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PROCEEDINGS

OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCLETY

OF LONDON,

1919, pp. 1-225,

with 10 Puatrrs and 72 TEx. FIGURES.

Z<ansonian Institags ~

: 2AaGAg, y
Wetional Muses: Y

PRINTED FOR THE SOCIETY,
SOLD AT ITS HNOUSH IN REGENTS PARK.
LONDON:

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

ease

OF THE

COUNCIL AND OFFICERS

OF THE

ZOOLOGICAL

SOCIETY

OF LONDON.

WSS):

Patron.
His Masnusry Tare Kine.

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

Tur Hon. Cecit Barine, M.A.
THE Lord CarMICHAEL,G.C.S.L.,
K.C.M.G.

Lr.-Conr. 8S. Monckton
MAN, M.D., F.R.S.

CHARLES DrumMmonp, 4Hsq.,
Treasurer.

Aurrep Hzra, Esq., Vice-

President.
Haroun 8. Fercuson, Esq.
Carr. Huan
M.A.
Aurrep H. Cocks, Esq., M.A.
Pror. James P. Hin, D.Sc.,
BERS:
WiiiiAM Huntsman, Esa.

Corr-

S. GLADSTONE, |

E. G. B. Mrapr-Watpo, Esq.

Pror. Ernest W. McBripe,
D.Sce., F.R.S., Vice-President.

Major ALpErr Pam.

P. Cuaumers Mircnuein, Esq.,
C.B.E., M.A., D.Sc., Li...
F.R.S., Secretary.

AvrRIAN D. W, Pouttock, Esq.

THe Lorp QUEENBOROUGH.

THe Marguts oF Stiico, F.S.A.
Vice-President.

Mayor Ricwarp 8. Taynor.

Antuony H. Wincrreip, Ksa.

ArrHour Situ Woopwarp,
Kiso) (HIND: RS: = Vace=
President.

PRINCIPAL OFFICERS.
P. Cuanmers Mircuety, C.B.E., M.A., D.Se., LL.D., F.R.S.,

Secretary.

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

D, Sera-Suira, Curator of Birds and Inspector of Works.

Lieut. Epwarp G. BouLmncrr, Curator of Reptiles.

Prof, H. Maxwett Lurroy, Curator of Insects.

Joun Barrow, Accountant.
W. H. Conn, Chief Clerk.

VW «LIST OF CONTENTS.

1919, pp. 1-225.

EXHIBITIONS AND NOTICES.

The Szcrerary. Report on the Additions to the Society's
Menagerie during the months of November and

Wecemiber elon cence tar re noncancer Se ey eae:

Mr. C. Davirs SHERBORN, F.Z.S. Exhibition of, and re-
marks upon, a letter, written in 1693 by Malpighi
LOM Ee Mat ewe Mabel § cnc i.tus ita h ss fue nase nas takes Sere:

Sir Douetas Mawson. Lantern exhibition of, and remarks
upon, Australasian Antaretic and Sub-Antaretic life.

The Secrerary. Report on the Additions to the Society’s
Menagerie during the month of January, 1919 ......

The Secrerary. Report on the Additions to the Society’s
Menagerie during the month of February, 1919 ......

Mr. F, Martin Duncan, F.R.M.S. Exhibition of, and re-
marks upon, a series of photographs and lantern-
slidesiot Marine Zooloonien a. acco. tenancies

Dr. F. E. Bepparp, F.R.8. Exhibition of, and remarks
upon, the tcetal Sperm Wrballes) {5.2..2-¢- 5:5 sceneeas ens:

Mr. R. I. Pococn, F.R.S., F.Z.8., Curator of Mammals.
Exhibition, illustrated by lantern-slides, to show
structural characters of the Felide .................005-

219

Tue Secrerary. Report on the Additions to the Society’s
Menagerie during the month of Mareh, 1919 .........

Mr. T. Gerrarp, F.Z.8S. Exhibition of, and remarks upon,
alseries of eadsrok VV eiier Ucar eames a serra cere rrr 4.

Dr. W. T Cauman, F.Z.S. Exhibition and detailed account
of various Marine Boring Animals ....................

Tus Srecrerary. Exhibition, on behalf. of Mi. George
Jennison, of a sevies of lantern-slides of a Chimpanzee
kept am! the! Open ait tae .e.. seats eee eceeeee ere eas

Tue Secrerary. Exhibition of, and remarks upon, two
plrotoemayolis) of ety byvaimeg ©) ical pit amerer ere e ener nee eee oi

Mr. E. G. Boutencr, F.Z.8. Exhibition of, and remarks
upon, British Rats and their varieties ..................

Lt.-Col. S. Moncxron Copeman, F.R.S. Exhibition of a
series of iantern-slides to illustrate his ‘¢ Experiments
Onn rare IDE nSTNNMAMTOM ” - sescancsacudodsoddscndooescaasdonoes

The Secrerary. Report on the Additions to the Society’s
Menagerie during the month of April, 1919............

Tue Secretary. Exhibition of, and remarks upon, the
photographs of the young living Okapi shown at the
previous Scientific Micebim ge.) Wali se aeetscneeRe nm seeeee ee

THe Secrerary. Report on the additions to the Society’s
Menagerie during the month of May, 1919

eee eer ees oce

Miss L. HE. Curresman, F.E.S. Exhibition of living speci-
mens of Light: giving Beetles from Cuba ...... reece eee:

Mr. EK. Htron-Auuen, F.Z.8., Exhibition of a series of
lantern-slides demonstrating results of experiments
on cultivation of Vernewilina

225

10.

PAPERS

. On the External Characters of existing Chevrotains.

By R. I. Pocock, F.R.S., F.Z.8. (Text-figures 1-5.)

. Report on Deaths of Animals in the Gardens in 1918.

By J. A. Murray, M.D., Acting Hon Pathologist to
CASO CVCEY | Atma teteae ates St eS AME RS PG |

On a Collection of Fishes from Lake Tanganyika, with
. Descriptions of three new Species. By G. A. Bou-
LENGER, F.R.S., F.Z.8. (Text-figures 1-3.) ............

. On the Skull and Affinities of Rana subsigillata A. Dum.

By Miss Joan B. Procrmer, F.Z.S8. (Text-figures
ROSE 2s) es ctarseean a Pa asegiiers nto scene see eae orate

. On the Breeding of Oryx gazella at Gooilust. By

Bel eb rANuNa MEAS.) (CBlatevl.)) fe sca.cedossess.cee

. A Comparative Study of certain Sense-Organs in the

Antenne and Palpi of Diptea. By K. M. Smrru,
A.R.C.S.,. D.1.C. With Appendix by Professor H.
Maxwetu Lerroy, F.Z.S. (Plates I-IV. and Text-
{SUSU SS 225 5) PR oredr eeieea orca oe aoc CRMC

. The Progressive Reduction of the Jugal in the Mam-

malia. By Lancetor T. Hoesen, B.A., BSe..........

. Descriptions of two new Lizards and a new Frog from

the Andes of Colombia. By G. A. Boutenerr, F.R.S.,
EZ. Chext four eStA soto)» sem scmscttccesarat ssc sueqane ss

A Unique Case of Asymmetrical Duplicity (Duplicitas
asymmetros) in the Chick. By Nort Tayre*, B.Sc.
(Lond.). (From the Zoological Department, Univer-
sity of London, University College.) (Plates I-III.
aman bext=troures: | de) ess geeees eves nwneecaceinasteiseisieeaes

Some Points in Insect Mechanics. By H. R. A. Mat-
LOOK, RS. ZS. Mext=tigures 1-8.) 21.2 2.526::

Page

11. On some Equatorial and other Species and Genera of
African Iehneumonine contained in the Collection of
the British Museum. By CraupE Mortey, F.E.S.,
1 RY ass a 2 ORME iat a rR eve te SR ee

12. Results of a Mendelian Experiment on Fowls, including
the Production of « Pile Breed. By J. 'T. Cunnine-
HAM. J MeAt.. SER ZS. “Blaite wh caanceee tes taatae eee ae

13. Some Points in the Anatomy of the Takin (Budorcas
taxicolor whiter). Based on the examination of a
specimen in the Zoological Society of London. By
Miss Karanern F. Lanper, M.S8c., F.Z.S. (Hon.
Acting Prosector to the Society). (Plate I. and
ence asus ERE ao. ciate Meant he ao) Mie eae ane ae thea

Alphabetical List of Contibutors

Gave rep cod eek ee it Oe RN OP ty WM ah ek UU eg? NW he Wee 2 Meee

Pave

Lay,

203

APA RHE rCAh List
OF TIE

CO NR OURS.

With References to the several Articles contributed by each.

(1919, pp. 1-225.)

Bepparp, Dr. F. E., F.R.S.
Exhibition of, and remarks upon, three fetal Sperm-

NUM TINSLVSIS Es Sere ae Mae oe woe ere ee a we

Briaauw, F. E., C.M.Z.8.
On the Breeding of Oryx gazella at Gooilust. (Plate I.)

Boutencer, EK. G., F.Z.S.
Exhibition of, and remarks upon, British Rats and

IS TIGRVAULCLITC SUNN 20 1 Poh. 8 41, Rami ens Gace Cae ga eS

BouLencer, G. A., F.R.S., F.Z.S.
On a Collection of Fishes from Lake Tanganyika, with
Descriptions of three new Species. (Text-figures 1-3.) .
Descriptions of two new Lizards and a new Frog from
the Andes of Colombia. (Text-figures 4 & 5.) ...........,

Catman, Dr. W. T., F.Z.S.
Exhibition and detailed account of various Marine

[SOAS NOVO IS, > ORAM aE. n> Me eee a

221

224

222

Vili
CaeEsmAN, Miss L. E., F.E.S.

Exhibition of living specimens of Light-giving Beetles
ibeoviet Ol Gl Meenmo ae Aree seen eG SU onan soln aoenene De cb

Corrman, Lt.-Col. 8. Monckton, F.R.S.

Exhibition of a Series of lantern-slides to illustrate his

“ Kixperiments on Sex Determaiinarnlonia eee

Cunninauam, J. T., M.A., F.Z8.

Results of a Mendelian Experiment on Fowls, including
the Production of a Pile Breed. (Plate I.) ...............

Duncan, F. Martin, F.R.M.S.

Exhibition of, and remarks upon, a series of photo-

graphs and lantern-slides of Marine Zoology ...............

Gerrard, T., F.Z.8.

Exhibition of, and remarks upon, a series of heads of
AVY oT 1G) eae eRe ERMC rete tra cranes bonadmenunoasnocae

Heron-Autey, E., F.R.S., F.L.S., F.Z.8.

Exhibition of a series of lantern-sliles demonstrating

results of experiments on cultivation of Verneuwilina ......

Hoesen, Lancetot T., B.A., B.Sc.

The Progressive Reduction of the Jugal in the Mam-

TVAULTS, A Age head cisctigs deiaheh ued clvichevee eke ee

Lanper, Miss K. F., M.Se.. F.Z.S. (Hon. Acting Prosector
to the Society.)

Some Points in the Anatomy of the Takin (Budorcas
taxicolor whitei). Basel on the examination of a
specimen in the Gardens of the Zoolugical Society of
London. (Plate I. and Text-figures 1-7.) ....,,......

Page

225

173

222

203

ix
Lerroy, Prof. H. M., F.Z.S.

Appendix to Paper by K. M. Smith on a Comparative
Study of certain Sense-Organs in the Antenne
and Palpi of Diptera. (Plates I-IV. and Text-
AUST Senki cwisneies cia aa cianint diepssree oa som vows hanes verses tei

Matuocg, H. R. A., F.R.S., F.Z.S.

Some Points on Insect Mechanics. (Text-figures 1-8.)

Mawson, Sir Dovenas.

Lantern exhibition of, and remarks upon, Australasian
Antarctic-andySub-Amtaretic Wife... bse napen eh seems

Mircueut, P. Cuaumers, O.B.E., M.A., LL.D., F.B.S.,
F.Z.8., Secretary to the Society.

Report on the Additions to the Society's Menagerie
during the months of November and December, 1918 ...

Report on the Additions to the Society’s Menagerie
duritmsithe month ole Jantiany sel QUO 2 oe veaecececanee ace:

Report on the Additions to the Society’s Menagerie
dining thie month of Hebruanry, NOUS nem ectseceen cece

Report on the Additions to the Society’s Menagerie
Guciae fhe month of March; UU Oi. ascwea seme dee saci

Exhibition, on behalf of Mr. George Jennison, of a

series of lantern-slidesofa Chimpanzee keptin the open air

Exhibition of, and remarks upon, two photographs of
Dy Nye OAD ae ceciaie <= nov cernisctle nee es eucooane menses nce ss

Report on the Additions to the Society’s Menagerie
Cuicimeihemmonti Of April RO ewes eces ct naceccem es

Exhibition of, and remarks upon, the photographs of
the young living Okapi shown at the previous Scientific
NCSU ee sch icc g saiuss 0% ans bale eect staan meta iol interna desc acl oyeis

Report on the Additions to the Society’s Menagerie

dimming themonbhrot Mai, ONO mies leacievelatnlerals-js.s min seeiee
Proc. Zoot. Soc.—1919. b

Page

Walt

219

223

Mortey, Ciaups, F.E.S., F.L.S., &e.

On some Equatorial and other Species and Genera
of African Ichneumonine contained in the Collection of
the sBritishwMiuseuin...<0etesck «1c Beets Lee ee

Murray, J. A., M.D., Acting Hon. Pathologist to the
Society.

Report on Deaths of Animals in the Gardens in
1 OES ae Sle aula a OA Ge Se ie tad eaten tee ae ee ee

Pocock, R. I., F.R.S., F.Z.8., Curator of Mammals.
On the External Characters of existing Chevrotains.
(Mext-fig ures, 1-5: ia. cenges seen ace cleat eee eee

Exhibition, illustrated by lantern-slides, to show struc-
tunallleharactersiot the selidcews se. 1.3 ere eeee ener rere

Procter, Miss Joan B., F.Z.8.
On the Skull and Affinites of Rana subsigillata A. Dum.
(Text-figures 1 & 2.)

Peewee eee eee re eas oases sessoresesresserseeseee

SHERBORN, C. Davizs, F.Z.S.

Exhibition of, and remarks upon, a letter written in
1693 by Malpighi to Dr. Mathew Faber ..................6.-

Smiru, K. M., A.R.C.S., D.I1.C.

A Comparative Study of certain Sense-Organs in the
Antenne and Palpi of Diptera. (Plates —-IV.and Text-
hears L4G.) 2. 5 odes wa Vsteca ne le eee

Tayuer, Nort, B.Sc. (Lond.).
A Unique Case of Asymmetrical Duplicity (Duplicitas
asymmetros) in the Chick. (Plates I-III. and Text-
Ais UTes) Mey 2.) wah ees Abid icy) «ACER REA ee een

Page

219

INDEX.

1919.—Pages 1-225.

[New names in clarendon type.

Systematic references in italics,

(z.8.U.) indicates additions to the Society’s Menagerie. |

Agama stellio (z. s. L.), 219.
Aglaojoppa glabrinotor, sp. n.,

138.

Amblyteles auricomus, sp. n.,

159.

—— spilopterus, sp. n., 158.

—— testaceator, sp. n., 160.

Anolis apollinaris, sp. n., 79.

Anthropopithecus troglodytes (z. 8. L.),
PNG),

AVES:

A unique case of Asymmetrical Du-
plicity in the Chick: structure, 83 ;
Results of a Mendelian experiment
on Fowls, including the production
of a Pile Breed: structure, 173.

Barichneumonconcinnator,sp.n.,
sess

—— fossifer, sp. n., 154.

—— planinotum, sp. n., 154.

Benecles dimidiatus, sp. n., 171.

politanus, sp. n., 171.

Bison americanus (4. 8. L.), 222.

Bos taurus (z. 8. L.), 222.

Calophasis mikado (z, 8. 1,), 222, 225.

Catadelphus rubricaput, sp. n.,
123:

Chasmias ruficaudator, sp. n.,
157.

Chysotis ventralis (z. s. L.), 221.

Clemmys caspica (z, s.L.), 221.

Coelichneumon cornellifer, sp. n.,
140.

— geminifer, sp. n., 142.

—— globulifer, sp. n., 142.

— scopulifer, sp. n., 140.

— striatifer, sp. u., 1438.

— sublunifer, sp. n., 141.

—— sulcifer, sp. n. 142.

—— thyridifer, sp. n., 148.

Corymbichneumon, ¢. n., 136.

----- Carinifer, sp. n., 136.

Cratichneumon testacecolor,
sp. n., 153.

Ctenichneumon castanopygus,
sp. n., 164.

Ctenochares gracilentor, sp. n.,
127.

—— microcephala, sp. n., 126.

Cygnus melanocoryphus (z. s. u.), 224.

Epijoppa carinifer, sp. n. 120.
—— pygidifer, sp. n., 120,

xii INDEX.

Epijoppa striatifrons, sp. n., 121.

triangulifer, sp. n., 121.

Equus asinus (2. 8. u.), 224.

—— prévyi (Zz. s. L.), 224.

Erythrojoppa nigripedalis, sp. n.,
125.

—— rufipedalis, sp. n., 124.

Felis bengalensis (z. s. u.), 222.

GEOGRAPHICAL:
Descriptions of two new Lizards and
a new Frog from the Andes of
Colombia, 79; Lantern-exhibition
of Australasian Antarctic and Sub-
Antaretie Life, 219.

Haplochilus dhonti, sp. n., 17.
Hyena crocuta (z. Ss. L.), 222.

Insncra :

A comparative study of certain sense-
organs in the autenne and palpi
of Diptera: structure, 31; Some
points in Insect mechanics: struc-
ture, 111; On some Equatorial and
other Species and Genera of African
Tchneumouins contained in the
collection of the British Museum :

117; On

Light-giving Beetles from Cuba,

structure, systematic,

exhibited, 225.

Lagenesta duplicator, sp. n., 146.

—— sinifer, sp. n., 145.

—— triangulifer, sp. n., 147.

—— triplicator, sp, n., 146.

Lama gluma (z. 8. L.), 224.

Lamprologus dhonti, sp. n., 19.

Leontocebus rosalie (z. 8. 1.), 220.

Leptothecus alutacefer, sp. n.,
150.

—— mesonotifer, sp. n., lol.

—— punctifer, sp. n., 151.

Lophophorus impeyanus (z. 8. L.), 222.

Lycaon capensis (%. 8. u.), 225,

Macropus bennetti (z.s.u.), 221, 224.
Mag wenga, ¢. n., 166.

— maculipennis, sp. n., 166.
Mamarra:

On the external characters of exist-
ing Chevrotains: structure, 1; On
the breeding of Oryx gazella at
Gooiluat: structure, 29; The Pro-
gressive Reduction of the Jugal
in the Mammalia: structure, 71 ;
Some points in the anatomy of the
Takin (Budorcas taxicolor wh:tet):
structure, 203; On three feetal
Sperm-Whales, exhibited, 221;
Lantern-Exhibition of structural
characters of the Felide, 221; On
heads of Waterbuck, exhibited,
222; On Marine Boring Animals,
exhibited, 222; On a Chimpanzee
kept in the open air, lantern-slides
exhibition, 223; Photographs of a
living Okapi, exhibited, 225, 225 ;
On British Rats and their varieties,
exhibited, 224.

Melanichneumon carinifer, sp.n.,

148.

—— glaucopterus, sp. n., 149.

—- melanopterus, sp. n., 148.

Miojoppa quadrilineola, sp. n.,
167.

Mouuvsca:

Photographs and lantern-slides of
Marine Zoology, 221.

Monopeltis capensis (Z. s. L.), 225.
MorpnoLoay. See Srrucrure.

Neotypus obscurator, sp. n., 168,
Nestor notabilis (z. s. u.), 219.
Nicoria trijuga (z. s. u.), 221.

Otaria californiana (z. s. L.), 224.
Ovis musimon (z. s. u ), 224.

Paratilapia lukuge, sp. n., 18.
PaTHOLOGY.
Report on deaths of animals in the
Gardens in 1918, 13.
Pionus fuscus (z. 8. u.), 224,

INDEX.

PIscEs :
On a collection of fishes from Lake
Tanganyika: structure, 17.
Platylabus atricinctus, sp. n.,
168.
mediorufus, sp. n., 169.
Poceocephalus meyeri (z. s. L.), 222

el al sl

Proctoporus bogotensis, sp. n., 80.

Ratufa maxima (z. s. L.), 222.
ReEpTILta :

On the skull and affinities of Rana
subsigiliata : De-
scriptions of two new Lizards and
a new Frog from the Andes of

structure systematic,

structure, 21;

Colombia :
mo:

Serpentarius serpentarius (z. Ss. L.), 225.

Spilichneumon didymatus,
sp. n., 161.

-—— triangulator, sp. n., 162.

——- unipunctor, sp. n., 162.

Stenichneumon ochraceator,
sp. n., 152.

STRUCTURE.

Mamnatra: On

Mammalia, 71; Some points in the
Anatomy of the Takin (Budorcas
taxicolor whitet), 208.

Reprint: On the skull and affinities
of Rana subsigillata, 21; Deserip-

_ tions of two new Lizards and a
from the Andes of

new Frog
Colombia, 79.

the external cha- |
racters of existing Chevrotains, 1; _
On the breeding of Oryx gazella |
at Gooilust, 29; The Progressive |
Reduction of the Jugal in the |

X111

Pisces: Ona collection of fishes from
Lake Tanganyika, 17.

Insecta: A comparative study of
certain sense-organs in the antennse
and palpi of Diptera, 31; Some
points in Insect mechanics, 111;
On some Equatorial and other
Species and Genera of African
Ichneumonine contained in the
collection of the British Museum,
117.

Aves: A unique case of Asymme-
trical Duplicity in the Chick, 83;
Results of a Mendelian experiment
on Fowls, including the production
of a Pile Breed, 173.

Sus scrofa (z. s. u.), 222

aos

Testudo elegans (z. s. u.), 221.

Trogus gryps, sp. n., 122.

Ursus arctos (z. 8. u.), 224.

Xanthojoppa areolator, sp. n.,
1338.

——— bipapillator, sp. n., 182.

—— collifer, sp. n., 132.

—— cothurnator, sp. n., 132.

—— debilitor, sp. n., 130.

——— explanator, sp. n., 1381.

—— geminator, sp. n., 133. ©

—— gracilator, sp. n., 1382.

-—— rotundator, sp. n., 130.

——- striator, sp. n., 132.

—— truncator, sp. n., 131.

Xenojoppa fossifrons, sp. n., 163.

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

PROCEEDINGS

OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

AQOQLOGICAL SOCIETY

OF LONDON,

1919, pp. 227-499,

witH 13 PuLArEs and 64 TEx-FIGURES.

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

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

OF THE

COUNCIL AND OFFICERS

OF THE

ZOOLOGICAL

SOCIETY OF LONDON.

ISI).

Patron.
His Masesty THe Kina.

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

Tue Hon. Cectt Barine, M.A.

THE Lorp CarmIcHAEL,G.C.S.L.,
K.C.M.G.

Lr.-Con. 8. Monckton Copr-
MAN, M.D., F.R.S.

Cuartes DrumMonp, Esq.,
Treasurer.
ALFRED HzRA,
President.

Capt. Haroxp 8. FERGUSON.

Carr. Hucu 8. GLADSTONE,
M.A.

Aurrep H. Cocks, Esq., M.A.

Pror. JAMES P. Hrut, D.S8c.,
F.R.S., Vice-President.

WitiiaAm Huntsman, Esq.

E. G. B. Mreaps-Watpo, Esq.

Esq., Vice-

Pror. Ernest W. McBripe,
D.Sce., F.R.S., Vice-President.

Masor ALBERY Pam.

P. CHaumers Mircoety, Ksq.,
C.B.E., M.A., D.Se., LL.D.,
E.R.S. , Secretary.

ADRIAN dD. W. Poutocg, DAR.

Tue Lorp QUEENBOROVGH.

THe Marquis or Srico, F.S.A.,
Vice-President.

Masor Ricuarp 8. Tayror.

AntHony H. WINGFIELD, Esq.,
Vice-President.

Arriur® Smira Woopwarp,
Esq., LL.D., F.R.S., Vice-
President.

PRINCIPAL OFFICERS.
P. Cuatmers MircuHey, C.B.M., M.A., D.Sc., LL.D., F.B.S.,

Secretary.

R. I. Pococn, F.R.S8., F.L.8., Curator of Mammals and
Resident Superintendent of the Gardens.

D. Sera-Suiru, Curator of Birds and Inspector of Works.

Lieut. Epwarp G. BouLencer, Curator of Reptiles.

Prof, H. Maxwety Lerroy, Curator of Insects.

F. Martin Duncan, F.R.M.S., Librarian.

Joun Barrow, Accountant.
W. H. Coin, Chief Clerk.

LIST OF CONTENTS.

1919, pp. 227-499.

EXHIBITIONS AND NOTICES.

The Secretary. Report on the Additions to the Society’s
Menagerie during the months of June, July, August,
AMG Sep RETIN be OND ioe. a. tees. acu cetera ee letieisisje sesreleins oe 463

Mr. Oupriztp Tuomas, F.R.8. Exhibition of three inter-
esting Mammals obtained by Dr. Aders, F.Z.S8., in
ATTN DET Aes ne NR SPY Salt) ease ya eydbseesle 464

Dr. A. SmiraH Woopwarp, F.R.S., and Prof. G. Exuior
Smits, F.R.S. Discussion on The Zoological Position
and Affinities of Tarsiws. (Text-figure 1.) ............ 465

Prof. J. P. Hitt, F.R.S. The Affinities of Zursius from
the Kmbryological Aspect. (Table, Plate I., and
[RRA LTICATICER TIERS) Rice MRR nieeE eh dened matadace-oceeeaposcdmor 476

Prof. F. Woop-Jonrs, D.Sc., M.B., F.Z.S. The General
IAMaCOMhyA Olt LAT SUS. 5. calm aus materi scence rice mare ater 491

Mr. R. I. Pocock, F.R.S., F.Z.8. Structure of Zarsius ... 494

Mr. J. T. Cunnincuam, M.A., F.Z.S. Development of
SLCER TINS WERE SOREN eA ob en oo bbe eh avouoseeeee 495

Dr. P. Cuatmurs Mircuett, F.R.S. Characters of Zarsius. 496

Prof. E. W. MacBrips, F.R.S., F.Z.8., Vice-President of
the Society. Summing up the Discussion on Zarsius. 497

Mr. F. Martin Duncan, F.R.M.S., F.Z.S. Exhibition of,
and remarks upon, a series of photographs showing
the actinic quality of the light from a living Pyro-
OOPS CBOE jsacnoengdnpsosooyogoonb de saubsoaovanasascososdD

Mr. EK. Heron-Auen, F.R.S., F.Z.8. Exhibition and de-
scription of a series of Skiagraphs of the Foramini-
ITEM OY FEXSTONOIS WGPNQOIUOER Goocncoac scone desc edenccassseecescos

Tur Secretary. Report on the Additions to the Society’s
Menagerie during the month of October, 1919

eescee ue

Sir Epmunp Gites Lopmr, Bt., F.Z.S. Exhibition of, and
remarks upon, a series of skulls of the Beaver.........

Page

498

499

14,

15.

16.

lie

18.

iS),

bo
bo

PAPERS.

Report on Methods of Rat Destruction. By E. G.
BoutencerR, F.Z.8., Curator of Reptiles, Zoological
Society of London. With an Introduction by P.
CyAumers Mircnett, C.B.E., F.R.8.. LL.D., D.Sc.,
Secretary to the Society ............ dtd sakieiqunseetteteaed:

On certain Features of the Otic Region of the Chon-
drocranium of Lepidosteus, and Comparison with
other Fishes and Higher Vertebrates. By Epwarp
PERE TE S PACHIS EE Aue, sync iain aeratlie mate wtee oe eae

A List of the Snakes of West Africa, from Mauritania
to the French Congo. By G. A. Bounencmr, F.R.S.,
HAS: (Vextttgurés: 1 ds 2.)) 5)... 2e0 Ny) es ee

A List of the Snakes of North Africa. By G. A. Bov-
TAIN GME, HHA EUS), (HANS steer te sisecss § argo a cosas se Aaa Baa Re:

A Description of New Species of Zeuglodont and
of Leathery Turtle from the Eocene of Southern
Nigeria. By C. W. AnpreEws, D.Sc., F.R.S. (British
Museum, Natural History). (Plates I. & II. and
Mx MOUEES lO. )- o canticesuguee voce ssececeh alas ute naepeeneae

On the Occurrence of Denticles on the Snout of Xiphias
gladius. By J. THornton Carter (Hon. Research
Assistant, Department of Zoology, University of
London, University College). (Plates I-III.) ......

. Crustacea from the Falkland Islands collected by Mr.

Rupert Vallentin, F.Z.S.—Part III. By the Rev.
Tuomas R. R. Steppine, M.A., F.R.S., F.L.S., F.Z.8.
(Plates IV. and Vext-figures1—8,) 210... . 0.2.10 -2-

. Field-Notes on some Mammals in the Bahr-el-Gebel,

Southern Sudan. By Major J. Stevenson Hamitron,
CaMEZES.. CC With, Chatts\in sett 125, 520) ccnsassn on serecs

. Descriptions of a new Snake from the Transvaal,

together with a new Diagnosis and Key of the Genus
Xenocalamus, and ef some Batrachia from Mada-
gascar. By The Hon. Paun A. Meruuen, M.A.,
EAS (ext tenure: Loh eevee tate. tcl st.coee asst cere

Page

227

245

309

321

341

93. On the Variation in the Number of Dorsal Scale-rows
in our British Snakes. By Miss Joan B. Procter,
BeEZSs a Lext-@uUres M3.) cosine cease or eeeecee wcntieer 357

24. On the Species of Balaninus occurring in Borneo
(Coleoptera, Curculionide). By Guy A. K. Mar-
SHAG DISe., UZ Sun (Plates mi ner lille) eae re erer eeeaer 365

25. On some new Fishes from near the West Coast of Lake
Tanganyika. By G. A. BouLengsr, F.R.S., F.Z.8.
(Lext-fig av esi6=10.) oc. secon cushions oe see eee eee eee 399

26. The Radula of the Mitride. By the Rev. A. H. Cooxs,
SiGe Das Legere (Uesxtyseetontes TIS),)) sacecocganoesbododoos 405

bo
“I

. Note on the Righting Reaction in Asterina gibbosa
Penn. By E.S. Russeny, M.A., B.Sc., F.Z.8. ...... 423

28. Experiments on Sex Determination. By Lt.-Col. 8.
Monckton Copeman, M.D., F.R.S., F.L.S. ............ 433

29, The Variations in the Digastric Muscle of the Rhesus
Macaque and the Common Macaque. By Cuas. F.
Sonntac, M.D., Ch.B., F.Z.S., Anatomist to the
Soctebyan (Mextatioures eco) eeme se eres been ene reee eee 437

30. On the Nematode Parasites of a Chapman’s Zebra. By
Miss M. Turner, B.Sc. (Text-figures 1-6.)............ 44]

31. The Development of the Mesenteries in the Actinian
Urticina crassicornis. By James F. Grmuit, M.A..,
IM. D:, DSc. (Rext=gures los) eee emeees ee eee eee 453

32. The Ciliation of the Leptomedusan JMelicertidiwm octo-
costatum (Sars). By James F. Gremuiuy, M.A., M.D.,
DiSe. si(Vext-toures6:) 2) 51). peewee eareeeene ce ctcce 459

Alphabetical Mistvol Contributors o--aeeeeeer eee es eeeee eres vil

Mirae) © ain dies rhe odidiasd Saco d eee aa sla a eee x1

ALPHABETICAL LIST
CONTRIBUTORS,

With References to the several Articles contributed by each.

1919, pp. 227-499.
pp

° Page

Aus, Epwarp Puetps, Jr., F.Z.S8.

On certain Features of the Otic Region of the Chondro-
cranium of Lepidosteus, and Comparison with other Fishes

andyLncher: Viertebratesie..8 .. abt. .0 ai cee as eceean adh cere 245
Anprews, C. W., D.Sc., F.R.S.

A Description of New Species of Zeuglodont and of
Leathery Turtle from the Eocene of Southern Nigeria.
(Plates I. & Il. and Text-figures 1-3.) 22... ...i...s.0.0005 309

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

Report on Methods of Rat Destruction. With an
Introduction by P. CHatmers Mircuety, C-B.E., F.B.S.,
Mi. DiSe:, Secretary to the Society |... /6.2.5........... 227

Bovuuencer, G. A., F.R.S., F.Z.8.

A List of the Snakes of West Africa, from Mauritania
to the French Congo. (Text-figures 1 & 2.) ............... 267

A List of the Snakes of North Africa..................... 299

On some new Fishes from near the West Coast of Lake

399

Hancanyika. (Lext-fisures GoM) jos. eace ssn

vill
Page
Carter, J. THORNTON.

On the Occurrence of Denticles on the Snout of Xiphias
gladius. (Plates 1.—QIL.) ........-....0.02.0-- 2000+ 0-0 =e ecae 321

Cooxg, Rev. A. H., Sc.D., F.Z.S.
The Radula of the Mitride. (Text-figures 1-18.) ..... 405

Copeman, Lt.-Col. S. Moncxron, M.D., F.R.S., F.Z.S.

Experiments on Sex Determination.....................-.- 433

Cunnineuam, J. T., M.A., F.Z.8.
Development of Warscismire reece eaee een ee Eee eee eere 495

Dunoan, F. Martin, F.R.M.S., F.Z.S.

Exhibition of, and remarks upon, a series of photo-
graphs showing the actinic quality of the light from a
livaime Panophenus waeecley. sat aa-e-res ee: pew eee eer ee rece cee 498

GeuuiuL, JAMES F., M.A., M.D., D.Sc.

The Development of the Mesenteries in the Actinian
Urticina crassicornis. (Text-figures 1-5.) .................. 453

The Ciliation of the Leptomedusan JMelicertadium
octocostatum (Sars). (Text-figure 6.) ...............-....00+ 459

Hamitron, Major J. Stevenson, C.M.Z.8.

Field-Notes on some Mammals in the Bahr-el-Gebel,
Southern Sudan. (With Chart.).............:.-:::0 cress 341

Heron-Auten, E., F.R.S., F.Z.S.

Exhibition and description of a series of Skiagraphs of
the Foraminiferan genus Vernewiline ........-..+-2000 eee 499

Hitt, Prof. J. P., F.R.S.

The Affinities of Tarsius from the Embryological
Aspect. (Table, Plate I., and Text-figures 1=5,) Coie. ak 476

15
Page

t=)

Lover, Sir EpMunp Gites, Bt., F.Z.S.

Exhibition of, and remarks upon, a series of skulls of
NS BSA CTPA age sl ua OG ORC e aE OER AEE eee 499

MacBrips, Prof. KE. W., F.R.S., F.Z.8., Vice-President of
the Society.

Summing up the Discussion on Varsius ............00.0.. 497

MarsHau, Guy A. K., D.Se., F.Z.S.

On the Species of Balaninus occurring in Borneo
(Coleoptera, Curculionide). (Plates I. & II.) ............ 365

Metuuen, The Hon. Paut A., M.A., F.Z.S.

Description of a new Snake from the Transvaal,
together with a new Diagnosis and Key of the Genus
Xenocalamus, and of some Batrachia from Madagascar.
(lexi fice We cine eitasse cise sopbode doonsoaioonmdeecebeusen 349

Mircuett, P. CHatmers, C.B.E., F.R.S., LL.D., D.Sc.,
Secretary to the Society.
Introduction to Report on Methods of Rat Destruction. 227
Report on the Additions to the Society’s Menagerie

during the months of June, July, August, and September,
RETR eee RRR tc ater ats afes'ais/rs'n's[o cels'eida's s/o de ateuna tarde Rae mete etm rae 463

ClOATAOUSIES Ot SIGHRSOUIS Sas nadhacconcnacosaendoacnd BO prt tr 496

Report on the Additions to the Society's Menagerie
durme thermonth of October; VOLS. a. 2-- tees: 499

Pocock, R. I., F.R.S., F.Z.8.
SELMChuURe Ok LUGSTUS .os.cesneees sateen eee Sak Ca, We ee 494

Procrer, Miss Joan B., F.Z.S.

On the Variation in the Number of Dorsal Scale-rows
in our British Snakes. (Text-figures 1-3.) ............... 357
Proc. Zoou. Soc.—1919. b

Page
RusseE.u, E. 8., M.A., B.Sc., F.Z.8S.
Note on the Righting Reaction in Asterina gibbosa
HERG MANAG gata tevaieietng ainie a emia sl tne are acleener aaieteke sales aa ere 423
Smirn, Prof. G. Etuiot, F.R.S8.
Discussion on the Zoological Position and ‘Affinities of
Marsius: \(Vext-feures) is veree aces ae eee et ee nee nena: 465

Sonnrac, Ouas. F., M.D., Ch.B., F.Z.S., Anatomist to the
Society.
The Variations in the Digastric Muscle of the Rhesus
Macaque and the Common Macaque. (Text-figures 1-5.) 437

SreBpBine, Rev. THomas R. R., M.A., F.R.S., F.L.S., F.Z.8.

Crustacea from the Falkland Islands collected by
Mr. Rupert Vallentin, F.L.S.—Part III. (Plates I.-IV.
EnOl Meanie Wexei)), Tannseobaoononocanabosdce oe eade ches sarees 327

THomas, OLDFIELD, F'.R.S.
Exhibition of three interesting Mammals obtained by
Dr. Aders, HiZ:Ss im Zamziloae oy avcrnnce case steer ceeeriae 464

Turner, Miss M., B.Sc.

On the Nematode Parasites of a Chapman’s Zebra.
(Rext-fioures 1 =6.) 5.0.0.0. -sccaueeemn een area eicert saree 44]

Woonv-Jonss, Prof. F., D.Sc., M.B., F.Z.8.
The General Anatomy of Tarsius............0..000ee seco 491

Woopwarp, Dr. A. Smiru, F.R.S.

Discussion on the Zoological Position and Affinities of
Varsius: “(Lext-figure 1.) 0........ceecaeaeere ae ee ee ee 465

INDEX.

1919.—Pages 227-499.

[New names in clarendon type. Systematic references in italics,
(z.8.L.) indicates additions to the Society’s Menagerie. |

ActTINIZ: Balaninus gyrosicollis, sp. n., 378,
On the development of the Mesen- | —— imitator, sp. n., 388.
teries in Urticina crassicornis: | —— longiclavis, sp. n., 594.
structure, 453; On the ciliation of | ———- moestus, sp.n., 381.
the Leptomedusan Melicertidium | ——- nigrocinereus, sp. n., 378.
octocostatum (Sars): structure, | —— nigrorufus, sp. n., 380.
459. —— pusio, sp.n., 391.

flurus fulgens (z. 8. u.), 463.
Ageleus flavus (z.8.L.), 463.
Allabenchelys dhonti, sp. n., 402.

—— quincunx, sp. n., 386.
— rufulus, sp. n., 377.
—— sellatus, sp. n., 395.

Anabas ctenotis, sp. n., 403.
Antilope cervicapra (z. 8. u.), 499.

Balaninus analis, sp.n., 387.
—— bilineatus, sp. n., 375.
—— bispilotus, sp. n., 371.
—— bryanti, gp. n., 369.

—— commodus, sp. n., 390.
—— consocius, sp. n., 354.
—— decemnotatus, sp. n., 389.
—— deceptor, sp. n., 389.
—— delicatulus, sp. n., 392.
—— discreticoxis, sp. n., 396.
—— eugenie, sp. n., 393.
—— excavatus, sp. n., 373.
—— excisipes, sp. n., 373.
——— glabricollis, sp. n., 370.
—— grypus, sp. u., 390.

—— semisuturellus, sp. n., 376.
—— sesquilineatus, sp. n., 574.
—— shelfordi, sp. n., 390.

—— subpartitus, sp. n., 380. ©
—— trinotatus, sp. n., 383.

— tumidirostris, sp. n., 382.
-— unifasciatus, sp. n., 384.
Barbus euchilus, sp. n., 400.
micchilus, sp. n., 401.

Bos grunniens (Z. 8. L.), 468.

Boselaphus tragocamelus (z. 8. u.), 499.

Cephalophus adersi, 464.

Cercocebus aterrimus (z. 8. u.), 464.

Cercopithecus erythrogaster (z. 8. L.),
464,

Colobus kirki, 464.

| Comatibis eremita (z. 8, L.), 464,

Xil

Connochsetes albojubatus (z. 8. u.), 463.
Cosmochelys dolloi, gen. et sp. nov.,
314,
Crotalus atrox (z. 8. L.), 464.
CRUSTACEA:
On Orustacea from the
Islands: systematic, 327.
Cylichnostomum zebre, sp. n.,
445.
Cynelurus jubatus (z. 8. L.), 463.

Falkland

EcuHInoDBRMATA :

Note on the Righting Reaction in
Asterina gibbosa Penn.: structure,
423.

Elephas maximus (2. 8. L.), 464.
EMBRYOLOGY:

The affinities of Zarsius from the

Embryological aspect, 476.

Felis leo (z. s. u.), 499.
pardus (z.s.L.), 464, 499.
FORAMINIFERA +
Exhibition of a series of Skiagraphs
of the Foraminiferan genus Ver-
neuilina, 499.
Francolinus granti (z. 8. u.), 463.

GEOGRAPHICAL:

A list of the Snakes of West Africa, |

267; A list of the Snakes of North
Africa, 299; On a new species of
Zeuglodont and of Leathery Turtle
from the Hocene of Southern
Nigeria, 309 ; On Crustacea from
the Falkland Islands, 327; Field-
notes on some Mammals in the

Southern Sudan, 341; On a new |

Snake, and Batrachia from Mada-
gascar, 349; On the species of
Balaninus occurring in Borneo,
365; On new Fishes from the

West Coast of Lake ‘langanyika, |

399.
Gephyromantis, gen. n., 351.
—— boulengeri, sp. n., 351.

Hippopotamus amphibius (z. 8, u.), 463.
Hyena hyena (z. 8. u.), 499.

|
(

INDEX.

INSECTA :

On the species of Balaninus occur-
ring in Borneo: structure: syste-
matic, 365; Exhibition of a series
of photographs showing the actinic
quality of the light from a living
Pyrophorus Beetle, 498.

Labeo dhonti, sp. n., 399.

MamMatta :
Report on Methods of Rat destruc-

tion: descriptive, 227; Field-
notes on some Mammals in
the Southern Sudan: structure,

341; Experiments on Sex Deter-
mination, 433; On the Digastric
Muscle of the Macaques : structure,
437; Discussion on the Zoological
Position and Affinities of Tarsius :
structure, 465, 491; Exhibition
and remarks on a series of skulls
of the Beaver, 499.
Mantidactylus argenteus, sp. n.,
3538.
Mo.tusca:
The Radula of the Mitride: struc-
ture, 405.
Morpnotoey. See Srructrure.
Musophaga rosse (z. s. L.), 463.

Nectarinia kilimensis (z. s. u.), 464.
Oyibos moschatus (z. s.t.), 499.

Pappocetus lugardi, gen. et sp.
noy., 309.
Paradisea minor (z. s. L.), 499.
PATHOLOGY:
Tixperiments on Sex Determination,
433; On the Nematode Parasites
of a Chapman’s Zebra, 441.
Petrodromus, 464.

| Phractura lukuge, sp. n., 402.

Phrynosoma cornutum (z.s. L.), 464.
Pituophis sayi (z. 8. u.), 464.
PISCcEs :
On certain features of the otic region
of the chondrocranium of Lepi-

INDEX.

dosteus: structure, 245; On the
occurrence of Denticles on the
Snout of Aphias gladius: struc-
ture, 321; On new Fishes from the
West Coast of Lake Tanganyika:
structure, 399.

Plethodontohyla tuberifera,

sp. n., 304.
Pseudoleistes virescens (z. 8. L.), 463.
Pterorhinus davidi (z. 8. u.), 463.

ReEpriqia :

A list of the Snakes of West Africa :
systematic, 267; A list of the
Snakes of North Africa: syste-
matic, 299; On a new species of
Zeuglodont and of Leathery Turtle:
structure, 809; On a new Snake,
and Batrachia from Madagascar:
structure, 349; On the number of
dorsal scale-rows in British Snakes:
structure, 357.

Rusa unicolor (z. s. t.), 499.

Xill

of Lepidosteus, 245; On the oceur-
rence of Denticles on the Snout
of Xiphias gladius, 321; On new
fishes from the West Coast of Lake
Tanganyika, 399.

Reptinia: On a new species of
Zeuglodont and of Leathery
Turtle, 309; On a new Snake, and
Batrachia from Madagascar, 349;
On the number of dorsal Scale-
rows in British Snakes, 357.

Inszcra: On the species of Balaninus
occurring in Borneo, 365.

Mortusca: The Radula of the
Mitride, 405.

EcrNoprrmara: Note on the Right-
ing Reaction in Asterina gibbosa
Penn., 423.

Actinta : On the development of the
Mesenteries in Urticina crassicornis,
453; On the ciliation of the Lepto-
medusan Melicertidium octocos-
tatum (Sars), 459.

StrRucTURE. Tanais nierstraszi, sp. n., 332.
Mamata: Field-notes on some | Tarsius, 465.
Mammals in the Southern Sudan, | Testudo elephantina (z.3..), 464.
341; On the Digastric Muscle of | Trachymantis, gen. n., 352.

the Macaques, 437; Discussion on

the Zoological Position and Affini- | Vinago calva (z.s. L.), 463.

ties of Tarsius, 465, 491; Exhibition

and remarks on a series of skulls | Kenocalamus transvaalensis,

of the Beaver, 499.
Pisces: On certain features of the

sp. n., 300,

otic region of the chondrocranium | Zamenis flagelliformis (z. 8. u.), 464.

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LES fOr vCON TNs,

1919, Parts I. & IL. (pp. 1-225).

EXHIBITIONS AND NOTICES.

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

of November and December ..............5 HO mS Ogee OCA pO Go.o Doles eo ac

Mr. C. Davis Sumrzory, F.Z.8. Exhibition of, and remarks upon, aletter written in 1693

by Malproaictio di Mathew. Haber es: ict tise! s oe tale re ala ele. to ens tala caries olittet eta arene ee

Sir Dovetas Mawson. Lantern exhibition of, and remarks upon, Australasian Antarctic

BNC SUPA MGATSEIC TLS 2-0. fw mac oe era leeat Mel Ae) =I) ge a ole a) dua lacw oseeala) < Seaape ee es ete a aed

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

Of@antanye TOU!” us Sake ogee ena acl aha Sg ee aac

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

Coty! Ei eV oNTD Fe ea prod AO) 8 fairey ce aan Doe i rma ene RAM cet AO CA a were ene cd

Mr. F. Martin Duncan, F.R.M.S. Exhibition of, and remarks upon, a series of photo-
graphs and lantern-slides of Marine Zoology

Dr. F. EH. Bepparp, F.R.S. Exhibition of, and remarks upon, the foetal Sperm-Whales ..

Mr. R. I. Pococs, F.R.S., F.Z.8S., Curator of Mammals, Exhibition, illustrated by lantern-
slides, to show structural characters of the Helidz

Ce ]

Tue Secretary. Report on the Additions to the Society’s Menagerie during the month of
March, 1919

Ce cr CY

Mr. T. Gzrrarp, F.Z.S. Exhibition of, and remarks upon, a series of heads of Waterbuck

Dr. W. T. Cauman, F.Z.8. Exhibition and detailed account of yarious Marine Boring
ANTIMNAlS e's. oie DOPE oC rem aN OE ria cidade sant cnn cidly Mcna a atic, © De eeiars

Tae Secretary. Exhibition, on behalf of Mr. George Jennison, of a series of lantern-
slides of a Chimpanzee kept in the open air

Tun Sucrutary. Exhibition of, and remarks upon, two photographs of a living Okapi ..
Mr. EK. G. Bounrnesr, F.Z.8. Exhibition of, and remarks upon, British Rats and their
WDE Hs Caen GMOS lb eee MEE OT coh Otcls GORA eC EO LORE eA Sao ome

Lt.-Col. 8. Monckton Coreman, F.R.S. Exhibition of a series of lantern-slides to illus-

trate his.” xperiments on) Sex etermimationiy meiner ieieetecaraiter es ete

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

of April, 1919 --..... atin) all esa'se Va aan anhaPat a 9 gat SWI ae OMT Le Bs See clade wens ee

Tne Secrerary. Exhibition of, and remarks upon, the photographs of the Young living

Okapi shown at the previous Scientific Meeting ..........25 ..ce+ee.0.c.+ se ueue

The Secrwrary, Report on the Additions to the Society’s Menagerie during the month
of May, 1919

Page

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PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF TIE

ZOOLOGICAL SOCIETY OF LONDON

PAPERS.

1, On the External Characters of existing Chevrotains.

By R. I. Pocoox, F.R.S., F.Z.S.
[Received February 4, 1919: Read February 18, 1919.}
(Text-figures 1-5.)

CONTENTS.

20
tic}

Lev]

Genrenrcenain esweene a rectencc det 2 ctiaeiric fasinat aya eas
@olovationweea aie ei mote sc cleiee ote tian curetneon se:
IRIAOEE NTN > edo ecton Soe ean OR ROE Nema oe Nae e eee aoe eee
JE YGRE | yl OSIRIS, 5 edhe er eis a al oe eo
Interramal gland

TERIA | AVE sbndoousdincs onaboonenees
Teanlandysubeaudall area. s.00/)..00..ss.ace.- cece. tes
eR SIM Meee iat MEE NY. ch ols caine tanits oath a eae ARE
EMITS PALE erase coed ened fave ees

Incidence of generic characters ...............6..0000
Wiaenoses ot thelgenetay. ..4.. cece once sseuel ce acesen noes

COON NT EP FB Bw De

Generic names. —Gray (Cat. Rum. Mamm. Brit. Mus. pp. 97-
99, 1872) referred the Chevrotains to two families: the Tragulide
for the Oriental species, and the Hyemoschide for the single
West African form. Furthermore he admitted two genera of
Tragulide, namely Meminna or, as it should have been, Memina
according to his original spelling of the name in 1821 (Med.
Repos. xv. p. 307) for the spotted Indian and Ceylonese species
meminna, and T'ragulus- for the unspotted species occurring to
the east of the Bay of Bengal.

The family Hyemoschidz does not appear to have been adopted
by later writers, and the genus constituting it has been usually

Proc, Zoou. Soc.—1919, No. I. l

2 MR. R. I, POCOCK ON THE

cited as Dorcatheriwm. Also no oneseems to have followed Gray
in granting generic status to meminna (cf. Lydekker, Cat. Ung.
Mamm. iv. pp. 261-298, 1915). But in 1916 Thomas (Ann.
Mag. Nat. Hist. (8) xviii. pp. 72-73) gave subgeneric rank to
meminuna, adopting for the species the name J/oschiola Hodgson,
as quoted by Gray, on the grounds of the preoccupation of
Memina or Meminna by G. Fischer in 1814. Furthermore he
showed, in opposition to Merriam’s opinion, that 7ragulus is
typified by one of the larger spotless Oriental species, and that
Hyemoschus should stand for the West African form, which may
be generically distinguished from the now extinct European
Dorcatheriwm.

In the following pages I adopt Thomas’s nomenclature but,
for what appear to me to be good and sufficient reasons, I give
full generic rank to Moschiola.

The materials on which this paper is based were fresh examples
of the following species: Hyemoschus aquaticus, Moschiola me-
minna, Tragulus javanicus, T'. kanchil, and 7’. stanleyanus.

Coloration.—As is well known, Hyemoschus and Moschiola difter
from Zragulus in retaining on the body a distinct pattern of
white spots which on the flanks fuse, or tend to fuse, into longi-
tudinal stripes and on the croup frequently exhibit a transverse
arrangement. The pattern recalls that of the Tragelaphine
Pecora; and in this connection it may be noted that Hyemoschus
is white on the front of the pasterns as in typically marked
éxamples of those antelopes.

There are also some interesting points connected with the pecu-
liar pattern of the neck and throat in these primitive Ruminants.
In Moschiola there is a continuous white median stripe of
considerable width running from the chin on to the abdomen.
From a point a little behind the corner of the mouth this white
median stripe gives off on each side a single white lateral stripe
which runs along the side or lower edge of the mandible and
ceases approximately on a level with the point of attachment of
the posterior edge of the ear. From its position this may be
called the mandibular stripe. Towards the middle of the throat
the median white stripe branches again, giving off on each side
a second, or jugular stripe which passes obliquely backwards for
a short distance in the direction of the shoulders, but is much
shorter than the mandibular stripe. Thus in WMoschiola the
mandibular stripe is comparatively long and the jugular stripe
short, and both arise from an uninterrupted median white band
traversing the throat from end to end and passing on to the chest,
with only a slight extension of pigmented hairs inwards towards
the middle line where the throat passes into the chest.

In Hyemoschus the same general arrangement prevails, but
both the mandibular and jugular stripes are much longer, the
former passing beyond the level of the ear well back on to the side
of the neck, and the latter reaching almost to the shoulder.
‘Moreover, there is no encroachment of pigmented hairs where

EXTERNAL CHARACTERS OF CHEYVROTAINS, 3

the white of the throat runs into that of the chest. Tragulus
differs both from Hyemoschus and from Moschiola in having the
mandibular stripe undeveloped or very short. ‘The jugular stripe,
however, is as long as in //yemoschus, but the dark band which
separates it from the median white band on each side is broader
posteriorly and is continued farther forwards and frequently
meets its fellow of the opposite side a little behind the line of
the angles of the jaw, thus dividing the white of the interramal
area from that of the throat. Moreover, the white area of the
throat, which is much broader behind than in front, is separated
from the white of the chest by a dark transverse band.

Text-figure 1.

A. Rhinarium of Hyemoschus aquaticus, from above.

B. 5 Moschiola meminna, from the side.

C. External ear of Hyemoschus aquaticus. t, tragus; 1, lappets overlying
the depression.

D. The same, with the external rim of the capsule cut open behind the
tragus, showng the auditory orifice. ¢, tragus turned forwards ;
1, lappets ; +, supporting ridges ; a, cut edge.

The Rhinariwm in the three genera is large, naked, and sculp-
tured by grooves into larger and smaller areas. Its dorsal surface
extends backwards some distance behind the posterior angle of
the nostrils, which are comparatively narrow and slit-like, but
not more valvular than in the Cervide. The area of the rhina-
rium beneath them is deep laterally and wide in front, wider
than the internarial septum and as wide as the anterior rim of
the lower lip (text-fig. 1, A, B). .

4 MR. R. I. POCOCK ON THE

The Facial vibrisse are generally well developed, consisting of
mystacial, submental, superciliary, subocular, and genal bristles,
two pairs cf genal tufts being frequently present. The superior
tuft is situated either high up about halfway between the eye
and the ear but at a lower level, as in an example of Tragulus
stanleyanus, ov below the corner of the eye, as in a specimen of
T. kanchil. The inferior tuft, represented by a single bristle,
arises some distance behind the corner of the mouth. “Both tufts,
however, are not always present. When they are present, the
full complement of vibrisse coincides with that of some small
Cervide and Bovide and, as I have elsewhere remarked, with -
that of typical Carnivora. The submental vibrisse are arranged
in definite lines. The interramal tuft, which is of special interest,
is described in the following paragyra aph.

The Interramal aerndeeston Moschiola (text-fig. 2, B) the chin
and interramal area are covered with hairs which are short on the
chin and gradualiy become longer on the throat, without showing
any sharp line of demarcation. The interramal tuft of vibrissve,
consisting of two pairs of bristles, sometimes with an additional
median one, is set in thé posterior half of the interramal area. In
Hyemoschus (text-fig. 2, A) the chin is scantily covered with fine
short hairs sharply defined from those of the throat, which are
longer than in Moschiola. The interramal vibrisse, consisting of
two or three pairs of bristles and one unpaired, making five or
seven in all, form a cluster as in Moschiola. In Tragulus (text-
fig. 2, C, D) the chin is even more scantily hairy, apart from the
submental vibrissee, than in Hyemoschus, but it is not naked as
described originally by Gray and recently by Lydekker in his
Catalogue, although it appears to be naked when examined by the
naked eye. Behind the chin there is a tongue-shaped area of
skin, covered, like the chin, with short hairs and extending back-
wards along the middle of the interramal area as far as the
interramal tuft of vibrisse which is set at its posterior edge.
This tract of skin overlies a cutaneous gland which is sometimes
so thick posteriorly that in profile view it forms a swelling
projecting well below the inferior edges of the mandibular rami.
The presence of this gland, which has been noticed by previous
authors, serves to distinguish Zragulus from Moschiola and
HHyemoschas.

In an adult male of 7. stanleyanus (text-fig. 2, D) there was
only a single pair of interramal vibrisse arising near the posterior
edge of this gland. In one example of 7. kanchil there were
five vibrisse arranged in a transverse row. In another example
of the same species there were three pairs of these vibrisse, and
in one example of 7’. javanicus the arrangement and number of
the vibrissee were the same as in the last-mentioned example
of T. kanchil. A second example of 7’. javanicus (text-fig. 2, C)
showed two pairs of vibrissze and one bristle in the centre of the
area circumscribed by them.

The external Har (text-fig. 1, C, D) is small in all cases, but

EXTERNAL CHARACTERS OF CHEVROTAINS. 5

noticeably larger relatively in JZyemoschus than in the Oriental
forms. It is attached to the head by a broad base, the narrow
elongated cup-shaped base of the typical Ruminantia, in which
the ears are highly mobile, being undeveloped. The tragus is

Text-figure 2.

!
HI j ya ii iN
i MW) aa A y
My fs Na
HR MN NN
Cc

\ S. SAT WN
RENAN
71\X\\ \ Sy

{ \ WA NN

Underside of head of Hyemoschus aquaticus (g immat.), showing tuft of
interramal vibrisse.

. The same of Moschioia meminna (& ad.).

- The same of Tragulus javanicus (g ad.), showing the interramal gland.

D. The same of Trragulus stanleyanus.

A.
B
Cc

merely a small excrescence, but it is better defined in Moschiola
than in Hyemoschus and Tragulus. Low down towards the
posterior edge of the pinna there is a pair of short strengthening
ridges, and in front of these towards the anterior rim of the

6 MR. R. I. POCOCK ON THE

pinna there is a definite pit or depression overlapped in front and
capable of being closed by two lobate thickenings, one above the
other, the upper being larger than the lower. The auditory
orifice, opening upon a thick ridge-like execrescence, is concealed
by the tragal ridge which forms the external border of the
capsule of the pinna.

Text-figure 3.

EARS

EA\\

AS \I\YE)

Nata iM
\

ZN NERA"
Yh, iy) i actos }
Na a!

A. Anal area of Moschiola meminna 6, showing the short tail and the hairy
scrotum.

B. The same of Tragulas javanicus , showing the long tail and the naked
scrotum.

C. The same of 7. javanicus 2, showing the naked area round and below
the vulva,

_ The ear of the Tragulide differs from that of the Pecora in
three particulars, namely, in the breadth of its basal attachment

EXTERNAL CHARACTERS OF CHEVROTAINS. 7

to the head, in the development of only two supporting ridges,
both placed near the posterior rim, and apparently in the pre-
sence of the depression overlapped by lobes in the position of the
supratragal ridge of the normal mammalian ear.

The Vail, as is well known, varies in length. In Moschiola
(text-fig. 3, A) it is very short. When depressed, its tip reaches
the scrotum * but does not conceal it, and the scrotum is covered
with hair. In. 7’ragulus (text-fig. 3, B, C) the tail is much longer
and covers the scrotum, which is naked except laterally at the
base. Similarly in the female the tail covers the genital orifice,
which is situated at the upper end of a large tract of naked skin.
In Hyemoschus the tail is at least as long as in Tragulus, but the
subeaudal area is not naked as in that genus but somewhat thinly
covered with fine white hairs.

The Legs of the Tragulide, as compared with those of the
Pecora, are remarkable for the shortness of the metacarpal area,
a primitive character recalling the condition seen in the Suide.
In Hyemoschus the legs and feet are relatively much stouter than
in 7ragulus and Moschiola, in which they are slender and delicate,
and in the African genus the interdigital integument forming
the floor of the interdigital depression reaches a little nearer to
the heels of the hoofs than in its Oriental allies. In the three
genera there is a smooth pad of moderately thickened, naked
skin on the posterior side of the hock (caleaneal area). This pad
is indistinctly defined in Tragulus, but is sharply defined in the
other genera.

In Hyemoschus (text-fig. 4,C, D) the metacarpal and meta-
tarsal areas are everywhere thickly covered with hairs. Those
on the back of the metacarpus grow downwards, whereas on the
back of the metatarsus they grow backwards on each side, forming
a median crest. ‘lhe back of the pasterns between the false hoofs
and the true hoofs is naked, except for a few hairs in the middle
line, and the walls and floor of the interdigital cleft in front are
also almost naked.

In Yragulus (text-fig. 4, B) the metacarpal and metatarsal
areas are everywhere comparatively scantily hairy, especially on
their posterior sides where the hairs are so short and sparse that
their surfaces have been described as naked. ‘The hairs cease
altogether some little distance above the false hoofs. Hence the
nakedness of the back of the pastern may be said to extend above
the false hoofs, a condition not observable in Hyemoschus. The
interdigital cleft in Zragulus is also naked to all intents and
purposes, as in Hyemoschus.

In Moschiola (text-fig. 4, A) the metacarpal and metatarsal
areas are as hairy as in Hyemoschus, but on the posterior side of
the metatarsus the hairs grow upwards from a line a little above

* 1t may be noted that the scrotum in these primitive Ruminants is sessile and
set high up on the perineal area just beneath the anus and is not pendulous between
the hind limbs as in the Pecora. Its position recalls that of the scrotum in the
Suidze and Camelidee

8 MR. R. I. POCOCK ON THE

the false hoofs where there is a definite parting, the hairs below
the line growing downwards. Moschiola, however, is distin-
guished both from Hyemoschus and Tragulus by having the
posterior sides of the pasterns and the interdigital clefts hairy
instead of almost naked.

Text-figure 4.

A. Posterior surface of hind leg of Moschiola meminna, the arrows showing
the direction of the hair-growth.

B. Posterior side of fore foot of Tragulus javanicus.

C. Posterior side of hind leg of Hyemoschus aquaticus.

D. Posterior side of fore foot of the same.

The Penis of an undetermined species of Zragulus was figured
and described by Léunberg (Nova Acta. R. Soc. Upsal. (3) xx.
p. 33, pl. 11. fig. 20, 1904). The penis of an example of 7’. stun-
leyanus (text-fig. 5, C, D) agrees tolerably closely with it except
that there is an additional coil on the spirally twisted terminal
portion, possibly due to its being more contracted. As in Linn-
berg’s specimen there is at the base of the spirally twisted
termination a well-developed lamina ending proximally in a free

EXTERNAL CHARACTERS OF CHEVROTAINS. 9

process, the two combining to give the incrassate appearance to
the distal end of the organ. Normally this lamina and _ its
process ave closely folded on the shaft of the penis, but they are
capable of being unfolded and spread (text-fig. 5, D).

Text-figure 5.

ALC ys

iy,

VS Yay ayes

A. Extremity of penis of Hyemoschus aquaticus, from the left side.
B. The same of Moschiola meminna.
C. The same of Tragulus stanleyanus, with apex coiled and lamina folded up.
D. The same, with apex partially uncoiled and lamina unfolded.
EH. The same of Tragulus kanchil folded up.
EF’. The same from the right side, with the apex and the lamina partially
unfolded.
Lettering :—d, distal, and p,’proximal portion of the lamina in the two
species of Tragulus.

The penis of Zragulus kanchil (text-fig. 5, EH, F) resembles in a
general way that of 7’. stanleyanus, but there are some well-

10 MR. R. I. POCOCK ON THE

marked differences. The spirally twisted portion is here and
there thickened and geniculate and the lamina is somewhat
differently shaped. When the penis is contracted the spirally
twisted portion and the lamina are tightly folded together into a
thickened knot.

The penis of Moschiolu. meminna was figured and described by
Gerhardt (Verh. Deutsch. Zool. Ges. xvi. p. 153, 1906). The
penis of an example I examined is in close agreement with it.
It ends in a long, many-coiled portion, with an acuminate tip
and there is no lamina (text-fig. 5, B).

In Hyemoschus (immature example) (text-fig. 5, A) I find the
penis to be in general agreement with that of JJoschiola, the
point being sharp and the lamina absent, but the spiral twists of
the terminal portion are not so numerous or so close, the twisting
resembling rather that of a ‘“ gimlet” than of a “corkscrew,” to
which the twisting seen in Moschiola may be compared.

The incidence of the characters above described may be epi-
tomised as follows :—

1. a. Body spotted and striped with white ............... Moschiola, Hyemoschus.
bs Bodysunspotted ] 4.8 ts oe eras Tragulus.
ec. Mandibular stripe moderately long, jugular stripe short... Moschiola.
d. Mandibular and jugular stripes both HOWE sasoos s05%0 ... Hyemoschus.
e. Mandibular ering short or absent, jugular stripe very
OT errata nee Ca ne rats i be Ee Ne une ie Oil IAS
2a a Thee end siieat Lalas NRE scare nae ‘Mosehiola, Hyemoschus.
6. Interramal gland present Sdesncec athens cencsncee eye tem cg elatss
3. a. Tail short, scrotum and inguinal region hainy Pe eee en oschioleas
6. Tail long, scrotum and inguinal region hairy .................. _Hyemoschus.
ec. Tail long, scrotum and in: suinal region TAK CMa eerie cee Tragulus.
4. a. Legs and feet stout and strong ............ 00000 ... Hyemoschus
b. Legs and feet fineand slender... Hee done | Moschiola, Tragulus.
ce. Legs and feet normally covered with hair ‘(apart from the
hock-pad) _... .... Moschiola.
d. Legs normally hairy, back of ‘pasterr ns and interdigital ‘cleft
almost naked . ... Hyemoschus.
e. Less very scantily hairy below knees and hocks ‘behind,
back of pasterns and interdigital cleft almost naked ...... Tragulus.

5. a. Penis simple, attenuated, spirally coiled distally, without

lamina ........ ... Hyemoschus, Moschiola.
b. Penis complex, ‘spirally ‘coiled distally but ‘provided with a

lamina normally folded up behind and beneath the twisted

Lerinmation ys. san heise Moesa de ohak esis ote Re ee Tragulus.

Collecting the characters under generic headings yields the
following diagnoses :—

Genus Tracuuus Pallas.

Body unspotted; mandibular white stripe short or absent,
jugular stripe very long; white of throat narrow, separated from
that of chest and almost or entirely separated from that of inter-
ramal area.

An interramal gland covered with almost naked skin extending
from the chin to the interramal tuft of vibrissz.

EXTERNAL CHARACTERS OF CHEVROTAINS. 11

Kars short.

Tail long, covering the scrotum and inguinal region, which are
naked.

Legs and feet siender, interdigital cleft and back of pasterns
almost naked, posterior side of legs below knees and hocks naked
just above false hoofs, very scantily hairy elsewhere.

Penis complex, with a lamina normally folded up, beneath and
behind the coiled terminal portion.

Genus Moscuioia (Hodgson) Thos.

Body spotted and striped ; mandibular white stripe moderately
long, jugular very short ; white on interramal area, throat and
chest forming a wide continuous band.

No interramal gland, the interramal area covered with hair,
from which the tuft of vibrissz arises.

Ears short.

Tail very short, only covering the anus and leaving uncovered
the scrotum and inguinal region, which are hairy.

Legs and feet slender, interdigital cleft and back of pasterns
and posterior side of limbs below knees and hocks normally hairy,
hairs on posterior side of metatarsal region growing upwards
from a parting just above the false hoofs.

Penis simple, attenuated, spirally coiled distally, without
lamina.

Genus HyEmoscuus Gray.

Body spotted and striped ; mandibular and jugular stripes long ;
white of throat continuous with that of interramal area and of
chest.

No interramal gland, the interramal area covered with hair,
from which arises the tuft of vibrisse.

Kars long for the family.

