arid the shoulder- -girdle and front limb are in good condition,
while remains of about 18 vertebre are seen. Wintor tunately the
matrix is very hard and little can be done in the way of develop=
ment. The specimen was found by Mr. Whaits on the farm
La-de-da, about 20 milés to the west of Beaufort West, and
probably belongs to the upper part of the Pareiasaurus- Zone.

The skull is in many respects very remarkable. The orbit is
exceedingly large and the temporal fossa deep and very narrow,
and the squamosal has a long descending process which brings
the quadrate below the back of the orbit. The face is very short,
and there appear to be no teeth in either upper or lower jaw.

The bones of the skull are not in good condition for showing

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sutures. The snout is missing in front of the nostril, but from
the shape of the lower jaw its length can be fairly well assumed.
The nostril is large and the distance between it and the orbit very
short. It is probable that the septomaxillary meets the lachrymal.
The suborbital and postorbital arches are slender. The frontal
and parietal regions are both fairly wide and there is a large oval
parietal foramen. The squamosal is somewhat like that of the
Dinocephalia, but is slender and the descending process very long.
There appears to be a distinct quadrato-jugal. The palate, so far
as preserved, agrees with that of the Therocephalia. There is a
pair of long slender prevomers, and the palatines and back of the
pterygoids are not unlike those of Seylacosaurus, but whether in
other respects the palate is Therocephalian or Dinocephalian the
evidence does not show. Certainly the palate is not the least
like that of the Anomodont. The lower jaw is short and tooth-
less. It agrees fairly closely with that of the Anomodont, but
there is a rudimentary coronoid process.

The shoulder-girdie has a large distinct precoracoid and coracoid
loosely articulated to each other. The scapula is somewhat like
that of the Dinocephalia and also a little like that of the Thero-
cephalian Jctidosuchus, but unlike that of the Anomodont. The
limb bones are long and slender.

Galeops is an entirely new type of Dromasaurian and represents
a new family, the Galeopide, characterised by the absence of
teeth and the presence of a small coronoid process. The other
family, which may be called the Galechiride, includes Galechirus
and Galepus, both with teeth and without a coronoid process.

Suborder THEROGCEPHALTIA.

ScYMNOGNATHUS WHAITSI, gen. et sp.n. (Pl. XC. figs. 4, 5.)

This interesting type was found by Mr. Whaits near Beaufort
West. Within quite a small area remains of four or five
animals were obtained. Unfortunately, most of the bones are
extremely weathered, and so infiltrated with lime as to be
practically limestone nodules so hard that any development is
almost impossible. Two skulls are sufticiently well preserved to
show the general characters of the genus, though it is impossible
to be sure of most of the sutures.

The new type is a very near ally of Gorgonops torvus Owen,
and it is only after considerable deliberation that I have decided
to place it in a new genus. With perfect certainty it can be
placed in the Family Gorgonopide. When Owen first described
Gorgonops in 1876 he unfortunately came to the conclusion that
the temporal fossa was roofed over as in Labyrinthodonts or
Pareiasaurus, and this mistake was also made by Lydekker in
1890, and by Seeley in 1895. On examining the type and, at
that time, only known specimen when in London three years ago,
I discovered that the temporal fossa is not roofed over, though
the parietal region is broad, and noted the observation in a paper

862 DR. R. BROOM ON

published in 1910*. Within the last few months Mr. Whaits
has fortunately discovered a good skull of Gorgonops which shows
that it has quite a large temporal opening, very similar to that
in the present genus.

In general appearance Scymmognathus differs from the more
typical Therocephalians mainly in having the intertemporal region
about as wide as the interorbital. The snout is long, the orbits
small, the temporal region fairly wide, and the squamosals more
powerfully developed than in most Therocephalians.

The total length of the skull is, in the type, 305 mm. In a
second but much crushed specimen the length is probably about
325 mm. From the front of the snout to the front of the orbit
the measurement in the type is 165 mm., and the length of the
orbit about 40mm. The interorbital width in the type is 70mm. :
in another very imperfect specimen it measures 78 mm., and ina
third about 76 mm. The narrowest part of the parietal region
measures in one of the larger specimens 78 mm.

The parietal foramen is situated well back and is 6 mm. in
diameter.

Though the limits of many of the bones cannot clearly be made
out, much of the cranial structure can be seen. The bones of the
snout seem to agree with those of the better known Thero-
cephalians, though the septomaxillary is relatively larger. The
jugal is unusually well developed and the postorbital is very large.
The postfrontal also seems to be much larger than in any other
type hitherto examined. The squamosal is more massive than in
most Therocephalians, and the quadrate is relatively small and
largely hidden by the squamosal. The occipital condyle is single.

The lower jaw is fairly similar to that previously described
and figured in Lycosuchus and Aloposaurus. The front of the
dentary is very deep and has a well-marked mental process.
Posteriorly the dentary has a short coronoid process and for a
considerable distance lies above the angular. The surangular is
much smaller than in other known Therocephalans.

There are five upper incisors, one large canine and three or four
molars. The incisors are long pointed teeth, oval in section, and
apparently without serrations. The space occupied by the five
incisors in three specimens 1s 37 mm., 39 mm., and 40mm. ‘The
space between the last incisor and the canine in four specimens is
18 mm., 22mm., 25mm.,and 26mm. The length of the canine
in seven specimens is 16 mm., 18 mm., 20 mm., 20 mm., 20 mm.,
20mm.,and 21mm. Behind the canine is a diastema varying from
7 mm. to 16 mm., followed by either three or four molars. The
number of molars apparently depends on age. When three molars
are present the distance occupied by them is 20 or 21 mm.: when
four are present they occupy 25 mm. to 29 mm.

1 3 or 4

3 or 4

The dental formula is, i. il

42 ¢

* “ Observations on some specimens of South African Fossil Reptiles preserved in
the British Museum.” ‘I'r. Roy. Soc. 8. Afr. vol. 1. pt. 1, 1910, p. 20.

NEW FOSSIL REPTILES. 863

Scymnognathus may prove to be related to Seymnosaurus.
The dental formule are practically the same in the two genera,
but Scymnosaurus seems to be a higher type of Therocephalian,
and the shape of the snout and lower jaw differs sufficiently
markedly to justify one in keeping the genera distinct.

ANLUROSAURUS STRIATIDENS, sp. n. (Pl. XCI. fig. 7.)

The imperfect snout which forms the type of this new species
was discovered by me at Kuilspoort, near Beaufort West, at a
horizon which is probably 500 feet above the town.

It is a smaller species than @. felinus, and in addition differs
from this and the other known species in the relatively small size
of the teeth, in the feeble development of the serrations, and in the
incisors and canine having feeble vertical ridges on the enamel.

So far as preserved there is a pretty close agreement in the
structure of the bones of the snout with those of previously
described species. The septomaxillary is larger than usual. The
mandible has a broad angular chin.

The incisors are small, rounded, pointed teeth, placed so near
each other as to be almost touching. Those whose crowns are
preserved (3rd, 4th, 5th) have the enamel folded into about half
a dozen vertical corrugations. ‘The 5th tooth has fine serrations
on its posterior edge. The space occupied by the five incisors is
16mm. The diastema between the 5th incisor and the canine is
7 mm.

The canine is long and slender. The base measures antero-
posteriorly 7 mm. and the height of the crown as preserved is
14 mm. It originally probably measured about 20 mm. The
molars are small with only faint indications of posterior serrations.
Two are well preserved and there are remains of the other two.
Probably 5 is the complete number, which would make the dental
formula agree with the other species of #lurosaurus. The four
preserved molars measure 10 mm., and the front one is only 9mm.
behind the canine.

PRISTEROGNATHUS PLATYRHINUS, sp.n. (Pl. XCI. fig. 8.)

The specimen which I take as the type of this new species is an
imperfect snout found by Mr. Whaits at Grootfontein, about 12
miles to the west of Beaufort West, and probably from the upper
part of the Pareiasawrus-Zone. The specimen consists of the
front half of the skull. It is broken into three pieces and the
upper nasal region is missing. The matrix is extremely hard and
ditticult to clear off, but most of the characters can be satisfactorily
made out.

In the large majority of Therocephalians the snout is deeper
than broad. In this specimen the snout is broad and flat, and the
lower jaw comparatively straight and with very little of the usual
upeurving in the canine and incisor region. The widest part of
the snout is immediately above the canine, where it measures
55 mm., and on the same plane the height of the snout is 40 mm.

864 DR. R. BROOM ON

There is very little crushing. Nearer the front of the nose the
height is only about 25 mm. and the width 45 mm. The
maxillary bone is rather deeply pitted. The palate is broad
and flat behind and in front slopes upwards to the opening of the
internal nares. The prevomers are narrow slender bones which
in section are seen to have thin vertical plates. The palatines
are apparently as in Scylacosaurus, but appear to approach each
other more closely in the middle line. The pterygoids have each
a very thin vertical plate which is closely placed against its
neighbour.

There are apparently 6 incisors, though only evidence of the
last five are preserved. The anterior ones are fairly round on
section, but the last three are more flattened and havea posterior
edge which is not improbably serrated. The teeth decrease in
size as we pass backwards, the 6th being only about half the size
of the 4th. The last four teeth occupy a space of 20°5 mm. and
the whole six probably 30 mm. ‘The diastema between the 6th
incisor and the canine is 9 mm. Thecanine at its base measures
10 mm. by 7mm. Behind the canine is a diastema of 7 mm.
followed by 8 pointed molars. Each of the anterior molars has
an antero-posterior diameter of nearly 3 mm., and the whole series
occupies a space of 29 mm.

The dentition of the lower jaw is mainly concealed in front.
Probably there are 3 incisors and 1 canine. The molars are well
shown on the right side and are 8 in number, and occupy a space
of 32mm. There is no evidence of any serrations, and were they
present the specimen would be cupecred to show some of them.
Stoo! ne:

There is some doubt whether this specimen belongs to the same
genus as Pristerognathus polyodon Seeley. The type is only the
front of a snout, and the number of molars is unknown, and only
a figure of the underside is given by Seeley. However, the two
specimens are probably from near the same horizon, and the
arrangement of the teeth, so far as known, is sufficiently close to
render it advisable to place this new species provisionally in
Seeley’s genus Pristerognathus.

The dental formula is probably i.

ALOPECORHINUS PARVIDENS, gen. et sp.n. (PI. XCI. fig. 9.)

This new genus is founded on an imperfect snout ee, by
Mr. Whaits at Beaufort West. It consists of the greater part of
the left maxillary and dentary and much of the right maxillary
and dentary.

It resembles Pristerognathus platyrhinus in the broad short
nose and: approximates in dental formula, but differs in having
a much more slender jaw and in the relatively smaller size of the
teeth, especially the molars, and in having a much shorter
precanine portion of the snout.

The anterior upper incisors are lost but there is evidence of the
last three. The whole series probably measured 19 mm, The

NEW FOSSIL REPTILES. 865

diastema between the last incisor and the canine is only 2 mm.
The canine measures at its base 6 mm. by 5mm. The molars
are at least 7 in number, and are small, pointed but apparently
unserrated teeth. The seven occupy 17°5 mm. Between the
canine and the first preserved molar is a diastema of 9 mm.

The lower incisors are preserved in section and are 4 in number,
the 4th being inside the line of the others. The canine is far
forward and small and rounded, the section measuring 3°5 mm.
in diameter.

The dental formula is probably i. _ é. 7 ann eS

P

IcTIDOGNATHUS HEMBURYI, sp. n. (PI, XCI. figs. 10, 11.)

This new species is founded on four imperfect snouts found by
Mr. H. J. Hembury at Beaufort West. There are minor
differences between the snouts probably due to crushing, to age,
and possibly to sex, but I believe they all belong to one species.
To avoid any possibility of confusion I shall take the skull
(fig. 10) as the type. It is the best preserved specimen but does
not show the molars satisfactorily.

As in Alopecorhinus the snout is broader than deep. This
is probably also the case in /ctidognathus parvidens, but the only
known specimen of this species is considerably crushed. From
the front of the snout to the front of the orbit the measurement
is 42 mm. ‘The antero-posterior diameter of the orbit is 18 mm.
The interorbital region is 19 mm. across. The measurement
across the snout at the canine region is about 27 mm.

The premaxillary is a very small bone forming the anterior and
lower margins of the nostril, It carries six small pointed, rounded,
smooth incisors. I fail to detect serrations on any of them. The
six incisors occupy a space of 10 mm.

The septomaxillary is unusually large and forms as large a part
of the facial surfaces as does the premaxillary. Thereis the same
foramen between it and the maxillary seen in typical Thero-
cephalians, but it is relatively smaller than in other forms.

The nasal bone is well developed and is interesting from the
fact that it is only very little broader behind than in front.

The maxillary is typically developed. As in so many Thero-
cephalians, the centre part of the bone is markedly pitted as if for
the accommodation of glands or sense organs. In the same region
there are numerous foramina passing into the bone, and some of
these foramina lead back into a canal in the bone. As there is
no large foramen in the maxillary bone which might be regarded
as the foramen for the maxillary branch of the Vth nerve, I
think it probable that this nerve subdivides in the maxillary
bone and comes to the surface by a number of small foramina,
and that the pits were for sense organs which were supplied by
this nerve. The tactile vibrisse of mammals, or the remarkable
sense organs of the beak of Ornithorhynchus may be the modified
homolognes of these supposed ancestral organs.

866 DR. R. BROOM ON

There are two canines—a very small anterior and a fairly large
long slender posterior one. The small canine has a diameter of
‘5 mm.: the large canine measures 3°5 mm. by 4mm. The two
are a little more than 2 mm. apart. The exact number of
molars is a little doubtful but appears to be 8, and they oceupy
16 mm. in one of the specimens. The molars appear to be
all short, smooth, pointed teeth without serrations.

Tn the lower jaw there are four incisors, a large canine, and eight

eo In
Ictidognathus parvidens there are certainly nine upper molars and

possibly ten, but as this species is certainly allied to J. parvidens
it seems better to keep both in the same genus.

- 6 2
molars. The dental formula would thus be 1. yy @ 7? M-

Suborder ANOMODONTIA.

* ENDOTHIODON WHAITSI, sp.n. (Pl. XCIII. fig. 18.)

This new species is only known by the skull, a few vertebre
and ribs, and a couple of limb-bones. Fortunately the skull is in
beautiful condition. The specimen was discovered by the Rev. J.
H. Whaits at Beaufort West. It isa near ally to HYndothiodon
uniseries Owen, and has the molars in a single row, but it
differs from EH. uniseries in a considerable number of points
besides in being nearly twice as large. In Hndothiodon uniseries
the greatest length of the skull is 360 mm.: in Hndothiodon
whaitsi the skull measures in length 570mm. With the ex-
ception of Oudenodon magnus it is the largest known Anomodont.
As I hope shortly to publish a full account of the genus Endo-
thiodon, 1 shall here merely give a preliminary description of this
interesting species.

The skull is narrow and deep. The greatest width across the
squamosals is probably about 360 mm., while the interorbital
region is only 140 mm. and the greatest width of the palate
150mm. There is a slight degree of crushing, but not such as to
make these measurements far wrong. ‘The notch in the pre-
maxilla for the point of the lower jaw is very deep and narrow.
The nostrils are large, measuring 65 mm. by 45mm. The nasals
do not overhang them as in #, wniseries, the whole width of the
nasals being only 85 mm. The orbit is situated 180 mm. behind
the front of the snout and measures from 60 to 65 mm. in
diameter.

The parietal crest forms a huge arch over the top of the head
about 300 mm. in length. Itis very narrow for its depth, and
is mainly formed by the postorbitals and the squamosals, the
parietals being relatively small. Near the union of the anterior
and middle third the two postorbitals are pushed apart by an
enormous development of the preparietal. This forms a pro-
minent boss 90 mm. in length and 50 mm. in width, and rising
20 mm. above the edge of the postorbitals. In itis situated the

* See also Addendum, p. 876.

NEW FOSSIL REPTILES. 867

parietal foramen, which is only about 10 mm. in diameter. This
is the more remarkable in that in Z. wniseries the foramen
measures nearly 20 mm. in diameter.

The zygomatic arch is very massive. The malar process of the
jugal is less marked than in the smaller species. The squamosal
extends forwards to below the postorbital arch. Anteriorly the
zygomatic arch measures 65 mm. in depth and posteriorly
80 mm.

The lower jaw is very powerful and measures 400 mm. in
greatest length.

* ENDOTHIODON PLATYCEPS, sp.n. (PI. XCIIT. fig. 19.)

This new species was also discovered by Mr. Whaits, near
Beaufort West. It is founded on a fairly complete but slightly
crushed and probably barely mature skull. The lower jaw is
complete, and the only important part missing from the skull is
the whole of the nasal region and the anterior part of the
frontal.

The greatest length of the skull is 275 mm., and the greatest
width about 215 mm.

The frontal region is flat, and the parietal crest instead of
vising up from this is continued straight back. There does not
appear to be a distinct postfrontal bone. The preparietal is large
and passes forwards for 35 mm. in front of the pineal foramen.
The foramen is 14 mm. wide and 8 mm. in antero-posterior
diameter. Behind it is a large boss formed by the preparietal.
There is nothing particularly noteworthy about the squamosals,
jugals, or occiput.

The dentition is better seen in the lower than in the upper
jaw. It is made up of aseries of 7 teeth in a fairly regular row.
Behind this the same row has 9 more teeth, but median to these
there appear others, so that the front half of the dentition is an
irregular single row, the back half an irregular double row.
Parts of the crowns of two teeth are preserved and there appear
to be no serrations on them. If this prove to be the case, this
species will require to be placed in a new genus, as /ndothiodon
bathystoma has the teeth markedly serrated in front and behind.

The lower jaw is pointed in front, but at the lower part of the
symphysis it is very broad. The back part of the dentary is
unusually slender. Hndothiodon platyceps difters from H. wniseries
and #. whaitsi in the double row of teeth behind and also in the
flat broad head; while from #. bathystoma it differs even more
markedly. The teeth are smaller than in the other species.

PRODICYNODON BEAUFORTENSIS, sp.n. (Pl. XCIII. fig. 21.)