Tail long, covering the scrotum and inguinal region, which are
hairy.

Legs and feet stout and strong, interdigital cleft and back of
pasterns nearly naked, posterior side of the limbs between the
knee and hock and the false hoofs normally hairy, hairs on the
back of the metatarsus forming a median crest, no parting above
the false hoofs.

Penis simple, attenuated, spirally coiled distally ; no lamina.

”

ON DEATHS IN THE SOCIETY’S GARDENS. iba;

2. Report on Deaths of Animals in the Gardens in 1918.
By J. A. Murray, M.D., Acting Hon. Pathologist
to the Society.

{Received March 4, 1919: Read March 4, 1919.)

As in previous years the main facts of the mortality among
the animals in the Society’s Gardens are summarised in the
Tables (I. and IT.) given below. In birds and reptiles (including
batrachians and fishes), the combined mortality statistics show
practically no change as compared with 1917 and previous years.
The higher mortality among mammals is mainly due to the
admission during 1918 of a large number of young monkeys
(M. rhesus) in which a heavy death-rate occurred. In addition,
a considerable number of the more easily replaceable animals were
sacrificed on account of the food-shortage occasioned by the war,

LW i En Ae = ess

Table II. gives the distribution of the more important causes
of death among the chief mammalian orders, in birds, and in
reptiles, batrachians, and fishes. In the case of the primates a
separate column has been reserved for the young J/acacus rhesus
admitted during the year. ‘This has seemed advisable to avoid
obscuring the details of the sufficiently severe losses among the
other monkeys, many of which had been in the Gardens for years.
In the case of acute infections of the respiratory tract, time has
not permitted the accurate separation of the cases into lobar and
broncho-pneumonia, capillary bronchitis, and acute congestion,
and they are therefore all included under the one heading of

14 DR. J. A. MURRAY ON

pneumonia. The considerable numbers in which no diagnosis
was made include, in addition to obscure cases, those which were
not examined, or were too decomposed to allow of a satisfactory
post-mortem examination.

Taste II.

MamMats.
|
= | | = |
= gee Erltses i| ' REPTILES
é | 2 | ape | 2s | Breps, || anp
$ |e |= |e = 5 | Total. FIsHEs.
Sh An Sot cami re |
S alds|/ae\ele
. General Diseases.
TuberGulOsis:-.Aeg.2-- ee nee oseee weet Woy 1 6 2 1 35 50 4
Miycosisgabecrretcst: aacicekh gasences pe cesche esses oe ee af 4 a2
Septicsomiiacpencestecessa-peesenaee: IS ecoml. alse 2a eal 8 3 11
INDSCESS een a Reni) ae 4 1 6 2 3
Berivonitismercoce cet aeetet eercie Coat eos le ail a tess Da Pa ES an
Helminthiasis .................. capt Woleies aN Reccel) ee lae z 5 6
INGTIBENSIS-e seneeonebosoebanneone abe : ue peel Wacren | eas Merete LE 2
ATES IND ree te nanan eneceeee th Osa rae ca| eae lhokee age esate 6 1
: Fee ules syctein | |
INTAIECIRISIS 5 peco00a ssecnanserios bo00sn 1 cia SRE RR RGU Ca sce 2 || J igaes| as
IBTEATO SNE Gououpeoosenoobsenacodes 8 34 12) 13)16| 14161 | 98 __ | 15
(Bdemavotiliuirasy ccc. ese ceecccee ails) omen eee sl eee etal lagers i fh 2 se
Bleurisy sh teres acre eee | | |
Dene ee eed a | ales eae ae
: he aey ne (ee cena |
Gastritis .. Be gaa Henoonee BeREe RAGE fk-obine |iessGu' llesoeae hous: (hace 1 ABS Serine vi.
Enteritis ..... ssuaeteasseni@e dd | adel iG: le Gelli aia eat oo 27 all 2 3
Tntestinal obstruction : PORE Hee 52s | peat [foc [Parent 1 3 1
Intussusception gens eee || eee asa eee leis 1 3 ee : me
Hepatitis: 120% caterers adec.seuel ees | 4-H cgi oe Bll eceay era ec 2) gal 8
amcneatiGis! 2. cone. s | Seales feces loco: 5 eel SB
. Urinary and Generative
Systems. | | |
INGO TENS couone: Beecuoaueneaduagabenal wenae erat IN Pee Bek 4
Cystitis, ............05. Searsddiceae lle FRET tired eS ulolang, sistema live ie se oe
Oar bi seca cnce Re ei wnsgete Nea ce Gee ee et 1
. Various. | | | | |
WNiews Giro witlhins ons sohecesveeeccec ee et Se tiestiel| ts estes | el [a eet ieeeeena al Pah Bek
Ghee aT a ee eet ee eee Se es Be | Ce |e omar Aaa 2
IBGE NVOYBH NEY=%2) cos dap nuance sosanoeca cee a eg eg alia ie 1 soi 2
Starvation and Malnutrition .... ... | .. | .. | .. |... |... dink a} 4 | 25
Injuries discovered post-Q 4 | i | 1
Sein % ve fe foe fone | oe Be
Killed by companions, yats, &e. ae: | Te els eal sk 19 2
Killed by order . pend Naki metal NE WApianef oc's | 17 5
Not diagnosed PN BCT 23 10d WES ee a) Mae Lal ao 58 89

Acariasis.—In addition to causing scabies when parasitic in
the skin of mammals, parasitic mites also occur in the respiratory
organs of birds with some frequency. The young Rhesus monkeys
referred to above were almost invariably found to harbour these
parasites in their lungs where they produced extensive lesions

DEATHS IN THE SOCIETY’S GARDENS. 15

consisting in connective tissue overgrowth around the bronchi,
local collapse of the lung, and sub-pleural dilatations of the
bronchioles and alveolar passages. In these cysts the mature
parasites are found, sometimes in large numbers. The excreta
of the mites, which contain very resistant doubly-refracting
crystals, seem to be the principal cause of the irritation. The
majority of these monkeys also suffered from intestinal nema-
todes, of which an Csophagostoma was the most frequent and
caused damage of varying severity to the colon. The larve of
this parasite apparently enter the body through the skin and
encyst in the wall of the large intestine causing multiple hemor-
rhages. When mature they burrow through into the lumen of
the gut, and the passage thus afforded to the micro-organisms
of the intestine occasionally leads to local and general peritonitis.

Atelectasis.—The two deaths in mammals ascribed to this cause
both occurred in new-born animals and present no points of
interest. The bird referred to this rubric was an adult Chilian
Sea-Hagle, in which sudden death was produced by the valvular
occlusion of the opening of an abdominal air-sac by a foreign
body. Very few respiratory movements apparently sufficed to
distend the air-sac and compress the lung above it so quickly and
completely as to cause death.

Pancreatitis.—This condition, which is an extremely rare cause
of sudden death in man, has been the cause of death six times—
four times in monkeys, once in a bear, and once ina bird. In
these, extravasations of blood of varying extent were found
throughout the gland. In addition, evidence of old inflamma-
tory changes have been found in the pancreas in a Californian
Sea-Lion and in a Slender Dog. In the latter, the condition
seems to have been due to stenosis of the pancreatic duct leading
to accumulation of inspissated secretion in all its branches, and
a very extensive overgrowth of the connective-tissue stroma of
the gland. The cause of the disease in monkeys has not been
cleared up and requires further study. The marine Carnivora
apparently share the enhanced liability to this rare and dangerous
disease with the Primates, a seal which died in 1917 showing it
in the acute stage. The Sea-Lion mentioned above had ap-
parently recovered from an attack.

New Growths.—Death could be ascribed to malignant new
growths in three cases :—In a Golden Eagle a large teratoma of
the testis, containing bone, islands of keratinising epithelium,
and masses of eosinophile leucocytes, was the seat of a fatal
suppuration. A Diamond Dove presented multiple white
nodules of lymphosarcoma in the liver. A Racoon died from
an enormous carcinoma of the thyroid gland. In none of these
were metastases observed in other organs. In addition, new
growths were observed in a number of animals dying from other
~eauses. A carcinoma of the liver was found in a Marsh-Buck
which died of septic pneumonia. This growth is interesting
because it is identical in structure with a type of cancer common

16 ON DEATHS IN THE SOCIETY'S GARDENS.

in the cow and sheep, and rare in other animals. A fibro-myoma,
arising from the circular muscular coat of the large intestine, was
discovered in a Hedgehog. It was loosely impacted in the pelvis,
and death was due to tuberculosis of the lungs and not to the
tumour.

Comparative Pathology of the Thyroid Gland.—The material
examined during 1918 has confirmed and extended the obser-
vations reported last year on the thyroid. In consequence of the
increased attention directed to this organ a number of cases of
thyroid enlargement have been found and studied. Senile cystic
goitre was observed in an Andaman Teal. Ina Nutmeg Finch
both thyroids were greatly enlarged, being twenty to thirty times
the normal size. ‘lhe microscopical appearance was that of ex-
ophthalmic goitre in man, the thyroid vesicles being filled with
masses of proliferated epithelial cells and practically no normal
vesicles with colloid could be seen. The Baska Tortoise which
died in 1918 also showed an enlarged thyroid. The gland had
undergone a tumour-like transformation, the epithelium of the
dilated vesicles being folded into papillomatous outgrowths. The
whole gland was increased in size, being three to four times as
large as that of another tortoise of nearly the same weight.
Neither in the Nutmeg Finch nor in the Baska Tortoise was
there any record of symptoms during life which could be ascribed
to the changes in the thyroid.

The accompanying table (Table III.) shows the influence of
meat and vegetable diet on the size of the thyroid gland very
clearly.

The animals are arranged in descending order as to weight and
the Herbivora are all much heavier than the Carnivora placed

TasieE ITI.

CARNIVOROUS. HERBIVOROUS.
Elephant-Seal oe if WasSiot land Tels eetiieie. eek eee ote
Lion CORDED Di Fei one sh aceneee CLOW) (Ghnw vers (GRw%)) snoeacacodossesos snosee OPO)
Tiger . Laue SOR Cpe a ae Malkin) ccoms atecre cetacean = OO)
Grizzly Bear... feana marcha 20) Zid eee See a ee cee: eer ene OO)
Himalayan Bear 2.4. see We S0 An0a) <i... SF cETE Seas BiO
CSPSIUNGM. “des nenncs ocdeboaraoseaovences JUL) Water- Buck .. i bot Sect)
Leopard (33 Ibs.) a ei iaeie ee amare) Maaco gest cocttnec cae suse seenage 2:0
Serval @blbs.) ccs 218 Capybara ..... Sei a tded 15)
Skunk ‘(72 lbs.) cs. ..00...cee 970 Poreupine (22 Ibs.) . Ak ea Peo

against them. Nevertheless, the latter have the heavier thyroids
with the exception of the Elephant-Seal and the Sea-Lion, which
live on a fish diet. The contrast in size of thyroid aia the
Elephant-Seal weighing 950 Ibs. and the Skunk weighing only
74 lbs. shows that it is not a structural peculiarity of Carnivora
as such, but a consequence of diet peculiarities, which manifests
itself in this way. Similarly, the Bears, Cat-bears, and Coatis,
which subsist on a mixed diet, do not aon the case re thyroids
of the=strict Carnivores,

‘
A}

ON FISHES FROM LAKE TANGANYIKA, ie

3. On a Collection of Fishes from Lake Tanganyika,
with Descriptions of three new Species. By G. A.
Boutencer, F.R.S., F.Z.S.

(Published by permission of the Trustees of the British Museum.)
[Received January 15, 1919: Read March 4, 1919. |
(Text-figures 1-3.)

When recently stationed in the Belgian Congo at Albertville,
M. Dhont-De Bie, who accompanied the late Dr. L. Stappers on
his 'Tanganyika-Mweru Expedition, made a collection of small
fishes which has reached me through the kind mediation of
Dr. L. Péringuey, Director of the South African Museum.

The collection, although a large one (1210 specimens), contains
representatives of but comparatively few species, most of the
specimens being referable to Z%lapia burtoni (780), Haplochilus
dhonti (210), and Stolothrissa tanganiee (94). One species,
Mastacembelus mellandi, is an addition to the fauna of Lake
Tanganyika, being previously known from the Solwin River
in Northern Rhodesia (Congo watershed), and three are here
described as new.

The species represented are the following :—

Leprpostrenip®. Protopterus cethiopicus Heck. (Albertville).

Ciurei%. Stolothrissa tanganice Regan (Lukuya River).

CuHaracinip&. Alestes vittatus Blgr. (Albertville).

Cyprinipz. Barbus serrifer Blgr. (Albertville), B. tanco-
pleura Bligr. (Albertville, Lukuga K.).

Cyrrinopontip®. Haplochilus dhonti, sp. nu. (Lukuga R.),
H. pumilus Blgr. (Albertville, Kalimié R., Lukuga &.), Lam-
prichthys tanganicanus Blgr. (Lukuga f.).

Cicntipe. Tilapia melanopleura A. Dum. (Lukuga R.),
7. burtoni Gthr. (Albertville, Kalimié R., Lukuga R.), 7. hori
Gthr. (Lukuga R.), 7. dardennii Blgr. (Lukuga R.), Petro-
chromis tanganice Gthr. (Albertville), Paratilapia lukuge, sp. 0.
(Lukuga R.), Simochromis diagramma Gthr. (Albertville), Lobo-
chilotes labiatus Blgr. (Albertville), Lamprologus dhonti, sp. n.
(Albertville), Plecodus paradoxus Blgr. (Albertville).

MastracEMBELIDE. MJastacembelus mellandi Blgr. (Lukuga R.).

Descriptions of the new Species.

HAPLOCHILUS DHONTI.

Depth of body 42 to 5 times in total length, length of head
4 times. Head flat above; snout shorter than eye; mouth

Proc. Zoou. Soc.—1919, No. Il. 2

18 MR. G. A. BOULENGER ON FISHES

directed upwards; lower jaw projecting; eye 3 times in length
of head, 3 postorbital part of head, 13 times in interorbital
width ; preorbital 3 diameter of eye. Dorsal 10, originating at
equal distance from head and from root of caudal, above anterior
third of anal, median rays longest, $ length of head. Anal 14—
15, rays as long as dorsals. Pectoral 2 length of head, reaching

Text-figure 1,

A ts es
SS LOIRE
<

Haplochilus dhoiti.

a little beyond root of ventral; latter small, mearer end of snout
than root of caudal. Caudal rounded, as long as head. Caudal
peduncle twice as long as deep. 29-31 scales in longitudinal
series, 16 round body in front of ventrals; no lateral-line pits.
A blackish lateral band.

Total length 35 millim.

210 specimens from the Lukuga River.

The nearest ally of this species appears to be H. myapose Blgr.,
from Zululand.

PARATILAPIA LUKUGA.

Depth of body 3 to 3] times in total length, length of head
3 times. Head twice as long as broad; snout with curved upper

Text-figure 2.

Paratilapia lukuge.

profile, rounded, broader than long, as long as postocular part of
head, shorter than eye, which is 24 times m length of head,
exceeds interorbital width, and equals 3 times preorbital depth ;

FROM LAKE TANGANYIKA. ‘ 19

mouth extending to below anterior border of eye; teeth very
small, in 2 or 3 series; 2 series of scales on the cheek, width of
scaly part 3 diameter of eye. Gill-rakers rather long, 16 or 17
on lower part of anterior arch. Dorsal XI-XII 13-14; spines
subequal from the sixth or seventh, which measures 3 to 3 length
of head, and equals longest soft rays. Anal III 9; third spine
2 length of head. Pectoral as long as head, extending a little
beyond vertical of origin of anal. Ventral produced into a fila-
ment, which extends beyond origin of anal. Caudal with deep
crescentic notch. Caudal peduncle 14 times as long as deep.

Scales denticulate, 34-36 a ; lateral lines — the upper
extending to the caudal peduncle, or to the root of the caudal.
Pale brown above, white beneath ; 4 to 6 dark spots on each side,
the first on the gill-cover, the last at the base of the caudal ;
dorsal and anal usually with dark spots or a dark longitudinal
band, the rays tipped with black.

Total length 63 millim.

Several specimens from the Lukuga River.

Closely allied to P. dewindti Blgr.

LAMPROLOGUS DHONTT.

Depth of body 32 to 32 times in total length, length of head
24 to 3 times. Head twice as long as broad, with convex upper
profile ; snout rounded, 13 times as long as eye, which is 4 to
43 times in length of head and equals interorbital width ; mouth

Text-figure 3.

AY)
ay

NY aa

Lamprologrs dhonti.

not extending to below anterior border of eye; 6 rather large
canine teeth in front of upper jaw and 4 in lower, followed by a
moderately broad band of minute teeth ; head entirely naked.
Gill-rakers very short, 6 on lower part of anterior arch. Dorsal
XVII-X VIII 8-9 ; spines increasing in length to the last, which
is a little less than 3 length of head ; longest soft ray § length of
head. Anal V—VII 6-7; last spine ? length of head. Pectoral

DH
al

20 ON FISHES FROM LAKE TANGANYIKA.

2 length of head. Ventral produced into a filament, extending
beyond origin of anal. Caudal rounded. Caudal pedunele as

long as deep. Scales 253 / ms ; lateral Imes 4. Greyish,
with a dark network on the body; soft dorsal and anal, and
caudal with small dark spots.

otal length 65 miilim.

Five specimens from Albertville.

This species should be placed between £. mocquardw Pellegr.,
from the Congo, and LZ. tretocephalus Blgr., from Lake Tangan-

yika.

ON THE SKULL AND AFFINITIES OF RANA SUBSIGILLATA. 2]

4. On the Skuli and Affinities of Rana subsigillata A. Dum.
By Miss Joan B. Procter, F.Z.8.

[Received January 17,1919: Read March 4, 1919.j
(Text-figures 1 & 2.)

Whilst making a study of the esteological characters in the
genus Hana, it was pointed out to me by Mr. Boulenger that
the skull of the West African Rana subsigillata A. Dum., to
which he was the first to draw attention *, might prove on closer
examination to be of such special interest as to justify a detailed
description. Mr. Boulenger, whom I have to thank for his kind-
ness in giving me every facility to make this study, regards this
frog as the monotype of a subgenus named by him Awbria. In
his paper 7, “‘ Apercgu des principes qui doivent régir la classifi-
cation naturelle des especes du genre Rana,’ some of the most
striking characters of this frog are briefly noticed, and its position
in the genus explained.

On raising the scalp, fer a superficial examination, the skull is
seen to approach that of Rana (Pyxicephalus) adspersa Tschudi
in several characters, which will be enumerated later. It is
strongly ossified, rather depressed, and broader than long, the
general shape being typieally frog-like. The interorbital portion
of the brain-case is slender. Seen in profile, the cranium slopes
upwards from the nasal region toa point in line with the posterior
orbital borders, from which there is an abrupt decline to the
foramen magnum (see text-fig. 1 c).

THE MEMBRANE-BONES OF THE CRANIUM.

The nasals are large, well-developed triangular bones, and
somewhat rugose. They meet each other in the median line
almost throughout their length, which is nearly two-thirds that
of the fronto-parietals. Anteriorly each is obtusely pointed ;
posteriorly they ferm short oblique sutures with the anterior
borders of the fronto-parietals, exposing in the centre a minute
diamond-shaped area of the ethmeid. The distal ends do not
reach the maxille proper, although they rest upon the maxillary
processes in conjunction with the palatine cartilages.

The vomers are oblique, presenting an acute angle backwards
and inwards, where they approach the proximal ends of the
palatines, and the parasphenoid. Anteriorly they are deeply
notched, the anterior processes reaching the maxille. These

* C. R. Ac. Sci. Paris, 165. (1917) p. 987.
+ Bull, Soc. Zool. France, xliui. 1918.

22, MISS J. B. PROCTER ON THE SKULL AND

bones overlie the subnasal lamine and the adjoining border of
the ethmoid. The vomerine teeth are arranged in a simple line
of four, springing from a prominent ridge on the outer oblique
edge of each bone.

The fronto-parietals are strongly ossified, and somewhat rugose
on the anterior surface. The sagittal suture commences almost
at their anterior extremities, but does not persist beyond a third
of their length. Their combined width, anteriorly, is about a
quarter of their length. At the postero-inner corner of the orbit
they form small sharp projections, and then reach double the
width. Posteriorly there is a slight sagittal crest, with two
oblique lateral wings which form the commencement of the
mastoid processes.

The parasphenoid is of the usual dagger shape, but rather
more shapely than that of 2. temporaria, and has an extremely
tapering point. The lateral limbs of this bone are partially
hidden beneath the superimposed inner limbs of the pterygoids.

THE CARTILAGE-BONES OF THE CRANIUM.

The ethmoid, as compared to that of 2. temporaria, is elongated.
In the specimen figured it is almost one-half the length of the
entire skull. Anteriorly it is trilobate, or fleur-de-lys-shaped ;
dorsally this portion is overlain by the nasals, and ventrally the
central lobe is partially obscured by the vomers and the proximal
ends of the palatines. The main tubular portion of this bone,
which reaches almost to the optic foramen, is covered by the
fronto-parietals, but is visible on the ventral side through the
semi-transparent parasphenoid. As already stated, only a minute
diamond-shaped area of the ethmoid is exposed to view on the
dorsal surface, where it is bounded by the postero-median notch
between the nasals anteriorly, and by the antero-median notch
between the fronto-parietals posteriorly. In the antero-inner
corner of the orbit the ethmoid is pierced on each side bya small
foramen for the orbito-nasal nerve.

The prootics.—The dorsal surface of each is largely covered by
the inner branch of the squamosal, and its inner borders underlie
the fronto-parietals. In the anterior wall below the flange of the
fronto-parietals is the foramen for the fifth and seventh cranial
nerves. The prootic forms the roof and the anterior wall of the
auditory capsule.

The exoccipitals meet each other in the median line both dor-
sally and ventrally. The prootic-exoccipital suture proceeds along
the crest of the mastoid process ; the fronto-parietal-exoccipital
suture is difficult to trace. Ventrally the anterior borders of
these bones are bounded by the parasphenoid. The exoccipital
condyles are well developed, and are visible from three aspects;
at the base of each are two foramina, the upper minute and the
lower larger and internally divided ; ‘these give exit to the ninth
and tenth cranial nerves.

AFFINITIES OF RANA SUBSIGILLATA. 23

THE CHONDROCRANIUM.

The nasal cartilages consist of a roundly-pointed nasal roof
above and a trifid sub-nasal lamina below.

Small triangular prerhenals are attached to the septum nasi.

The palatine cartilages extend from the ethmoid to the maxille,
and are not peculiar in any way.

Owing to the thickness of the fronto-parietals and the extreme
delicacy of the chondrocranium, J have not satisfied myself as to
the size and exact positions of the fontanelles. It is clear, how-
ever, that there is a large median fontanelle, the anterior portion
of which is bordered by the ethmoid, which at this point has a
shightly bilobular tongue-shaped area carved out of it. The
posterior wall of the otic capsule is cartilaginous; it is bounded
by the parasphenoid below and by the prootic above.

The columella auris is strongly developed. The inter- and
medio-stapedial portions are not unusual in any way; the extra-
stapedial section is strongly developed and has a fan-shaped
terminal of considerable size, which is applied in an inverted
position to the tympanic membrane. The squamosal, at the
junction of the interior limb with its stem, forms a deep arch
over this delicate cartilage, and also gives support to the annulus
tympanicus, which is somewhat funnel-shaped, broader than deep,
and slightly notched above.

THE MAXILLARY ARCH.

The palatines are straight, semi-transparent bones, placed at
right angles to the axis of the skull. They do net meet each
other.

The sguamosals differ widely from those of Rana (sensz stricto).
They are enormously developed. The zygomatic branch, slightly
rugose, forms a suture with the maxilla which is prolonged for-
ward to the naso-palatine bar. his process tapers considerably
at its anterior end, and is about half the length of the entire
skull*; the suture is about feur-fifths of this length. The inner
limb, just over one-third the length of the zygomatic, with which
it forms a right angle, is superimposed on the prootic, of which
but a small border is exposed on each side. The posterior limb,
or stem, of this bone is somewhat oblique, flat, and rounded at
its distal end; it is applied to the pterygoid and the quadrate
cartilage.

The pterygoids are the most remarkable bones in the skull.
The inner processes overlap the lateral processes of the para-
sphenoid almost to their junction with the blade-like portion.
At the junction of the anterior and posterior limbs, and opposite
the root of the interior limb, is a large rounded process, which,
when the skull is in its natural position, is directed outwards and

%* Processes measured from their junction with the shaft of the bone, not from the
right angle where they join each other.

24 MISS J. B. PROCTER ON THE SKULL AND

downwards *; when the jaws are closed the outer face of this
process is presented to the coronary process of the angulo-splenial,
which it resembles in size and shape. This process seems to be
very rare in the Anura, and until new has remained unnoticed.

The quadrate cartilage projects in two strong condyles, well
beyond the extremities of the pterygoid, squamosal, and quadrato-
Jugal bones. It 1s of massive build, and in the specimen figured
strongly ossified. ts

The premaxille have rather long processes. Ventrally, at the
suture which they form with each other, there is a shallow oval
pit.

The maaxille are wide, strong, and slightly rugose. Ventrally,
and close to the sutures which they form with the premaxille,
are round shallow concavities. . Posteriorly each is slightly
bifid, the inner border of the inner limb completing the squamoso-
maxillary suture, and the outer and longer branch forming a
wedge-shaped suture with the quadrato-jugal.

The quadrato-jugal is a small short bone; its suture with the
maxilla is very dithcult to trace in old specimens.

THE MANDIBULAR ARCH.

Meckel’s cartilage, where it articulates with the quadrate, ends
in a prominent down-curved knob.

The angulo-splenial leaves much of Meckel’s cartilage exposed
to view; its coronary process is well developed, and, as before
stated, lies face to face with the peculiarly similar process of the
pterygoid, from which it is only separated by the elevator
temporalis muscle.

The dentary at its distal end is raised into a slight tooth-like
projection, which, when the jaws are closed, fits into the shallow
pit at the anterior end of the maxilla which I have already
mentioned.

The mento-meckelian bone forms a similar but larger projection
which fits into the median pit situated across the premaxillo-
premaxillary suture.

THE HYOID APPARATUS.

As will be seen from text-fig. 1 d, the hyoid apparatus, though
differing slightly from that of fh. temporaria, is not peculiar in
any way.

SUMMARY AND CONCLUSION AS TO THE AFFINITIES OF
RR. SUBSIGILLATA.

From the above description it will be seen that the skull of
Rt. subsigillata differs almost in every particular from that of
R. temporaria, which, being the type of the genus, it is customary
to take as a standard.

* Owing to this oblique position it is slightly foreshortened in text-fig.la, b; a
better idea of its size can therefore be obtained from 1 ce, where it is seen projecting
below the maxilla.

————— ST ee

AFFINITIES OF RANA SUBSIGILLATA, 25

Its most salient characters are :—

1. The prolonged squamoso-maxillary suture, a feature which
it shares with J, adspersaw alone. (See text-figs. 1 a, ¢;
2 a, b, ©, €.)

2. The fourth pterygoid process, which it appears to share
with &. kuhlii alone, although a faint indication of this
process is sometimes met with in 2. adspersa. (See text-
figs. 1 b,¢; 2 d.)

3. The marked overlapping of some of the membrane-bones,
the most important of which are :-—

a. The interior branch of the pterygoid on the transverse
limb of the parasphenoid ; an overlap only otherwise
found in 2. adspersa.

b. The interior limb of the squamosal on the prootic, as in
R. adspersa, R. grunniens, Rk. macrodon, ete.

c. The nasals and fronto-parietals almost entirely obscuring
the ethmoid, as in &. adspersa and KR. tigrina. (Text-
figs. 1 and 2.)

4. The slight pitting of the nasals, dorsal plane of the fronto-
parietals, zygomatic processes of the squamosals, and the
maxille; a character carried to excess in R. adspersa, in
the adult of which these bones are covered with granular
asperities. (Text-figs. 1 a, ¢; 2 6, ¢, e.)

As regards the squamoso-maxillary suture and the pterygoid
overlap, it is an extraordinary thing that, whereas in no other
species of the genus Rana are these characters exhibited even in
old age, in &. subsigillata and L&R. adspersa they are well marked
even in specimens under a year old, in which the frontals are
still separated from the parietals (see text-fig. 2 a). This
seems to me to greatly enhance the importance of these features,
proving them to be no recent modification, and to isolate
completely these two species among all others of the genus.”

It is thus clear that the two species are closely related, a fact
which seems conclusively proved by the development of the skull
in the young ft. adspersa.

Specimens of this frog at about one year of age (text-fig. 2 a)
resemble in every cranial character the rather older young of
R. subsigillata. They have the frontals yet separated from the
parietals by oblique sutures ; the sagittal suture is complete, and
a moderate portion of ethmoid is exposed above. At this stage
the skull shows no rugosities. At the age of about two years
(text-fig. 26) the skull shows signs of pittings, and the fronto-
parietals close in and expose slightly less of the ethmoid. At
the age of about three years (text-fig. 2c, d) the skull con-
forms so exactly to that of R. subsigillata that (except for the
fourth pterygoid process) to describe it would be to recapitulate
the greater part of this paper. It has, however, slightly longer
and more closely-set preemaxillary processes, and consequently a
more pointed nasal roof, less triangular nasals, less developed
extrastapedials, and a less prominent quadrate.

26 MISS J. B. PROCTER ON THE SKULL AND

In later life (text-fig. 2.) the cranial characters of R. adspersa
become more pronounced, the whole skull broadening and assum-
ing a more convex and massive form, and the slight rugosities of
the halfgrown becoming marked excrescences in old specimens.

Although, therefore, these two species differ so widely in the
adult state, especially in external characters (such as general
shape, glandular folds, metatarsal tubercle, ete.), they bear
important fixed characters in common, which isolate them com-
pletely from all other species of the genus Rana. The subgenus
Aubria Blgr. might therefore be united with the subgenus Pyai-
cephalus Tsch. In this I find that I am not much at variance

Text-figure 1.

Skull of Rana subsigillata. (Nat. size.)

a. Dorsal view. c. Lateral view.
b. Ventral view. d. Hyoid.

with Mr. Boulenger, who has kindly allowed me to quote the
following remarks from his unpublished monograph on the Frogs
of Africa :-—“ Although I am at a loss to guess on what grounds
Giinther (Cat. Bat. Sal. p. 7) reached the conclusion, from the
original description of Aug. Duméril. that R. sabsigillata appears
to belong to the genus Zomopterna (= Pyxicephalus), it is un-
doubtedly a fact that, in spite of its very different appearance, it
is more nearly allied to R. adspersa (the type of Pyxicephalus)
than to any other, as the cranial characters show.”

After reviewing the cranial and other characters of these two

AFFINITIES OF RANA SUBSIGILLATA. 20

species and comparing them with, for instance, those of R. brevi-
ceps and £. delalandii, it is indeed difficult to understand how the
older authors (Tschudi, Duméril & Bibron, Giinther, Peters, etc.)
can have placed them together in one genus (Py«icephalus or
Tomopterna) merely on account of that worthless character, the
shape of the metatarsal tubercle. On trying to make use of their
comparative descriptions of Rana and Pyxicephalus one feels the
need of applying St. Jerome’s dictum: ‘“‘ Major styli pars que
delet quam que seribit.”

Text-figure 2.

Skull of Rana adspersa.

. At about one year. Dorsal view. (Nat. size.)
; two years. Dorsal view. (Nat. size.)
€. 5 5, three years. Dorsal view. (Nat. size.)
d. 5, 5, three years. Ventral view. (Nat. size.)
e. Skull of an old specimen. Dorsal view. (Half nat. size.)

oak

It seems probable that both species are derived from the
R. tigrina group of Rana, s. str., many peints about the latter's
skull confirming this view *.

* The proportions and positions of the nasals, frento-parietals, and ethmoid;
length and strength of the zygomatic process of the squamosal ; length of the immer
limb of the pterygoid and its suture with the parasphenoid.

“Wl Wel AOA Te

“6161

Sudoy [BWAOUge YIIM a[BWiey y[Npe pus B

S Ze

azeb xAfi1Q jo j[Bo

THE BREEDING OF ORYX GAZELLA AT GOOILUST. 29

5. On the Breeding of Orya gazella at Gooilust.
By F. H. Biaauw, C.M.Z.S.

[Received February 20, 1919: Read March 18, 1919.]
(Plate 1.*)

This species of Antelope is a scarce animal in Zoological
Collections in Europe, and therefore some details about its
breeding may be of general interest.

I got my pair in the summer of 1913, and although the im-
porter was very mysterious about its origin, it soon became
evident that the animals originated from the Kalahari, and had
reached me via the Zoological Garden of Pretoria, where they
had been kept some time.

Dwing a trip to South Africa in the spring of 1914 I saw
photos of animals of this species shot in German South-West
Africa, and a settler from that country told me that Oryx gazella
was fairly numerous in the southern parts of that colony to his
own knowledge.

The pair of animals at Gooilust did extremely well, and I soon
was able to let them out together in an enclosed field. Although
they fought at first—and these fights looked rather formidable
on account of the tremendously long and needle-sharp horns—I
soon found that the fighting was more play than anything more
serious, and so I could envi the sight of it without much
apprehension.

The animals, although in perfect health, did not breed at first,
and therefore, ‘having the chance of obtaining another female in
the spring of 1916, I acquired it. This female, although well
built in other respects, had deformed horns, which were rounded
as in Oryx algazel, so that I could not admire her very much.
However, I put all three animals in the field, and during the
course of the summer both females were covered by the male
and became pregnant. In the spring of 1917 the female with
the deformed horns, which had been looking in bad health for
some time, gave birth to a dead calf which was nearly fully
developed ; and as she did not seem to recover entirely, I later on
separated her from the others for fear of contagion.

On the 2nd of June of the same year my old female gave birth
to a splendid calf, which appeared quite strong and well.

This calf may be described as follows :——

General colour a rufous sand-colour. A black tip to the tail.
A dark streak from the eyes downwards, losing itself before
reaching the bottom of the jaw. There is a black tip to the
mane. The swelling which the adults have under the throat is
very conspicuous in the newly-born calf, and the hair there is
lengthened, forming a throat-mane. Inside of ears full of long
hairs, forming a fringe which projects out of the ear-shell. On
the outside margin of the ear a very thin black edge.

* Vor explanation of the Plate see p. 30.

30 THE BREEDING OF ORYX GAZELLA AT GOOILUST.

The horns became visible when the calf was three days old.
Perhaps they were visible from the first, but as during the first
two days I could not get near enough I cannot be positive about
this.

At the age of six days, under certain lights, the black mark on
the upper part of the fore legs of the adults became visible like a
dark shadow, especially on the inner side of the leg.

At six weeks old some of the face-markings of the adults
became slightly visible, and both the dark markings above the
knee and hocks. Horns 4 inches long.

The calf did not follow the mother, as I have found so many of
the larger antelopes do, but it kept itself hidden in a long tuft
of grass as a fawn would do. From time to time the mother
would come to it to give 1t milk, or if something came near that
would frighten it, the calf would keep still until it was quite near,
and then would make a sudden rush in the direction of its dam.
When the object of its fright was gone, it would again hide,
generally in the same tuft of long grass. Unfortunately, I have
not been able to rear the calf to maturity.

In the second half of July the weather, which had been warm
and dry, suddenly changed, getting cold and very wet. Of course,
everything was tried to induce the animals to enter into their
house, but it was several days before we succeeded. When at last
it was accomplished, the old female was suffering from a slight
diarrhea, and after a couple of weeks it died. An attempt was
made to rear the calf on cow’s milk, but this was not a success,
and on the 4th of August it died unfortunately. Although the
immediate cause of death seemed to be the chill occasioned by the
bad weather, I suspect the female with the abnormal horns, which
got ill first, to have contaminated the herd, as after a while she,
although not having been exposed to the bad weather, developed
similar symptoms and died, and the same happened to the bull.

It was not possible to get a photo of the calf with its mother,
as she would always put herself between the calf and the
photographer.

It is, I suppose, unnecessary for me to say that Orya gazella is
one of the most beautiful antelopes in existence, and that every
effort should be made to prevent this splendid animal being
exterminated in its native country. In German South-West
Africa these antelopes were preserved in large extents of country.
I trust that under new rule the same excellent measures will
continue.

These Antelopes are, I believe, still fairly numerous in some
parts of the Kalahari, but I have not been able to ascertain
that they are protected there. Measures in that direction would
certainly be very welcome to all lovers of the splendid South
African fauna.

EXPLANATION OF PLATE I,

Oryx gazella. Young calf and adult female with abnormal horus. i

ah oi nae

ms Zo S, IG, SMUT Pl. i.

ine, al,

Wié, Bs Wi. Be

SENSE-ORGANS OF DIPTERA.

Bile & iametsson, Ltd.

I 4, So AGI, SWOT, ell, i.

WIE, Be Hn@, Oo

SENSE-OREANS OF IDINEWE IVa,

Bale & Danielsson, Ltd.

Im 2, So WIG, SMWUIS, iPal, WN.

Fig. 9.

SENSE-ORGANS OF DIPTERA.

Bale & Danielsson, Lt

¥
oe

Ps 4 Sa IIB. SION! IPI, IW,

Fig. 11.

Fig. 12.

SENSE-ORGANS OF SDIPT ERA:

Bale & Vanieisson, Ltd,

SENSE-ORGANS IN ANTENNA AND PALPI OF DIPTERA, 31

6. A Comparative Study of certain Sense-Organs in the
Antenne and Palpi of Diptera. By IR M. Siri
A.R.C.8., D.I.C.* With Appendix by Professor H.
Maxwe.u Lerroy, F.Z.S.

[Received August 18, 1918: Read February 18, 1919.]

(Plates I-IV. ¢ and Text-figures 1-43.)

INDEX. Page
Tmtrodwctroniw. eeu catia here actrees ene ane etdeeatat Ou
Material and Method! ANA ta oo) | al
General Description of the Siousce lOreans endear review... 83
Chief Modifications of the Sense-Pits ............sceeeesses0) O84
Situation of the Sense-Organs.. aoe 35
Correlation between Form of Senee: Onsale pa Tani
of the Flies.. Ma tre SMe nce NDHs eres anv ROR mara cee MOO?
Function ......... 36
Systematic Description of sn Shes Qusaroast m Grew various
Families . baritone AN cata LA a pred emer TUS 17 6
HV) OS) NG UO. Coals Ae dem arya ebro ta a Pe ER 06
RELELCINCE SHARMA ee ae aee RAR oR Ree cea cee Be Bhan HOO

(1.) Inrropuction.

This piece of work was begun at the suggestion of Prof. H. M.
Lefroy and Mr. F. M. Howlett. It was originally to consist
merely of a comparison of the antennal sense-organs of Calh-
phora vomitoria and Musca domestica, but was enlarged to cover
as many types of Diptera as possible. It was decided to make a
comparative study of these sense-organs in the different families
and to correlate if possible their structure with the habits of the
insects.

Several workers have picked out insects apparently at random
and figured their antennal sense-organs, but so far as [ am aware
no one has yet made a systematic study and figured the type of
sense-organ in each family. This I have endeavoured to do, and
though it may not have led as yet to discoveries relating to the
function of these organs, it should at least have some anatomica!
and histological value.

(II.) Maveriat anp Meruop.

All the specimens examined were caught by myself in this
country. They were all taken alive and put direct into the
fixative. Several fixing fluids were tried: Carnoy’s Fluid, second

* Comiunicated by Prof. I], Maxweit Lerroy, F.ZS.
7 For explanation of the Plates see p. 69.

32 MR. K. M. SMITH ON CERTAIN SENSE-ORGANS

formula (consisting of Absolute Alcohol 6 parts, Glacial Acetic
Acid 1 part, Chloroform 3 parts), also Flemming’s Fluid, Cor-
rosive Sublimate, and Bouin’s Fluid. The one that gave the
best results was Carnoy’s Fluid.

The entire insects, or, in the case of large flies, the head only,
were left in this fixative for at least twenty-four hours, and were
then washed thoroughly in 90 °/, alcohol. The heads were cut

into longitudinal and transverse sections by the method of either —

double embedding in. wax and celloidin or embedding in wax
alone; for most purposes 1 found that wax alone sufficed. In
this latter case, after washing well in 90 °/, alcohol to remove
all traces of the fixative, the heads were left in absolute alcohol
for twenty-four hours; they were then put into a mixture of
equal parts of absolute alcohol and chloroform, thence into pure
chloroform, thence into a mixture of chloroform and wax, and
finally into pure wax. The changes must be as gradual as

ossible or shrinkage will occur. Much depends upon the time
the object to be cut is left in the wax; the chitin will become
hard and brittle if left in the wax too long, and a similar result
occurs if not left in long enough. No definite time can be given;
it varies from two hours with fragile insects to twenty-four hours
or more with very large flies with thick chitin, such as Asilids
or Muscids. 6 was found to be a good thickness at which to
cut the sections, and all the drawings and photographs are made
from sections of this thickness. Various stains were tried, those
giving the best results being Delafield’s Hematoxylin, Heiden-
hain, and Hematein. Most of the figures are drawn from
sections overstained in Delafield’s Hematoxylin and washed
out in Acid Alcohol. All the drawings were made at the level
of the microscope stage with the camera lucida at varying
magnifications.

The great difficulty in carrying out this piece of work has lain
in the tendency of the chitin, and with the chitin the delicate
structures underneath, to break up under the razor ; therefore a
good razor with a good edge is essential, and a wax of medium
hardness, melting-point 56°C., is the best. Owing to this
difficulty it is necessary to cut very large numbers of sections
and to select the best; over three hundred slides had to be
prepared in order to obtain the results here given.

The species of flies examined have been exactly determined
whenever possible. Owing, however, to various causes complete
identification of the material has not always been feasible, and
in some cases only the genus or family can be given. Acknow-
ledgments are due to My. C. G. Lamb, to Mr. C. J. Wainwright,
Mr. F. W. Edwards, and Rev. A. E. Eaton, for determining
certain species. I desire to thank Mr. Hugh Scott for much
useful assistance and advice, and Mr. Cecil Gunns, Chief Labo-
ratory Assistant at the Imperial College, for his valuable help in
making the photomicrographs.

—————

IN THE ANTENN# AND PALPI OF DIPTERA. 33

(III.) GeyxeraL DESCRIPTION OF THE SENSE-ORGANS
UNDER REVIEW.

The sense-organs of the antennz and palps of Diptera here
discussed * are composed of elements of the same general type as
those described in many other insects. These elements consist
each of a large, modified, hypodermal cell, above which is a very
thin-walled chitinous process, rising from the thicker chitin of
the general surface; and of a nerve- cfibre, which runs close up to
the base of, if not actually into, the chitinous process. The large
cells were taken by some earlier writers to be nervous elements
directly connected with the nerve-fibres, but Berlese (1) regards
their function as glandular and considers that the nerve-fibres
are in close apposition to, but not directly connected with, them.

Berlese (1) distinguishes several types of these chitinous sensory
processes (‘“‘sensilli”) including the following :—(i.) :Trichoid,
with the base sunk below the general surface of the chitin
but with the apex projecting above; (ii.) Basiconic, arising
directly from the general surface ; (iii.) Caloconic, with the base
sunk in a pit and with the apex not reaching the level of the
general surface. Wheeler (2) records the occurrence of all these
types in ants, From my observations it appears that in many
Nemocera the sensory processes are scattered singly on the
surface and are referable to one or other of the above ty pes.
In some Psychodide a modification is found: the processes are
very large and long, either spiral in form or bifid or triradiate,
arranged in pairs, a single pair to each joint of the flagellum of the
antenna (see text-fig. 3). In Cecidomyide also, there are peculiar
“looped hairs” and other structures which may be modifications
of simple sensory processes: see Felt (3), ete.

In the species of Bibio and the Mycetophilide which I have
examined, and in all the other families of Diptera studied, there
occur the compound sense-organs or ‘ sense-pits” which are the
special subject of this paper. These may be regarded as com-
posed of a greater or smaller number of the sense-organs outlined
above, united together, sunk in pits of very varying form or size,
and modified in various other ways (see below).

In describing a typical sense-pit, Pl. LV. fig. 11 is a good ex-
ample to take, being a photomicrograph of a transverse section of a
sense-pit in the antenna of Sarcophaga carnaria, magnified about
600 diameters. There is first a somewhat large opening in the
chitin leading down into the pit itself; in Sarcophaga carnaria
this opening is wide and leads abruptly to the sensory processes,
but in some species, e.g. certain Muscide, this opening leads into
a long channel lined by chitin which sometimes exhibits spiral
or convoluted folds or even a series of communicating ridges like
basket-work (Jfusea domestica). The floor of the pit consists of

* No attempt is made in this paper to deal with the organs known as Chordo-
tonal Organ or Johnston’s Organ, situated near the base of “the antenna in certain

Diptera.
Proc, Zoon. Soc.-— 1919, No. ITT. 3

34 MR. K. M. SMITH ON CERTAIN SENSE-ORGANS
a very thin chitinous membrane which is produced into the
sensory processes much in the same fashion that fingers arise
from a glove. These sensory processes vary much in shape;
in the particular insect chosen as an example they are bottle-
shaped, with the portion resembling the neck of the bottle
produced to a great extent. In some cases the abrupt change
from the thick surface chitin to the chitin lining the sense-pit is
extraordinarily marked, e.g. Pl. I. fig. 1. Beneath the floor of
the pit is a rounded mass of large radiating cells, each cell sepa-
rating from its fellow as it approaches the base of the pit and
running to its corresponding sensory process, see Pl. IV. fig. 11.
The whole mass of cells is embraced by a branch of the large
antennal nerve; this appears black in the photograph, Pl. IV.
fiomlal eve

Although, as mentioned above, the view of most workers has
been that these large cells are themselves nervous elements,
Berlese (1) asserts that this is a mistaken view, and that they
are glandular, while the nerve-fibres run between them. He says
that they secrete a fluid which fills the sensory processes, comes
through the chitin to the exterior, and bathes the whole surface
of the pit. He quotes Von Rath as agreeing with this view,
also Erichson (1847) and Suley (1891) as having affirmed the
presence of a “ humour” secreted osmotically. Packard (4) also
states that these sensory processes are filled with a serous fluid
and are definitely olfactory.

(IV.) Carer MopiricaTIons OF THE SENSE-Pits.

As stated above, the sense-pits are modified in form in various
ways. The following examples of this may be mentioned. In
the Stratiomyide (text-figs. 10-12) the processes are in groups,
each group arising from a common foundation, and having much
the appearance of a partly closed hand. In the Sy: phide the
chief distinguishing feature lies in the large number of sensory
processes and the great size of the pit ; also in this family appears
a further modification in that there are two kinds of pits, one
consisting of a simple inpushing filled with coarse chitinous rods
(PI. 1. fig. 2), and with few or none of the thin-walled sensory
processes, the other consisting of the large and beautiful type of
pit figured for many of the Syrphide, Hristalis tenax, Xylota syl-
varum, and others. A somewhat similar difference in the size
of the sense-pits occurs in the Muscide, and here also a further
modification is found, namely in the fusion of three or four
or sometimes more of the larger pits; e.g. Pl. II. fig. 5, in which
the pits are double. In some species of the Muscidz and some
Anthomyide, the entrance to the pit is elongated into a channel
whose wall consists of spiral folds or basket-work of chitin
(text-fig. 38). The greatest variation occurs however in the size,
shape, and number of the sensory processes themselves, which a
glance at the Plates will at once make clear,

IN THE ANTENNZ AND PALPI OF DIPTERA. 35

(V.) SrruaATION OF THE SENSE-OrGANS.

In those families of Nemocera which have no antennal sense-
pits, but only scattered sensory processes, these latter are found
on a number, if not on all, of the joints of the flagellum. In the
Bibionide, which have true pits, these were seen on a number of
joints of the flagellum (as well as on the palpi: see below), but
in the Brachycera and the Cyclorhaphous families antennal sense-
organs were only observed on the terminal joint *. As to the
palpi, pits were seen to be present on these organs in Myceto-

hilidee, Bibionide, and Therevide; while special sense-organs
of a somewhat different form are described in certain Stratio-
myide, Asilide, and Dolichopide. Wesché (5) also describes
pits in the palps of Rhyphide, Simuliide, Empide, and Pipun-
culide. He considers that when the antennal sense-organs are
not highly developed, those on the palpi are so, to compensate
for the deficiency. This is certainly true in some cases
(e.g. certain Mycetophilide, Therevide, and the families alluded
to by Wesché); but in some forms rather complex organs are
resent in both antenne and palpi, e.g. in Bibio marci, certain
Stratiomyide, Asilide, and Dolichopide.

(VI.) CoRRELATION BETWEEN ForM oF SENSE-ORGANS
AND Hapsirs OF THE FLIES.

It was hoped that after a systematic examination of the
antennal sense-organs of the Diptera, some definite correlation
between the form of these organs and the habits of the insect
would be found. Such correlation, however, is not apparent.
Wesché (5) mentions the case of Gastrophilus equi, the horse-bot,
coming up against the wind straight towards a horse, and points
out that this insect is well provided with antennal sense-organs ;
but so far as I bave investigated, the Syrphide have by far the
most complex and perfect sense-pits of all the Diptera, and it is
not easy to see how their mode of life calls for this development.
There is also the case of Musca domestica and Calliphora vomi-
toria ; the former is supposed by some entomologists to find its
food chiefly by sight and the latter chiefly by smell, yet there
is very little difference in the antennal organs of the two, except
that those of J/. domestica are slightly more complex, though of
course smaller in proportion.

In some flies possessed of very large eyes and keen sight,
e.g. Asilids and Empids, the sense-organs seem slightly smaller,
but in the case of the Muscide both eyes and sense-pits are large.
The Asilids and Empids are predaceous, but their sense-pits show
no apparent modification correlated with this fact. As to flies of
parasitic habits, I have examined no Pipunculids, but Wesche (5)
found in a species of this family no pits in the antenne but a pit.

* The flagellar “ complex” of the Brachycera is here treated as a single joint.
%

36 MR. K. M. SMITH ON CERTAIN SENSE-ORGANS

in either palp. In various Tachinids examined the pits are very
similar to those of non-parasitic Muscids. Of the Hippoboscide,
Ornithomyia appears to be poorly provided with antennal sense-
organs, having only a few thin sensory processes sunk in shallow
depressions, text-fig. 43.

(VII.) Function.

At different times very many entomologists have attempted to
settle the vexed question as to whether the antenne bear the
olfactory organs or no, by mutilation of the antenne or by
painting them with gum or some similar substances. It seems
to me that experiments like this cannot definitely decide this
question, relying as they do largely on statistics and there being
of necessity so great an element of chance. McIndoo (6), in his
paper on the olfactory sense of Hymenoptera, concludes that this
sense must be looked for elsewhere than in the antenne. He
says: “ It is seen that about one-quarter of all the workers who
have experimented on insects with mutilated antenne assert that
these appendages do not bear the olfactory organs.” On page 295
he states further: ‘It is now generally believed that the
antenne bear the organs of smell, but as all the antennal organs
are covered with a hard membrane the objection has been raised
that such organs cannot receive olfactory stimuli.” I do not
think the term ‘‘ hard membrane” is applicable to the antennal
organs of the Diptera at any rate, as the sensory processes in the
pit are so excessively thin-walled as only to be visible under quite
a high power of the microscope. Berlese (1) states that a fluid
is secreted which passes through these processes and bathes the
interior of the pit. Packard (4) also says that these processes
ave filled with a serous fluid and are definitely olfactory.
Personally 1 have never been able to find any traces of this
fluid.

W.M. Barrows (7), in his experiments on antenne, found that
gum on the antenne of Drosophila ampelophila does not keep out
odours. He etherised some flies and cut off the terminal antennal
segment (and he declares that the ether did not affect his
experiments). He writes:—“ It seems certain that the sense
of smell is absent or at least greatly reduced in those flies that
have lost the terminal joint of the antenne.” I emphasize the
word “ terminal,” because in all flies other than Nemocera it is
only in the third (7.e. terminal) joint that the sense-pits occur.
Wesché (5), who has studied this subject in some detail, writes:
“A number of experienced entomologists have separately come
to the conclusion that auditory organs exist in the antenne of
many species, and the deep pits or cavities in the antenne
of Muscids are thought to be such. This part then may be
a tactile, an auditory, or an olfactory organ in different species,
and it is probable that in many instances all three senses are
located, perhaps not exclusively but in part, in the antenne.

ours 5

iN THE ANTENNA AND PALPI OF DIPTERA. 37

Further, I found on the third joint of the antenne of Gastio-
philus equi a larger number of sense-pits than on any of the
flies mentioned (Helophilus pendulus, Hchinomyia fera, Thelaira
nigripes, Calliphora erythrocephala, Stratiomys chameleon), and
of a different structure.” As to the pits of G. equi being of a
different structure, I think Wesché is mistaken because, so far
as I could determine, they were very similar to the sense-pits of
other Diptera; they were certainly numerous, though not in
my opinion as numerous as those of the Muscids. Wesché also
says: “I think that when the antenne are not particularly
sensitive the palpi have these structures to compensate.’ This
statement is borne out only partially by the results of my
investigation (see above). Wesché states further: “ We thus
see that the palpi like the antenne can bear organs of three
senses—touch, taste, and smell, but I do not think that any
one palpus has more than two of these senses developed at the
same time.” He concludes: ‘“ (i.) Both the antenne and palpi
of insects are capable of receiving the stimuli of several senses ;
(ii.) Their capacities differ greatly in different species and conse-
quently a general rule is an impossibility.”

Packard (4) states definitely that these organs are olfactory
and agrees with Berlese (1) in saying that the pit is bathed in
a fluid.

Wheeler (2) describes what he calls olfactory and gustatory
sensilli on the third joint of the antenne. Hewrites: ‘It seems
to be impossible to distinguish between the sense-organs in insects,
although it may be asserted that the organs of smell are situated
mainly or exclusively on the antenne, whereas those of taste are
found on the mouth-parts, especially on the maxille and Jabium
and their palpi.” As stated above, I have not examined the
organ situated near the base of the antenna in certain Diptera,
known as the “ Chordotonal Organ” or “ Johnston's Organ ” and
which is regarded as auditory by most workers.

(VIII.) Sysremaric Description OF THE SENSE-ORGANS
IN THE VARIOUS FAMILIES.

ORTHORHAPHA. NEMOCERA.

Tipunip# (Text-figs. 1, 2). Species examined: Pachyrrhina
histrio F. (C. G. Lamb det.), and one undetermined species.
One kind of special sensory structure or ‘“ Sensillus ” was found
to be present; this is of the type described by Berlese (1) as
“ Trichoid,” that is, sunk in a pit but projecting up to or beyond
the general surface level. The shape of this sensillus varies
slightly in the two species examined, in one case being narrowed
to a point and having a distinct cap (text-fig. 1), The sensilli
are scattered more or less indiscriminately over the surface of
the antenne. In the palps no particular sensory apparatus was
apparent,

Packard (4) states: ‘The olfactory pits of Tipulids seem to

38 MR. K. M. SMITH ON CERTAIN SENSE-ORGANS

have a different structure to those of the other Diptera as the
external passage is closed.” I think Packard is mistaken here,
for my examination of the antenne of the Tipulide revealed no
pits at all but only the ‘‘ Trichoid ” sensilli of Berlese.

Text-figure 1. Text-figure 2.

Transverse sections through the middle cf the antenne of an undetermined
Tipulid and of Pachyrrhina histrio. X 980.

Except where otherwise stated, all the sections are through the third joint
of the antenne.

EXPLANATION OF LETTERING on the figures:—Ap., aperture of pit: B., base of

hair; C., cells; Ch., chitin; H., hairs; W., nerve; Nw., nuclei; Sp., sensory
process or sensillus; 7’, top of second joint of antennee.

PsycHopip# (Text-fig. 3). The form examined was Psychoda
albipennis Zett., determined by Rev. A. E. Eaton.

In this insect the apparent sensory apparatus consists of a pair
of pseudopodia-like processes on opposite sides of each joint of
the flagellum. These processes are very large for the size of the
insect and are triradiate. They are exactly comparable to the
sensilli of Tipulide, consisting as they do of an outpushing of
very thin chitin, only in this case they are carried to an enormous
length and are three-branched. There is also a very thick ring
of long hairs situated at the base of each joint. In text-fig.3 only
their bases (B) are shown so that they should not block out the
triradiate processes. In the palps there is no special sensory
apparatus visible.

Specialised processes or ‘“ chete” of this type have been
described in several other Psychodide. In brunettia superstes
Ann,, they are single, large, and spivally curved; and processes
of this form are also present in several other species belonging to
more than one genus (see Brunetti, ‘ Fauna of British India,’
Dipt. Nematocera, 1912, pp. 198-9, 248, &c., and pl. 4). Eaton
also refers to cheetz of somewhat the same type in Z'elmatoscopus

IN THE ANTENN® AND PALPI OF DIPTERA. 39

soleatus (Knt. Mo. Mag. 1893, p. 126), and to S-shaped ones
being present on each alternate joint of the flagellum of Psychoda
revisende (in litt., 23. 1.1918). A pair of small bifid chet is
present on each joint (except the last) of the flagellum in

Text-figure 3.

Segment of the flagellum of the antennae of Psychoda albipennis, showing the
triradiate process of one side in plan. X 980, from a whole antenna in cedar-
wood oil.

Psychoda bengalensis Brun., and P. nigripennis Brun. (see
Brunetti, op. cit. p. 199), but no case of a triradiate process
has hitherto been recorded. Some species of Phlebotomus also
have pairs of shorter or longer geniculated spines on each joint
of the flagellum (see Brunetti, op. cit. p. 200, and Newstead,
Bull. Ent. Research, v. 1914, figs. 4, 9, 10, &c.).

Cunicip& (Text-fig. 4).
In text-fig. 4, which is a transverse section near the tip of the

antenna of Culex pipiens, may be seen the sensilli, in this case
not deeply sunk as in Tipulidze, but of the basiconic type.

Text-figure 4.

Transverse section through the middle of the antenna of Culex pipiens. X 980.

40 MR. K. M. SMITH ON CERTAIN SENSE-ORGANS

CECIDOMYID&.

I have been unable to examine any of these insects myself, but
Felt (3) describes the peculiar looped hairs or circumfili on the
antenne; he compares them to the ‘apparently fleshy hypo-
dermal structures protruding from relatively large symmetrically
placed orifices on the antennal segments of Campylomyzariinz
and of certain Chironomids.” As I have not been able to
examine any Cecidomyide I ‘cannot speak with certainty, but
possibly Felt is not quite correct in describing these processes
as “fleshy hypodermal structures protruding from relatively
large symmetrically placed orifices.” What he takes for an
orifice may be the sudden change from the thick chitin of the
antennal surface to the excessively thin chitinous outpushing,
which is comparable to one of the sensory processes of a sense-pit
enormously enlarged. This sudden change from very thick to
very thin chitin certainly gives the impression of an orifice, but
under a high power it will be seen at once not to be so. I have
not found any of these very large chitinous processes in the Chiro-
nomids, but they are very evident in the Psychodide (see above).

Text-figure 5. is Text-figure 6.

Text-fig. 5.—Longitudinal section through the first and second joints of the
palp of an undetermined Mycetophilid. X 980.

Text-fig. 6.—Another view of the same. > 980.

ee:

hin

i teed

IN THE ANTENNA AND PALPI OF DIPTERA. 4l

MycerrorHiLip& (Text-figs. 5, 6).

Turning to the palps we find an entirely new modification of
the sensilli; instead of occurring singly each one sunk in its
own pit (Tipulide), or arising singly from the surface (Chiro-
nomidz), they are grouped together and sunk in a large pit some
distance below the chitin in the tissue of the palp. This isa
** sense-pit,” comparable to those of the later families. The pit
lies in the second joint and there is one in each palp. The sen-
sory processes are long and thin and spatulate. Text-fig. 6 is a
drawing of a transverse section showing the pit and its opening ;
text-fig. 5 is from a longitudinal section, showing the pit but
not its orifice.

Brpion1p& (Text-figs. 7, 8).

In some species of this family the multiple groups of sensilli,
or sense-pits, occur in both antenne and palps.

Text-figure 7.

Longitudinal section through the antenna of Bibio marci. X 600.

42 Mi. K. M. SMITH ON CERTAIN SENSE-ORGANS

In Sibio sp. the sensilli in the antenne, though increased in
size, still occur singly or in pairs. Text-fig.7 is a drawing of a lon-
gitudinal section of the antenne of Bibio marci 2; in this species
sense-pits are present, though the sensory processes are not very
numerous nor very deeply sunk, one sense-pit occurs in each

Text-figure 8.

Tiansverse section through the palp of Bibio marci. X 600.

segment of the antenna. In the palp the number of pits is
greater, and most of the upper surface of the palp in the region
near the head is broken up into sense-pits {text-fig. 8).

Wesché (5) describes sense-pits in the second joint of the palpi
of Bibio hortulanus.

SIMULIIDA.

I have not examined any members of this family, but Wesché
(5) deseribes pits in the palpi of Stmuliwm reptans.

RHYPHIDA,

Wesché (5) describes sense-pits in the palpi of Rhyphus
fenestralis.

IN THE ANTENNA AND PALPI OF DIPTERA, 43

ORTHORHAPHA. BRACHYCERA.

Leprip# (Text-figs. 9-16).

In Leptis scolopacea (C. G. Lamb det.) there are two types of
sensory processes, large ones arising directly from the surface and

smaller ones of somewhat irregular shape sunk deeply in pits
and not reaching to the surface, 7. e. the cceloconic type. In

Text-figure 9.

A. Wu.

Transverse section of the palp of Leptis tringaria. X 980.

L. tringaria (C. G. Lamb det.) one type of sensilius only was
seen, 7. e. celoconic. In the palp there is a well-marked sense-
pit, and beneath it a mass of radiating cells with a few large
nuclei (text-fig. 9). The sensory processes are long and some-
what spatulate.

SrratioMyip® (Text-figs. 10-12). Species examined deter-
mined by C. G. Lamb.

Text-fig. 10 shows a longitudinal section through one side of the
antenna of Oxycera trilineata F. In this species there are not
pits sunk below the surface, but groups of large sensory processes
projecting above the chitin and protected by large and thick
chitinous spines. These sensory processes resemble two hands
placed palm to palm with the fingers directed slightly inwards ;
in some cases, however, there are only two or three processes.
These “ hands” are duplicated on the other side of the antenna.

MR. K. M. SMITH ON CERTAIN SENSE-ORGANS

Text-figure 10.

Portion of a longitudinal section through the antenna of Oxycera trilineata.
X 980.

Text-figure 11. Text-figure 12.

Text-fig. 11.—Portion of antenna of Microchrysa polita in plan. X 980.
Text-fig. 12.—Longitudinal section through the top of the palp of Oyacera
pulchella. X 980.

IN THE ANTENNA AND PALPI OF DIPTERA. 45

Text-fig. 11 shows a portion of the antenna of Microchrysa
polita L., showing the “‘ hands” in position; the thin chitin which
forms the base of the structure is represented by cross-hatching.

Much the same state occurs in the palp. Text-fig. 12 shows
a longitudinal section through the top of the palp of Oxycera
pulchella Meig., with one ‘‘ hand” and one large single sensory
process.

Wesché describes very similar sense-organs in the antenne of
Stratiomys chameleon.

TaBpanip# (Text-fig. 13). Species examined determined by
C. G. Lamb.

In the antenne of the members of this family studied is found
a return to the less complicated form of sensory structure.
There are two kinds of processes, a large straight type sunk singly
in a small pit, but projecting far beyond the level of the chitin

Text-figure 13.

Part of a transverse section of the antenna of Tabanus bromius. XX 980.

(text-fig. 13): this is the trichoid type of sinsory process. There
are also small shouldered processes, sunk singly very deeply in the
tissue of the antenna. There is little or no difference between
the sensory processes of Tabanus bromius L.(Q ) and Hematopota
pluvialis L.(Q). Ihave been unable to find any special sensory
structure in the palps of this family.

Asitip (Text-figs. 14-16). Species examined determined by
C. G. Lamb.

In this group we return once more to the actual sense-pit.
Text-fig. 14 shows a longitudinal section through the antenna
of Laphria marginata; the chitin is enormously thick and the
entrance to the sense-pit is guarded by thick knobs and pro-
jections of chitin and interspersed with small pits containing one
or more sensory processes of varying shapes. ‘The processes in

Text figure 14.

Part of a longitudinal section through the antenna of Laphria marginata. X 980.

Text-figure 15.

Part of a transverse section through the antenna of Neoitamus eyanurus. X 980.

SENSE-ORGANS IN ANTENNA AND PALPI OF DIPTERA, ~ AT

the pit itself are long and regular, each one rising from a thick
chitinous ring or collar. Text-fig. 15, from a transverse section
through the antenna of Neoitamus cyanwurus Loew, shows a pit
widely open to the exterior, with short strong processes each
again rising from a collar, and having a ridge along its point.
Note the bulge i in the hypodermal layer formed by the columnar
cells, called glandular by Berlese. Text-fig. 16 shows a transverse
section through the palp of Laphria marginata L.; the whole of
one surface is broken up into long slender processes, not sunk in
pits but arising directly from the surface. Note the thickness of
the chitin in these insects.

Text-figure 16.

Transverse Section through the palp of Laphria marginata. X 980.

THEREVIDH (Text-fig. 17).

There are, so far as could be ascertained, no pits in the antenna
in this family. In the palp of Thereva nobilitata (text-fig. 17) is
a more complicated condition. There is a large sense-pit widely
open to the exterior, with long spatulate sensory processes ; also
here and there are single trichoid processes.

Empip#. Species studied determined by C. G. Lamb.

So far as could be ascertained there are no complicated sense-
pits in the forms examined. No special sensory apparatus was
visible in the palps of species examined,

48 MR. K. M. SMITH ON CERTAIN SENSE-ORGANS

Text-figure 17.

Part of a transverse section through the palp of Thereva sp. X 980.

Wesché (5) describes a large sense-organ on the palpi of an
Empid, Ocidromia glabricula. ‘This species I have not examined.

DotrcHorop1p® (Text-fig. 18). Species studied determined by

C. G. Lamb.
Text-figure 18.

Part of a transverse section through the antenna of Pecilobothrus nobilitatus.
X 980.

IN THE ANTENNH AND PALPI OF DIPTERA. 49

Text-fig. 18 shows a drawing of a portion of a transverse
section of Pesilobothrus nobilitatus L. Two distinct kinds of pro-
cesses are present, large finger-shaped processes scattered in large
numbers on the surface, and a smaller kind broad at the base and
tapering to a fine point. Turning to the palp, there are again
no pits and only one type of sensory process, this is in the shape
of a curved finger. In Dolichopus brevipennis Meig., the sensory
apparatus was very similar.

CYCLORHAPHA. ASCHIZA.

PIPUNCULIDA.

T have not had the opportunity of examining this group, but
Wesché (5) describes Pipunculus zonatus as having small and
characterless antenne but a well-marked sense-organ in the tip
of each palp.

Syrpuip# (Text-figs. 19-27).

This is by far the most interesting family of all the Diptera,
so far as the sense-pits are concerned. The Syrphide show an
enormous specialisation in the shape and complication of their

Text-figure 19,

Part of a transverse section through the antenna of Syrphus sp. X 980.

sense-pits, and the pits in different species of the family show
a wide differentiation. All seusory processes and sense-pits are
confined to the antenne, none being present in the palpi, so far
as could be seen. ‘Text-fig. 19, part of a transverse section of the
antenna of Syrphus sp., shows a somewhat deeply sunken pit
lined at each side with stiff chitinous hairs; at the base are two
bottle-shaped processes.
Proc. Zoou. Soc.—1919, No. IV. 4

50 MR. K. M. SMITH ON CERTAIN SENSE-ORGANS

Text-figure 20.

Transverse section of the antenna of Rhingia campestris. X 600.

Text-figure 21.

Transverse section {of antenna of Volucella bombylans. X 600.

IN THE ANTENNZ AND PALPI OF DIPTERA. 51

Text-fig. 20, from a transverse section through the antenna of
Rhingia campestris Meig., near the articulation, shows two sense-
pits, one on each side of the antenna; they are widely open and are
guarded by stiff hairs, the sensory processes are small and regular.
The bases of the pits approximate very closely to each other and
are separated by a large branch of the antennary nerve which
closely embraces each pit. In Volucella bombylans (text-fig. 21)
only one kind of sense-pit is found; this has a somewhat narrow

Text-figure 22.

Transverse section through antenna of Volucella inanis. X 600.

opening and is deeply sunk, the channel is guarded by a few
stiff hairs and the sensory processes are small and regular, Text-
fig. 22 shows a drawing of a transverse section of the antenna
of Volucella inanis. The section does not pass through the
opening of the pit. The processes here are of a different type,
they are rather short and sharply pointed, broadening towards
the base, and each process is separated from its fellow by a large
knob or block of chitin: cf. Rohler on Volucella (8).
A®

52 MR, K. M. SMITH ON CERTAIN SENSE-ORGANS

We come now to Hristalis tenax, the Drone Fly; this is a very
interesting case, possessing as it does two types of sense-pits
utterly different from each other. One type may be called the
Complex Pit and the other the Simple Pit. Pl. I. fig. 1 is from a
photograph of the former. It illustrates the very abrupt ending
of the thick chitin and the very.regular shape and character of
the whole sense-organ. Each cell can be seen running up to its

Text-figure 23,

Semi-dia sy: mmatic longitudinal section through the antenna of Vy/ota sylvarum.
x 128.

corresponding sensory process, and the whole organ is isolated by
a space running round the base of the cells and cutting it off from
the rest of the tissue. Pl. I. fig. 2 is from a photograph under ~
a low power of the simple type of pit. It is merely a large in-
pushing of the chitin widely open to the exterior, and instead of
the delicate sensory processes found in other sense-pits, there are

large and coarse rods of chitin, Also there 1s no abrupt thinning

IN THE ANTENNA AND PALPI OF DIPTERA. 53

of the surface chitin but it continues round the base of the pit
unchanged.

Turning to another species, Yylota sylvarum, a somewhat
similar condition is found. In this case, however, the simple pit
has no rods of chitin inside, but a few small sensory processes,
and is comparable to the sense-pits in many of the foregoing
families. The complex pit of Xylota, however, is very large and
beautiful ; it possesses an enormous number of sensory processes

Text-figure 25.

Text-figure 24.

Text-fig. 24.—Oblique section through the top of a sense-pit in the autenna of
Xylota sylvarum. X 980.

Text-fig. 25.—Semi-diagrammatic longitudinal section through the antenna of
Sericomyia borealis. X 128.

and almost fills the third joint of the antenna. Text-fig. 23 is a
semi-diagrammatic representation of a longitudinal section of
this joint. On the left side of the figure will be seen the large
and beautiful pit with its opening and opposite to it the small pit.
As the large pit is followed downwards it is seen to divide into
two ; the chitinous lining of the pit thickens between the pro-
cesses. PI. I. fig. 3 is from a photograph of a transverse section

Text-figure 26.

Transverse section of the antenna of Sericomyia borealis. X 600.

Text-figure 27.

Another transverse section of the same. > 600.

SENSE-ORGANS IN ANTENNA AND PALPI OF DIPTERA. 55

through the antenna of the same insect, taken under a low power.
It illustrates how far the pit extends into the antenna and the
large number of chitinous processes. Text-fig. 24 is an oblique
section through the roof of the pit slicing partly through the
chitin and exposing the pit itself. The chitin is perforated and
through each perforation can be seen a sensory process. Seri-
comyia borealis shows a somewhat similar condition. Text-fig. 25
is a semi-diagrammatic representation of a longitudinal section
through its antenna. The only difference from the condition in
AX. sylvarwm lies in the fact that the pit does not bifurcate. Text-
fig. 26 is from a transverse section of the antenna of S. borealis,
showing the large pit dissociated from the rest of the tissue, with
a cell running to each process. The pit is surrounded by a nerve
and all the nuclei are confined to the parts of the cells remote
from the sensory processes. Text-fig. 27 is a section through the
opening of the large pit and the nerve is shown encircling the
large cells.

CYCLORHAPHA. SCHIZOPHORA.
Supsip# (Text-figs. 28, 29.)

Text-fig. 28 is a drawing of a transverse section of the antenna
of a large undetermined Sepsid showing a sense-pit with its large

Text-figure 28.

Transverse section through the antenna of an undetermined Sepsid. X 600.

cells and the nerve encircling them. The sensory processes are
bottle-shaped and drawn out toa point. Text-fig. 29 is a seml-

56 MR. K. M. SMITH ON CERTAIN SENSE-ORGANS

diagrammatic drawing of a longitudinal section through the
antenna of a smaller Sepsid, showing the very large nerve
running to the sense-pit. The sensory processes are triangular.

Text-figure 29.

Semi-diagramimatic longitudinal section of the antenna of a small undetermined
Sepsid. XX 600.

SAPROMYZIDH (Text-fig. 30).

Text-fig. 30, longitudinal section of the two antenne of Sapro-
myza sp. On the outside of each antenna is one large widely

Text-figure 30.

Longitudinal section of the two antenne of an undetermined Sapromyzid.
X 600.

open pit, and on the inside of each a number of rather irregularly
shaped pits grouped somewhat closely together. The figure
shows the large nerves running to the pits.

IN THE ANTENN AND PALPI OF DIPTERA. 57

Opvomyzip# (Text-fig. 31).

Text-fig. 31 is a longitudinal section of an antenna of Opomyza
germinationis (F. W. Hdwards det.}; there is one sense-pit,
rather long and narrow, lined at the sides with stiff hairs and
having a few sensory processes at the bottom.

Text-figure 31.

Longitudinal section of antenna of Opomyza germinationis. X 600.

CHLOROPID# (Text-fig. 32).
'f'ext-fig. 32 shows a transverse section of the antenna of an

Text-figure 32.

Transverse section of the antenna of an undetermined Chloropid. X 600.

58 MR. K. M. SMITH ON CERTAIN SENSE-ORGANS

undetermined Chloropid. I have been unable to find any of the
true sense-pits in this species, and there appears to be only one
inpushing of the chitin filled with stiff hairs.

Hetomyzip# (Text-fig. 33).

The general surface of the antenna of Z'ephrochlamys rufiventris
is beset with large finger-like sensory processes spread about in
great profusion and interspersed between very large chitinous

Text-figurej33.

Longitudinal section through the second and third joints of the antenna of
Tephrochlamys rufiventris. X 600.

spines. Text-fig. 33 is a longitudinal section through the second
and third joints of the antenna of the same insect showing the
antennary nerve passing from one joint to another and also a pit
of simple type.

CorDYLURID& (Text-fig. 34).

Text-fig. 34, from a transverse section through the base of the
third joint of the antenna of Scatophaga lutaria, shows a com-
paratively large pit. The sensory processes are finger-shaped
and very numerous.

IN THE ANTENN& AND PALPI OF DIPTERA. 59

Text-figure 34.

Transverse section through the antenna of Scatophaga lutaria. XX 600.

AntHomy1D& (Text-figs. 35-37).

Text-fig. 35 is from a transverse section through the antenna of

Text-figure 35. Text-figure 36.

Text-fig. 35.—Transverse section of the antenna of Fannia sp. X 600.
Text-fig. 36.—Part of a longitudinal section of the antenna of Fannia sp. X 600-

60 MR. K. M. SMITH ON GERTAIN SENSE-ORGANS

Funnia. This shows one large pit and one small one; the sensory
processes vary in shape, those in the large pit are pointed while
those in the smaller are like a curved finger: this is unusual in
the same insect ; there are also large thin-walled processes (Sp.)
arising directly from the general surface. Text-fig. 36 is from a

Text-figure 37.

Transverse section of the antenna of an undetermined Anthomyid. X 600.

longitudinal section through the antenna of an undetermined
species (either Mydwa or Phaonia). It illustrates a sense-pit and
long, rather sickle-shaped processes, very profusely scattered on
the general surface. Text-fig. 37 is a transverse section of the
antenna of a large Anthomyid, showing a pit and its contained
processes.

Muscip# (Text-figs. 38 & 39 and Pls. II., II., IV. figs. 4-10).

The antenne in this family possess by a the eee number of
pits found in any of the Diptera examined ; they do not approach
those of the Syrphids in beauty and complication but are rather
small and regular. As far as can be ascertained there are no
solitary sensory processes on the surface of the chitin, but the
sense-pits vary greatly in size, some being very small, with only
two or three processes. Text- -fig. 38 shows part of a transverse
section through the antenna of Jusca domestica. It illustrates a
very beautiful pit, with complex chitinous ‘ basket-work” in the
long channel leading inwards from the orifice. The sensory pro-
cesses are finger-shaped and regular; the whole organ with its
large radiating cells gives somewhat the appearance of a fan. A

IN THE ANTENN® AND PALPI OF DIPTERA. 61

small pit with three sensory processes is shown on the right of
the figure.

Pl. II. fig. 4 is a photomicrograph of a transverse section
of the antenna of Calliphora erythrocephala; this shows the cells
running each to its long, straight, sensory process. PI]. II. fig. 6,
a transverse section of the antenna of the same insect under a
low power, shows one large pit with a number of small ones cut

Text-figure 38.

Part of a transverse section of the antenna of Musca domestica. X 980.

at different levels and mostly opposite to the large pit. Pl. II.
fig. 5, a longitudinal section, shows two large pits with their
openings and the beginning of a third pit, the large antennal
nerve being conspicuous. C’. vomitoria was also examined and
found to be very similar. For a detailed account of this latter
insect, see Rohler (9).

Pl. III. fig. 7 is a photomicrograph of a transverse section taken
under a very high power, of a specimen of Lucilia having the
thoracic chetotaxy of LZ. sericata; it shows the long chitinous

62 MR. K. M. SMITH ON CERTAIN SENSE-ORGANS

hairs at the opening and the short strong sensory processes
very wide at the base.

Pl. III. fig. 9, a photomicrograph of the antenna of Phormia
coerulea (=grenlandia) cut near the junction of the second and
third joints, illustrates the nerve and the variations in shape of
the sensory processes. PI. ITT. fig. 8, a photomicrograph of a lon-
gitudinal section through the antenna of the same insect, shows
the position of the sense-pits ; they are clustered together at the
end of the under surface nearest the head and are eleven in
number.

Pl. IV. fig. 10, 2 photomicrograph of a transverse section of
the antenna of Pollenia rudis: one pit is visible. The third joint
appears to be reinforced in the centre by a strong band of chitin
running parallel to the outer walls, which appears black in the
photograph.

A form which I have not examined, but in the antenne of
which funnel-like pits have been described (see Deegener (10) ),
is Muscina stabulans.

Text-figure 39.

Part of a transverse section of antenna of Mesembrina meridiana. X 600.

Text-fig. 39 shows a pit in the antenna of another Muscid,
Mesembrina meridiana L. (C. G, Lamb det,). It illustrates the

IN THE ANTENN&Z AND PALPI OF DIPTERA. > AGS

long channel running down to the base of the pit (cf. 1/. domes-
tica, text-fig. 38). There are a few short sensory processes, with
here and there a chitinous spine.

SARCOPHAGID 2.

PI.IV. fig. 11 is a photomicrograph of a transverse section of the
antenna of Sarcophaga carnaria. Each cell may be seen running
to its corresponding sensory process. The processes are broad at
the base, changing abruptly to a fine point. The nerve (J), which
shows black in the photograph, encircles the whole sense-organ.

Tacuinip& (Text-figs. 40, 41, and Pl. IV. fig. 12).

Text-fig. 40 shows a transverse section through the antenna of
an undetermined species. It illustrates the peculiar shape of the
sensory processes and the large nerve running to the sense-pit.

Text-figure 40.

Transverse section of the antenna of an undetermined Tachinid. > 980,

Pl. IV. fig. 12 is a photomicrograph of a transverse section of the
antenna of another undetermined species. In this, the sensory
processes are long and thin; the nerve is seen enclosing the whole

64 — MR. K. M. SMITH ON CERTAIN SENSE-ORGANS

sense-organ. ‘Text-fig. 41, a longitudinal section of the antenna
of Compsilura concinnata Meig. (= Phorocera serriventris Rond.)
2 (C. J. Wainwright det.), shows the nerve running up from the
head and branching in the third joint, and the large number of

Text-figure 41.

5 Coa TT

Longitudinal section of the antenna of Compsilura concinnata. X 128.

pits of various sizes on each side of the antenna. Mr. Wain-
wright informs me that this insect is a parasite of many hosts,
mostly Lepidopterous, but including also some Sawflies.

Cisrrip# (Text-fig. 42).

Gastrophilus equi: the surface of the antenna of this species
(text-fig. 42) is beset with small sensory pits with long slender

IN THE ANTENNE AND PALPI OF DIPTERA. 65

processes, the pits being placed close together. The hairs on the
surface are very numerous and give a matted appearance.

Wesché (5) pays great attention to this fly and considers it
the most highly sensitive of the Diptera, but with this I do not
agree (see above).

Text-figure 42.

Part of a transverse section of antenna of Giastrophilus equi. 600.

PUPIPARA.

Hippososcip& (Text-fig. 43). Ornithomyia avicularia L. (C.G.
Lamb det.): this species has not the true sense-pit, but slight
depressions in the surface with a few thin sensory processes.

Text: figure 43.

Sp.

Transverse section of the antenna of Ornithomyia avicularia. X 600.

Proc. Zoot. Soc.—1919, No. V. 5

66 MR, K. M. SMITH ON CERTAIN SENSE-ORGANS

APPENDIX.

This paper is the outcome of inquiry into the habits of the
House-fly which was initiated at the Zoological Gardens in 1915.
Miss Lodge’s paper on the behaviour of flies has appeared in the
‘Bulletin of Entomological Research,” November 1918; this
paper is complementary to it, and in trying to arrive at an inter-
pretation of these sense-organs, we have also further unpublished
work on the behaviour of flies towards chemicals which aids us
in understanding the function of the antenne. The structure of
the sense-organs described above sbows a general similarity in
fundamental design but marked variation in detail. Speaking
broadly the third joint of the antenne is alone concerned, it is
set over its surface with simple sensitive bairs, 1t contains pits or
cavities which have a more complex structure; these pits are as
a rule protected or situated on the inner surface of the antenna
and the whole joint is set on to the second in a curiously complex
fashion. Further, there is in some species an inner chitinous
skeleton, which has probably to be considered in conjunction with
the elaborate conical structure of the base of the sensory joint.

In considering the function of this antenna, one point sug-
gested itself which we may first dispose of: some flies have the
curious habit of hovering, 7.e. of resting poised in the air,
stationary, with a wing velocity adjusted to maintain them so;
it seemed possible that the antenne functioned as a speed-
perception organ, relative not to space but to air: that is, the
fly was able to feel if it were progressing against the air and
when sensation was nil, was stationary. As these organs are less
well developed in the Hover-fly section of the Syrphide than in
the *‘ Bee-fly” section, this is not supported. Another possible
view is that they ave auditory in function, and some of the more
complex pits suggest the auditory organs of Locustide ; but there
is little reason to believe that any flies respond to sound. I think
that a great deal of further observation will be required to
ascertain if some of the Syrphide especially do not have auditory
pits, but I think it is not the explanation of the function of the
sense-organ generally.

A third point that seems to be clear is that the antenne and
pits are not modified in the sexes: it has long been known that
the females of some Fruit-flies secrete essential oils or complex
aromatic ketones or aldehydes which are limited within the
species and which attract the males in a very wonderful manner:
in such cases males must have extreme delicacy of sense percep-
tion; but females have equally perfect sense-organs (if these
organs are olfactory in function) and they are not secondary
sexual characters, as they occur equally well developed in both
sexes of each species. Miss Lodge’s observations show that flies
with the antenne cut off are unable to retain their balance and

IN THE ANTENNA AND PALPI OF DIPTERA. 67

readily fall over. I think the structure of these organs shows
that the mere cutting of these large nerves must lead to very
great nervous disturbance, and there does not seem to be any
very good reason to believe that they function in control of the
legs.

An enormous number of observations with flies shows that they
are extremely sensitive to the presence of certain compounds.
If sugar is put out in a room in which there are many flies, the
sugar is presently covered with flies; if with the sugar is mixed
a chemical substance, it is possible to determine if the flies avoid
it, or if it is attractive to the flies. The only compounds to which
House-flies are sensitive are the volatile constituents of essential
oils, the volatile organic compounds which are soluble in fats,
and those compounds which give off ammonia. The compounds
are generally those which are fat-solvents, and, apart from the case
of Musca, the most active are the constituents of essential oils.

The antenna contains (1) a large nerve network, (2) secreting
cells and frequently, (3) fat cells. Apparently the hairs over the
main surface are filled with secretion, probably of a fatty nature,
which picks up essential oils readily. <A fly that is a flower-
feeder is attracted by the traces of essential oils secreted by the
flowers, and directs its course to the flower: arriving there, its
large outer surface is probably saturated with the absorption of
the perfume; it has still to locate the honey-secreting glands and
it is then that the pits come into play. The pits are generally
guarded, often on the inner unexposed surface of the antenna,
and only very great concentration of the perfume affects them ;
but they are extra sensitive and aid the fly in locating the source
of the perfume in the flower. Flies not only have to find food in
flowers, but also to find breeding-places : for some of these, other
kinds of odours are certainly the attraction ; and some flies have
two kinds of pits, one kind of which, I suggest, function for the
food, the other function for the breeding-place. J/usca, for
instance, is definitely attracted by ammonia, whether from the
manure heap, or from chemical compounds that slowly disengage
ammonia: it has two kinds of pits. It is a pure supposition,
that one of these pits locates one class of compound, another
locates another class ; but it is borne out by the occurrence of the
two kinds in Fristalis, which is a flower-feeder and which breeds
in decaying organic matter. The explanation that is suggested
of these organs is that they are purely olfactory, that the general
surface of the antenne acts for delicate perceptions, that the pits
come into play when the concentration of the absorbed liquid
has dulled the simple organs on the outer surface, and that the
final location of the source of scent is due to the protected pits.
Further it is suggested that the presence of two kinds of pits in
some species is correlated with the dual perception in the
female of food and of breeding-place, in the male, of food and of

the female.

68 MR. K. M. SMITH ON CERTAIN SENSE-ORGANS

There remains the curiously complex cone on which the
third joint is set: two explanations are possible: one is that it
isa method of rigidly fixing the third joint so that it cannot
damage the nerve by bending or twisting motion; the other is
that the concentration in the third joint of the antenna of
volatile liquids expands it, puts pressure on the cone and gives
the fly the impulse to go where the excess of absorbed light will
vapourise, so relieving the pressure and also renewing the
sensitiveness of the organ. I put this forward as a mere
suggestion, but I think that some explanation of this curious
structure will be found and that any one who is further investi-
gating the behaviour of flies will find it useful.

There is obviously scope for much detailed observation and
for correlated examination of the sense-organs. Mr. Smith’s
paper offers a fine basis for this work, and if observation of habits
can be correlated with investigation of structure, we should get
much further in our understanding of the senses and sense-
reaction of flies.

H. M. Lerroy.
Imperial College of Science.

REFERENCES.

(1) A. Bertese.—Gli Insetti, vol. i. 1909, pp. 610-633, 693-695.

(2) W. M. Wueeter. —Ants, 1910, p. 59 sqq.

(3) E. P. Fevr.—-Cireumfili of the Cecidomyiide. 23rd Rep.
State Entomologist, being N.Y. State Mus. Bull. 124,
Albany, 1908, pp. 305-7.

(4) es a Su of Entomology, 1898, pp. 264—-
(5) W. Wescut.—Some new sense-organs in Diptera. Journ.
Quekett Club, (2) vol. ix. 1904, pp. 91-104, pls. 6, 7.

(6) N. E. McIypoo.—The olfactory sense of Hymenoptera.
Proc. Ac. Nat. Sci. Philadelphia, vol. Ixvi. 1914, pp. 294-
341, pls. 11, 12.

(7) W. M. Barrows.—The reactions of the Pomace fly, Droso-
phila ampelophila Loew, to odorous substances. Journ.
Exp. Zool. vol. iv. 1907, pp. 515-537.

(8) E. Router.—Zur Kenntniss der antennalen Sinnesorgane
der Dipteren. Zool. Anz., vol. xxx. 1906, pp. 211-219,

(9) E. Router.—Beitrige zur Kenntniss der Sinnesorgane der
Insecten. Zool. Jahrb., Anat., vol. xxii. 1905, pp. 225-
285, pls. 15, 16.

(10) P. Dercener.—Sinnesorgane, being Chap. 3 of C. Schréder’s
‘Handbuch der Entomologie,’ vol.i. pp. 140-233 (Lief. 1-2,
1912-13).

IN THE ANTENNZ AND PALPI OF DIPTERA. 69

“EXPLANATION OF THE PLATES.

Except where otherwise stated, all the sections are through the third joint

of the antenne.

EXPLANATION OF LETTERING :—Ap., aperture of pit; B., base of hair; C., cells;
Ch., chitin: H., hairs; N., nerve; Nuw., nuclei; Sp., sensory process or sensillus ;
T., top of second joint of antenne.

Fig.

So OU

Fig.

Welle oS

Fig. 10.
11.
12.

Pruate I.

. Part of a transverse section of the antenna of Evistalis tenax. > 600.
. Transverse section of the antenna of Wristalis tenax, showing the “simple”

pit. xX 128.

. Transverse section of antenna of Xylota sylvarum. X 128.

Prats II.

. Transverse section of the antenna of Calliphora erythrocephala. X 600.
. Longitudinal section of the antenna of C. erythrocephala. X 600.
. Transverse section of the antenna of the same insect. 128.

Prare IIT.

. Transverse section of the antenna of Lucilia sericata. xX 980.
. Longitudinal section of the antenna of the same insect. > 600.
. Transverse section of the antenna of Phormida coerulea. X 60.

Prats IV.
Transverse section of the antenna of Pollenia rudis. X 600.
Transverse section of the antenna of Sarcophaga carnaria. X 600.
Transverse section of the antenna of an undetermined T'achinid sp. X 600.

‘a
ane
us

ON THE REDUCTION OF THE JUGAL IN MAMMALIA. a

The Progressive Reduction of the Jugal in the Mammalia.
By Lanczrtor T. Hoesen, B.A., B.Sc.*

[Received February 27, 1919; Read April 8, 1919.)

Although the character of the zygomatic arch is at once a most
diagnostic feature of the Mammalia as a class, and one which
undergoes profound modification in the individual orders and
families, no adequate general account of it has been given since
Slade t contributed to our knowledge of the features it exhibits
in response to the functions it is called upon to perform.
Significant as it is, however, to appreciate the manner in which
the development of the zygomatic arch depends on the energy
and character cf the masticatory process, such treatment does not

xhaust the interest of its structure. Cases occur abundantly
where, though otherwise similar in curvature, resistance, and
functional significance, it is fundamentally dissimilar in the part
played by its component elements ; and a careful study indicates
a tendency for the suppression of the jugal element independently
manifested along totally divergent lines of mammalian descent.
Moreover, while the manner in which the zygomatic arch 1s
adapted to the requirements of different types of mammalian
organization leads to considerable diversity in its structure, there
are some striking particulars of resemblance where its phy s10-
logical importance may be totally different; and the student of
morphology cannot fail to give consideration to any details
of contrast and comparison which do not seem to be called into
being by the conditions of environment and in consequence
capable of being attributed to convergent evolution. Certain
points which remain therefore to be set forth or emphasized seem
to justify a brief survey of the role played by the malar among
the mammalia, though it may necessitate repeating imatter
already published.

In no group of Placentals except the Carnivora does the

zygomatic arch show the same uniformity of structure as in
the ] Marsupials, and when the varied types of dentition and diet
exemplified by the latter group are taken into account, this fact
should merit serious attention in discussing the possible diphyletie
origin of the group. It may at least be inferred that the type of
structure which characterizes the Marsupials approximates closely
to the ancestral condition. In all living Metatheria the jugal
plays a conspicuous part in its formation—even in the singular
genus Notoryctes. It is a robust structure usually curving out-
wards conspicuously. The relations of the Jugal are in general
the same: it extends from the lacrymal antero-dorsally at the
border of the orbit to the glenoid postero-ventrally. The.

* Communicated by Mr. H. W. Unruanx, B.A., B.Sc., F.Z.S.
* Slade, D. D. “The significance of the Jugal Arch.” Pr. Amer. Phil. Soe.
XXXiv., 1895.— Cf. also Hallman, E., “ Die Vergl. Osteol. die Schlafenbeins.”

72 _ MR. L. 1. HOGBEN ON THE

squamosal, however, always occupies a large proportion of the
dorsal side of the arch ; and it is perhaps not strictly accurate to
state * that the jugal always actually participates in the formation
of the glenoid cavity, because though the latter may be compact,
as in Phascolarctus, its limits are not always clearly defined. Not
infrequently it is a large smooth area, tapering off imperceptibly
to the flattened ventral rim of the arch, and when the condyle of
the mandible is small only a portion of this surface is strictly
articular. It is certain for example that the head of the mandible
in its normal rotation does not come into contact with the jugal
in the Rat Kangaroo, Hypsiprymnus rufescens. The postorbital
process of the zygomatic arch is but slightly indicated, indifferently
by the squamosal or jugal or both among closely % allied genera, and
the same variability in the relation of the different elements of the
zygomatic arch to the eye-socket is encountered among the various
orders of the Placentals. Widely as the structure of the zygo-
matic arch differs among the Placentalia, tle part played by the
jugal in its composition varies more remarkably. In a large
number of genera that occupy a somewhat isolated position, and
appear to have diverged from the main lings of mammalian
phylogeny at an early date, and also in some cases in the less
specialized members of the larger groups, the jugal displays what
may be regarded as the ancestral condition, extending from the
lacrymal antero-dorsally to the glenoid postero-ventrally. But
more generally it becomes displaced by the encroachment of the
squamosal and maxilla, its reduction being sometimes accompanied
by a strengthening of the zygomatic arch or in others by a
weakening, and frequently without any evident modification
-either of its contour or its rigidity.

In the Rodentia the zygomatic arch is always comparatively
well developed; and no student of this group can fail to be
impressed by the highly characteristic modifications it undergoes
among the various families. Brandt +, following Waterhouse,
paid particular attention to the character of the zygomatic arch
in his classification of the Rodents; and it may be said that his
system would have approximated more nearly to those of sub-
sequent investigators {, if he had studied the relation of its
constituent elements in more detail. Thus the Bathyergide
placed by him with the Myomorpha, the Anomaluride and
Haplodontide in the Sciuromorpha, should be separated from
those groups on account of the relations of the jugal bone if for
no other reason.

* Weber, Max: “es erstreckt sich bis zur Fossa glenoidea und bildet deren
Aussenflache.” Einfithrung in die Anatomie systematik der recenten und fossilen
Mammalia.

+ Brandt, J. F. “Untersuchungen tiber die craniologischen Entwickelungsstufen
und die davon herzuleiteten Verw andtschaften und Classificationen der Nager der
Jetztzeit, mit besonderer Beziehung auf die Gattung Castor.” Mém. Ac. Sci. St.
Pétersboure, 1855.

t Winge, H. “Jordfundne og nulevende Gnavere.” HK Museo Lundi, 1888.

Tullberg, T. “‘ Ueber das System der Nagethiere.” Upsala, 1899.

REDUCTION OF THE JUGAL IN MAMMALIA. 73

In the Sciuride and Castoride the jugal is least reduced,
forming alinost the entire arch and extending from the lacrymal
antero-dorsally to participate in the formation of the glenoid
postero-ventrally. In most of the remaining families the Jugal 1s
invaded by the maxilla from before and the squamosal from behind.
Tn a number of families the jugal still meets the lacrvymal : such are
the Pedetide and Anomaluride, Dipodide, Bathyergide, and, in
marked contrast with the other Hystricomorphine families, the
Chinchillide. In the Hystricide, Erethizontide, and the Caviide
the zygomatic process of the maxilla encroaches further and
occupies a large portion of the arch excluding the junction of the
jugal with the lacrymal; but the jugal nevertheless takes part in
the formation of the glenoid cavity, although in the larger Cavies
it is covered over on the external side of the arch by a superficial
extension downwards of the squamosal. This is conspicuously the
ease in Hydrocherus, in which genus the jugal appears on the
outer side to be reduced to a narrow yertical wedge of bone
between the zygomatic processes of the maxilla and squamosal,
albeit in curvature and general characteristics the zygomatic arch
conforms in a striking manner to the hystricine type. In fact,
Hydrocherus, which in several respects apart from its great size
appears to be one of the most highly specialized members not only
of the Caviide but of the Rodents, illustrates forcibly the general
tendency among the Mammalia for the displacement of the jugal
without apparent reference to function at all. The most extreme
reduction of the jugal is met with in the Murine forms, where the
whole arch is very slender and largely composed of the maxillary
zygomatic process which approximates closely to that of the
squamosal. In the Duplicidentata, on the other hand, the jugal
forms almost the entire arch extending behind but not actually
uniting with the glenoid. There is rarely any marked post-
orbital process in the zygomatic arch in Rodents.

Among the S. American Edentates the zygomatic arch is
complete only in the Dasypodide, where considerable differences
are exhibited even among members of the same genus. In
Dasypus sexcinctus, Chlamydophorus, and in Priodontes gigas the
jugal curves downwards and articulates with the squamosal by a
horizontal fissure; and in the first two genera there is a slight
indication of the downwardly directed process so characteristic of
the Sloths and Glyptodontide. In Zatusia hybrida the jugal also
eurves downwards, but articulates with the squamosal vertically.
In TVatusia novemcincta the arch is represented by a broad
straight bar mainly composed of the jugal element. In the
remaining recent families the jugal fails to meet the zygomatic
process of the squamosal, though in the Bradypodide it is a large
bone characterized by a ventral prolongation. Inu the Myrmeco-
phagide the jugal is slender and often styliform, while the
zygomatic process of the squamosal is represented by a blunt
projection except in Myrmecophaga tamandua, where it is drawn
out into a tapering process that almost meets the jugal. There

74 MR. L. ’. HOGBEN ON THE

is evidently within the Xenarthra a tendency for the reduction of
the jugal along several genetic lines; yet despite the variety of
structure displayed by the zygomatic arch among EHdentata vera,
there is one feature common to all the coups of which it is
composed: the jugal, however much reduced from behind, not
only meets the lacrymal but extends well in front of the orbit
though less conspicuously than in the Ungulates. Insignificant
as this might at first appear, it is striking in view of the extra-
ordinary ‘diversity exhibited in the habit, diet, general
organization, : and in other characters of the zygomatic ar on itself ;
and there is every reason to regard it as a common pices
peculiarity.

Whether this conclusion is correct or not, the Xenarthra are in
this respect sharply differentiated from the two spurious groups
of Edentata, Pholidota and Tubulidentata. In the Manide the
zygomatic acebs being incomplete, superficially resembles that of
the Myrmecophagidee ; ; but there is a zygomatic process to the
maxilla and the jugal is absent. If it were present, it would
oecupy the middle rather than the anterior portion of the arch as
in the Myrmecophagide. In Orycteropus.the zygomatic arch 1s
complete ; and the maxilla forms at least half of it, reducing the
jugal to a slender rod between it and the well- developed ZY gO-
matic process of the squamosal. It is interesting in relation to
Elliot Smith’s view * that the Tubulidentata show affinity with
the Protungulate stock, to note that im all modern Ungulates
and Hyrax the jugal suffers no displacement on account “of the
maxilla.

Among the Carnivora there is striking uniformity in the
dhemecies of the zygomatic arch which, as Slade observes, main-
tains in all outstanding particulars the same features as those
exhibited by the Creodonta. It is a robust structure curving
upwards generally from behind. The jugal forms a large part of
the arch and may extend back to the glenoid as in Lutra: in
front, it never extends beyond the anterior boundary of the orbit ;
but it articulates with the lacrymal universally in the Fissipedia.
On the internal surface of the arch the zygomatic processes of
the squamosal and jugal sometimes meet. There is a postorbital
process usually on the jugal, and sometimes on the squamosal.
The Pinnipedia display the same general characteristics except
that the lacrymal is absent. Significantly enough, the jugal
extends back to the rim of the glenoid in the two families least
specialized to aqui: atic life--the Otariide: and Trichechidee ; where-
as in the true Seals (Phocide) the extent of the jugal both at its
anterior and posterior extremities is in a marked manner more
restricted by the growth of the maxilla and squamosal, though it
cannot truly be siid that the zygomatic arch is essentially different
in other respects.

* Smith, G. Elliot. “The Brain in the Hdentata.’ Trans. Linn. Soc. London,

vil., 1899. —Cf. also Wortman. ‘The Ganodonta and their relations to the
Edentata.” Bull. Amer. Mus. Nat. Hist. ix., 1897.

STat eae

OS a yO

REDUCTION OF THE JUGAL IN MAMMALIA. 75

The jugal bone in the Ungulates articulates with the lacrymal,
extending beyond the anterior border of the orbit, as in the
Xenarthra, and constituting an appreciable part of the facial
region of the skull. The zygomatic arch is universally complete,
and at its posterior end is nearly always flattened. Among the
Tapiride and Rhinocerotide the jugal is least reduced in the
Perissodactyle forms, where the orbit is not enclosed. In the
Equide the posterior extremity of the jugal does not extend
vertically behind the orbital ring which is completed by the
junction of the frontal with the squamosal. In the Pecora,
Vragulina, and Tylopeda the zygomatic arch is so similar as to
separate these three groups widely from the Suina. The orbit is
enclosed by the union of the jugal with the frontal, and the
former does not extend far behind it postero-ventrally. But in
the less specialized Suina, where there is no separation of the
orbit from the temporal fossa, the jugal may extend back to the
glenoid cavity as in Sus and Dicotyles. Here the same tendency
for the displacement of the jugal by the zygomatic process of the
squamosal manifests itself, as has been referred to already in
Pinnipedia. In Hippopotamus alone does the enclosure of the
eye-socket occur—and not invariably throughout the genus ; but
the structure of the orbital ring when complete differs in this case
from that of the higher Artiodactyla in that the jugal and the
frontal meet one another on the anterior rim of the orbit internal
to the lacrymal bone.

The jugal is not prolonged into the facial region beyond the
lacrymal in the Hyracoidea, the condition of the zygomatic arch
being similar to that which obtains in the Marsupials and some
Rodents. The jugal enters largely into the formation of the
glenoid cavity : in no mammals do the maxilla and squamosal play
less part in the formation of the arch; and if the modern
Hyracoidea provide any indication of the type of zygomatic arch
prevalent in the protungulate stock, although it may be admitted
that the extension of the jugal into the facial region is a secondary
modification, the considerations stated with regard to the reduc-
tion of the jugal both in the Perissodactyles and Artiodactyles are
nevertheless greatly reinforced.

The progressive reduction of the jugal is clearly illustrated in
Andrews’s work on the ancestry of HKlephants. In the modern
forms the maxilla is extended backwards in a stout zygomatic
process excluding the union of the jugal with the lacrymal; and
the squamosal approaches it upon the dorsal side of the arch. The
jugal, which is reduced to a narrow segment interposed between
them, sends back a slender limb postero-ventrally which takes part
in the formation of the glenoid. The participation of the jugal in
the jaw articulation does not appear to be a secondary condition,
for, as Dr. Andrews* observes, in JMJeritherium ‘‘the jugal is
large and forms the-greater part of the zygomatic arch. Posteriorly

* Andrews, C. W. Tageblatt des V. Int.-Zool. Cong. Berlin, No. 6. Phil. Trans.
vol. 196.

76 MR. L. T. HOGBEN ON THE

it runs back beneath the zygomatic process as far as the glenoid
cavity in the formation of which it takes a small share.”

There is little to add to accounts previously given relating to
the jugal arch in the Sirenia and Cetacea. In the Sirenia the
arch is massive, particularly in Manatus, and the jugal curves
deeply downwards in Halicore. The malar suffers little displace-
ment on account of the maxilla and it extends postero-ventrally
near the glenoid. In Janatus the jugal almost meets the frontal
behind the orbit. In the Cetacea the jugal joins the lacrymal
when the latter is present; and the separation of the orbit from
the temporal fossa is achieved as in the Equide by the union of
the squamosal with the frontal.

Winge™, in his recent revision of the Insectivora, lays stress on
the early separation of the Dermoptera, Cladobatide, and Macro-
scelidee from the other families of the order; but again overlooks
the morphological value of the zygomatic arch. For nothing
more emphatically distinguishes the Tupaiide (Cladobatide,
Winge) and Macroscelide than the relations of the Jugal. In all
the remaining families it is greatly reduced, so that the arch is
frequently incomplete owing to its total suppression as in the
Soricide, Centetide, Solenodontid, and Potamogalide. In the
Evinaceide, Talpide, and Chrysochloride, it is present asa slender
element inserted in the middle of the arch between the zygomatic
processes of the maxilla and squamosal; never does it join with
the lacrymal. In marked contrast, on the other hand, the relations
of the jugal in the Tupatide and the Macroscelide are essentially
sinilar to the condition characteristic of the Metatheria, Hyrax,
and certain Rodents: that is, it extends back to the glenoid
ventrally and reaches forward to meet the Jacrymal on the
anterior border of the orbit. In theaberrant genus Galeopithecus
the jugal is even more powerful and contributes a large share
in the formation of the glenoid cavity.

In the Chiroptera the zygomatic arch is complete; but the role
of the jugal is greatly restricted, as in the Erinaceide. The
maxilla displaces the jugal, not as is usually the case on the
ventral side, but dorsally ; and sometimes comes into contact with
the squamosal element on the upper border of the arch.

The enclosure of the eye-socket, which has been independently
eftected in certain of the more specialized members of other orders,
has become a fixed character in the Primates through the union of
the postorbital process of the jugal with the frontal, so that the
former has acquired a new role. The jugal does not usually
extend far behind its junction with the frontal in the Lemurs and
in the Anthropoidea; but in certain genera of the Prosimiz it
may extend back to the glenoid, as e.g, Chiromys, Indris, and
Tarsius. A more rigid distinction—not previously mentioned—
between the Lemuroidea and the higher Primates than the
complete enclosure of the orbit by the ingrowth of the jugal to

* Winge. ‘ Udsigt over Insektaedernes indbyrdes Slaegskab.”’ WVidensk. Meddel.
f, Dansk Naturhist. Forening Kjébenhavn, 68. 1917.

REDUCTION OF THE JUGAL IN MAMMALIA. 77

meet the alisphenoid, is to be found in the fact that the jugal
articulates with the lacrymal in the former but not in the latter
group. It is frequently stated, even by such authorities as Weber*,
that the Catarrhina are distinguishable from the Platyrrhina by
the junction of the jugal with the parietal in the last named: but
this is not by any means a universal rule. In Mycetes the jugal
does not meet the parietal ; and a specimen of Ateles ater in the
Birkbeck College collection shows the jugal joining the parietal
on one side of the skull but not on the other.

A comparison of the skull in the Anthropoidea with that of
forms like Chiromys and Jndris suggests that even in the
Primates, in which the role of the jugal has become relatively
more important on account of its inclusion in the orbit, it has not
escaped the general tendency towards reduction by the invasion
of the squamosal or maxilla or both apparently developed in-
dependently along the various lines of mammalian phylogeny, as
indicated in the previous survey. It is without doubt true that
so far as the curvature and strength of the zygomatic arch is
concerned, its development, as Slade observes, ‘‘ depends upon the
energy and character of the masticatory process.” Nevertheless, if
the problem is approached in the light of modern evolutionary
theory with the mind less taxed with the necessity of discovering
utility in every structure, it is difficult to reveal any general
teleological significance in the part individually contributed by
the various elements of which it is composed. Seeing that in a
diversity of isolated genera among the Placentals exhibiting every
possible variety of diet and habit, and also in some of the less
specialized representatives of the larger groups themselves, the
jugal displays essentially the same relations as in the Metatheria,
namely, extending postero-ventrally from the glenoid to the
lacrymal antero-dorsally, it is hardly possible to agree with Weber
that the jugal was.small in the earliest Mammalia (op. cit.) as in
the Insectivora of to-day: on the contrary, there can be little
doubt that this represents the ancestral condition retained by the
class till a date later than that at which the modern lines of
Mammalian descent had become differentiated. In some cases the
jugal has become effectively eliminated although the arch is
eomplete and even robust, as in the Monotremest where the
malar is vestigial or absent, at least as a separate ossification.
The reason for this progressive reduction of the jugal quite
irrespective of the form and function of the arch, as is most
emphatically demonstrated by a comparison, for instance, of the
skulls of Hrethizon and /ydrocherus, remains obscure. Neverthe-
less, the manner in which it is effected among the different orders
of Mammalia is so characteristic and, in a sense, conservative that
similarity in the morphological relations of the jugal in Mammals

* Weber, Max. LEinfiihrung in die Anatomie und Systematik der recenten und
fossilen Mammalia. Jena, 1904.

+ Vide Sixta: “Untersuchung iiber den Bau des Schaedels von Monotremen und
Reptilien,” Zeitschr. f. Morph. und Anthropol. ii., 1900,

78 ON THE REDUCTION OF THE JUGAL IN MAMMALIA.

that exhibit different habits of nutrition should prove of greater
service for the elucidation of phylogenetic problems and taxo-
nomical difficulties than has hitherto been recognized.

A similar shrinking of the postorbital due to genetic factors at
present unknown would appear to account for the disappearance
of that bone in the Theromorpha. For clearly the suppression of
the postorbital cannot be assigned to any teleological reason, since,
as has been indicated above, the separation of the orbit from the
temporal fossa has been reacquired— presumably to meet an
evident physiological need—by entirely different devices in several
orders of modern Mammalia independently.

ON TWO NEW LIZARDS AND A NEW FROG FROM COLOMLIA. 79

8. Descriptions of two new Lizards and a new Frog from
the Andes of Colombia. By G. A. Bounencrr, F.R.S.,

ie Zas
[ Received March 7, 1919: Read April 8, 1919. ]

(Published by permission of the Trustees of the British Museum.)
(Text-figures 4 & 5.)

A collection of Reptiles and Batrachians from Bogota and
neighbourhood, sent to me for identification by Bro. Apollinaris-
Maria, of the Instituto de la Salle, Bogota, contained examples of
three new species, of which descriptions are here given. The
types are preserved in the British Museum.

ANOLIS APOLLINARIS, Sp. n.

Head nearly once and a half as long as broad, scarcely longer
than the tibia; forehead concave; no frontal ridges; upper
head-seales rather small, strongly keeled ; supraorbitals enlarged,
separated by two or three series of scales ; occipital not enlarged ;

Text-figure 4.

Anolis apollinaris.

canthus rostralis short and strong; canthal scales four; loreal
rows five; 9 or 10 upper and as many lower labials; ear-opening
oval, half the diameter of the eye-opening. Body compressed ;
dorsal and lateral scales very small, granular, strongly keeled,
larger on the vertebral region; ventral scales feebly keeled,
larger than largest dorsalis. The adpressed hind limb reaches
the eye; digital expansions strong; 25 lamelle under phalanges
II and ITI of the fourth toe. Tail cylindrical, covered with small
strongly keeled scales, its length more than twice that of head

80 MR. G. A. BOULENGER ON TWO NEW

and body. Dark olive above and on the sides, with a fine
blackish network; head and a vertebral band pale, the latter
with narrow transverse processes ; small round light spots on the
sides and tail; fore-arm, tibia, and lower parts pale green.

Watal deme thier. pens. .... 396 mm.
(15 Bete [ae eer ae ea te 26
IVa) Or end steerer 17
BOG Gat ermect nears 80
Hore slimmest eee 46
Helsing lange ioe eer 81
TDA kere gee ee meee ae 25
HDS [Sem ReMi te eee = Ae | eta ccaineaal 290

A single female specimen from near Bogota.

PROCTOPORUS BOGOTENSIS, Sp. n.

Head rather large, with obtusely pointed snout and swollen
temples; limbs moderately developed, meeting when pressed
against the body. Frontonasal as long as broad, as long as and
broader than the frontal, which is twice as long as the suture

Text-figure 5,

Proctoporus bogotensis.

between the frontoparietals and a little shorter than the inter-
parietal; latter widening posteriorly, nearly the same length as
the parietals, which form an oblique suture with the large upper
temporals ; a pair of large occipitals; four supraoculars, three in
contact with the frontal; nasal divided ; no loreal; a series of sub-
orbitals; three upper labials to below eye; one anterior and three
pairs of large chin-shields, the first two pairs forming a suture;

LIZARDS AND A NEW FROG FROM COLOMBIA, 81

_ 9 vows of scales between the latter and the collar; 8 collar-shields.
Dorsal scales twice to twice and a half as long as broad, tri-
earinate, forming uninterrupted series across the back ; 31 scales
from occiput to sacrum inclusively. Ventral scales, except the two
outer rows on each side, as long as broad or a little longer than
broad, in 10 longitudinal and 23 transverse series. 32 scales round
the middle of the body. 5 large preanal shields (2). 6 femoral
pores on each side. Tail thick, scales as on the body, but dorsals
shorter and ventrals narrower. Brown above, with five black
longitudinal streaks beginning on the head; throat white, spotted

-with black; belly black, with small white spots.

Total length (tail reproduced) ...... 115 mm.
ECL od WI on ee ae ten Meyer inventory asIAS 13
Wircitihpotelneadhemeemaeuce as secs neta 9
From end of snout to fore limb.. ... 20

: As GMM casa. Seashee 60
Tittonbes Invenio} Benet oBacend cease nares 16
TB bina diitrnhO eas ae Renan nanmoseesooe hates 19

A single male specimen from Bogota.

HYLIXALUs GRANULIVENTRIS, Sp. n.

Tongue distinctly nicked behind. Head as long as broad ;
snout rounded, projecting beyond the mouth; canthus rostralis
distinct ; loreal region feebly oblique, feebly concave; inter-
orbital region as broad as the upper eyelid; tympanum distinct,
half the diameter of the eye. Fingers moderate, first and second
equal, the tips dilated into small discs; subarticular tubercles
well developed. Toes half-webbed, the web extending as a fringe
to the discs, which are larger than those of the fingers; sub-
articular tubercles feebly prominent; an oblique fold on the
distal half of the tarsus, extending along the inner side of the
first toe ; an oval inner and a round outer metatarsal tubercle.
The tibio-tarsal articulation reaches the anterior border of the
eye; tibia a little less than half the length of head and body.
Upper parts smooth, with a few small, feebly prominent warts on
the head and back; throat and belly strongly granulate. Upper
parts and throat blackish; belly and lower surface of limbs
white. Male with an internal vocal sac.

From snout to vent 22 millim.

A single male specimen from Bogota.

Proc. Zoou. Soc.—1919, No. VI. 6

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ASYMMETRICAL DUPLICITY IN THE CHICK. 83

9. A Unique Case of Asymmetrical Duplicity (Dupli-
citas asymmetros) in the Chick. By Nort Tayier,
B.Se. (Lond.)*. (From the Zoological Department,
University of London, University College.)

[Received April 8, 1918: Read May 13, 1919.]

(Plates I-III. f and Text-figures 1 & 2.)

INDEX. Page
I. Inrropucrory Nore... sae Be enormennnccactec date:
Il. Descriptive. The mor eonoleay of Binstodenm EB erpncnchis cee eee iam OAL
A. General appearance of the Blastoderm and orientation ......... 84
B. Detailed description of the relation of the two embryonal
formations and the structure of the lesser (Rudiment 9) .. 85
C. The morphology of the Beene: embryonal formation (Rudi-
ment @)| Mccain eseesaceee 86
D. Summary of ihe Sacipall mor mpliclogieal Pimonan Wis ae the
greater embryonal formation (Rudiment @)........................ 91
HIT. Discusston .......... seat ceutisemmeseMaare MO
Part A. The Pranplincenects of ualyaonel for ieconte (0s cir a ied beeen fo)
1. The condition of the gut... ABS SSIS 5 LEE ISS TOS
2. The condition of the heart- primordia sins . «94
3. The anterior portions of the fore-gut and vascular system | eee
Part B. The origin and relationship of the two embryonal rudiments... 98
IV. TeRatToLoGicaL SIGNIFICANCE AND CONCLUSIONS ......... = LOZ
1. Bearing on the nature of unequal monstrosity, “ Dekeiie
asyimmetros ” pach sea sets seoake ora deer omer ees ee eee ate tea LOD
2 A Conelisinnsiwe sieeve es ee ein ol tenella he LOG,

I. Inrropuctory Nore.

The following paper consists of two main parts, firstly the
description of the blastoderm, and secondly a discussion as to its
mode of origin and development. Its bearing on teratological
problems is also very briefly considered.

Hitherto little attention has been paid to the abnormal products
of morphogenesis, largely, no doubt, owing to the scantiness of our
knowledge of the causal factors of development; nevertheless, it
would seem that, in view of the important bearing that the
abnormal must have in the interpretation of the normal, it is
deserving of more attention than it has hitherto received. It was
thought, therefore, that it might be quite worth while to take
such a case as the present and subject it to a somewhat detailed
consideration from two aspects. Firstly, with respect to the
morphogenetic problems which it might itself present, considered
in relation to what is known of the factors of normal development,
and secondly with respect to its own bearing on our knowledge
of the latter and on some of the general problems of teratology.

* Communicated by Prof. J. P. Hux, F.R.S., F.Z.S.
+ For explanation of the Plates see p. 109.
6*

84 MR. NOEL TAYLER ON A UNIQUE CASE OF

I have been unable to find any described instance of any
blastoderm closely resembling it, and it appears of some interest
in connection with several morphogenetic questions, in particular
with the problem of the factors concerned in the early develop-
ment of the anterior portion of the embryo and also with that of
the localisation of the prospective embryo upon the blastoderm.

It also has a bearing on at least two teratological problems,
the mode of origin of double monstrosity, and the nature of that
particular and little studied type of duplicity in which the two
components are unequally developed—the Duplicitas asymmetros
of Schwalbe.

IJ. DEscRIPTIvE.

The Morphology of Blastoderm E, and the embryonal formations.

A. General appearance of the Blastoderm and orientation.

The blastoderm described in the following pages was very kindly
placed at my disposal some time ago by Prof. J. P. Hill, F.R.S.,
to whom I wish to express my thanks for his assistance in reading
and criticising this paper. When the blastoderm came into my
hands it was in an unstained condition, and after being stained
with borax carmine and examined as a whole mount it was cut
into serial sections for more detailed study.

Unfortunately I am unable to state the time of incubation, of
which there is no record.

In the first place I shall give a brief description of the gross
morphology of the blastoderm, and then after reference to the
structure of the lesser rudiment and to the relations of the two
embryonal centres as determined from a study of the sections,
shall proceed to a description of the morphology of the greater
embryonal rudiment.

Pl. L. fig. 1 is a reproduction of a photograph of the blastoderm,
which it will be seen is oval in shape, 12 mm.in length by 7 mm.
in breadth.

Situated in the median longitudinal axis of the blastoderm is
the more completely developed formation, which will henceforth
be designated as Rudiment a or alternatively the greater or
major embryonal formation. It was found to possess 17 pairs of
mesoblastic somites, and is normally developed as regards its
posterior region.

In Pl. I. fig. 3 the abnormality of the anterior region, which
is chiefly evident in the position of the anterior intestinal portal
and in the condition of the Fore-Brain, is apparent.

In the anterior prolongation of the long axis of Rudiment a
and situated rather over 2 mm. from the anterior limit of its
Fore-Brain, is the lesser or minor embryonal formation which
will be designated in the sequel as Rudiment j3 (see Pl. I. fig. 1).

This lesser embryonal formation may be concisely described as

ASYMMETRICAL DUPLICITY IN THE CHICK. 85

a primitive-streak-like rudiment, and its longitudinal axis as
defined by its “primitive groove ” is, with reference to the con-
tinuation of the longitudinal axis of Rudiment a, bent to the
left at an angle of about 30°.

Between the two embryonal formations there runs what appears
in the whole mount as a more darkly staining strand (PI. I. fig. 1)
which will be more minutely described in the sequel.

The longitudinal axis of Rudiment a will be used as a basis
for the orientation of the whole blastoderm ; thus, when we speak
of the posterior end of the Rudiment 6 we imply that portion
of it situated proximally or nearest to Rudiment a.

B. Detailed description of the relations of the two Embryonal
Formations and the structure of the lesser (Rudiment /).

Reference may now be made to the structure of the lesser
rudiment, which is more properly described as “ primitive streak-
like” than as a primitive streak.

It will be seen from the figure that it consists of two main
parts, a rather diffuse and somewhat darkly staining oval area,
and the more deeply staining primitive streak proper (PI. I. fig. 1,
Rud. (2) with its more translucent primitive groove.

The dimensions of the dark oval area, which the sections show
to consist of thickened mesoblast, are about 78u x 64, while the
length of the “ primitive groove” is about 28u.

Pl. IIL. fig. 2 represents a section of Rudiment (3 in the region
of the primitive groove. It will be seen, that it is quite com-
parable to the sections of a rather advanced primitive streak.

The three primary germ-layers are well defined and separate
except in the region of the “primitive groove” (Pr.g.) where
the ectoderm and mesoderm are in continuity. One receives the
impression from these sections that there is, in this region, a well
marked proliferation of the median ectoderm as in the normal
primitive streak.

Peripherally the mesoderm splits into somatic and splanchnic
layers on either side (PI. II. fig. 2, Som.mes., Sp.mes.), the ccelom
thus formed being portions of the parietal cavity.

The median unsplit mesoderm extends back from the region
of the ‘‘ primitive groove ” for some distance as will be seen from
Pl. I. fig. 1; in section, it is seen to become free from the super-
imposed ectoderm, and this fact suggests the question as to
whether or not we might be justified in regarding it as of the
nature of a head-process. If we are so justified, it would become
necessary to regard Rudiment (6 as orientated in the reversed
direction to Rudiment a, 7. e., the two embryonal formations
would be placed head to head (-><—) and not in the same
direction (—> —>).

It may be noted that cases have been recorded of two opposing
embryonal formations placed upon a single germinal area. Such
a case was described by Mitrophanow (8), in which the two

86 MR. NOEL TAYLER ON A UNIQUE CASE OF

embryonal rudiments were represented by two primitive streaks
about equally developed; while other and older stages have been
recorded by Kaestner and others (6, 7).

Under the circumstances however, the question does not seem
one of great importance, and it seems better to refrain from any
definite decision as to the orientation of Rudiment 3.

No structure seems to be present which can undoubtedly be
regarded as a “ primitive knot.”

“We may now turn to the morphological relationships of the
two embryonal formations. These two structures occurring in a
single blastoderm, the three primary germ-layers of each and
those of the extra embryonal portion of the blastoderm, are of
course continuous each with each. It may also be noticed that,
whilst the sinus terminalis is perfectly regular in outline, the area
pellucida shows signs of division into two regions of greater trans-
parency (see P]. I. fig. 1) in relation to the two embryonal forma-
tions, since the blood islands of the vascular area are concentrated
somewhat in the regions posteriorly and postero-laterally to
Rudiment 3. It is of more importance, however, to find that
there is what may be described as a definitely differentiated
morphological connection between the two embryonal formations,
viz., the rather darkly staining and irregular strand which in the
photograph of the whole mount (PI. I. fig. 1) appears to run in the
long axis of the blastoderm between the two embryonal rudiments.
In the sections this is found to consist of a narrow and somewhat
irregular band of unsplit mesoderm, thicker and broader in the
region of the Rudiment (3 and becoming more tenuous and
irregular as it comes into proximity to Rudiment a.

Its somewhat irregular and broken appearance in the whole
mount is seen from tke sections to result from the presence of
vessel-like lacune in its substance, spaces, however, which are
devoid of blood corpuscles.

On either side of this median strand the mesoderm splits into
somatic and splanehnie layers, and it would seem that the extra-
embryonal ecelom situated laterally to Rudiment (6 and to the
mesodermal strand is continuous with the parietal ccelom of
Rudiment a.

C. The Morphology of the greater Embryonal Formation
(Rudiment a).

Proceeding to a consideration of the structure of Rudiment a,
the morphology of the posterior region may be briefly dismissed
as it is that of a normal embryo of 17 somites, PI. I. fig. 3 and
text-fig. 1 represent a semi-dia grammatic figure of the morphology
of the anterior region, while transverse sections thr ough the more
important regions are shown in Plates II. and III. figs. 4 & 5.
The positions of these transverse sections are indicated in text-
fig. 1.

"In section D, p. 91, is given a concise > summary of the more im-
portant morphological abnormalities which occur in Rudiment a,
and this is subdivided into four categories, viz., those affecting

ASYMMETRICAL DUPLICITY IN THE CHICK, 87

the general proportions of head, the nervous system, the gut, the
vascular system, and celom,

Text-figure 1,

Diagram of a longitudinal horizontal section.

a. The general proportions of the anterior end and its relations to
the extra-embryonal portions of the blastoderm.

There is a complete absence of any anterior amniotic fold, as
is shown in the sections, Pl. III. fig. 4, Hl and H2, The lateral
limiting sulci are fairly well marked (lat.l.sul.). There is no
head-fold and consequently no anterior limiting sulcus; the ecto-
derm of the anterior end of the head of the embryo is continuous
with that of the blastoderm immediately in front of the head
without being recurved back under the latter, so as to delimit
it from the blastoderm as is normally the case at this stage of
development.

Whilst the cranial flexure is, in a normal chick of 15-17 somites,
well marked, consisting of a bending of the antero-posterior axis
of the head, and there has also occurred its partial revolution
about the dorso-ventral longitudinal plane, Pl. I. fig. 3 indicates
at once that the antero-posterior axis of the head of Rudiment a
is perfectly straight, while the sections show that there is little
sign of any rotation (Pl. III. fig. 4, H1-E 5). The slight obliquity
of the dorso-ventral plane, which is seen in section E 4 (fig. 4),
is in the reverse direction to normal, and is no doubt the con-
sequence of the morphological asymmetry of the head which has
yet to be described.

Reference must now be made to the general dimensions and

‘88 .MR. NOEL TAYLER ON A UNIQUE CASE OF

proportions of the head, features of considerable theoretical
interest. Pl. I. figs. 1 and 3 show well what we may briefly
characterise as the “shortness” of the head, and they further
emphasise the fact that in Rudiment a the brain does not play
the important part in moulding the general contours of the head,
which is the case in the normal chick, and which is largely
consequent upon the considerable dilatation of the brain-vesicles.

This abbreviation of the anterior region of Rudiment a is due
to two main causes, viz., the under-development of the Fore-
Brain and the condition of the heart and fore-gut.

In the normal chick of 16-17 somites the anterior intestinal
portal is situated some distance posteriorly to the auditory vesicles.
In Rudiment a, on the contrary, the anterior intestinal portal
is anterior to these (PI. 1. fig. 3 and Pl. II. fig. 5, KE 8, 7. and
r.aud.ves.).

Now it is obvious that to describe the facts in this way is really
to beg the question, for one is assuming that the auditory vesicles
are normal in position.

An alteration in the relative positions of auditory vesicles and
intestinal portal might result either from an abnormal develop-
ment of the brain (for the auditory vesicles are normally placed
in relation to the neuromeres of the hind-brain) or from an actual
displacement of the anterior intestinal portal (¢.e., an under or
over-development of the fore-gut), or it might be associated with
a combination of both these factors.

By comparison of the measurements of Rudiment a with those
of normal embryos, it seems clear that in the case of the former
each of these factors plays a part in the production of the
condition described.

Measurements of Normal Embryos.

No. Ent. (Length (Length
Designation. Somites. Length. Ant. End—Aud. Ves.) Ant. Bada. I.P.)
IN GY eee 16 o’5 mm. 1°6 mm. 2-1 mm.
INGE penne 16 6 mm. 1-5 mm. 21 mm.
ING Sire 15 6 mm. 1-4 mm. 1-9 mm.
INA ure , 17 6 mm. 1-4 mm. 2 mm.
Mean of Measurements of Normal Embryos.
5-9 mim. 1:47 mm. 2:02 mm.
(nearly)
Measurements of Embryo a of Blastoderm E.
16 6 mm. 1:3 mm. 1-2 mm.

These measurements are reproduced above, and they require
but little comment. The mean of the measurements taken from
the extreme anterior end to the plane of the auditory vesicles is
just under 1°5 mm., while the mean of the measurements from
the anterior end to the anterior intestinal portal is 2°02 mm. In
the case of Rudiment a the former measurement is 1°3 mm., the
latter 1-2 mm.

Hence the intestinal portal is situated no less than *8 mm.
more anteriorly than in the normal embryo, while the length of

ASYMMETRICAL DUPLICITY IN THE CHICK. 89

the head, as defined by the distance between its anterior end and
the auditory vesicles, is barely ‘2 mm. less than normal.

It is therefore clear that the abbreviation of the head of
Rudiment a is largely apparent, resulting from the abnormally
anterior position of the intestinal portal; but is in part real,
since the head, as more strictly defined by the position of the
auditory vesicles, measures ‘2 mm. less than normal.

b. The nervous system.

The nervous system is, posteriorly to the auditory vesicles,
quite normal, its development only becoming abnormal in the
anterior region.

All the chief divisions of the brain are clearly recognisable
(Pl. I. fig. 3; Pls. IT. & IIT. figs. 4, 5), and the five neuromeres
of the Hind-Brain are quite distinct. Cranial flexure being
absent, the neural axis is, of course, straight.

The degree of abnormality increases as we pass _ postero-
anteriorly and is greatest in the Fore-Brain. This abnormality
is of two kinds: firstly, a general under-development of the Fore-
Brain, and, to a less extent, of the Mid-Brain; secondly, a
definite bilateral asymmetry of the Diencephalon, evident pavrti-
cularly in the optic evaginations.

The Mesencephalon appears somewhat compressed in the
sagittal plane. The Telencephalon is very small and its cavity
relatively minute. (PI. I. fig. 3 and Pl. IIL. fig. 4, H 1 and E 2,
Tel.).

The asymmetry of the Diencephalon was a striking feature of
the whole mount (PI. I. fig. 3, Dien.) which gave the impression
that the right optic evagination is entirely wanting, the left being
relatively well developed and almost making contact with the
ectoderm. Reference to the sections through this region (PI. III.
fig. 4, E3 and E4) shows that the cavity of the Diencephalon is
extremely irregular in shape. The left optic evagination is well
marked (l.op.). The Diencephalic wall in the corresponding
position on the right side shows no such definite evagination,
though dorso-laterally and ventro-laterally it is bent outwards
somewhat suggesting commencing evaginations. The more dorsal
outbulging, which is the larger of the two, would seem to represent
the right optie vesicle, but if so it is very much less developed
than the left, the brain in this region being markedly asymmetrical
(fig. 3). A median ventral downgrowth of the diencephalic floor
is well marked even in the whole mount (fig. 3, infidep.) ; in trans-
verse section (fig. 4, E 4, infdep.) it is seen to approximate to a
median dorsal thickening of the gut-entoderm. It is probably
infundibular in nature. Further anteriorly the floor of the
diencephalon becomes very thick (PI. ITT. fig. 4, E 3).

ce. The fore-gut.

Owing to the anterior position of the intestinal portal the fore-
gut is relatively very short. Throughout its length it is dorso-
ventrally compressed. The hyoid pouch almost makes contact
with the ectoderm (PI. 11. fig. 5, E 7, l.My.P.).

90 MR, NOEL TAYLER ON A UNIQUE GASE OF

The condition of the anterior portion of the gut is of extreme
interest, and here, as in the case of the other organ-primordia,
deviation from the normal is very great.

In the first place is to be noted the apparent absence of any
sign of an oral plate, the formation of which, in the absence of
any head-fold, is of course impossible. The gut is remarkable
by reason of its extension forward to the prechordal region (see
Pl. I. fig. 7B). The anterior extremity of the chorda comes into
relation with the dorsal wall of the gut, which latter then effects
contact with the well-developed infundibular downgrowth of the
Diencephalon (fig. 7 B, inf.dep.); this contact extends for some
distance, throughout which the mid-dorsal wall of the gut is
markedly thickened, showing some signs of cellular proliferation.

The most surprising circumstance is, however, that in front of
this point, at which one would expect the gut to terminate, it
extends for a considerable distance ventrally to the Telencephalon
(Pl. III. fig. 4, E2, and Pl. I. fig. 7 B, Ant.eaxt.¥.@.), from which
it is separated by a large vessel formed by the union of the two
apparently dorsal portions of the mandibular aortic arches. It
is only in the extreme anterior region of the Telencephalon
(Pl. III. fig. 4, E1) that the fore-gut has disappeared.

ad, Heart, vascular system, and celom.

The Heart, as seen in the whole mount, is characterised by its
relative shortness and broadness (PI. I. fig. 3, Ht.). It presents an
outwardly-directed convexity on either side. The sections show
that the two endocardial tubes have completely failed to fuse,
and that they are, for the greater part of their extent, widely
separated. Only immediately in front of the anterior intestinal
portal do they even approximate in the mid-line, and even here
no fusion, sensw stricto, has occurred. The right heart-tube is
somewhat shorter, and for the greater part of its extent its
diameter is considerably less than that of the left (Pl. ITI. fig. 5,
E5, and Pl. II. fig. 5, E 6).

It will be seen that the condition of the heart resembles that
which is sometimes described in an otherwise normal chick under
the name of “ double-heart,” and which was elucidated by Camille
Dareste (2), who showed it to result from the failure of the two
heart-tubes to fuse as normally occurs.

Tracing the heart-tubes anteriorly they do not fuse beneath
the ventral wall of the gut to form the ventral aorta as in the
normal chick, but in the region of the infundibulum, where they
have become very small, they become continuous laterally with
two large vessels lying on either side of the upper surface of the
gut (as will be seen in the sequel, it is probable that these two
dorso-ventral vessels running on either side of the gut are not
morphologically the mandibular arches as one would at first sight
suppose).

Anteriorly these two vessels (the right of which is the larger)

ASYMMETRICAL DUPLICITY IN THE CHICK, 91

pass round the median infundibular-gut junction (E 3) and fuse
to form a large median aortic space lying between the Telen-
cephalon and the prechordal prolongation of the gut (PI. IIT.
fig. 4, E.2, Ant.cor.Sp.). Posteriorly the two vessels (Pl. ITT.
fig. 4, E 3, R.Lat.aor.Sp., L.Lat.aor.sp.) become continuous with
the dorsal aorte: (E 5-E6, Dor.Aor.), which pass back on either
side of the chorda in the normal manner.

The myocardium is characterised by its width; in front of the
anterior intestinal portal there is a ventral myocardial septum.

With respect to the pleuro-pericardial ccelom, this is remark-
able for its extreme forward extension immediately beneath the
einbryo, the lateral amniocardiac vesicles opening into each other
and forming a median ccelomic space which extends in front of
the anterior limit of the embryonal formation proper. This
forward extension is permitted by the absence of the head-fold
(Pl. ILI, fig. 4, H1 and E 2, Ant.p.p.c.; text-fig. 2, Ant.p.p.c.),
and in the sequel it will be seen to be a point of considerable
morphogenetic significance.

Text-figure 2.

Transverse section through the extreme anterior region of embryonal
formation «,

For explanation of letters see p. 109.

D. Summary of the principal morphological abnormalities of the
greater Hmbryonal Formation (Rudiment a).

A. General proportions of anterior region and its relations to
the extra-embryonal portions of the blastoderm.
I. Absence of an anterior amniotic fold.

Il. Absence of a head-fold.

Ifl. Absence of cranial flexure.

IV. Abbreviation of head proper as indicated by the distance
between the anterior end and the plane of the auditory
vesicles.

92 MR. NOEL TAYLER ON A UNIQUE CASE OF

B. The nervous system.

I. Smallness and compression of the anterior two divisions
of the brain, the Telencephalon and the Diencephalon.
TI. Reduction of the optic evagination on the right side.

C. The gut.

I. Its shortness, in consequence of the position of the
anterior intestinal purtal.
II. Absence of an oral plate.
III. Thickening and proliferation of entoderm of the gut-roof
contiguous to the infundibulum.
TV. The abnormal prechordal prolongation of the gut.