This small species is founded on an imperfect skull obtained by
me at Kuilspoort at about the same horizon at which T'aognathus
megalodon was found. The specimen consists of the crushed ant-
orbital portion of a small skull with the front two-thirds of the

* See also Addendum, p. 875.

868 DR. R. BROOM ON ,

lower jaw in position. At first sight the skull might readily be
taken for that of a small Oudenodon, but there are two marked
differences. There are a number of small maxillary teeth, and
the lower jaw ends in front in a pointed beak which fits into a
deep depression in the premaxillary exactly as in Hndothiodon.
There is no tusk.

The nostril is 8 mm, in length by 7 mm. in depth. The
orbit is 12 mm. behind the nostril. The inter-orbital width is
probably 14 mm. Close to the outer edge of the maxillary bone
are at least two small smooth pointed molars, and other molars
are arranged in a row further in.

The lower jaw is almost typically Endothiodont,

The only form nearly related to the present one is that de-
scribed by me eight years ago as Prodicynodon pearstonensis. The
two agree in the arrangement and structure of the molars, but
differ markedly in the proportions of the head. P. pearstonensis
has a much broader snout and an enormous premaxillary. It
also has the orbit much further back. As neither type is in
good condition it 1s impossible to affirm with certainty that the
two species belong to the same genus, but at present it will be
eonvenient to keep them together.

When Prodicynodon was first described, and until recently, the
presence or absence of a tusk was believed to be a generic dis-
tinction. Dicynadon and Oudenodon were helieved to be distinct
genera, For the last five years the evidence has been steadily
aceumulating in favour of the tusk being merely a sexual character,
and now the evidence seems to be conclusive. Mr. Whaits has
collected a large number of the common little Beaufort West
Endothiodont Dialurodon whaitsi, and tusked and_ tuskless
specimens seem to be about equally common, while there are no
other characters to separate the specimens. Mr, D. M.S. Watson
has succeeded in obtaining two specimens of Oudenedon bolorhinus
recently described by me from Kuilspoort, at the same locality
as afforded the type, but one specimen is tusked and the other
tuskless,

This conelusion will necessitate the giving up of the genera
Oudenodon and Opisthoctenodon and placing the species of these
old genera under Dicynodon and Pristerodon, Fortunately very
little confusion will result,as I have for years assumed the
possibility of the tusk being merely sexual,

DIcYNODON LATICEPS, sp, n. (Pl, XCII. figs, 12, 13.)

This new species of Dicynodon is founded on a beautiful skull
obtained by Mr. J. H. Whaits on the Nieuwveld. With the
exception of the lower jaw being missing and the tips of the
maxillaries with most of the tusks being broken off, the skull may
be regarded as perfect. It belongs to the very unusual broad-
headed variety of which only a few specimensare known. When
viewed from above the resemblance is so close to Dicynodon
tigriceps Owen as to suggest that it might be a young specimen,

NEW FOSSIL REPTILES. 869

but the palatal view shows that the tusk is differently placed and
relatively very much larger.

The greatest length of the skull measured obliquely from the
snout to the back of the squamosal is 270 mm., and the greatest
width across the squamosal is also 270 mm.

The snout is very short and the nostrils completely roofed over
by the projecting nasals. The breadth across the nasals is 83 mm.
The premaxillary is broad and shallow. The maxilla isshort and
also shallow. ‘The tusk is large and situated right beneath the
orbit, and directed downwards. In diameter it measures 25 mm.
by 28 mm. at its base. The frontal region is broad and flat, the
narrowest part between the orbits being 63 mm. If there is a
distinct postfrontal it is very small. The parietal foramen is
large, measuring 15 mm. by 10 mm. The preparietal lies mainly
in front of it. The parietals are unusually large, and the posterior
branch of the postorbitals more slender than in most species.
The postorbital forms the front half of the long slender post-
orbital arch. The squamosal extends forwards below the post-
orbital arch and has a large articulation with the maxilla. The
pterygoids where they meet are broad and, except for the median
ridge, flat. Considering the great width of the skull the quadrates
are not far apart.

The general structure of the skull will be better understood
from the illustrations given.

DicyNopon Psirracops, sp.n. (PI. XCII. fig. 17.)

In working at the troublesome genus Dicynedon we have con-
stantly been in doubt as to whether the small Dicynodon specimens
are distinct species or only younganimals. Frequently they agree
sufficiently in general shape and structure as to suggest the
probability of their being young = specimens of Dicynodon
leoniceps or some other large species. But this is certainly not
always the case. Many specimens of Dicynodon jouberti are
known from the Pareiasawrus horizon and all small, while no
large Dicynodon is known to occur in the same zone. At Beaufort
West a considerable number of specimens of a small Dicynodon
also occurs, but there is no evidence of any large Dicynodon
having lived at the period. The large Anomodonts are the
Endothiodons. There is thus satisfactory evidence that the
common Dicynodon of the Beaufort West commonage is an adult
animal, and apparently a new species. ‘The best specimens are a
good skull with much of the skeleton and a fairly good skull with
nearly the complete skeleton. I take the latter as the type.
Both specimens were obtained by Mr. J. H. Whaits.

The greatest length of the type skull is 102 mm., and of the
second specimen 112 mm. The width of the type across the
squamosals is about 66 mm. Across the maxilla the maximum
width is 42 mm. The interorbital width is 20 mm., and the
intertemporal 18 mm.

The following are the most noteworthy characteristics of the
Proc. Zoon, Soc.—1912, No. LVIII. 58

870 DR. R. BROOM ON

species. The nasals are so narrow that the nostrils look almost
directly upwards. The upper part of the nasals is thickened,
and the prefrontal region of the orbital margin is also elevated.
The frontal region is broad and flat. The parietal foramen is
situated in an elevated pyeparietal. The postorbitals approach
each other behind the foramen and nearly touch, forming a
parietal ridge. The tusk is small and directed forwards and
downwards. The lower jaw has the front portion unusually
broad and deep. The foramen behind the dentary is very
small.

The whole skeleton from the snout to the end of the tail
probably measures 500 mm.; the humerus measures 50 mm. and
the femur 58 mm.

DICcYNODON LUTRICEPS, sp. n. (PI. XCII. figs. 14-16.)

The type of this new species is an imperfect skull found by me
at Kuilspoort, Beaufort West district. The skull has lost the tip
of the beak, the postorbital and zygomatic arches, and there is
about 20 mm. missing from the postpterygoid and from the
parietal region so that the contact between the occipital and
anterior portions of the skull is lost, but otherwise the skull is
complete.

The most noteworthy characters of the type are the relative
shortness of the beak, the broad concave frontal region, and the
broad flattened intertemporal region, the upper surface of which
is almost entirely formed by the postorbitals. In a maxillary
from the same locality, and believed to be of the same species, the
tusk is feeble and directed downwards and slightly forwards.

The greatest length of the skull from the snout to the back of
the squamosal is about 195 mm., and the greatest width across the
squamosals is 145 mm. From the front of the beak to the orbit
is probably about 48 mm., and the antero-posterior diameter of
the orbit is about 43 mm. The interorbital width is 35 mm., and
the intertemporal region 29 mm.

The type specimen is tuskless. The caniniform process is
directed downwards and forwards and has a marked low outer
ridge which passes upwards towards the jugal arch. Below the
nostril is an anterior ridge parallel to the other and forming a
well-marked valley between the two. The nostril is fairly large
and the nasal is considerably thickened above it, forming an
overhanging supranasal ridge.

The prefrontals are small, but the frontals are well developed.
Posteriorly they enclose between them the small preparietal and
meet the anterior ends of the parietals. The postfrontals are
long and narrow. ‘The preparietal is small and seems to lie
entirely in front of the pineal foramen. The postorbitals are
very large where they overlap the parietals, but the postorbital
arch is unusually feeble.

The parietals are large and powerful, but are almost completely

a

NEW FOSSIL REPTILES. 871

hidden by the postorbitals. There has been a little doubt as to
whether the pair of bones usually supposed (Seeley, Broom, etc.)
to be parietals are really the parietals, or whether the median
bone in which lies the pineal foramen, and usually called the
preparietal (Seeley, Broom) or interparietal (Newton), may perhaps
be the true parietal. This latter view has recently received the
support of Jaekel. The median bone, called for convenience
preparietal, is met with in most Anomodonts. In some (/ndo-
thiodon) it is very large; in others (Cistecephalus) it is quite
absent. The size of the paired bones depends to a considerable
extent on the development of the preparietal. In Cistecephalus
there can hardly be any doubt that the large pair of bones behind
the frontals are the parietals. They have the same relations to
the squamosals, interpavietal, frontals, and postorbitals as the
parietals have in most reptiles, and there can be, I think, no
reasonable doubt but that these bones are homologous with the
parietals of the mammals. When the preparietal appears and
the intertemporal region becomes narrowed the parietals are
much reduced in front, but posteriorly the relations to the
squamosals, interparietal, and postorbitals remain constant. In
Endothiodon the preparietal is so large that the parietal seems to
be completely separated from it by the frontal. In Dicynodon,
as exemplified by this skull, the parietals still meet the frontals.
What the preparietal is, is not clear. It certainly is not the
interparietal. I am inclined to look on it as a neomorph
developed in connection with the pineal eye. There is no trace
ot it known in Dinocephalians, Dromasaurians, Pelycosaurs,
Therocephalians, or Cynodonts, though in some of these the
pineal eye was probably as large as even in Hndothiodon.

There is little of special note in the palate or occiput.

The horizon from which the specimen was obtained is probably
about 300 feet above that of Beaufort West.

EMYDOPS MINOR, gen. et sp.n. (Pl. XCIII. fig. 20.)

When Owen, in 1876, described the specimens of Cistecephalus
in the British Museum, he named one species Wistecephalus
arctatus and referred two specimens to it. The type differs very
considerably from Cistecephalus microrhinus, the type species of
the genus, and Lydekker in his Catalogue places C. arctatus
doubtfully under Cistecephalus. On more than one occasion I
have also expressed the opinion that C. arctatus does not belong
to Cistecephalus.

Recently, I discovered at Kuilspoort a small imperfect skull
which apparently belongs to the same genus as Owen’s C. arctatus,
though a distinct species. Pretty certainly the genus is not
Cistecephalus, and the question arises, is 1t Ouwdenodon, or rather
Dicynodon? The only specimens known are tuskless, and there
are apparently no molar teeth. In the imperfect state of the
specimens it is impossible to clearly differentiate the genus from

do*

872 DR. R. BROOM ON

Dicynodon at present, but there seems little doubt they represent,
if not a distinct genus, at least a subgenus. The more note-
worthy characters are the wide parietal region with large
parietals, slender postorbital arch, and feeble beak.

The length of the skull is probably about 45 mm., and the
ereatest breadth about 30 mm. The orbit measures 12 mm. in
diameter. The intertemporal region is 14 mm. across.

The beak is in very imperfect condition, and little can be made
out with certainty as to its structure. It may be stated with
confidence that 1t was short.

The frontals are large, and form an interorbital region 10 mm.
wide. From near the supraorbital margin to the anterior end of
the parietal there runs backwards and slightly inwards a shallow
groove. ‘There is a moderate sized triangular postfrontal and
a large median preparietal. ‘This latter seems to lie entirely in
front of the pineal foramen. The parietals are large and form
the greater part of the broad intertemporal region. The post-
orbital is long and slender. It forms a feeble postorbital arch
and the inner margin of the temporal fossa. The squamosal is of
the typical Anomodont type. Its zygomatic position extends
forwards to below the orbit. The articular region is badly
preserved.

The lower jaw is very like that of Oudenodon, but the beak
portion is small, and probably little more than the symphyseal
region was covered with horn.

Suborder CYNODONTIA.

IcTIDOPSIS ELEGANS, gen. et sp.n. (Pl. XCIII. fig. 22.)

This new genus and species is founded on a nearly perfect little
skull obtained at Harrismith, Orange River Colony. It is a very
near ally of Vythosaurus larvatus Owen, but is much smaller, and
differs in the number of molars and in other cranial characters.

In general shape the skull agrees fairly well with Vythosaurus.
The orbit is near the middle of the skull and relatively larger than
in the better known genus, while the jugal arch is more slender.
In /ctidopsis the snout is shorter, and the molars are 6 in number
instead of 7.

The greatest length of the skull is probably 63 mm. and the
width is 42 mm. ‘The interorbital width is 12°5 mm.

The premaxillary bone is badly preserved, but it 1s manifest that
there are four mcisors. The first three incisors are moderately
round, but the last is more flattened. There do not appear to
be any posterior serrations.

The maxillary is relatively shorter than in Vythosaurus, and
deeper. Above the canine, and in front of the lachrymal are
little elevations of the bone. On the canine elevated area are
three small foramina and near the root of the 3rd molar two other

en ee

NEW FOSSIL REPTILES. 873

foramina, while two more are near the anterior end of the bone.
As there is no single large supra-maxillary foramen, it is probable
that all these small foramina are for branches of the maxillary
nerve. The canine is long and slender. It is ridged somewhat
after the manner of the canine of the cat. Behind it are 6 molars.
The 1st is small, pointed, and without any cusps. The 2nd, 3rd,
4th, and 5th all closely resemble one another. ‘There is a large
pointed median cusp and a small anterior and posterior cusp.
While essentially similar in type to the molars of NVythosaurus
they differ in that the anterior and posterior cusps are relatively

smaller. The 6th molar is a small tooth, and unfortunately the
crown has been lost from both sides of the skull.

The incisors measure about 6°5 mm. Behind the last incisor
is a diastema of 4mm. The canine measures antero-posteriorly
about 2°5 and its height is 7 mm. Ata distance of 1-5 mm.
behind the canine is the Ist molar, and the whole series of six
occupies the space of 13 mm.

Only a small fragment of the septo-maxillary is preserved, but
it manifestly forms part of the face, and was probably as in
Nythosaurus.

The nasal is moderately wide in front, but narrows on passing
backwards, and then near its middle it becomes about twice as
wide as in front. Round the bone near where it meets the
maxilla is a series of three or four foramina.

The lachrymal forms the front of the orbit and, as in Vytho-
saurus, it is larger than the prefrontal.

As in Vythosaurus and most Cynodonts the prefrontal meets
the postorbital, shutting out the frontal from the orbital margin.
The postorbital forms about half of the postorbital arch and
overlaps a small part of the parietal.

The parietal is large, and there is an obvious pineal foramen.

The squamosal is like that of Wythosaurus, except that in
Ictadopsis there is a much more prominent auditory groove.

The fractured edge of the occipital crest shows the interparietal
distinct from the parietals, and the lateral bone, which I believe
to be the opisthotic, distinct from the parietal, the interparietal,
and from the squamosal.

The occiput and palate have not been cleared.

The dentaries are in position, but the posterior bones of the
jaw have been detached and displaced, probably by insects before
the skull was fossilized.

The type of Jctidopsis elegans was found at Harrismithinassociation
with Zystrosaurus. Unfortunately the geology of Harrismith is
unknown. Some of the first specimens of South-African Dinosaurs
were got there by Mr. J. M. Orpen in 1853, and there can be no
doubt that these are from the Red Beds of the Stormberg Series.
It is not a little startling to find that the same commonage
yields fossils which in Cape Colony belong to a horizon about
3000-4000 feet lower than the Red Beds. Probably the Molteno

874 DR. R. BROOM ON

Beds and the Burghersdorp Beds are greatly thinned out, or
possibly there is an unconformity.

Some years ago I divided the Upper Beaufort, or Triassic Beds,
into three zones: (1) the Lystrosawrus-Zone, (2) the Procolophon-
Zone, and (3) the Cynognathus-Zone. While these zones in the
main hold good, the limits of them are still unknown. In the
extensive Jystroswauruce beds of Colesburg, Middelburg, and
Cradock no Procolophons or Cynodonts are known; but Mr. D.
M. 8S. Watson has recently found Lystrosaurus associated with
Cynodonts to the west of Burghersdorp, and a similar association
we now know occurs at Harrismith. Again, while no Lystro-
saurus or Cynodont remains are certainly known from the
Procolophon beds, the Procolophon-like genus Thelognathus occurs
at Aliwal North with Cynodonts, and possibly Procolophon itself.
It seems not improbable that later work will further subdivide
the Triassic Beds, each zone counting from the point where a
new type begins, but before we can make any further advance
with confidence we require to have a much fuller knowledge of
the distribution of the fossils of the Upper and Middle Trias.

In the meantime I think we are safe in stating that Vytho-
saurus, Ictidopsis, and probably Galesaurus come from an older
zone than the Cynognathus beds. None of the known specimens
have been found near Burghersdorp, nor have any traces of
Lystrosaurus ever been found, from which we may conclude that
Lystrosaurus and probably these small Cynodonts became extinct
before Cynognathus appeared.

NyTHOSAURUS BROWNI, sp.n. (Pl. XCIII. fig. 23.)

This new species is founded on an imperfect lower jaw obtained
by Mr. Alfred Brown at Aliwal North. A large part of both
dentaries is present, but most of the symphyseal portion 1s missing
with the canines and incisors. Five molars are well preserved on
the left side and three on the right.

While in a number of respects the specimen differs from
Nythosaurus larvatus Owen, it seems probable from the position
of the symphysis that there were seven molars, and as in general
structure the molars agree with those of Vythosawrus larvatus, we
may consider it as probable that the species belongs to this genus.

The dentary differs from that of Vythosaurus larvatus in haying
a more slender horizontal ramus, in having a more marked angle,
and in the ascending ramus passing more upwards.

The molars preserved are probably the 3rd to the 7th. All
have three cusps. What is apparently the 3rd molar has the
median cusp short, and the others only feebly developed. In the
Ath and the other later molars the middle cusp is about twice as
long as the anterior and posterior cusps. In the second last
molar there is an additional small anterior cusp which gives it
four cusps, but there appear to be only three cusps in the last
molar. The five molars occupy 13°5 mm.

¥
:
:
t
«

NEW FOSSIL REPTILES. 875

Though the crowns of the molars resemble considerably those
of the Triconodont mammals there is the marked difference that
in Vythosaurus there is no trace of a cingulum, and there is

apparently only a simple root to each tooth.