D. Heart, vascular system, and ccelom.

J. No fusion of heart-tubes.
II. Small size of right heart-tube.
III. Median aortic space beneath the Telencephalon.
IV. Width of pericardium.
V. Anterior extension in mid-line of pleuro-pericardial
ceelom.

Ili. Discussion.

Part A. The Morphogenesis of Embryonal Formation a.

It would appear advisable, before proceeding to a considera-
tion of the ultimate origin of the two embryonal rudiments
upon this blastoderm, to discuss in some detail the morphology
of Embryonal Formation a, the more completely developed of
the two rudiments.

Such a discussion will, I believe, result in fairly definite conclu-
sions as to the primary factors responsible for the abnormal devel-
opment of Rudiment a, and indirectly to the causes responsible
for the presence upon the blastoderm of the second Embryonal
Rudiment (().

There appears to me ample justification for the view that many
of the abnormalities of Rudiment a, particularly those affecting
the vascular system and gut, are secondary, in the sense that they
are consequent upon an inhibition of the normal development of
the extreme anterior portion of the embryo, in particular of the
medullary plate region.

For this abnormal arrest of the growth of the anterior region
of the medullary plate the only adequate explanation would
appear to be the existence of some germinal defect, which would
therefore be the primary modification in the sense indicated
above.

That this is indeed the case seems the only view consistent
with the occurrence of the following abnormalities affecting the

ASYMMETRICAL DUPLICITY IN THE CHICK. 93

anterior portion, particularly the medullary plate region of the
embryo :—

a. The absence of a true head-fold.

paignages 2 », an anterior amniotic fold.

Cot “5 5, cranial flexures.

d. The shortness of the brain resulting mainly from the
under-development of Telencephalon and Diencephalon.

e. The condition of the optic vesicles.

The assumption will therefore be made in the following
pages that the primary factor responsible for the abnormal
development of Rudiment a was this abnormal arrest of the
growth of the anterior portion of the medullary plate, and the
attempt will be made te demonstrate how the other structural
modifications are necessary but purely secondary consequences cf
this initial disturbance of the ‘ causal harmony of development ”
(Driesch).

1. The condition of the gut.

In the first place, we may consider the condition of the fore-
gut, particularly with reference to the anterior position of the
anterior intestinal portal.

It may be recollected that in the normal chick two main
factors are responsible for the formation of the fore-gut :—

1. A definite forward growth of the brain.

2. The backward growth of the posterior margin of the
splanchnopleuric head-fold owing to pleuro-pericardial
ceelomic expansion.

It is generally agreed that the initiation of fore-gut formation
is due to the development of the head-fold, which latter itself is
due to the more rapid growth of the anterior part of the
medullary plate.

With regard to the precise importance of these two factors
there has been considerable difference of opinion, and it may
now be accepted as established that the mechanism of fore-gut
formation differs considerably in the various groups of verte-
brates. With regard to the condition in the chick embryo,
Robinson (18) considered that the former factor was that mainly
if not solely operative. Other authors (Graper, Rouviére, 4, 14)
have contended that there occurs also a backward growth of the
anterior intestinal portal, and it may be regarded as established
that under normal conditions both these factors play a part in
the formation of the fore-gut in the chick (a view which it will
be seen is supported by a consideration of this embryo). On the
other hand, in the marsupial Dasywrus, Miss Parker (11) has
shown that the second factor plays a far more fundamental réle.

Since in the major rudiment of blastoderm E the former factor
was largely absent, it is not surprising that a condition of affairs
has arisen analogous in many respects with that in Dasyurus,

94 MR. NOEL TAYLER ON A UNIQUE CASE OF

where, according to Miss Parker, the initiation of head-fold
formation is due to the forward growth of the brain-plate, whilst
the expansion of the pericardium is the main factor responsible
for the extension of the fore-gut.

A feature which is of undoubtedly great interest is that in an
embryo in which there can have been very little active forward
growth (7. ¢., in contra-distinction to general or interstitial
growth) of the medullary plate, relatively to the line separating
embryonal and extra-embryonal areas, which notably Robinson
considered as the main factor concerned in fore-gut formation,
there should be present a well-marked gut extending anteriorly
up to the anterior limit of the Telencephalon.

That a true head-fold could never have been present is fairly
evident from the sections (Pl. III. fig. 4, H1, H2, E3), and
accordingly it may be concluded that the second factor (pleuro-
pericardial coelomic expansion) must have been that which was
mainly if not solely operative in the formation of the entire
extent of the fore-gut. This is borne out by the presence of the
pleuro-pericardial ccelom, separating somatopleure and splanchno-
pleure in the extreme anterior region of the embryo, in front
even of the anterior limit of the gut itself (fig. 4, E1, and text-
fig. 2), a condition which is not found, of course, in the normal
chick embryo. It is of importance, too, to note that the slight
inturning of the ectoderm beneath the extreme tip of the embryo
does not extend to anywhere near the anterior limit of the gut
(fig. 4, E1 & E2; Pl. I. fig. 7 B).

It is probable, therefore, that in this embryo even the initiation
of fore-gut formation was due to the formation of the pleuro-
pericardial celom. There must have been an extension in
towards the middle line, and a subsequent union of the amnio-
cardiac vesicles, thus producing a condition somewhat similar to
the pleuro-pericardial canal of mammals. Expansion of the
pleuro-pericardial cavity so formed in the posterior direction
would result in the formation of a splanchnopleuric or cardiac
fold and a fore-gut.

Now it will be recollected that the apparent length of the
fore-gut as determined by the position of the anterior intestinal
portal is 1:2 mm., as compared with a normal length of about
2°1mm. This seems to indicate that while in normal morpho-
genesis the active forward growth of the brain plays a part in
the increase in length of the fore-gut, the absence of this factor
in specimen E must have influenced unfavourably the action of
pleuropericardial expansion, resulting in an unduly slow pro-
gression backwards of the cardiac fold, and an abnormally short
fore-gut, which is therefore a secondarily induced condition
resulting from the primary abnormality of the medullary plate.

2. The condition of the heart-primordia.
The earlier processes in the development of the heart stand
in intimate causal relationship to fore-gut formation, and one

Se eer

ASYMMETRICAL DUPLICITY IN THE CHICK. 95

naturally turns therefore to a consideration of the condition of
this organ.

The two heart-tubes lie side by side beneath the fore-gut, and
are widely separated for the greater part of their length, actual
fusion having occurred at no point. No doubt this condition
results from the shortness of the fore-gut and the correlated great
width of the pericardium (Pls. II. & III. fig. 4, E5, fig. 5, E 6, E 7),
since this is clearly in harmony with the view expressed by Miss
Parker (11) that the approximation of the heart-tubes is due to
the growth in length and decrease in width of the pericardium
which, under normal circumstances, occurs at this stage.

In this connection the phenomenon of so-called “double heart,”
which sometimes occurs in otherwise apparently normal embryos,
seems of interest. In the few examples of this condition which
I have had the opportunity of examining, the “ duplicity ” of the
heart, really, of course, a non-fusion of the two heart-tubes,
has been associated with an unusually anterior position of the
cardiac fold, a condition which we should expect to be always
present if the above view as to the cause of the approximation
and fusion of the heart-tubes be true.

It has been mentioned that in this specimen the right heart-
tube is of considerably less diameter than the left, while in the
normal chick the reverse is the usual condition; it has been
suggested by Miss Parker (11) that this condition is a precocious
indication of heart-curvature, the lesser limb becoming the
concave border of the heart. With reference to the apparent
anomaly of specimen HE, it may be recalled that (a) occasion-
ally in the chick, flexure of the heart is to the left instead of to
the right; (6) that the condition here is possibly a functional
or physiologically induced asymmetry, which seems to be the
more likely explanation.

In this connection it is of interest to note that Wilder (19)
has pointed out that a second cause of “deformity,” at least as
regards bilateral symmetry or equality of components, and one
that is specially operative in assisting the secondary deformation
of a diplopage, is found in the striving among the parts during
growth for the best physiological efficiency. This is particularly
notable in the unequal degree of development often seen in the
duplicate systems of organs (particularly those functional from
an early period, e.g., the circulatory and digestive systems)
possessed by a diplopage.

This inequality of the heart-tubes may perhaps be regarded,
then, as an instance of the capacity of the various organs of the
body to undergo morphological changes in response to altered
physiological conditions, comparable, say, with such a common
pathological condition as the cardiac hypertrophy and dilatation
induced by valvular incompetence. The inducing cause would be
the primary asymmetry of the head and fore-brain (see Pls. I.
and IT. figs. 3, 4, H3 & E4), owing to the under-development of
the fore-brain and optic evagination on the right side.

96 MR. NOLL TAYLER ON A UNIQUE CASE OF

Congestion has taken place on this side, as is evinced by the
greater size of the right mandibular arch. The greater resistance
to the flow of the blood on the left side, owing to the smaller size
of the vessels, has led to a congestion in the left ventral aorta and
heart-tube, hence its greater diameter.

3. The anterior portions of the fore-gut and vascular system.

There yet remain for consideration the anterior portion of the
fore-gut and its relations to the central nervous system.

Firstly, with reference to the absence of an oral plate, a glance
at the figure of the longitudinal section (Pl. I. fig. 7 B) will make
quite clear the impossibility of a fusion between the anterior gut
entoderm and adjacent ectoderm such as normally occurs, since
here the pleuvo-pericardial cavities, which normally fuse in the
mid-line posteriorly to the oral plate (fig. 7 A, p.p.c.), in this
case extend to the anterior end of the fore-gut and to some con-
siderable distance in front of it (Pl. II. fig. 4, H1-E 4, Pl. 1. 7B,
p.p.c.), indeed for some distance in front of the anterior end of the
embryo itself: this is a definite median eceelom, formed presumably
by the fusion in the mid-line of the lateral ccelomic cavities.

There is, indeed, no sign of any thickening of the floor of the
fore-gut in the region where one would expect fusion between
entoderm and ectoderm, were it possible to take place.

It will be remembered, however, that there is a very marked
thickening and proliferation of the median gut-roof adjacent to
the infundibular region of the fore-brain (PI. III. fig. 4, H 3, H4,
Pl. I. 7 B), and if this entodermal thickening be regarded as the
protochordal plate complex, we can, I think, consistently inter-
pret the condition of the fore gut and its relations to nervous
and vascular systems with reference to the normal morphological
relations of these parts.

In Pl. I. fig. 7B is given a semi-diagrammatic reconstruction
of a median longitudinal section of the major embryonal formation
of Blastoderm E alongside, for the purpose of comparison, of a
similar sagittal section of the anterior region of a normal embryo
of about the same age (fig. 7 A).

It will be recollected that in the normal chick the entoderm of
the roof of the extreme anterior end of the fore-gut is some-
what thickened in the vicinity of the termination of the chorda
and the floor of the diencephalon, a slight diverticulum in this
region forming the so-called prechordal gut or Seessel’s pocket.
while an entodermal thickening situated ventrally comes into
relation with the ectoderm to form the oral plate. The depres-
sion of the oral plate beneath the ventral surface of the head is
increased as development progresses by the growth of the brain
and the occurrence of the cranial flexures. ‘The two lateral com-
ponents of the mandibular aortic arch meet in the mid-line just
behind the posterior limit of the oral plate to form the median
ventral aorta (fig. 7 A, X’).

ASYMMETRICAL DUPLICITY IN THE CHICK. oy

Now, on the assumption that in Rudiment a the dorsal
thickening of the gut-wall in the infundibular region represents
the protochordal plate, interesting deductions follow as to the
morphological relationships of the associated parts. In the first
place, it follows that that portion of the thickened gut-roof
immediately in front of the point where the latter makes contact
with the infundibular region of the Diencephalon must be
regarded as the equivalent of the entodermal moiety of the oral
plate of the normal chick: and further, that portion of the
thickened entoderm immediately posteriorly to this as the mor-
phological anterior limit of the gut, 7.e., the prechordal gut
(Seessel’s pocket) of the normal chick (PI. I. fig. 7 A).

If this interpretation be correct, we must conclude that the
singular anterior prolongation of the gut, which is well shown
both in the transverse and longitudinal sections (fig. 4, E 2, and
fig. 7B, Ant.ext..G'.), is morphologically (2. e., in reference to
the normal condition) a part of ventral gut entoderm, and that
that portion of the gut-roof which extends from the point of con-
tact with the infundibalum to its topographical anterior end Z
(fig. 7 B), and which is therefore apparently dorsal, is morpho-
logically ventral, being the equivalent of that part of the gut-
floor of the normal embryo which extends from the posterior
limit of the oral plate to, say, the point Z' (fig. 7 A).

The dispositions of the anterior portions of the aortic system
can only, it seems to me, be consistently interpreted on this
view. The median vessel between the telencephalon and the
anterior prolongation of the gut (fig. 4, E 2, Ant.aor.sp.) is
formed by the union of the continuations of the two dorsal
aortee which pass round the infundibular-gut connection ; ventro-
laterally it is continuous with the anterior continuations of the
two lateral heart: tubes.

Now there seems to be no room for doubt that this median
vessel X (fig. 7 B) must be regarded as morphologically the
median ventral aorta X' (fig. 7 A), the portions of doxsal aortze
which pass round the infundibular-gut connection to open into
it represent the mandibular arches, while a most significant fact
is the absence of any fusion between the anterior continuations
of the two heart-tubes on the apparent or topographical ventral
side of the gut.

The portion of the gut-wall with which this median aortic
space is in immediate relation, though apparently dorsal, corre-
sponds, it must be recollected, to that part of the entoderm
which forms a part of the floor of the gut of the normal chick.

These modifications of the normal morphology of the head no
doubt result from the absence of the normal growth zones of the
anterior end of the medullary plate and brain.

1. Primarily, the absence of normal head-fold focaetion
inhibited the possibility of any conjunction of the entoderm of
the fore-gut with ventral head-ectoderm to form an oral plate.

2. The presence of a considerable part of what should normally

Proc. Zoo. Soc.—1919, No. VII. 7

98 MR. NUEL TAYLER ON A UNIQUE CASE OF

have constituted the anterior floor of the gut, beneath the fore-
brain as a prechordal prolongation of the gut, resulted from the
fact that gut-formation was effected by the formation and expan-
sion of the pleuro-pericardial coelom alone and not by head-fold
formation. This is evinced by the anterior extension of the
median pleuro-pericardial cavity (fig. 7 B, Ant.p.p.c.).

3. The retention of this original condition was no doubt a
result of the absence of the normal expansion and flexure of the
fore-brain. This latter, in the normal embryo, it must be recol-
lected, conditions, largely, the depression of the oral plate beneath
the ventral surface of the head; and had the growth of the
fore-brain occurred in the usual manner in this case, no doubt it
would have tended (though it seems doubtful whether it would
have succeeded in effecting this) to drag round at least a por-
tion of the apparently dorsal wall of the pre-infundibular gut
(fig. 7 B, Ant.evt./.G.) into a ventral position.

Part B. The Origin and Relationship of the Two Embryonal

Rudiments.

There seem good grounds, then, for the conclusion that the
structure of embryonal Rudiment a is only explicable as resulting
from a very. early germinal defect, affecting in particular, it
would seem, the anterior medullary plate region. Turning now
to the anterior rudiment, designated in the descriptive part as
Rudiment 3, this can only be described as primitive streak-like,
so far does it deviate from the normal. There would seem here
to be no question but that it must have taken origin from a very
defective germinal centre compared with that from which a
normal embryo is derived.

Blastoderm E, then, must have been characterised from a
very early stage by the presence of two germinal centres, both
defective presumably, in the sense that they must have lacked
some portion of the germinal or formative material present in
the normal germ.

It is proposed to consider here, as briefly as possible, the
probable mode of origin of this digerminal blastoderm.

In the first place, it may be emphasised that while all di-
embryonal blastoderms must have been characterised at an earlier
stage by the presence of two germinal centres, Specimen EH
differs from the great majority of those hitherto described in the
important respect that each of these germinal centres must have
been defective, and in this section we shall be concerned almost
solely with demonstrating that this defectiveness results from the
mode of origin of the two centres, and is only explicable on the
supposition of a certain mode of origin.

Twinning or duplicity is of two main types, to which the
terms Dizygotic and Monozygotie have been applied, practically
all the multitudinous theories which have been suggested falling
into one or other of these two categories. ‘The former signifies

ASYMMETRICAL DUPLICITY IN THE CHICK, 99

the development in proximity of two embryos, each derived from
a fertilised ovum. In this type secondary fusion may or may not
occur, The latter signifies the development of two more or less
complete individuals from a single fertilised ovum, and involves, of
course, a process of fission or separation of the original germ into
two more or less autonomous centres of embryonal formation.
Hither type may be specific, as in the armadillo, or merel ae
abnormal and sporadic, as in the chick and man. Both types
have been experimentally induced in lower forms, the former in
particular by Driesch and Metschnikoff (3, 9), who have sue-
ceeded in producing a single larva from the fusion of two eges
ov young larve, the latter by numerous investigators.

As to which mode of twinning is responsible for the production
of the diembryonal blastoderms which are of so common occur-
rence in the chick, there would seem to be no general agreement,
both seem possible @ priori, and the conclusions of the majority of
authors hitherto, seem to be mainly surmise. Advocates of the
former theory in one or other of its numerous forms have been
the Hertwigs, Weidman and Tur, Broman, Windle, Panum,
Klaussner and Marchand, and of the latter numerous authors
including in recent times, Bateson, Kaestner, Wilder, &c.

The present specimen seems of extreme interest as leaving no
room for doubt between these two alternatives.

The following features of the blastoderm, considered in con-
junction, seem to indicate such a close morphogenetic relation-
ship between the two embryonal rudiments as can only be
construed as due to their having taken origin from an originally
single germinal centre, presumably by some process of fission or
division which resulted in the physiological autonomy of the
two secondary centres.

1. The regular outline of the blastoderm as a whole, which shows
no sign of division into two portions.

2. The continuity of the germ-layers in the various regions of
the blastoderm, and the continuity of the lateral eeelomic
spaces in the region of Rudiment #8 with those in the region
of Rudiment a.

3. The development of the two embryonal rudiments on a single
longitudinal axis. The axis of the “ primitive groove” of
Rudiment 7 is situated on the anterior continuation of the
median axis of Rudiment a (see PI. I. fig. 1).

Further, along that portion of the longitudinal axis be-
tween the two rudiments the median band of unsplit meso-
derm forms what may be regarded ina sense as a differentiated
connection between the two rudiments, forming a conclusive
indication that they are indeed developed on what is strictly
a single primary morphogenetic axis. In other words, the
two embryonal rudiments must have arisen from the original
centre subsequently to the laying down or, at all events, to

the determination of the primary longitudinal axis.
7*

100 MR. NOEL TAYLER ON A UNIQUE CASE OF

It may here be noted that this blastoderm would seem to
fit in well with C. M. Child’s theories of development and
reproduction by the physiological? isolation of parts (1).

4, It has been pointed out above that the primary nature of the
abnormality of the two rudiments is a lack or deficiency,
and it would seem that this deficiency must be regarded as
reciprocal. While the posterior parts of Rudiment a are
quite normal, the amount of disturbance increases as we
pass anteriorly. From section A of the discussion 1t was
concluded that the seat of the primary disturbance was the
extreme anterior region of the embryo, in particular of the
medullary plate. The inference seems fairly natural that
Rudiment 3 represents that portion of the germinal substance
which was lacking in the extreme anterior portion of Rudi-
ment a. Expressed in terms of Child’s theories, one would
say that that portion of the germ from which Rudiment @
developed came to lie outside the range of dominance of the
primary centre, and thus becoming physiologically isolated,
commenced to develop as an independent centre, According
to Child (1), “ Parts of the individual may come to lie
beyond the range of dominance in consequence of increase
im size of the whole, of decrease in range and degree of
dominance by decrease in the metabolic rate in the dominant
region, of decrease in conductivity of the paths of correla-
tron, and of the direct local action of external factors which
merease the independence of subordinate parts.” The factors
responsible for the initiation of the secondary centre of
embryo formation on blastoderm E must presumably have
been somewhat of the nature of the second and fourth of
these, or of a combination of the two. The more likely
explanation would seem to be that the primary disturbance
imvolving the anterior medullary plate region, or more
strictly (since it must have occurred at a very early period)
the apical region of the major axis, led to the defective
development of the anterior portion of the central nervous
system, with the secondary modifications of Rudiment a
discussed in Part A above. On the other hand, the abnormal
development of the anterior end of Rudiment a would have
been accompanied by a decrease of “ metabolic rate,” and
lence in the range of dominanee with the consequent
isolation of Rudiment 5 as a secondary germinal centre.

5. While, as emphasized in 3 supra, the two rudiments clearly
lie on a single long axis, it will be noticed that this axis
curves somewhat to the left in the region of Rudiment /.
It seems possible that this fact may be correlated with
the morphological asymmetry of the head of Rudiment a,
which is more defective on the right side (see Pls. I. & IIT.
tigs. 3,4, E3 and E.4). These two facts suggest an obliquity
of the original disturbanee in relation to the major axis.

ASYMMETRICAL DUPLICITY IN THE CHICK, 101

There are two features of the blastoderm which seem to
require further brief comment :-—

a. The significance of the primitive streak-lke character of
Rudiment /3.

b. The relatively great distance separating the two embryonal
rudiments.

a. Experimental work on the localization ef the prospective
embryo upon the unincubated blastoderm and the relations
of the primitive streak to the future embryo have shown
that the primitive streak furnishes material for the develop-
ment of those regiens of the latter lying posteriorly to the
heart, while, on the contrary, “the material of the primi-
tive streak does not enter into the fermation of the brain”
(Peebles, 12).

But obviously it dees not follow that the converse must
hold, ¢. ¢., that material from which the brain normally
arises, may not under certain circumstances have the power
of giving rise to a primitive streak-like structure or tissue.
Judging from the analogy of lower forms, one might
expect that it would have this power, though it is important
to recollect that, as Child points out, the limitation of the
capacity of reconstitution, 2. e., ef the potentiality of the germ,
is more or less progressive from lower to higher forms.

In any case, owing to the conditions under which the
development of the chick takes place, its experimental
demonstratien weuld be a matter ef extreme difticulty.
Nevertheless, the primitive streak-like nature of Rudiment /3
appears witheut deubt indicative of its physiclegical isola-
tion, 2. €., that it is developing, or “endeavouring” to develop,
if one may be permitted this teleological but expressive
phrase, as a whole.

For, to pursue the analogy with lower forms, gastrulation
and primitive-streak formation resemble each other in this,
that they are each, the one in the heloblastic, the other in
the meroblastic egg, the earliest fundamental morphogenetic
process by which the long axis of the body is determined.

Hence the earliest obvious act, by which a physiologically
isolated part of a holoblastically developing embryo is seen
to be developing as a whole, is the more or less successful
attempt at gastrulation.

In the same way the formation of a primitive streak-like
structure—however imperfect—by a group of blastomeres
upon a blastoderm must surely be regarded as indicative of
their physiological isolation, that is, of an attempt to develop
as a whole, however limited their capacity to do so.

6. With respect to the second point, @ priori, there would
appear no grounds for supposing that the effect of the
initial disturbance would be limited to the embryonal

102 MR. NOEL TAYLER ON A UNIQUE CASE OF

portions of the blastoderm. At the early stage at which the
disturbance must have occurred, there can hardly have been
any sharp distinction between the embryonal and extra-
embryonal portions of the blastoderm: hence the presence
of a second centre of embryonal formation clearly implies the
existence of a region of blastodermie extension and growth
in correlation with it, quite distinct from that related to the
main embryonal rudiment (Rudiment a). The existence of
such a centre also explains the great length of the blasto-
derm as a whole, composed as it is of two adjacent areas of
blastodermic extension lying along one longitudinal axis, and
further explains the great separation of the two rudiments,
since the rate of extension of that portion of the blasto-
derm lying between them must of necessity be considerably
greater than in any other area, for here there is increase of
blastodermic extent in relation to each of the embryonal
formations.

LTV. TERATOLOGICAL SIGNIFICANCE.

1. Bearing on the nature of unequal monstrosity,
‘“* Duplicitas asymmetros.”

Unequal double monsters are those in which the two com-
ponents show marked dissimilarity in the degree of their
development. They are best known in mammals, particularly in
man, in which numerous types have been recorded. There
appears, however, to be little knowledge as to the earlier stages
of these forms of double monstrosity, hence little would seem to
be definitely established as to their mode of development, though
there has been considerable speculation on these questions. Very
few cases of unequal double monstrosity in birds seem to haye
been put upon record, and here, too, little is known as to their
mode of origin and development. Kaestner, one of the foremost
students of double monstrosity in birds, remarks (6, 7) that,
‘* Ueber die embryonale Entwicklung parasitiren oder, wie man
sie auch nennt, asymmetrischen Doppelbildungen wissen wir
wenig”; while Wilder (18), speaking of mammals, said: “ To
my knowledge this form of monster bas never been studied for
the purpose of testing whether or not the two components were
ever originally physical duplicates.”

The accurate definition of unequal double nionstrosity is a
matter of some difficulty in the absence of knowledge as to their
mode of origin and development.

Geoffroy St. Hilaire, who placed them in the second order of
his group of ‘“ Monstres Doubles,’ characterised them as
follows (15) :-—

‘** Monstres doubles parasitaires ou composes de deux individus
trés-inégaux et trés-dissemblables un complet ou presque com-
plet, analogue & un autosite; l’autre non seulement beaucoup
plus petit, mais trés-imparfait, analogue 4 un omphalosite ou

A Ae tor

ASYMMETRICAL DUPLICITY IN THE CHICK, 103

méme & un parasite, par conséquent incapable de vivre par lui-
méme, et se nourissant aux dépens du premier dont il n’est
physiologiquement qu’un simple appendice.”

According to Wilder (18): ‘These monsters consist of two
components of very unequal development, the one (autasite)
being normal or nearly so, and the other (parasite) quite incom-
plete and attached to the first as a dependent growth usually
adhering to some point on the ventral side.”

Schwalbe (16) defines them as: ‘“ Die asymmetrischen Doppel-
bildungen, die zweite grosse Hauptgruppe der Doppelmissbildun-
gen, sind dadurch characterisiert, dass ein Individualteil eine
bedeutend geringere Ausbildung aufweist (Parasite) als die
andere, vollig ausgebildete (Autosite).”

Kaestner (6, 7) more briefly speaks of them as, ‘‘ Doppel-
bildungen mit unsymmetrisch gelegenen oder ungleichmiassig
entwickelten Komponenten.”

It is instructive to compare these definitions: the latter three,
those of Wilder, Schwalbe, and Kaestner, it will be noted, are
based purely on morphological data, in striking contrast to the
original definition of St. Hilaire, which was based also on a
physiological conception, that of nutrition. This latter is now
clearly inadmissible, and in any case was only rendered possible
by the lack of early stages. The only strictly valid definition
and classification of unequal monstrosity would be based not on
physiological, or even on morphological, but on morphogenetic
data, in other words on ontogeny, just as there is only one natural
classification of animals, that based on phylogeny.

In other words, the only rational or natural classification must
be based on a consideration of past history, and through that of
causality.

Now the fundamental principles which are raised by a con-
sideration of the morphogenesis of asymmetrical duplicity appear
to be first, the relationship of the two centres at their time of
origin, and secondly, the possibility and extent of a subsequent
modification of these relations due to secondary causes, which may
or may not act in such a way as to obscure the original condition.
Obviously the rational classification of double monstrosity will
only be founded on the fullest possible enlightenment as to the
former of these questions, which in turn can only follow from a
knowledge as to the extent and nature of secondary modification.
‘The essential problem was appreciated by Wilder in the passage
quoted above, ‘this form of monster has never been studied for
the purpose of testing whether or not the components were ever
originally physical duplicates.”

The question is, after all, merely an aspect of a wider one which
has been stated as the “ ontogenetic permanency of teratological
organisation.” With reference to this, Stockard (17), speaking of
the cyclopean condition, says: ‘‘ the cyclopean defect is present
from the first in the same condition as it will continue throughout
development.”

104 MR. NOEL TAYLER ON A UNIQUE CASE OF

With regard to the origin of unequal duplicates, most writers
hitherto seem to have inclined to the opinion that they were
originally physical duplicates, the asymmetry resulting from
secondary deformation. ‘Thus Wilder in his earlier paper (18)
awlvanced this opinion, basing it partly on the fact that where it
is possible to determine the sex of the ‘‘ parasite,” ‘it seems
always to be the same as that of the autosite.” In his later paper
he seems to have modified somewhat his earlier views, “ all are
not necessarily deformed.”

The discussion may no doubt become somewhat pedantic unless
it be constantly borne in mind that the numerous morphological
types of uneyual duplicates must almost certainly differ con-
siderably in their ultimate origin. ‘ Fiir jede Forme, eine
Spezieluntersuchung eintreten ” (Schwalbe).

On the other hand, were we in a position to establish a classifi-
evtion of unequal monstrosity based upon their morphogenesis,
the most fundamental distinction that could possibly be made
would almost certainly be that between those which did and those
which did not arise from like centres, assuming for purposes of
argument that both of these types do in fact occur.

Bearing in mind that the present discussion is concerned with
the avian blastoderm, we may venture, then, to postulate three
fundamental problems raised by a consideration of unequal
monstrosity.

1. Can the two embryonal centres of an asymmetrical duplicate
be explained as arising ultimately from a single, normal centre 2
In other words, is their nature dizygotic or monozygotic %

2. At their time of origin are the two centres like or unlike ?
3. In the latter case—

(a) What are the causal factors responsible for the dissimilarity
of the centres ?

and

(6) What is the relation between the dissimilarity of the
centres and the production of the two centres from an
originally single centre ?

In other words, is there any direct causal connection
between duplicity and asymmetry ?

Now, clearly, could a definite answer be given to these questions
in every well defined case of asymmetrical duplicity that is placed
upon record, we should be far advanced towards a comprehension
of the main types of these, and of their morphogenesis.

The answer to these questions, as far as the blastoderm described
in this paper is concerned, seems fairly definite, if the conclusion
arrived at in section B supra be valid. There it was concluded :

1, That the blastoderm was certainly monozygotic.

ASYMMETRICAL DUPLICITY IN THE CHICK, 105

2. That the two embryonal centres were, from the earliest
period, dissimilar.

3. That the dissimilar potentiality of the two centres almost
certainly resulted from their mode of origin, z.¢., from a
qualitatively unequal division of the ‘‘ formative substance”
of the original germ.

Here, then, we have at least one well-defined morphogenetic
type of asymmetrical duplicity, that which owes its essential
characteristics to the production of two daughter centres of unequal
potential from, in the first place, a single and presumably normal
centre. It seems impossible that the two embryonal rudiments
on this blastoderm can ever, to adopt Wilder’s terminology, have
been ‘ physical duplicates.” Z

It is not possible in this paper to enter into a comparative
discussion of unequal monstrosity, even in the very restricted
field of avian teratology. In few cases have instances of unequal
monstrosity in birds received sutliciently detailed description to
make this a fruitful undertaking, however desirable. But at
least this may be said :—

That there are at least one or two cases of unequal mon-
strosity which must have resembled Blastoderm E in their mode
of origin so closely as to belong to the same morphogenetic type.
Such would appear to be—

(a) A goose blastoderm described by Grundman (5) and quoted
by Schwalbe (16).

(b) Three chick embryos described by Dareste (2) and also
quoted by Schwalbe.

(c) Possibly a blastoderm described by Kaestner (6) consisting
of a normal chick embryo of 17 (?) somites, laterally to
which occurred a primitive streak-like rudiment, closely
resembling, according to his figure, Rudiment ~ of
Blastoderm EK. ‘There was apparently no sign of any
differentiated morphological connection between the two
rudiments.

The blastoderms described by Grundman and Dareste resemble
each other and differ from Blastoderm E in that the lesser
embryonal rudiment has attained a considerably higher degree of
development than is the case in the latter,

In all these cases the lesser rudiment is characterised by the
extreme deformation and lack of development of the head and
by the absence of the heart. In none of these cases is there any
description of sections. Fortunately, however, I happen to have
in my possession a blastoderm which appears to resemble these
closely, and as it appears not devoid of interest, 1 hope shortly to
have the opportunity of describing its structure.

It may be noted here that Blastoderm E appears unique among

106 MR. NOEL TAYLER ON A UNIQUE CASE OF

described cases of Duplicitas asymmetros in an important and
significant respect. That is the fact that the greater embryonal
formation (“ autosite”), as well as the lesser, exhibits extreme
defects of organisation. It is unnecessary to comment further
here upon this interesting fact, for its morphogenetic significance
was more or less directly considered in Part III. One may
merely emphasize here that it is this characteristic which directly
and indirectly affords strong evidence as to the mode of origin
of the blastoderm, and further, that while this fact is morpho-
genetically of the utmost significance, it indicates no fundamental
difference between this specimen and other forms of asymmetrical
duplicity as regards their ultimate mode of origin.

Apart from the blastoderm referred to above, there are a
number of cases on record, the significance of which appears far
more uncertain; it would seem difficult even to arrive at a definite
conclusion as to whether they can be regarded as monozygotic in
origin. Such, for instance, are blastoderms upon which occur two
embryos, both of which are morphologically normal, but which
are very unequal in their degree of development as indicated by the
number of somites. Such forms, it would seem, admit of equally
plausible explanation, on either the Dizygotic or Monozygotic
hypothesis. In the former case the explanation of the difference
_of age of the two rudiments is clear; in the latter they might
perhaps be regarded as arising from a merely quantitative unequal:
division of the germ, such that each centre received its full
complement of ‘formative substance,” yet the two centres were
quantitatively unlike in such a way as to result in a retarding of
the development of one of them.

If this mode of origin be regarded as likely, we then appear
to be able to distinguish three main morphogenetic bupes of
duplicity :

Type 1. In which the original disturbance of the germ results
in the production of both quantitatively and qualitatively
like daughter centres, giving rise to the equal duplicity,
the well-known type of Duplicitas symmetros.

Type 2. In which the two daughter centres are quantitatively
unlike, but qualitatively similar, giving rise to two embryos
both morphologically perfect but unequal in their stage of
development.

Type 3. In which the daughter centres are both qualitatively
and quantitatively unlike, giving rise to such a form as
Blastoderm E, in which either, or even both, embryonal
eentres exhibit gross abnormalities of structure, apparently
always of the nature of a defect of organisation.

The question of modifications and deformations of a secondary
nature is too complex and lengthy to be dealt with here; it may,

ASYMMETRICAL DUPLICITY IN THE CHICK, 107

however, be again emphasized that it is of extreme importance in
the elucidation of the ultimate mode of origin of asymmetrical
duplicates.

2. Conclusions.

1. Blastoderm E appears unique among hitherto described
chick blastoderms exhibiting asymmetrical duplicity, in that
both, not merely one, of the embryonal formations exhibit gross
morphological defects.

2. The morphology of Blastoderm E seems only to be explicable
on the monozygotic theory of origin, ¢.e., that both centres origi-
nated through some kind of disturbance, the exact nature of
which is at present obscure, from a single and possibly normal
germ.

3. The two centres of embryonal formation, to which the
original disturbance gave rise, must have been from their origin,
qualitatively and quantitatively, of unlike potential.

4. The primary modification induced in the greater embryonal
formation resulted in the inhibition of the normal growth of the
anterior portion of the nervous system and of the formation of
the head-fold.

From this primary modification, indirectly followed various
secondary modifications, due to the causal correlation of the
various organ-primordia of the embryo.

5. While in the normal chick both the anterior growth of the
medullary plate and the pleuro-pericardial expansion are operative
in the formation of the fore-gut, the importance of the latter
factor is indicated by the condition of embryonal formation a, in
which, though no true head-fold could have been present, there
is nevertheless a well-developed fore-gut.

6. Blastoderm E appears of extreme interest with reference to
the localisation of the ‘formative material” in the early chick
blastoderm. For while it has been experimentally demonstrated
that the material of the primitive streak does not enter into the
formation of the brain, it appears that, on the other hand, the
material from which the anterior region of the medullary plate
normally arises may under certain circumstances have the power
of giving rise to a primitive streak-like mass of tissue.

7. Blastoderm E seems to have an important bearing on the
problem of the origin and nature of asymmetrical duplicity.

There has hitherto been difference of opinion as to whether
the two members of an asymmetrical duplicate could or could
not have been originally (to use Wilder’s term) “physical
duplicates.”

Opinion hitherto bas inelined to the view that in the majority
of eases this was the case, the inequality of the two components
resulting from secondary causes. Authors, however, have recog-
nised the difticulty of the problem and the lack of any conclusive
evidence,

108

MR. NOEL TAYLER ON A UNIQUE CASE OF

It is of extreme interest, therefore, to find in Blastoderm EK a
case of unequal duplicity in which there can be no room for doubt
that the two embryonal formations were from the first unlike,
the asymmetry therefore being intimately bound up with the
actual origin of the two centres of embryonal formation from a
single centre, and not resulting from secondary modification
during the course of development.

(oe) at G Or ad

Literature referred to.

. Cuitp, C. M.—Individuality in organisms. Univ. of Chicago

Press, 1915.
Dareste, C.—Recherches sur la production artificielle des
monstruosités. Paris, 1891.

. Driescu, Hans.— Die Verschmelzung der Individualitit

bei Echiniden Keimen.” Arch, fiir Ent. Mech. der Org.,
Bd. x., 1900.

. Grirer, L.—Arch. fir Ent. Mech. der Org., Bd. xxxii.,

LSU

. GrunpMANn.—Anatomischer Hefte, 1900.
. Kagstner, S.— Arch. fiir Anatomie, 1899.
. Karstner, S.—Die Ents. d. Dopp. des Mensch. u. d. Hoh.

Wirbeltiere, 1912.

. MrrropHanow, Paut.—‘‘ Notes Embryologiques et Térato-

géniques.—II{. Note sur un blastoderme double de la
poule.” Arch. fur Ent. Mech. der Org., Bd. i., 1895.

. Merscunixorr, E.—Embryologische Studien an Medusen.

Wien, 1886.

. Newman, H. H.—The Biology of Twins. Univ. of Chicago

Press, 1917.

. Parker, K. M.—‘“‘ The early development of the Heart and

anterior vessels in Marsupials with special reference to
Perameles.” Proc. Zool. Soc. 1915.

. Persies, FLorence.—“ Localization of the chick embryo

upon the blastoderm.” Journ. Exp. Zool. vol. i., 1904.

. Ropirnson, A.—Journ. Anat. Phys. vol. xxxvii., 1902.

. Rovyrkre, H.—Journ. de Anat. xl., 1904.

. Sr. Himarre, Georrroy.—Traité de Tératologie, 1832-37.

. ScuHwaLBe, E.—Die Morphologie der Missbildungen des

Menschen u. der Tiere. II. Tiel. Die Doppelbildungen.
1907.

. Srockarp, C. R.—Amer. Journ. Anat. vol. vi., 1906-07.
. Witvrr, H. H.—Amer. Journ. Anat. vol. 111., 1904.

9 99 99 9 vol. Vi1ll., 1908.

ASYMMETRICAL DUPLICITY IN THE CHICK.

109

EXPLANATION OF THE PLATES.

REFERENCE LETTERS.

A.I.P. =anterior intestinal L.op. =left optic evagination.
portal. Mesenc. = mesencephalon.
Ant.aor.sp. =anterior aortic space. Mes.som. =mesoblastic somites.
Ant.ext..G.=anterior extension of | Om.mes.v. |=omphalomesenteric vein.
fore-gut. O.p. =oral plate.
Ant.p.p.c. =anterior extension of | P.p.c. =pleuro - pericardial
pleuro - pericardial cavity.
cavity. Pr.g. =“ primitive groove.”
Ch.d. =chorda dorsalis. R.aud.ves. =right auditory vesicle.
Dor.aor. =dorsal aorta. R.Ht.T. =right heart-tube.
Dien. = Diencephalon. R.Hy.P. =right hyoid pouch.
End. =entoderm. R.lat.aor.sp. =right lateral aortic
FB. =fore-brain. space.
F.G. =fore-gut. R.mes.pro. =right mesodermal (?)
H.B. =hind-brain. proliferation.
He. =heart. Rud. a, a, :
Inf.dep. =infundibular depression. | Rud. 3 } —Rudiments « & 3.
Lat.l.sul. | =lateral limiting sulcus. Som. =somatopleure.
L.aud.ves. =Jeft auditory vesicle. Som .mes. =somatic mesoderm.
L.Ht.T. =left heart-tube. Sp. =splanchnopleure.
L.Hy.P. =left hyoid pouch. Sp.mes =splanchnie mesoderm.
L.lat.aor.sp. =lett lateral aortic space. Tel. ==Telencephalon.

Pl. I. fig. 1. Photograph of the Blastoderm, stained with Borax carmine and
mounted in balsam.

Pl. II. fig. 2. Transverse section of the primitive streak-like Rudiment 3.

Pl.I. fig. 3. Semi-diagrammatic representation of the anterior (cephalic) region of
embryonal formation @,

Pls, I., LI. figs. 4 & 5. Transverse sections of the anterior (cephalic) region of
embryonal formation a.

Kl.
K2.
K3.
K4,
Ko.
K6.

7.
K8.

In the plane of the telencephalon.

Posterior to E 1 in the region of the anterior extension of the fore-gut.

In the anterior region of the diencephalon.

Through the diencephalon, showing the left optic evagination, and the

infundibular depression.

In the region of the mid-brain, showing the dorsal aorte and the

heart-tubes.
Posterior to E 5.
In the region of the hind-brain.

Through the plane of the auditory vesicles and the anterior intestmal

portal. (See also text-fig. 1, p. 87.)

Pl. I. fig. 7. A. Diagrammatic representation of a median longitudinal section of
the anterior region of a normal chick embryo of 18 somites (after Lillie,

“ The Chick,” fig. 67).

For comparison with B.

B. Diagrammatic reconstruction of the median longitudinal section of the

embryonal formation a.

For comparison with A.

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pe ety eS iis “a8 ee : on ts ie ete
: : ee a i

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jeeiert eI ag
ae bee

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SOME POINTS IN INSECT MECHANICS. Latial

10. Some Points in Insect Mechanics.

Bye: A. MAUnOCK, HAR. S...F'Z.S.
[Received March 18, 1919: Read March 18, 1919.]
(Text-figures 1-8.)

In books on Natural History and on the anatomy of animals
the descriptions are chiefly concerned with generic or specific
characters, and little attention is paid to the mechanical features
of the various parts of the organisms. ‘The mechanics of joints
and muscles, however, are not without interest, although the
similarities are so close over a wide range of orders that they do
not—with few exceptions—form a sufficient basis for purposes of
classification.

Taking the whole range of animal life there is a broad dis-
tinction, in the mechanical sense, between those types which have
internal skeletons (7. e., vertebrates) and those where the skeleton
is external (7. e., insects and arthropods). In both the chief use
of the skeleton is to form a more or less rigid base for the attach-
ment of the muscles (especially the muscles of locomotion), but
the form of the joints on which the muscles operate is very
different in the two cases.

Joints may be conveniently classified by their degrees of
freedom. The most general freedom which any two connected
parts of a structure can have with reference to one another is
that each can be both relatively displaced, and also turn about
three axes at right angles to one another. With joints only the
rotational freedom need be considered, and therefore the greatest
number of degrees of freedom for a joint is three. Such joints
are met with in the case of vertebrates, as, for example, at the
shoulder, which allows the arm to be raised or lowered in a for-
ward or sideways direction, and also to turn about its own axis
relatively to the shoulder. The elbow-joint has two degrees of
freedom, namely, so as to alter the angle between the fore-arm
and the humerus, and to turn the wrist about the mean axis
of the radius and ulna. The last two joints of the fingers are
-examples of joints with one degree of freedom only.

The jointed parts in vertebrates are kept in position by what
may be compared to an elastic stocking composed of ligaments,
and the working surfaces of the bones are kept from actual
contact by a thin cushion of cartilage. The elasticity of the
connection makes the “degree of constraint” (7. e¢., the limits
which the motion, other than that appropriate to the degree of
freedom, cannot exceed) rather lax, and for this reason the verte-
brate-joint can bear accidental strains without the injury which a
more rigid constraint would induce.

The animals whose skeleton is external are not so favourably
circumstanced in this respect. None of their joints have more
than two degrees of freedom, and in general only one. The

112 MR. H. R. A. MALLOCK ON

attachment of the jointed parts is made by a very short, flexible
ligament which joins processes on each, and may be compared
with two tubes connected as shown in text-fig. 1, the tubes
representing the skeleton of the part. Such a joint has only one
degree of freedom, and the constraint is very close *.

The close constraint imposed by these connections makes the
joints rather liable to damage from accidental forces, and it
seems probable that the limit to the size to which articulate
animals can be developed is ina great measure determined by the
brittleness which necessarily follows this type of construction.

Text-figure 1.

Diagram of typical Arthropod joint.

When a moving body is suddenly stopped, as when striking
the ground after a fall, the forces called into play are propor-
tional to the mass of the body, and if the force is so applied as to
tend to turn a limb in any direction except that in which it is
properly intended to move, the reaction at the joint, i.¢., the
force tending to break the ligament, will be, for similar structure,
in the ratio of the mass to the square of the linear dimension—
or, in other words, the chance of breakage decreases directly with
the size.

The great difference between the size of the largest articulate
and largest vertebrate rather bears out this view.

The heaviest land + arthropod—a beetle—may weigh something
over an ounce, and an elephant 4 or 5 tons. Some of the aquatic
articulates—crayfish, I believe—weigh 30 pounds, and of the
aquatic vertebrates, whales approach 100 tons.

Where more than one degree of freedom is required in the limb
of an articulate, this object is generally attained by placing two
or more joints of one degree of freedom in close succession, and
in nearly all cases the legs are thus constructed, the joints of
the coxe, trochanter, and femur being close together with their
axes mutually set at right angles. Text-fig. 2 shows diagram-

* T am informed by Dr. Calman that in the leg-joints of some crustaceans one
of the connections sketched in text-fig.1 is absent, so that in these cases the joint

allows partial freedom of two degrees.
+ Excluding Land-Crabs.

SOME POINTS IN INSECT MECHANICS, 113

matically the typical arthropod leg. The leg is supposed to be
extended into a straight line, though in most cases the natural
limitations of the motion of the joints would prevent such an
extension being actually accomplished. In insects with folding
wings two degrees of freedom are obtained in the same way by
two closely approached joints at the base of the wing.

The locomotive muscles of insects offer many points of
mechanical interest. As far as the legs are concerned, the same
forms of muscles and muscular attachments are found not only
in insects but in all arthropods with well-developed joints, and
this can be readily examined in the legs of the large crustaceans ;
but in insects which have a head, thorax, and abdomen, the
chief locomotive muscles are all contained in the thorax, which
may be compared with the engine-room of a ship, the head corre-
sponding to the conning tower, and the abdomen to boiler-room,
stokehold, aud repair outfit.

Text-figure 2.

Diagram of typical Arthropod leg:
(a) coxa, (5) trochanter, (c) femur, (d) tibia, (e) tarsi.

It was said at the beginning of this note that the mechanical
details of animals with external skeletons rarely offered characters
adapted for the purposes of classification.

Amongst insects, however, the mechanism of the wing-muscles
does definitely separate one small order, viz., the dragon-flies,
from all the others. Dyagon-flies or their allies are, I believe,
geologically the oldest form of insect known, and the action of
their wing-muscles is the simplest. It is represented diagram-
matically in text-fig. 3.

Each of the group of muscles which act on the wing is attached
at its lower end to processes of the thorax, and is capped by a
chitinous cone, rather like an extinguisher, terminating in a fine
ligament. These ligaments lead respectively to points on either
side of the wing-joint; thus the contraction of one set of muscles
(represented by A in the figure) raises the wing, while the other
group, B, depresses it.

In all other Orders of flying insects the arrangements are much
Proc. Zoou, Soc.—1919, No. VIII, 8

114 MR. H. R. A. MALLOCK ON

more complex. The muscles are never directly attached to the
wings, bué to various portions of the thorax, quite distant from
the wing-joint, and deformation of the thorax caused by their
contraction acts indirectly on the wings. It would requive more
elaborate figures than can be given here to make this action
clear, but text-figs. 4,5, and 6 indicate roughly the sections (in
the three principal planes) of the thorax and groups of muscles.
The dotted lines in 5 and 6 show the kind of deformation
produced by the more or less horizontal and vertical groups. —

The question arises as to why has this complicated and indirect
method prevailed,

Text-figure 3.

Diagrammatic half cross-section.

Action of wing-muscles of Dragon-flies.

If the problem were set of designing a mechanism for flapping
wings the dragon-flies solution would certainly be the first to
suggest itself; yet it evidently must have some disadvantages
since it has not been generally adopted.

Amongst many other curious mechanical devices found in
insects the folding of the wings is worth attention. The folding
here referred to is not the mere alteration of the attitude of the
wings at rest and in flight; but an actual folding of the wing
itself, after the manner of a fan. Here, again, more elaborate
figures than can be given in this place would be required to show
the complexity which is found in some of these folded wings.

The simplest is that used by some of the Hymenoptera, where
a single longitudinal fold is formed from the base to near the tip.
This, while not altering the length of the wing in repose,
diminishes its width. In the majority of flying beetles the length
of the wing in the folded position is reduced. This is effected by
a joint on the leading edge of the wing from which the nervures
on the folding part of the wing-membrane radiate. The wing so

SOME POINTS IN INSECT MECHANICS. -

Text-figure 4.

Head

Typical arrangement of flight-muscles in other Orders :—=

(4) half cross-section of thorax ; (5) longitudinal section of thorax ;
(6) half plan-section of thorax.
g*

115

116 SOME POINTS IN INSECT MECHANICS.

folded can then be completely covered by the elytra. The muscle
operating the folding part is contained in an enlarged nervure at
the base of the wing.

Some of the beetles with short elytra and some of the Ortho-
ptera (notably earwigs) have two such joints on the length of the
wing so that the latter when folded is only one third as long as
when extended.

The folds in such wings are extremely complex and can be
more or less represented by a fan at the end of a handle, half the
length of the fan itself. The wing-membrane must be supposed
to be continued from the fan proper to the base of handle. Each
rib of the fan is jointed at its half length, but the axis of these
joints is in the plane of the fan and at right angles to the rib.
See text-fig. 6.

Text-figure 7.

ZN
Zi

Text-figs. 7 & 8.—To illustrate the triple folding of wing of Earwig.
The letters mark identical points in the expanded and folded conditions.

In folding, the fan is first shut and the leading edge turned
through nearly 180°, but before this motion is completed the
half-way joint comes into action so that the outer edge of the
membrane is now under the joint of the fan. See text-fig. 7.

In the case of earwigs, at any rate of the species found in
England, it seems more than doubtful whether the wings are
ever used. The thorax only contains traces of flight muscles, and
I have not been able to distinguish any folding muscles in the
wings themselves.

The ancestors of the present species probably had working
wings which for some reason have now fallen into disuse. }

ON AFRICAN ICHNEUMONIN® IN THE BRITISH MUSEUM, 117

11. On some Equatorial and other Species and Genera of
African Ichneumonine contained in the Collection
of the British Museum. By CraupE Mor ey, F.E.S.,
F.ZS., &., Framlingham, England.

[Received April 11, 1919: Read June 17, 1919.]

Only a small part of the very numerous specimens now lying
idle and neglected in our National Museum is here treated of
—in fact, only species referable to the first (Ichneumonine)
of the five known Subfamilies of Ichneumonide have been
touched upon, and of that Subfamily no more probably than a
tithe. One of the interesting points of the present paper is the
discovery in Equatorial Africa of genera, though no new species
known hitherto only from Asia; and this goes to strengthen a
possible former connection of the two faunas, by way of the
Chagos Archipelago and the Seychelle Islands, suggested by me
(Trans. Linn. Soc. xv. 2, 1912, p. 170). But, although identical
species are captured at Lyndhurst and Simla, none of the African
ones to the south of the Sahara—except such stragglers as Bassus
letatorius—appear to be quite the same as upon the northern
coasts of the Mediterranean; indeed, a few forms, e. g., the
genera Magwenga and Skiapus, are exclusively Ethiopian, while
the larger Xoridides and Cryptides are among the finest and
most beautiful in the world. All, or nearly all, the insects of
the present paper are contained in the material collected by
S. A. Neave in East Central Africa during recent years;
wherever there is reference to other entomologists their names
are inserted in full.

CLASSIFIED CATALOGUE

of Species and Genera herein described.

ICHNEUMONID.

IcHNEUMONING.
J oppides.

Epizoppa Morl.
. verecunda Tosq.
. fumosa Morl.
. pygidifer, sp. n.
. dimidiata Morl.
corrugata Tosq.
. eucelea Morl.
. carinifer, sp. n.
. triangulifer, sp. n.
. varlabilis Morl.
. pilosa Cam.
. Striatifrons, sp. n.
12. rubricata Mort.

Tro@us Panz.
1. gryps, sp. n.

=
BODONIANB wD H

CATADELPHUS Wesm.
1. anceyi Berth.

2. nigrocyaneus Tosq.

3. rubricaput, sp. n.

ErytTHRoJoppa Cam.
1. rufipedalis, sp. n.
2. nigripedalis, sp. n.

LEPTOPHATNUS Cam.
1. ruficeps Cam.

CTENOCHARES Forst.

1. blandita Zosq.
2. testacea Szépl.

3. microcephala, sp. n.

4. gracilentor, sp. n.

IscHnoyoppa Kriech.
1. luteator Fab.

XANTHOJOPPA Cam.

. lutea Cam.

. debilitor, sp. n.

. rotundator, sp. n.
. truncator, sp. n.

. Inermis Vor.

. explanator, sp. n.
. gracilator, sp. n.

. Striator, sp. n.

. bipapillator, sp. n.
. collifer, sp. n.

11. cothurnator, sp. n.
12. geminator, sp. n.
13. areolator, sp. n.

HorprisMENUS Grav.

1. fulvator, sp. n.
2. cibdelus Tosq.

=
SOON OWE Cob et

CoRYMBICHNEUMON, g. n.

1. carinifer, sp. n.

118 MR.

A@LAosoPPA Cam.
1. johannis Cam.
2. glabrinotor, sp. n.
3. comptulus Tosq.

Ca@LicuNnEumon Thoms.

. scopulifer, sp. n.
. cornellifer, sp. n.
. maculiscutis Cam.
. natalensis Cam.
. sublunifer, sp. n.
. geminifer, sp. n.
testaceus Cam.

. globulifer, sp. n.
. sulcifer, sp. n.

. thyridifer, sp. n.
. striatifer, sp. n.

BOD OMNIA O Se whe

as et

Ichneumonides.
Oxypygini.
Lagenesta Cam.
1. sinifer, sp. n.
2. duplicator, sp. n.
3. triplicator, sp. n.
4. triangulifer, sp. n.
(monitor, sp. 11.)

MELANICHNEUMONThoms.

1. carinifer, sp. n.
2. melanopterus, sp. n.
3. glaucopterus, sp. n.

LertotmEcts Cam.
1. alutacefer, sp. 1.
2. punctifer, sp. n.
3. mesonotifer, sp. n.

STENICHNEUMON, Thoms.

1. ochraceator, sp. n.

CRATICHNEUMON Thoms.

1. testacecolor, sp. n.

BaRricuNeumMonN Thoms.

. concinnator, sp. v.
. incubitor Linn.

. planinotum, sp. n.
. sexalbatus Wesim.
. fossifer, sp. n.

. mundatus Tosq.

Oe Go bo

IcunEuMON Linn.
1. rubrornatus Cam.

CrasMias Ashin.
1. glaucopterus Mor.
2, ruficaudator, sp. n.

Amblypygini.

AMBLYTELES lVesm.

. spilopterus, sp. n.

. auricomus, sp. n.

. fulvocaudatus Tosq.
. hegatorius Fab.

. maculicaudis Cam,
. testaceator, sp. n.

—"

Oe Oo bo

ICHNEUMONID.
ICHNEUMONIN A.
JOPPIDES.
EIPIJOPPA.

CLAUDE MORLEY ON AFRICAN .

SPILICHNEUMON Thoms.
1. didymatus, sp. 1.
2. unipunctor, sp. n.
3. triangulator, sp. n.
XENOJOPPA Cam.
1. fossifrons, sp. n.
(kali, sp. n.)
CrENICHNEUMON Thoms.
1. castanopygus, sp. n.

Heresiarchini.
MAGWENGA, gen. nov.
1. maculipennis, sp. n.

Miosoppa Cam.
1. quadrilineola, sp. n.

Listrodromini.

NEotypus Forst.
1. obscurator, sp. n.

Platylabini.

Pratyiabus Wesm.
1. atricinctus, sp. n.
2. mediorufus, sp. n.
3. ceta Morl.

4. vallatus Morl.

Pheogenini.

BrneEcies Cam.
1. dimidiatus, sp. n.
2. politanus, sp. n.

Morley, Revis. Ichneum. iv. 1915, p. 49.

Vosquinet in 1896 described two species of the Fabrician genus
Joppa from Africa; and these, upon examination, I found to
differ to such an extent from the more typical Neotropical forms
of that genus that I erected a new one for them under the
above name, and in it added six other kinds that appeared then
to be undescribed. One or two of the characters at that time
ascribed to the group must be deleted in order that it may
include other species, which, while agreeing in all the more
salient features, differ herein: the scutellum, though sufficiently
convex, is not invariably subpyramidal nor always deeply punc-
tate, and in one of the following new kinds the peculiarly
glabrous and glittering metanotal areola is replaced by one of
the more common dull and rectangular form. As a whole, the
genus is remarkable in its brilliant metallic coloration and the

ICHNEUMONIN IN THE BRITISH MUSEUM. 119

rugosity of the surface, often combined with totally nigrescent
wings. One specimen I have seen was labelled ‘ Mackayella rufa.
Type”; but this appears to bea MS. name. ‘The known species
are as follows :—

Table of Species.

(10) 1. Anterior wings more or less broadly infumate.
2. Infumation of wings dense throughout; head red.
(6) 3. Abdomen fusiform; hypopygium not covering
terebral base.
(5) 4 Mesonotum and scutellum red; hind tarsinormal. 1. vereewunda Tosq.
(4) 5. Mesonotum red, scutellum blue-black; tarsi

incrassate ........... Deacon res 2. fumosa Morl.
(8) 6. Abdomen Sabeyinabsear bypopyaginm covering
terebral base ............ . 3. pygidifer Morl.

(2) 7. Infumation of wings extends ao ae oulize 5;
head not red.

(9) 8. Facial orbits and anus not oe hind femora

TIAN ECL. Sooeandooaonece 4, dimidiata Morl.
(8) 9. Facial orbits and anus pales inal Teas: entively
blocky svcd nan Re 5. corrugata Tosq.
(1) 10. Anterior wings hy Ate or Sabhyaline dhesustout
(12) 11. Mesonotum metallic blue; wings subhyaline ...... 6. eucaelea Morl.

(11) 12. Mesonotum entirely rufescent; wings hyaline.

(22) 18. Antenne and legs mainly black ; mesonotum dark
red; anus white.

(19) 14. Inner orbits more or less broadly, and anterior
coxee, white.

(18) 15. Base of metanotum longitudinally striate on either

side.
(17) 16. Scutellum convex, sapine, mupeeates mesouotum

SHUT sosnocuoo asses : soouaoccodescedasa Za QUE Wilorls
(16) 17. Scutellum pyramidal, vet regose ; esoh Otani

bituberculate . at - 8. triangulifer Morl.
(15) 18. Base of anatanonditn math Ri ‘ails Monest dived

SULIAUC HERE e veces. 9. variabilis Morl.

(14) 19. Inner aulbiie aed anode Coxe ceaiite:
(21) 20: Metanotal areola nitidulous; second segment not

pale .. Bones j 10. pilosa Cam.
(20) 21. Mie tan otal aes Gene aaonal eeemee icatit

NNW, Sooncadoccoe 11. striatifrons Morl.
(13) 22. Antenne, legs, Briloreccthorar nice. eae abdonnen

Coie te 12. rubricata Morl.

1. VERECUNDA Tosq.

This, the type of the genus, was described by Tosquinet (Mém.
Soc. Ent. Belg. v. 1896, p. 101) from the Cape; it appears to be
not uncommon, and I have already recorded it from Natal,
Rhodesia, British Hast Africa, Uganda, and Nyasaland. Addi-
tional material is from Mlanje in Nyasaland at 2300 feet on
September 5, 1913, and the valley of the Kola River, near
EK. Mount, Chiperone, in Portuguese Hast Africa, at 1500 to
2000 feet on 3rd April, 1913.

120 MR. CLAUDE MORLEY ON AFRICAN

2. FUMOSA Morl.

This species usually has the pronotum and propleure red, but
the latter is occasionally, like the prosternum, black-blue; its
anal white markings are not infrequently obsolete. It is appar-
ently somewhat local in Central Africa; I have recently seen
females taken at Mlanje in Nyasaland on 5th and 25th Septem-
ber and 7th November, 1913, by Neave; and at Salisbury in
Mashonaland in September 1913, as well as at 5000 feet there
during June 1900, by Marshall.—The var. apicalis certainly does
not merit specific rank, for I have seen a 9 of intermediate form
between the typical one and this variety, which, while showing
the apically clear wings of the latter, bears no white markings
at all, and, moreover, has the mesonotum and part of mesopleurie
of the dull ferruginous coloration found in £#. triangulifer,
though the abdomen is of normal breadth and much broader
than in that species. It was taken at Mlanje in Nyasaland,
between September and February, 1914, by J. B. Davey.

3. PYGIDIFER, Sp. n.

A stout and somewhat large, bright blue species with the head,
pronotum, and propleure red; centre of the long flagellum
broadly, second segmental plica and the anus from middle of the
sixth segment, white; and the mandibles and remainder of
flagellum black. Abdomen gravid with the second and third
segments longer than broad, and the hypopygium covering base
of terebra. Length,23mm. 9 only.—The resemblance of this
species to #, fumosa is remarkable in the structure and coloration
of the head, thorax, legs, and wings, of which the last are some-
what more ample though equally infumate; but the sub-
cylindrical abdomen is utterly different in shape and anal
structure. The second segment is distinctly and the third
slightly longer than broad, with the former deeply emarginate
in the centre of its apex and the latser more broadly on either
side of its base; apex of second and whole of the following
segments glabrous and strongly nitidulous, as also is the convex
venter; hypopygium large, apically simply rounded and covering
base of the almost concealed terebra. So extraordinary is this
structure that a new genus would be requisite for the present
species were it not that both oxypygous and amblypygous forms
occur in the allied genus Protichnewnon.—The type was captured
at Entebbe in Uganda during August, 1911, by C. C. Gowdey.

7. CARINIFER, Sp. n.

A dull ferruginous and coarsely sculptured species with the
palpi, mandibular base, frontal orbits, flagellar band, anus from
fifth segment, ventral plica and more or less of the anterior with
apex of the hind coxee beneath, and inner side of anterior femora,
white; mandibular apex, flagellum, apex of prosternum and base
of mesosternum, and remainder of anus from base of fourth

ICHNEUMONINA IN THE BRITISH MUSEUM. 121

seement, hind trochanters with their tarsi and more or less of
their tibie, black; wings hyaline or slightly flavescent, with
stigma black. Length, 12-14mm. 92 only.—Known by its
distinctive coloration, the but slightly convex and glabrous
scutellum which bears a few scattered punctures, the simple
mesonotum, by the longitudinal striation at metanotal base, and
especially by the shining longitudinal carina on centre of both
mesonotum and second segment.—Probably not rare: Mlanje at
2300 feet, on 13th June and 8th October, 1913, in Nyasaland ;
on the S.E. slopes of Mount Kenya at 6-7000 feet, between
3rd and 12th February, 1911, in British Hast Africa; at Western
Ankole at 4500 to 5000 feet in mid-October, 1911 (Weave), and
an unlocalised specimen (ex coll. Cameron) from Uganda.

8. TRIANGULIFER, Sp. n.

A dull ferruginous and coarsely sculptured species with flagellar
band and anus from fifth segment alone, white ; mandibular apex,
flagellum, whole sternum indefinitely, a discal mark at base of
fifth segment and nearly the whole legs, black; wings hyaline
with stigma black. Length, 15-17mm. ¢ 9 .—Much narrower
than /. fumosa and constantly larger than 2. carinifer, from the
latter of which it differs in little but its less profuse markings,
subpyramidal and closely rugose scutellum, and in having the
mesonotum slightly elevated tuberculiformly on either side of its
apex, the centre of which is triangularly depressed. The typical
3 has the facial orbits broadly but not the frontal, base of
mandibles and of palpi, the anterior coxe beneath and their
femora internally, stramineous; flagellum immaculate black.—
All the specimens I have seen were captured by Neave at Mlanje
in Nyasaland, at about 2300 feet, during the first half of June
and early in September, 1913.

10. proosa Cam.

Henicophatnus pilosus Cam. Ann. 8. Afr. Mus. v. 1906, p. 166,
2; Epioppa nigricoxata, Morl. Rev. Ichn. iv. 1915, p.53, ¢ @.

I have examined Cameron’s type, from the district of King
William’s Town, in the Cape Town Museum.

11. STRIATIFRONS, sp. n.

A short and somewhat stout species with the scape, head,
except the black mouth and clypeal apex, pro- and mesothorax
with scutellum, crimson; metathorax and basal segment bright
blue, remainder of abdomen violaceous with anus, apex of second
segment both above and below, and apex of the first below only,
pale stramineous; legs entirely and flagellum, except its central
stramineous band, black; metathorax coriaceous, with fine carine
and the areola not more shining. Length, 13mm. 92 only.—
Similar in size and sculpture to H, nigricoxata Morl. (Revis.
Ichn. iv. 1915, p. 53), but with the frons, which is utterly

JQ) MR. CLAUDE MORLEY ON AFRICAN

glabrous in Z. verecunda, stoutly trans-striate throughout, the
rugose scutellum but simply convex, the metanotal areola half as
long again as broad, rectangular, flat and not shining, and the
second ‘segment like the anus apically white-—Hntebbe in Uganda
during the first half of September, 1911.

TROGUS.

Panzer, Krit. Revis. 11. 1806, p. 80; Forster, Verh. pr.
Rheinl. xxv. 1868, p. 188.

The Zrogus group of Joppid genera consists of Dinotomus,
Catadelphus, Trogus, and Automalus. Dalla Torre’s Catalogue of
1902 listed thirty-three species of the main genus, but seven
of these belong to Dinotomus and one (teste Viereck, Proc. U.S.
Nation. Mus. xlvi. 1913, p. 369) to <Aglacjoppa, and a ninth
proved to be synonymous; but none of them have hitherto been
known from Africa. Automalus, with its single palearctic and
single Alaskan species, is equally unknown; but the widespread
Dinotomus lapidator Fab. extends to Algeria from the novth.
Catadelphus was represented by a couple of kinds, both described
in 1896, but not since recorded. In 1844, Wesmael split off
Automalus from Trogus on account of its straight and not
centrally angled clypeus, and ten years later he erected Cata-
delphus for the reception of Ichnewmon arrogator, because its
scutellum was simply convex and not pyramidal as in the more
typical species. Several of Cameron’s Indian Joppides also
belong to the present group, but there will at present be need to
consider no more than one of them.

1 GRYes) sp. mi.

An extremely large and stout, blue-black species, with the
flagellum except apically and legs except basally, conspicuously
pale. Head, thorax, and apical attenuation of flagellum black,
with the centrally elevated face and sometimes the temples
badious; clypeus apically truncate, labrum strongly exserted.
Mesonotum and scutellum very dull and dead black with the base
of the former longitudinally impressed, the latter not strongly
elevated and laterally carinate only to its centre; metanotum
rugose with areola triangular, costule strong though irregular,
and a basal prominence on either side. Abdomen black or blue-
black and smooth with basal segment dull, finely punctate, and
centrally bicarinate; terebra stout and hardly exserted. Legs
long and stout, bright testaceous with coxze, trochanters (front
ones badious) and onychii alone black. Wings large and ample,
strongly and evenly fumate, with blue reflection; discoidal cell
narrow and parallel-sided ; nervelet small; areolet higher than
broad, elongate-triangular and coalescent above. Length, 26 mm. ;

exp. al. 48mm. Q only.—Larger than Catadelphus nigrocyaneus
with dull mesonotum, no ae flagellar band, &c. ; most closely
allied to the Sonoran T. atr ooceruleus Cresson. —Gowdey found

ICHNEUMONINE IN THE BRITISH MUSEUM. 123

the type at Entebbe in Uganda on 17th June, 1913; and Neave
took another female, with entirely black (probably owing to
grease) abdomen, at Mlanje in Nyasaland, on 8th February,
1913.

CATADELPHUS.
Wesmael, Bull. Acad. Sc. Belg., Annexe, 1854, p. 134.

Not more than half a dozen species are yet described, but these
are distributed over southern Europe to the Red Sea, the United
States, Central America, and the northern half of Africa.

Table of African Species.

(2) 1. Alar areolet pentagonal; legs black, tibiae and

panel ne dete nesien eeden. Ppaseee eee atest eicraaen ecient Ieeanceym berths
. Alar areolet triangular; hind femora and tibie

black or red.
(4) 3. Head quadrate and black; hind femora and tibis

—
=
WH
to

THOUNVOWE coo ono donass coon sansbasoderdvossedsousosoDy ced xavbes ea OUR OEOrenS MMOS
(3) 4. Head transverse and red; hind femora and tibix
[SRY co sorigcsedeno onoenedd bsosco don pdeOpobsaGebnan doc ssonaO mes AeE ULE ney none Noy.

3. RUBRICAPUT, sp. n.

A large, blue-back insect with the anus and centre of flagellum
white-marked. Head indefinitely rufescent, small and constricted
behind the prominent eyes ; face closely punctate, transverse and
not higher than the sparsely punctate and apically truncate
clypeus; mandibles aciculate- punctate, with their margins
smooth and teeth black; frons centrally sulcate and somewhat
elevated. Antenne black and rather strongly inflated beyond
the central white band which is broad, but incomplete below.
Thorax black and closely punctate; mesonotum, especially
laterally, smoother with elongate and subparallel notauli; meta-
notal carinz entire, areola narrow and obsoletely discreted from
the basal area, apically elevated and emitting costule from its
centre ; petiolar and dentiparal area obliquely substriate ;
spiracles elongate, apophyses wanting. Scutellum black, convex
and smooth, with large and sparse puncturation; its lateral
cavine vallate and extending to near apex. Abdomen deplanate,
elongate fusiform and blue with purpurascent reflection ; post-
petiole shagreened and centrally aciculate, second and third
segments very finely punctate and dull with gastrocceli of former
deeply impressed, though hardly as broad as the striate and
transimpressed intervening space; remainder nitidulous with
dise of seventh, and apex of sixth discally, flavous ; venter plicate
throughout, with terebra slightly exserted and spicula red. Legs
slender and black (femora et tibize anticorum, articulis 3-5
tarsorum posticorum desunt). Wings evenly nigrescent through-
out, with brilliant blue iridescence; stigma, nervures, and the
punctate tegule, black ; areolet triangular, subcoalescent above ;
basal nervure continuous through median, nervelet obsolete.

124 MR. CLAUDE MORLEY ON AFRICAN

Length, 23mm. @ only.—Very like Catadelphus migrocyaneus
Tosq., though with the head not quadrate and entirely red, the
anus white, &e.—The type occurred to Delmé-Radcliffe at Msozi
in Uganda during February 1903.

EGRYTHROJOPPA.
Cameron, Ann. Nat. Hist. ix. 1902, p. 146.

Head with the labrum exserted, and clypeus apically truncate
base of flagellum usually broadly rufescent, its centre in 2 some-
what strongly dilated. Base of metanotum deeply discreted and
vertically elevated ; areola small and tuberculiform, not laterally
carinate nor depressed; aree entire; petiolar area basally
carinate, narrow and a little explanate throughout; apophyses
wanting. Scutellum pyramidal and in ¢ discally acute; higher
than mesonotum and laterally carinate at least basally, with
its basal declivity in ¢ subvertical and the apical oblique.
Abdomen elongate and usually narrow, about double length of
thorax; postpetiole distinct; discal strize of second segment
centrally elevated; venter with segments two to four plicate and
the apical obtuse, nearly as long as penultimate. Legs elongate,
with tarsi spinulose ; hind femora not extending to apex of third
segment. Areolet broadly triangular, coalescent above; nervelet
often distinct; recurrent nervure centrally, and radius basally,
sinuate; basal nervure subcontinuous through median.—The
great majority of Peter Cameron’s genera are unintelligible
without reference to the types; those of Hrythrojoppa are in the
British Museum, and his four Indian species seem to represent
the opposite sexes of but two, The following insects are suffi-
ciently homogeneous, though the abdomen is a good deal broader
and more deplanate than in those formerly described.

1. RUFIPEDALIS, sp. n.

Dull pale ferruginous, with the abdomen blue and wings
evenly nigrescent ; base of propleurz black-marked. Head pos-
teriorly buccate and nearly as broad as the often testaceous eyes ;
ocellar region circumsuleate ; frons concave and radiately striate
from scrobes; face closely and confluently punctate, centrally and
laterally elevated ; clypeus large, distinctly and evenly punc-
tate, laterally elevated, with its. apex truncate and stoutly mar-
eined. Antenne subdilated beyond centre and apically attenuate;
fourteen basal flagellar joints bright testaceous and remainder
black. Thorax robust and stoutly shagreened ; notauli short and
distinct, sternauli wanting; speculum glabrous and impressed,
finely striate below; metathorax evenly coriaceous with complete
are and strong carine; petiolar area very narrow, trans-striate
and extending nearly to base; spiracles linear, apophyses want-
ing. Scutellum strongly convex and almost pyramidal, discally
obtuse, shining and pilose with a few sparse and fine punctures.

ICHNEUMONINZE IN THE BRITISH MUSEUM. 125

Abdomen elongate-fusiform, deplanate, bright metallic blue, with
petiole alone (though to a variable extent) rufescent ; postpetiole
broader than long, finely shagreened with deep apical and lateral
punctures; second and third segments closely punctate, former
not basally striate, its gastroceeli large and transverse; venter
plicate throughout; hypopygium remote from the subexserted,
black terebra. Legs stout and somewhat short, brick-red with
only the onyches black; hind coxe with stout scopule. Wings
ample and evenly nigrescent, with stigma black; areolet trian-
gular, coalescent above; nervelet obsolete. Length, 20-22 mm.
© only.—Apparently common in Central Hast Africa; many
taken at Mlanje during May, June, September, and December,
1913; inthe Ruo Valley at 2000 feet in December 1913; and
to the S.W. of Lake Chilwa, during January 1914, in Nyasaland ;
as well as at Masongaleni at 3000 ft. in March 1911, in British
Kast Africa (Veave) ; and near Chirinda Forest in Gaza Land,
during March 1907 (Marshall).

2. NIGRIPEDALIS, sp. n.

Dull, pale ferruginous with the abdomen blue and wings
evenly nigrescent ; base of mesopleure sometimes black-marked.
Length, 19-21mm. og 2.—This species is certainly distinct
from the last, yet the 2 differs solely in having the hind legs
(except usually their trochanters and disc of cox) and the inter-
mediate tarsi, black; the flagellum immaculate black and the
scutellum pyramidal with its disc acute. The ¢ agrees there-
with, but the scutellar disc is subspinately produced.—Entebbe
early in September 1911, and on W. shore of the Victoria
Nyanza at Buddu at 3700feet in September 1911, in Uganda ;
and at Mlanje, during the first half of June 1913, in Nyasaland.

LEPTOPHATNUS.
Cameron, Ann. 8, African Museum, v. 1906, p. 165.

The deeply impressed basal metanotal sulcus places this genus
incontrovertibly in the Joppides.

]. RUFICEPS Cam,

This striking species was originally described from Cape
Colony; but it has a wide range. I have recently examined
- examples of both sexes (Cameron knew only the Q, loc. cit., and
I brought forward the ¢ in the Annals of the 8. Afr. Mus. 1917)
from Barberton in the Transvaal, where P. Rendall found it
about 1890 (ex coll. Distant); from Mlanje on 25th February,
1913, in Nyasaland ; from the Siroko River near the west foot
of Mount Elgon at 3600 feet in mid-August, 1911; on the west
shore of the Victoria Nyanza at 3700 feet near Buddu during the
following September; on the Semliki Plains near the south

126 MR. CLAUDE MORLEY ON AFRICAN

shore of Lake Albert at 2200 feet at the end of November, 1911 ;
and between Jinja and Busia in some forest east of Busoga at
about 3800 feet at the end of July 1911, in Uganda; as well as
on the southern slopes of Mount Hlgon at some 5500 feet early
in the preceding June (eave), in British East Africa,

CrENOCHARES.
Forster, Verh. pr. Rhein]. xxv. 1868, p. 191.

The limits of this genus are, owing to the paucity of its original
description, at present somewhat too elastic; but there can, L
think, be little doubt that the two new species here brought for-
ward are at least more closely allied therewith than any other.
Five kinds belonging here were known to me in 1915; the late
V. Szépligeti described another from East Africa (Bull. Mus.
Pavis, 1907, p. 137) and four more in the Kilimanjaro Expedi-
tion, which was published in 1910; I brought forward another
from South Africa in 1916 (Ann. 8. Afr. Mus. xv. p. 371), and
Matsumura has recorded the genus from the Island of Sachalin
(Journ. Coll. Sapporo, iv. 1911, p. 94), off the east Asian coast.

1. BLANDITA Tosq.
W. Haygarth has found this species at Durban in Natal.

2. TESTACEA Szeépl.

A common species in South Africa, recently captured by Bell
Marley at Stella Bush and Howick in Natal, and P. Rendall met
with it at Barberton in the Transvaal (in coll. Distant); but
rarer farther north, for Neave could find only a single female,
with flavidous scutellum and apically black hind femora, on the
S.E. slopes of Mount Kenya at 6000-7000 feet during February
1911, in British East Africa.

3. MICROCEPHALA, sp. n.

A clear testaceous insect, with the face and vertical orbits
indefinitely stramineous; flagellum and hind tibie black with the
8th to 22nd joints of the former, and base of the latter along
with their whole tarsi, white. Anterior tarsi nigrescent. Scu-
tellum dull, apically a little constricted, laterally carinate as far
as apex; apophyses small, but distinct ; gastrocceli large and not
white. Wings flavescent-hyaline with areolet as broad as high, ©
and not coalescent above; stigma ferruginous. Length, 15 mm.
2 only.—Very like C. testacea but with the head smaller, abdo-
men stouter and immaculate; antennz longer and very broadly
white-banded, stigma darker and the hind legs quite differently
coloured, with only the onychii black. [Os. In this unique I
have taken the left hind tarsus ag normal; curiously, the right
one has the apical two-thirds of its basal jomt nigrescent.| The

ICHNEUMONINA IN THE BRITISH MUSEUM. MOAT

typical female was taken in Uganda, Southern Toro, Fort Portal
Road, at some 4000 feet, sounds the end of October 1911.

4, GRACILENTOR, sp. 0.

A slender, testaceous species with the head, antenne, and anus
all white-marked black; hind tarsi nigrescent throughout. Head
transverse-oval and posteriorly nearly as broad as the not very
prominent eyes; flavous with the occiput, vertex, and frons black ;
frontal orbits to vertex citrinous; occiput abruptly declivous be-
hind eyes and, like the centrally carinate frons, closely punctate ;
face and clypeus flat, closely and confluently punctate, not dis-
creted and the latter centrally depressed before its apical margin ;
labrum exserted, mandibles very slender with the teeth piceous.
Antenne filiform and serrate, black with scape and underside of
basal half rufescent ; flagellar joints 16-23 white; apices desunt.
Thorax cylindrical and substramineous with no notauli, apophyses
nor are; areola represented by two slight longitudinal carine.
Scutellum dull, but slightly convex, carinate to apex. Abdomen
sublinear, much longer than head and thorax; basal segment
linear with the obsoletely punctate-aciculate postpetiole hardly
broader; gastrocceli very small; anus black from near base of the
fifth segment with the seventh, apex of sixth and the stout
valvulee, white. Legs slender and somewhat elongate. Wings
narrow, flavescent-hyaline; stigma luteous; areola pentagonal,
broad above; radial cell elongate and narrow. Length, 14-15 mm.
3 only.—The outline of the abdomen is curiously like that of
Mesoleptus males.—Found at Howick in Natal about 1904, by
J. P. Cregoe.

IscHNOJOPPA.
Kreichbaumer, Entom. Nachr. xxiv. 1898, p. 82.

This distinct genus needs more revision than I am at present
able to effect. It was erected for the reception of a species of
Joppid with elongate metapleural spiracles, occurring in both
India and Senegal; and I have shown that Chinese, Bornean,
and Australian examples are not structurally distinct (Revis.
Ichn. iv. 1915, p.97). Dr. A. Roman finds that /schnws melano-
pygus Holmgren and its variety belong here (Entom. Tidskr.
xxxi, 1910, p. 172); and it seems nearly certain that the two
kinds described under Jschnus by Tosquinet in 1896 must also be
included, on account of their elongate thoracic spiracles—thus
Africa will be left with but a single /schnus-species (I. trochan-
teratus Burgst, Tijds. v. Entom. lv. 1912, p. 267; Tunisian
Hym. 1913, p. 15, 9). Szépligeti brought forward two new
species of Jschnojoppa from Kilmanjaro in 1910, and two more
from Java (Leiden Mus. Notes, xxix. p. 235) in 1908: also, I
have added a couple (Ann. 8. Afr. Mus. 1916-17), giving the
total of nine known kinds, of which seven are African. Some
synonymy may be expected.

128 MR. CLAUDE MORLEY ON AFRICAN

1. tureator Fab.

Ichneumon luteator Fabricius, Ent. Syst. Suppl. 1798, p. 222;
Bodargus rufus Cameron, Journ. Str. Br. R. Asiatic Soc. xxvil.
1902, p. 53.

In Africa this species is already recorded from Sierra Leone,
Congo, Uganda, Nyasaland, British Kast Africa, German (olim !)
East Africa, Rhodesia, and Zululand. Additional material is
from Kayena in Cape Colony during October 1916 (lL. Pévin-
guey); the east slopes of the Aberdare Mountains at 7000 to
8500 feet in British East Africa at the end of February 1911;
and a male (with the vertex black and external radius remark-
ably straight) from Western Busoga, between Kakindu and the
S.E. shore of Lake Kioga at 3500 feet on 22nd of the following
August, in Uganda.

Neave also took a couple of males, which I shall here merely
term VAR. wnicolor, nov., though they are pretty certainly of
specific rank, for their coloration is testaceous with nothing but
flagellum (excepting the normal white band) and mandibular
teeth black; further, the abdomen is a little rounded laterally, a
good deal less parallel-sided than the typical form, the external
radius is nearly straight and the areolet much narrower, nearly
coalescent above. These occurred near Kampala on the Kampala-
Jinja Road, which is partly forest, during July 1913; and in the
Durru Forest, Toro, at the end of October 1911, both at 4000
feet, in Uganda. With them, Neave sent home a single female,
which may or may not belong here, for (though the radius is
sinuate and abdomen linear) only the mandibular teeth, part of
flagellum, and the anus from base of fifth segment, are black.
This was found on the S.E. slopes of Mount Kenya early in
February 1911, at fully 6000 feet, in British Hast Africa.

XANTHOJOPPA.
Zanthojoppa Cameron, Ann. Nat. Hist. vii. 1901, p. 378.

This genus of testaceous insects was erected for the reception
of a single Khasian species, and its author added five more in
the same Magazine during 1903 and 1907, all from eastern
India. In 1906 Cameron erected another name for a South
African species (Anisogoppa, Ann. 8. Afr. Mus. v. p. 168), which
differs so little that I ventured to synonymise them, when bringing
forward a new kind (XY. inermis Morley, ib. cit. xv. p. 358) in
1916 from Cape Colony. J now find the genus to be well repre-
sented in Central Africa by numerous specimens and a dozen
somewhat closely allied species, which may be recognised by :—
Head somewhat small, not broader than thorax and never
buccate; posteriorly narrower than the internally entire eyes;
ocelli always, but their intervening space rarely, black; clypeus
neither short nor discreted from the deplanate face, apically sub-
truncate; cheekselongate. Antenne nearly always white- banded

ICHNEUMONIN& IN THE BRITISH MUSEUM. 129

in both sexes, long and slender, of g setaceous and distinctly
serrate, of 9 but slightly dilated centrally and apicaily acuminate.
Not: ame wanting, mesonotum always shagreened and dull; meta-
notum also usually shagreened, ares wanting with areola at most
indicated ; costulze and spivacular carina alnrare wanting. Scu-
tellum convex, rarely pyramidal, always laterally and usually
also apically carinate. Abdomen elongate and in both sexes
subeylindrical, dull and always immaculate testaceous, with
hypopygium remote from terebral base. Legs slender and longer
than in the Asiatic species, testaceous with the hind ones rarely
black- and white-marked; tarsal claws simple. Wings ample
and always flavescent, with stigma testaceous.—Super ficially this
genus is similar, especially i in the shape of the capital vertex, to
festaceous species of Adoplismenus, from which the shagreened
exoskeleton, lack of conspicuous apophyses, paucity of metanotal
carine, and very much duller body sufficiently separate it, though
the scutellar structure is alike.

Table of African Species.

(22) 1. Flagellum white-banded in both sexes; metanotal
areola obsolete or wanting.
(21) 2. Head not abruptly declivous posteriorly, temples
distinct ; tarsi not white.
(10) 3. Apex of scutellum depressed and not at all cari-
nate.
(5) 4. Areola nearly circular; hind tarsi and tibie both
[SRYO oa aren auecncad toot 1. lutea Cam.
(4) 5. Areola rectangular or dentine at Faoee ‘ind lone
infuscate towards apices.
(7) 6. Smaller; stramineous; scutellum nel carinate
laterally ..
(8) 7. Larger; hagtageense : sovireliininn eiaonale erate
laterally,
(9) 8. Seutellum apically rounded ; postpetiole abruptly
explanate... Eh cats 3. rotundator Morl.
(8) 9. Seutellum oie ens nostaetale sradnally
explanaiicleeeente tec: 4. truncator Morl.
(3) 10. Apex of quia ales sta! andl enntinatie (pncrelhe
out.
(20) 11. Face regularly punctate; scutellum not longer than
basally broad ; hind tarsi mainly testaceous.
(19) 12. Mesonotum apically simple; postpetiole not stout.
(14) 18. Face distinctly punctate; metanotum dull and
not striate; its transcarina wanting ........... 5. inermis Morl.
(18) 14. Face obsoletely punctate; metanotum oni sal
not dull, its transcarina distinct.
(18) 15. Frontal orbits not pale; metanotal trans-striation

. debilitor Morl.

weak.
(17) 16. Postpetiole subcircular and abruptly explanate ... 6. explanator Morl.
(16) 17. Postpetiole slender and gradually explanate ...... 7. gracilator Morl.
(15) 18. Frontal orbits stramineous; metanotum strongly

striate . aa wesseeeeeess, 8. Sériator Morl.
(12) 19. Ntesonotume Evel saaeltexensice _postpetiole

Stoutier..aee. tecceseeseese 9. bipapillator Mori.

Proc, Zoo. Soc. a0) No. IX. 9

130 MR. CLAUDE MORLEY ON AFRICAN

(11) 20. Face irregularly punctate ; scutellum longer than
basally broad; hind tarsi entirely black ......... 10. collifer Morl.
(2) 21. Head abruptly declivous behind eyes, temples
obsolete; hind tarsi white-banded .................
(1) 22. Flagellum not white-banded ; metanotal areola
well defined.
(24) 23. Metanotum laterally black; areola dull; legs all
MESLEKECIORIS: “oon sacnbenoe Abo capcostue noo Sap hoa dé cagunEnHnED ac
(23) 24. Metanotum immaculate; areola shining; hind
(PTS, [AGI ccoaosoco con eau ocd anu Liscessessssseee 13. aveolator Morl.

11. cothurnator Morl.

12. geminator Morl.

1. LuTEA Cam.

Only known from the south of the Continent, in the Cape and
Natal; and apparently not extending far north, since I have seen
none of the present genus with entirely black hind tibize from
Rhodesia, &e. Cameron’s generic diagnosis is drawn entirely
from the 2, as I have pointed out (Ann. §. Afr. Mus. xvii. 1917,
p. 195). I possess the species from Zululand.

2. DEBILITOR, sp. n.

A small, pale, debilitant form, differing from the remainder of
the genus in its paler coloration, proportionately shorter antennz
of but 8 mm., the peculiarly small head with black ocellar region,
the shorter legs and, especially, in the weak lateral scutellar
carine, which do not extend to the apex. Length, 10 mm.
3 only.—The unique type was captured at Howick in Natal by
J. P. Cregoe, and presented to the British Museum in 1904,

3. ROTUNDATOR, Sp. n.

This is the first of a series of extremely closely allied species,
which I find to differ inter se in nothing but the facial punctura-
tion, mesonotal colour and apical contour, mesonotal and scutellar
and postpetiolar structure, as well as occasionally the colour of
the hind legs.—The present insect has the face obsoletely punc-
tate; mesonotum apically simple, with its sides rarely infuscate ;
metanotum dull and not striate, its transcarina wanting, and
areola both rectangular and obsolete; scutellum longer than
basally broad with its apex depressed, rounded, and not carinate ;
postpetiole laterally curved and abruptly explanate basally ; legs
testaceous, with the hind tarsi infuscate towards their apices.
Length, 12-15 mm. 6 2.—It is recognised by the scutellar and
postpetiolar structure, and by having the radial nervure distinctly
more reflexed than that of its congeners.—Quite the commonest
species of the genus in British Hast Africa, where S. A. Neave
found males between 7000 and 8500 feet on the east slopes of
the Aberdare Mountains towards the end of February 1911;
at 5000 to 6000 feet on the south slopes of Mount Elgon
in the middle of June; and 500 feet higher among some
forest on the Nandi Plateau at the end of the preceding May.
But the majority are from Uganda, where C. C. Gowdey took it
in the Mabira Forest at Chagwe in the middle of July 1911;

ICHNEUMONIN® IN THE BRITISH MUSEUM. 131

Neave in the Durro Forest at Toro, on Mount Kokanjero to the
S.W. of Elgon, at Mbarara in southern Toro, the Bugoma Forest
at Unyoro, and in northern Buddu at altitudes ranging from
3500 to 4500 feet during July and October.

4, TRUNCATOR, Sp. 0.

-Face obsoletely punctate; mesonotum apically simple; meta-
notum dull and not striate, with both areola and transcarina
wanting ; scutellum not longer than basally broad, apically de-
pressed and there abruptly truncate though not carinate ;
postpetiole slender and gradually dilated; legs testaceous, with
hind tarsi infuseate towards their apices. Length, 12-14 mm.
3 9. The only species with apex of scutellum truncate.—I have
seen but a single pair, of which the male is from the Mabira
Forest by Chagwe in July, and the female from Mbarara in
southern Toro during October, both at about 3800 feet in
Uganda.

5. inermis Morl.

Face evenly punctate ; mesonotum apically simple; metanotum
dull and not striate, areola rectangular and obsolete, transcarina
wanting; scutellum not longer than basally broad, apically
elevated and carinate throughout; postpetiole slender and
gradually dilated ; legs testaceous, hind tarsi infuscate towards
their apices. Length, 12-l4mm. ¢ 2.—This is the first of a
homogeneous group of species with the scutellum itself somewhat
flat, though the stout carina, which entirely surrounds its apex,
is distinctly elevated and conspicuous ; from its allies, the rectan-
gular and dull areola will distinguish it.—It was first described
by me in the female sex from Cape Colony, whence it appears to
extend as far as Ugandaand British East Africa, in both of which
its range is co-extensive with that of XY. rotwndator; i have seen
ten examples from the Mabira Forest, the Budongo Forest, near
Unyoro, Entebbe, lala in the Maramas District at 4500 feet,
and from the S.E. slopes of Mount Kenya between 6000 and 7000
feet during March, June, July, February, and December, 1911
to 1914.

6, EXPLANATOR, sp. n.

Face obsoletely punctate; mesonotum apically simple; meta-
notum shining and trans-striate, with both areola and transcarina
wanting; scutellum not longer than basally broad, its apex
elevated and carinate throughout ; postpetiole subcircular and
abruptly explanate basally; legs testaceous, with hind tarsi
infuscate towards their apices. Length, 14mm. ¢ only.—Dis-
tinct in the finely trans-striate metanotum which it shares with
the next species, and unique in the abrupt dilation of the post-
petiole.-—I have only seen the type of this species, which was
captured in the Gold Coast by W. P. Lowe and presented to the
British Museum in 1911.

132 MR, CLAUDE MORLEY ON AFRICAN

7. GRACILATOR, sp. n.

Face obsoletely punctate; mesonotum apically simple; meta-
notum striate and not dull, with areola wanting and transcarina
distinct though fine; scutellum not longer than basally broad,
apically elevated and carinate throughout; postpetiole slender
and very gradually explanate towards its apex; legs testaceous,
hind tarsi infuscate towards their apices. Length, 12-14 mm.
3 9. Differs from the last species solely in the presence of the
apical metanotal transcarina and the very slender postpetiole.—
A common species, of which I have seen a dozen examples from
the Durro Forest, Mabira Forest, and Buddu on the west shore
of the Victoria Nyanza at 3500 to 4500 feet during July, Sep-
tember, and October, in Uganda; and from a few miles east of
Mumias in the Maramas District of British East Africa at the
latter altitude in the middle of June 1911.

8. STRIATOR, Sp. Nn.

Very closely allied to the next species and differing from it
only in its simple mesonotal apex, its flavidous frontal orbits,
basally rufescent flagellar disc and somewhat shorter wings; from
this and the whole remainder of the genus it differs in having
the head fully as broad as the thorax and the metanotum strongly
trans-striate from its apical transcarina to base. Length, 15 mm.
? only.—The type, which alone I have seen, was captured on
17th February, 1911, by W. A. Lamborn, in Lagos.

9, BIPAPILLATOR, Sp. n.

Face evenly punctate ; mesonotum apically elevated on either
side into an obtuse tubercle; metanotum shining and trans-
striate, areola rectangular and obsolete, transcarina distinct ;
scutellum not longer than basally broad, apically elevated and
carinate throughout; postpetiole stout and gradually dilated ;
legs testaceous, with hind tarsi nigrescent. Length, 15-17 mm.
g only. Instantly known in this genus by the apically bituber-
culate mesonotum ; the large size, dark hind tarsi, and laterally
somewhat rounded abdomen are also distinctive.— The largest
species of the genus and apparently rare in Uganda, whence I
have examined four examples, taken by C. C. Gowdey, all at
Chagwe in the Mabira Forest between the 17th—20th July, 1911,
and on 3rd July, 1913.

10. COLLIFER, Sp. Nov.

Face irregularly punctate; mesonotum apically simple; meta-
notum dull and not striate, areola rectangular and_ obsolete,
transcarina wanting; scutellum distinctly a little longer than
basally broad, apically elevated and carinate throughout; post-
petiole slender and gradually dilated towards apex; legs testaceous,
with hind tarsi entirely nigrescent. Length, 15 mm. ¢ 9.—A
very distinct species, differing from all the above in its irregularly

IGHNEUMONINZ IN THE BRITISH MUSEUM. 133

punctate face, subvertical first recurrent nervure, and in having
the head behind the eyes constricted in a straight line, which
renders the eyes peculiarly prominent.—Apparently rare and
confined to Uganda, whence I have seen both sexes taken in the
Mabira Forest at 3500 to 3800 feet during July, and the Eee
Forest at 3400 feet during mid-December 1911, by 8. A. Neave

and about Entebbe during the middle of March 1914, by C. C.

Gowdey.

11. COTHURNATOR, sp. n.

A testaceous species, with both antenne and hind legs white-
banded black. Instantly recognised from the remainder of the
genus by the pure white third “and fourth (and in ¢ fifth) joints
of the hind tar si, which are peculiarly slender ; the whole thoracic
sculpture, including that of the scutellum, is as in XY. inermis,
but the abdomen is more fusiform and less elongate with its disc,
especially apically, more finely punctate and almost nitidulous;
the capital structure approaches that of XY. collifer, though the
temples here are very much shorter and the occiput falls away
immediately behind the black ocellar region. Length, 12 mm.
6 @.—The distribution seems distinct from that of the fore-
going species, for those I have examined are from 4500 to 5000
feet at Western Ankole in mid-October 19115; and the Siroko
River, near the west foot of Mount Elgon at 3600 feet, during
the preceding August; in Uganda.

12. GEMINATOR, Sp. 0.

This and the following species are so different from all the
above in their deeper ferruginous coloration, black antennz,
shorter and stouter legs, in the rather broader temples, much
longer upper mandibular tooth and more robust outline, that
they will probably not be found congeneric. X. geminator is
easily recognised by the longitudinal black mark occupying the
confluent external and dentiparal metanotal areze ; it is similar to
the next species but the legs are testaceous throughout, the
metanotal transcarina is strong with well-defined, subeirenlar and
dull areola, and no trace of costule. Length, 12-13 mm. ¢
only.—-‘Two males were captured on the N.W. and at Buddu on
the W. shores of the Victoria Nyanza at about 3700 feet by
Neave, in the middle of September 1911.

13. AREOLATOR, Sp. nD.

A ferruginous species with the legs neither elongate nor
slender, upper mandibular tooth long and transcarina of the
immaculate metanotum weak; flagellum entirely black, setigerous
and somewhat stout. Differing from the last species in the
possession of a well-defined elongate and shining areola with
some trace of costule ; and from the remainder of the genus in
having the extreme base (only) of the hind tibie, along with the

134 MR. CLAUDE MORLEY ON AFRICAN

whole of their tarsi, dead black. Length, 12-13 mm. ¢ only.—
Taken with Y. inermis and XY. rotundator in the Mabira and
Budongo Forests of Uganda at rather more than 3500 feet
during July and December.

HoPLisMENUS.

Gravenhorst, Ichn. Euvop. 11. 1829, p. 409; Wesmael,
Nouv. Mem. Ac. Brux. 1844, p. 13.

Head slender, not tumidous, somewhat narrowed behind the
eyes and towards the mouth; clypeus large, apically truneate ;
labrum usually very shortly exserted. Antenne slender, seta-
ceous; of ¢ subserrate, of 9 a little dilato-explanate, beyond
their centre. Thorax subcylindrical, discally gibbulous ; meso-
notum somewhat convex, with distinct notauli; metathorax with
basal sulcus profound, and the apophyses acute; spiracles linear
or elongate-oval. Scutellum strongly elevated, abruptly declivous
apically. Abdomen subfusiform ; hypopygium not covering base
of the distinctly a little exserted terebra. Legs somewhat slender.
Wings normal ; areolet pentagonal.

The bidentate metathorax, exserted terebra, &c., lend members
of this genus decided Cryptid facies ; but the males bear only the
faintest traces, and the females none at all, of sternauli. To
Platylabus it also bears a superficial resemblance but may be
distinguished therefrom, besides the major points above indicated,
by the straight and slender hind tibie and by the pentagonal
areolet. This genus has been placed by all former authors among
the Oxypygini, and so recently as March 1914 Herr Pfankuch
told me (in lit.): “It is my mind that Hoplismenus belongs to
the Ichneumonides”; but the conformation of the metathorax
and scutellum,as also the flagellar structure, ally it with the
Trogoide of Forster, and it was there treated of in my
‘Tchneumonologia Britannica’ in 1903; among the Oxypygini
it were certainly aberrant. Perhaps Acanthejoppa Cameron
(Entom. xxxv. 1902, p. 109) will prove synonymous, in which
case thirteen Asiatic species will be added; but the majority of
these have the head a great deal larger and more buecate than
any Hoplismenus of my acquaintance—though that its author was
ignorant cf the present genus is proved by his inclusion therein
of his H. ceylonicus (Spolia Zeylanica, ili. 1905, p. 100), which is
a EHupalamus and the male of his Melanichneumon kandiensis
(loc. cit. p. 99).

Africa may be considered the home of Hoplismenus, since more
kinds are known thence than from the remainder of the globe;
these include Jchnewmon dentatus Smith (Trans. Entom. Soe.
London, 1874, p. 391, n. 12), as I am able to state from an
examination of the Japanese 2 type. Kriechbaumer described
two species from Hast Africa in 1894; Tosquinet knew eleven
in 1896; Pie found another in 1897; while Szépligeti brought
forward no fewer than nineteen additional ones in 1910 from

ICHNEUMONIN® IN THE BRITISH MUSEUM. 135

Kilimanjaro, some of which may belong to Hoplojoppa, as was
indicated by Dr. Roman (Entom. Tidskr. xxxi. p. 169) in 1910.

1, FULYVATOR, Sp. n.

Pale testaceous with only part of flagellum, apices of mandibles
and of hind onyches, black. Head obliquely constricted and
narrow behind the very prominent eyes, whence the occiput falls
abruptly away ; frons obsoletely punctate, trans-substriate before
the ocelli and centrally sulcate ; face and clypeus nearly strami-
neous, flat and obsoletely punctate, with apex of latter truncate
and only laterally margined ; labrum exserted and cheeks long.
Flagellum slender and filiform, black with a central white band,
its base rufescent and apex attenuate; of ¢ subserrate, of 9 dis-
tinctly a little dilated, beyond centre. ‘Thorax cylindrical ;
mesonotum dull and shagreened, with slight notauli ; metanotum
shining and evenly punctate with complete, but fine, arez ;
areola subrectangular, half as long again as broad, extending
nearly to base, and emitting conspicuous costule from its basal
third; spiracular are trans-striate; apophyses large, vertical
and acuminate. Scutellum much higher than metathorax, its
dise closely punctate and not strongly elevated, but both laterally
and apically stoutly vallate. Abdomen dull; basal segment very
slender with the shining and laterally punctate postpetiole very
little explanate ; gastrocceli wanting, thyridii obsolete and their
intervening space not striate; venter plicate throughout ; anus
of 2 acute, with the rufescent terebra distinctly a little exserted.
Legs long and slender; calcaria elongate, coxee not scopulate.
Wings flavescent hyaline; stigma and subcosta testaceous, ner-
vures darker; areolet pentagonal, subcoalescent above ; nervelet
wanting ; basal nervure continuous through median. Length,
11-13 mm. ¢ 9.—Similar to H. fulgens Tosq., but with the
metanotal arez distinct and scutellum carinate throughout its
apex.—I have seen half a dozen examples of both sexes from the
Tero Forest, 8.E. of Buddu, at 8800 feet, towards the end of
September; from Fort Portal Road, Mbarara, Southern Toro, at
about 4000 feet, late in October; from the Durru Forest, 'Toro,
at 4000-4500 feet, a few days later, in Uganda; and from Ilala
in the Maramas District, 14 miles east of Mumias at 4500 feet,
during the middle of June, in British Hast Africa,

2. CIBDELUS Tosq.

No one has referred to this species since the ¢ was brought
forward (Mem. Soe. Entom. Belg. v. 1896, p. 52, n. 3) from
Dinko, Scioa, in Abyssinia, where “Ragazzi captured it in 1887.
The 2 is a very dull, claret-red coloured insect, with the meta-
thorax entirely, mesopleurze except above, hind tarsi, prosternum
partly, and apical half of flagellum, black; the frontal orbits
triangularly, flagellar band, pronotal margin, and anus white.
It is at once recognised by the distinctly and evenly punctate

136 MR. CLAUDE MORLEY ON AFRICAN

frons; by the shining and very sparsely punctate scutellum, of
which the lateral carine extend nearly to the apex ; the rugulose
and white-pilose metanotum with elongate and subhexagonal
areola, which in the Q is dull and only basally aciculate ; and
by having the anus from apex of the nigrescent fifth segment
clear white. Length, 14 mm.—Captured between Jinja and
Busia in some forest to the east of Busoga at 4000 feet, towards
the end of July 1911, in Uganda.

CoRYMBICHNEUMON, g.n.

This new genus differs from Celichnewmon Thomson solely in
having the scutellum spherical; it stands out from the thorax
in the form of a one-third embedded ball, and is not laterally
carinate. The only specics yet known is remarkable for being
of sombre coloration, with both the -scutellum and mesonotum
conspicuously pale stramineous.

‘© Monobasic.”

1. CARINIFER, Sp. n.

©. A stout, claret-coloured insect with the mesonotum and
scutellum dull pale lemon-yellow. Head dull, closely punctate,
sometimes black, and very short behind the eyes; palpi pale
flavous, cheeks buccate, clypeus broad and apically truncate,
hardly discreted from the deplanate and centrally subelevated
face. Antenne black, filiform and stout, with a 10-jointed
central white band; basal flagellar joint nearly as long as the
two following, which are apically subexplanate. Thorax stout,
very finely punctate, black or with only the frenum and tegule
black ; mesonotum, base of propleure above, and callosities below
radix pale flavous; metathoracic carine strong and entire; areola
aciculate, twice as long as broad, extending to base and emitting
costule slightly beyond its centre; spiracles elongate, apophyses
wanting. Scutellum pale flavous, globulai and glabrous; post-
scutellum small and concolorous. - Abdomen stout, subdeplanate
and immaculate; basal segment broad and apically explanate
with postpetiole punctate-aciculate; second segment deplanate
and finely punctate-aciculate, with large gastroceeli; second and
third ventral segments plicate; anus acute and hypopygium
remote from base of the black and distinctly a little exserted
terebra. Legs stout and elongate, indefinitely varied with black ;
hind coxe finely punctate, simple; their tarsi setiferous beneath,
with large and simple claws. Wings evenly infumate-flavescent ;
stigma and nervures black ; areolet no higher than broad, broad
above ; discoidal cell subparallel above and below.— ¢ differs in
having all the above red parts black; the pale flagellar band
further from the base; the abdomen and _ scutellum more
deplanate; the metanotum laterally punctate, with no costule ;
the front tibiz anteriorly testaceous and the anterior coxe,

fing Sot eh

ICINEUMONIN IN THE BRITISH MUSEUM. Ata

underside of scape and whole face with mouth-parts, pale flavous.
Length, ¢ Q 14-18 mm.—A couple of females were captured by
Neave in Nyasaland (on the Mlanje Plateau at 6500 feet on
November 10th, 1913) and in British East Africa (on the 8.E.
slopes of Mount Kenya at between 6000 and 7000 feet early in
February 1911); the single male, I have reason to believe, is that
recorded by the late Colonel C. T. Bingham under the erroneous
genus Lactolus, in his Report of the Ruwenzori Expedition,
No. 13, Hymenoptera (Trans. Zool. Soc. Lond. xix. pt. 2, 1909,

. 179); it was captured at Salt Lake on Ruwenzori at some
7000 to 8000 feet in 1895 by Scott Hlliot.

AGLAOJOPPA.
Cameron, Ann, Nat. Hist. vil. 1901, p. 381.

In my Revision of the [chneumonide (iv. 1915, p. 112), this
genus is recognised as sufliciently distinct from Celichnewmon ;
and Tables are given of the European, Oriental, and American
species, though none were known from Africa till 1917, when the
Ann. 8. Afr. Mus. published one that is not dissimilar to the
two following kinds.

1. JOHANNIS Cam.

Ichneumon johannis Cameron, Records Albany Museum, 1.
1905, p. 245, ¢.

2. A dull, black species with the mesonotum, scutellum, and
sides of thorax sanguineous-red ; vertical and a line at the external
orbits, flagellar band, postscutellum, apical angles of the three
basal and whole of the 6th—7th segments, with inner side of the
front tibie, white, Head posteriorly short, but nearly as broad
as the eyes; frons closely and not finely punctate; face and the
undiscreted clypeus closely punctate, apex of latter truncate and,
like apex of cheeks, rufescent; labrum exserted and flavidous.
Thorax not very dull, very finely punctate, with elongate notauli;
metanotum deplanate; areola rectangular, longer than broad,
subglabrous, emitting strong costule from a little beyond its
centre ; spiracles elongate; spiracular and lateral aree alone red.
Scutellum deplanate, glabrous, sparsely punctate, carinate only at
its base, apically broad and truncate. Abdomen subovate and
short ; basal segment apically badious with postpetiole punctate,
its apical angles and those of the two following segments white,
with the third except narrowly in its centre apically white
throughout, as also is the anus. Legs normal, badious; hind
coxe mutic. Wings hyaline, stigma black, areolet higher than
broad and broad above, nervelet nodiform. Length, 13 mm.
Remarkable for the glabrous and rufescent callosity at the juxta-
antennal orbits —This female was captured at Salisbury in
Mashonaland by Marshall during March 1900; the male type,

138 MR. CLAUDE MORLEY ON AFRICAN

which I have seen, is from the Cape; and W. E. Jones has
subsequently found the species at Mfongosi in Zululand.

2. GLABRINOTOR, Sp. D.

Very like the last species, but with the head posteriorly bueccate
and fully as broad as the eyes, not at all black but bright red
with only the orbits of its utterly glabrous frons and the palpi
white. Thorax strongly nitidulous, with a white callosity beneath
radices ; mesonotum extremely isolatedly punctate with but short
notauli : metathorax deplanate with areola sublinear, fully
thrice longer than broad, glabrous and emitting weak costule
from beyond its centre; external and lateral are badious.
Scutellum glabrous and glittering with some half dozen pune-
tures, and laterally carinate to its centre. Abdomen fusiform
and apically acute; postpetiole badious, sparsely punctate, with
no carine; fourth and following segments nitidulous; apical
angles of the second narrowly and whole of the 6th—7th white.
Legs normal; anterior femora and tibie internally whitish ;
hind coxal scopule large. Length, 13 mm. @ only.—The
sublinear metanotal areola is not unlike that of the genus Crato-
joppa Cam. (Ann. Nat. Hist. vii. 1901, p. 281), in which the
scutellar cavine are wanting, the areolet coalescent above, the
head large, &e., which characters do not here obtain.—Found by
Neave in Nyasaland to the 8.W. of Lake Chilwa on 16th January,
1914.

3. COMPTULUS Tosq.

Ichneumon comptulus Tosquinet, Mém. Soc. Entom. Belg. v.
T8936; pote Ov

3 2. Klongate, punctate, black and somewhat dull. Head
quadrate, a little shining, finely punctate; black with all the
orbits, except in ¢ the vertical, somewhat broadly white; face
broader than long, transversely striate-punctate; clypeus hardly
discreted, its apical margin rounded and narrowly, or in ¢
laterally, white. Antenne short; of 2 a little dilated centrally,
attenuate apically, black, and white-banded; of ¢ also black,
servate, sed fracte. Thorax rosy, punctate, with sternum black ;
an elongate line before and a linear callosity below radices, and
in ¢ a pronotal mark, white; metathorax convex, strongly punc-
tate, with long griseous pilosity and complete arez, of which the
areola is elongate, apically rounded and more finely punctate,
of § centrally nigrescent. Scutellum somewhat convex, smooth
or in ¢ punctate, laterally carinate nearly to apex, which in ¢d is,
along with carine at base and whole postseutellum, white.
Abdomen very closely punctate, longer than head and thorax,
of ovate, of ¢ cylindrical ; black with apices of the second and
third segments narrowly rufescent and the seventh broadly in its
centre, along with the ¢ first apically, and three following on
either side, white; terebra a little exserted, ¢ valvule black

see

ICHNEUMONIN® IN THE BRITISH MUSEUM. 139

and concealed. Legs short and stout, black with front tibie dull
stramineous; anterior legs of 2 obscurely red, of ¢ with coxe
white. Wings subhyaline, with an indefinite cloud across the
areolet; stigma, nervures and tegule black, radix whitish ;
areolet irregularly pentagonal, nearly coalescent above ; nervelet
small; lower basal nervure distinctly postfureal. Length, ¢ 9
15 mm.—It was originally described from Togoland, but two
males taken by Neave differ so little that I have ventured to
regard them as the opposite sex of Tosquinet’s species. They
occurred in the valley of the South Rukuru River between
June 20th and 27th, 1910, at 3000 feet in Nyasaland.

Ca@LICHNEUMON.

I do not think any species have yet been recorded from Africa
(excepting my note in Revis. Ichn. iv. 1915, p. 121) under this
name ; but seven of those described by Berthounieu (Ann. Soc.
Ent. France, 1894) under the genus Jchnewmon from Algeria
seem to belong here, with C. hemorrhoidalis Grav., C. rudis
Fonse., and C. sinister Wesm., which extend from southern
Kurope. I find Platylabus maculiscutis Cam. and Lchneumoin
natalensis Cam. to also belong here ; and the dozen species is
completed by one brought forward in the Ann. 8. Afr. Mus.
by me in 1917, Probably some of Tosquinet’s and Kreich-
baumer’s Jchnewmones will also have to be included, when the
types are re-examined. Holcichnewmon Cameron (Ann. Mus.
Transvaal, ii. 1911, p. 175) is synonymous.

Table of some African Species.

(8) 1. Black or castaneous, with pale flavous-marked
thorax.

(3) 2. Hind cox stoutly scopulate; scutellum glabrous. 1. scopulifer Morl.
(2) 38. Hind coxe mutic; scutellum distinctly punctate.
(5) 4. Metanotal areola longer than broad; scrobes
tuberculates. crass. ce ceesrsets ce ecacntereereeese eae | Ze cormellafer Mors
(4) 5. Metanotal areola not longer than broad; scrobes
simple.

(7) 6. Abdomen and hind legs red ; petiolar area black . 3. maculiseutis Cam.
(6) 7. Abdomen mainly and hind legs black; petiolar

axeaypale-marked) jy...0.a. ce «ca--- ss-easoseeeeseeeen 4) Ratdlensis Cams
(1) 8. Testaceous; rarely with head and thorax black-

marked.
(12) 9. Mesonotum nitidulous; head and thorax pale

stramineous-marked.
(11) 10. Postpetiole aciculate; gastrocceli large; scutellum

WiARTEE) BL OAe Haas. niles dashigats-Sulen da ah nea sae eae | Ooe SAUD Leave fere Mon,
(10) 11. Postpetiole punctate; gastrocceli small ; seutellum
TREE Senne 6. geminifer Morl.

(9) 12. Mesonotum dull and shagreened; rarely dull
flavous-marked.
(14) 13. Areola narrow and elongate; lower basal nervure

Postiuncaly atx, oes caet sesshre coon dpa easnreeenm «coumadin LeStaceue Cam.

140 MR. CLAUDE MORLEY ON AFRICAN

(18) 14. Areola broad, not elongate; lower basal nervure
continuous.

(20) 15. Second abdominal segment neither striate nor

black-banded.
(17) 16. Scutellum convex and subcircular; areola glabrous 8. globulifer Morl.
(16) 17. Scutellum deplanate and elongate; areola dull and

shagreened.
(19) 18. Second segment transversely impressed ; gastro-

UPN OLSON Ey Se pnosceseav ens ous osenaddonode dic ccaaddccdons — Ga SUleafeEP Ailloyell
(18) 19. Second segment entire; gastrocceli large and trian-

gular . 10. thyridifer Morl.

(15) 20. Second segment black-banded and strongly striate. 11. striatifer Morl.

1. SCOPULIFER, Sp. n.

A claret-colowred and somewhat shining female, with the
occiput and frenum indefinitely black; all the orbits but the
temporal, the mandibular base, sides of clypeus, flagellar band,
prothoracic margin, lines before and beneath radix, a central
mesonotal mark, two more on mesopleure, sides of petiolar area,
the scutellum and postscutelluim, stramineows-white. Head pos-
teriorly short, but as broad as the eyes; frons distinctly and face
not closely punctate, the latter transverse, centrally prominent
and not discreted from the apically truncate clypeus, which is
centrally subdentate. Thorax shining, with distinct notauli;
metathorax evenly punctate and not dull; areola obviously
longer than broad, parallel-sided, glabrous, apically incomplete
and emitting weak costule from its centre. Scutellum deplanate,
subquadrate, glabrous with a very few fine and scattered punc-
tures, apically truncate and not at all laterally carinate. Abdomen
elongate fusiform with apices of segments a little prominent ;
three basal segments closely punctate and somewhat dull; post-
petiole and base of second segment punctate-striate, the former
basally bicarinate; gastroceeli large and deeply impressed ;
terebra black and a little exserted. Legs indefinitely nigrescent-
red with apices of front coxe broadly, and of the intermediate
narrowly, white; hind coxal scopule large. Wings hyaline,
stigma and tegule black, radix white; areolet pentagonal, broad
above; nervelet wanting, second recurrent nervure externally
produced at its centre. Length, 12mm. 9 only.—Captured at
Mlanje on 15th August, 1913, in Nyasaland.

2. CORNELLIFER, Sp. 0.

A black species with abdomen except petiole, metathorax
except petiolar area, mesopleure and most of the legs, red; all
the orbits, cheeks, face except a central didymate mark, clypeus,
mandibles, the @ flagellar band, prothoracic margin, lines before
and beneath anterior radix and behind the posterior, a central
mesonotal and two mesopleural marks, scutellum, postscutellum,
apical angles of the basal segment minutely, anterior coxe and
trochanters entirely and the hind ones discally, stramineous ;
3 also has the basal segment except its angles black, and the

ICHNEUMONIN. IN THE BRITISH MUSEUM, 141

petiolar area stramineous. Remarkably like the last species but
differing in its colour, puncturation, especially of the frons and
mesonotum and scutellum ; in the much stronger costule, at least
basally black first segment with its pale apical angles; in the
indication of a nervelet and the simple second recurrent nervure ;
in its somewhat stout facial tooth between the scrobes ; and also
in the lack of 2 coxal scopule. Length, 14-15 mm. ¢ 2?.—
The type is from Blantyre in Nyasaland on 9th June, 1910
(Dr. J. HE. S. Old); and the male occurred in the Gold Coast
about the same time (IV. P. Lowe).

3. MACULISCUTIS Cam.

Platylabus maculiscutis Cameron, Ann. 8. African Mus. v.
1906, p. 179, 3; Ichnewmon transvaalensis Cameron, Ann. Mus.
Transvaal, ii. 1911, p. 174, @.

An examination of both types convinces me of the above
synonymy. The ¢ is from Klerksdorp in November 1890, and
the @ is labelled ‘‘ Fonteinen, 24th April, 1905,” both in the
Transvaal. A pair of homotypes were taken in South Africa

so long ago as 1845, and in the Ruo Valley, Nyasaland, at
2000 feet on October 25th, 1913.

4, NATALENSIS Cam.

Ichneumon? natalensis Cameron, Ann, 8. African Mus. v, 1906,
p- 160, 3.

The details contained in the above Table are drawn from the
type in the British Museum. Another dg, also labelled “ type”
by Cameron, is in the 8. African Museum ; taken at Malvers in
Natal by Barker.

_0. SUBLUNIFER, Sp. 0.

A small, fulvous and not testaceous female with the occiput,
vertex, frons, and scrobes to radicule, antennz except underside
of scape and their white band, mesonotum and a mark in each of
the external and dentiparal ares, black; remainder of the head,
prothorax, elongate callosities before and beneath radices, two
discal mesonotal lines, mesopleurz except speculum, scutellum
except lateral carinz, postscutellum, anterior coxe and trochanters
and dise of hind cox, very pale stramineous. Head transverse,
narrower than eyes and posteriorly but slightly emarginate : face
closely but weakly, and frons obsoletely, punctate; clypeus sub-
discreted, apically truncate and centrally a little impressed.
Mesonotum very closely punctate and not shagreened, with
distinct though fine notauli; metanotum shining, finely punctate-
aciculate, with petiolar area long and broad, discreted ; areola
proportionately short, subsemilunate, broader than long, basally
rounded and emitting weak costule from its centre. Scutellum
deplanate, glabrous, laterally carinate to near its truncate apex,
before which it is traversed by a few longitudinal striations.

142 MR. CLAUDE MORLEY ON AFRICAN

Abdomen immaculate fulvous, fusiform; postpetiole centrally
very finely aciculate, laterally punctate; gastroceli large, deep
and transverse, with the narrow intervening space evenly punc-
tate; terebra black and exserted. Legs normal; hind cox
mutic. Wings hyaline; radix and tegule stramineous, stigma
castaneous ; areolet small and pentagonal, nervelet nodiform ;
basal nervure continuous through the median. Length, 7 mm.
© only.—Captured at Northern Buddu at 3800 feet during the
middle of September 1911 in Uganda.

6. GEMINIFER, Sp. Nn.

A rufescent-testaceous female, very like the last but larger with
the mesonotum nitidulous and only sparsely punctate, its dise red
and sides occasionally black ; the scutellum is rufescent with its
lateral carine and instriate apex stramineous; the metanotal
areola, though similarly shaped, is much larger; the abdomen
testaceous with poustpetiole evenly and distinctly punctate
throughout; gastroceeli small and much narrower than their
intervening space; terebra basally rufescent ; cox testaceous,
with no stramineous markings. Length, 12-14 mm. 2 only.—
A couple of females (one with the metathorax testaceous and
only geminate spots as in the Jast species; the other with the
external and dentiparal arez black and a stramineous pleural
streak) at Mlanje in September and October 1913 at 2300 feet
in Nyasaland. Also found by W. E. Jones, during March 1917,
at Mfongosi in Zululand.

7. TESTACEUS Cam.

Holcichnewmon testaceus Cameron, Ann. Mus. Transvaal, ii.
LOM yp: iia, os

I have examined this unique, bred in the Transvaal, and find
it belongs here.

8. GLOBULIFER, sp. n.

Males differing from the next species (C. sulcifer) solely in
their general flavous coloration, lack of metapleural marks ;
in the subglabrous postpetiole, simple second segment with
normal gastroceeli; in the black ocellar region, hind tarsi and
hind tibize, with pale calearia; in the subcircular metanotal
areola, the apically coalescent alar areolet; and in the sub-
conically elevated scutellum with long black pilosity. Length,
13mm. od only.—A couple of males were found at Kibwezi at
3000 feet, at the beginning of April 1911, in British East Africa;
and in the Tero Forest, S.E. of Buddu, at 3800 feet, at the end
of September 1911, in Uganda.

9. SULCIFER, Sp. Nn.

A very dull, rufescent-lestaceows species with only the face
subflavidous ; flagellum except the central white band, sides of

ICHNEUMONIN® IN THE BRITISH MUSEUM. 143

mesonotum, a common streak in the external and dentiparal
ares, alone black; apices of the hind tibie and their calearia
nigrescent. Head obsolete behind the eyes; frons and the
centrally elevated face hardly punctate; clypeus not discreted,
apically truncate; mandibles not apically darker. Mesonotum
dull and shagreened, with short notauli; metathorax dull ee
dentiparal are transaciculate; areola dull and shagreened,
little longer than broad and rounded at both extremities : sotals
utterly “wanting. Scutellum elongate - triangular, deplanate,
shining and sparsely punctate, with no pilosity, laterally mar-
gined to near apex. Abdomen elongate-fusiform, unicolorous ;
basal segment closely and evenly punctate; the second trans-
versely impressed throughout at its basal fourth; gastrocceli very
small; terebra black and a little exserted. Legs somewhat short
and not slender, coxal scopule wanting. Wings flavescent-
hyaline; radix, tegule, and stigma entirely pale testaceous ;
areolet pentagonal and broad above; nervelet and another in
second recurrent nervure, short. Length, 12 mm. @ only.—
Several females captured in Britisi East Africa (on the Yala
River at the southern edge of the Kakumga Forest, at about
5000 feet, towards the end of May 1911) and Uganda (in the
Durru Forest, Toro, at 4000-4500 feet, at the end of October
1911; and at Fort Portal Road, Mbarara, in Southern Toro, at
some 4000 feet, a day or two earlier). Also taken by G. F. Leigh
on Ist March, 1910, at Pinetown in South Africa.

10. THYRIDIFER, sp. n.

A very dull, rufescent-testaceous female with only the face and
two elongate vittz on the black mesonotum subflavidous. Very
similar to the last species, but differing in a few essential details;
ocellar region and centre of occiput confluently, mesonotum
centrally as well as laterally, scutellum centrally, basal suleus
only of metathorax, with whole of the hind tibize and tarsi,
black; metanotal areola smaller, emitting distinct though weak
costule ; centre of scutellum more closely punctate and sub-
convex ; postpetiole obsoletely punctate-aciculate ; gastrocceli
deeply impressed, somewhat large and as broad as their inter-
vening space ; areolet coalescent above,with the second recurrent
nervure simple. Length, 14 mm. 92 only.—Harrar in Abys-
sinia, during 1912 (Collector ignot.).

11]. STRIATIFER, sp. Nn.

Rufescent-testaceous with the face and orbits indefinitely, meso-
notal margins and whole hind tarsi, flavidous ; ocellar region
circularly, flagellum except the white band, sides only of meso-
notum brovdly, whole external arez, spiracles of basal segment,
a transfascia before centre of the second segment and at base of
the third, black. Head posteriorly strongly emarginate, with
obsoletely punctate frons; flagellum serrate. Anterior margin

144 MR. CLAUDE MORLEY ON AFRICAN

of mesonotum sulcate and strongly elevated ; metanotum punc-
tate, with the dentiparal aree aciculate and areola hexagonally
quadrate, emitting strong costule from its centre. Scutelium
subquadrate, deplanate, elevated from base to the truncate apex,
laterally margined throughout, and much higher than metanotum.
Abdomen dull and closely punctate with the postpetiole, second
segment to near its apex and the third to centre, longitudinally
striate ; gastroceli deeply impressed. Areolet triangular, almost
broader than high, coalescent above. Length,11 mm. ¢ only.—
Doubtless this species were more correctly placed (on account of
its distinctively striate basal segments, the apices of which may
be said to be slightly produced laterally) in one of the more
typically Joppid genera, though its facies are sufficiently
analogous with the present one.—A single male was captured

at Chagwe in the Mabira Forest at some 3500 feet during the
middle of July 1911, in Uganda.

Tribe [CHNEUMONIDES.
Subtribe Oxypygini.

LAGENESTA.

Head large and not strongly constricted posteriorly ; occiput
margined ; labrum strongly prominent, clypeus apically truncate ;
mandibles bidentate, the upper tooth the longer. Antenne stout,
dilated beyond their centre. Metanotum transversely punctate,
with no distinct ares ; petiolar region transversely carinate below
its centre. Seutellum deplanate and not laterally carinate.
Abdomen neither punctate nor striate, with sides of segments
not angularly prominent apically ; apical third of petiole broadly
dilated; gastroceli large and stoutly striate, but not deeply
impressed ; ventral plica strong on second and third, weak on
fourth and fifth, segments. Legs stout with tarsal joints spinose,
explanate and basally constricted; hind femora extending to
fourth segment. Areolet large, pentagonal, and broad above,
emitting recurrent nervure from its centre; radial nervure
apically reflexed.

The above are the characters assigned to this genus ky Cameron,
at the time of its erection (Ann. Nat. Hist. vii, 1901, p. 376) ;
but they are entirely drawn from the female of one species, of
which he knew the male later (lib. cit. xi. 1903, p. 185) and the
only other published species four years afterwards (Zeits. Hym. u.
Dipt. v. 1905, p. 86, 9 ). These are all from the Hills of northern
Eada, ° whence I have ween a third kind*; and have no doubt

* LAGENESTA MONITOR, sp. n.

A cylindrical, brick-red and cyaneous-black species, with infumate wings. Head
cubical and broader than thorax, with the frons to ocelli and mandibular apices
alone black; occiput strongly margined ; vertex centrally rugose, laterally smooth
and behind the eyes strongly elevated into stout and obtuse spines; frons rugose
and centrally produced into a stout spine between the scrobes; temples longer than

ICHNEUMONIN® IN THE BRITISH MUSEUM. 145

that the original account of the genus will have to be sufficiently
modified to embrace the following additional African kinds, which
I have compared with the types of both Cameron’s species. He
himself, while placing it in the Joppides, was doubtful of its
position and remarks that it “does not fit well into any of
Kriechbaumer’s groups of the Hemijoppine”; as a matter of
fact, the total lack of basal metanotal sulcus in the genotypical
specimen, with its very flat scutellum, places it (I consider,
certainly) in the Ichneumonides. Of palearctic groups, it is
most closely allied to Melanichnewmon and especially Hupalamus.

1. SINIFER, sp. n.

A large and stout, dull black species, with head and mesonotum
shining ; wings evenly nigrescent, head entirely red, antenne and
hind tarsi white-banded. Head and scape rich red and glabrous ;
former strongly buccate behind eyes and nearly as long as broad,
posteriorly strongly emarginate, with occiput bordered; scrobes
externally produced below; face and clypeus sparsely and coarsely
punctate, the latter hardly discreted, apically truncate, laterally
rounded and elevated; mandibles stout and sparsely punctate, with
teeth black. Antenne subfiliform; of 9 very slightly dilated
beyond centre, with the 10th to 18th flagellar joints white; of ¢
subserrate, with the 19th to 25th flagellar joints white. Thorax
very stout and black with prothorax and in Q apices of both
scutellum and postscutellum with most of frenum, rufescent ;
mesonotum glabrous, with a few irregular fine punctures; notauli
short and very deeply impressed; metathorax evenly scabriculous
throughout, not basally suleate; of § with petiolar carina alone
distinct; of 92 with areola weak, elongate-oval, longer than
broad. apically acuminate, with its carine weaker than the
centrally emitted costule ; petiolar area short and not discreted;

eyes; face quadrate, longitudinally punctate and laterally sulcate before the elevated
orbits; clypeus continuous, smoother, apically broadly rounded with its angles
strongly elevated and centre minutely bidentate; labrum concealed, cheeks not
elongate, lower mandibular tooth slightly the shorter. Antenne setaceous, not very
slender, red with the apical third black and central joints subserrate. Thorax
cylindrical, dull and closely punctate, brick-red with all the sutures black; meso-
pleure, dentiparal arese and the parallel-sided areola all apically produced;
metanotum deplanate and double as long as petiolar area; basal area obsolete,
costule wanting: spiracles linear. Scutellum and postscutellum red, evenly
punctate and subdeplanate, with basal fovea and carine black. Abdomen parallel-
sided and slender, double as long as thorax, obsoletely punctate, subnitidulous and
black, with venter concolorous and only the second and third segments strongly
plicate; basal segment glabrous and red, with obsolete punctures and a discal
longitudinal fovea; thyridii wanting, valvule exserted. Legs elongate and some-
what slender ; deep red with hind tibie and tarsi alone black ; hind femora extending
only to apex of second segment. Wings ample and evenly infumate; radix and
tegule red, stigma and nervures black; areolet exactly pentagonal, broad above,
emitting the broadly bifenestrate recurrent nervure from its centre; discoidal cell
nearly parallel-sided ; nervelet pellucid, lower basal nervure postfurcal ; nervellus
intercepted at its lower third. Length, 22 mm. ¢ only. Type in Mus. Brit.
Assam.

Proc. Zoou, Soc.—1919, No. X, ; 10 5

146 MR. CLAUDE MORLEY ON AFRICAN

spiracles large and apophyses wanting. Scutellum deplanate,
glabrous with a few fine scattered punctur es, laterally carinate to
near apex. Abdomen dull and elongate- fusifor m; basal segment
shagreened ov in ¢ nearly smooth, with postpetiole strongly
explanate and flat; second segment not at all striate, its gastro-
coeli deep and a little narrower than the intervening space ; plica
on second to fourth ventral segments, and dise of 2 seventh
segment, entirely white; terebra subexerted, black. Legs normal ;

inner side of anterior tibie and of front femora, with whole hind
tarsi from second segment, white; hind cox of 9 with large,
brown scopule ; claws large. Wangs deeply infumate throughout,
with cyaneous reflection ; areolet triangular and coalescent above ;
nervelet short; basal nervure continuous through median; first
recurrent nervure (of Iehn. Brit. i. 1903, p. xxxvi) sinuate at
apex of anal nervure. Length, 16-17 mm. ¢g 2.—The ¢ was
found in some forest land between Jinja and TMbyagoe east of
Busoga, at about 4060 feet on 28th July, 1911, in Uganda; and
females at Mount Mlanje on 29th November, 1912 and again on
Ath February, 1913,in Nyasaland. W. EH. Jones took another 9
during March 1917, at Mfongosi in Zululand.

2. DUPLICATOR, Sp. 1

A large, rich crimson species, with slightly flavescent-hyaline
wings; frons triangularly and broadly to below ocelli, the 9 10th
to 16th flagellar joints, the sixth and seventh segments discally,
ventral plica and apices of all the ventral segments, white ;
mandibular apices, remainder of flagellum and the whole of third
to fifth segments, black; hind coxal scopule strong, but not large.
In sculpture this species differs from the last in little but its
rugulose metanotum with discreted petiolar area, much shorter
capital vertex, distinctly postfureal lower basal nervure and
broader areolet which is not laterally coalescent above.—The ¢
also has the whole clypeus, face, centre of external orbits, under-
side of scape, pronotal margin, a linear callosity below radix, the
postscutellum, apical margins of the second and third segments
narrowly, and the three apical joints of its basally black hind tarsi,
white. Length,17 mm. 6¢ 9.—The female is from 2300 feet ae
Mlanje on 21st October, 1913, in Nyasaland ; and the male was
found at South Kavirondo at 4200 feet early in May 1911 in
the Upper Kuja Valley of British Kast Africa. In Zululand,
Mfongosi, South African Museum.

3. TRIPLICATOR, Sp. ll.

So similar to /. duplicator as to need no detailed description ;
therefrom I am able to distinguish it only by its rather less clear
red coloration, smaller size, more slender and more parallel-sided
abdomen, much narrower frontal white markings, the entirely
erimson third abdominal segment, and the utterly hyaline wings
with their areolet quite coalescent above and basal nervure exactly

ICHNEUMONINA IN THE BRITISH MUSEUM, 147

continuous through the median. Length, 15-l16mm. ¢ 9.—
Apparently not an uncommon species about Mlanje, where a
series was captured during January, February, and March, 1913

and 194.

A, TRIANGULIFER, Sp. 1.

A stout, shining, black female with the scutellum and its frenal
sides, basal fovere and centre of mesonotum throughout, meso-
sternum except centrally at the base, meso- and meta-pleure
entirely, rosy; both clypeus and frons except longitudinally in
their centre, central flagellar band except below, postscutellum,
apices of first and sixth segments narrowly, of the second broadly
on either side and whole of the seventh with most of venter,
anterior coxse and (excepting a discal streak) trochanters, with
the four apical joints of hind tarsi, white; anterior legs mainly
pale, hind femora and tibie rufescent black. Wings hyaline;
hind coxal scopule large. Mesonotum glittering and only sparsely
punctate ; scutellum absolutely glabrous, laterally black carinate
to near its truncate apex; metanotum evenly and scabrously
punctate with no are but an elongate-triangular (wedge-shaped)
portion, gradually contracted apically, glabrous and nitidulous;
pleural carina strong. Postpetiole deplanate, explanate and
shagreened. Stigma black; areolet coalescent above; lower basal
nervure a little postfurcal. Length,15 mm. @ only.—A con-
spicuous species with, I believe, unique metanotal structure ; and
obviously belonging to the present genus by the analogy of the
puncturation and frontal-colour distribution, besides more per-
tinent characters, though superficially very ike Melanichneuwmon.
-—It was found in the Durro Forest, Toro, at fully 4000 feet,
between 23rd and 29th October, 1911, in Uganda.

MELANICHNEUMON.

This genus was divided from the broad division, Zehnewmon in
the Wesmaelian sense, by Prof. Thomson so long ago as 1893
(Opuse. Entom. xvill. p. 1954); but was therein again merged
by both Berthoumieu in 1894 and Schmiedeknecht in 1903.
Thomson’s groups are perfectly natural and are now, all too slowly,
coming into general recognition: they were adopted by me in
1903. Consequently, we must bear in mind that in the case of
the present and few following genera, other species may already
exist under the old broad heading of /chnewmon. Smits van
Burgst has recorded (Tunisian Hymen. 1913) three European
species of Melanichnewmon extending to North Africa; I have
described two new kinds (Ann. 8, Afr. Mus. 1916); and
Dr. Roman has shown (Zool. Bidr. Upsala, i. 1912, p. 262) that
TIchneumon leucophthalmus, 'Thunberg 1822, belongs here and
that its male is J. fimbriator, Thunb.