AppEenpuM (29th July, 1912).—Since the above paper was
read evidence has been obtained which shows that the genus
Fndothiodon must be subdivided. Seeley twenty years ago
placed Hnrdothiodon uniseries in a distinct genus, Hsoterodon, and
until quite recently I have followed him in this. A fragmentary
maxilla recently found by Mr. Whaits shows that Seeley was
correct. In Hndothiodon bathystoma the teeth have long pointed
crowns with coarse serrations both in front and behind. In
Endothiodon uniseries the teeth have flattened crowns serrated
only behind, and in Hndothiodon platyceps there are no serrations
on either side. This latter point has been confirmed in a second
specimen. Until the crowns of the teeth of Hndothiodon whaitsi
are known we may provisionally place it with Hndothiodon unt-
series. ‘The group would thus be formed of

Endothiodon bathystoma Owen.
Esoterodon uniseries Owen.

Esoterodon whaitst Broom.
Emydochampsa platyceps Broom, gen. nov.

EXPLANATION OF THE PLATES.
PuLatE XC.

Side view of snout of Taurops macrodon Broom. Nearly 4 nat. size.

. Side view of tooth of probably Hecasaurus priscus Broom. Nat. size.
This tooth, though not associated with the type of Hecasaurus priscus,
is from the same horizon, and as it is the tooth of a Dinocephalian of
the sizeof Hecasaurus it very probably belongs to this genus and species.

Fig. 3. Upper view of tooth of probably Hecasaurus priscus Broom. Nat. size.

Fig. 4. Side view of skull of Scymnognathus whaitsi Broom. A little over 4 nat.

size. The skull is much flattened and distorted by crushing.

Fig. 5. Lower jaw of Scymnognathus whaitsi Broom. + nat. size. Though this

jaw is rather smaller than that of the type specimen it almost certainly

belongs to this species. It may have belonged to a young animal or to a

female. ‘The specimen is considerably crushed.

Pig.

Fig.

ire

Puate XCi.

Fig. 6. Side view of the skull of Galeops whaitsi Broom. Nat. size. The
specimen is slightly crushed and considerably weathered. ‘he orbital
margins and the preorbital portion of the skull and the lower jaw are the
outer surface of the bones viewed from within. The back portion of the
skull shows the quadrate and parts of the squamosal and opisthotic. A
sclerotic plate is seen in the orbit.

Fig. 7. Side view of snout of lurosaurus striatidens Broom. Nat. size. The
specimen is considerably crushed.

Fig. 8. Side view of snout of Pristerognathus platyrhinus Broom. 4%; nat. size.

9. Under view of snout and lower jaw of Alopecorhinus parvidens Broom.
42 nat. size.

Fig. 10. Upper view of imperfect skull of TIctidognathus hemburyi Broom.
Nat. size.

Wig. 11. Under view of snout of Ictidognathus hemburyi Broom. Nat. size.

876 PROF, S. J. HICKSON ON THE

PratEe XCII.

Fig. 12. Upper view of skull of Dicynodon laticeps Broom. 3% nat. size.

Fie. 13. Under view of snout of Dicynodon laticeps Broom. 5 nat. size.

Fig. 14. Side view of skull of Dicynodon lutriceps Broom. 7 nat. size. The
contact between the front part of the skull and the occiput is missing,
but the relation of the parts must be almost as restored.

Fig. 15. Upper view of skull of Dicynodon lutriceps Broom. About +5 nat. size.

Fie. 16. Palatal view of skull of Dicynodon lutriceps Broom. About 3; nat. size.

Fig. 17. Upper view of skull of Dicynodon psittacops Broom. About 5 nat. size.

Prate XCIII.

Fig. 18. Side view of skull of Hndothiodon whaitsi Broom. About 3 nat. size.

Fig.19. Upper view of lower jaw of EHndothiodon platyceps Broom. About 75
nat. size.

Vig. 20. Upper view of skull of Himadops minor Broom. # nat. size.

Fig. 21. Side view of skull of Prodicynodon beaufortensis Broom. About 7% nat.
size.

Fig. 22. Upper view of skull of Ictidopsis elegans Broom. % nat. size.

Fig. 23. Side view of left dentary of Nythosaurus browni Broom. % nat. size.

53. On the Hydrocoralline Genus, Hrrina. By Professor
S. J. Hickson, F.R.S., F.Z8., The University of

Manchester.
[ Received April 29, 1912: Read June 4, 1912.]

(Plates XCIV.—XCVI.)

INDEX.
Page
Revision of the genera Hrrina, Labiopora,
ESF OCC) CER” cise heen bciconbed Hap eeneeiida ale ar@andana aco iMcny
Description of new species ........... 882

Notes on the structure of Hrrina nove zelandie . 884

Note on geographical distribution ..................... 886
Description of old specieS ..........s00seeeeeeeeeeeeeess 887
General remarks on structure ...............0cceceeeseee 891
Dist of Known Species ......-.....ccncosseeeeneereeseencee 898
Additiowall motes eri... scc-weseee.deeeheseeeeeeeereeeeeaeee BOO
Diteratinre 28 ee a es Seen een Oa
Explanation of the Plates ...........:..0ceeseseseeeeees 895

Four years ago Professor Benham sent to me some specimens
of Stylasterina from New Zealand, with a request that I would
name them for him. At the same time he forwarded some notes
and drawings which have proved to be of considerable assistance
to me in working out their details. I am also indebted to
Professor Dendy for the loan of another specimen also from New
Zealand ; to the late Mr. Morgan, of Worthing, for the loan of a
specimen from an unknown locality; and to Mr. Gilchrist for
a specimen from the Cape of Good Hope.

All these specimens clearly belong to one of three genera,
Errina, Labiopora or Spinipora, as they exhibit the characters
that these genera exhibit in common and by which they can be
separated from other Stylasterina. These characters are: (1) a

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HYDEROIDS OF THE GENUS HRAINA-.

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HYDROCORALLINE GENUS ERRINA. 877

style in the Gasteropores ; (2) no style in the Dactylopores; and
(3) the Dactylopores, or some of the Dactylopores, guarded by a
grooved lip or spine—called by Moseley the “ nariform process.”
To any one acquainted with Moseley’s classical memoir of the
Stylasterina (8), and with the subsequent literature, which is not
very extensive, it might seem a simple matter to determine
whether the specimens about to be described belong to any one
of the three genera or not; because, according to Moseley’s
descriptions, the following characters were diagnostic :—

Errina, with only one kind of Dactylopore.

Labiopora, with two kinds of Dactylopores. The larger kind
of dactylopore with a nariform process * and arranged in
rows. The smaller kind of dactylopore, without a nari-
form process, arranged between the rows.

Spinipora, also with two kinds of Dactylopores. The larger
kind of dactylopore with long grooved spines, not arranged
in rows. The smaller kind of dactylopore at the base of
the larger ones.

Moseley’s memoir was published in 1881, and since that date
only four new species of Hrrina and one new species of Labiopora
(Z. moseleyi) have been described; but owing to the rarity
of these Hydrocorallines in the seas that have been recently
investigated, very little progress has been made in our knowledge
of them.

The specimens from New Zealand and elsewhere that I have
examined convinced me that a thorough revision of the genera
was necessary, and consequently the task of naming Professor
Benham’s specimens has taken me much longer than J anticipated,
The general results of my investigations have been to show that
the limits or frontiers between the three genera are ill-defined
and that it is necessary to accept von Marenzeller’s (6) proposal
to unite Labiopora with Hrrina.

In the first place, I have found, as von Marenzeller (6) has
done, that the presence of two kinds of dactylopores is not a very
reliable character; because in some forms that are otherwise
closely related the dactylopores of the smaller kind that have no
grooved spines may be numerous, scarce, or altogether absent
(e. g., Hrrina nove zelandie). A genus such as Labiopora
cannot therefore be absolutely separated from Hrrina by the
character of the dimorphism of the dactylozooids.

Moreover, the arrangement of the grooved spines in definite
rows is another character that is subject to considerable variations
and cannot be relied upon for diagnostic purposes. The arrange-
ment of these processes seems to be correlated in some way with
the method of growth of the hydrophytum as a whole, and that
is again, I believe, dependent upon the conditions of the en-
vironment.

* The term “nariform process’ introduced by Moseley is not very convenient, and

I have consequently used the the expression “grooved spine” for the ccoenosteal
processes that shelter the dactylozooids.

878 PROF, S. J. HICKSON ON THE

The characters that appear to me of more importance and to

signify a more profound differentiation are based upon :—
1. The texture of the surface of the corallum.
2. The aspect of the grooves in the nariform processes.

Moseley described the surface of Hrrina as composed of a
coinpact, hard, glistening, white, caleareous tissue, and he adds
that the canals generally are in this genus larger in proportion to
the size of the zooids than in most other forms, and the mesh-
works formed by them are comparatively wide open. Associated
with these two characters we usually find that the surface is
marked by shallow longitudinal grooves perforated by a series
of small apertures through which the vertical canals pass. These
cceenosteal pores are well defined in the species of this group that
I have examined, and in the type specimens of E. labiata and
LH. ramosa they are about ‘05 mm, in diameter. The surface of
Spinipora is hard and compact as in Errina, and it is also
perforated by well defined ccenosteal pores.

In the original description of the genus Labiopora, Moseley
describes the ccenosteum as being “‘ minutely reticulate in tex-
ture,” and in the description of Labiepora moseleyi Ridley (10)
also describes the surface as being ‘‘ minutely reticulate.” In all
the specimens I have examined there is a very marked contrast
between the surface of the specimens now included in the Zabio-
pore group of species and of those I propose to retain in the
Hrring group.

I should prefer to describe the surface of the former group of
species as being “ granular” rather than “reticulate.” In Hrrina
(Labiopora) capensis it is coarsely granular and in all the other

species of the group it is minutely granular (cf Pl. XCVI.
figs. 14 & 15). Below the surface, the ccenosteum is minutely
reticulate, being perforated by a network of small-meshed canal
passages (fig. 14), in contrast with the wide-meshed canal
passages of the Hrrina group.

As regards the grooved spines. In the following species that
I have examined the groove in the spine that protects a dactylo-
pore is turned towards the apex of the branch on which it is
situated: HH. labiata, HL. horrida, and EL. ramosa. The groove has
the same aspect in Z. glabra and HL. carinata, if we may judge
from the figures given by Pourtales (9).

In the only two specimens of the genus Spinipora that are
known the grooves also turn towards the apex of the branch
(Pl. XCV. fig. 8). In the Zabiopora group of species there is
considerable variation in the direction of the grooves. In some
of them, all or nearly all the grooves are turned away from the
apex (Pl. XCVI. fig. 11), but in others the grooves are turned
in all directions, the grooved spines forming irregular clusters on
the surface (Pl. XCVI. figs. 12 & 13). The same arrangement of
the grooves occurs in Hrrina gracilis and in Hrrina macrogastra,
according to von Marenzeller, who writes “‘ Ihre Oeffnung ist
nach hinten gerichtet, selten seitlich” (5), and also in Hrrine
jissurata of Gray (2).

~~.

HYDROCORALLINE GENUS ERRINA, 879

Taking these characters as guides it seemed to me that the
diagnostic characters of the three genera might be stated as
follows :—

(a) Ceenosteum hard and compact, perforated by well-
defined ccenosteal pores. Grooved spines turned
towards the apex of the branch.

1. With short grooved spines and only one kind of

daicty lope nee ny Ae) 225/00) shi Reena Errina.
2. With long grooved spines and two kinds of
GaiCtylOMOMestMaan seas cide Medes ct ttdae teeta or Spinipora.

(6) Coenosteum granular and reticulate, without well-
defined coenosteal pores. Grooved spines turned
away from the apex of the branches or irregularly
TIAGO CK Trac vaaeRieeate tase Sot uwMioe sled shlcroiald ectuanclanle Labiopora.

If the three genera be joined together to form a subfamily, the
Errinina, this subfamily might be defined as follows :—

Hydrophytum arborescent and irregularly flabelliform, gastero-
pores and dactylopores not arranged in cyclosystems. Gastero-
pores with a large brush-like style. Dactylopores without a
style. Some of the dactylopores protected by a grooved spine
(narial process) on the surface of the coenosteum,

This arrangement of the genera, however, breaks down on
further analysis, and I see no cther course than to arrange all the
species in three groups under the one generic name Lrrina.

The genus Hrrina was founded by Gray in 1835 for a species
of coral found in the Mediterranean Sea and formerly called
Millepora aspera by Linneus.

As von Marenzeller has pointed out, Gray’s description of the
spines in this species as *‘Superne longitudinaliter fisse” is not
consistent with the description of the species known to Linneus
and Esper.

I have examined the type specimens in the British Museum,
and have found that Gray’s description is not correct. The
spines in these specimens are irregular in arrangement, but where
they are isolated and not in clusters the groove is directed away
from the apex. Moreover, the character of the surface of the
ccoenosteum, the presence of a few small dactylopores without
grooved spines, and other features prove that this species is more
closely related to the type species of Labiopora than it is to any
of the other species of Hrrina.

According to the system I had, at first, proposed the type
species of Hrrima would thus be a species of Labiopora and
Moseley’s Hrrina ramosa would become the type species of the
genus. 3

Such a proposal, therefore, would not only be contrary to the
rule of zoological nomenclature, but it would also be extremely
inconvenient. Moreover, one species at least (#. macrogastra)
would occupy an intermediate position, having a surface similar

880 PROF. S. J. HICKSON ON THE

to that of Hrrina and grooved spines similar to those of
Labdiopora.

A plea might still be made to keep the genus Spinipora distinct,
but I am convinced that intermediate forms will be found
between the deep-sea species of Hrrinw and the only known
species of Spintpora, and that sooner or later it will be found
impossible to keep it apart from the others.

In the following pages, therefore, I have regarded all the
species that have been attributed to the three genera as belonging
to the one genus, Hrrina of Gray, but for convenience of welievemce
I have added after the generic name (Labiopora) or (L.) in the
case of those species that were formerly described as belonging to
the genus Labiepora and to others that belong to that group of
species, and (Spinipora) or (S.) in the case of Hrrina echinata, the
only known species of the Spinipora group.

Genus Errina Gray.
With the characters of the subfamily Errinina (p. 879).
The “ Hrrina” group of species.

Cenosteum hard and compact, perforated at the surface by
well-defined ccenosteal pores usually arranged in rows in shallow
longitudinal surface-grooves. Gasteropores with or without a
scale. All the dactylopores protected by short grooved spines
(narial processes) with the grooves turned towards the apex of
the branch.

The only species of this group that have been sufficiently well
described to make identification possible without reference to the
type specimens, are Hrrina labiata Moseley and H. ramosa
Hickson & England and #. horrida H. & HK. Other species
are LHrrina carinata Pourtales and #. pourtalesii Dall.
Pourtales in 1871 described three species, which at first he placed
in the genus Hrrina but subsequently transferred them to a new
genus, Lepidopora. These species were referred back again to
Errina by Moseley. Their names are /#. glabra, E. cochleata, and
EH. dabneyi. ‘These three species were distinguished from Hrrina
by the presence of a lip or lid-like process similar to that of
Cryptohelia hanging over the gasteropores. Wrrina fissurata of
Gray may have been a specimen of Labiopora, but as the original
specimen hag been lost it is useless to speculate on its supposed
affinities.

The species may be arranged as follows :—

* q@. Gasteropores with a definite lip or scale :—
Li. glabra, EL. cochleata, H. dabneyi, and EL. ramosa.
6. Gasteropores in the angles formed by the branches :—
L. horrida.
e. Gasteropores without scales and distributed on the surface
of the coenosteum :—
L. labiata, FE. carinata, FL. pourtalesii (?).

* See Note p. 894.

HYDROCORALLINE GENUS ERRINA. 881i

As regards the distribution it may be said that all the species
are inhabitants of deep water (?. e. 50-600 fathoms). Being
deep-sea species they are probably widely distributed, but at
present H. pourtalesii, H. ramosa, and HL. horrida have only been
found in the Pacific Ocean and Malay Archipelago, and the
remaining species in the Atlantic Ocean.

Spinipora group of species.

Coonosteum hard and compact, perforated at the surface by
well-defined ccenosteal pores usually arranged in shallow branching
grooves irregularly arranged on the surface. Gasteropores
without a scale. Dactylopores of two kinds. The larger kind
being guarded by long grooved spines (narial processes) crowded
and overlapping on the terminal branches, often worn down
short and separated by considerable intervals on the stems and
older branches (Pl. XCV. fig. 8). All the grooves of these
spines turned towards the apex of the branch. Smaller dactylo-
pores not protected by grooved spines, but scattered between and
on the projections that guard the larger dactylopores.

The Spinipora group is represented by only one species,
Errina (S.) echinata Moseley, and this species has been found off
Rio de la Plata in 600 fathoms by the ‘ Challenger ’ Expedition,
and off Providence Island in the Indian Ocean in 75 fathoms by
Professor Stanley Gardiner (4).

Labiopora group of species.

Ceenosteum granular and minutely reticulate, without clearly
defined ccenosteal pores. Gasteropores without scales but some-
times protected by short grooved projections or “lips.” Dactylo-
pores protected by grooved spines of variable lengths, the grooves
turned away from the apex of the branch or irregularly in all
directions. Some of the dactylopores usually without grooved
spines.

The species of this group that have been described already
are Hrrina aspera Gray, Hrrina (Labiopora) antarctica Gray,
Errina (L.) moseleyit Ridley, and Hrrina gracilis von Maren-
zeller.

Errina aspera was the name given by Gray to the Linnean
species Millepora aspera from the Mediteranean Sea. Hrrina (L.)
antarcticé was originally described by Gray as a Polyzoon
(Porella antarctica), but was subsequently redeseribed and figured
by Moseley as the type species of Labiopora. The type specimen
was found off the Hast coast of Tierra del Fuego, but a second’
specimen was discovered by the ‘ Alert’ Expedition in 30 fathoms
off 8.W. Chili and described by Ridley (10). Hina (L.) mose-
leyi was found by the ‘ Alert’ Expedition off the same cosat in
2-10 fathoms. Hrrina gracilis was found off the pack ice in the
Antarctic Ocean in deep water (von Marenzeller) (6).

I have had an opportunity of carefully examining a piece of

882 PROF. S. J. HICKSON ON THE

the type specimen of Hrrina (L.) antarctica and comparing it
with the specimens from various localities mentioned at the
beginning of this paper. I have found, as I expected, the same
difficulties in the determination of species that are met with in
the systematic zoology of other zoophytes. My impression is
that all the specimens from New Zealand belong to one distinct
species, that the specimen from the Cape of Good Hope belongs
to another distinct species, and that both these species are distinet
from the four species that have already been described. But
there is so much variation in the specimens from New Zealand
that it is clearly desirable to have a careful description of each.
T have also added for convenience’ sake a new description of the
two earlier species for comparison.

Errina (LABIOPORA) NOVH ZELANDI#. (Facies Ramosa.) (PI.
XiODVi te noi Plex NEES fig. 9>)

This specimen was obtained from Preservation Inlet, W. coast of
South Island of New Zealand, in about 3 fms. of water, and was
lent by the Canterbury Museum to Prof. Benham. The colony
is flabelliform, with profuse ramification but without anastomoses.
The terminal branches are usually delicate. This may be ex-
pressed in figures by saying that at a distance of 3 mm. from the
extremity of a terminal branch the diameter may be not more
than 1mm. The larger branches are slightly compressed in the
plane of the flabellum ; the others circular in section.

Colour : salmon- pink*,

Surface minutely granular, substance of the ccenosteum
minutely reticulate.

Grooved spines (narial processes) numerous, arranged in rows,
rarely in clusters. The groove, in nearly all cases, turned directly
away from the apex of the branch.

Gasteropores more numerous on one side of the flabellum than
on the other, sometimes provided with a lip. Diameter of gas-
teropores 0°27 mm.

Large dactylopores 0°06 x 0:16 mm. (The large kind of dactylo-
pores of the genus are protected by the grooved spines, and the
measurements given indicate approximately the width x depth of
the groove at its deepest part.)

Small dactylopores rare or absent. The small dactylopores are
often difficult to determine until the coral is thoroughly cleaned
by boiling in eau de javelle. JI have examined and re-examined
a small. branch thus cleaned and can find no small dactylopores,
but as I have only a small amount of material at my disposal,
and as Benham states in his MS. notes that the small dactylo-
pores are “ rare,’ I cannot deny their existence.

* There are so many shades of red to be found in corals that I have used the
technical term which expresses the shade of red that comes nearest to that shown
by this coral,

HYDROCORALLINE GENUS ERRINA. 883

Errina (LABIopORA) NOV# ZELANDIZ. (Facies Benhami.)
(PI. XCIV. figs. 1 & 2;.Pl. XCVLI. fig. 13.)

This specimen was also found in Preservation Inlet and lent to
Prof. Benham by the Colonial Museum. One branch of this
specimen was well preserved in spirit. Thecolony is flabelliform,
with profuse ramification and abundant anastomoses. The
terminal branches are thick, the diameter of such a branch at a
distance of 3 mm. from the extremity being about 3mm. All
the branches are approximately circular in section.

Colour : salmon-pink.

Surface minutely granular and substance minutely reticulate.

Grooved spines numerous, arranged roughly in rows. These
spines are more numerous and longer than in the facies ‘‘ Ramosa.”
They are, moreover, frequently arranged in clusters, so that they
have the appearance of ‘branched spines” (fig. 13). The
grooves are in general turned away from the apex, but when
the projections are clustered they turn in all directions.

Gasteropores equally numerous on the two sides of the flabellum.
Without a lp. Diameter varying considerably from 0:13 mm.
to 0°17 mm.

Large dactylopores 0:06 x 0°16 mm.

Small dactylopores not infrequent, 0°05 mm. in diameter.

Errina (LABIOPORA) NOV ZELANDI®, (Facies Dendyi.)
(Pl. XCIV. fig. 4.)

This specimen was obtained in Milford Sound, W. coast of
South Island, and was lent to me by Professor Dendy.

It is not very profusely branched, but probably has a flabelli-
form mode of growth. ‘The terminal branches are delicate and
of approximately the same diameter as.those of facies “‘ Ramosa.”
There are no anastomoses in the specimen.

Colour: salmon-pink.

The surface is minutely granular and the substance minutely
reticular.

Grooved spines not very crowded but quite irregularly disposed,
not in rows, never in clusters. The groove in all cases turned
away from the apex of the branches.

Gasteropores on both sides of the branches but rather more
numerous on one side than the other. Usually guarded by a
small lip. Diameter 0°22 mm.

Large dactylopores 0:06 x 0-11 mm.

Small dactylopores 0:05 mm. in diameter, usually guarded by a
shallow collar or lip.

A feature of this specimen that should be mentioned is the
suppression of the spines on the larger branches. Only the
terminal branches are echinate.

884 PROF. §. J. HICKSON ON THE

Errina (LABIOPORA) Nova ZELANDIZ. (Facies Cooki.)
(PITACYV. figs 55 Pl PO Vals testa Oils iale2/3)

This specimen was obtained from the cable in Cook Straits
between the two islands of New Zealand, and was lent to
Professor Benham by the Colonial Museum. According to
Professor Benham’s notes, two specimens were obtained, one
being 50 mm. in height x 70 mm. across, and has six main
branches; the other is smaller, 30 mm. in height x60 mm.
across. The general form of the ccenosteum is flabellate, the
main axis flattened but the branches circular in section. The
branches do not anastomose in the specimen examined. Only a
small piece of one of these colonies was sent to me, and from
that I have drawn up the following notes.

Colour: pure white.

Surface and substance as in the other facies.

Grooved spines not very crowded and not arranged in definite
rows, frequently in clusters with the grooves pointing in all
directions (fig. 12).

Gasteropores evenly distributed on both sides of the flabellum,
without any lip or collar, 0°22 mm. in diameter.

Large dactylopores 0-06 x 0:08 mm.

Small dactylopores rare or very rare, 0°09 mm. in diameter.

In comparing these four facies of the species, several points of
interest may be observed.

They all agree in the general texture of the ccenosteum, and
they all have a more or less flabellate form of growth.

As regards the size of the hydrophytum as a whole, it is
impossible, owing to the broken condition of all the specimens
examined, to give exact measurements. A specimen of uncertain
facies in the Colonial Museum is 90 mm. x 70 mm. (according to
the MS. notes of Professor Benham), the specimen of the facies
‘“‘Cooki” was 50 mm. in height x 70 mm. in expanse. Judging
from these figures and from the size of the branches of the other
specimens, it seems probable that the normal size of a full-grown
specimen of the species is not more than 100 mm. x 100 mm., or

that, in words, it is a coral that does not normally attain a very
large size.

Of the other characters, perhaps the most important one to
consider is the dimorphism of the dactylopores, because this cha-
racter has been used as a diagnostic character for the separation
of the genera Labiopora and Hrrina. In the facies ‘‘ Benhami ”
and ‘ Dendyi” there are clearly many small dactylopores lying
on the general surface of the ccenosteum between the grooved
spines and distinct from the larger dactylopores. In the facies
‘““ Ramosa” no such dactylopores could be found in the specimen
J examined (although Professor Benham says they are rare), and
in the facies ‘‘ Cooki” they are certainly very rare. With the
many points of resemblance in form, colour, size of pores, etc.
between the two facies ‘“‘ Ramosa ” and ‘ Benham,” it would be

HYDROCORALLINE GENUS ERRINA. 885

very rash to propose that they should be separated into distinct
species on account of this one character.

It seems to me, therefore, that the presence of small dactylo-
pores in addition to the dactylopores of the ordinary type, in
other words, the character of dimorphism in the dactylopores, is
not a character that should be regarded as absolutely diagnostic
either of the genus or of any one of its species.

As regards the gasteropores there seem to be some variations.
In “ Benhami” the gasteropores vary considerably in diameter
from 0:13-0:17 mm., but in “‘ Ramosa,” ** Cooki,” and ** Dendyi”
they are more constant in diameter, being 0°27 mm. in the first
named, and 0°22 mm. in the latter. It is probable that these
figures are not of much value for systematic purposes. There
are many technical difficulties in the way of making accurate
measurements of the mouths of a large number of gasteropores
on any single specimen, and unless the average diameter of a
large number of gasteropores of one specimen can be compared
with similar averages from other specimens of the same facies or
species, the figures given simply represent a statement of fact
concerning a given specimen. The real value of the figures I
have given is that they prove that the diameter of the gastero-
pores is a variable quantity and cannot be used, except in a very
general way, as a guide to the determination of species. One
point of rather special interest is that in the facies “‘ Ramosa ”
with slender terminal branches the gasteropores appear to be
actually larger than they are in ‘ Benhami” with thicker terminal
branches. This seems to indicate that there is no relation
between the thickness of the branches and the size of the gastero-
pores, since the expectation would be that the stouter terminal
branches would bear the larger gasteropores.

The presence of a raised margin on one side of the gasteropores
in some of the specimens of this species is a feature of some
general interest. Im some of the gasteropores of the facies
‘“‘Ramosa” these processes are of considerable size, and bending
over the pore have an appearance very similar to the lid of a
Cryptohelia. In ‘** Dendyi” they are rudimentary, but in the
other specimens they are absent.

The presence of a definite lip or scale on the edge of the gastero-
pore, it must be remembered, was the principal character relied
upon by Pourtales for the separation of the genus Lepidopora
from the genus Hrrina, but if we accept Moseley’s view that the
species of Lepidopora should be incorporated with Hrrina, then
we have a parallel series of variations as regards this character in
the Hrrina group, to that in the Labiopora group.

At one time I thought that the difference in the length of the
grooved spines (nariform processes) might be a useful character
for the separation of the species in this genus. In all the speci-
mens, however, I found that the projections on the young actively
growing terminal branches are longer than they are on the older
branches, and consequently there is a difficulty in fixing a standard

Proc. Zoou. Soc.—1912, No. LIX. 59

886 PROF, S. J. HICKSON ON THE

of measurement for comparison. Taking the measurement of a
few of the projections at a distance of about 3 mm. from the apex
of the branches, I have found that the average is in ‘ Benhami”
0:7 mm., and in “ Dendyi” 0:4 mm., the projections i in “ Ramosa ”
and ‘*Cooki” being intermediate in size between these two
measurements. The differences between these averages are so
small, and the difficulty of avoiding a relatively large error in the
calculation is so great, that the measurements are of no more
scientific value than to express roughly the general impression of
observation that the projections are longest in ‘“ Benhami,”
shortest in “‘ Dendyi,” and of medium length in other specimens.
The longest projections in ‘‘ Benhami” are, however, not simple
nariform processes as they are usually in the other facies, but
groups of two, three or four of these processes clustered together
(fig. 13). The clustering together of the grooved projections
may be seen in some of the other facies, such as ‘‘ Ramosa” and
“ Cooki,’ but it is never such a pronounced feature as it is in
“* Benhami.”

Lastly, a word about geographical distribution. All the
specimens were dredged off the coast of New Zealand, and two of
these four were found in the same bay (Preservation Inlet).
There is every reason to believe, moreover, that they were all
found in shallow water. In my opinion these facts have some
weight in determining the question whether the specimens
should be placed in one species or in several species. In a rare
genus such as Hrrina, species found at widely separated localities
will in all probability be affected by their isolation and show
differences that entitle them to rank as distinct species, but
there is much less probability that the genus would be able to
develop or to maintain specific differences in the same waters.
Unless, therefore, a very clear case is made out that the dif-
ferences between the specimens of the genus Hrrina from New
Zealand waters are constant or of fundamental importance, the
most convenient as well as most scientific course to pursue is to
place them together in the same species.

Errina (LABIOPORA) CAPENSIS, sp. n. (PI. XCV. fig. 7;
Pl. XCVI. fig. 15.)

This species is represented in my collection by three broken
terminal branches. The largest piece is 36 mm. in length, the
diameter at the base is 7 mm., and the diameter of the branch
3 mm. from the apex is 45 mm. In the largest specimen of
the New Zealand species (‘‘ Dendyi”) the diameter at the base
(evidently the base of attachment) is 6 mm., and the diameter of
a branch 3 mm. from the apex is only 2 mm.

From these facts it seems probable that in this species the
hydrophytum reaches to much greater dimensions than does that
of the New Zealand species.

The branches terminate in blunt, slightly flattened and ex-
panded extremities.

HYDROCORALLINE GENUS ERRINA. 887

The hydrophytum is probably flabellate in growth, the ramifi-
cation not very profuse and anastomoses rare.

Colour: salmon-pink,

Surface coarsely granular and substance coarsely reticulate.

As this species seems to approach the Zrrina group in some
respects, attention may be called to the striking difference there
is between the coarsely granular character of its surface and the
fine smooth porcellanous character of the surface of the Hrrina
group.

Grooved spines very short, numerous, quite irregular in arrange-
ment and never in clusters. The form of the spine is that of a
shallow semicircular ridge open on the side turned away from the
apex.

Gasteropores equally distributed on both sides of the branches,
never provided with a lip. Diameter 0°3 mm.

Large dactylopores about 0°25 mm. in diameter.

Small dactylopores:—It may be open to discussion whether
there is or is not any true dimorphism of the dactylopores in this
species. It has been shown that in one of the specimens from
New Zealand (‘“‘ Dendyi”) the small dactylopores are provided
with shallow collars. In the Cape specimen some of the dactylo-
pores are considerably less in diameter than the majority, but
they are provided with exactly the same kind of semicircular
ridge as the larger ones. The question of dimorphism, therefore,
resolves itself in this case into a question whether the essential
feature of the dimorphism of the dactylopores consists in their
size or in the presence of a grooved spine. Ridley states that in
Errina (Labiopora) moseleyi the dactylopores are of the usual
uniform size, and the two kinds can only be distinguished by the
presence or absence of nariform processes. These facts seem to
emphasize the conclusion that the so-called dimorphism of the
dactylopores is not really a feature of very great importance, and
to suggest as a probability that the small dactylopores are the
shelters for young dactylozooids which in their later stages of
growth increase in size and become protected by a grooved spine.

The specimen described and figured by Gray (1872) as Hrrina
jissurata, from the Antarctic seas, was apparently very closely
related to this species. Unfortunately the specimen has been
mislaid (jide Moseley) and cannot therefore be re-examined, but
the figures show a similar robust habit of growth and short semi-
circular grooved spines with the grooves all turned away from the
apex of the branch.

Errina (LABIOPORA) ANTARCTICA Gray.

Porella antarctica Gray.

Labiopora antarctica Moseley.

Labiopora antarctica Ridley.

The type-specimen was found off the Falkland Islands,
54° 27'8., 59° 40’ W., in 45 fathoms. 1

A second specimen, attributed to this species by Ridley, was
found in Trinidad Channel, 8.W. Chili, in 30 fathoms.

Dos

888 PROF. S. J. HICKSON ON THE

Hydrophytum flabellate in growth. There is no statement to
the effect that the branches anastomose. No record of size
beyond the statement that it is smaller than L. moseleyi.

Colour: bright crimson, with the compressed forked tips paler.

Surface minutely reticulate. Grooved spines arranged in rows,
but not in clusters, on both sides of the branches, all turned away
from the apex of the branch.

Gasteropores without a lip, 0°22 mm. in diameter (in the Chili
specimen).

Large dactylopores 0:09 x 0:2 mm.

Small dactylopores 0°08 mm., without any lip or collar.

Errina (LABIoporA) MOSELEYI Ridley.

The single specimen of this species was found at Port Rosario,
S.W. Chili, 2-10 fathoms.

Hydrophyture flabellate in growth. Anastomoses frequent.
95 mm. in height x 185 mm. in width.

Colour : ‘ vermilion.”

“An anterior clearly distinguishable from a posterior surface,
by the development on it of numerous tubercles, chiefly in the
terminal branches, which are very slightly indicated in the latter.”
Surface minutely reticulate. Grooved spines not arranged in
definite rows nor in clusters.

Gasteropores without any lip, 0°32 to 0°35 mm. in diameter.

Large dactylopores 0°1 to 0:14 mm. in longitudinal diameter.

Small dactylopores without lips or tubercles, of about the same
size as the large dactylopores.

Errina (Lapropora) AspERA Linn. (Pl. XCV. fig. 6.)

I have examined the type-specimen of this species in the
British Museum on which Gray (1) founded the genus Hrrina.
There can be no doubt that the affinities of the species with the
genotype of Labiopora are closer than they are with Moseley’s
Errina labiata. It belongs to the ZLabiopora, and not to the
Errina group of species.

One of the colonies in the British Museum is 80 mm. in height
by 85 mm. in width. It forms a flabellum with a clear difference
between the anterior and posterior surfaces. The branches
terminate in fine points and do not anastomose. The branches
are about 3 mm. in diameter at a distance of 3 mm. from the
apices. The surface is minutely granular. The grooved spines
sometimes occur in clusters, but when solitary the grooves are
turned away from the apex of the branch. There are a few small
dactylopores without spines and some with small or rudimentary
spines. This specimen was dredged off the coast of Sicily.

A little while ago a specimen of a Stylasterine coral was sent
to me by the late Mr. John Morgan of Worthing. It was
purchased in a sale and there was no record of its locality.

At first I thought it should be placed with the other specimens

HYDROCGORALLINE GENUS ERRINA. 889

in my collection in the species Hrrina (L.) nove zelandie, but,
on comparing it with the specimens in the British Museum, I
came to the conclusion that it is probably related more closely to
Erriva (L.) aspera. Itis moderately branched and roughly flabelli-
form in growth. ‘The terminal branches are fairly thick, being
about 2—2°25 mm, at a distance of 3 mm. from the apex. The
branches do not anastomose.

The surface and substance of the cenosteum are minutely
granular.

The grooved spines are very crowded and clustered on the
terminal branches, and not arranged in rows (fig. 6). The
grooves are turned in all directions, but the majority of them
away from the apex of the branch.

The gasteropores are equally distributed on both sides of the
flabellum, and are without any lip or collar. Size 0-13-0:17 mm.
in diameter.

The large dactylopores are very variable in size, ‘09 x -09 mm.
to -06 x 0:11 mm.