10"

148 MR. CLAUDE MORLEY ON AFRICAN

1. CARINIFER, Sp. Nn.

A stout and closely punctate but not very dull, black female
with no paleantennal band; frontal and centre of external orbits,
labrum, mandibles and cheeks, basal half of flagellum indefinitely,
and the anterior legs except basally, fulvidous ; elongate callosities
before and beneath radix, a circular central mesonotal mark, whole
scutellum and postscutellum, alone stramineous. Head trans-
verse; face and clypeus strongly and evenly punctate, not
disereted, the latter slightly produced centrally. Flagellum
filiform, neither explanate nor apically attenuate. Notauli short
and deep; metathoracic carinee very strong, with the external
dentiparal outwardly curved at its base; areola subaciculate,
basally glabrous, rounded and there touching postscutellum,
apically subtruncate ; spiracles large, apophyses wanting. Scu-
tellum deplanate, glabrous with a few fine scattered punctures,
not laterally margined. Abdomen stout and fusiform, immaculate
black ; basal segment stoutly bicarinate to the closely and evenly
punctate postpetiole; gastrocceli large and broader than the shortly
striate intervening space; anus nitidulous and obsoletely punctate;
the third and obscurely ochraceous second ventral segments
plicate, hypopygium remote from terebral base. Legs stout and
not short, black with anterior femora, tibize and the front tarsi
clear red; coxe not scopulate. Wings hyaline; areolet pen-
tagonal, broad above ; internal cubitus straight, with no nervelet ;
stigma castaneous-black ; lower basal nervure distinctly post-
fureal. Length, 17 mm. @ only.—This species has much the
facies of Hepiopelmas leucostigmus, Grav., with the anus more
acute and hypopygium remote.—Harrar in Abyssinia during

May 1911 (Collector ignot.).

2, MELANOPTERUS, Sp. n.

An extremely dull and closely shagreened, black species with
only the laterally carinate scutellum nitidulous; flagellar band
alone white; wings strongly infumate. Head transverse, con-
stricted behind the eyes; frons evenly, face and the undiscreted
clypeus closely punctate, with apex of the latter a little rounded ;
labrum and palpi obscurely rufescent. Flagellum centrally sub-
explanate and apically attenuate. Notauli short and deeply
impressed; metanotum with only the petiolar carine strong ;
areola and costule obsolete, the former shaped as in the last
species, though sculptured as remainder of metanotum ; spiracles
large, apophyses short and acute. Abdomen fusiform and im-
maculate; postpetiole punctate-shagreened ; gastrocceli wanting,
thyridii shining, intervening space not striate; venter plicate
throughout. Legs normal; coxe not scopulate ; front tibie and
tarsi obscurely rufescent. Wings nigrescent throughout; areolet
coalescent above ; cubital nervure curved, with no nervelet ; basal
nervure continuous through the median. Length, 14 mm,
2 only. —The outline is similar to that of J/. ’ewcomelas Gmel.,

ICHNEUMONIN# IN THE BRITISH MUSEUM. 149

but the head is posteriorly shorter.—The type was captured
on Mount Kokanjero to the 8.W. of Elgon at 6400 feet on
8th August, 1911, in Uganda.

3. GLAUCOPTERUS, Sp. Nn.

A somewhat dull and evenly punctate-shagreened, black male
with the laterally carinate scutellum more shining ; front tibiz
alone internally whitish; wings flavescent. This might well be
the alternate sex of the last species, with which the sculpture and
colour mainly accord, were it not that the mandibles are rufescent,
the metanotal areola and costule are strong, the apophyses
wanting, the abdomen subcyaneous with gastrocceli deeply im-
pressed though narrower than the substriate intervening space,
and the wings deeply clear flavescent with only their apices
infumate, the radix and costa, stigma and nervures clear flavous,
and the lower basal nervure distinctly postfureal. Length, 13 mm.
3 only.—Very similar, superficially, to the males of my Pimpla
glaucoptera (Revis. Ichn. iii. 1914, p. 68) from Uganda and
British EK. Afvica.—Taken on a plateau of Mount Mlanji at about
6500 feet on Ist May, 1910, in Nyasaland.

LEPTOTHECUS.

Head with the clypeus apically truncate and labrum prominent ;
cheeks usually strongly buccate. Antenne broadly pale-banded and
not considerably dilated beyond their centre. Thorax unusually
cylindrical ; metathorax large and distinctly longer than broad,
apically somewhat straight and abruptly declivous; areola
elongate-hexagonal, fully twice longer than broad, apically trun-
cate and often basally incomplete ; apophyses spinate. Scutellum
deplanate. Abdomen slender, longer than head and thorax,
apically narrowed from base of fourth segment, with seven visible
dorsal segments ; petiole slender and elongate, becoming gradually
a little explanate towards its apex; pygidium large and two-
thirds the length of the penultimate; ventral plica obsolete ;
terebral valvule strongly prominent, remote from hypopygium
and as long as the white-marked two apical segments. Legs with
tarsi spinose ; hind legs much the longest, with their tibiz basally
constricted. Wings with areolet pentagonal and constricted
above; nervelet distinct.

In his erection of this genus, Cameron (Entomologist, xxxvi.
1903, p. 240) correctly places it in the Oxypygini, and an exami-
nation of the genotype enables me to emend it slightly, as above
given. Its author considered that it ‘‘may be known by the
elongated spined median segment, with its elongated coftin-shaped
areola, confluent with the lateral aree at the base; by the long
projecting ovipositor ; and by the smooth impunctate abdomen,
with its small gastroceeli.” It has the facies of Stenichnewmon
Thoms., but the face is not apically constricted, the cheeks are
buccate in the typical species and the juxta-coxal is entirely

150 MR. CLAUDE MORLEY ON AFRICAN

disereted from the pleural area ; the dull and shagreened thorax,
of both the known kinds (lib. cit. p. 260 et Zeits. Hym. u. Dipt. v.
1905, p. 142) from northern India, is similar to that of S. ochropis
Gmel. Though not agreeing in every particular with the
diagnosis of Leptothecus, which was based on a single Oriental
female, the following kinds are sufficiently homogeneous and
differ from Flyaldenenims so little (besides the er of basal
metanotal suleus) that I should expect to find it mixed among
Szepligeti’s conception of that genus.

1. ALUTACEFER, Sp. 0.

A very dull and alutaceous, black species with sparse pure
white, and the thorax with red, markings. Head short and not
extending posteriorly behind the eyes; ocelli at vertex of the
abrupt occipital declivity ; face closely punctate, its centre and
the continuous clypeus more coarsely punctate, centre of the
latter slightly emarginate with its glabrous lateral angles white
and subelevated ; palpi, except their apical joint, and the exserted
labrum white; frons transversely pure white throughout.
Antenne attenuate, and a little explanate, beyond their central
partial white band. Thorax deplanate and rosy-red with pro-
thorax, sides of mesonotum and dise of metanotum black; notauli
very small, speculum not shining; areola double as long as
broad, basally acuminate but not téaching base of metathorax,
with weak costulee ; petiolar area short, basal area tr langular and
hardly carinate ; spiracles elongate, apophyses strong and acute.
Scutellum red, deplanate, somewhat elongate and laterally strongly
carinate to the flavidous apex. Abdomen with apices of the two
basal segments, whole of the sixth and seventh, and of the second
to fourth ventral ones, white; terebra black and distinctly
exserted, with its extreme base covered by hypopygium ; first
segment evenly punctate and somewhat narrow, thyridii of the
second lateral and longitudinally linear. Legs elongate and
slender; anterior brick-red with coxe and trochanters white,
their tarsi and the imtermediate femora nigrescent; hind legs
black, with the second to fifth tarsal joints pure white. Wings
hyaline and not broad; tegule and costa black, stigma ferru-
ginous; areolet as broad as high, laterally coalescent above ;
discoidal cell narrow and subparallel, with obsolete nervelet ;
basal nervure subcontinuous.— ¢ differs in having part of the
facial and external orbits, whole clypeus but not centre of the
frons, white; the flagellum is setigerous and more elongate ;
the apical half of the metanotum ee nd-red, with white genital

valvule. Length, 11-13 mm. 6 9.—Var. ¢. Head _ black
with the palpi "and frontal orbits alone white. -Amother 3 has
the legs basally nigrescent-1ed.— Males were captured in the Tero
Forest of Uganda during July 1912 (C. C. Gowdey). Females at
Unyoro about 3400 feet in the Budongo Forest of Uganda in
mid-December 1911. Two additional females, also from Uganda,
have the centre of scutellum and a dot on the postscutellum

ICHNEUMONIN® IN THE BRITISH MUSEUM. 151

white, the metathorax entirely dull rosy and its arez obsolete ;
they occurred at Entebbe during September 1911 and at some
3500 feet in the Mabira Forest, Chagwe, in the middle of the
preceding July; and Jones took a d at Mfongosi in Zululand,
January, 5. African Museum.

2. PUNCTIFER, Sp. Nn.

A very dull and closely punctate, black male with sparse white
marks, and the thorax mainly red. Extremely similar to the
last species but with the mesonotum punctate, in place of
alutaceous; the head immaculate ; cheeks narrow, clypeus
extremely small; palpi and the entire scape alone white ;
flagellum longer and not setigerous; notauli deeply impressed
and elongate; areola small, apophyses obsolete; petiole lineav ;
genital valvule, anterior legs basally and the hind tarsi, black ;
nervelet wanting. Length, 13 mm. <d only.—Found by
C. C. Gowdey in the Tero Forest during July 1912, in Uganda.

3. MESONOTIFER, Sp. n.

A very dull and alutaceous, black species with sparse pure
white, and the thorax with red, markings. Head by no means
short, though obliquely constricted, behind eyes; cheeks broad ;
face deplanate, definitely and somewhat confluently punctate, not
discreted from the more sparsely punctate and apically truncate
clypeus; mandibles slender and black ; palpi, clypeus, and the
glabrous frons pure white. Antenne elongate and subexplanate
beyond the mainly white central band, apically attenuate, with
the eight basal flagellar joints elongate. Thorax rosy-red with
prothorax, mesonotum laterally, frenum and disc of metanotum
basally from the small but acute apophyses, black ; mesonotum
apically produced at its centre, notauli short ; metanotum closely
punctate with areola elongate-triangular, apically attenuate and
not extending to base, emitting weak but entire costule from
near its base; spiracles elongate. Scutellum convex, shining
and glabrous with a few sparse punctures, its sides carinate to
near apex whici is, like the postscutellum, flavidous. Abdomen
elongate with apices of two basal segments and base of the third
narrowly, with whole venter, testaceous; seventh and apex of
sixth segments white ; thyridii of the second small and laterally
linear; terebra black and shortly exserted. Legs slender and
strongly elongate, nigrescent with apex of hind metatarsus to
middle of its onychium white, hind trochanters beneath and
apices of their cox pale testaceous; anterior legs internally
rufescent or testaceous; claws large and simple. Wings hyaline
and not broad; tegule and costa black, stigma ferruginous ;
areolet no higher than broad, laterally coalescent above, emitting
recurrent nervure before its centre; lower basal nervure post-
fureal.— g differs in its white mandibles, immaculate black
clypeus and frons, more slender flagellum; deep and crenulate

152 MR, CLAUDE MORLEY ON AFRICAN

notauli, extending to scutellar fovea; sublinear and strongly
white-pilose basal segment ; large and black genital valvule; and
in having the hind tarsi white, with only the metatarsal base
and very small claws black. Length, 15-17 mm. ¢ 9.—The
typical female was taken at Masaka in Uganda early in
November 1913 (C. C. Gowdey). Cotypes are from the Tero
Forest of Uganda in July 1912, along with a couple of males ;
another male at Mwera, near Entebbe, on 26th of the next month
(Gowdey). In the middle of August 1911, on the Siroko River
at the west foot of Mount Elgon at an altitude of 3600 feet, also
in Uganda.
STENICHNEUMON.

Van Burgst has recorded three European species of this
Thomsonian genus from northern Africa, and Cameron has
attributed to it, with a query, another species from the south of
the Continent (Records Albany Mus. 1. 1905, p. 229). It would
appear uncommon here, and I have seen but one species :—

1. OCHRACEATOR, sp. n.

An extremely dull, pale ochraceous male with indefinite black
markings, and the hind legs dark-lined with white tarsi. Head
strongly transverse and abruptly declivous immediately behind
eyes; occiput and centre of frons nigrescent ; face superficially
and clypeus very sparsely punctate, the latter not discreted and
apically truncate. Flagellum black, with joints 14 to 24 white;
relique absent. Thorax dull and alutaceous, higher than broad ;
metanotum and mesonotum with basal marks, scutellar carine
and the postscutellum, indefinitely nigrescent ; areola obsolete,
obliquely and irregularly striate, subhexagonal, emitting obsolete
costule; petiolar area shagreened, with no carine; spiracles
elongate, apophyses absent. Scutellum deplanate and punctate,
apically truncate, with an elevated carina throughout its margin.
Abdomen indefinitely black with apices of the segments narrowly,
valvule and venter, pale. Legs elongate; hind ones with base
of tarsi, inner side of tibia and a discal line on femora, sub-
nigreseent ; remainder of hind tarsi white. Wings hyaline;
tegule and radix pale, stigma and nervures black; areolet small
and pentagonal, emitting the strongly sinuate recurrent before its
centre; nervelet wanting; basal nervure continuous through
the median. Length, 1] mm. ¢ only.—Captured in Lagos,

Western Nigeria, on 13th February, 1911, by W. A. Lamborn.

CRATICHNEUMON.

No species of this Thomsonian genus have yet been reported
from Africa, apart from such as may be included in the main
genus Ichnewnon by Tosquinet, &e. I consider the following is
sufticiently typical to be placed here, though several of its features
are abnormal.

————eeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEEE—eeEeEeEeEeEeEeEEEy

ICHNEUMONIN# IN THE BRITISH MUSEUM. 153

1, TESTACECOLOR, sp. n.

A stout and dull, clear testaceous female with only the sternum,
frons and frontal orbits substramineous ; flagellum alone black,
white-banded ; wings flavescent. At once known in the present
genus by its unique coloration; the aciculate-glabrous, and
laterally punctate, postpetiole ; the small, subquadrate and nearly
glabrous metanotal areola, with or without distinct central
costule ; by the pale stigma; and by having all the simple coxe
curiously deplanate beneath. Length, 13-14 mm. @ only.—
The alutaceous thoracic sculpture relates this species to (. rubri-
cosus Hlmer., but the head is very broad and distinctly broader
than thorax.—Found at Unyoro in the Budongo Forest in the
middle of December at 3400 feet; and at Chagwe in the
Mabira Forest in the middle of July at 3500 feet, in Uganda,
during 1911.

BARICHNEUMON.

Berthoumieu has described three species from Algeria; Burgst
has discovered that three of the palearctic kinds extend to ‘Tunis;
and I have brought forward (Ann. 8. Afr. Mus. xv. 1916, p. 364)
another from the Cape. This Thomsonian genus will probably
be found here over at least the more temperate regions.

1. CONCINNATOR, sp. n.

A small and evenly punctate, black male with vertical dots and
base of scape beneath obscurely, and all the tibie and femora
clear, fulvous; wings subinfumate throughout. Head not
strongly constricted posteriorly; frons and face closely, the
apically a little rounded clypeus very sparsely, punctate. Fla-
gellum short and setaceous, hardly longer than head and thorax,
apically pale beneath. Thorax stout, cylindrical, shining and
evenly punctate ; notauli indicated; metanotal areola triangular,
its basal carina convex and apical concave, emitting strong
costule a little beyond its centre; petiolar area elongate, fee
creted with very strong lateral carine; spiracles small and
elongate, apophyses wanting. Scutellum punctate and shining,
only basally carinate. Abdomen cylindrical and dull; basal
segment stout, gradually explanate, with the postpetiole sub-
bicarinate and evenly punctate ; gastroceeli small and superficial,
intervening space not striate; ventral segments two to four
plicate, valvule black and not exserted. Legs neither short nor
slender; all the femora and tibie fulvous, with the front tibie
internally paler. Wings somewhat small, evenly brunneous
throughout ; stigma black; areolet obliquely pentagonal, broad
above, emitting the sinuate recurrent nervure beyond its centre ;
lower basal postfurecal. Length, 8mm. dg only.—The sculpture
and disposition of the wing nervures are very like B. gemellus
Grav., though the whole puncturation is less close and more
distinct.—Two or three males were found in Abyssinia, probably
about Harrar, in 1910 (Collector ignot.).

154 MR. CLAUDE MORLEY ON AFRICAN

2, iNcuUBITOR Linn.

A female of this species, differing from the somewhat un-
common palearctic form with entirely black legs only in its lack
of. white flagellar and anal markings, was discovered on the
western slopes of Mount Kenya, on the Meru-Nyeri Road at
between six and eight and a half thousand feet, in the middle of
February 1911, in British Hast Africa.

3. PLANINOTUM, sp. n.

A closely punctate, black female with flavous markings and the
abdomen mainly red. Head constricted and narrow behind eyes;
mandibles, palpi, labrum, lateral angles of the apically truncate
clypeus, and frontal orbits nearly to the concolorous vertical dot,
pale stramineous; frons sparsely and finely punctate; face evenly
punctate, centrally elevated and apically discreted. Antenne
centrally white-banded and apically attenuate. Thorax very
closely punctate, dull with pronotum, callosity below and an
elongate one before radix, flavous; notauli fine; metathorax
shagreened, with the broad and undiscreted petiolar area apically
transaciculate ; areola quadrate, parallel-sided, deplanate and
extending to base of metanotum, with weak lateral carine and
no costule; spiracles linear, apophyses obsolete. Scutellum
nitidulous, very finely punctate, not margined, clear flavous with
its basal third and the postseutellum fulvous. Abdomen elongate
and parallel-sided, with anus from base of sixth segment more or
less definitely black; postpetiole deplanate, finely and irregularly
punctate; gastroceeli transverse and deeply impressed, their
intervening space much narrower than central area of post-
petiole; terebra black and a little exserted. Legs normal, with
hind tarsi slender; indefinitely nigrescent red, with hind coxe
and femora darker; hind coxe finely punctate beneath, with
very large scopule. Wings hyaline; radix and tegule fulvous,
stigma red; areolet pentagonal, as broad as high and somewhat
small. Length, 9 mm. @ only.—From all species known to
me of this genus, to which the pale vertical dots and punctate
petiole show that it belongs, it differs in its nearly smooth meta-
notum with the parallel-sided areola extending to base; the
central abdominal segments, also, are more deplanate than is
usually the case.—A couple of females were received from Algeria
about 1849, I believe from Francis Walker.

4, SEXALBATUS Wesm.
A single typical male of this palearctic species was captured in
Abyssinia, probably about Harrar, in 1910.

5, FOSSIFER, Sp. 0.
A closely punctate, black male with profuse stramineous-white
markings, and the abdomen and legs mainly red. Head trans-

a ae

ICGHNEUMONIN IN THE BRITISH MUSEUM. 155

verse, Short and not strongly constricted posteriorly ; stramineous
with mandibular apices, 2 dot at the genal orbits, whole occiput
with ocellar region and centre of frons to serobes, black; frons
subglabrous; face and clypeus distinctly punctate, with the centre
of latter verry conspicuously impressed longitudinally at its apex.
Antenne normal, immaculate. Thorax black with prothoracic
margin, prosternum, linear callosities before and beneath radices,
a broad mesopleural streak, two discal mesonotal vittee and apex
of metathorax on either side, white; notauli short and deeply
impressed ; areola cordiform, subglabrous, a little longer than
broad, not extending to base, and emitting strong costule ;
spiracles linear, apophyses wanting. Scutellum convex, margined
to its centre ; stramineous, with base of its carine and two dots
on postscutellum concolorous. Abdomen red with the basal
segment alone black; postpetiole convex, evenly and closely
punctate, with no carine ; gastroccell deeply impressed, large and
broader than the striate inter vening space; valvule fulvous,
Legs normal; dull red ; tarsi, hind ie and most of their coxee
black; anterior and dise of hind coxe whitish; front tibie
inter nally pale. Wings normal; tegule and stigma black, radix
whitish; areolet shghtly higher chan broad, coalescent above.
Length, 11 mm. go only. At once known by the definite
thoracic white markings and by the clypeal fossa.—A couple of
males from Uganda: one at Chagwe in the Mabira Forest at
some 3500 feet in the middle of July 19115; and the other
between the Seziwa River and Kampala at the same altitude
towards the end of the following month.

6. muNDATUS Tosq.

This is probably not an uncommon species in Central East
Africa, since I have seen several males, but nothing that I can
assign as its female. The peculiarly elongate form and dull red
coloration are distinctive; and the punctate postpetiole assigns
it to the present genus, though Tosquinet (Mém. Soc. Entom.
Belg. v. 1896, p. 31) described the male under /cehnewmon, in its
Wesmaelian sense. The extent of black markings is variable :
the thorax is usually black with a flavous pronotal callosity, and
the mesonotum, excepting a discal vitta, red; sometimes the
flavous 1s become red; at others the black Benita mesonotal line
is lacking, or the spiracular and dentiparal arez are also red,
with or without the propleure. Also all the coxe and trochanters,
or as in the type only the anterior coxee above, are black.—The
specimens from which these details are drawn come from Higo
Samula in Abyssinia on 30th Pee 1911 (#. J. Stordy);
Abyssinia, probably about Harrar, in 1910 (Collector ignot.) ;
and one, which considerably extends its eae range, from the
valley of the Rukuru River between 20th and 27th utie. 1910,
at 3000 feet, in Nyasaland. Another male, differing solely in its
black legs with only the anterior tarsi red and inner side of front
tibiz flavous, was found on the west slopes of Mount Kenya, on

156 MR. CLAUDE MORLEY ON AFRICAN

the Meru-Nyeri Road at some 7000 feet, about 20th February,
ONE

IcGHNEUMON.

This genus is nowadays used in the sense to which it was
sgeunichal by Professor Thomson of Lund (Opuse. Entom., 1893,
p- 1911), though the difficult task of assigning to their correct
subgenera she. very numerous species, that: were therein included
in the earlier times and have been therein placed by careless
authors during the recent ones, is by no means yet completed ;
and this is as true of Africa as elsewhere, being especially
applicable to Tosquinet’s descriptions of 1896. In fact, of the
fifty kinds still placed here from Africa, such things as T. intra-
torius of Fabricius (17938), 7. apicalis of Wiedeman (1824), I
desjardinit of Brullé (1846), and JZ. frontalis of Guérin (1846 ;
nec Fourcrey in 1785) are unrecognizable without reference to
the scattered and probably lost types! Hence some synonymy
is sure to arise. My own experience goes to show that this
genus 1s but poorly represented in the Ethiopian fauna, and that
such forms as oceur have usually extended from the somewhat
broad distribution of palearctic kinds throughout the southern
Mediterranean shores.

1. RUBRORNATUS Cam.

The male only of this conspicuous insect was brought forward
by Cameron (Records of the Albany Museum, i. 1904, p. 141)
from the Cape, whence I have not received it; nor, cacionely
enough, from any central part of the Continent. But about
Harrar in Abyssinia it must be of comparatively frequent occur-
rence, for fully half a dozen were comprised in a small collection
there made in 1910; and these males show considerable constancy
of markings in respect to the flavous and black, though the
peculiarly characteristic brick-red of the basal moiety of the
second segment may be half obscured by black as in the southern
type, black only narrowly at the sides or so broadly suffused with
that colour that the rufescence is traceable only between the
deeply impressed gastrocceli. This male is obviously allied to
the abundant palearctic J. sarcitoriws Linn., which extends to
Algeria; and, judging by analogy, I here assign to it the
following female :—

A very stout and closely punctate, dark crimson female, with
ovate abdomen ; central flagellar band, whole scutellum, apices
of second and third, and dise of the sixth segments, pale flavous ;
remainder of abdomen from base of third segment, small apical
marks before its pale band on dise and sides or else the whole
centre of second segment, flagellar apices, sometimes disc of hind
coxee, and the whole thorax except mesonotum or also except
metanotuin indefinitely, black. Mesonotum with a very faintly
nigrescent longitudinal central vitta; stigma fulvous. Exactly
resembling J. sarcitorius in both structure and sculpture; identical

.
|
:

—

ICHNEUMONIN& IN THE BRITISH MUSEUM. 157

in those of the metanotum, postpetiole, coxal scopule and basal
nervure; though a little stouter. Length, ¢ 2, 11-12 mm.—-
Of this distinct female I have seen two examples, differing only
in the extent of rufescent coloration on the metanotum and
second segment, which were respectively captured so far apart as
Alaba in Abyssinia on 15th November, 1911, by R. J. Stordy,
and Durban in Natal during 1904, by J. P. Cregoe.

CHASMIAS.

The right of this genus to distinction from Jchnewmon has
constantly been a moot point since its erection under the name
Chasmodes by Wesmael in 1844 (nec Cuvier, 1836); it was re-
named by Ashmead in 1900. We must, I think, look to the
Kthiopian fauna to supply us with a good basis of generic dis-
tinction, since less than half a dozen species have yet been
described altogether: one of these is from Abyssinia and was
discovered by Tosquinet.

1. GLAUCOPTERUS Morl.

This was recently brought forward (Ann.§8. African Mus. xv.
1916, p. 865) by me from the Cape; but a second female, differing
in nothing but its black mesonotum, anus and hind tibia, was
captured in Abyssinia, probably about Harrar, during 1910,

2. RUFICAUDATOR, Sp. 0.

A very strongly elongate and somewhat dull species, with the
head except mouth and top of face, base of antennze beneath,
anus from apex of fourth segment and anterior tibie, brick-red ;
centre of flagelluin very broadly but indefinitely flavous; seutellum
alone stramineous. Head posteriorly as broad as the eyes, with
cheeks and temples buccate; frons, the centrally elevated face,
and the clypeus evenly and distinctly punctate, apex of the last
glabrous and sinuate with its centre conspicuously produced.
Thorax cylindrical, with superficial notauli ; metathorax closely
and evenly punctate with fine carine; areola subhexagonal,
apically a little constricted, extending to base and emitting
costule from its centre; spiracles elongate, apophyses wanting.
Scutellum subconvex, glabrous, with some punctures. Abdomen
linear, narrower than thorax and just double length of head and
thorax, dull and very closely punctate, with the postpetiole
evenly punctate on either side of its glabrous and nitidulous
centre; second segment fully twice as long as broad, with the
deeply impressed gastrocceli narrower than their intervening
space; fourth and fifth segments discally quadrate; valvule red
and hardly exserted. Legs not long, tarsi slender; front femora
internally and their tarsi also rufescent. Wings subhyaline, with
stigma black; areolet as broad as high, not laterally coalescent
above; nervelet elongate but evanescent. Length, 18 mm.

158 MR. CLAUDE MORLEY ON AFRICAN

3 only.—The type occurred by the Yala River on the southern
edge of the Kakumga Forest at between 4800 and 5300 feet
towards the end of May 1911, in British Hast Africa.

Subtribe Amblypygini.
AMBLY'TELES.

This genus, as established by Wesmael (Nouv. Mém. Acad,
Bruxelles, 1844, p. 111) has, like his Zehnewmon, been subdivided
by Thomson into three genera, of which the typical one is well
represented by thirty-seven African species, all from the north
or south of the Continent except three ; Tosquinet had one from
Sierra Leone ; Szépligeti described one (Bull. Mus. Paris, 1907,
p. 187) from East Africa; and another, captured by the Kil-
manjaro Expedition, in 1910, for the synonymy of which ¢f. Roman
(Entom. Tidskr. xxxi. 1910, p. 144). Setanta Cameron (Ann.
Nat. Hist. vii. 1901, p. 483) is already known to be identical
with the present genus.

1, SPILOPTERUS, sp. 0.

A stout, ferruginous species with the metathorax, mesopleure
and most of abdomen, black; apex of wings broadly nigrescent.
Head posteriorly buccate and broader than the eyes; mandibles
stout, punctate, and apically black; cheeks buccate; face and
clypeus rugulosely punctate, obsoletely discreted, with the latter
laterally elevated and apically truncate; vertex trans-striate and
not broad. Antenne attenuate and hardly darker apically, of 2
slightly explanate beyond their centre. Thorax dull and coarsely
punctate ; frenum black; notauli apical and small; pleure finely
punctate, with the speculum dull; metanotum evenly granulose
throughout, with no carine, its basal suleus somewhat deeply
impressed, spiracles elongate, and apophyses wanting. Scutellum
convex, closely and deeply punctate, lateraily strongly carimate
to near apex. Abdomen elongate-fusiform and finely punctate
with the fourth and foliowing, usually all the dorsal, segments
black; postpetiole and anus from base of third segment
shagreened, or the former in ¢ shining and smoother ; gastro-
celi very small; hypopygium extending to apex, terebra very
slightly exserted. Legs clear red with the posterior cox and
centre of their femora more or less broadly black ; tarsi and hind
tibie stout. Wings flavescent with whole apices of the front
ones abruptly nigrescent from the recurrent nervure and areolet,
which is subcoalescent above; tegule and the basally paler
stigma fulvous. Length, 12-14mm. ¢ 9.—Found in scattered
forest on the Nandi Plateau at about 6000 feet during early June
1911; and on the southern foot and slopes of Mount Elgon at,
5100-5800 feet between the 8th and 13th of the same month, in
British East Africa.

ICHNEUMONINA IN THE BRITISH MUSEUM. 159

2. AURICOMUS, Sp. n.

3. A handsome, aureo-flavous species with profuse black
markings and evenly nigrescent wings. Head short, hardly
narrower than the eyes, with the posterior margin subtruncate ;
face finely, with its centre and clypeus much more coarsely,
punctate; mandibles narrow and punctate, labrum exserted,
clypeus centrally a little produced; frons longitudinally in the
centre and back of head, below the occipital margin, black.
Antenne hardly longer than half the body, stout and attenuate
throughout, immaculate orange with the joints short. Thorax
closely and finely punctate, with all the pleure more sparsely
punctate and shining; whole frenum, basal scutellar fovea
broadly, propleure both above and below the collar, mesopleurz
and -sternum except the radical callosities, an apical line and
small mark above cox, metathoracic base and petiolar area,
black ; mesonotum indefinitely nigrescent and discally pubescent,
notauli apical ; metanotal carine entire; areola square, glabrous,
glittering and spinately produced basally in the centre ; spiracles
elongate and apophyses wanting. Scutellum a little convex,
shining, finely and not closely punctate. Abdomen dull and very
closely punctate with golden pilosity, stronger on the fifth and
following segments ; second to fifth segments except their apical
margin regularly and sides irregularly, and whole venter, black ;
first segment punctate and, except apically, indefinitely rufescent ;
gastroceeli shining, narrower than the simple intervening space ;
second and third ventral segments plicate, the last not centrally
produced. Legs somewhat stont and not elongate, with only the
lower side of hind coxee and trochanters partly black; claws
simple. Wings evenly infumate throughout; tegule and radix
flavous, stigma and costa black; areolet broad above and a little
produced apically below ; radius apically reflexed.—The ¢ differs
very slightly in having the sternum, pleure, and metathorax
immaculate black, as also are the posterior coxe; but the frons
and occiput are pale; coxe not scopulate. Length, 14 mm.
3d 9.—The typical male was found at Nairobi during June or
July, 1912, by Dr. A. D. Milne; the only female I have seen is
labelled with the MS. and most inappropriate name “/ulgidipennis,
Cam., Type. Uganda,” and lacks abdomen. .

3. FULYOCAUDATUS Tosq.

This male has not been mentioned since first described from
Hadda Galla‘in Abyssinia by Tosquinet (Mém. Soc. Entom. Belg.
v. 1896, p. 76). It is an extremely conspicuous species in its
black wings and body with orange antennze, red head and anus,
and fulvous legs. he present example of but 11 mm. is smaller
than the type and has the areola as long as broad, distinctly
hexagonal, and not ‘a bords arrondis”: it is from Peter
Cameron’s collection, simply labelled “ Erythria.”

160 MR. CLAUDE MORLEY ON AFRICAN

4, necarorius Fab.

This is a very variable species, first described by Fabricius
(Entom. Syst. ii. 1798, p. 141), and the subtropical specimens I
have seen differ somewhat from the common Kuropean form,
which has not. been recorded from Africa. The female type of
Eristicus iridipennis Cameron (Records of the Albany Museum,
i. 1904, p. 142), from the Cape, exactly agrees with the type,
which I have examined in the Calcutta Museum, of Pompilus
divisus Smith (2nd Yarkand Mission, Hym. 1878, p. 14; Water-
house, Aid to Ident. Ins. ii. 1885, pl. 169. fig. 2, 9), from
Kashmir, and I have seen it also from Nepal. These I have no
hesitation in synonymising with A. negatorius var. nubilus
Berthoumieu (Ann. Soc. Entom. France, 1895, p. 640); Smith’s
name has priority for the varietal form.

5. MACULICAUDIS Cam.

The male type of Pseudamblyteles maculicaudis Cameron (Ann.
Ss. Afr. Mus. v. 1906, p. 164) could be placed with equal propriety
in Ichnewmon, s.st7., or the present genus, from which Psewdam-
blyteles Ashmead, differs solely in the instable character of lacking
apophyses. It is not a good genus and must be abandoned. The
type, recorded from Cape Colony, is labelled ‘* Kimberley.”

6. TESTACEATOR, Sp. n.

A clear testaceous and closely punctate species with only the
central -flagellar band and all the orbits white, base of flagellum
red, its apex and the tarsal pulvilli alone black. Head posteriorly
as broad as eyes, with both the temples and cheeks buccate; frons
and face closely and evenly punctate; clypeus glabrous with a
few large punctures, not discreted but foveate on either side
before its lateral margin, apically a little produced obtusely on
either side of centre; mandibles bidentate, subparallel-sided,
apically rufescent. Antenne filiform and not apically attenuate,
with basal flagellar joints elongate. Thorax stout, dull and very
closely punctate ; notauli obsolete, speculum convex and shining;
metanotal carine fine and distinct; areola hexagonal, hardly
longer than broad, not quite extending to base and emitting
costule before its centre; petiolar area discreted, centrally
pavallel-sided and there transaciculate ; spiracles linear and
apophyses obsolete, obtuse. Scutellum deplanate, glabrous, and
nitidulous with half a dozen deep punctures, laterally carinate
to its centre. Abdomen sublinear and immaculate, with anus
nitidulous; basal seement slender, centrally punctate, postpetiole
very finely aciculate throughout with no carine; gastrocceli
superficial and striate, their intervening space aciculate; venter
convex throughout, terebra black and hardly exserted. Legs
normal, immaculate; hind cox simple. Wings silaceous-hyaline;
radix and tegule testaceous, costa and nervures black, stigma
luteous; areolet pentagonal and somewhat small, laterally

ICHNEUMONINA IN THE BRITISH MUSEUM. 161

constricted but not coalescent above; radius not apically curved,
nervelet obsolete, basal nervure continuous through the median.
Length, 12 mm. @ only.—The abdominal structure is similar
to that of A. subsericans Grav. Found at Salisbury in Mashona-
land by G. A. K. Marshall during March 1900.

SPILICHNEUMON.

No African species have yet been assigned to this genus, which
was divided from Amblyteles by Prof. Thomson (Opuse. Entom.
xix. 1894, p. 2087); and it has been but little employed by
systematists. The range, however, is much broader than has
hitherto been suspected ; and I find that Cameron’s two genera,
Haliphera (Kutomologist, 1903, p. 237) and Hutanyacra (Trans.
Ent. Soc. Lond, 1903, p. 227) represent respectively the females
and males thereof. Further, Cameron mistook Barichnewmon,
among the Oxypygini, for the present genus.

1. DIDYMATUS, sp. nN.

An elongate black species with profuse whitish markings; the
legs and abdomen, except its base, red with the segmental angles
whitish. Head transverse and obsolete behind the eyes; outer
orbits broadly, inner to vertex, whole face, and mouth-parts
except apices of the slender mandibles, flavescent white; clypeus
broad, apically truncate, and not basally disereted from the evenly
punctate face; occiput shining, pilose and finely punctate.
Antenne half length of body, attenuate, with the joints short
and subserrate; scape white beneath. Thorax shining and
finely punctate, discally deplanate; notauli elongate but super-
ficial; pronotal margin, prosternum, broad callosities before
radices and a line below them, a mesosternal spot and a transverse
mark below hind radices, a discal mesonotal spot and the whole
dentiparal ares, white; mesopleure and a mark above the meta-
pleure red; upper metanotal carinw strong, areola glabrous and
subquadrate with its base centrally elevated and extending to
postscutellum; spiracles elongate, apophyses wanting. Scutellum
and the glabrous postscutellum flavous; former deplanate,
glabrous, sparsely and evenly }.unctate, not margined. Abdomen
deplanate, subparallel-sided, somewhat dark red with sides of
the four basal segments apically, and apices of the basal or two
basal narrowly, whitish; first segment black with the post-
petiole evenly and somewhat diffusely punctate; gastrocceli
narrower than the intervening space; hypopygium spinately
produced ; terebra concealed. Legs somewhat slender; anterior
coxe, and marks above and below the black hind ones, whitish ;
hind tarsi infuscate. Wings hyaline; tegule white, costa black,
stigma ferruginous; areolet higher than broad, not coalescent
above.—The ¢ has the antenne longer and more strongly
serrate; basal segment red before its pale apex; mesopleure,
metapleural mark, and apical margins of the four basal segments,

Proc. Zoot. Soc.—1919, No. XJ, 11

162 MR. CLAUDE MORLEY ON AFRICAN

flavescent. Length, 13-15 mm. 6 2.—The type was captured
at Mlanje on 4th October, 1913, at 2300 feet, in Nyasaland; and
the androtype between Jinja and Busia in some forest land east
of Busoga on 28th July, 1911, at 4000 feet, in Uganda.

2. UNIPUNCTOR, Sp. n.

An elongate black male with profuse whitish markings; the
legs and abdomen. except anus, red with segmental apices whitish.
Extremely like S. didymatus but with the mandibles and palpi
black, the frontal and external orbits immaculate; basal two-
thirds of flagellum red, pleurze and mesonotum immaculate ;
areola finely sculptured and apically stramineous; scutellum
convex ; basal segment red, the fifth and sixth black, with only
their apex whitish, and the fourth basally nigrescent; tegulee,
coxe and trochanters entirely black; areolet broader and very
broad above. Length, 13 mm. 6 only.—The type was captured
at Harrar in Abyssinia, 1912.

3. TRIANGULATOR, Sp. ND.

A black male with profuse whitish markings; the legs and
abdomen red, with segmental apices whitish. Very similar to
the last species and differing from S. didymatus in the convex
scutellum, sculptured metanotal areola, black external orbits and
mouth-parts, and in the rufescent flagellum. It is distinct in
having the areola remote from metanotal base, and its apex
flavous, which colour extends thence obliquely to the obsolete
apophyses; the abdomen is not black-marked, and the stigma is
black. Length,9 mm. ¢ only.—The type is from Queenstown in
Cape Colony, where it was captured during 1907 by EK. 'T. Wells.

XENOJOPPA.

The discovery of this genus in Africa is as interesting as that
of Lagenesta and other genera that have hitherto been regarded
as purely Oriental ; it is even more so, on account of its peculiar
specialization, for it is the only one known to me among the
Ichneumonides that shares coxal dentition with the Phzogenides.
Xenojoppa was published by Cameron in 1902 (Entomologist,
xxxv. p. 179) upon finding that his original name for the genus,
Magrettia (Ann, Nat. Hist. vii. 1901, p. 480) had already been
employed by Brunner v. Wattenwyl in the latter’s ‘Monographie
der Stenopelmatiden und Gryllacriden’ (Verh. z.-b. Ges. Wien,
Xxxvill, 1888, p. 285). ‘Two species were brought forward by its
author (lib. cit. p. 481 et xii. 1903, p. 569); and I have seen a
third *; all from Northern India. The following considerably
extends its southern range.

* XENOJOPPA KALI, sp. noy.
A shining and metallic, particoloured species. Head buccate and black, with the
inner ocular orbits flavous and most of the mouth rufescent; clypeus not discreted.
Antennz setaceous, not elongate, with the seven central flagellar joints white.

ICHNEUMONINA IN THE BRITISH MUSEUM. 163

1. FOSSIFRONS, sp. n.

A stout, tricoloured female: head and thorax pale stramineous
with occiput through ocellar region and centre of frons to
scrobes, antennz (except a discal central seven-jointed white
band), and dise of thorax mainly, black; abdomen and legs clear
red with only apices of sixth and seventh segments subnigrescent
before the narrowly white anus, and tarsi discally infuscate.
Head posteriorly as broad as eyes, with both temples and
cheeks buceate; frons depressed throughout, glabrous and glit-
tering, slightly transaciculate below ocelli, with the elevated
orbits incurved below ocelli; face and clypeus transverse, not
disereted, distinctly and in centre rugosely punctate, with apex
of the latter centrally a little concave, not dentate; labrum
exserted. Thorax with pleure and sternum flavidous ; mesonotum
shining, sparsely and coarsely punctate, black with elevated
callosities before and below radices and two discal vitte flavous ;
notauli elongate, fine and subparallel; metathorax dull and
punctate-aciculate, with petiolar area strongly concave, centrally
stramineous and very long; areola proportionately short, trans-
verse, extending to base, apically concave and laterally rounded,
emitting no costule; spiracles elongate, dentiparal arez produced
but with no apophyses, externally stramineous. Scutellum
slightly convex, very coarsely punctate, laterally finely carinate
to the excised apex; flavous with a black longitudinal streak
down its centre to near apex ; postscutellum flavous. Abdomen
closely punctate and dull; basal segment shagreened, with sides
of the broad postpetiole distinctly punctate; gastrocceli striate
and deeply impressed, broader than the aciculate intervening
space; hypopygium extending to apex, terebra black and hardly
exserted. Legs normal, anterior paler, claws minute; anterior
tarsi subdilated, hind cox subacutely dentate on their inner
side. Wings hyaline, radix and tegul fulvous, costa and stigma
black ; areolet triangular, coalescent above, distinctly produced
externally, broader than high; nervelet, and another in second
recurrent, distinet ; discoidal cell subparallel, lower basal nervure
vertical and very slightly postfureal. Length, 13mm. @ only.

Thorax czerulescent and metallic, with its disc green and reticulate ; notauli wanting ;
areola small and quadrate, emitting the entire costulz from near its apex; petiolar
area narrow and parallel-sided. Scutellum stramineous and subglabrous, elevated
and laterally margined throughout. Abdomen fusiform and brilliant blue with
basal sezment fulvous, its apex and the apices of the second and fifth to seventh
segments pale stramineous ; first segment convex and nearly smooth, the second a
little dull with longitudinal striation and large gastrocceli; terebra not exserted, its
base covered by hypopygium; venter black with the two basal segments white and
alone plicate. Legs clear fulvous, with only the anterior coxz white and
hind tarsi black. Wings slightly clouded; stigma black, radix and tegule testa-
ceous ; cubital nervure, bearing very slight nervelet, straight and subparallel with
the anal; areolet not coalesced above, radius apically retlexed. Length, 14 mm.
2 only.—Extremely distinct from both Cameron’s species in its conspicuous
coloration, black face, narrower abdomen, and in the straight inner cubital nervure
with sessile areolet and apically curved radius. India bor.; Dehra Dun and, I
believe, Assam. fs
11

164 MR. CLAUDE MORLEY ON AFRICAN

—The peculiar prolongation of the elevated frontal orbits below
the ocelli is remarkable.—Taken at Entebbe in Uganda between

10th and 20th March, 1914, by C. C. Gowdey.

CTENICHNEUMON.

The third of the subgenera into which Thomson split the
Wesmaelian genus Amblyteles (Opuse. Entom. xix. 1894, p. 2083).
I have already assigned to it one South African species (Ann.
S. Afr. Mus. xv. 1916, p. 365); three of the paleearctic kinds
that extend to Algeria fall herein ; and Dr. Roman has recently
shown that the Egyptian A. tauricus Kriechb. (Kntom. Tidskr.
xxxl. 1910, p. 157, ¢ 2) is also referable to it.

1. CASLANOPYGUS, Sp. n.

A slender and somewhat small, black male with the antenne
(except apices), anterior tibiz and femora, face (except its stra-
mineous orbits) and clypeus, fulvous; and the anus from base
of fifth segment, crimson; wings evenly infumate. Head small
and transverse, posteriorly, obliquely constricted; frons and face
subconfluently punctate, clypeus irregularly punctate, short and
impressed in centre of its truncate apex; labrum exserted.
Thorax immaculate black, closely punctate and not dull; notauli
deeply impressed; metathorax convex, with stout carinz ; areola
subtriangular, apically truncate, basally constricted and acuminate
at postscutellum ; spiracles elongate, apophyses wanting. Scu-
tellum deplanate, apically aciculate and laterally carinate to
centre. Abdomen sublinear and dull with second segment striate;
postpetiole abruptly explanate, closely aciculate, with spiracles
prominent and its apical angles punctate ; gastroceli deeply
impressed and somewhat large; valvule red, extending to apex.
Legs slender and black with the front tarsi, tibie, and apical
half of femora internally, pale; tarsi setiferous. Wings small,
infumate throughout; tegule and stigma black; areolet exactly
pentagonal, broad above; nervelet wanting, lower basal nervure
slightly postfureal. Length, 11-12mm. ¢ only.—Closely allied
in coloration to Amblyteles castanopygus Steph.—Males were
taken on the western foot of the Aberdare Mountains, at an
altitude of 8300 feet, on Ist March, 1911, in British E, Africa.

Subtribe Heresiarchini.

The following genera are here grouped under this subtribe in
a somewhat different sense from that intended by its erector in
Smith’s ‘Insects of New Jersey,’ 1900, p. 567; though hardly
from that of Wesmael when erecting the typical g genus Her esiarches
(Mém. couron. Acad. Belg. 1859, p. 93), since the latter makes no
reference to the shape of the face which, on the contrary, is
shown (as figured) to be not transverse. Its meeting
spiracles are linear, and the restriction of the group in my
‘British Ichneumons’ of 1903 was for local purposes only.

ICHNEUMONINA IN THE BRITISH MUSEDM. 165

As a matter of faet there appears to be but a single foundation
which consists solely of the simple mandibles, ending in a single
point or tooth in place of the bidentate apex usual in Ichneu-
monide, to distinguish this group from the Oxypygini; and I do
not find that justoren endlls characters (Proc. U.S. Nation. Mus.
1900, p. 12), ‘‘ Head, viewed from in front, broader than long ;
occiput strongly concave, the temples broad, full,” no repetition
of which is made at lib. cit. p. 20, are constant. At all events
the following genera all have the face longer than broad and
bear facies of Oxypygini, usually with the remote hypopygium of
that subtribe. Some local cause for which we are at present
unable to account—not improbably peculiar ease of emergence
from the host-pupa—has eradicated the lower mandible, and on
that account thrown these genera in a sufficiently convenient
group, differing in nothing but the mandibular structure from
pe iunieins and Oxypygini*. In all other respects M/yermo,

Chiaglas, ete., might be considered true Stenichnewmon-species,
while Fileanta ruficauda Cam., from India, is almost a true
Amblyteles palliatorius Grav., from Kurope.

Neither Wesmael nor ibe phone recognized the present
Subtribe; but Schmiedeknecht in 1902 adds to the already
known four genera Myermo, Fileanta and Setanta from India, of
which I have shown (cf. Amblyteles supra) the last to be a mere
synonym.

List of Heresiarchid Genera.

STENopvonTUS Berthoumieu, Ann. Soe. Kntom. France, Ixv. 1896, p. 346= Grat-
hoays Wesmael, Nouv. Mém. Acad. Brux. xviii. 1844, p. 165 (nec Westw.
1842).—Kuropa, Africa bor.

Herestarcues Wesmael, Mém. couron. Ment Belg. viii. 1859, p. 93.— Europa.

RuexipErRMuS Forster, Verh. preuss. Rheinl. xxv. 1868, p. 192 (species eastat.).—
Huropa.

Pragrotrypees Ashmead, Proc. U.S. Nation. Mus. xxiii. 1900, p. 20.—Amer. bor.

Gyroponra Cameron, Ann. Nat. Hist. vii. 1901, p. 485.—India.

FILeaAnta Cameron, Ann. Nat. Hist. vii. 1901, p. 525.—India.

Myerrmo Cameron, Ann. Nat. Hist. vii. 1901, p. 523.—India, Burma.

Mrosoppa Cameron, Zeitschr. Hymen. u. Dipt. 11. 1902, p. 891.—India.

ALGATHIA Cameron, Zeitschr. Hymen. u. Dipt. i. 1902, p. 392.—India, Burma.

Crraguas Cameron, Ann. Nat. Hist. ix. 1902, p. 152 et 1. c. xx. 1907, p. 81.—India.

Na#narta Cameron, ae Nat. Hist. xi. 1903, p. 313.—India.

Casprpina Cameron, Trans. Entom. Soc. Lond. 1903, p. 219.— Assam.

TrRIProgNaTuus B ees Genera Insect. xviii. 1904, p. 49.

MacropHatnus Cameron, Rec. Albany Mus. i. 1905, p. 232 et Ann. 8. African
Mus. v. 1906, p. 175.

RossetLa Cameron, Ann. South African Museum, v. 1906, p. 176.—Africa mer.
AtLonotTus Cameron, Ann. Nat. Hist. xx. 1907, p. 29.—Borneo.

ORTHOGNATHELLA Szépligeti, Faun. S.W. Australiens, i., ix. 1908, p. 3820.—
Australia.

HereEstaRcnorpes Brethes, Ann. Mus. Buenos Aires, xix. 1909, p. 51.— Argentina.

* In the European fauna, such thines as Ichnewmon rufidens Wesm. and
Amblyteles wriguttatus Grav., fall into the present category.

166 MR. CLAUDE MORLEY ON AFRICAN

MAGWENGA, gen. nov.

Mandibles unidentate, apically acuminate, not externally sul-
cate ; cheeks and temples elongate and quite straight, not buceate ;
frons centrally carinate; occiput and juxta-scrobal orbits acutely
margined. Flagellum filiform and apically strongly attenuate,
its basal joints elongate. Metathoracic spiracles linear and
elongate; metanotum with neither carine nor are, its centre
slightly elevated. Scutelluin laterally carinate throughout. Post-
petiole not at all sculptured, with no apical fovea; gastroccell
large ; apex of second segment centrally emarginate ; hypopygium
remote from anus. All the claws stoutly pectinate. Wings
hyaline with a central fascia and apical spot nigrescent.— Nearest
to Caspipina, but the cheeks are not buccate, the flagellum not
compresso-dilated ; metanotum excarinate; postpetiole neither
basally elevated nor apically foveate; basal nervure net con-
tinuous ; tarsi pectinate and wings maculate.

1, MACULIPENNIS, sp. n.

A dull black female, with the hind coxe red; the wings
centrally black-banded, with a circular apical spot. Head long,
glabrous and sparsely punctate, behind the eyes parallel; clypeus
and face deplanate, hardly discreted, with confluent and some-
what close punctures, apex of former truncate and laterally mar-
gined; palpi white; cheeks longer than breadth of the white
base of mandibles. Antenne black with their centre, except
laterally, white-banded. Thorax stout, convex and closely punc-
tate, discally smooth; notauli obsolete, speculum small and
elittering ; metathorax discally smooth and shagreened, its base
depressed but not suleate; petiolar area triangular and laterally
striate, very small; apophyses wanting. Scutellum large, smooth
with a few punctures, broadly margined, with extreme apex
aciculate. Abdomen elongate-fusiform, roughly punctate, with
anus from fifth segment and the postpetiole glabrous; petiole, or
at least its base, red; gastrocceli large and double breadth of their
intervening space; terebra hardly exserted. Legs stout; an-
terior with coxe, inner side of tibize and of their femoral apices,
flavidous; hind trochanters and their indefinitely scopulate
coxe bright red; third and fourth hind tarsal joints white.
Wings hyaline and not broad ; areolet not quite coalescent above,
lower basal nervure strongly postfurcal ; a central broad infumate
band from the basal to centre of the disco-eubital nervures and a
round concolorous spot on apex of radius; hind wing also
infumate in radial cell. Length, 10-llmm. @ only.—These
remarkable females are from 3700 feet at Buddu on the west
shores of the Victoria Nyanza, on 19th September; and from
3800 feet to the south-east of Buddu, in the Tero Forest, on
26th September, 1911, in Uganda.

ICHNEUMONINA IN THE BRITISH MUSEUM. 167

Mio0s0PPaA.

Head not broader than thorax, roundly constricted posteriorly ;
temples shorter than eyes; face subdeplanate and not discreted
from clypeus ; cheeks not short, mandibles gradually constricted
to apex, palpi slender and elongate. Antenne stout and not
longer than body, broadly white-banded, and beyond their centre
compresso-dilated. Mesopleurze obliquely carinate above their
centre; metathorax gradually and roundly curved, with its
lateral carine entire ; petiolar area parallei-sided, narrow and
extending to basal third of metathorax ; areola obsolete, incom-
plete, remote from metanotal base; spiracles suboval and thrice
longer than broad; apophyses wanting. Scutellum deplanate,
stoutly carinate laterally to near its centre; postscutellum
basally bifoveolate. Abdomen with first longer than second
segment, and gradually explanate from petiole; second and third
segments closely punctate; gastroceeli superficial, Jonger than
broad with the inter vening space closely striate; second and
third ventral segments plicate ; ; terebral valvule exserted, as long
as two anal segments. Basal nervure not continuous through
median ; “arerancllet indistinet.—-I place this genus [of which ie
above is Cameron’s diagnosis (Zeitschr. Hymen. u. Dipt. 11. 1902
p. 391), slightly emended by an examination of the genotype |
next the Platyurini, with which the structure of the basal
segment 1s in exact agreement; and J am not sure that it were
aa better actually merged therein, in spite of the mandibular
conformation. The Simele known species from India was
described, loc. cit., incorrectly as female.

1. QUADRILINEOLA, sp. 0.

A dull, black and closely punctate male with the legs except
basally, antenne except disc of scape, mandibles except apices,
and the palpi fulvous ; facial orbits shortly and two facial vitte,
a small callosity before radices, a dot at apical angles of post-
petiole, scutellum except extreme base, and the postscutellum
flavous. Clypeus apically truncate and a little reflexed ; frons
and face closely punctate and pale pilose; areola short, sem1-
circular, and apically incomplete; scutellum shining, sparsely
punctate, and subconvex ; gastrocceli small and valvulz luteous ;
coxe and trochanters black; tarsi simple; areolet very broad
above; radius sinuate. Length, 1lmm. 6 only.—The sculp-
ture and outline are similar to Cratichneumon annulator Fab.,
rather than Platylabus pedatorius Fab., both black males of like
form and structure; the distribution of the pale marks is
peculiar.—The type was taken at Deelfontein in South Africa
about 1903 by Col. Sloggett.

Subtribe Listrodromini.
NEOTYPUS.
To the two species of this genus, which is likely to find its

168 MR. CLAUDE MORLEY ON AFRICAN

headquarters in Africa, and was erected by Forster (Verh. pr.
Rheinl. 1868, p. 194), that have already been recorded by Kriech-
baumer and myself, I can now add a third.

1. OBSCURATOR, sp.

A small, dull, black Hane with the thorax and petiole entirely
red. Head transverse and narrow behind eyes; frons closely
punctate and apically trans-striate; face finely trans-striate,
centrally elevated; clypeus glabrous, broad, uneven, bifoveate,
apically emarginate and centrally subreflexed ; labrum concealed,
mandibles slender and duli testaceous. Antenne elongate, slender,
apically attenuate, with the basal joints elongate and centre
white banded. Thorax short and stout, closely and evenly
punctate, with deep notauli and dull speculum ; petiolar area
covering two-thirds of metathorax, discreted, its central area
deeply concave and trans-striate; areola proportionately short,
transverse-semilunate, not extending to the depressed base,
emitting weak costule from centre; spiracles oval and oblique,
apophyses stout and obtuse. Scutellum red, convex, dull, closely
punctate, laterally margined to near apex. Abdomen ovate,
dull, very finely punctate; basal segment red and linear, abruptly
explanate at the black postpetiole, which is finely punctate-
aciculate with its apical angles and those of the second segment
white-dotted ; gastroceeli very small; apex of fifth segment
narrowly, of sixth breadly and whole of seventh, white; terebra
black, not exserted. Legs black and not short, the anterior
tibie and front femora mainly testaceous; coxe simple, claws
pectinate. Wings hyaline, stigma and nervures black; areolet
broad above, nervelet distinct, basal nervure continuous through
the median. Length, 8mm. @ only.—The type occurred at
Mlanje on 26th May, 1913, in Nyasaland.

Subtribus Platylabini.

PLATYLABUS.

To my Table of the southern African species of this genus
(Ann. 8. Afr. Mus. xvi. 8, 1917, p. 201) I am now enabled to

add a couple of very distinct species, and the alternate sex of
another.

1. ATRICINCTUS, Sp. n.

A rufescent-testaceous species with profuse black markings,
and the setaceous flagellum pale banded. Head slightly broader
than thorax, and but little narrowed behind the strongly pro-
minent eyes; ocelli and mandibular apices alone nigrescent ; face
and mouth flavidous, the former obsoletely punctate and not
discreted from the apically truncate clypeus; labrum exserted,
lower mandibular tooth very small. Thorax dull, with the
mesonotum shagreened and notauli distinct ; petiolar area, post-
scutellum, and dise of frenum abruptly black: areola as long as

ICHNEUMONIN4! IN THE BRITISH MUSEUM. 169

broad, basally rounded, emitting fine costule from its centre;
basal area, distinct, spiracles elongate, apophyses wanting. Scu-
tellum dull and somewhat elevated with dark lateral carinz to
near apex. Abdomen cylindrical and dull with base of segments
two to five, and centre of the second, black-banded; basal
segment smooth, shining and basally whitish ; thyridii transverse-
linear, with the intervening space very narrow; valvulz obsolete.
Legs slender with apices of the hind coxe and of their femora
nigreseent, of their tibie infuscate; hind tarsi stramineous
throughout. Wings normal and flavescent, with stigma and
tegule flavous; areolet as broad as high, subcoalescent above,
emitting recurrent from its centre; basal nervure continuous
through median. Length,10mm._ <o only.—The black-banded
abdomen and metathorax are distinctive-—One male at Fort
Portal Road, Mbarara, in Southern Toro, at 4000 feet, on 22nd
October, 1911, in Uganda.

2. MEDIORUFUS, sp. n.

A pale testaceous species with sparse black and white markings;
thorax tricoloured. Heal transverse and abruptly constricted
behind the prominent eyes; frons smooth with its base, like the
elevated ocelli, black: all the orbits broadly, mouth and face.
stramineous, the last deplanate and obsoletely punctate, not
discreted from the apically truncate clypeus, which is centrally
impressed; lower mandibular tooth and the occipital margin
strong and nigrescent;. ¢ with centre of frons and whole oeciput
black. Antenne setaceous, black and a little incrassate beyond
their broadly white-banded centre ; basal joints rufescent beneath.
Thorax dull; mesonotum laterally deep black, which colour is
hounded internally by a narrow flavous longitudinal band;
elongate callosities before radices clear stramineous, notauli
short and deeply impressed; metanotum shining and sparsely
punctate; areola cordiform, longer than broad and apically sub-
constricted, emitting strong costule before its centre; petiolar
area transaciculate, spiracles elongate, apophyses wanting.
Scutellum margined laterally to apex and, like the postseutellum,
nitidulous and stramineous. Abdomen dull, discally deplanate,
of 2 elongate-fusiform ; the four basal segments closely sculp-
tured, with postpetiole and base of the second centrally aciculate ;
thyridii triangular, not small and hardly narrower than the
intervening space; anus from base of sixth segment, valvule
and the slightly exserted terebra, black ; seventh segment of both
sexes discally white; ¢ with dise of third and fourth segments
black-marked. Legs slender and normal, with the hind tarsi
apically nigrescent and their tibie straight. Wings hyaline,
tegule and stigma testaceous ; areolet as broad as high, coalescent
above and emitting recurrent slightly ae its centre; basal
nervure continuous. Length, 9-10 mm. ¢ 9 .—The tr icoloured
mesonotum and anus are remarkable.—The type occurred on

14th July, 1910, at Blantyre, in Nyasaland, to Dr. J. KE. 8. Old;

170 MR. CLAUDE MORLEY ON AFRICAN

the androtype was captured at 6400 feet, on Mount Kokanjero,
to the south-west of Elgon, in early August 1911, in Uganda. A
rather differently coloured female is labelled simply ‘“ Erythria,”
and shows no structural modifications.

3. cETA Mori.

A female of this species which I brought forward in Ann.
S. Afr. Mus. xvi. 8, 1917, p. 203, with the capital disc entirely
red, was taken at St. James, in Cape Town, on 24th October,
1911, by K. H. Barnard.

4. vVALLATUS Morl.

I was only able to indicate the male of this species (Ann.
S. Afr. Mus. xv. 1916, p. 370). The @ differs in having the
ocelli alone black, the six basal flagellar joints apically sub-
nodulose and the first pale beneath; the stramineous thoracic
markings are obsolete and mesonotum totally black; the areola
not broader than long; the scutellum is testaceous, postpetiole
nigrescent, terebra black and but slightly exserted.

Fort Portal Road, at Mbarara, Southern ‘Toro, at some 4000 feet,
on 22nd October; and on the north-west shores of the Victoria
Nyanza, at 3800 feet, in the middle of September 1911, in
Uganda.

Subtribe Pheogenini.
BENECLES.

Head fully as broad as thorax; apex of clypeus broadly and
centrally impressed, its sides obliquely suleate above; mandibles
stout and parallel-sided with the upper tooth acute and the lower
the shorter; temples small and obliquely constricted. Antenne
longer than body with scape distinctly shorter than basal
flagellar joint, which is much longer than the second. Meta-
notum slightly impressed basally and not produced apically, with
complete areze and no apophyses; areola broader than long and
subconstricted towards its truncate and strongly carinate apex ;
spiracles small and subcireular. Scutellum evenly subconvex and
basally carinate on either side; postscutellum stout. Abdomen
elongate and exactly parallel-sided from the postpetiole to the
exserted terebra; gastroceli large, oblique, deeply impressed
and remote from base; lunule large, anus pale-marked, terebra
reflexed and sometimes subvertical. Hind legs much longer
than the anterior. Areolet pentagonal, constricted above; nerve-
let wanting; basal nervure continuous through the median ;
nervellus intercepted far below its centre.

“The small, round metathoracic spiracles refer this genus to
the Pheogenint. In Ashmead’s ‘ Classification of the Ichneumon
Flies’ it would come in near Herpestomus. Characteristic
is the long, sharp pointed apical and indistinct subapical tooth
of the mandibles,” says Cameron, in erecting this genus
(Entom. xxxvi. 1903, p. 260) on a single Indian individual.

Pe oe Te ee

ICHNEUMONINA IN THE BRITISH MUSEUM. Wt

I consider the genotype more remarkable in its large and remote
gastroceeli, subcubical head, which is no narrower than the thorax,
the reflexed terebra, and the very broad cheeks which are
anteriorly hardly narrower than the eyes. In all species the
metathoracic spiracles are suboval.

1. DIMIDIATUS, sp. n.

A black species with the thorax and petiole red, anus and
trochanters white. Head transverse, deep black, and posteriorly
narrow ; frontal orbits broadly, and centre of the external ones
narrowly, white; face and clypeus black, deplanate, closely punc-
tate, discreted, with the latter broad and apically truncate; palpi
white. Antenne filiform, broadly white-banded with the basal
joints rufescent beneath. Thorax cylindrical, dull and brick-red
with pronotum white and prosternum nigrescent ; mesonotum
closely shagreened, with elongate but superficial notauli; meta-
thorax somewhat short, closely punctate; areola as broad as
long, apically truncate and basally rounded, emitting distinct
central costule ; basal area entire, the petiolar subvertical and
transaciculate ; apophyses wanting, spiracles exactly ovate. Scu-
tellum shining, quadrate, sparsely punctate and laterally margined
throughout. Abdomen deep black, with the first segment brick-
red; three basal segments dull, the first convex and shagreened
with prominent lateral tubercles; thyridii transverse-linear,
intervening space very narrow : segments five to seven deplanate,
quadrate and strongly nitidulous, the seventh white; terebra
slender, black, slightly exserted. Legs black with the anterior,
except basally, subtestaceous; all trochanters, except apices of
hind ones, pure white and calearia subconcolorous ; hind coxe
simple. Wings hyaline and somewhat small; tegule and stigma
dull ochraceous; areolet pentagonal and not large; basal nervure
continuous. Length, 9 mm. 9 .——Certainly allied to the genus
Dicelotus in its abdominal structure.—Taken at Durban during

1902 hy F. Muir, ex coll. D. Sharp.

2. POLITANUS, Sp. 0.

A black species with most of the thorax and legs testaceous,
and the anus white. Very similar to the above species in struc-
ture, but with the basal segment black ; the whole metanotum
with frenum and a central spot at mesonotal apex black; the
external orbits broadly, with the face and mouth except longi-
tudinally in the centre, white; the legs are not white-marked
and the areola is a little transverse. Length,7mm. Q only.—
Captured at Western Ankole during the middle of October
1911, at between 4500 and 5000 feet, in Uganda. Pheogenini
appear rare in Tropical Africa; of the twenty species recorded
from the Continent, no more than four are known south of the
Sahara, besides those here described.

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ay pat thieale bh PRES pty tip Bera 7 oe atte tea oF HEEB vil noepa Abeae E
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P. Z. S. 1919, CUNNINGHAM, PI.

Bale & Danielsson, Imp.

HYBRID COMMON FOWLS.

RESULIS OF A MENDELIAN EXPERIMENT ON FOWLS. N73

12. Results of a Mendelian Experiment on Fowls, includ-
ing the Production of a Pile Breed. By J. T.
CunnincHam, M.A., F.Z.S.

[Received April 29,1919: Read May 27, 1919.]
(Plate 1.*)

In the ‘ Proceedings’ of this Society for 1912 there was pub-
lished a paper by me under the title ‘‘ Mendelian Experiments
on Fowls,” in which I recorded certain results of a cross between
a male black-red Gallus bankiva and a Silky hen. The object of
the present paper is to describe the later generations obtained
from this cross, and especially to describe in detail how it
produced a “pile” type of colour which bred true. In my
former paper I mentioned the fact that the two white cocks
of the first brood of the F, generation at the age of 41 months
showed a yellowish-orange band of colour across the loins, and
the second of them had also some very pale patches on the neck
ventrally and above the eye on each side. In April 1916 I
exhibited at a meeting of the Zoological Society a number of
skins of the fowls bred from this cross, including specimens
of the pile birds of both sexes, but hitherto I have not recorded
in detail the characters observed in successive generations of the
crossed birds, as I propose to do in the present paper.

All the generations were descended fiom a single pair of the
F, generation, hatched from the original crossed pair at the
Zoological Gardens. To distinguish the generations, broods,
and individuals I have adopted the following formule: the
number indicating the generation is placed below the line after
the letter F, it 1s followed by a Roman numeral indicating which
of the successive broods of that generation the bird belongs to,
then follows an Arabic numeral indicating the particular bird
of the brood, and lastly the symbol of sex. Thus F,V 19 isa
hen numbered | in the 5th brood of the 4th generation.

In my former paper, which dealt only with the F, generation,
I pointed out that the only two whites of the first brood showed
a trace of colour, whereas the original silky of the crossed pair
had no colour. Both these whites were cocks, and F,I 1 ¢
showed a band of faint orange-yellow colour across the loins,
while F,1 2 showed a similar hand, still fainter, together with
some very pale patches on the neck ventrally and over each eye.
This condition was noted on September 30, the birds having
been hatched on May 15 (age 43 months).

There were two others, both white, in a second brood, date of
hatching not recorded. Of these F,IT 2 also showed a tinge
of yellowish colour across the loins. F,I1 19 was given away
at an early age, and no pigment on this hen was noted, although,
considering that at the time I was not aware of the sexual

* For explanation of the Plate see p. 202.

174 MR. J. T. CUNNINGHAM ON RESULTS OF A

difference in this pigment, it is not certain that pigment was
entirely absent.