The small dactylopores are very numerous, 0°06 mm. in
diameter, and provided with small curved lips.

The specimen is like the type in being white in colour,

Mr. Morgan’s specimen differs from the type in having rather
more slender branches, in having the pores equally distributed
on the two surfaces of the flabellum, and in the presence of
numerous small dactylopores.

Errina (LABIOPORA) GRACILIS von Marenzeller.

Several specimens of this species were found by the ‘ Belgica’
Expedition attached to the swabs when dredging off the pack ice
in the region of 71°S. and 88° W., z.e. about 20 degrees west
and 15 degrees south of the Straits of Magellan. The depth
is not recorded by von Marenzeller, but it is probably between
500 and 600 metres.

The hydrophytum is flabellate in growth, with well-marked
anterior and posterior surfaces. One of the specimens, which
proved to bea female, was 25 mm. in height and 30 mm. in width.
Another, which proved to be a male, was 100 mm. by 140 mm.

The surface of the ccenosteum is finely wrinkled, and marked
with transverse and longitudinal ridges (Kammschen). There are
apparently no well- marked comnosteal pores.

Grooved spines not arranged in definite rows, but in irregular
clusters or singly. The grooves of the grooved spines turned
away from the apex of the branches or sideways.

Gasteropores with a lip, :015 mm. in width.

Large dactylopores sheltered by spinous projections provided
with a deep groove.

Small dactylopores with or without a lip.

Colour: white or brownish.

There can be little doubt that this species belongs to the

890 PROF. §. J. HICKSON ON THE

Labiopora group. It is true that the description given by von
Marenzeller of the surface of the ccoenosteum does not agree with
that of the other species of the group, but it does not agree either
with the description given of the surface of the coonosteum of the
Hrrina group. The absence of well-defined ccenosteal pores, the
grouping of some of the grooved spines in clusters, and the direc-
tion of their grooves—all point to the affinities of the species with
the Labiopora group.

It may be remarked that this is the only species of the group
that oecurs in deep water.

Review of the Labiopora Group of Species.

The careful examination of the specimens belonging to the
Labiopora group leads me to the conclusion that there are very
few characters that can be used with much confidence for the
separation of species *. However, it may be convenient for the
present to recognise six species :—

Hrrina (Labiopora) aspera Linn. Mediterranean Sea.

Lrrina (Labiopora) antarctica Gray. Chili and Falkland Islands.
30 to 45 fathoms.

Lrrina (Labiopora) moseleyi Ridley. Chili. Shallow water.
Lrrina (Labiopora) nove zelandie Wickson. New Zealand.
Shallow water.
Lrrina (Labiopora) capensis Hickson. Cape of Good Hope.
30 fathoms.
Lirrina (Labiopora) gracilis von Marenzeller. Antarctic Sea.
Deep water.

Of these six species Hrrina (Labiopora) capensis appears to be
the most sharply defined. It probably attains to a much larger
size, has more robust branches terminating in blunt and some-
what flattened extremities. Its substance is coarsely reticular
and its surface coarsely granuiar. The grooved spines are short
and semicircular in shape. All the dactylopores are guarded by
these spines.

The other five species are very closely related. Hyrina (L.) ant-
arctica appears to be distinguished from the others by the grooved
spines being arranged in definite rows, and Hrrina (L.) moseleyi
by the differentiation of an anterior from a posterior surface
of the flabellum. Of Hrrina (L.) nove zelandie all that ean
be said is that 1t appears to be a very variable species which
does not exhibit any one particular distinguishing feature.
Errina (L.) gracilis is distinguished from the others by the
texture of the surface of the ceenosteum, and the colour is not
red but white or brownish. Hrrina(L.) aspera has a close resem-
blance to some of the facies of Hrrina (L.) nove zealandie, but
it is always white in colour, in this respect resembling the facies
* Cooki.”

* See Note, p. 894.

HYDROCORALLINE GENUS ERRINA. 891

General Remarks on Structure.

Gasteropores.—Before concluding this account of these species
I may refer briefly to one or two results of a negative character
that I have obtained. It occurred to me that if the speci-
mens J examined belonged to different species, there might be
some measurable difference to be observed in the styles of the
gasteropores. The style is in all cases lke an elongated and
sharply pointed brush, but careful and constantly repeated com-
parisons of the styles of the gasteropores of different specimens
revealed no characters by which they could be distinguished.
The type of style is the same in all the specimens, and although
the brush is more slender in some cases than in others, there is a
greater range of variation in this respect in the gasteropores of
a single specimen than there is in the gasteropores of different
specimens taken at random (Pl. XCV1. figs. 14 & 15).

The way in which it is possible to study the shape of the
gasteropore styles is to make a vertical fracture in a plane parallel
with the long axis of a branch. Ina large percentage of such
fractures the whole length of at least one gasteropore with its
style will be exposed.

In such fractures it may be observed that some of the gastero-
pores extend as far down as the axis of the branch, but in others
they extend only a short distance below the surface. In the long
gasteropores the style can be traced right down to the base of the
pore, but it is usually supported by one or, in some rare cases,
more than one, tabula (fig. 14) similar to the tabulee described by
Miss England (5) in the gasteropores of Spinipora. In the short
gasteropores the style ends abruptly at the base of the pore and
is not supported by a tabula (fig. 15).

In order to make any scientific use of this character in the
classification of the species 1t would be necessary to make a large
number of fractures, and for that purpose I have not sufficient
material.

The results I have obtained on the few fractures I have made
are as follows :—

In Errina (L.) nove zelandie facies ‘“‘ Ramosa” and facies
‘“Cooki” the gasteropores (I have observed) are long and exhibit
atabula. In the same species facies “ Benhami” the gasteropor es
are short, and the same is the case in the facies “ Dendyi.” In
Mr. Morgan’s specimen of Hrrina (L.) aspera the gasteropore is
also short. In Krrina (L.) capensis the gasteropores are short,
extending only a little way beneath the surface. As to this
character in the gasteropores of the other species I have no
evidence to bring forward.

My conclusions, from this evidence, are that the length of the
gasteropores and the presence or absence of a tabula are not, at
present, characters of any value for purposes of classification.

Gonophores.—All the specimens | have examined show ampulle.
They can usually be seen on the terminal branches as shallow

892 PROF. 8. J. HICKSON ON THE

convexities on the surface, sometimes between the spines, some-
times carrying the spines with them. They appear to be always
absent in the older branches and stems. Our knowledge is still
so very imperfect concerning the growth and relations of the
gonophores in the Stylasterina that it is not reasonable to use any
differences that may be observed in the characters of the ampulle
for purposes of classification.
The questions to be answered are the following :—

1. Are there any constant differences between the male and the
female ampullee ?
Is there any correlation between the sex of the colonies and
characters of the general structure of the coenosteum ¢

These two questions could be answered without much difficulty
by the examination of a number of preserved specimens of the
same species from the same locality.

In Professor Benham’s notes on the structure of a specimen
which probably corresponds with rrina (Labiopora) nove
zelandic facies “‘ Ramosa,” I find the following remark :—“ Each
ampulla seems to have a small pore (?dactylopore) on its surface.”
In the specimen I have examined of this species I cannot find
these small pores on the ampulle. Von Marenzeller also remarks
that the male ampulle exhibit small dactylopores “in niedrigen
Spitzchen.”

In a previous paper (3) I have shown that, in the ripe male
gonophores of Distichopora and Allopora, the spermatozoa are
discharged by a spout-like seminal duct. In the case of the female
gonophore, however, the only way of escape of the embryo is by the
rupture of the whole surface of the ampulla. It seems possible,
therefore, that the pore on the ampulla mentioned by Professor
Benham may be for the opening of the seminal duct and that the
specimen may be a male. On the other hand, it must not be
assumed that when the surface of a specimen is marked by large
shallow depressions having the size and appearance of ruptured
ampull, the specimen is necessarily a female colony, because in
the specimen of Hrrina (Labiopora) capensis in which these
shallow depressions are very well-marked I found to my surprise
that it was a male colony.

As regards the structure of the gonophores there is very little
to be said at present. The male gonophores of Hrrina (Labiopora)
capensis exhibit a well-developed spadix (manubrium), and in this
respect the genus seems to resemble Distichopora and to differ
from Allopora a. The only other spirit specimen I have received
was a small branch of Hrrina (Labiopora) nove zelandic facies
‘* Benhami,” and this proved to be a female. The specimen was
not, however, in sufticiently good condition to enable me to study
the structure of the gonophore.

Cenenchymal Canals.—From the two spirit specimens I have
ascertained certain isolated facts which may be of some use when
our knowledge of the genus is extended.

HYDROCORALLINE GENUS ERRINA. 893

According to Ridley the conenchymal canals of Z. moseleyi and
L. antarctica ave 0:035 to 0°07 mm. in diameter, the meshes
between the canals are about the same diameter in ZL. moseleyi,
but in LZ, antarctica they vary from the same diameter up to
0-14 mm. Ridley does not state how his measurements were
obtained, consequently the results I have obtained cannot be
compared with his results with any degree of precision.

I have measured the diameter of a number of canals that have
been cut transversely in my sections at a level corresponding
with the base of the dactylozooids, and I find that the average
diameter of these canals is in Hrrina (Labiopora) nove zelandie
0-03 mm. and in Hrrina (L.) capensis 0'05 mm, As the specimens
must have undergone very considerable contraction during de-
calcification and imbedding, I have regarded the measurements of
the meshes as worthless. All that can be said is that in both
species the meshwork of canals is close and elaborate.

In the ‘ Challenger’ monograph Moseley gives as a character
of the genus Hrrina that the number of tentacles on the gastero-
zooids is four. In the Siboga species of Hrrina the number of
tentacles varied from four to five.

In Hrrina (Labiopora) nove zelandiw 1 have found some diffi-
culty in counting the number of tentacles, but judging from a
single series of sections, there are not more than four. In the
specimen of Hrrina (Labiopora) capensis, however, a very large
number of gasterozooids can be clearly seen without decalcification,
and in these the number varies from four to six. From this fact
it is obvious that the number of tentacles on the gasterozooids
cannot be relied upon as a generic or even as a specific character.

List of the Species of the Genus Krrina.

A. Coenosteum hard and compact, perforated by well-defined
ceenosteal pores. Grooved spines turned towards the apex
of the branch.

1. With short grooved spines and only one kind of dactylopore.
The #rring group.
Lrrina labiata Moseley.
Errina ramosa Hickson & England.
Errina horrida Hickson & England.
Errina carinata Pourtales ¢
Errina pourtalesii Dall ¢
*Hrriva glabra (= Lepidopora glabra Pourtales).
*Hrrina cochleata (= Lepidopora cochleata Pourtales).
*Hrrina dabneyi (= Lepidopora dabneyi Pourtales).

2. With long grooved spines and two kinds of dactylopores.

Errina echinata (= Spinipora echinata Moseley).

* See Note, p. 894.

894 PROF. 8. J. HICKSON ON THE

B. Ceenosteum granular and reticulate, without well-defined
coenosteal pores. Grooved spines turned away from the
apex of the branches or clustered and irregularly placed.

The Labiopora group.
Errina aspera Linn.
Errina fissurata Gray.
Errina antarctica (= Labiopora antarctica Gray).
Errina moseleyi (= Labiepora moseleyi Ridley).
Errina nove zelandice Hickson.
Errina capensis Hickson.
Errina gracilis von Marenzeller.

The following species has the grooves turned away from the
apex as in the Labiopora group, but has well defined ccenosteal
pores :—-

Errina macrogastra von Marenzeller (7).

| Additional notes on Professor Hickson’s paper on the
genus Hrrina, received August 14th, 1912.

1. The adjective ‘‘ Hydrocoralline” is used in the title of the
paper at the suggestion of the Secretary of the Society for the
convenience of Zoologists who do not possess a special knowledge
of these Coelenterata. I have in several previous papers declared
my opinion that to unite Millepora with the Stylasterina in one
Order, the Hydrocorallina, is unsound. J have found no reason
to change my mind in that respect. The word “ hydrocoralline ”
may still be used, however, to signify the corals that are Hydrozoa
in very much the same way as the words “corals,” ‘“ zoophytes,”
“worms,” are used, without any strict systematic significance.

2. Since the paper was read I have been able to examine a
number of specimens in the possession of the Muséum d’ Histoire
Naturelle, Paris, for which I am indebted to the kindness of
Prof. Joubin.

In this collection there are some specimens which I have
identified as Hrrina dabneyi Pourt. from the Azores. An
examination of these specimens convinced me that they belong to
the Labiopora group of species, as they exhibit all the charac-
teristic features of that group. From this it seems probable that
the other two species, W. glabra and L. cochleata, that were in-
cluded by Pourtales in the genus Lepidopora should also be
transferred from the Hrrina to the Labiopora group. |

Literature.

(1) J. E. Gray.—(No title.) Proc. Zool. Soc. 1835, p. 85.
(2) J. EH. Gray.—Notes on Corals from the South and Antarctic
Seas. Proc. Zool. Soc. 1872, p. 744.
(3) S. J. Htcxson.—The Gonophores of Allopora and Disticho-
pora. Quart. Journ. Microscop. Sci. vol. xxxu. 1891,

p- 382.

HYDROCURALLINE GENUS ERRINA, 895

(4) 8. J. Hickson & Hrten M. Enciuanp.—The Stylasterina of
the Indian Ocean. ‘Trans. Linn. Soc. xu., 1909.
(5) S. J. Hickson & Henen M. Eneranp.—The Stylasterina of
the Siboga Expedition, 1905.
(6) E. von Marenzevuer.—Madreporaria and Hydrocorallia.
Résultats du Voyage du §.Y. ‘ Belgica.’ 1903.
(7) E. von MarenzeLLer.—Stein- und Hydro-Korallen. Bull.
Mus. Comp. Zool. xliii. 2. 1904.
(8) H. N. Mosetey.—Zoological Collections of H.M.S. * Chal-
lenger,’ vol. 11. 1881, p. 50 seq.
(9) L. F. pp PourtaLes.—Deep Sea Corals. Illustrated Cata-
logue Museum Comp. Zoology, Harvard, 1871.
(10) S. O. Ripteyv.—Zoological Collections of H.M.S. ‘ Alert.’
Proc. Zool. Soc. 1881, p. 105.

EXPLANATION OF THE PLATES.
Prats XCIV.

Fig. 1. Hrrina (Labiopora) nove zelandie (facies Benhami) ; a portion of a branch
showing the grooved spines arranged in clusters or with the grooves
turned away from the apex. X 45 diam.

Fig. 2. Another photograph of the same facies (Benhami) showing a number of
closely packed anastomosing branches. In this view it will be seen that
although the greater number of the grooves are turned away from the
apex of the branches (the top of the photograph), in places where they are
clustered they are turned in all directions. X 4°75 diam.

Fig. 3. The same species (facies Ramosa). ‘The grooved spines are not so crowded
and rarely form clusters. Nearly all the grooves are turned away from
the apex. > 9'3 diam.

Fig. 4. The same species (facies Dendyi). The grooved spines are still more scattered
and nearly all the grooves are turned away from the apex. in this
photograph several gasteropores can be seen. XX 9°3 diam,

PratE XCV.

Bie. 5. Hrrina (Labiopora) nove zelandie (facies Cooki.) In this facies many of
the spines are in clusters, consequently the grooves appear to be turned
in all directions. > 45 diam.

Fig. 6. Lrrina (Labiopora) aspera. In this specimen the spines are rather
water-worn, but frequently occur in clusters and the grooves point in all
directions. XX 45 diam.

Kig. 7. Hrrina (Labiopora) capensis. The spines are very short and the grooves
all pomt away from the apex. In this photograph the style can be seen
in many of the gasteropores. X 45 diam.

Fig. 8. Hrrina (Spinipora) echinata. In this species, as in the Hrrina group of
species, the grooves are turned towards the apex. X 45 diam.

PratEe XCVI.

Fig. 9. Errina (Labiopora) nove zelandie (facies Ramosa). Slightly enlarged,
showing the profuse ramification without anastomoses, and branches
terminating in delicate points. Drawn by Prot. Benham.

Fig. 10. Hrrina (Labiopora) nove zelandie (facies Cooki). Slightly enlarged,
showing the method of ramification and branches terminating in blunt
points. Drawn by Prof. Benham.

Fig. 11. A terminal branch of the same species (facies Cooki), showing the dactylo-
pores (D) protected by the grooved spines and the gasteropores (G).
<6diam. Drawn by Prof. Benham.

Fig. 12. A portion of a larger branch of the same species (facies Cooki), showing the
grooved spines in clusters. X ca. 6diam. Drawn by Prot. Benham.

896 MR. E. DUKINFIELD JONES ON

Fig. 13. Errina (Labiopora) nove zelandie (facies Benhami). One of the clusters
of four grooved spines as seen from above. X ca. 12 diam.

Figs. 14 & 15. These two drawings are intended to show the difference in texture
between Hrrina (L.) nove zelandie (fig. 14) and Hrrina (L.) capensis
(fig. 15), as seen in vertical section of the ccenosteum. Another point
of difference between the two specimens is that in the former (fig. 14)
the gasteropore penetrates into the depths of the branch and shows a
tabula (¢) in its course; whereas in the latter (fig. 15) the gastero-
pore ends abruptly a little way below the surface. This difference,
however, cannot be regarded as a specific distinction. G, gasteropore ;
D, dactylopore; St., style. Both figures X 35 diam.

54. Descriptions of new Butterflies of the Genus Thecla
from 8.E. Brazil. By E. Duxinrimip Jonss, F.Z.8.,
F.E.S.

[ Received May 18, 1912: Read October 29, 1912. |

(Plate XCVII.*)

INDEX.
Systematic : Pages
Thecla, 13 new species Of ....0.......0eccccseveeeeee 896-902

The species described in the present paper were captured by
myself and the types are in my own collection.

THECLA HAMILA, sp.n. (Pl. XCVII. fig. 1.)

Allied to 7’. gispa Hew.

Male. Upper side rich purplish blue; costa and termen of fore
wings narrowly black, wider at apex: base of costa greenish ;
a lar ge black silky spot with blue reflection beyond cell : cilia
incl Hind wings: costa and termen narrowly black; inner
margin greyish black, rather broad at tornus; tails black with
white tips; a few white scales on inner side of lobe; some
greyish green at tornus above the black margin; submedian hairs
greenish blue; head black; thorax greenish blue; abdomen
purplish blue. Under side dull stone-green suffused with black,
a pale green reflection over all. Fore wings: a medial row of
diffused light stone-green spots from costa to vein 2, the spot
above vein 3 distal; an indistinct subterminal diffused pale green
band; termen black; cilia black with pale green band at apex.
Hind wings darker; a broad diffused black medial shade, very
dark below costa; a narrow dark postmedial shade and broad
subterminal shade; a medial series of green spots between the
veins from inner margin to vein 5, those above veins 2 and 3
distal; termen and cilia black.

Expanse 34 mm.

Hab. Castro, Parana, Brazil.

* For explanation of the Plate see p. 902.

IDS, Se Ie OCG WANE,

Horace Knight del.et lith. West,Newman chr.
NEW SPECIES OF THECLA FROM 5. E. BRAZIL.

NEW BRAZILIAN BUTTERFLIES. 897

THECLA FANCIA, sp.n. (Pl. XOVII. fig. 2.)

Allied to 7’. punctwm H.-S.

Male. Upper side bright metallic blue with a greenish reflection
on basal half of wings; costa of fore wings narrowly, apex and
termen broadly black; a brown circular brand on upper angle of
cell; hind wings outwardly broadly black; tails tipped with
white. Under side yellowish grey: fore wings, a postmedial
band of indistinct brown lunular spots outwardly bordered with
white from costa to vein 2; hind wings: postmedial band inwardly
dark brown, outwardly white, angled on veins | and 2, inwardly
oblique from 3 to 4, placed more distally from 4 to 6 and above 7 ;
a small orange spot at tornus and a larger one between veins 2
and 3.

Expanse 30 mm.

Hab. Castro, Parana, Brazil.

THECLA SICRANA, sp. n. (Pl. XCVII. fig. 3.)

Male. Upper side dull metallic blue; costa and termen of fore
wings narrowly black ; submedian hairs on hind wings greyish ;
tornus dull orange; tails and cilia black tipped with white.
Under side bright green; inner margin of fore wings bluish
grey; hind wings: an indistinct postmedial series of lunular spots
inwardly black, outwardly white; dark red-brown subterminal
spots above veins 1, 2, and 3, the one above 2 larger; tornus dark
red-brown; cilia black tipped with white. Palpi white with
some lateral tawny scales, 3rd joint black; frons brown, with a
few metallic green scales; a white point between antennex;
abdomen bluish grey above, white beneath,

Expanse 30 mm.

Hab. Castro, Parana, Brazil.

THECLA BERTHA, sp. n. (Pl. XCVII. fig. 4.)

Female. Upper side smoky brown, inwardly suffused with
bluish grey ; fore wings irrorated from median nervure to inner
margin with light blue ; cilia smoky brown interrupted by white
between the veins; in the hind wings the irroration covers the
cell also; tornus tawny, with a dark spot on lobe; tails black
with white tip; cilia outwardly whitish. Under side bright
green; basal half of submedian area of fore wings brownish grey,
outer half light grey; a rather broad clear white postmedial
band with indistinct dark central line from costa to vein 2; cilia
reddish brown, tipped with white between the veins. Hind
wings: a nearly straight white medial line defined by maroon on
both sides from vein 8 to vein 1, angled inwards at origin of
veins 3 and 4, broken at vein 2, being placed more distally below
the vein, angled inwardly on submedian fold ; a large white spot
slightly irrorated with maroon above upper angle of cell; three
spots in cell; one spot above vein 2 and one above vein 1;
a terminal series of lunular maroon spots irrorated with white,
fusing at the veins, the one above vein 2 being produced inwardly,

898 MR. E. DUKINFIELD JONES ON

forming a large maroon spot without irroration; cilia reddish
brown, tipped with white between the veins.

Expanse 30 mm.

Hab. Castro, Parana, Brazil.

THECLA SCHAUSA, sp.n. (PI. XOVIT. fig. 5.)

Male. Upper side: fore wings black; cell and submedian area
to near tornus metallic purplish blue; extreme edge of costa
tawny; a small dark grey brand beyond cell; cilia grey; hind
wings metallic purplish blue; costa and apex black; termen
narrowly black; tornus tawny red; cilia grey. Under side:
fore wings brown; a purplish reflection on medial area below
costa; a postmedial wavy dark red line from costa to vein 2;
a terminal band of dark red irrorations wide at apex and tapering
down to vein 2; termen tawny red: bind wings purplish brown ;
an inner and a medial line of red irrorations; outer half of wing
somewhat greyish, very thickly irrorated with red; a diffused red
subterminal band; termen tawny red. Frons tawny red; palpi
white mixed with brown; pectus white.

Female. Very similar to the male; the blue of the wings is
duller; cilia almost white ; termen of hind wings more broadly
black.

Expanse, male 17 mm., female 17 mm.

Hab. Castro, Parana; Sao Paulo, S.E. Brazil.

THECLA JAPOLA, sp.n. (Pl. XCVII. fig. 6.)

Allied to 7. gaina Hew.

Female. Upper side dull steel-blue ; costa, apex, and terminal
area of fore wings broadly greyish black; a dark diffused spot
beyond cell; basal half of extreme costa tawny ; cilia outwardly
brown. Hind wings: costal and terminal areas greyish black,
lighter than fore wings; an indistinct subterminal row of dark
lunular spots, conspicuous above veins | and 2; a red spot on
lobe ; tails black, tipped with white; termen black, preceded by
whitish line; cilia black at base, white at centre, brown at tips.
Under side brownish grey; fore wings: an inwardly diffused
reddish-brown postmedial band, outwardly white, straight from
costa to vein 2; terminal area pale grey with two diffused brown
bands, the inner broad, the outer narrow ; termen dark brown ;
cilia light brown ; hind wings: an inwardly reddish-brown, out-
wardly white medial band straight from costa to vein 2, angled
on veins 1 and 2 forming a very acute W; terminal area whitish
grey with an inner brown band, broad and diffused at costa,
narrow and distinct at tornus, bent strongly towards base on
inner margin, followed at tornus by orange-red to a black spot on
lobe, some white at base of lobe ; an orange-red lunular spot with
black point above vein 2; termen dark brown; cilia lighter with
pale band.

Kixpanse 28 mm.

Hab. Castro, Parana, Brazil.

NEW BRAZILIAN BUTTERFLIES. 899

THECLA FERNANDA, sp.n. (PI. XOVII. fig. 7.)

Allied to 7. nubitlum H. H. Druce.

Male. Upper side dark blackish brown ; a red spot on lobe and
a few white scales at tornus; tails black tipped with white.
Under side yellowish grey-brown ; fore wings: a dark postmedial
band outwardly bordered with white, nearly straight from costa
to vein 2; hind wings: postmedial band inwardly shaded with
red and outwardly with white, angled on veins | and 2, proximate
between veins 3 and 4, straight from 4 to 7, distal above 7
forming conspicuous spot at costa. Differs from 7. mubilwm in
the band on fore wing being straighter, that on the hind wing
less acutely angled on veins | and 2 and in the distal spot at costa,
also in absence of discocellular bar.

Expanse 29 mm.

Hab. Fernandes Pinheiro, Parana, Brazil.

THECLA NoRA, sp.n. (Pl. XCVII. fig. 8.)

Male. Upper side dark metallic lilacine blue; costa and termen
of fore wings narrowly black, broader at apex; a large black
brand beyond celi; cilia black at base, whitish metallic blue at
tips; hind wings: costal area greyish brown; a fine black
terminal line; cilia black with bluish-white band. Under side
brownish grey : fore wings suffused with steel-blue on submedian
area; postmedial band inwardly dark brown, outwardly white
from costa to vein 3, below vein 3 the brown changes to blue ;
a very indistinct subterminal line, inwardly white, outwardly
brown; cilia brown; hind wings: postmedial band dark brown,
shaded inwardly with orange and outwardly with white, strongly
angled on veins 1 and 2, outwardly displaced between 4 and 5;
a series of subterminal lunular spots shaded inwardly with white,
followed by orange spot at tornus, a dark spot irrorated with
white below vein 2 and an orange spot containing black point
below 3; a narrow white terminal line; cilia dark brown with
bluish-white tips.

Expanse 24 mm.

Hab. Castro, Parana, Brazil.

THECLA MOLENA, sp.n. (PI. XCVII. fig. 9.)

Closely allied to 7’. lemona Hew.

Male. Upper side dull dark metallic blue; fore wings: outer
half of costa, apex and termen black; a large oval black brand,
crossed by blue discocellular bar; extreme costa tawny at base;
hind wings: termen narrowly black; tails black tipped with
white; a red spot on lobe. Under side brown with a lilacine
reflection on terminal area of fore wings and the whole of the
hind wings. The bands and lines are much the same as in
7. lemona, but they are much more diffused and subdued; this
difference and the lilacine reflection readily distinguish the
species. 7’. lemona is also a larger insect than the present.

900 MR. FE. DUKINFIELD JONES ON

Female. Upper side brown, suffused with lilacine on basal area.
Under side as in male, but the lilacine reflection is much less
pronounced,

Expanse, male 24 mm., female 24 mm,

Hab, Castro, Parana, Brazil.

THECLA ESMERALDA, sp. n. (PI. XCVII. figs. 10, 11.)

Close to 7. janias Cr., but differs in the very much wider
black margins of the upper side and in the postmedial line, and
in the orange spot in place of the black spot on the under side in
janias.

Male. Palpi and head vivid metallic green; legs green; tarsi
inwardly whitish and banded with black; thorax, abdomen, ‘and
wings vich purplish blue. Fore wings broadly bordered with
black on costa and termen, very broad at apex; a small circular
dark grey brand. Hind wings broadly black on costal and apical
areas, narrowly from vein 4 to tornus; a green spot on the black
at tornus; tails black tipped with white. Under side: fore
wings vivid metallic green, inner margin grey; cilia brown:
hind wings vivid metallic green; a fine black wavy postmedial
line, most distinct at inner margin; some lunular subterminal
spots near tornus, that above vein 2 being bright orange; some
white scales at tornus. )

Female. Thorax green; wings and abdomen brown, with slight
lilacine grey tint on hind wings and basal half of fore wings.
Under side as in male, but the postmedial line clearer and the
orange spot above vein 2 much larger, there 1s also some orange
at tornus; cilia inwardly black, outwardly whitish.

Expanse, male 27 mm., female 25 mm.

Hab. Castro, Parana, Brazil.

THECLA CASTRENA, sp.n. (Pl. XCVIT. figs. 12, 13.)

Allied to 7. phrosine H. H. Druce.

Male. Upper side dark blackish brown ; a few lilacine scales
at base and along inner margin of fore wings; a slight lilacine
reflection on hind wings; termen dark and cilia white in both
wings. Under side: fore wings pale lilacine blue; apex and
inner margin whitish; a lilacine reflection over the whole ;
a medial V-shaped mark on submedian fold; a slightly wavy
red-brown postmedial band from costa to vein 2, followed by
white spot above vein 6; a fine black terminal line, becoming
brown at apex ; cilia white at tornus, shading to brownish at apex.
Hind wings white with lilacine reflection ; a bright orange-tawny
irregular postmedial band of elongated spots, placed distally helow
veins 1, 3, 5, 6 and 8, and proximally below 2, 4 and 7; a sub-
terminal row of minute tawny and black spots between the
veins; a fine black terminal iine, shading to brown at tornus ;
cilia white.

Female. Upper side the same as the male, except that the cilia

NEW BRAZILIAN BUTTERFLIES. 901

are brown. Under side yellowish grey, the hind wings lighter
than fore wings; bands as in male, but the postmedial is out-
wardly shaded with white and there is a subterminal series of red
spots from vein 2 to 6; termen brown ; cilia brown: on the hind
wing the subterminal spots are much larger than in the male,
termen brown, inwardly shaded with white, cilia greyish
white.

Expanse, male 25 mm., female 24 mm.

Hab. Castro, Parana, Brazil.

THECLA CAMPA, sp.n. (PI. XCVII. fig. 14.)

Near 7’. biblia Hew.

Male. Upper side dull blackish blue, shading to black on outer
half of fore wings; cilia brown ; hind wings; termen narrowly
black ; fringe of abdominal fold white; cilia black, with white
band at tornus, outwardly brown at apex. Under side light
green with a light lilacine-blue reflection; cilia of fore wings
light brown ; hind wings: an obscure greyish-white discocellular
bar; an obscure sinuous greyish-white postmedial line, more
clearly defined from inner margin to vein 3; a black spot at
tornus with some white scales above and below it; a minute
black point below vein 2 and a small orange-red spot ringed with
black below vem 3; termen near tornus bluish white. Palpi
white with green reflection, third joint black above, white
beneath ; frons metallic green; back of head and tegule dark
brown ; eyes surrounded with white; body blue-black above,
creamy white beneath.

Expanse 22 mm.

Hab. Castro, Parana, Brazil.

THECLA DATITIA, sp.n. (Pl. XCVII. fig. 15.)

Very near 7. tadita Hewitson, but differing from it in the
possession of tails, the narrow black margins on upper side of
the hind wings, and in the terminal rows of lunular.spots and the
black spots on the under side of the hind wings.

Male. Upper side: fore wings smoky black; a very dark
medial suffusion from costa to vein 2; outer two-thirds of cell
filled with raised brown scales; a submedian patch of lilacine
blue from base to near tornus; termen dark; cilia greyish ;
hind wings lilacine blue, broadly black on costa and apex;
a series of diffused black subterminal spots, confluent below apex ;
termen dark ; cilia greyish ; tails dark with white tips. Under
side grey with light brown bands. Fore wings: a light brown
discocellular bar; a well-defined light brown postmedial band,
diffused on inner side; a narrower and wavy subterminal band
followed by a row of diffused spots; termen light brown; cilia
grey at base, brown at tips. Hind wings: base light brown
irrorated with grey; a light brown discocellular bar; a dark
narrow postmedial band, diffused on inner side and followed by

Proc. Zoou. Soc.—1912, No, LX. 60

902 MR. R. LYDEKKER ON

narrow grey line and broad light brown suffusion ; a medial spot
ringed with grey at costa; a terminal series of diffused lunular
spots, containing a black spot above vein 2.

Expanse 23 mm.

Hab. Fernandes Pinheiro, Parana, Brazil.

EXPLANATION OF PLATE XCVII.

Big. 1. Thecla hamila 3. Fig. 9. Thecla molena @.
2. » jfancia 6. 10. » esmeralda 3.
3. » sierana g. 11. iF 6 :
A. » bertha 2. 12. >» castrena 6.
5. » schausa 6. 13. a Bs 5
6. » japola 9. 14. » canpa o.
ie » fernanda ¢. 15. > © datitia So:
8. » nora 6.

55. The Bornean Bantin.

By R. Lypexxer, F.R.S., F.Z.S.*
[ Received October 1, 1912: Read October 29, 1912. ]
(Text-figures 123-125.)

That the Bantin of Borneo should be subspecifically distinct
from the typical Bos sondaicus of Java is what might naturally
be expected, and the only wonder is that its distinction has
not long since been recognized and a name assigned to the
local race. This, however, is not improbably due to the fact
that, so far at least as this country is concerned, the Javan
Bantin is an exceedingly rare animal in museums, the only com-
plete adult example in the British Museum (Nat. Hist.) being
the mounted skin of a bull obtained by exchange with the Leiden
Museum in 1846, and now, as might have been expected after
long and unprotected exposure in the galleries at Bloomsbury, in
an exceedingly dilapidated condition. The Museum also possesses
a fine mounted head of a bull, presented in 1904 by Baron L. T.
H. van Heckeren-tot-Walien; and also the skeletons of a bull
and cow, with the horns, obtained from Leiden at the same time
as the mounted skin, the skull of the former being exhibited in.
the Pavilion at the further end of the Lower Mammal Gallery.
In addition to these is the mounted skin of an immature male,
and also a young skull, with horns—from the Lidth de Jeude
collection—both of which present all the characteristics of the
Javan race.

On the other hand, the Museum has a considerable series
of skulls and horns of the Bornean Bantin presented by Mr.
H. B. Low in 1880 and 1887, as well as two presented by
Mr. W. B. Pryer in 1886, all these being from North Borneo.
Sir Edmund Loder also possesses at least two frontlets and

* By permission of the Trustees of the British Museum.

THE BORNEAN BANTIN. 903

horns of subadult Bornean Bantins in his private museum at
Leonardslee.

As regards the colour of the Bornean Bantin I know nothing
definite ; but, in the absence of any statement to the contrary,
I presume it is approximately similar to that of the typical
Javan race; that is to say, adult bulls are blackish brown or
black with a large white rump-patch and white ‘“ stockings,” while
the cows and young bulls have the dark area rufous or chestnut.

In all the above-mentioned Javan specimens in the Museum
the horns are of the type of those shown in text-fig. 123, which

Text-fig. 123.

Skull and horns of male Javan Bantin (Bos sondaicus) in the
British Museum.

represents the skull of a bull obtained from Leiden in 1846. At
their origin the horns are directed mainly outwards, but at rather
more than half their length they curve suddenly inwards with
a somewhat forward and finally a backward inclination. This

60*

904 MR, R. LYDEKKER ON

causes them to have a relatively large maximum span and a cor-
respondingly small tip-to-tip interval, as is indicated in the table
of measurements given below. In consequence, I take it, of the
outward and somewhat backward direction of the basal portion
of the horns, the frontal region of the skull is somewhat convex,
while the intercornual ridge has a distinct prominence in the
middle line.
Text-fig. 124.

Skull and horns of male Bornean Bantin (B. sondaicus lowi), from the
Rejang Valley, in the British Museum.

In the Bornean Bantin, as typified by the largest of the skulls
presented by Mr. Low to the British Museum (text-fig. 124), the
horns, which are relatively stouter, are less curved and have a

THE BORNEAN BANTIN. 905

more upright direction, arising at an angle of about 45° from the
ends of the frontal ridge of the skull, and finally curving inwards
to a less extent than those of the typical race, the terminal
portion thus being shorter and the maximum span less. Similar
characters are shown in the two frontlets with horns in the

. Text-fig. 125.