My view concerning this trace of pigment in otherwise white
recessives was that the segregation of the colour-character was not
complete, but that a trace of the pigment-character passed over
to the recessives. If this happened at every generation the
amount of pigment should increase, and in order to test this I
mated the slightly pigmented recessive with a coloured dominant
bird, The birds of F, were mated thus:

(1) F,I 136 white x F,I 6@ coloured and
F,I 89 coloured.
(2) F,I 23 white x F,I 7@ coloured.

The two females of the first mating both had light-coloured
heads, while No. 7 in the second mating had a black-coloured
head. his is an individual difference in the colour of the
dominants which is further discussed below.

F. I was from the second mating shown above, and consisted
of four chicks hatched, three “ white ” or recessive, one coloured.
This, of course, shows that the hen parent was heterozygous for
colour, as if she had been a pure dominant all the chicks would
have been coloured. In this brood I noticed for the first time a
peculiarity of the recessive chicks in the down stage. The
coloured chicks, as in other black-red breeds, are longitudinally
striped: a broad dark stripe runs along the mid-dorsal line, and
on each side ig a narrower dark lateral stripe which on the head
passes through the eye, the intermediate parts and ventral down
being of a lighter brown. In the recessive chicks there are white
bands where the darker bands oceur in the coloured chicks, and
yellow down on the rest of the body. As will be seen later, this
agrees with the “pile” character in the fledged and adult birds.

The first brood of F,, hatched May 13, were carefully examined
on June 23, when they were 6 weeks old and nearly fledged.
All three vecessives had colour on the throat and breast, but in
different degrees: No. 2 had most colour, not only brown colour
quite distinct but also black feathers here and there; No. 3 had
less colour, and No. 4 least. At this date there was no coiour on
the back.

F, 11, hatched about May 23, 1912, consisted of 5 chicks,
4 coloured, 1‘ white” or recessive. This last showed very
slight but distinct colour on the throat on June 29, when
5 weeks old, and no colour on the back. As in Brood I, at
6 weeks of age, this is the condition when the chicks are first
fledged, when they have developed the first adult plumage.

In brood F,III, hatched May 28, there were 11 chicks,
6 coloured and 5 white. Of the latter, all except No. 7 showed
some brown on throat and breast on July 17, when they were
7 weeks old. A slight but distinct trace was afterwards seen on

the throat of No. 7.
At this time I knew nothing about the colour-characters of

MENDELIAN EXPERIMENT ON FOWLS. 175

pile fowls, and did not suspect that sex had an important
influence on the development of the small amount of colour
which I was observing on the white or recessive chicks. In the
second generation F, I 1&2 were both males, and I made a
note that the colour on the back of these seemed to get a little
deeper as they grew older. F,II 1 was a white female and
Fi IT 2 a white anal. The former was given away at an early
age without any record being made of thie presence of colour on
ne breast; F, II 2 was killed on November 8, 1912, when it was
about 5 months old, and it is recorded in my notes that it had
a tinge of yellow over the back (which includes the dorsal part of
the folded wings), but that the colour was very slight.

In the three broods of F, above described the ‘two whites of
the first brood were both males, the single white of the second
brood was also male, of the oe wbthas in the third brood
FIT 7,8,&11 were males, 9 & 10 females. On August 25 I
first observed with certainty that in the males the co date dis-
appeared on the throat and breast as it developed on the back
and loins, while in the females it continued to increase somewhat
both in depth and extent on the throat and breast. At this date
in FI 2¢ the brownish colour was much diminished on the
br east, and there was quite a deep buff on the back and shoulders.
[a3 i 3.6 the colour on the back was not so deep. F,I 43 was
killed on J uly 23, its skin was preserved, and is still in my posses-
sion showing a narrow band of rather deep brown extending from
the throat to the breast, and scarcely any yellow on the back, At
this date this brood was 33 months old, and, as I found the young
fowls were sexually mature at 5 months of age, it is evident that
this change of colour in the cocks from the breast to the back
takes place at puberty, while before that time, in the first adult
plumage, the cocks resemble the hens. On ne same date the
only white in the second brood, F, IL 5, had slight brown on
throat and breast, rather deep buff on back and shoulders.
In the third brood which, of course, was younger, the cocks
showed colour coming on back, and a little still on the breast,
except No. 7, which had least colour of all the five whites. The
hens 9 & 10 had brown on throat and no colour on_ back.
In August of the following year I recorded in my notes the fact
that of the surviving white, 7. e., piles of this generation, the two
cocks, F,I 2&3, Taal buff colour on the eth: but no 26 lone on
the breast, while ‘the hens, F, I 9& 10, had brown on the breast,
but no Bolen on back or loins. Two further points are to be
noticed in F,, (a) that the amount of colour in the recessives for
plumage colour varies in the individuals though present in all,
(6) that the maximum amount of colour is distinctly greater
than in F,. The total number of F, chicks hatched was 28,
11 pile, 12 coloured.

H, LO3y ands Heal ol:
The first brood of this generation was hatched April 11, but all

176 MR. J. T. CUNNINGHAM ON RESULTS OF A

the yellow chicks in it died in the first week, so there were no
recessives In plumage-colour among the survivors. The parents
of the second brood were F, 1 2&3 x F,Il 2&49, the cocks
recessive, the females welleured. As the coloured hens were
known to be heterozygous for plumage-colour, it does not much
matter for this character which of the cocks or hens were parents
of individual chicks. F,II 5 was recessive male, F, Il 6 &7
recessive females. These all had some brown on the throat in
the first mature plumage in different degrees, most in F, II 5
in which it was considerable. ‘There were seven chicks in ihe
brood, the other four being coloured, but Nos. 1 & 2 died in
the first week. No. 3, a cock, had plumage of remarkably rich
colour, a port-wine red on the saddle and shoulders, rich orange
neck and saddle-hackles, a deep glossy black on breast and tail.

Altogether nine broods were hatchel in F, the parents in
each case being one pile and one coloured. Of these Fs,
16 were pile and 25 coloured, a proportion not approaching very
closely to the equality expected on the Mendelian theory. All
the piles or recessives had more or less colour on throat or
throat and breast, some only a little, while F, V 19, hatched
June 13, had in July, when about 5 weeks old, deep and con-
tinuous ‘brown on throat and breast and extending in traces
over the whole ventral surface. The mother of this one was
F, IIL 9¢, which had, at any rate when examined the previous
year, only a little brown on throat. In the hens the amount
of brown usually increases somewhat as they grow older.

The birds of F, kept for breeding were :—

F IL 36d, coloured —very richly coloured.

Jp lu 5d, Le

F,IV 39, coloured.

FE. V 19, pile—much brown on breast and abdomen.
F, VIIL 69, coloured, silky.

if VAI aS colour af, silky.

F,VUI 8d, coloured normal.

Seven broods were hatched from these, forming the F, gene-
ration. Of these, F,III, F,1V, and F, V were offspring oe
parents similar in zolenr. The parents of E. IV were F II 5¢
pile and F, VIII 49 pile. This mating was made to ascertain if
any segregation took place between the Brow and the white of the
pile when they were bred together. Hight chicks weie hatched
and they were all pile. There were four cocks and four hens.
and in the sexually immature condition, after they were fledged,
all had more or less brown on the breast, though the amount of
colour varied from “considerable” in F_1V 3 to very slight
in F,1V 7& 8, both 9. This shows, 'T think, that the piles
breed true, that there is no segregation between the brown and
white.

The four broods—F, 1, F, 11, FB, VI, and F, Vil—were off-

spring of matings of pile with coloured as in previous generations,

MENDELIAN EXPERIMENT ON FOWLS. ~ 177

all the coloured being heterozygous and carrying pile. ‘The
total number of ads in these four broods was 26, of which
12 were pile and 14 coloured. In Broods I & II only one chick
survived to maturity in each, and that in Brood I was coloured.
F.Il 39 was a hen with much brown on breast. The male
parent of Er namely F, VII 24, had also, before the sexual
change of colour took pl: ce, much brown on breast. In the piles
of the other two broods there was brown on the breast of the
hen or yellow on the back of the cocks in all cases, but no very
‘distinct evidence of increase over the previous generation.
Owing to my visit to Australia in 1914, I was not able to
examine these two broods for colour till they were three or four
months old, when the males were already losing the colour on
the breast and developing it on the back, and the colour on the
back is never more than light yellow, while in the hens it is a
much deeper brown on the breast.

F, ILI was the offspring of F, VIII 6@ and F, VIII 7 3, both
dark-coloured birds with silky plumage. As these were both
heterozygous for plumage-colour there were some piles among
the offspring, the numbers being 5 coloured and 3 recessive. Of
the latter one died in the down stage, and the other two were
both males which had brown on the breast when first fledged.
F. V was the offspring of F, VIII. 83 x F, IV 39, both coloured.
Only one I, V 7 was pile, and this when first fledged had mode-
rate brown. on the breast.

The birds kept from I, for breeding again were :-—

F, Il 39, pile with much brown on breast.
F,V 54, coloured.

F, Ill 1 & 29, dark silky.

F, VI 164, coloured, dark.

F, VII 1, coloured.

F WAIN SG 49, coloured, light.

F. LN OF pile with brown on breast.
F. VI 22, coloured dark.

F. VI 39, coloured lhght.

F. Well She pile.

F, V 69, coloured light, silky.

F, Generation, 1915

In the F, generation eleven broods were hatched. Of these
six were from matings of pile with coloured, and of the chicks
produced by these 24 were coloured and only 9 pile. This is very
far from the expect ation of equality, and the difference is erate
than in F, or F,. In I, several of the broods were small,
though fertility bad deere: ased, and it is possible that more of oh
recessives were either unfertilized or died before hatching. In
all these piles of I°, as well as in those produced by matings of
heterozygous coloured birds together, brown colour was present
on the breast in the first feathering, disappearing in the cocks as

Proc. Zoou. Soc.—191Y9, No. XII. 12

178 MR. J. T. CUNNINGHAM ON RESULTS OF A

it developed on the saddle. I have no record that any general
increase in the pigmentation was noticed, but on the other hand
no individuals are noted as having slight or very slight colour on
the breast ; the records in my notes are either simply brown on
breast, or brown on breast moderate.

The Pile Coloration in Fowls.

I think the above evidence is sufficient to show that the
recessives from this cross of Black-red with White Silky were not
pure white, but developed in 6 generations into Pile fowls. The
Pile, a word which is probably related to pied and magpie, is
described by Mr. John Douglas, in Lewis Wright’s ‘ Book of
Poultry,’ 1885, as a variety of Game-fowls. According to the
description, it is white where a Black-red is black: neck and
saddle-hackles light chestnut-red, back chestnut and claret-colour,
shoulder covert and bow of the wings rich claret-red, breast white
or laced with pale chestnut, abdomen and thighs white, tail white
or with a slight tick of black on the sickles. This applies to the
cock; in the hen the head is light golden chestnut, breast a
rich chestnut right up to the thro at, thighs almost pure white,
hackles white faced with yellow- chestnut, back creamy white
with a shade of gold, tail white,

The word chestnut here evidently means much the same colour
as I have called brown, which on my birds is a light reddish
brawn. Nothing is said in the description about the immature
cocks having brown breasts, while in the adult the breast is white,
but this is naturally to be expected considering that the pile is the
absence of black. In the cock of black-red fowls the breast and
tail are black; in the hen the black is chiefly on the back, head,
and tail, and almost absent on the breast and abdomen. Before
sexual maturity the cock in its first plumage naturally resembles
the hen. At the same time it is to be noted that in Mr. Douglas’s
description black is not entirely absent from the pile, and it was
not entirely absent from those which I bred, an occasional black
feather or one partially black having been very frequently
observed.

Mr. Douglas remarks that Piles breed true to colour, as I have
shown was the case in my experiments, but that now and then a
cross of the black-red is thrown in to give hardness of feather,
after which every black-red produced from the cross should be
destroyed. This seemed to imply that the pile was dominant and
that when they were bred together after the cross they produced
dominant piles and recessive black-reds. As this was so contrary
to my own experiments, [ wrote to Mr. I’. Smalley of Silverdale,
Lancashire, an experienced breeder of Game-fowls and well
acquainted with Mendelism, who very kindly answered my
questions. He informed me that Pile is quite dominant to Black-
red so far as Game-fowls are concerned, whether the cross be
made with the Pile male or Pile female. Mr. Smalley also stated

MENDELIAN EXPERIMENT ON FOWLS. 179

that according to theory Pile first came from crossing Black-red
with white, but that such a cross cannot now be made because
there are now no pure white Game, all our whites being ‘“ sports”
from Brown-reds, which bred inter se, always breed white, but
which if put to any other colour would at once produce “ colour”
and most likely that colour would be Pile.

We have now to consider the theory or explanation of the
origin of the Pile coloration. My experiments show that a
recessive pile is produced by crossing recessive White Silkv with
dominant Black-red, and it therefore seems probable that a
dominant Pile would be produced by crossmg dominant white
with Black-red, or perhaps other colour. Such crosses I have not
yet tried, but hope to do so in future. With regard to my
recessive Pile it would seem that the simplest explanation is that
segregation is not complete or perfect, but that the chromosomes
of the white parent, originally without any colour factor, in the
segregation of gametes in the heterozygote take with them some
of the substance of the colour-bearing chromosomes, and so cause
the development of colour. It must be remembered that the
Black-red itself is pied: that is to say, in the cocks the black and
red are in separate areas, almost completely exclusive of one
another ; although in the hens these colours are more mingled, but
with reddish brown preponderating on the breast and abdomen.
The black segregates more completely than the red, but not quite
completely, since there are black feathers or traces of black in the
Piles. It is impossible to suggest that the black is dominant and
the red recessive, for in that case the F, would be black and white
pied. The recessive pile then, having a certain amount of
dominant red or reddish brown with recessive white, might be
expected to segregate, and the effects of such segregation would
be visible when the piles were bred inter se. hat such a
segregation does not occur completely is shown by the fact that
all the offspring produced by pile x pile have some amount of
colour, but the fact that this amount of colour varies in the
individuals may indicate that a varying degree of segregation
occurs, and probably by selective mating a pure white could be
again produced.

The fact, however, demonstrated in the experiments of Bateson
and Punnett * that when the Silky is crossed with a certain other
recessive white, completely coloured black-red birds are produced
in F, just as crossing of two white races of Sweet Pea may
produce completely coloured flowers, suggests that the origin of
the recessive pile in my experiments may require a more com-
plicated explanation. According to Bateson, the colour must be
due to two factors X and Y, and when either of these is absent
the bird is white. In the case of the two recessive whites pro-
ducing colour when mated, one has one colour-factor absent, the
other the other, so that when fertilization takes place the two
factors X, Y are both present and fully developed colour results,

* Bateson, ‘Mendel’s Principles of Heredity,’ 1909, p. 108,

180 MR. J. T. CUNNINGHAM ON RESULTS OF A

The absent factors being represented by small letters, one of the
white parents is Xy, the other xY, or, rather, these symbols
represent the constitution of the wametes, the par ent birds them-
selves must be XXyy or xxYY. The Silky in my experiment
then must contain one factor for colour and its gametes must be,
say, Xy. The Black-red, on the other hand, must contain both
colour- factors, and its constitution must be XXYY and its gametes
must be XY. The F,, then, with colour dominant will be XXYy ;
and the gametes of this will be XY, Xy. The fertilizations in F,
will therefore be XYXY, XYXy and XyXy

The result, it will be seen, is much the same as if the colour was
due to a single factor, the first combination giving pure dominants,
the second dominant (¢.e., coloured) birds carrying recessive
white, and the third recessive white like the original Silky.
The hypothesis of two colour-factors and the presence of one of
them in the Silky thus in no way helps to explain the appearance
of the pile-colour in the later generations of my cross: whether
there is one colour-factor or two, the appearance of colour in the
recessive could not occur if segregation were complete, as Mende-
lians assume it to be. Segregation would result in the recessive
in F, or any other generation having a total absence of one colour-
factor, whether X or Y, and the occurrence of the colour in the
pile shows that some portion of the missing colour-factor has
passed from the dominant to the recessive—in other words, has
not been completely segregated.

Differences in the Coloured Birds.

Although the production of the pile coloration is the most
important of the results of this cross which I have to record, I
propose to discuss peculiarities in the heredity of other characters
and consider how far they depart from expectation according to
Mendelian theory. One of these consists of variations in the
depth of colour in the coloured dominants. ‘These variations did
not form a continuous series, but divided themselves into two
distinct types—a darkand a light. I noticed this first in some of
the coloured birds of F,, wher F,I 6 & 8@ are noted as having
light-coloured heads, F,I 7 as having a black head. In later
generations I noticed the whole plumage of heus with black head
was darker than in those with light head, but I found it difficult
to distinguish the two types in the cocks when adult, though it
was evident enough in the first mature plumage in both sexes.
In ¥, TX, the parents, of which were F,I 16 coloured x F, III
10 Q pile, there were 5 coloured birds to one white, and all the
coloured were of the lighter type, but whether the male parent
was of this type was not noted.

I tried to acertain whether the lighter type was recessive to
the darker. The parents of F, III were both dark silky, and of
the 5 coloured birds in this brood two were of the darker type
with black crests, two were light, and one was intermediate in
general colour, but without the black crest. The parents of F, V

MENDELIAN EXPERIMENT ON FOWLS. 181]

were both of the light-coloured type, and the brood consisted of
6 coloured birds and one pile. The 6 coloured were all of the
light type, the character showing in the chicks in the down as
well as in the mature plumage. In both these broods there were
recessive piles, 3 in F, III, lin F,V. Therefore the parents
must have been carrying the recessive white or pile, and in the
pavents of F. III there was segregation not only between colour
and pile, but between light and dark colour. The brood F, VI
was the offspring of F, V 12 pileand F, VIII 7 darksilky. In
this brood theve were five coloured birds and four pile. Of the
former two were accidentally killed during: my absence from
home, so that I was unable to examine their characters, one was
a male, concerning which there is no entry in my notes to show
whether it was of the light or dark type, one was a dark female
with black head, and one was a female of the light-coloured type.
It is evident therefore that in this case segregation of light and
dark took place in the coloured male parent when mated with
the recessive pile—in other words, the dark type was carrying
the recessive light colour, as well as the recessive pile. On the
other hand, F, VII and F, X are broods in which a light-coloured
parent (female in one case, male in the other) mated with pile
produced only the light type in the coloured offspring.

We may conclude, then, that in the dark type segregation may
take place, while the lighter type is recessive and breeds true.
Whether the original black-ved cock of this cross was heterozygous
with respect to darkness of colour, or the lighter type arose de
novo from the dark type, is another question. Mendelians would,
of course, assume that the light type was a unit or factor which
always existed, as they do not admit the origin of new factors,
but their assumption seems to me unjustified. The facts as
described in my experiments may be explained on Mendelian
principles by the hypothesis of epistatic and hypostatic factors as
applied by Bateson to the colours of mice*. We may assume
that in the dark-coloured type there is a factor, D, which causes
the development of the dark colour and is, absent in the light
type. If we write D for the factor which determines the darker
colour and © for that which determines colour as distinguished
from absence of colour, using as usual the small letters for the
absence of these factors, then a dark-coloured bird carrying both
light colour and white (or pile) may be represented as

DdCe and the pile as ddce.

The gametes of the dark bird will be
DG, dG, De, de, while those of the recessive will be all de.

The fertilizations will be

IDC deed. cccseiostinsate dark-coloured offspring.
GN OAC(G ape tae ae Ee are light im at
Died Pcl. 562 eee. recessive white or pile.
Merde; ey ies nde: Se . aa

* “Mendel’s Principles of Heredity,’ 1909, p. 78.

182 MR. J. 1. CUNNINGHAM ON RESULTS OF A

All this is, of course, regardless of the hypothesis that the
colour in itself is due to two separate factors, X & Y. The third

of the above fertilizations implies that the factor for darkness.

might be carried by the recessive without the factor for colour.
If so, light-coloured birds mated with pile might produce dark-
coloured offspring. I have not observed such cases in my results.
On the other hand, another possibility is perhaps worth con-
sidering—namely, that the factor determining the darker colour
is only quantitatively different from that causing the lighter.
The darker colour may be due to an additional colour-factor
segregating separately. Thus, if we substitute Ce for Dd in the
above scheme, we shall have as the ote of a dark-coloured
bird mated with recessive, CCec, 2 Cece, ecec—in other words,
dark, light, and recessive in the proportions 1, 2, 1 instead of
1,1,2. Which of these agrees better with the actual results of
experiment, the numbers reared by me were too small to decide.

Pigmentation of Skin.

In my previous paper I described the occurrence of a certain
amount of pigmentation in the skin and internal membranes of
one of the birds otherwise recessive for this character— namely,
F,If 23d. In subsequent generations this impurity of the
recessives with respect to this character was constantly observed,
the amount of pigmentation varying in different individuals, in
some cases being greater than in the F, generation. Thus, in
F,IV 22, which died on October 14th shen about 5 weeks old,
the skin of the abdomen was dark, almost black, lighter over
sternum, dark again over crop and ventral side of neck, There
was a good deal of pigment round the eye and on lower eyelid.
The shoulder-joint, elbow-joint, and wrist-joint appeared almost
black on dorsal side, and the bones of the leg and wing were
dark. Slight pigment was visible in the peritoneum over the
gizzard, but none in or on the ovary.

In FI, of six chicks five were recessive in skin-pigmentation
and one pigmented. In al] the recessives there was pigment in
the skin of the abdomen and round the eye. F,IV 19 and
FIV 29 were killed in September, 1913, and I skinned them
after cutting the feathers short to show the surface of the skin,
but the dry skins now do not show the pigment distinctly, as it
was in the fresh state. JI noted from the examination of
F.IV 29 immediately after death that the abdomen was quite
dark, and that the pigment was not only in the skin and
peritoneum but in the connective tissue of the abdominal wall.
There was pigment over the gizzard, the oviduct on the left and
the vestigial right oviduct were conspicuously black, and also the
ureters appeared as black streaks.

In the later generations all the recessives had pigment in the
abdomen. It is clear that in this character, as in that of
plumage-colour, pure recessives and therefore complete segrega-
tion did not occur, and also that although individuals varied the

MENDELIAN EXPERIMENT ON FOWLS. 183

amount of skin-pigment in some cases increased. Conversely, it
was noticed in some cases that the amount of pigment in
heterozygous individuals was diminished—for example, F, VII 1
was pigmented like all the rest of that brood, but when killed
and plucked in August the following year the breast and back
were scarcely darker than in a recessive. Both the parents of
F, were pigmented, but if one of them were a heterozygous
dominant, half the offspring would also be heterozygous, and this
was probably the condition “ot Valli

Structural Characters.

The three characters hitherto considered are colour-characters
due to pigment in the plumage or in the skin and connective
tissue. I have now briefly to review the structural characters, of
which there are five—namely, silky plumage versus normal, comb,
crest, feathering of legs, double hallux.

In the 10 eds of E, generation there were three with silky
plumage and seven normal, in the first brood the numbers were
two silky to six normal, the proper propor tion of one recessive to
three dominant. In F, 23 birds altogether, all were normal,
although one parent of some of the br oods was silky—the other
must therefore have been pure dominant. In later generations
recessive silkies were again obtained. In two silkies of LAYS 1
thought the silky character was not perfect, the proximal ‘parts
of the lar ge wing-feathers being as in normal plumage, and only
the ends Noose-barbed But I was not able to make a careful
comparison with the pure silky, so that I have no distinct
evidence of imperfect segregation in this character, and in any
ease 1t would be difficult to be certain about a slight degree of
the normal character in silky plumage.

In my previous paper I suggested that the form of the
posterior end of the rose-comb was connected with the presence
or absence of the crest, and accordingly these two characters may
here be considered together. J stated in the previous paper that
when the crest is present and large the posterior end of the
rose-comb is truncated and trifid. EF, III 11 ¢ seemed to con-
tradict this rule, for it had a vose-comb pointed behind toeether
with a crest. But in this case the crest was slight, and the point
behind was not so perfect as in cases where the erest is absent.
F,IV 12 was another similar case, the rose-comb being pointed
behind and the crest very slight. In subsequent generations
many cases occurred in which “the crest was absent, and these
must, of course, be regarded as pure recessives.

But the crest, when present, showed great variation in size
and dev elopment, noted by me as very slight, moderate, and full.
It may be assumed by Mendelians that the ‘full crests are pure
dominants and the lower degrees are variations in heterozygotes.
I was unable to investigate “this by breeding specially the indi-
viduals with minimum snort of crest, either with one another
or with those without crest (vecessives). The great variation in

184 MR. J. 1. CUNNINGHAM ON RESULIS OF A

size of crest is, however, « fact worth recording, and is anything
but simple dominance in the heterozygote. The single combs I
obtamed appeared to be pure recessives, but there were scarcely
any singles in the later generations.

Similar remarks apply to leg-feathering. There are many
cases of total absence, and these may be regarded as pure
recessives. But, again, there are individual variations, such as
very slight, shght, Sagi enemie. and complete. There ave also three
cases, F Ways da). IME AQ" arin FIV 16, in which there was only
an infinitesimal trace of featheri ing on the legs. The parents of
the last both had “clean ” legs, 7. e. no leg-feathering at all, and
all the others of F, IV were destitute of feathers on the legs.
The case is interesting as showing that a very minute degree
of a dominant character may appear in the offspring of two
recessives. In previous experiments with Japanese Long-tailed
fowls I noticed a similar minute trace of feathers in chicks of
this breed, which normally has no feathers on the legs. These
facts tend to support the view that minute degrees of a character
may occur in individuals which are not heterozygous, and that
such a character as complete leg-feathering is not necessarily an
indivisible unit.

The irregularity of dominance in the double hallux was
mentioned in my previous paper. The later generations show
that the heredity of this character is so irregular that it is
impossible to distinguish the recessive from the heterozygote.
The parents of F. Eells for example, both had normal toes on both
feet, and of the ‘eleven chicks ten had normal toes and one had
the double hallux on both feet. It is evident that one or both of
the parents was heterozygous for this character with the normal
character dominant. If the chick with double hallux is a
recessive, both of the parents must have been heterozygous, and
in that case the number of recessives should have been 1:3
instead of 1: 10.

The parent of F, [and of F, IT all had normal toes, yet in the
former brood two out of six had the double hallux, in the latter
all seven had normal toes. ‘The female parent of F, VI had a
double hallux on both feet, the male parent had normal toes, and
six of the seven surviving chicks had a double hallux on both
feet, but in one of them the hallux was only slightly cleft, and
in the 7th the hallux was double only on the left foot. This
case would agree with theory if the double hallux in the one
parent was a pure dominant and the normal feet in the other
pure recessive. In other cases it is impossible to tell whether a
normal is pure recessive or heterozygous. In several cases both
parents with normal toes give chicks all normal,e.g. F, VIII,
BMV COR VG Ee s yensky Sut, the parents of which were both
from F, VILL. produced three chicks with normal toes to four with
the double hallux. It would seem not merely that the normal
may be dominant, but that segregation sometimes occurs and
sometimes does not.

185

NDELIAN EXPERIMENT ON FOWLS.

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J TS OF A

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— Sey

GHAM ON RESULTS OF A

. CUNNIN

MR. J. 1

188

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189

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191

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| quoo Jo yurod $4sai0 4yysg | “yueso1g ‘ajaj dog a[qnocy yr *paquiod ‘asoy | “[BULO Ny ‘pamopog =“ & 9

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‘ayy. SOT TIT WX peanoyjoo P | Tq squerg “XT a OVAL

MR. J. T. CUNNINGHAM ON RESULTS OF A

192

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“4SBolq TO UMOIG YOUTT

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193

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MR. J. T. CUNNINGHAM ON RESULTS OF A

194

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MR. J. T. CUNNINGHAM ON RESULTS OF A

196

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|
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197

MENDELIAN EXPERIMENT ON FOWLS:

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MR. J. 1. CUNNINGHAM ON RESULT

198

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199

MENDELIAN EXPERIMENT ON FOWLS.

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OT “heyy petoyeH
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MR. J. T. CUNNINGHAM ON RESULTS OF A

200

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201.

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Proc. Zoou, Soc.,—1919, No. XIV.

202 RESULTS OF A MENDELIAN EXPERIMENT ON FOWLS.

EXPLANATION OF PLATE Tf.

Fig. 1. Ventral surface of hen produced from cross between silky hen and black-red
Bankiva cock, showing “pile” coloration. The dorsal surface is white
without any of the reddish brown colour. The hen represented was
F; IL 3 & in the record of the pedigree, z.¢., a hen of the second brood of
the 5th generation. It was killed Dec. 12,1915, when 1 year and 8 months
old.

Fig. 2. Dorsal surface of cock from the same cross, showing the ‘‘ pile” coloration
in the male. The colour is very slight compared with that of the female,
and consists of a slight yellow tinge across the Joins and on the upper
wing-coverts. The rest of the body is pure white. The specimen was
¥, V 12 in the pedigree, é.¢., a cock of the fifth brood of the 6th genera-
tion. It was killed on Dec. 31, 1915, when 7 mouths old.

Ke Zo So OIG. IE/AINIDIEIR, Pl, Ml.

Photo. D. Seth-Smith, F.Z.S.
POSTERIOR PORTION OF TONGUE OF TAKIN (nat. size),

Gale, Scas and Danielsson, utd

ON THE ANATOMY OF THE TAKIN. 203

13. Some Points in the Anatomy of the Takin (Budorcas
tavicolor whitei). Based on the examination of a
specimen in the Gardens of the Zoological Society of
London. By Miss Karaneen F. LAnper, M.8e.,
I’.Z.8. (Hon. Acting Prosector to the Society).

[Received May 13, 1919: Read May 27, 1919. ]
(Plate I.* and Text-figures 1-7.)

The animal in question was a male from North-west Bhutan,
which was presented to the Zoological Society of London on
June 22nd, 1909, and died in the Gardens on May 7th, 1918.

Unfortunately at that time it was impossible for any full
examination of the anatomy to be made; a few notes were taken
of the parts too large for preservation and some of the smaller
organs were preserved in formalin. The account here given is,
therefore, very incomplete, but may serve for comparison when
next a Takin is available for examination.

Hodgson, when giving the first description of this animal,
considered that its nearest affinity was with the Gnus, but that
it would probably be placed between the Gnus and Musk Oxen in
a classificatory scheme. Matschie created the group Ovibovine
for Budorcas and Ovibos as a result of examination of the
external characters, skull and metacarpus, and these two animals
have usually been regarded as closely allied. Dr, Chalmers
Mitchell has pointed out that they have in common the curious
formation of the naso-frontal suture. In them a triangular
process of the frontal fits deeply between the triangular proximal
ends of the nasal bones, whereas in the majority of Ruminants
the nasal bones project into the frontal area without divarication.

The Goral and Serow (Nemorhedus goral and Capricornis
bubalinus) have also been grouped with the Chamois (Rupricapra
tragus) and RKocky-Mountain Goat (Zaploceros montanus) to form
the Rupricaprine Section of the Bovide, and in some of the skull
characters the Goral approaches the Takin as much as anything.

So far as I am aware no account has yet been given of the,
anatomy of the soft parts of the Rupricaprine section, Various
anatomists have published accounts of the anatomy of the other
Ungulata with which Budorcas has at one time or another been
compared, In particular, the papers by Lonnberg on Ovibos and
Connochetes have been consulted for comparative purposes,

Mr. R. 1. Pocock recorded the external characters of the Takin
described in this paper, both before and after death. In order to
render this account as complete as possible I quote verbatim his
observations published in the ‘ Annals and Magazine of Natural
History ’ (6) :—

“The rhinarium is continued inferiorly to the edge of the

* For explanation of the Plate see pp. 204-6.

204 MISS K. F. LANDER ON THE

upper lip as a narrow mesially grooved strip, which is longer than
in Vemorhedus owing to the greater depth of the upper lip.
Laterally an area of naked skin, narrower than in Vemorhedus,
is continued with a bold curve beneath the widely expanded
nostrils, and curving round their posterior extremities passes into
the dorsal portion of the rhinarium, which is much shorter from
before backwards than in Vemorhedus, being considerably more
overgrown with hair.

“The feeé resemble in essential particulars those of the dried
example figured and described in 1910, except that on the fore
foot there is no trace of the transverse ridge of integument just
where the hair of the pastern ceases in the interungual space.
There is no trace of definite pedal gland, although the hair at the
bottom of the interdigital depression in front is stuck together
with secretion indicating activity of the skin at that spot. The
hind foot is like the fore foot.

“There is no trace of preorbital gland or of inguinal glands in
the ordinary sense of that term; but the two mammez on each
side, set as far out from the middle line as the outer edge of the
scrotum, are close together, one in front of the other, in the centre

of a distinct swelling like a small udder. When the skin is cut
away, this swelling is seen to be caused by a blackish glandular
mass like a small bunch of grapes, and blackish secretion could
be squeezed through a single pore on the posterior teat with the
use of considerable pressure. This unusual condition of the
mammary gland in the male is worth putting on record, although,
pending the examination of other specimens of Budorcas, it must
be regarded, I think, as pathological in one individual.

“The penis is provided with a pendulous prepuce, three inches
long, rising from the abdomen six inches in front of the scrotum.
Just within the orifice of the prepuce the skin is highly glandular
and overgrown with long hairs, which protrude from the aperture
to form a tuft three or four inches long. The glans penis is
apically attenuated and provided with a straight, moderately
stout, urethral prolongation projecting some little way beyond
the tip of the glans. Except for the greater elongation of the
free portion of the urethral canal, the glans penis is very like that
of Nemorhedus.”

Mr. Pocock points out that some marked differences may be
found in these respects between Budorcas and Ovibos, particularly
in the strong development of the rhinarium in the former and
its great reduction in the latter, and in the prolongation of the
urethral canal beyond an attenuated glans in the penis of the
Takin, while in the Musk-ox the canal is not produced beyond
the glans, which is blunt at its end.

In the presence of a preorbital gland and hairs between the
hoofs Ovibos differs from Budorcas, as also in the arrangement
of the four mamme, the absence of a protruding tuft of hairs
from the prepuce, and the presence of longitudinal ridges in its
cavity.

ANATOMY OF THE TAKIN. 205

The tongue of this Takin measures 270 mm. in length, and in
front, where it is broadly rounded, 60 mm. in breadth, narrowing
to 50 mm. in the middle and expanding again posteriorly to a
width of 65 mm. Its dimensions thus correspond almost exactly
with those of the adult male Musk-ox described by Lonnberg.
160 mm. from the anterior border is a- transverse depression,
slightly convex anteriorly, behind which the tongue shows the
usual convexity of the posterior part of this organ. The position
of the transverse groove, more than half way back on the tongue
surface, is a point in which the Takin differs from the Musk-ox,
agreeing with the Gnu.

On the anterior flat portion filiform and fungiform papille are
found. These extend on to the under surface for a distance of
5 mm. back from the tip, but over the lateral border, 40 mm.
back, they cover 15 mm. of the lower aspect. In this respect
Budorcas resembles the Musk-ox, Gnu, Reindeer, Sheep, and
Goat, but differs from Bos. As in Ovibos and Connocheetes, the
fungiform papille are very numerous on the under surface.

The filiform papille are small and flat in the centre, only just
distinguishable to the naked eye and very closely set. Towards
the sides they become more filiform and rather less flattened, and
the same process of alteration in form takes place from before
backwards, those at the sides being always more filiform than the
central ones at the same transverse level.

80 mm. from the tip the lateral papille are distinctly elongated,
tapering and recurved, a similar elongation of the central ones
being found 40 mm. further back. In the transverse groove the
papille are 2mm. in length, and 1 mm. broad at the base, tapering
to a fine point. In the greater narrowness and pointedness of
the filiform papille the Takin is more like Bos than Ovis, Ovibos,
or Antilope cervicapra.

The fungiform papille are small and scattered in the anterior
central region; they have to be looked for here, while on the
sides and at the tip they are the most noticeable structures on
the tongue surface and are set about 3 mm. apart. In the
middle of the tongue they become more prominent, but there are
not many in the region lying between the groove and a line
7 mm. from the anterior border. At the sides and back of the
central region these papille are about 3 a millimetre in diameter.
Around the groove they are more numerous and measure 1 mm.
across. In Bos, Ovis, Capra,and Capreolus fungiform papille are
entirely absent on the central part of the anterior half of the
tongue. The presence of small scattered fungiform papille in
this region is characteristic of Ovibos, Connochetes, and Rangifer.

In the posterior part of the tongue behind the sulcus the
filiform papille are long and recurved in the central region,
becoming gradually shorter, thinner, and more hairlike laterally
until on the lateral margin they are merely raised dots.
Proceeding backwards the same transition takes place, but rather
more rapidly. The fungiform papille increase in size as they
pass backwards, becoming 2 mm, in diameter at-the back,

206 MISS K. F. LANDER ON THE

A central strip, 10 mm.in breadth, is sharply differentiated
from the rest of this part by being entirely devoid of fungiform
papillae, and by the character of the filiform papille. It gives
the impression of being bordered by an irregular row of fungiforms.
The filiform papille only maintain their long pointed character
for a distance of about 20 mm. back from the transverse groove,
becoming progressively smaller and more scale-like for a further
35 mm. back. Behind this point they again increase in size, but
retain their scale-like form; some are as large as 3 mm. in
diameter, and in form thev are either circular or very slightly
pointed ; they are very hard and horny. At the lateral margins
of the strip these pass rather abruptly inte the long pointed form
typical of the rest of the central region of the posterior half of
the tongue. (Cf. Plate I.)

The well-defined central strip of Baudorcas is not described for
any other animal, but in Rangifer and Capreolus the centre of
the back of the tongue is said to be destitute of fungiform papille.
In Ovibos only the anterior one-third of this central region is
devoid of fungiform papille. This part of the tongue seems to
be more hike that of Bos than is the tongue of the Musk ox.

Papille are present on the lateral aspect of the posterior part
of the tongue in its anterior half, @. ¢., as far back as the row of
cireumvallate papille. Papille are stated to be present on the
lateral aspect of this part in the Musk-ox and some of the
Cervidee, but not in Connochetes, Ovis, or Capra.

It is in some eases very difficult to determine whether a given
papilla on the back of the tongue is of the fungiform or an the
circumvallate variety, as Linnberg found in Madoqua saltiana.
Counting the doubtful ones as circumvallate there are twenty-
three of these on the left side and twenty on the right, arranged
in four irregular rows. The fourth or outermost row appears
only at the anterior end of the group. About fourteen of these
are almost certainly circumvallate, and the number is thus
comparable with that found in Bos (10-17) and Ovibos (12-15),
but if the greater number be counted the Takin approaches the
Sheep (18- “D), Capra (16-17), or Connocheetes (20) in this respect.
The presence of four rows brings Budorcas into line with the
Sheep ; Ovibos, Connochetes, Bos,and Rangifer present only two
rows. The grouping into rows is, however, very ill-defined in the
Takin, and the papille are very variable in size. They form the
usual V-shape, the arms of the V reaching forwards to a point
about half-way back along the posterior portion of the tongue.

Along the side of the attached portion of the organ runs a low
fold of mucous membrane extending forwards to a distance of
80 mm. from the tip. It bears several ‘long pointed denticulations,
which look upwards and backwards.

The hyoid bone is well-developed. The body is square, 30 mm.
in size, bearing a small ridge on its anterior surface. The distance
between the facets for the ceratohyals is 30 mm. The thyrohyal
is 70 mm. in length and runs horizontally outwards * to meet

* Note: For the purposes of this description the larynx is regarded as being
vertical in its long axis, as when the head of the animal is held high, and as figured.

ANATOMY OF THE TAKIN. 207

the superior cornu of the thyroid cartilage. The ceratohyal is
40 mm. long, and runs forwards, outwards, and upwards to
articulate with the epihyal, 25 mm. in length. ‘This runs
backwards, upwards, and outwards and bears the stylohyal,
which passes backwards, eutwards, and very slightly upwards for

Text-figure 1.

Lateral Aspect of Hyoid and Larynx of Takin,

100 mm. before bifurcating widely into two terminal processes.
The superior of these continues on the curve of the bone and
ends bluntly truncated ; the inferior is short and bluntly rounded
and projects slightly forwards.

The bone is thus very like that of os, except for the

208 MISS K. F, LANDER ON THE

preponderance of the ceratohyal over the epihyal in length.
The hyoid of Ovibos is not described by Lonnberg.

The laryna, like that of the Musk-ox, is very elongated, owing
to the expansion of the thyroid cartilage. Its maximum length
is 200 mm. and its maximum transverse vertical diameter 105 mm.
The expansion consists of a hollowing out and backward extension
of the anterior part of the thyroid cartilage, which measures .
180 mm. along its convexity, while the posterior border, from
superior to inferior cornu, is only 60 mm. long. The amount of
extension may be judged from the fact that the distance between
the lowest (7.e., most posterior) point of the thyroid to the
insertion of the posterior cornu is 105 mm., while from the same
point to the antero-inferior surface on a vertical plane is 70 mm.

The superior cornu is 25 mim. in length, and runs obliquely
upwards and backwards, thus presenting an intermediate
eondition between the superior cornu of Ovibos, which passes
nearly vertically upwards at right angles to the long axis of the
larynx, and that of Bos or Capra, which runs parallel with that
axis. (Text-fig. 1.)

Text-figure 2.

Arytenoid Cartilage of Takin.

It is thus clear that the expansion of the thyroid cartilage
met with in the Musk-ox, and approached in the larynx of the
Blackbuck and Saiga, is carried very much further in the Takin.
In the latter the bulbous extremity of the expansion is carried
back far beyond the tip of the posterior cornu. In Ovibos the
extremity is on a level with the tip of the cornu and the
cartilage on the whole has a squarish appearance ; in Budorcas
it is about three times as long as broad. In Capra, Cervus, and
Capreolus the length of the dorsal part of the cricoid is greater
than the length of the thyroid; in Ovibos the cricoid length is
72 per cent. of the thyroid length; in Connochetes 80 per cent. ;
in Bos they are about equal, but in the Takin the percentage is
only 33.

The inferior cornu of the thyroid in Budorcas is very much

ANATOMY OF THE TAKIN. 209

shorter and less curved than in Ovibos ; it measures 50 mm. and
is only slightly arched.

The cricoid is of the usual form ; the expanded posterior plate
is 60 mm. in depth and 55 mm. across, It presents a median
posterior keel nearly 10 mm. in height. Owing to the extension
of the anterior part of the thyroid the ring is very obliquely
sloped, the anterior part lying at a lower level than the lowest
part of the posterior part. (‘l'ext-fig. 1.)

The arytenoids are massive, 85 mm. in total length. The free
apex is expanded into a thin, curved, and fenestrated plate. The
vocal process is a rounded knob-like eminence. The muscular
process is a long stout bar extending downwards and forwards
for 40 mm. from the body. It is thus only 5 mm. shorter than
the expanded free portion. (Text-fig. 2.)

Text-figure 3.

(a) Anterior Aspect of Lungs. (0) Transverse Section of Trachea.

The epiglottis is bluntly rounded at the apex, 50x 35 mm.
in size. Only about one-half of it is free. The surface is pitted.
It is therefore similar to the epiglottis of the Gnu and Musk-ox.

The tracheal rings are irregularly imbricate. Their posterior
ends are separated by a distance of 30 mm. and there is thus no
posterior keel, in contrast to the trachea of Bos and Connochetes.
There are 15 rings above the eparterial bronchus and then
4 succeeding ones before the bifurcation. The absence of keel
gives the trachea of Budorcas a resemblance to that of Ovibos,
but there is no dorso-ventral flattening. The ends of the rings
in Budorcas project dorsally, whereas those of Ovibos appear to

210 MISS K. F, LANDER ON TUE

bend over, the terminations facing each other. The shape of the
cavity 1s practically circular in the former animal and oval or
kidney-shaped in the latter. (Text-fig. 3 0.)

The Jungs are pyramidal in shape. The left lung shows no
fissures whatever ; the right lung has a well-marked upper lobe,
very much smaller than the lower, but the fissure marking this
off is incomplete at the upper border. (Text-fig. 3 a.)

There is a well-developed azygos lobe, measuring 130 x 80 mm.

The left lung of Ovibos is described as remarkable among the
Ruminants for its simplicity, the upper lobe sitting with a broad
base on the lower; the left lung of Budorcas is still more
strikingly simple. The right lung is also markedly more simple
than in other Ruminants.

ALIMENTARY SYSTEM,

The paunch consists of the usual two unequal sacs, lined by a
mucous membrane presenting flattened, tongue-shaped papille,
whose maximum length is 15 mm. and breadth 4mm. These
gradually become confluent, presenting a moss-like surface, and
run into a reticulum, the cells of which measure about 20 mm.
in diameter and are bounded by walls as high as 4 mm. or 5 mm.
Secondary and tertiary ridges are also present.

The paunch is thus very like that of Ovibos and distinctly
different from that of Connochetes, in which the papille are
conical with blunt ends and there is a sharp dividing line between
paunch and reticulum ; the saes of the paunch also are-subequal
in size in Connochetes.

Sixty-one folds may be counted in the psalterium; many of
them are mere ridges; twenty-one of them form high folds.
This is the same number as that named by Lénnberg for Ovibos.

In the abomasum seventeen folds were found, the animal thus
agreeing roughly with Bos (14-16) and Antilope cervicapra (19),
but having more folds than Ovis or Capra (13-16) or
Connochetes (12-13), and fewer than Quvibos.

The small intestine measures 103 ft. in length, the large gut
38 ft., while the cecum measures 2 ft. 3 ins. The colon is of a
perfectly simple tubular type, with a uniform muscular coat, no
sacculations and no appendices epiploices.

The small intestine is thus 2:7 times the large in length, a
figure exactly corresponding to that given for the Blackbuck, and
comparable with that of Ovibos (2°6) and various antelopes (2:3
to 2°7). In Ovis and Capra the small intestine is fully three
times as long as the large, and in Connochetes the figure is nearer
four, as it is in Bos.

Unfortunately no observations could be made on the grouping
of the intestinal loops.

The liver is divided into right and left lobes, the former
occupying three-fourths of the whole area of the anterior surface.
This lobe is squarish in shape, with rounded corners, and is
13 inches in breadth, while the oval left lobe measures 3 inches.

ANATOMY OF THE TAKIN. 211

These proportions present a marked contrast to those of Ovibos,
in which the right and left lobes are almost equal in size. In
Gos and Ovis the right lobe is somewhat larger than the left.
(Text-fig. 4a.)

Text-figure 4.

(a) Anterior, (6) Pssterior Aspect of Liver.

The Spigelian lobe is practically undifferentiated, as in
Connochetes ; it can be distinguished in the livers of the other
animals mentioned for comparison,

The lobus caudatus is club-shaped, with its narrow end
reaching out to the right margin.

The quadrate lobe is square in shape, 44 inches in breadth
(text-fig. 4).

The gall-bladder is definitely bilocular and lies in the fossa

212 MISS K. F. LANDER ON THE

between the quadrate and right lateral lobes, extending for some
distance below the inferior margin. The bile-duct is highly
valved. The gall-bladder thus seems distinctly different in form
from that of Ovibos; it is also situated rather nearer to the right
margin than in the Musk-ox. On the whole it is more like that
of Los or Connoshetes.

The testis is oval, or, with the epididymis, spindle-shaped.
The total length of the testicle is 85 mm., of which 21 mm.
represents globus minor and 8 mm. globus major. The width is
40 mm. and thickness 30 mm. ‘Ihe head of the epididymis
enwraps the anterior free border of the gland for the upper halt
of its length. It is only connected with the tunica albuginea of
the testis around its periphery; the rest of the expanded head
can be separated from the testis by tearing the lax areolar tissue
between them. The organ is enclosed in a thick tunica vaginalis

Text-figure 5.

Testis of Takin.

of the usual type, reflected on to the epididymis along the whole
length of its head and body. ‘The globus minor forms a free
bulbous projection, 20 mm. in breadth, 15 mm. in depth and
thickness. ‘The epididymis (in formalin) is salmon-pink in colour,
marked by numerous very tortuous superficial veins. The tunica
albuginea is bluish white; the gland shells out of this capsule
with the greatest ease, being separated from it by a number of
large tortuous veins which run mainly in a longitudinal direction
and make marked grooves on the surface of the soft brown testis.
The gland and capsule are only connected by a very small strand
at the upper pole. (Text-fig. 5.)

The vas deferens ascends beside the body of the epididymis,
accompanied by two large veins. Above the caput it is joined
by a large pampiniform plexus, forming a spermatic cord 17 mm.
in thickness.

ANATOMY OF THE TAKIN. 213

The brain is 119 mm. in total length, of which 104 mm.
represent the length of the cerebral hemispheres. Its breadth
is 91 mm., and height 58 mm. It is thus distinctly small when
compared with the size of the animal. 1

The olfactory bulbs are not present, and there is a wide saw-
cut across the frontal region; otherwise it is in excellent
condition. It has been preserved in formalin for twelve months.

Text-figure 6.

A.P.S.= Anterior Perforated Spot. O.N.=Optic Nerve.

Dioptograph outlines of Lateral and Superior Aspects of Brain.

The optic lobes are completely concealed and the projecting
occipital poles of the hemisphere rest on the anterior aspect of
the cerebellum, which is sloped to receive them.

The hemispheres are well fissured, the sulci being deep and
complicated and almost all of them having numerous hidden sulci
on their walls which run from the surface perpendicularly inwards
to the floor of the main fissure,

914 MISS K. F. LANDER ON THE

The “ pseudo-Sylvian ” fissure extends on the lateral aspect for
a distance of 30 mm. It is 16 mm. deep on the infero-lateral
margin and conceals a good deal of operculated cortex marked by
hidden perpendicular sulci, converging towards a central point
near the highest part of the fissure. 8 mm.and 15 mm. from its
upper extremity the fissure gives off small branches which run
backwards and forwards respectively.

The two terminal sulci are deep and well-marked. The
anterior one commences on the inferior aspect of the frontal pole
and runs downwards and backwards for 15 min. and then turns
backwards to pursue a course parallel with the rhinal fissure. It
is 5mm. deep. The posterior terminal sulcus is only 15 mm. long.
Between these and the rhinal fissure is a gyrus of 6 mm. bre adth,

The notch on the rhinal fissure, said to represent the feline
‘“* pseudo Sylvian,” lies 6 mm. behind the confluence of the
anterior and posterior terminal sulci and is 10 mm. long.

There is a typical Ungulate suprasylvian arch, 16 mm. deep,
with a horizontal ramus 30 mm. long and vertical rami 13 mm.
in length. The gyrus surrounding the ‘‘ Sylvian” fissure is in
most places over 12 mm. in width, but the termination of the
fissure reduces it to 5 mm.

The posterior branch runs back to the occipital pole. It is
prolonged forwards beyond the posterior vertical limb of the
suprasylvian, reaching to within 5 mm. of the “ Sylvian ” fissure.
A shallow groove passes over the intervening gyrus, connecting
the two sulci.

The sulcus obliquus of Holl is represented by a triradiate sulcus
of which one branch runs forwards 15 mm. above the rhinal
fissure to a distance of 7 mm. from the “Sylvian”; another
upwards to 7 mm. from the posterior branch of the suprasylvian ;
while the third passes backwards and downwards to a point
5 mm. from the hind end of the rhinal on the inferior aspect.
Also belonging to this complex is a small isolated sulcus, .7 mm.
long, lying between the anterior ramus of the triradiate and the
forward prolongation of the posterior branch of the suprasylvian.

A small curved sulcus perpendicularis, 8 mm. long, lies between
the “Sylvian” and suprasylvian.

The diagonal sulcus is 22 mm. long and lies in the usual
position round the end of the anterior vertical limb of the
suprasylvian suleus. Its extremities both curve upwards. A
gyrus of 4 mm. width separates the two sulci. Its accessory
sulcus appears to be absent.

The orbital sulcus commences on the upper aspect of the frontal
pole, 7 mm. from the supero-inesial margin. It first forms a
curve, convex laterally, and then turns over the anterior margin
and runs backwards to join the anterior terminal suleus at its
angle. The curved part is wide, but rather shallow, presenting
a trench-like appearance; the hinder part is 10 mm. deep. A
short cross-branch connects it with the front end of the rhinal.

From the junction of the anterior vertical and horizontal limbs
of the suprasylvian a well-marked transverse fissure runs straight

ANATOMY OF THE TAKIN. 215

up to the supero-mesial margin. Into this, half way up, runs
the coronal, which passes straight backwards from the tip of the
frontal pole, a distance of 45 mm.

The calcarine, intercalcary, and cruciate sulci form one
continuous furrow, cutting the supero-mesial margin 40 mm.
behind the frontal pole and running downwards and forwards on
the lateral aspect for 16 mm. Its end is separated from the
coronal sulcus by a gyrus of 5 mm. breadth.

The lateral sulcus starts 5 mm. behind the transverse sulcus as
a short straight sulcus, 15 mm. long. The hinder end of this is
embraced by the widely bifurcated anterior end of another longi-
tudinal sulcus which pursues an undulating course backwards for
40 mm., ending by bifurcating widely 9 mm. in front of the
occipital pole. This gives off four short branches in its anterior

Text-figure 7.

Dioptograph outline of Mesial Aspect of Brain.

$.C.Q.=Superior Corpus Quadrigeminum. ©O.C.=Optic Chiasma.

half, two up and two down. ‘The latter reach to within 5 mm.
of the suprasylvian ; elsewhere the gyrus separating the two sulci
is 13 mm. broad. The lateral sulcus, as in most Ungulata, is
obliquely placed, so that it approaches the mesial margin more
nearly at its anterior end.

The ectolateral sulcus lies between and parallel with the
preceding and the posterior branch of the suprasylvian. It is
20 mm. long and its posterior bifurcation forms a vertical sulcus,
25 mm. in length, running parallel with the hind margin of the
hemisphere. Across the anterior end of the ectolateral sulcus
lies a vertical sulcus 20 mm. long separated from it by 3 mm.

A small curved entolateral sulcus, concave mesially, 15 mm, in
length from tip to tip, lies between the lateral sulcus and the

216 MISS K. F. LANDER ON ‘THE

supero-mesial margin. In front of this, lying mainly on the
mesial aspect, is an oblique straight sulcus, 25 mm. in length.

The rostral sulci are well marked; in front of the cruciate-like
sulcus lies an H-shaped sulcus, and parallel with the anterior
margin of the brain runs a curved sulcus 30 mm. long and 5 mm.
deep.

Sulei subcingulti are represented by a longitudinal sulcus,
broken in the middle, placed between the intevcalary sulcus and
the corpus callosum. The anterior part, 25 mm. long, curves
over the genu and bifurcates widely, the lower limb continuing
the curve, the upper running upwards between the two genual
sulci. Behind the splenium lies a vertical sulcus which is a
branch of the calearine. This latter is 28 mm. long and 10 mm.
deep. The intercalary is 15 mm. deep at its commencement ;
it arises from the calcarine about the mid-point of the latter.

The rhinal fissure is well marked posteriorly ; in its depths is
a good deal of buried cortex. It turns round the occipital pole
and runs up nearly to the posterior end of the calearine. The
hippocampal convolution is 5mm. broad anteriorly and below
the “Sylvian” fissure expands into a broad triangular mass,
40 mm. along the base and 25 mm. from base to apex. Near its
posterior angle this is marked by a horizontal sulcus 10 mm. long.
he apex forms a well-marked ‘“ pseudo-temporal lobe,” extending
downwards and forwards over the crus cerebri and optic tract to
the anterior perforated spot.

The supra-callosal gyrus is 8 mm. in breadth and subdivided
by a well-marked small vertical sulcus into a larger outer mass,
or gyrus of Andreas Retzius (hippocampus nudus), 5 mm. in
breadth, and a smaller pyriform dentate fascia, the narrow end
of which lying behind the splenium is 2 mm. wide, the broader
end 5mm. ‘he sulcus limitans is also well marked.

The anterior corpus quadrigeminum measures 15 mm. in length
by 12 mm. in breadth, while the posterior is 7 mm. in antero-
posterior diameter and 9 mm. transversely.

So far as I am aware the brains of the Musk-ox and Gnu have
not been described. For comparison, therefore, two brains from
the Society’s collection have been used, those of a Barbary sheep
and an Anoa. The latter is characterised by Elliot Smith as the
simplest and most generalised Ox-brain. The three brains are
very nearly the same size and have been preserved in formalin
for about the same time.

The brain of Budorcas appears much simpler than that of Ovis,
owing chiefly to the absence of the numerous small isolated sulci
or secondary branches of the main sulci which are found in the
sheep. The cruciate upturning of the intercalary sulcus is rather
less marked in Ovis and the transverse sulcus does not meet the
suprasylvian. The arrangement of the orbital is rather different ;
this sulcus does not join the anterior terminal but the rhinal in
Ovis. The Sylvian appears to rise from the rhinal and the
diagonal is distinctly U-shaped, embracing the anterior vertical

ANATOMY OF THE TAKIN. Neff

limb of the suprasylvian suleus. These last two features are not
constant in the ovine brain, however, and are not seen in the
sheep brain figured in the Catalogue of the Museum of the
Royal College of Surgeons.

Where the brain of the Takin differs from that of the sheep
it agrees with the bovine type represented by the Anoa. The
arrangement of orbital, anterior terminal and rhinal, and the
relative positions of the transverse, coronal and suprasylvian sulci
are almost identical in these two, but the cruciate suleus is even
less marked in the Anoa than in the sheep. The development of
this suleus seems to be partly associated with the position of the
coronal sulcus with reference to the supero-mesial margin. In
the Anoa the gyrus between them is less than 3 mm. in width,
in the sheep 7 mm., but in the Takin 16mm. There is thusroom
for a long ‘cruciate’ sulcuson the lateral aspect in the Takin,
for a very short one only in the sheep, and in the bovine brain
the cruciate upturning is confined to the mesial aspect.

Budoreas difters markedly from the bovine type in the absence
of any approach to the curious “double” form of Sylvian fissure
found in the ox-like brain.

The anterior corpus quadrigeminum is as much larger in the
Takin than in the sheep as it is larger in the sheep than in the
Anoa, but the difference is slight and may be partly due to
flattening consequent upon preservation. The lody is distinctly
triangular in outline in mesial section in the Takin, oval in the
sheep, and more flattened still in the Anoa.

A similar progressive change is seen in the relative position of
the supra-callosal gyrus and splenium. In the Anoa there is
hardly any cortex directly behind the splenium ; the whole mass
hes underneath the corpus callosum; in Qvis the small pointed
extremity of the hippocampus nudus lies in a line with the middle
of the splenium; in Budorcas,as stated, the gyrus is 5 mm. broad
horizontally behind the splenium.

The occipital pole is more pointed and projects rather further
backwards over the cerebellum in the Takin and Anoa than in
the sheep.

It seems hardly advisable to attempt to draw far-reaching
conclusions of systematic importance from the scanty material at
present available, more especially since there are no data on record
of the anatomy of the other members of the Rupricaprine section.

Support is given to those who hold the aflinity of Budorcas
with Ovibos to be a close one; the two animals differ in but few
points of their soft anatomy, and in many cases they share
characters which differentiate them from other ruminant
Artiodactyla.

The suggested relationship between Connocheetes and Budorcas
is not borne out; the resemblances between the two animals are
few and far between and are mostly points of detail and minor
importance.

Between Gos and Ovis, Budorcas seems to hold an intermediate
Proc. Zoou. Soc.-—i919, No. XV. 15

218 ON THE ANATOMY OF THE TAKIN.

position, agreeing now with one and now with the other, but a
definite pronouncement as to the systematic position of the Takin

ong the Artiodactyla would be unjustifiable until many more

observations on the anatomy both of Budoreas and of allied
Ruminants have been placed on record.

BIBLIOGRAPHY.

Lonnpere: On the Soft Anatomy of the Musk-Ox. P.Z.S.
1900, p. 152.

On the Soft Anatomy of Connochetes Gi.
Kongl.. Svensk. Vet.-Akademiens Handlingar,
Band 35, No. 3.

Material for the Study of the Ruminants.
Nova Acta Reg. Soc. Scien. Upsal., Series IIT.
vol. xx.

. Pocock: The Serows, Gorals, and Takins of British India.

Journ. Bombay Natural History Soc. vol. xix.

p. 807; and vol. xxii. p. 317.

Ov Gs On the Specialised Cutaneous Glands of Ruminants.
PEAS: VOLO pssao:
GF mr, On some External Characters of Ruminant
« Artiodactyla. Ann. Mag. Nat. Hist. (9) i.
pp. 136, 140.
7. Hopeson: On the Takin of the Hastern Himalayas. Journ.

Asiatic Soc. Bengal, 1850, p. 65.

. CHALMERS Mircuetit: P. Z.8. 1907, p. 467.

. Matscuin: Sitz.-Ber. Gesells. Naturf. Berlin, 1896, p. 30.

Garrop: On the Anatomy of the Ruminants. P.Z.S8.

eH 08 25

. Murre: On the Anatomy of the Saiga. P. Z. 8. 1870.

. Sisson: Veterinary Anatomy.

. Exuior Suir: Catalogue of Physiological Series: Museum
Roy. Coll. Surgeons, 1902.

Karly notices of the Takin and its varieties are found also :—

. Lypexker: P. Z. 8. 1908, p. 795.

15. He The Field, 1907, vol. 110, p. 887.

16. As The Field, 1908, vol. 111, p. 790.

17. = Cat. Ung. Mammals Brit. Mus. vol. i. pp. 210,
lg WMS

18. THomas: On the Duke of Bedford’s Expedition in N. China.

ASCO Myo

19. Wuire: P. Z. 8. 1908, p. 668.
20. Bartey: Journ. Bombay Nat. Hist. Soc. vol. xvi. 1907,

p. 842, vol. xxi. 1913, p. 1069.
. BentoHam: Rec. Ind. Mus. 1909, p. 249.
. Hume: P. Z.S8. 1887, pp. 483-6.

AUSTRALASIAN ANTARCTIC AND SUB-ANTARCTIC LIFE. 219

EXHIBITIONS AND NOTICES.

February 4th, 1919.

Dr. 8. F. Harmer, F.R.S., Vice-President,
in the Chair.

The Srcrerary read the following reports on the Additions
made to the Society's Menagerie during the months of November
and December, 1918 :—

NOVEMBER.

The registered additions to the Society’s Menagerie during the
month of November were 433 in number. Of these 3 were
acquired by presentation, and 430 were deposited.

The following may be specially mentioned :—

1 Kea Parrot (Westor notabilis), from New Zealand, presented
by Lady Ian Hamilton, on November 27th.

DECEMBER.

The registered additions to the Society’s Menagerie during
the month of Deeember were 58 in number. Of these 45 were
acquired by presentation, 12 were deposited, and 1 was received
in exchange.

The following may be specially mentioned :—

1 Chimpanzee (Anthropopithecus troglodytes), from Sierra Leone,
deposited on December 6th.

A collection of 32 lizards, including 8 Starred Lizards (Agama
stellio), from Salonika, sent by Capt. W. D. Motton and G. H.
Colt, F.R.C.S.

Mr. C. Daviss SHerporn, F.Z.S., exhibited and made remarks
on a letter written in 1693, by Malpighi to Dr. Mathew Faber.

Sir Doveras Mawson gave a lantern exhibition of Australasian
Antarctic and Sub-Antarctic Life, and made the following
remarks :—

The immense area of the Southern Seas supports abundant
marine life which, if not utilized by man directly, is indirectly
converted into useful products, such as Seals and Penguins. The
tameness of these creatures makes them an easy prey to man,
who finds therein remuneration from the marketing of blubber,

220 THE SECRETARY ON ADDITIONS TO THE MENAGERIE,

oils, and skins. This traffic is on the increase, and it is quite
certain therefore that unless the killing of these creatures is
controlled, and regulations enforced for their proper protection,
the species in many instances will rapidly become extinct—a fate
that has already overtaken the valuable animals in many Sub-
Antarctic lands.

Even Macquarie Island, a dependency of Tasmania, has been so
worked by New Zealand sealers that it has also suffered. Never-
theless, it still abounds in most types of Sub-Antarctic life, and
is the most ideal spot in those seas to be retained as a National
Reserve for the protection and propagation of the various species
of Penguins and Seals. This little island, which lies buried in
mist and fog amidst the turmoil of the great rolling seas that
sweep unchecked around the Globe in those latitudes, is one of the
wonder spots of the world, for to this ocean sanctuary flock the
seal and bird life of millions of square miles of the surrounding
waters. It is the great focus of such life in the Autralasian Sub-
Antarctic, and an indescribable attractive foree impels the land-
seeking life in those wide seas towards its shelter. The Penguins
throng the beaches, and it is one of the few havens left to the
great elephant-seals. Seal and bird life, with which the island
still teems, has been greatly restricted as a result of the slaughter
by the sealers. Within five years after the discovery of the
island, the Fur-Seal was almost exterminated, and the species
is now virtually extinct. A species of flightless Parrot is now
non-existent, while the noble King Penguin has dwindled in
numbers until it is now represented only by one small rookery at
Lusitania Bay.

Turning to the Australasian Antarctic shore, the great abun-
dance of seals, penguins, and whales will sooner or later attract
exploiters, and it is well that the responsible Governments should
legislate ahead in the interest of the continuance of the species.

February 18th, 1919.

A. Smrra Woopwarp, Hsq., LL.D., F.R.8., Vice-President,
in the Chair.

The Secretary read the following Report on the Additions
made to the Society’s Menagerie during the month of January
1919 :—

The registered additions to the Society’s Menagerie during the
month of January were 42 in number. Of these 21 were acquired
by presentation, 9 were deposited, 10 were received in exchange,
and 2 were purchased.

The following may be specially mentioned :—

2 Lion Marmosets (Leontocebus rosalie), from §$.K. Brazil,
presented by Sir George Noble, Bt., F.Z.8., on January 25th.

STRUCTURAL CHARACTERS OF THE FELID. DAI

2 Bennett’s Wallabies (Macropus bemnetti), from Tasmania,
purchased on January 25th.

2 Caspian Terrapins (Clemmys caspica), from Palestine,
presented by Major EH. EH. Austen, D.8.O., F.Z.S., on January
loth.

March 18th, 1919.

ALFRED Ezra, Esq., Vice-President
in the Chair,

The Secretary read the following Report on the Additions
made to the Society’s Menagerie during the month of February
1919 :—

The registered additions to the Society’s Menagerie during the
month of February were 42 in number. Of these 12 were
acquired by presentation, 18 were deposited, and 12 were
purchased.

The following may be specially mentioned :—

1 Sallé’s Amazon (Chrysotis ventralis), from St. Domingo,
presented by Lady Edith Windham, on February 21st.

1 Starred Tortoise (Vestudo elegans) and 1 Ceylonese Terrapin
(Nicoria trijuga), from. Trincomali, presented by Mr. Edward
Canham, on February 14th.

Mr. F. Martin Duncan, F.R.M.S., exhibited a series of photo-
graphs and Jantern-slides of Marine Zoology, and drew attention
to the great economic importance of marine biological investiga-
tion to the successful continuance of our sea- fishing industries.

April 8th, 1919.
Dr. S. F. Harmer, F.R.S., Vice-President,

in the Chair.

Dr. F. E. Bepparp, F.R.S., exhibited and made remarks on
three foetal Sperm-Whales, drawing attention to the smallest
foetus exhibited, which measured four and a half inches in length.

Mr. R. I. Pocock, F.R.S., Curator of Mammals, gave an
exhibition, illustrated by lantern-slides, to show some of the
structural characters by which the genera of the Felidae may be

229, ON MARINE BORING ANIMALS

distinguished from each other ; special attention being drawn to
the formation of the feet in the Cheetah (Acinonyzx), to the modifi-
cations of the hyoidean apparatus in the Lions, Tigers, Leopards,
and Jaguars (Panthera), and to the position of the partition in
the auditory bulla in other genera.

April 29th, 1919.

Prof. Ernest W. MacBripz, D.Sc., F.R.S., Vice-President,
in the Chair.

The Srcrerary read the following Report on the Additions
made to the Society's Menagerie during the month of March
1919 :—

The registered additions to the Society’ Menagerie during the
month of March were 147 in number. Of these 120 were
acquired by presentation, 8 were deposited, 5 were received in
exchange, 10 were purchased, and 4 were bred in the Menagerie.