B

A & B.—Frontlets and horns of two Bornean Rantins (B. sondaicus lowit)
in the collection of Sir E. G. Loder.

collection of Sir E. G. Loder, represented in text-fig. 125, these
also exhibiting the relatively smaii size of the horns which
appears to characterise many Bornean Bantin heads. The skull
of the Bornean Bantin is characterised by the flatness of the
forehead and the straight intercornual ridge.

906 MR. E. G. BOULENGER ON THE

The dimensions, in inches, of the specimens represented in
text-figs. 123 & 124 are as follows :—

B. sondaicus. B. s. lowi.

Length of horn on outer curve......... 25 214
Basalvorrthvot ditto, asst s.ssee tees 11} 123
Maximum span of horns ............... 28 2335
Tip-to-tip interval of ditto ............ 16 13
Length from intercornual ridge of

skull to tip of nasals .......... een 16 133
Minimum width of skull between base

of hownecorelamdioroitn sas eeeet ee a i

These differences fully justify the right of the Bornean Bantin
to racial distinction, and as none of the comparatively few
synonyms of Bos sondaicus appears referable to that race, I
propose that it should be known as B. sondaicus lowi; the skull
and horns represented in text-fig. 124 (B. M. No. 87.2.10.4)
being the type.

It may be added that both the Burmese Bantin or Tsaine, for
which I proposed the name of B. s. birmanicus in the Society’s
‘Proceedings’ for 1898, p. 277, and the Siamese Bantin, for
which I have suggested the name of B.s. porteri, Ibid. 1909,
p. 669, have horns of the general type of those of the typical
race, although those of the Siamese race are more heavily wrinkled
at the base than any Javan horns in the British Museum.

Information is still required as to whether the Bantin occurs
in Sumatra.

56. Notes on the Breeding of the “ Millions” Fish (Grar-
dinus peciloides). By Epwarp G. BouLunesr, F.Z.S..,
Curator of Reptiles.

[Received October 18, 1912: Read October 29, 1912. ]

The little fish Girardinus peciloides, popularly known as
‘‘Millions” in Barbados on account of the enormous numbers
frequenting all the shallow pools in that island, is said to be
of great practical value on account of its devouring mosquitoes ;
the absence of malaria, which is so prevalent in the neighbouring
islands, is believed to be due, as first pointed out by Mr. C.
Kenrick Gibbons, to the presence of this fish, for the insect con-
veying the malarial disease breeds only in shallow pools, where
these fish abound and, it is well to add, have no competitors,
G. peciloides being the only freshwater fish inhabiting Barbados.
For this reason, the Society arranged with the Colonial Office to
keep a large stock and to supply specimens to various tropical
stations in the British Empire. The fishes have bred on numerous
occasions in small aquariums in the Reptile and Tortoise Houses,

BREEDING OF GIRARDINUS PCSCILOIDES. 907

and I have thus been able to observe certain facts in relation
to their breeding habits, to one of which in particular I wish to
draw attention, namely that in some cases the male breeds before
the assumption of its secondary sexual characters. Before dealing
with this matter it is necessary to mention briefly the sexual differ-
ences, which are very marked. Females, which may measure up
to 25 mm. in length, are simply of a dull olive-grey in colour,
usually with a dark patch above the anal fin, while the males,
which do not attain quite so large a size, are most conspicuously
ornamented with red, blue, violet and yellow, with a dark ocellar
spot situated in the middle of the body, in front of or beneath
the dorsal fin, and another at the base of the caudal fin, these
spots appearing at least fourteen days before the brilliané hues.
Males are to be further distinguished from females by the
position and prolongation of their anal fin, which is transformed
into an intromittent pairing organ.

The male is remarkably active, and is perpetually courting the
female, going through all kinds of antics in front of her, and as
breeding goes on al through the year, at least in captivity, the
latter is in an almost permanently pregnant condition, and within
a fortnight of having brought forth a brood (such a period
representing the duration of the gestation, at a temperature
of over 70°), once more brings a generation into the world.
It should be borne in mind, however, that a single impregnation
is sufficient for the fertilization of several broods, the embryos of
the second and third generations being already in an advanced
condition when the first young are born. Each brood comprises
from five to as many as twenty-five fish.

The young fishes grow rapidly : about 4 mm. long at birth, they
double that length in a fortnight or so, when their sex can be
determined by the shape and position of the anal fin; it is not,
however, until at least six weeks after birth that the males
become adorned with the brilliant colours which characterise the
adult, this period again varying according to the temperature of
the water.

The first brood, containing a male which bred before the
assumption of its secondary sexual characters, was produced on
March 11th of this year, and numbered eleven. As is almost
invariably the case, a number of weaklings died within a few days
of their birth, and eight fish remained at the end of a fortnight,
when I found the sexes to be equally divided, which is remark-
able, for as a rule females outnumber males by about three to one.
On May 7th, six young ones were born of this brood, and at the
time none of the males had developed their adult coloration, the
only markings distinguishing them from the females being the
spots beneath the dorsal fin and at the base of the caudal, and it
was in fact not until six days later that one of these acquired its
full sexual livery. The others followed suit on May 17th and
18th. These broods were kept at an average temperature of 80°.

In another case, a brood of fourteen fishes, born on June 5th,

908 THE SECRETARY ON ADDITIONS TO THE MENAGERIE.

and of which only seven survived, two males and five females,
reproduced their species on September 29th and 30th, the adult
coloration of the males appearing only on October 7th and 16th.
When we consider the fact that these fishes, owing to the warmer
tanks being already occupied, were kept at the low average
temperature of 65°, we must conclude that, like their growth, the
period of gestation must have been slow, and that the male
undoubtedly bred at least a month before acquiring its full
secondary sexual characters.

A parallel case is well known in the male Salmon, which
occasionally becomes sexually mature in the parr condition,
while the Wrass (Corts julis) and the Dragonet (Callionymus
lyra) have been observed by Mr. H. W. Holt also to become
sexually mature prior to having fully developed their adult
characters. In the two latter cases, however, the question is
one of degree only, the characteristic male livery not being
entirely absent, as in Girardinus.

EXHIBITIONS AND NOTICES.
October 29, 1912.

Prof, E. A. Miycuin, M.A., F.R.S., Vice-President,
in the Chair.

The Srcrerary read the following report on the Additions that
had been made to the Society’s Menagerie during the months of
May, June, July, August, and September 1912.

May.

The registered additions to the Society’s Menagerie during the
month of May were 448 in number. Of these, 207 were acquired
by presentation, 60 by purchase, 43 were received on deposit,
72 in exchange, and 66 were born in the Gardens.

The number of departures during the same period, by death
and removals, was 203.

Amongst the additions special attention may be directed to :—

A large collection of Mammals and Birds from Nepal, including
1 Indian Rhinoceros (2hinoceros unicornis), 1 Kiang (Zquus
kiang), 2 Barasingha Deer (Cervus duvaucelli), 2 Sambur Deer
(Cervus aristotelis), 3 Hog-Deer (Cervus porcinus), 2 Monaul
Pheasants (Lophophorus impeyanus), 2 Cheer Pheasants (Catreus
wallichi), and 2 Wedge-tailed Fruit-Pigeons (Sphenocercus sphe-
nurus); presented on May 21st by H.M. Tue Kine.

A pair of European Bison (Dison bonasus), from Russia, pre
santed by H.G. the Duke of Bedford, K.G., on May 15th.

———

THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 909

1 Black Leopard (felis pardus), from Assam, purchased on
May 21st.

1 Branded Palm-Civet (Arctogalidia stigmatica), from Sarawak,
new to the Collection, presented by Granville J. Altman, Hsq.,
F.Z.S., on May 16th.

1 Civet (Viverra civetia), from EK. Africa, and 2 Egyptian
Mongooses (Mungos ichnewmon), from Zagazig, presented by
H.E. Hassan Hasib Pasha on May 21st.

1 Ural Owl (Syrniwm uralense), from Northern Europe, pre-
sented by E. G. B. Meade-Waldo, Esq., V.P.Z.S., on May 8th.

A collection of rare Reptiles from India, received in exchange
on May 21st, containing, amongst others, 2 Banded Kachugas
(Kachuga dhongoka) and 3 Variegated Snakes (Zamenis fascio-
latus), new to the Collection, and 2 Gharials (Gavialis gangeticus).

JUNE.

The registered additions to the Society’s Menagerie during the
month of June were 511 in number. Of these 305 were acquired
by presentation, 66 by purchase, 50 were received on deposit,
13 in exchange, and 77 were born in the Gardens.

The number of departures during the same period, by death
and removals, was 205.

Amongst the additions special attention may be directed to :—

A collection of Malayan Animals presented by the Government
of the Federated Malay States and other Donors on June 24th,
including 1 Indian Elephant (Hlephas indicus), 1 Tiger (Melis
tigris), 1 Clouded Tiger (felis nebulosus), 1 Binturong (Arctictis
binturong), 2 Argus Pheasants (Argusianus argus), 4 Rufous-tailed
Fireback Pheasants (Aconws erythrophthalmus), and 1 Malayan
Peacock-Pheasant (Polyplectron bicalcaratum), besides the
following, which were all new to the Collection, viz. :—1 Short-
tailed Mongoose (Jungos brachyurus), 10 Charlton’s Tree-
Partridges (drboricola charltoni), 1 Water-Cock (G@allicrea
cinerea), 2 Water-Snakes (Acrochordus javanicus), 1 Bowring’s
Gecko (Hemidactylus bowringii), 1 Bridled Gecko (/Hemidactylus
Jrenatus), and 1 Leschenault’s Gecko (Hemidactylus leschenaultii).

2 Malayan Tapirs (Vapirus indicus) 5 2, from Palenibang,
Sumatra, received in exchange on June 22nd.

1 Grys-bok (Nototragus melanotis) 3, from Cape Colony, pre-
sented by Dr. Louis Péringuey, C.M.Z.8., on June 3rd.

1 Shoebill (Baleniceps rex), from the Sudan, presented by
Lt.-Gen. Sir Francis Wingate, G.C.V.O., on June 29th.

2 Ocellated Turkeys (Meleagris ocellata), from Central America,
deposited on June 24th.

2 Chestnut-bellied Rock-Thrushes (Petrophila erythrogastra),
from the Himalayas, new to the Collection, received in exchange
on June 17th.

2 Brown-backed Robins (Thamnobia cambaiensis), from India,
new to the Collection, purchased on June 18th.

910 THE SECRETARY ON ADDITIONS TO THE MENAGERIE.

1 Temminck’s Robin (Hrithacus komadori), from Japan, new to
the Collection, presented by Wilfrid Frost, Esq., on June 14th.

2 Grey Thrashers (Z'oxostoma cinerewm), from Lower California,
new to the Collection, presented by H. D. Astley, Esq., on
June 15th.

6 Chilian Siskins (Chrysomitris uropygialis), new to the Col-
lection, presented by Arthur C. Macdonald, Esq., on June 26th.

2 Himalayan White-throated Ground-Thrushes (Geocichla cya-
nonotus), bred in the Menagerie.

12 Russell’s Vipers (Vipera russelli), born in the Menagerie.

JULY.

The registered additions to the Society’s Menagerie during the
month of July were 289 in number. Of these 155 were acquired
by presentation, 13 by purchase, 57 were received on deposit,
25 in exchange, and 39 were born in the Gardens.

The number of departures during the same period, by death
and removals, was 301.

Amongst the additions special attention may be directed to :—

] African Buffalo (Bubalus caffer) 3, from the Ishasha River,
Uganda, presented by Capt. E. H. Reid on July 29th.

1 Vaal Rhe-bok (Pelea capreolus) 3, from Basutoland, pre-
sented by M. H. Bosworth Smith, Esq., on July 6th.

2 Yellow-headed Wagtails (Motacilla citreola) and 1 Brown
Shrike (Lanius cristatws), from India, new to the Collection,
presented by W. J. C. Frost, Esq., on July 25th.

1 Changeable Hawk-Hagle (Spizaétus limnaétus), from India,
new to the Collection, deposited on July 25th.

1 Hemprich’s Gull (Larus hemprichi), hatched in the Menagerie
on July 22nd.

1 Black-and- White Cobra (Vata melanoleuca), from Dunkwa, Gold
Coast, presented by Dr. H. G. F. Spurrell, F.Z.8., on July 16th.

1 Snake-Fish (Polypterus senegalus), from the Upper Nile,
presented by the Rev. Archibald Shaw on July 9th.

AUGUST.

The registered additions to the Society’s Menagerie during
the month of August were 226 in number. Of these 74 were
acquired by presentation, 40 by purchase, 55 were received on
deposit, 9 in exchange, and 48 were born in the Gardens.

The number of departures during the same period, by death
and removals, was 285.

Amongst the additions special attention may be directed to :—

1 White-thighed Guereza (Colobus vellerosus), from West
Africa, presented by Miss Jean Aylwin on August 30th.

1 Roebuck fawn (Capreolus capreolus), from Ross-shire, pre-
sented by the Karl of Altamont, F.Z.S., on August Ist.

1 Mount Morrison Laughing-Thrush (Zvrochalopterum morri-
somanum), from Formosa, new to the Collection, deposited on
August 5th.

MR. R. LYDEKKER ON GAZELLA HAYI. 911

1 Magpie-Tanager (Cissopis leveriana), 3 Australian Crimson
Finches (% coohmia phaéton), 2 Peale’s Parrot-Finches (Lrythrura
pealet), and 6 Douglas Quails (Lophortyx douglasi), all bred for
the first time in the Menagerie.

1 Matamata Terrapin (Chel ys fimbriata), from Brazil, presented
by Oswald Allen, Esq., on August 10th.

12 Six-banded Cyprinodons (Haplochilus sexfasciatus), from
rast Africa, new to the Collection, presented by Dr. H. G. F.
Spurrell, EAS, on August 21st.

SEPTEMBER.

The registered additions to the Society's Menagerie during
the month of September were 227 in number. Of these 59 were
acquired by presentation, 18 by purchase, 20 were received on
deposit, 24 in exchange, and 106 were born in the Gardens.

The number of departures during the same period, by death
and removals, was 247.

Amongst the additions special attention may be directed to :—

4 Pelzeln’s Gazelles (Gazella pelzelni), presented by Dr. R. KK.
Drake-Brockman, F.Z.S., on September 26th.

1 Pelzeln’s Gazelle (Gazella pelzelni), presented by A. Gibbs,
EKsq., on September 26th.

1 Capybara (Hydrocherus hydrocherus), presented by Sir Walter
Egerton, K.C.M.G., F.Z.8., and J. J. Nunan, Esq., on September
2nd.

1 White-tailed Gnu (Connechetes gnu), presented by the
Government of South Africa on September 28th.

1 Kordofan Girafte (Giraffa camelopardalis antiquorum) 6,
presented by Lt.-Gen. Sir Francis Reginald Wingate, G.C.V.O.,
K.C.M.G., C.B., D.S.O., on September 13th.

1 Brown Tyrant (/yiarchus tyrannulus), from Uruguay, new
to the Collection, presented by Mrs. EH. F. Dickinson on
September 3rd.

1 Great Indian Bustard (Hupodotis edwardsi), new to the
Collection, presented by Sir John Hewitt, K.C.S.I., on September
30th.

2 European Bee-eaters (Werops apiaster), received in exchange
on September 11th.

2 Snake-headed Fishes (Ophiocephalus punctatus), from India,
new to the Collection, received in exchange on September 7th.

‘* GAZELLA HAYI” = GAZELLA FUSCIFRONS.

Mr. R. Lypexxer, F.R.S., F.Z.8.*, contributed the following
note :-—

‘“‘ As the result of an unfortunate accident, namely the trans-
position of the registration labels of two gazelles received
simultaneously at the British Museum, I find that I have
described and figured a specimen of the Seistan Gazella fuscifrons

* By permission of the Trustees of the British Museum.

919 MRS, R. HAIG THOMAS ON THE

as a new African species under the name G. hayi (Proc. Zool.
Soc. 1911, p. 961, text-fig. 193A). My apologies are due both
to the Society and to naturalists in general for this most unfor-
tunate mistake. :

The text-figure of the so-called G. hayi will serve as an
illustration of the male of G. fuscifrons. The gazelle collected
by Mr. Hay in Algeria is, I believe, G. isabella.”

The Rev. T. R. R. Stepsine, M.A., F.R.S., F.Z.S., communi-
cated a memoir on the Crustacea Isopoda collected by the
‘Porcupine’ Expedition in 1869-1870, in which he described
one new family, two new genera, and four new species.

This memoir will be published in the ‘ Transactions’ in due
course.

Mrs. Rosz Haic Tuomas, F.Z.8., exhibited the eggs of Phasi-
anus formosanus, P. versicolor, and their F.1 and F. 2 offspring,
and made the following remarks :—

“ Last February I exhibited to the Society a series of Pheasant
skins of an experimental cross between a Formosan female and a
versicolor male, illustrating the transmission by the male parent
to his female offspring of most of the female characters of his

Text-fig. 126.

Eggs ot P. versicolor, F.1, Fo.X Ve., F.2, Fo.X Ve.X Ve., and P. formosanus.

race: further evidence of this is now exhibited, namely, the eggs
of the two parent species and those of their F.1 and F. 2 offspring.
It is plain that a considerable difference in dimensions exists
between the eggs of P. formosanus, the larger, and P. versicolor
the smaller of the two: the expectation was that the eggs of the
offspring would resemble in size the egg of the female parent
species rather than that of the male parent species ; but, on com-
paring them, it is clear that the eggs of F.1 and F. 2 are simply

EGGS OF PHEASANTS. 913

P. versicolor. A photograph (text-fig. 126) by Mrs. Peter Haig
Thomas of the four eggs shows very accurately the comparative
Sizes.

The measurements of these eggs are as follows :—

Girth of length. Girth of breadth.

P. formosanus ......... 5; ins. 412 ins.
JE GORSUGNOP o4..204008. 4 Ale ,, 44 ,,
Fl Bo aVies eee. Ale ,, 45. ,,
Ha Hos GiViey<aVier: Ale ,, 45. ,

showing a slight increase in bulk in the eggs of F. 1’ and F. 2
over that of the versicolor.

Another character of which notice was omitted in February
was the iris colour: unfortunately F.19 died eggbound last May
before it occurred to me to make an examination. The male
and female versicolor differ in the colour of the iris, as do also
the male and female formosanus. The male versicolor has a pale
yellow iris, the female a soft brown centre with an outer ring of
pale yellow: the male formosanus has a red-brown iris, the female
a dark brown centre with outer ring of red-brown. Holding
formosanus, versicolor, and F. 1 hens close together, it is seen that
F. 1 has the female versicolor iris with the ring-colour a very
slightly deeper shade of yellow. Again, outstretching the wings of
these three hens side by side, it is observed that besides the secon-
daries and primaries the two groups of major coverts overlying
these feathers in F.1 9 also correspond exactly in pattern and
colour with the same groups in the versicolor 9 .”

At the conclusion of the Scientific Business, Sir Epmunp G.
Lover, Bt., V.P.Z.S., showed a large number of slides, including
a fine series of coloured subjects, and a number of living speci-
mens, to demonstrate the capacity of the new electric lantern
which he had recently presented to the Society.

a7

ah
ic ve.
‘

Aan)

No. 112.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*
October 29th, 1912.

Prof. EK. A. Mincuty, M.A., F.R.S., Vice-President, in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The Secretary read a Report on the Additions that had been
made to the Society’s Menagerie during the months May to
September, 1912.

Mrs. Rosze Haig Tuomas, F.Z.8., exhibited the eggs of
Phasianus versicolor, P. formosus, and of the F, and F, offspring of
an experimental cross between a male P. versicolor and a female
P. formosus. She drew attention to the resemblance in size of
the eggs of the offspring and of the male parent species, whereas
the expectation was a likeness to those of P. formosus, thus
showing the descent through the male to his female offspring of
the small egg of his species.

Mr. R. Lypexxer, F.R.S., F.Z.8., contributed a note pointing
out that, owing to an unfortunate mistake in labelling, the
Gazelle he described in the Society’s ‘ Proceedings’ for 1911 asa
new species, Gazella hayi, was really G. fuscifrons.

Mr. LyDEKKER also described the Bornean Bantin as a new
race to be distinguished from the typical Javan Bantin by the

* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent's Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications ; but it may be obtained on the
day of publication at the price of Starpence, or, if desired, sent post-free for
the sum of Sta Shillings per annum, payable in advance,

A2

more upright direction and less outward curvature of the horns
of the male.

Mr. EH. G. Boutencrr, F.Z.8., Curator of Reptiles, read a
paper dealing with the breeding-habits of the “ Millions” Fish
(Girardinus peciloides) from observations made in the Society’s
Gardens, and recorded cases of the male of this species breeding
before assuming the livery of its sex. The author drew attention
to parallel cases among fishes, in which, however, except in one
case, the question was one of degree only.

The Rev. T. R. R. Srespine, M.A., F.R.S., F.Z.8., read a paper
entitled “‘The Crustacea Isopoda of the ‘ Poreupine’ Expedi-
tion.” In the tribe Flabellifera, family Gnathiide, two new species
were assigned to the genus Gnathia, and a new genus was
instituted to receive a new species called Akidognathia edipus.
In the tribe Asellota a new family, Thambematide, was proposed
for a new genus with a new species named Zhambema amicorum.
Gnathia cristatipes, sp. n., owed its name to the striking processes
on the second and third joints of the second pereopods, its head
in dorsal view being almost circular. G'nathia schistifrons, sp. n.,
had a head much wider than long, witha rounded notch in the
middle of the front, a peculiarity which it shared with G. ceca,
Richardson, 1911, without sharing the blindness of that species.
Akidognathia edipus, gen. et sp. n., deserved its specific name alike
by the gouty appearance of the second perzopod and the problems
which required solving in regard to its mouth-organs; while,
contrary to the custom of the family, the adult male appeared
to have at least one pair of maxille. In the Asellota Thambema
amicorum, gen. et sp. n., seemed to be anomalous in the tribe by
having only four pairs of pleopods in the male and no uropods;
its narrowly cylindrical shape was also unusual.

This memoir will be published in the ‘Transactions’ in due
course.

Dr. F. EH. Bepparp, M.A., F.R.S., F.Z.S., Prosector to the
Society, read a paper on the Anatomy and Systematic Arrange-
ment of the Cestoidea, in which he gave an account of six species
of Tapeworms from Reptiles, belonging to the genus Jchthyotenia

(Sala)

Mr. E. Duxinrietp Jonss, F.Z.S., F.E.S., communicated a
short paper containing the descriptions of thirteen new species of
Butterflies of the genus 7’hecla which he had collected at various
localities in 8.E. Brazil.

ae
43

LANTERN DEMONSTRATION,

At the conclusion of the Scientific Business, Sir Epmunp G.
Lover, Bt., V.P.Z.S., showed a large number of slides, including
a fine series of coloured subjects, and a number of living speci-
mens, to demonstrate the capacity of the new electric lantern
which he had recently presented to the Society.

The next Meeting of the Society for Scientific Business will
be held on Tuesday, November 12th, 1912, at half-past Hight
o'clock p.m., when the following communications will be made :—

H. R. Hoae, F.Z.8.
Some Falkland Island Spiders.

Bruce F. Cummines.

On some Peints in the Anatomy of the Mouth-parts of the
Mallophaga.

FB, F. Larpraw, M.A., I.Z.8.

Contributions to a Study of the Dragonfly Fauna of Borneo.
—Part I. The Corduliine: the Genus Amphicnemis.

a JAMESON, M. D.Sc. 5 dedi D., FZ. ZS ! 5., and WILLIAM

ia TO
On some ee oF I the noe “Duck (Bdemia nigra) and
their relation to the Pearl-inducing Trematode in the Hdible

Mussel (A/ytilus edulis).

G. A. Boutuncur, F.R.S., I.Z.S.

Descriptions of Three new Fishes discovered in the Gold
Coast by Dr. H. G. F. Spurrell, M.A., F.Z.8.

The following papers have been received :—

G. ave SMITH, M A., and H. J . SCHUSTER, , M.A., D. Sce., E.ZS.

~The Genus Enaceus, or ne Land ae of Australia,

44
The Hon. Paun A. Menmuny, FLAS.
Description of an Amphipod belonging to the Family
Talitrids from the Woodbush, Transvaal.
Dr. F. D. WEtcH.

Observations on living Specimens of the Capped Langur
(Semnopithecus pileatus).

Communications intended for the Scientific Meetings should
be addressed to
P. CHALMERS MITCHELL,
Secretary.
ZOOLOGICAL Society oF LonpDoN,
Recent’s Park, Lonpon, N.W.
November dth, 1912.

No. 1138.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON,*

November 12th, 1912.

Dr. A. Smrra Woopwarp, F.R.S., Vice-President, in the Chair,

The Minutes of the last Scientific Meeting were confirmed.

The SecRETARY read a report on the Additions that had been
made to the Society’s Menagerie during the month of October,
1912.

The Secretary exhibited a photograph of the Hainan Gibbon
_ (Aylobates hainanus) in natural colours, taken by Messrs. Elliott
& Fry, by the Lumiere process.

Mr. G. A. Boutenesr, F.R.S., F.Z.S., exhibited a specimen,
155 mm. long, of the African Cichlid Perch (Hemichromis bimacu-
latus Gill), which had recently died in Capt. Vipan’s aquarium.
The specimen was remarkable for its large size. Although the
species is common over a considerable part of Africa, and hundreds
of specimens have been collected in various localities, no wild
example is known to exceed a length of 100 mm.

This Hemichromis is tinged with bright red about the head
and on the fins, and, as was noticed in the ‘ Fishes of the Nile,’
p- 462, this red is soluble in spirit, which, a few hours after the
immersion of the fish, acquires a bright orange-red colour.

Mr. Epwarp GERRARD exhibited the skull of an Indian
Rhinoceros (AAinoceros unicornis) which had recently died in
the Society’s Gardens. He pointed out that abscesses had been
formed at the base of the lower incisors, which had been much

* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent’s Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications; but it may be obtained on the
day of publication at the price of Stxpence, or, if desired, sent post-free for
the sum of Sia Shillings per annum, payable in advance.

46

rubbed down so that the nerves had become exposed. The
animal must have suffered great pain, and this no doubt was the
cause of his dashing his head against the walls and bars of his
den in the violent manner he at times did.

Mr. EK. G. Boutencer, F.Z.8., Curator of Reptiles, exhibited a
living specimen of the Salamander, Amblystoma tigrinum, which
he had obtained from an Axolotl placed in special conditions
with the object of forcing the metamorphosis.’

Mr. D, Seru-Suiru, F.Z.8., Curator of Birds, gave a lantern
demonstration of photographs of the young of Hemprich’s Gull
(Larus hemprichi), the Black-necked Swan (Cygnus melano-
coryphus), and the White Stork (Ciconia alba) which had been
hatched in the Society’s Gardens during the present year. He
also showed slides of the nest, in a tree near the Apes’ House,
made by the Orang-utan which escaped from its cage on
November 3rd, 1912.

Mr. H. R. Hoce, M.A., F.Z.8., read a paper entitled ‘Some
Falkland Island Spiders,” based on a small collection of Spiders
formed by Mr. Rupert Vallentin during a two years’ stay in the
Falkland Islands,

Of some of the species there were a fair number of specimens,
but they comprised only six species of Spiders and one of the
allied suborder Opilio.

The species were all apparently new, but the genera were all
to be found either in Patagonia, Tierra del Fuego, or the islands
about Cape Horn. The ancestors of the Spiders might all have
been transported aerially at an early period and therefore afforded
no evidence for or against a former land-connection, but in the
event of the latter there should be many more species. The
Opilio might have been conveyed under the bark of floating trees.

Mr. G. A. Bounencer, F.R.S., F.Z.S8., read a short paper con-
taining the descriptions of three new Fishes which had been
discovered by Dr. Spurrell in the vicinity of Bibianaha, near
Dunkwa, Gold Coast, and presented by him to the British
Museum.

A paper was communicated by Dr. H. Lister Jamuson, M.A.,
Ph.D., F.Z.S., and Dr. Wititiam Nicort, M.A., D.Sc., F.Z.S.,
which contained an account of some parasites of the Secoter Duck
(demia nigra), and discussed their relation to the pearl-inducing
Trematode in the Edible Mussel (/ytilus edulis).

A paper was received from Mr. F. F. Larpiaw, M.A., F.Z.S.,
dealing with some Dragonflies from Borneo belonging to the
subfamily Corduliime, and to the genera Disparonewra and

AT

Amphienemis of the subfamily Agrioninz, with an account of a
number of new species.

The next Meeting of the Society for Scientific Business will
be held on Tuesday, November 26th, 1912, at half-past Hight
o'clock P.m., when the following communications will be made :—

Conve Surra, M.A. . and _E, H.. J. . SCHUSTER, BUNS) D.S8e., F.Z 1.0.

“Ais er Te or the Land Capen of Teese

Ee iG ene oE Te in Sic a Contribution to
Paleohistology.

C. L. Boutencer, M.A., D.Sc., F.Z.8.

Report on the Myzostomida collected by Mr. Cyril Crossland
in the Red Sea in 1905,

The Hon. PavuL A. METHUEN, E.Z. 5.

“Deseription of an Amphipod belonging to the Family
Talitride, from the Woodbush, Transvaal.

Bruce F. Cummines.

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

The following papers have been received :—

G. P. FAaRRAN.

Plankton from Christmas Island, Indian Ocean,—II. On
Copepoda of the Genera Oithona and Paroithona,

48

E. EH. Bepparp, M. A., D.Se., , FR.S., PAS

Contributions to he Anatomy and. Speman Arrangement
of the Cestoidea.— VIII. On some Species of J chihyotonia and
Ophidotenia from Ophidia.

H. L. Hawetns, M.Sc., F.G.S.
The Anterior Ambulacrum of Hchinocardiuwm cordatum Penn.,
and the Origin of Compound Plates in Hchinoids.
H. B. Preston, F.Z.8.
Diagnoses of new Species and Varieties of Agnathous
Mollusca.
The Deut] Buffalo st aie Nigeria: with a Revision of
the Dwarf Buffaloes of Western Mists,

Communications intended for the Scientific Meetings should
be addressed to
P, CHALMERS MITCHELL,
Secretary.
ZOOLOGICAL Society oF Lonpon,
ReceEnt’s Park, Lonpon, N.W.
November 19th, 1912.

44,

45,

46,

47.

48.

ass),

50.

51.

52.

— Pape continued).

A Note on the rare British Nudibranen s1ancockia eudactylota Gosse. By Sir Cuaruus
parma @vie en © Nae He Aisa, CetemPEXOX X Vi). 5 wrelate telnleatem acne craw. csv eens

The North Rhodesian Giraffe. By R. Lyprxxer, F.R.S., F.Z.8. (Pl. LXXXVL) ..

On Antler-Growth in the Cervide, with special reference to Haphurus and Odocoileus
(Dorcelaphus). By R. I. Pocock, F.R.S., F.L.S., F.Z.8., Superintendent of the
Gardens and Curator of Mammals. (Text-figs. 108-112.) ............2. 201s sees

Polychxta from the Pacific Coast of North America.—Part I. Surrurip#, with a
Revised Table of Classification of the Genus Spirorbis, By Hutzn L, M. Prxet,
B.Sc., F.Z.S., Demonstrator of Zoology and Reid Fellow, Bedford College, University
Ototsoncdonsey (Ge NONNGV le 1a NENONGIONG ) Sora rasiaretain wleicloyeje) cies sjel sie elloletnieysieletels\niciesels

The One-sided Reduction of the Ovaries and Oviducts in the Amniota, with Remarks
on Mammalian Eyolution. By Hans Gapow, M.A., Ph.D., 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. By Frank H. Bepparp,
M.A., D.Sc., F.R.S., F.Z.8., Prosector to the Society. (Text-figs. 113-121.) .......

On Two new Trematode Parasites from the Indian Cobra. By Witttam Nicott,
M.A., D.Sc., M.D., F.Z.8., Lister Institute of Preventive Medicine, London. (Text-
ING, WOPL) 8 Gane pa hoo 0s ONO Res omnes aquOTe Oe ool ode CM OOO EEIAG agoooeDoLeto de

Statistical Note on the Worm Parasites collected from the Animals dying in the
Zoological Gardens, from December 1910 till April 1912. By Wuu1am Nicott,
M.A., D.Se., M.D., F-Z.S., Lister Institute of Preyentive Medicine, London ........

On some new Fossil Reptiles from the Permian and Triassic Beds of South Africa.
By he DROOM, Disa, OMGA Se - (Pisa XC: XCM) ves caiere «sole ciore of ovelaiv: slemeteks ears
53. Ou the Hydrocoralline Genus, Hrrina. By Professor 8, J. Hickson, F.R.S., F.Z.S.,
The University of Manchester. (Pls. XCIV.=XOVI.) 2.000.000. cise nes cones
54. Descriptions of new Butterflies of the Genus Thecla from §.E. Brazil. By E. Duxry-
Tb) WOKE WAS, IMIS. GEE OO WING) secu nn dosdddecoouldacdsoucbboon Gs bo°
55. The Bornean Bantin. By R. Lyprexxnr, F.R.S., F.Z.8, (Text-figs. 123-125.) ......
56. Notes on the Breeding of the “ Millions” Fish (Girardinus pwciloides). By Epwarp
Gy) BOunENGER, We Zcs,,.© orator Of Reptiles \fety (ieee csi nee vn eles ane merely e mee ce
SUES): ooo sabcdondo co omusuonH Onn OOd:.o0 DOOD a na Oe cee reer a SBODC
Mist) Ole COUMeI game OMeenemire rats. ors) sceiersie eiepetic. hecsteaasn.er¥naiare: v'd vie sisi mpeiwiei quale Sat atete :
ITE Ore Ole SSG Sb .co Jb Os on SReRU Uh SojoUA DOU oon moo a gmonem shot m ae.0n.0.5 aeEe
Alphabetical List of Contributors .......... .. Peete te one clan fares Pol aoner Petre ftene ee Sint siere
iNew, Generics Derm Seestaar Ueuas eric dsis wccre aiaters fe sikeioieioia/oce sie ais ofetaheiste Sidoueteiete: ¢ oisaaacoenenenaes
Imdexiof Seientine sNanmesie srreterta cs aia stelniaimon cate s)ere cueteleleiei7- Serpsleuredsoneees «cicero tarsi aya:
Hridex of TlustPations) settee esis. wales Seale os etsy tau Oe PRSISEEE «elaine aid erate Shee

784

808

822

LS EOF aP LA Paige

1912, Part LV. (pp. 757-913).

Plate Page
EXERT Dipreropeliis Wenundo aedaas cae srt es) steele eee 763
XX). “Hancockia eudactylord, te. «+ oe es)ce tine, ee ae onlay 770
LXXXVI. Giraffa camelopardalis thornicrofti ...+....++.+00-- 771
LXXXVII. Y
PX exeV North American) Serpalidee . wv. jes) teehee se eran 784 ty
LXXXIX. ar
XC. \
XCI. | ;
‘ South African Fossil Reptiles ....................-. 859 ¥
XCII. | . ml
XCTII. } : :
XCIV. j
XCV. ¢ Hydroids of the Genus Errina ........ Raat ee eal pel eye 876
XCVI. . | ;
XCVII. New Species of Thecla from 8.H. Brazil .............. 896
NOTICE.

The ‘ Proceedings’ for the year are issued in fowr parts, paged consecutively, ; { j
so that the complete reference is now P. Z. 8.1912, p.... The Distribution oH ;
is as follows :— at

Part I. issued in March.
SEN Ei Bape June.
ope aU ay September. |

spat oRLAV ger cri December,

‘ Proceedings, 1912, Part III. (pp. 505-756), were published on :
August 29th, 1912.

The ‘ Abstracts of the Proceedings,’ Nos. 112 & 113, are
contained in this "Gy .

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