The following may be specially mentioned :—

2 Spotted Hyenas (Hyena crocuta), from Africa, purchased on
March 22nd.

1 Leopard Cat (Melis bengalensis), 2 Pandas (dlurus fulgens),
2 Malabar Squirrels (2atufa maxima), from India, purchased on
March 27th.

1 Wild Boar (Sus scrofa), from Crécy, presented by Gen. Sir
Henry Rawlinson, on March 12th.

An American Bison (Bison americanus), from North America,
and one English Park Bull (Bos tawrus), from Woburn, deposited
on March 24th and March 26th.

2 Meyer's Parrots (Pewocephalus meyer), from 8.K. Africa,
presented by the Marques of ‘'avistock.

1 Mikado Pheasant (Calophasis mikado), from Mt. Arizan,
Formosa, received in exchange.

3 Himalayan Monauls (Lophophorus impeyanus), purchased.

Mr. T. GerrarD, F.Z.S., exhibited a series of heads of Water-
buck (Kobus), collected by Dr. Digby, and drew attention to
points of interest relating to the variation in size and shape of
the heads.

Dr. W. T. Cauman, F.Z.8., exhibited and gave a detailed
account of various Marine Boring Animals, including Chelura,
Pholas, Teredo, etc., and drew attention to the economic
importance of the scientific investigation of such Marine

a Sse

ON A CHIMPANZEE KEPT IN THE OPEN AIR. ONS

Animals in relation to the serious damage caused by them to the
timbers of wooden ships and to piers, and to the masonry of
breakwaters and similar constructions.

In the absence of Mr. Grorge JENNISON, the SECRETARY
exhibited a series of lantern-slides of a Chimpanzee, the property
of Dr. K. Butter, of Cannock, Staffordshire, which had been
successfully kept alive in the open airin England for eight years.
In a short note accompanying the slides Mr. Jennison gave the
following particulars :—

““Members of the Society will be interested in the unique
success of Dr. K. Butter at Cannock, Staffordshire, in keeping a
Chimpanzee in the open air in England without artificial heat for
eight years. ‘The doctor, who isa keen naturalist and the possessor
in normal times of a fine and varied collection of animals and
birds, purchased Antony the chimpanzee in question (Anthro-
popithecus troglodytes niger) at Liverpool in 1910. He was told
that the animal was from the Congo region and was three years
old, but as it weighed only 14 lbs. it was probably younger.
It weighed Jast summer 84 lbs. Antony was kept during the
whole period in a brick building facing south-west, slept in straw
and had no artificial heat, summer or winter; he made good use
of the swings and perches in the exercise pen and took great
interest in trying, not always unsuccessfully, to break them.
He was very fond of Dr. Butter’s dog and delighted to play with
and pull it about, and he would also play for hours with the
tame ocelots. Ladies and strange animals he disliked, but
without showing any vindictive feeling towards them; injuries
he resented and remembered.

As is the case with most chimpanzees he recognized few
masters, only the doctor and the chauffeur could command his
obedience ; with them he could be trusted at liberty and naturally
enjoyed much freedom. His dietary, in which there was no
meat, consisted of milk, milk and bread, tea, coffee, cocoa which
he liked very sweet, bread with jam or black treacle, nuts,
locust beans of which he ate great quantities, all kinds of fruit,
carrots, turnips, potatoes raw or boiled in their jackets, and
water to drink. Asa luxury he had grapes or raisins and sweets
of all kinds, especially toffee and chocolates.”

May 13th, 1919.

Prof. Ernest W. MacBring, D.Sc., F.R.S8., Vice-President,
in the Chair.

The Secrerary exhibited two photographs of a living Okapi,
and stated that the animal had been in the possession of its

22.4 THE SECRETARY ON ADDITIONS TO THE MENAGERIE.

present owners for a period of over three years. The photographs
showed thatit was a young animal, and that probably, as in the
case of the Giraffe, the Okapi does not reach its adult stage until
five or six years of age.

Mr. E. G. Bouuencer, F.Z.S., exhibited a series of living
specimens of British Rats and their varieties, and stated that
during the past four years there was evidence that the so-called
Old English Black Rat had increased in numbers.

Lt-Col. S. Monckton Coprmman, I'.R.8., exhibited a series of
lantern-slides made from carefully prepared charts to illustrate
his ‘ Experiments on Sex Determination.”

May 27th, 1919.

A. Smiru Woopwarp, Hsq., LL.D., F.R.S., Vice-President,
in the Chair.

The Srcrerary read the following Report on the Additions
made to the Society’s Menagerie during the month of April
TSS) Ss

The registered additions to the Society’s Menagerie during the
month of April were 148 in number. Of these 50 were acquired
by presentation, 20 were deposited, 1 was received in exchange,
and 77 were purchased.

The following may be specially mentioned :—

5 Californian Sea-Lions (Otaria californiana), from California,
purchased on April 19th.

1 Brown Bear (Ursus arcios), from Murmansk, presented by
the Captain and Officers of H.M.S. ‘ Heellent’ on April 30th.

1 Mouflon (Ovis masimon) (Sardinia); 6 Llamas (Lama
gluma) (Peru), bred in Europe; 1 Hybrid Zebra and Donkey
(Lquus grévyi x H. asinus), bred in Europe, deposited by H.G.
The Duke of Bedford, K.G.

2 Bennett’s Wallabies (J/acropus bennett?) (Tasmania), bred in
Sussex, presented by Sir Edmund Loder, Bt., on April 16th.

1 Dusky Parrot (Pionus fuscus) (Guiana), received in exchange
on Apvil Ist.

5 Black-necked Swans (Cygnus melanocoryphus) (Southern
South America), purchased on April 17th.

EXPERIMENTS ON CULTIVATION OF VERNEUILINA. 225

The Secrerary exhibited, and made some additional remarks
upon the photographs of the young living Okapi that were shown
at the previous Scientific Meeting.

June 17th, 1919.

Prof. Ernest W. MacBripz, D.Sc., F.R.S., Vice-President,

in the Chair.

The Secrerary read the following Report on the Additions
made-to the Society's Menagerie during the month of May
1919 :-—

The registered additions to the Society’s Menagerie during the
month of May were 169 in number. Of these 52 were acquired
by presentation, 15 were deposited, 5 were received in exchange,
95 were purchased, and 2 were born in the Menagerie.

The following may be specially mentioned :—

1 African Hunting-Dog (Lycaon capensis), 3 (5S. Africa),
purchased on May 7th.

1 Mikado Pheasant (Calophasis mikado), deposited by H.G.
The Duke of Bedford, K.G., on May 14th.

1 Secretary-bird (Serpentarius serpentarius), from South Alrica,
purchased on May 7th.

1 South-African Amphisbeena (J/onopeltis capensis), new to the
Collection, from South Africa, deposited on May 28th.

Miss L. EK. Currsman, F.E.S., exhibited some living specimens
of Light-giving Beetles from Cuba.

Mr. EK. Heron-Auien, F.R.S., exhibited a series of lantern-
slides demonstrating some of the results obtained in the
experiments he had been carrying out in collaboration with
Mr. ArtHur Haruanpd, F.R.M.S., on the cultivation of Verneui-
lina polystropha Reuss, in hypertonic sea-water and gem sand.

Proc. Zoot. Soc.—1919, No. XVI. 16

No. 190.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON."
March 18th, 1919.

ALFRED Ezra, Hsq., in the Chair.

The SecrErary read a Report on the Additions to the Society’s
Menagerie in the month of February 1919.

Mr. H. R. A. Matzocr, F.R.S., F.Z.S., gave an account of his
investigations on ‘Some Points in Insect Mechanics,” illustrating
his remarks with lantern-slides and diagrams.

Mr. R. I. Pocock, F.R.S., F.Z.S., communicated a paper by
Mr. F. E. Buaauw, C.M.Z.8., “On the Breeding of Oryx gazella
at Gooilust.”

Mr. F. Martin Duncan, F.R.M.S., exhibited a series of photo-
graphs and lantern-slides of Marine Zoology, and drew attention
to the economic importance of marine biological investigation.

* 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 subseribe to the Publications; but it may be obtained on the
day of publication at the price of Sixpence, or, if desired, sent post-free for
the sum of Stx Shillings per annum, payable in advance.

The next Meeting of the Society for Scientific Business will
be held on Tuesday, April 8th, 1919, at 5.30 P.m., when the
following communications will be made :

Dr. F. E. Bepparp, F.RB.S., F.Z.S.
Exhibition of, with remarks on, three fatal Sperm-Whales.
Lancetor T. Hoesen, B.A., B.Sc.

The Progressive Reduction of the Jugal in the Mammalia.
(Communicated by Mr. H. W. Unthank, B.A., B.Sc., F.Z.8.)

G. A. Boutencrr, F.R.S., F.Z.S.

Description of Two new Lizards and a new Frog from the
Andes of Colombia.

The following have been arranged :—
April 29th, 1919. Dr. W. T. Cauman, D.S8c., F.Z.8.
Exhibition of Marine Boring Animals.
May 13th, 1919. Lt.-Col. S. Moncxron Copeman, F.R.S., F.Z.8.

Experiments on Sex Determination (illustrated by lantern-
slides).

The Publication Committee desire to call the attention of
those who propose to offer Papers to the Society, to the great
increase in the cost of paper and printing. This will render it
necessary for the present that papers should be condensed, and
be limited as far as possible to the description of new results.

Communications intended for the Scientific Meetings should
be addressed to

P. CHALMERS MITCHELL,
Secretary.
ZOOLOGICAL Society OF Lonpbon,
ReceEnt’s Park, Lonpon, N.W. 8.
March 25th, 1919.

No. 191.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

April 8th, 1919.

Dr. 8. F. Harmer, F.R.S., F.Z.S., Vice-President,
in the Chair.

Dr. F. E. Bepparp, F.R.S., F.Z.S., exhibited and made re-
marks on three fcetal Sperm-Whales, drawing attention to the
smallest foetus exhibited, which measured 44 inches in length.

Mr. H. W. Untuans, B.A., B.Sc.,‘F.Z.8., gave a résumé of
Mr. Lancelot T. Hogben’s Paper on ‘‘The Progressive Reduction
of the Jugal in the Mammalia.”

In the absence of Mr. G. A. BouLencer, F.R.S., F.Z.S., his
communication on ‘fT wo new Lizards and a new Frog from the
Andes of Colombia” was taken as read.

Mr. R. I. Pocock, F.R.S8., F.Z.8., Curator of Mammals, gave
an exhibition, illustrated by lantern-slides, to show some of the
structural characters by which the genera of Felide may be
distinguished from each other, special attention being drawn to
the formation of the feet in the Cheetah (Acinonyx), to the
modifications of the hyoidean apparatus in the Lions, Tigers,
Leopards, and Jaguars (Panthera), and to the position of the
partition in the auditory bulla in other genera.

* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent’s Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications ; but it may be obtained on the
day of publication at the price of Sixpence, or, if desired, sent post-free for
the sum of Stax Shillings per annum, payable in advance,

The next Meeting of the Society for Scientific Business will
be held on Tuesday, April 29th, 1919, at 5.30 p.m., when the
following communications will be made :—

Dr. W. T. Carman, D.Sc., F.Z.8.

Exhibition of Marine Boring Animals.

Noeu Tayter, B.Se.

A unique Case of Asymmetrical Duplicity in the Chick.

The following communication will be taken on

May 13th, 1919. Lt.-Col. S. Moncxron Copeman, F.R.S., F.Z.8.
Experiments on Sex Determination (illustrated by lantern-
slides).
The following Papers have been received :—
Gro. JENNISON, Esq. :
~~ «4 Chimpanzee in the Open Air in England.”
CraupE Mor tey, Esq., E.ZS.

Equatorial and Other Species and Genera of African
Ichneumonide.

The Publication Committee desire to call the attention of
those who propose to offer Papers to the Society, to the great
increase in the cost of paper and printing. ‘This will render it
necessary for the present that papers should be condensed, and
be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should
be addressed to
P. CHALMERS MITCHELL,
Secretary.
ZOOLOGICAL SOCIETY OF LONDON,
Regent’s Park, Lonpon, N.W. 8.
April 15th, 1919.

No. 192.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

April 29th, 1919.

Prof. Ernest W. MacBrips, D.Sc., LL.D., F.R.S8., Vice-President,
in the Chair.

The SECRETARY read a report on the Additions to the Society’s
Menagerie in the month of May, 1919.

Mr. T. Grerrarp, F.Z.S., exhibited and drew attention to some
special points of interest in a series of heads of Waterbuck
(Kobus), collected by Dr. Digby.

Dr. W. T. Cauman, D:Sc., F.Z.S., exhibited and gave a de-
tailed account of various Marine Boring Animals, drawing atten-
tion to the economic importance of the scientific investigation of
such forms of Marine Animals in relation to the serious damage
caused by them to the timbers of wooden ships and to piers, and
to the masonry of breakwaters and similar constructions.

‘The Srcrerary read a communication, illustrated by lantern-
slides, from Mr. Guo. JeNNIsoN, on “‘ A Chimpanzee in the Open
Air in England,” drawing attention to the fact that the animal

* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent's Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications ; but it may be obtained on the
day of publication at the price of Sixpence, or, if desired, sent post-free for
the sum of Six Shillings per annum, payable in advance,

had lived in a healthy and vigorous condition for a period of
some eight years in the private grounds of its owner, Dr. John
K. Butter, of Cannock, Staffordshire.

The next Meeting of the Society for Scientific Business will
be held on Tuesday, May 13th, 1919, at 5.30 p.m., when the
following communications will be made :—

Lt.-Col. 8. Moncxron Copeman, F.R.S., F.Z.8.
Hxperiments on Sex Determination (illustrated by lantern-
slides).

Nort Tayier, B.Sc.

A unique Case of Asymmetrical Duplicity in the Chick.

The following Papers have been received :—

CuaAuDE Mortey, F.ZS.

_ Equatorial and other Species and Genera of African
Ichneumonide.

Ernest . Unwin.

On the Structure of the Respiratory Organs of the Terrestrial
Isopoda.

J.T. Cunninenam, M.A.

Result of a Mendelian Experiment on Fowls, including the
Production of a Pile Breed.

C. W. Anprews, D.Sc., F.R.S., F.Z.8.

A Description of New Species of Zeuglodons and Leathery
Turtle from the Eocene of Southern Nigeria.

The Publication Committee desire to call the attention of
those who propose to offer Papers to the Society, to the great
increase in the cost of paper and printing. This will render it
necessary for the present that papers should be condensed, and
be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should
be addressed to

P. CHALMERS MITCHELL,
Secretary.
ZOOLOGICAL Society oF Lonpon,

Recent’s Park, Lonpon, N.W. 8.
May 6th, 1919.

No. 193.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.”

May 13th, 1919.

Prof. Ernest W. MacBrips, D.Sc., LL.D., F.R.S., Vice-President,
in the Chair.

The Srcrerary exhibited two photographs of a living Okapi,
and stated that the animal had been in the possession of its
present owners for a period of over three years. The photographs
showed that it was a young animal, and that prebably, as in the
case of the Giraffe, the Okapi does not reach its adult stage until
five or six years of age.

Mr. E. G. Boutencsr, F.Z.8., exhibited a series of living
specimens of the British Rats and their varieties, and stated that
during the past four years there was evidence that the so-called

Old English Black Rat had increased in numbers.

Prof. J. P. Hitz, F.R.S., F.Z.S., gave a résumé of Mr. Noel
Tayler’s communication on ‘‘ A unique Case of Asymmetrical
Duplicity in the Chick,” and illustrated his remarks by a series
of lantern-slides.

* 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 Szxpence, or, if desired, sent post-free for
the sum of Six Shillings per annum, payable in advance.

Lt.-Col. 8. Mowcxron Coprman, F.R.S., F.Z.8., read a paper
on “ Experiments on Sex Determination,” and illustrated his
vemarks by a series of lantern-slides of carefully prepared charts.

The next Meeting of the Society for Scientific Business will be
held on Tuesday, May 27th, 1919, at 5.30 p.m., when the
following communications will be made :—

J.T. Cunnineuam, M.A., F.Z.8.

On Result of a Mendelian Experiment on Fowls, including
the Production of a Pile Breed.

Miss KarHuren F, LAnper, B.Sc., F.Z.8.

Some Points in the Anatomy of the Takin (Budorcas taxicolor
whiter).

Epwarp PuHetps Auuis, F.Z.8.

On certain Features of the Otic Region of the Chondro-
cranium of Lepidostews, and Comparison with other Fishes and
higher Vertebrates.

The following Papers have been received :—
CLAUDE Mor :ey, F.Z.S.
Equatorial and other Species and Genera of African
Ichneumonide.
C. W. Anprews, D.Sc., F.R.S., F.Z.S.

A Description of New Species of Zeuglodons and Leathery
Turtle from the Hocene of Southern Nigeria.

The Publication Committee desire to call the attention of
those who propose to offer Papers to the Society, to the great
increase in the cost of paper and printing. This will render it
necessary for the present that papers should be condensed, and
be limited so far as possible to the description of new results.

‘Communications intended for the Scientific Meetings should
be addressed to

P. CHALMERS MITCHELL,
Secretary.
ZOOLOGICAL Society oF Lonpon,

Recent’s Park, Lonpon, N.W. 8.
May 20th, 1919.

No. 194.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*
May 27th, 1919.

A. Smith Woopwarp, Hsq., LL.D., F.R.S., Vice-President,
in the Chair.

The Secrerary read a Report on the Additions to the Society’s
Menagerie in the month of April, 1919.

Mr. J. T. Cunninenam, M.A., F.Z.8., communicated his paper
on the “ Result of a Mendelian Experiment on Fowls, including
the Production of a Pile Breed.”

Miss Katuuren F, Lanper, B.Sce., F.Z.8., described some Points
in the Anatomy of the Takin (Ludorcas tawicolor whitet), and
illustrated her remarks by a series of lantern-slides.

In the absence of the Author, Mr. EK. Punters Aus, F.Z.S.,
his communication ‘“ On certain Features of the Otic Region of

* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent’s Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications; but it may be obtained on the
day of publication at the price of Sixpence, or, if desired, sent post-free for
the sum of Six Shillings per annum, payable in advance.

the Chondrocranium of Lepidostews, and Comparison with other
Fishes and higher Vertebrata,” was taken as read,

The Secrerary exhibited, and made some additional remarks
upon the photographs of a young living Okapi that were shown
at the previous Scientific Meeting.

The next Meeting of the Society for Scientific Business will
be held on Tuesday, June 17th, 1919, at 5.30 p.m., when the
following communications will be made :—

F.R.MS.

Exhibition of Lantern-slides illustrating the cultivation of
Verneuilina polystropha Reuss., in hypertonic sea-water and
gem-sand.

CiaupDE Mortey, F.Z.8.
Equatorial and other Species and Genera of African
Ichneumonide.
C. W. Anprews, D.Se., F.R.S., F.Z.8.

A Description of New Species of Zeuglodons and Leathery
Turtle from the Eocene of Southern Nigeria.

G. A. BounencEr, F.R.S., F.Z.8.

(1) A List of the Snakes of West Africa from Mauritania to
the French Congo.

(2) A List of the Snakes of N orth Africa,

The Publication Committee desire to call the attention of
those who propose to offer Papers to the Society, to the great
increase in the cost of paper and printing. This will render it
necessary for the present that papers should be condensed and
be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should
be addressed to

P. CHALMERS MITCHELL,
Secretary.
ZOOLOGICAL SOCIETY OF LONDON,

Recent’s Park, Lonpon, N.W. 8.
June 3rd, 1919.

‘ BUN Dany am REA AIM RB ROD NADA OL, Kraan ae le ee a
ie Sa say AY MaHE Roe PES AEN 2 io. 1 SOS 1 ae :

\s -
uy i i Pie

apy

No. 195.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.
June 17th, 1919.

Prof, E. W. MacBrinpg, F.R.S., F.Z.S., Vice-President,
in the Chair.

The Secretary read a Report on the Additions to the Society’s
Menagerie during the month of May, 1919.

Mr. J. T. Carter, F.Z.8., gave a réswmé of his paper on the
‘Occurrence of Denticles on the Snout of Xiphias.”

Miss L. E. Currsman, F.E.S., exhibited some living specimens
of Light-giving Beetles from Cuba.

Dr. C. W. Anprews, F.R.S., F.Z.8., communicated his paper
on ‘“ New Species of Zeuglodons and a Leathery Turtle from the

Kocene of Southern Nigeria.”

Mr: Epwarp Heron-A.uen, F.R.S., F.Z.S., described the ex-
periments he had been carrying out in collaboration with Mr.
ARTHUR HARLAND, F.R.M.S., on the cultivation of Vernewilina
polystropha Reuss in hypertonic sea-water and gem sand, and
illustrated his remarks with a series of lantern-slides.

* 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 Siwpence, or, if desired, sent post-free for
the sum of Six Shillings per annum, payable in advance.

In the absence of the Authors, the following communications
were taken as read :—Mr. CuaupE Mortey, F.Z.S., ‘*‘ Equatorial
and other Species and Genera of African Ichneumonide.” Prof.
G. A. Boutencer, F.R.S., F.Z.S., (1) ‘“‘ A List of the Snakes of
West Africa from Mauritania to the French Congo,” and (2) “A
List of the Snakes of North Africa.” Rev. THomas R. R. STEs-
BING, F.R.S., F.Z.S., ‘‘ Crustacea from the Falkland Islands
collected by Mr. Rupert Vallentin: Part JIT.”

This Meeting closes the Session 1918-1919. The next Meet-
ing of the Society for Scientific Business will be held on Tuesday,
October 21st, 1919, at 5.30 p.m.

The following communications have been received :—
B. L. Buarta, M.Sc.

Notes on Indian Ciliate Protozoa.

E. DuKINFIELD Jonzs, F.Z.S., F.E.S.
_ Descriptions of New Moths from South-Hast Brazil.

The Publication Committee desire to call the attention of
those who propose to offer Papers to the Society, to the great
increase in the cost of paper and printing. This will render it
necessary for the present that papers should be condensed and
be limited so far as possible to the description of new results.

Communications intended for the Scientitic Meetings should
be addressed to

P. CHALMERS MITCHELL,
Secretary.

ZOOLOGICAL Socinry oF Lonpon,
Recentr’s Park, Lonpon, N.W. 8.
June 23rd, 1919.

iY elves
ee oe
Pate

Exhibitions and Notices (continued).
5 Page
Miss L. EH. Cnuzsman, F.E.S. Exhibition of living specimens of Light-giving Beetles
Lieven, (CUR ecte ie ase occa Sie id Gi Gc ie gh bees ACG ERS PGI oR SER ETE CREE a Ni ena ar ae)

Mr, E. Heroy-Attzy, F.Z.S8. Exhibition of a series of lantern-slides demonstrating results
Omexperiments on cultivationlob, Menneutlina . ie 2 she sce ec ee ee core cece ence 225

PAPERS.

1. On the External Characters of Existing Chevrotains. By R. 1. Pocock, F.R.S.,, F.Z.S.
Glier Mea tres LD: nays terse pte earcostenceetecacetestecMebeyede ei reese) ote caters abbas caatatclerarc lose 1

2, Report on Deaths of Animals in the Gardens in 1918, By J. A. Murray, M.D.,
Welune: Hon. Pabholomist tor the: SQUlety vasiene 0. leisis ele'enthse0 os. s te + eitacae es cc 13

3. On a Collection of Fishes from Lake Tanganyika, with Descriptions of three new
Species, By G. A. Bounenaur, F.R.S., F.Z.8. (Text-figures 1-3.) .............. 17

4, On the Skull and Affinities a Rana oat A. Dum. a Miss Joan B. Procrsr,
E.ZS. (Text-figures 1 & 2. SSELE THEO ORME OS ORR RRERRNERS Fl OL ane ICR ETRE OC oe 4L

. On the Breeding of Oryx gazella at Gooilust. By F.E. Buaauw,C.M.Z.S. (PlateI.). 29

6, A Comparative Study of certain Sense-Organs in the Antenne and Palpi of Diptera.
By K. M. Smirn, A.R.C.S., D.C. With Appendix by Professor H. Maxwunu
Lurroy, F.Z.8. (Plates I-IV. and Text-figures 1-48.) 20... cee cece ee eee eee 31

7. The Progressive Reduction of the Jugal in the Mammalia. By Lancenor T. Hoesen,
TEL TRISTE aa coro ora ted ns og SP) Ue Ora Od eyo RA rape Ge i RNR Sa Mane AR Oe 77

8. Deseriptions of two new Lizards and a new Frog from the Andes of Colombia. By
G. A. Boutananr, F.R:S:, H:Z:8.: ((Dext-feures 4) 65 '5,)/.).csc.les denacies ob ves oe 79

9, A Unique Case of Asymmetrical Duplicity (Duplicitas asymmetros) in the Chick. By
Norn Taytor, B.Sc. (Lond.). (From the Zoological Department, University of
London, University College.) (Plates I-III. and Text-figures 1 & 2.)............ 83

10. Some Points in Insect Mechanics, By H. R. A. Matuocn, F.R.S., F.Z.8. (Dext-
EV URUES LO) Paaten ay Shai ats “inl ips jnga’ls Chases peat RRR MER no Roem tach OI eee EAS erratd aes, Ge era EL TE

11. On some Equatorial and other Species and Genera of African Ichneumoninz con-
tained in the Collection of the British Museum. By Cravpm Mortey, F.E.S.,
i VAiShy CxO) 6.6. oboe op oom anu OOOO He Bees ode co Oso odoOGs onan doSdn en aoae 117

12, Results of a Mendelian Experiment on Fowls, including the Production of; a Pile
Breed. By J. 0. Ounningnam, M.A., F.Z.S: *(Blate T.) cc ieee cee oe eee LFS

15. Some Points in the Anatomy of the Takin (Budorcas tawicolor whitet). Based on the
examination of a specimen in the Zoological Society of London. By Miss Karuienn
F. Lanner, M.Sce., F.Z.8., Hon. Acting Prosector to the Society. (Plate I. and

{RORUESTeaTIR ONES (2) is Samora a Te Blea cig bit enS 4 6 60a REDO HOC EL CENCE 203
SUAGIEOD RIE coin. cle Slo gteel diay fro PRM CMO MONE CTR minis char ee cp Erin asian ole ie neat ops Mae ARE 1
List of Council and Officers .............. Tee Pha RCS NG AN ey cee ete ae a aie Ll oo rl
HOTT rOtmOOMUbe My simeyvremergc ey foie cies assy abel de oi mebrahaanvaue naDhads Sura anemone (2 sepa anni oa a aa we (eae lil
Sone Lea Pret OG OOMUMDULONS: «(24's ale nel Tals a Gisiods'. Cote uieiacdtiastewaian weft ox tls.< sere vii

PLATES.

1919, Parts I. & IT. (pp. 1-225).

Plate Page
Briaauw: I. Young calf of Oryx gazella and adult female uae | 29
abnormal bhonnst «ss sete ee ee sean vee eee :
Smit: I.
U. : ‘ars
Tit. Sense-organs of Diptera ......-....... Bil
IV.-
TAYLER: is |
ie AT cdloublesChiekemubiny Ota. ete ere ese Sere
Tre!
Cunsincuan: ~L. Alybrid'Common Bowls 2.3... <2) 0s shes ce an es
Miss Lanper: I. Posterior portion of Tongue of Takin ............ 203,
NOTICE.

The ‘ Proceedings’ for the year are issued in fowr parts, paged consecutively,
so that the complete reference is now P. Z. 8.1917, p.... The Distribution
is usually as follows, but on account of war conditions Parts I. & IT. are
issued together: —

Part I. issued in Mareh.
If. - June.
es Mince Ly A sate September.
IV. i December.

‘Proceedings,’ 1918, Parts III. & VI. (pp. 197-810), were published
together in March, 1919.

The Abstracts of the ‘ Proceedings,’ Nos. 190-195, are
contained in this Part.

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY
OF LONDON

1919.

PARTS III. & IV.

CONTAINING Paces 227 to 499, witH 13 Puates
AND 64 TExt?-FIGURES.

FEBRUARY 1920.

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

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

[Price Twenty Shillings. ]

LIST OF CONTENTS:

1919, Parts III. & IV. (pp. 227-499).

EXHIBITIONS AND NOTICES.

The Secretary. Report on the Additions to the Society’s eee ie during the months
of June, July, August, and September, 1919 ....

Mr, Oxuprietp Tomas, F.R.S. Exhibition of three interesting Mammals obtained by
ID rh NGL esh Asha Aisha Zep Iy AMOS a Hoda Mamma arco TOAACODAOCabOOKo LU SO OC Apes cst,

Dr. A. Suirn Woopwarp, F.R.8. Discussion on The Zoological Position and Affinities
Oh Lansiisnma((hext-tloume Us) ceva scistaterencis = open aie eeccieisiomnectars «ae See ace etanene ae

Prof, G. Exxior Situ, F.R.S. The Zoological position of Tarsius

Prof. J. P. Hint, F.R.S. The Affinities of Tarsius from the Embryological Aspect.
(Table, Plate I., and Text-figures 1-5.)

Prof. F. Woop-Jonzs, D.Se., M.B., F.Z.8. The General Anatomy of Tarsius

Mr. R. I. Pocock, F.R.S., F.Z.S. Structure of Tarsius .......0. Henke SOMO Oo we aetie eats
Mr. J. T. Connineuam, M.A., F.Z.8. Development of Tarsits....0+..-+..eeseses oats
Dr. P. Cuatmurs Mitcnert, K.R.S. Characters of Tarsius ....... SRO ee wioreleese

Prof, H. W. MacBripz, F.R.8., F.Z.S., Vice-President of the Society. Summing up the
Discussion on Tarsius

eo ee eve eee cece es FS ee ee sere re ee ee ee eter senor eseeoescs 0

Mr. F. Martin Doncay, F.R.M.S., F.Z.8. | Exhibition of, and remarks upon, a series

of photograpls showing the actinic quality of the light from a living Pyrophorus
Beetle

CPs cece eeseec oe ® Fee ese ent 2 OF FESS eevee: -Feseseee eter eeOFeeereseseeet tM ouee

Mr. E. Heron-Atxen, F.R.S., F.Z.S. Exhibition and description of a series of Skiagraphs
of the Foraminifera genus Verneudlina

Ce ceore =e cere oceorecerr ee PF ee eoere OG ones ee

The Secretary. Report on the Additions to the Society’s Menagerie during the month
Of October; 1919 Aik easton eee aE RECO eoae Heh wens eae seater

Sir Epwunp Gitns Lopsr, Bt.,; F.Z.S. Exhibition of, and remarks upon, a series of skulls
of the Beaver

ee rece eee e rere ee oe ee ooese th es cere ee ee Pose @eecs et ee ee eoeeneeee ee

PAPERS.

14. Report on Methods of Rat Destruction, By E. G. Bourxncsr, F.Z.8., Curator of
Reptiles, Zoological Society of London. With an Introduction by P, Cuamurs
Mircnei1, C.B.E., F.R.8., LL.D., D.Sce., Secretary to the Society .......... anit
15, On certain Features of the Otic Region of the Chondrocranium of Lepidosteus, and

aia pees with other Fishes and Higier Vertebrates.

By Epwarp Preps Atis,
Jr, F.Z,S..

Page

463
464

465
465

476
491
494
495
496

497

498
499
499

439

245

Contents continued on page 3 of Wrapper.

Een,

ZOOLOGICAL SOCIETY OF LONDON.

Tas Society was founded in 1826 by Sir Sramrorp Rarr.zs,
Mr. J. Saprnz, Mr. N. A. Vieors, and other eminent Naturalists,
for the advancement of Zoology and Animal Physiology, and for the
introduction of new and curious subjects of the Animal Kingdom,

and was incorporated by Royal Charter in 1829. [!

Patra.

HIS MAJESTY THE KING,

COUNCIL.
HIS GRACE THE DUKE OF BEDFORD, K.G., F.B.8., President.

Tae Hon. Cucitt Barine, M.A. E. G. B. Mrapz-Waxpo, Hse.

Tux Lorp Carmicuazt, G.C.S.I1., || PRror. Ernst W. McBripg, D.Sc.,

K.C.M.G. F.R.S., Vice-President.
Lr.-Cox. 8. Moncxron-Corrmay, || Mason Atzurr Pam.

M.D., BBS. P. CHatmers Mrircuert, Ese.,
Cuartes Drummonp, Ese., C2B Ee MEAG! W)Scs,. bb!

Treasurer. F.R.S., Secretary.

Aurrep Ezra, Ese., Vice-Pre- || Appian D. W. Porzocr, Ese,

sident. Tue Lorp QuEENBOROUGEH.

Harorp 8. Fereuson, Esa. Tun Marevis or Srigo, F.8.A.,
Carr. Huen S. Grapstone, M.A. Vice-President.

Aurrep H. Cocks, Ese., M.A. Masor Ricuarp S. Tayzor.
Pror. James P. Hiiz, D.Sc, || ANtHony H. Winerterp, Ese.

BRS. ARTHUR SmitH Woopwarp, Esw.,
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4 |

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P. CHALMERS MITCHELL,

Secretary.
Regent’s Park, London, N.W. 8.
September, 1919.
MEETINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON
FOR

SCIENTIFIC BUSINESS.

1919.
TuEsDAY, APRIL ......... So
af IMGANY, ch 2m Sas 13 and 27.
- JUNE. eeeanede 17.
Ws OcTOBER yy. se Bile
Ss NOVEMBER...... 4 and 18.

The Chaar will be taken at half-past Five o'clock precisely.

LOGE OGICAR SOCIETY. OF LONDON.

LIST OF PUBLICATIONS.

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P. CHALMERS MITCHELL,
Secretary.
Regent's Park, London, N.W. 8.
September, 1919.

These publications may be obtained at the Socizry’s OFFICE
or through any bookseller.

PROCEEDINGS

OF THK

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

Or THK

ZOOLOGICAL SOCIETY OF LONDON.

PAPERS.

14. Report on Methods of Rat Destrustion. By EH. G.
BouLencer, F.Z.8., Curator of Reptiles, Zoological
Society of London. With an Introduction by
PiCaamimrs. Marormin (Cobb. oR. S., iLeD., DiSex
Secretary to the Society.

[Received September 6, 1919: Read October 21, 1919.]

I. LyrropuctTion.

The destruction of Rats had a direct interest for the Zoological
Society, as the abundant supply of food and shelter in the
Zoological Gardens not only maintained a large indigenous
population, but attracted rats from all the neighbourhood. In
the beginning of 1919, the Council were of the opinion that it
would be of interest to visitors, and of educational value, to
arrange an Exhibition in the Gardens illustrating the natural
history of Rats and Mice found in this country, the damage that
they do to food, property and health, and the chief devices
employed for their destruction. It was also decided to associate
with the Exhibition an enquiry into the efficacy of these devices,
with the object of recommending methods that were proved to be
simple and practical. As the scheme appeared to be of public
utility, it was agreed to seek advice and co-operation from the
Board of Agriculture and Fisheries, and from the Local Govern-
ment Board, as these Government Departments were known to
be taking an active interest in what had become a national
problem. The Board of Agriculture had already collected mucn

Proc. Zoou. Soc.—1919, No. XVII. 17

228 MR. E. G. BOULENGER: REPORT ON

information with a view to the introduction of legislation on the
subject, and the Local Government Board had a large experience,
especially on the relation of Rats to public health. Both Depart-
ments gave us immediate encouragement and assistance. Each
made a grant of £100 towards the expenses of the investigation.
They placed all the information in their possession at our dis-
posal, gave us invaluable advice as to the method of arranging
the exhibition and conducting the enquiry, and made our work
easier and more productive. Throughout the progress of the
enquiry, Mr. F. A. Falford of the Board of Agriculture, and
Mr. EK. C. Read, the practical expert appointed to assist him,
were in constant co-operation with us. The Council of the
Zoological Society delegated Mr, KH. G. Boulenger, Curator of
Reptiles at the Gardens, to arrange the exhibition and conduct
the research, with Mr. W. C. Harris as clerical and general
assistant, and the necessary attendants. Various members of
the Council with special scientific or practical knowledge,
Mr. R. [. Pocock, F.R.S., and other members of the staff, and a
number of private Fellows of the Society gave great assistance.
Special thanks are due to Dr. C. J. Martin, F.R.S., and
Mr. A. W. Bacot of the Lister Institute, to Dr. H. B. Newham
of the London School of Tropical Medicine, Dr. W. J. Howarth,
Medical Officer of Health for the City of London, Mr. W. Dalton,
Mr. Max Baker,and Miss Frances Pitt for the loan of specimens,
diagrams, etc. for the Exhibition ; to a very large number of
private persons and firms who lent apparatus, models, or samples of
poisons, traps, ete.; to Dr. J.8. Warrack, Deputy Medical Officer
of Health for the Port of London Sanitary Authority, for facili-
ties to study the methods employed on ships: to Mr. J. Horne,
Resident Manager of the Wood Lane Depot of the Royal Borough
of Kensington, for facilities to carry out experiments at that
depot; to Messrs. Lawson and Co. Ltd. of Bristol, Messrs. Boots
Ltd., and the Proprietors of the Ratinol Company, for supplying
large quantities of material for the experimental tests.

A separate file was kept for the records of each appliance,
preparation, or method. The rats in the Gardens gave the
opportunity for a large number of tests. Visits were made to
shops, warehouses, farms, houses, rnbbish-dumps, and so forth,
so as to get information relating to different conditions. In
Mr. Boulenger’s Report, the practical conclusions arrived at are
given first, and are followed by details regarding the methods
found to be most successful, and those which yielded more
uncertain or unsatisfactory results.

It is fortunate that the experiments point to the efficacy of
means that are simple to use, not costly, nor, in the torm recom-
mended, dangerous to human beings or stock. The best bait is
bread, the best gas is sulphur-dioxide, the best poison, Squills,
and the best trap a device not dangerous to other animals. With
regard to Squills, experimental research of a kind outside the
scope of our enquiry is required. The substance is obtained from
the common Mediterranean bulb, Scilla maritima, and is supplied

METHODS OF RAT DESTRUCTION. 229

by a few Firms in various forms, all of which are not efficacious.
It is important that a chemical investigation should be made, to
«letermine the active agent in the plant, and the best means
of preparing and supplying it.

It is vital to the success of any attempt to reduce seriously the
rat population of the country, that the work should be carried
out simultaneously over large areas. Rats are migratory
animals and will readily desert a locality which they find to be
uuwholesome, for an area in which measures are not being taken
against them. They are also prolific, and will rapidly multiply
beyond the capacity of areas in which they are undisturbed, and
reinvade areas in which they had been reduced. ‘These are
matters for administrative action, and the provisions of the new
Rat Destruction Bill, if carried out rigorously, should prove
successful. It is also to be remembered that the winter season,
now approaching, is the best time to make the attack. The
numbers of rats are naturally at their lowest, and the scarcity of
food in their outdoor haunts drives them to take baits more
readily, and to approach traps less warily.

II. Report on Mernops or Rar Desrrucrion.

By E. G. Bounencrr, F.Z.8., Curator of Reptiles, Zoological
Society of London.

In connection with the Rat Exhibition held this sumimer in
the Society’s Gardens, investigations were actively pursued on
various methods of rat destruction, and the following conclusions
were arrived at :—

1) That where the method necessitates baits being put down,
the food offered should differ from that which forms the staple
supply of the rats.

That dry bread is always accepted. That oatmeal, cheese, and
tallow are also attractive baits, and that fish, lard and dripping,
so frequently recommended, usually find favour only when no
other food is available.

That faint traces of the oils of Rhodium and Aniseed, so
commonly used to attract rats, instead of unproving the bait
have the contrary effect.

That the tastes of the Old Hnglish Black Rat (Ratius rattus)
and the Common Brown Rat (Rattus norvegicus) are identical, at
least in their surroundings in this country.

(2) That when rats are present in large numbers, and where
it is not practicable to use gus, poisoning is the best and cheapest
method to adopt for their destruction.

That of all the poisons we experimented with, Squill poison,
the extract of the bulb of the Mediterranean plant Scilla
maritima, which in the small quantities used in rat destruction
is harmless to domestic animals, gave the most satisfactory

results. hat it may be used with greatest success in the liquid
ae

230 MR. E. G. BOULENGER : REPORT ON

form, bread being soaked in a solution of the poison mixed with
equal parts of mille.

That good results may also be obtained when Barium Carbonate
is used in conjunction with Syuill Since Barium Carbonate has.
a corrosive action on the mucous membrane of the stomach,
compelling the rodents to leave their holes in search of water,
the device of putting down bowls of liquid squill in the vicinity

of the area treated should be resorted to in order that the rats.

may take more poison, thus ensuring their destruction.

(3) That the destructive power of virus is unreliable, and that
better results are to be obtained by the use of Squill, Barium
Carbonate, and other poisons.

(4) That trapping, provided the correct types of traps are
employed, is to be recommended at all times of the year, and
should be resorted to with special energy during the winter and
in the periods between the poisoning campaigns, in order to
destroy the surviving rats. That by persistent and skilful
trapping the numbers of rats, even in badly infested localities,
can be greatly reduced and kept under control.

That rats like passing through passages showing light at the

far end, provided the middle of the passage is not dark. That of

all the traps we tested, a wire tunnel-shaped cage-trap, known
to us as the Brailsford trap, and which embodied the above
principles, gave the best results. In the type in question, which
we regret is no longer being manufactured, the open doors at

each end shut when the rat steps on a platform in the centre of
the passage. With regard to this trap the results differed
according to the height of the doors and passage, and that when
these were eight inches high the largest percentage of captures

was obtained. The width was found immaterial, provided of

course it allowed for the free passage of the rat.
That the ordinary steel Gin trap and the “ Nipper,” a break-
back trap with a moveable platform, were the only other traps to

give very successful results, and with these the highest percentages.

of captures were obtained when they were covered with a wire
tunnel of the Brailsford pattern, but without the doors, the traps
being substituted for the platform.

That it is superfluous to avoid handling traps on the assump-
tion, often entertained, that rats are detracted by the odour
of man.

(5) That hunting with dogs and ferrets is a good method to

adopt for the reduction of the rat population, 340 of the 1076 rats
caught by us between the months of April and September being
captured by the aid of ferrets and a single dog.

(6) That gassing,an expensive method, has a distinct advantage

over all others in ae it kills not only the adult and half-erown.
rats, but also the newly-born in their nests.

That Sulphur Dioxide is the best gas for use in rat destruction,
and for killing rats on ships or in confined spaces it has no rival.

MELHODS OF RAT DESTRUCTION. OST

That it may also be used with some success in fumigating rat-
holes in the open.

In connection with the above conclusions I have the following
observations and recommendations to make :—

(a) Owing to rats when active measures are taken against
them frequently migrating to neighbouring farms and estates,
co-operation in their destruction is imperative, and the whole of
a rat-infested area should be treated on the same day, and not in
parts at different times as is at present generally done.

Oy 2 rick surrounded by 53 feet high galvanized iron sheetings
dug 23 feet into the ground, is just as effectually protected, aad
at a amel smaller cost, than when built on a massive non-portable
staddle.

(c) At present the giving of rewards for rat-tails is indulged in
by some and not by others. As those who do so cease when they
discover they are called upon to pay for their neighbours’ rats,
the payment of such rewards should be made compulsory for all
employers, at any rate during the winter months when scarcely
any young are about, but when every adult at this time repre-
sents a potential increase in the numbers for the near future.
This would go a long way towards keeping the vermin in check.

(@) In view of the recent undoubted increase in the numbers
of not only the Common Brown Rat but also the Old English
Black Rat, all ships should at regular intervals be subjected to
treatment with Sulphur Dioxide gas by means of Sulphur
Candles or a Clayton or some similar apparatus.

(e) The formation of a research school in the country, where
further experiments on rat-destruction could be continued
throughout the year, is desirable. That at this school the various
poisons recommended should be made up and supplied direct te
the rat-ofticers appointed by the County authorities; such officers
being from time to time required to attend a course there, and
be made acquainted with the latest developments.

It should be borne in mind that ou investigations were con-
<lucted at a time of the year—May to September—when plenty
of food is available for the rats, and that some of the methods
which resulted in failures would possibly have succeeded if
applied during the winter months.

The successes obtained deserve therefore, I think, special
consideration.

We realized that the first step towards finding an effective
method of rat-destruction, whether by means of poisons or traps,
was to ascertain what baits the rodents found most attractive, as
no device dependent on food being taken could, however ingenious,
otherwise prove a success. ‘The baits referred to in the table
below were given to rats in captivity and in the wild state, and
were nearly all put down together on over fifty different occa-
sions. In the case of the rats kept in captivity, the animals

232 MR. E. G. BOULENGER: REPORT ON

were changed from time to time in order to preclude any one set:
being more favourably inclined to any special diet. It will be
observed that plain bread was preferred to anything else, and it
was found more attractive than the staple food-supply in.
slaughter-houses, granaries, and fat-factories. All mealy sub-
stances were liked, especially oatmeal. With regard to fats, lard
and dripping were seldom taken, but tallow was readily eaten..
Contrary to the statements in many books on the subject of rat-
destruction, meat, except tripe, and fish,even when smoked, were.
only accepted when no other food was available.

Food faintly flavoured with oils of Rhodium and Aniseed,.
instead of improving the bait, had the contrary effect.

Although the Black Rat (fattus rattus) is supposed to have a
predilection for a fruitarian diet, this was not the case with the
individuals we experimented upon, their tastes not differing from
those of the Common Brown Rat (Ratéius norvegicus), bread and
other mealy substances being always preferred to fruit and vege-
tables. The results of the tests on the rats in captivity and on those
living in freedom being almost identical, they have been tabulated
together. The percentages given below are based on the ratio.
between the attractiveness of the various baits and plain bread—
the favourite food, which is represented by the figure 100 :—

Bread (plain) sees ace ew elOOL, |r SA le viaaa cies eeu Renee 0
Do. (faintly flavoured with Cabbawe ssi ssid sms oausearae 20
Rhodium) ............ 40 Banana aaeeoe eeu cop iets ee ae OO)

Do. (faintly flavoured with Walloweck sree wece ceen ce mTO
Aniseed) .............. 30 Drippinloy tees sc cee eae ee ae)

Do. (soaked in milk) ......... 60 Tardy .seusceing Sat calneanaes cemmeeees anny
Toone aandl MAVEN oooansensacoccsooc | GO) TRONS CTL eae OS age SU TG)
Oatmeal... eee 80 Tripe Ey RDU POGUE Bic SERRE
Barley eee ees 80 TSBCOL MEN Wee ree aaa heen. serrata ca Os
(OPES head geeGean a s5ee can eteccaunens saben 0) Teer Reha ee ae ee
Mini Ze ecdes. wecldcnboeeneseeao sa 0 With (iesh) eto
Den pileMicall yee ae treme ceetcs 10 “° Des (smoked) 6.0 te, eon
Malte CU UAL pean nae PE ae JBVMGANEP TEAS: cocosdonesocoacancsa GLO
Potatoes (taw) ....-...-----.....--. 10 it H@iieae We dvehiee ne ietaee tree

Do. (mashed with milk) . 20 |

Poisons.

Where rats are present in very large numbers under conditions
which precluded gas being used, poisoning was found to give the
most satisfactory results. Although the use of poisons involves
less trouble and expense than gassing and trapping on a large
scale, there has been much objection to the method owing to the
danger to man and domestic animals, and to the possibility of
the rats dying under floorings. With regard to the first objec-
tion, the danger has perhaps not been much exaggerated, for
although we have ourselves had no such unfortunate experiences,
we have heard of numerous cases where pigs and other valuable

MELHODS OF RAT DESTRUCTION. 233

animals were killed in addition to the rats as a result of treat-
ment with phosphorus, arsenic, and strychnine. As to the
second objection, in the course of our numerous tests, conducted
both in urban as well as rural districts, we did not receive a
single complaint with respect to rats dying under the floorings of
buildings, and the possibility of this occurring appears to be
more remote than has been suggested.

We spent some time experimenting with both Phosphorus and
Arsenic, but it was realized from the first that the dangerous
nature of these poisons, unless they proved of very exceptional
value, precluded their being recommended for general use.
Phosphorus, although obviously easy to detect, proved to be an
attractive bait, and far more so than arsenic; unlike the latter,
however, it does not, especially in summer, retain its toxic
properties for more than a few days.

Below are given tables showing the results of our tests and
investigations with phosphorus and arsenic preparations of
various kinds. In these, and in our tests and investigations
with other poisons, we have termed the result a ‘‘ Success”
where a definite reduction in the number of rats was recorded.
In the cases marked with an asterisk we were not present at
the test, but the facts were ascertained from reliable witnesses.

Phosphorus.
| jl a
Nature of Premises a2 ee yeult R E
am Laer amount 0 tesult. temarks.
: Bait used.

|
1. Military Camp 15 lbs. Success.| A very large number of dead rats
and Farm. | were collected on the day following |
| the test. This, a very badly in-
fested area, was almost completely |

cleared of the vermin.

2. Farm. 2 Ibs. Success.) Reduction in number of rats was |
| recorded.

3*. Gardens. | 2 Ibs. Success.| The rats were entirely eliminated. |
| This Garden had been treated for a
couple of months previous to the }
test with Virus, but with no
| resuit.

4*, Farm. 3 Ibs. Success. A reduction in the number of rats |

resulted. |

5. Part of Small 10 Ibs. Failure. | Although this town was badly in-|

Town. fested only a minute quantity of
the poison was taken. Subsequent
treatment with extract of Squills
was a success. |

6. Rubbish Dump. 5 Ibs. | Success., A number of dead rats were collected

234 MR, E. G. BOULENGER: REPORT ON
Arsenic.
Nature of Premises | Approximate

: = : ; sult. ‘ks.
oilocalitic } rmount of | Result Remark
° Bait used.

1, Private 2 Ibs. Success. | The rats vanishel for a period of 3 or
Residence. 4 weeks, after which they reappeared
but in small numbers.

2, Rubbish Dump. 4 lbs. Failure. | Only a small quantity of the bait
was taken and no dead rats were
discovered.

3. Baker’s shop. 3 Ibs. | Failure. | Although most of the bait dis-
appeared no reduction in thie |}
number of rats was recorded.

4. Rubbish Dump. 10 Ibs. | Success. | Most of the bait was taken on this
oceasion and a large number of
dead rats were collected.

|
|
|
|
|
|
|
|

5. General Stores. | 4 \bs. Failure. | The bait was not taken, i

With Strychnine, which is acknowledged a most effective poison,
we nade no experiments, as, owing to its deadly action, its whole-
sale use could not under any circumstances be recommended.

Much time was spent testing various preparations of which

Barium Carbonate formed the basis, this rat-poison having been
selected as the most suitable by the United States Depart-
ment of Agriculture and the Indian Government. Although
13 or 2 grains suffices to kill a rat, Barium Carbonate is more or
Jess harmless to domestic animals, cats and chickens withstanding
10-15 grains, and an average-sized dog over 100 grains. It has
also the advantages of being cheap, tasteless, and therefore easily
made attractive by mixing with a suitable bait: further, its
corrosive action on the mucous membrane of the stomach induces
the rats to leave their holes in search of drink. Although the
successes obtained with Barium Carbonate were not so decisive
as those resulting from treatment with Squill—a poison we refer
to Jater,—with one exception a definite reduction in the number
of rats resulted from our tests, and we satisfied ourselves that it
was as effective as the more dangerous poisons, such as phosphorus
and arsenic. In most cases the poison was put down with tallow-
fat mixed in equal proportions, and smeared on bread. ‘To make
certain of the destruction of all the rats leaving their holes in
search of water it was found advisable to place within their reach,
on the day following the treatment, bowls containing a solution
of squill and milk; the rats being thus made to partake of more
poison in their efforts at relief.

—

METHODS OF RAT DESTRUCTION. 235

Barium Carbonate.

| |

iy, . Approximate |
hata ee Prewiats amount of | Result. | Remarks.
| ese Bait used. }
|
| 1. Surgical Gut 2 Ibs. Success. All the bait was taken and the rats
Store. | disappeared for a month.
4 |
|
| 2. Ditto. 2 Ibs. | Do. | As the rats returned treatment was
| again applied in the store which
| was once more cleared.
| 3. Sweet Factory. 2 lbs. |Failure.| No definite improvement was re-
| corded, although nearly all the bait
| | was taken.
| 4, Garden. 1 lb. | Success.| The rats disappeared entirely for a
| couple of months.
Is. Knacker’s Yard. 5 lbs. Maas _ These premises were so seriously in-
| | | fested that a clearance was out of
| the question, the yard being strewn
| with bones to a depth of 6 feet.
| The rats were to be seen in
enormous numbers in the daytime.
| All the bait was immediately
| taken.
| |
6. Canteen. be | Success. A decrease in the number of rats was
| recorded.
s |
| | |
| 7. Granary on 5 lbs. Success.) Do.
| Wharf. |
|

Other preparations which we tested contained Plaster of Paris,
Magnesium Sulphate, Croton, and Squill. With the exception
of the last-named we found that. all these, however attractively
disguised, were only taken if no other food was available, and
their use as a raticide could therefore be disregarded. Squill,
which is obtained from the bulb of a plant (Scilla maritima)
occurring in great abundance on the Mediterranean coasts, can
be used either in the form of a powder or of a solution, and in
France and its Colonies it has for some years been known to ke
an effective rat-poison. In this country it has been little used,
as until quite recently the only proprietary preparations con-
taining the extract of the plant have been in the hands of
companies which likewise sold virus, and its use was only recom-
mended in the event of the virus proving a failuve. Although
extremely toxic as far as rodents are concerned—the minimum
lethal dose for a rat being only half a grain—Squill is compara-
tively harmless to domestic animals.

As will be seen below, a very large number of tests were made

236

MR. E. G. BOULENGER: REPORT ON

with Squill preparations, some in the liquid, others in the powder
form. In the liquid form the Squill was mixed with an equal
proportion of milk into which 81]bs. of bread were soaked for

every gallon of the solution.

As a powder it was given mixed

with tallow, or tallow and oatmeal, and smeared on bread. With
one exception, however, our successes were obtained when the
liquid Squill was mixed with bread and milk, and it is in this
form that we recommend its use.

Squill.

Nature of Premises
or Locality.

Approximate

amount of
Bait used.

Result.

Remarks.

1. Store.

2. Farm.

3*, Private Resi-
dence.

. Part of Town.

. Poultry Yard.

6. Poultry Yard,

. Farm.

2 Ibs.

10 Ibs.

1 lb.

20 lbs.

3 Ibs.

Success.

This store, in which the rats were to

be seen in broad daylight, was yery |

badly infested, and for 3 months
previous to our test an average of
50 rats a week were caught with
traps. As a result of the treatment

a large number of dead rats were |

discovered and the store was en-
tirely cleared of the rodents.

A number of dead rats were collected

in the course of the week following ;
the treatment, and a diminution }

in their numbers was recorded.

The rats were entirely cleared from
the premises.

A number of dead rats were col-
lected on the day followmg the)

application, and a diminution in
their numbers was recorded. ‘This
town had previously been treated
with virus ard phosphorus with
no results.

A decrease in the number of rats
was recorded, and a number of
carcasses were found on the pre-
mises on the day after the poison
had been put down.

No rats have been seen in this yard
since the test was made. Virus
had previously been used without
SUCCESS.

This farm was very badly infested,
and the rats were in evidence in
large numbers in the daytime.
The day following the treatment a
very large number of carcasses
were collected. The farm is now
comparatively free of the vermin.

MELHODS OF RAT DESTRUCTION. 230

|
Approximate |
amount of | Result. Remarks.
Bait used. |

Nature of Premises
or Locality.

8. Garden. 3 Ibs. | Success.) Large number of dead rats found |
in the week following the treat-
ment. A great improvement was

| effected.
9*. Farm. 8 Ibs. | Failure.) The bait was not taken.
10. Farm. & Ibs. | Success. A number of dead rats was collected

| on the day following the treatment,
| and a diminution in their numbers |
was recorded.

sy F H . . i
11*. House and 4 lbs. | Failure. | The bait was not taken.
Stables. | |

12. Town. 150 Ibs. | Success.| This test proved a great success, |

| dead rats being found in very large
/ quantities in all parts of the town
| | on the two days following the |
| treatment. The rats have been

| practically exterminated.

13. Embankment. | 4 lbs. | Success.) The rats were entirely cleared from
| this embankment.

14, Residence. 2 Ibs. Failure. Only a very slight diminution in the |
number of rats was recorded.

|
15. Food Store. 4 lbs. Failure.| Most of the bait was taken, but no |
good results were obtained.

Tn tests numbers 13, 14, and 15 squill-powder was laid down.
In all other cases the solution was used.

As a result of our investigations, we have no hesitation in
coming to the conclusion that Squill-solution is the most effective
rat-poison, and recommend its use in preference to Barium
Carbonate for the following reasons :—

-{1) That for rodents it is three times as toxic.
(2) That it is even less harmful to most domestic animals.

The one point against the use of Squill lies in the fact that at
present it is Eoetion moze expensive than most other poisons,
The plant grows, however, in extraordinary abundance on both
the African and European coasts of the Mediterranean, and
therefore no ditticulty should be experienced in obtaining a large
supply at a reasonable cost.

Virus.

The discovery some time back of a bacillus pathogenic to
rodents led to a new method for the destruction of rats and mice.
Following this discovery, the virulence of the organism was

238 MR. E. G. BOULENGER: REPORT ON

raised by artificial methods in the laboratory and a number of
Rat Viruses were placed on the market.

The manufacturers of these claimed that the bacillus was so
virulent that it would kill rodents in about ten days, and that
within a month the disease would spread with fatal results to the
whole of the rat—or mouse—population of the area under treat-
ment. The results of recent investigations on the efficiency of
some of the viruses on the market have been disappointing.
Bainbridge * has experimented on a very large number of rats in
captivity, and the results of his experiments showed that the
destructive power of all the viruses he tested was inconstant,
the death-vate in the different experiments varying from 20 to
50 per cent. Further, according to this experimentator the
presence of agglutinins 1 in the serum of the rats which survived
after being fed on virus gave reason to suppose that a certain
proportion acquired immunity and were therefore unlikely to
succumb to a second infection.

Experiments with the rat-viruses conducted some years ago
during the outbreak of plague in San Francisco also gave poor
results.

It must be recognized that, if generally successful, this method
of exterminating rats and mice by spreading among them a
disease, not affecting Man and domestic animals, could not be
improved upon, and I was therefore anxious to give a thorough
test to all the viruses on the market in the hope that the results
of the experiments under natural conditions would differ from
those obtained in the laboratory.

We made in all 7 tests with different virus preparations, and
of these one only was an unqualified success; three were partial
successes, a very slight reduction in the number of rats being
recorded, and three were absolute failures. A number of reliable
witnesses who visited the Exhibition in the course of the summer
informed us of their experiences, and we ascertained the successes
in these cases to amount to about 33 per cent.

Trapping.

Although we found that trapping alone will not always rid us
of the rats where they are present in very large numbers, never-
theless if the best traps are used skilfully and persistently the
vermin can be greatly reduced. Owing to the abundance of food
in our Gardens, the locality is not one where much success would
be expected. Nevertheless, in the four months referred to in
this report, we caught with traps alone 736 rats. Of the many
different traps laid down, three types accounted for 85 per cent.
of the total catch. Some of the traps that proved to be failures
were those we found most frequently in use in the rat-infested
localities which we visited, while the most successful type of
trap, several of which had been laid down in our Gardens for

* Journal of Pathology, vol. xiii. 1909.

° MEYHODS OF RAT DESTRUCTION. 239
several years, is now, so far as we can make out, no longer being
manufactured. The latter trap, known to us as the Brailsford
‘Trap, consists of a long, narrow wire cage with doors at each end,
which, when the trap is set, remain open, leaving a direct passage
through. A platform in the centre, where the bait is placed, is
connected by a spring which when trodden upon releases the
doors, imprisoning the rat or rats. Such traps were put down
on 687 occasions and accounted for the capture of 209 rats. Two
makes of this trap were formerly sold, the one in which the
height of the passage measured 44 inches, the other in which it
measured 7 inches. As the Jatter gave far better results than
the former, in order to ascertain the proportions which would
give the best results, traps of different height and width were
tested, with the result that the type with the greatest height
was Tound most successful.

The following are particulars of these tests :—

Percentage of rats caught in traps'42 high (25., 19.
y
9? 29 oe) 99 7 9 AO Ord 28.
7 ©
” ” 7) o) 8 Spe isreteateys 34.

The width, provided it allows for the free passage of the rat,
makes no difference.

Other cage-traps tested included such well-known types as the
ordinary Cage Trap with bait hook, and with a platform con-
nected by a spring with the door, the Eelpot or Wonder, the
Round Wire, and the “‘ Mysto.” We also experimented with a
number of new types sent to us, and with Barrel traps and
Stockades.

The ordinary Cage Trap.—YVhis trap, which is sold at most
ironmonger shops, we saw in use on many occasions in the eourse
of our visits to rat-infested localities, but were invariably in-
formed that it caught no rats. This was likewise our experience
with the type without the bait-hook. When, however, it was
provided with the platform which when trodden on closed the
door, more satisfactory results were obtained, 16 rats being
caught with 166 of these traps.

The Eelpot or Wonder Trap, which permits the free entry of
the rats whilst preventing their exit, is divided into two com-
partments connected with a trap-door. This trap has, I know,
frequently been responsible for the capture of numbers of rats,
With us it did not prove a great success. The keeping of a
female in the trap and baiting with the female genital organs
having been said to entice the males, both these devices were
resorted to, but with negative results.

The Round Wire Trap is a simple type, the rats having an
entry through a hole in the centre, but being prevented from
escaping by the presence of a number of fine loose wires which
hang from the sides of this aperture and close round tbe hole
when the rat climbs up in order to escape. For some weeks we

240 MR. E. G. BOULENGER: REPORT ON

placed one of these traps, measuring 94 inches in height, in
various localities in the Gardens frequented by the rodents, but
without success. At a later date we found that this type was
being used with good results in a rat-infested store, and on
compar ing 1t with the one we had been using found that it differed
in measuring only 7 inches in height, and a similar one was
procured foe our use. If not very successful it nevertheless
caught a number of rats, and obviously therefore, as was the case
with the Brailsford trap, it proved specially efficacious only
when conforming to certain measurements. With 124 traps of
this kind (7 inches in height) 12 rats were captured.

The “Mysto” Trap consists of a metal structure connected
with a tank full of water, The bait is placed just inside the
‘door, which closes on the entry of the rat. In order to effect its
escape the rat climbs up the cage to a hinged platform, which
collapses, precipitating the rodent into the tank. The collapse of
this platform automatically opens the door of the cage and the
next rat is thus enabled to enter and repeat the perfor mance.
Although I have been informed by some users of this trap that
it gave good results, the one which we tested for several weeks
was responsible for the capture of one rat only.

A number of new types of cage-traps were sent us and given a
trial. Some were most ingenious, but as they failed to catch
rats 1t is unnecessary to allude to them.

It was found that several of the cage-traps we tested were
made of too light wire, with the result that the captured rats
were able to fore the wires apart and escape.

Barrel traps have frequently been asserted to be very effective,
and we therefore experimented with these both in the Gardens
and in other localities where rats were numerous. One of the
methods we employed was to fill the barrel up to within a foot of
the top with earth, covered with sawdust and chaff, and upon
this placed various bait. For some days we allowed the rats to
feed from the barrel. When we found that most of the bait had
been taken for several nights in succession, water was substituted
for the earth, also covered with a layer of sawdust and chaff;
and we awaited results. None were, however, forthcoming, the
rats obviously suspecting a deception. The other method con-
sisted of covering the barrel with brown paper, upon which the
bait was placed. After the food had been taken for a few days
a cross was cut in the paper in order that the rats might fall
through into the barrel. This answered very much better, and
a number of rats were captured by this device.

The ordinary Stockade trap consists of a wooden stockade
about 4 feet high constructed to enclose a space about 10 feet
square. On one side is a door a foot wide and high, which is
open and closed by a cord and pulley. The Stockade having
been baited and the door kept open, the rats are for a week or so
allowed to feed and to run in and out. Subsequently the
operator secretes himself and at a suitable moment releases the

METHODS OF RAT DESTRUCTION. 241

cord controlling the door. This contrivance having proved
success we were anxious to give it a trial. The weak part, how-
ever, of this trap appeared to be the amount of time that would
be wasted by the man controlling the sliding-door, and therefore,
in order to dispense with the human element, with the help of
my friend Mr. Mark Barr, I worked out a scheme which enabled
the open door, which rested on a hinge, to fall by being drawn
towards an electro-magnet, the latter being in contact with a
number of batteries and an alarm-clock. ‘Thus if we wished the
door of the trap to be shut at a certain time all that was
necessary was to set the alarm for that time, when the current
acting on the magnet released the door. |With this a number of
rats were captured. During the winter months, when less food
is available, it may possibly give still better results.

The steel Gin trap has for years been used with success, and is
the type generally popular with the gamekeeper. Nine hundred
and seventy-three of these traps laid down in our Gardens were
during the four months in question responsible for 173 rats, a
percentage of 17. Considering that we obtained a higher per-
centage with the Brailsford trap, we presumed that the gins
would be specially successful if laid in the centre of any passage
showing light at the far end, and we consequently experimented
by covering them with large drain-pipes cut longitudinally in
half. This experiment was, however, a failure. Ata later date
we continued the experiment on the same lines and had _ holes
bored in the top of the pipes in order to allow for some light to
enter. This was slightly more successful than when the middle
of the pipe was in complete darkness, and a few rats were
caught. The results, however, could not compare with those
obtained when the traps were put down in the usual manner.
As the Brailsford traps continued to give such satisfaction, L
ordered a number of covers exactly similar to those of the cage-
trap, but without doors, bottom or centre platform, in order to
ascertain whether, when the gin-traps were covered with these
wire-passages, a higher percentage of successes would be attained.
The result was highly satisfactory, the gins, when placed under
these cheap wire-covers catching almost as many rats per trap as
the Brailsford and a higher number than when placed in runs
in the ordinary way.

Break-back traps.—With one exception these gave no good
results. The exception was a type known as the “ Nipper,” and
with 1962 of these traps 237 rats were caught. This, a very
inexpensive trap, is most suitable for putting down indoors or
under ‘cover, as in wet weather the wood is liable to warp and
render the appliance useless. We would suggest to the makers
that a similar trap be made of metal for outdoor use. As in the
case of the Gin traps, the “ Nippers,” when placed under wire-
covers, were responsible for a higher percentage of captures.
The figures below show results obtained with 100 Gin and
100 Nipper traps when laid down in the ordinary manner and

ee, MR. Lk. G. BOULENGER: REPORT ON

when covered with a wire tunnel 8” high by 4” wide by 24” or
36” long.

Number of vats caught with 100 Gin traps laid down in

the ordinary manner ...... 19
Do. do. 100 Nipper traps do. ...... 14
Do. do. 100 Gin traps covered with

WAURS WOUNITEIS scocassccoocodoson 28.
Do. do. 100 Nipper traps do....... alt

An additional advantage in the use of these wire covers lies in
the fact that the Gin and Break-back traps can be laid down
in poultry-runs without fear of injury to the full-grown birds.

It having constantly been stated that traps should be handied
as little as possible, and that in setting them gloves should be
worn ov the hands rubbed in earth, we experimented with a very
large number of traps to ascertain whether rats are really
influenced by the human odour and are alarmed thereby. On
fifteen occasions we laid down in a rat-infested locality outside
the Gardens 20 traps, 10 Gin traps and 10 Nipper traps, half of
these being carefully handled with gloves rubbed in earth and
faintly scented with aniseed, the other half being freely handled
with the naked hand. Asa result of this experiment we satisfied
ourselves that there is no necessity for attempting to disguise the
human odour, 36 rats being caught with the traps handled in
the ordinary way and only 31 with those laid down with gloves.

The following table shows the number and type of traps laid
down between the months of April and September with the
percentages of captures :

Type. THnes set, [Rats eaughe, Percentage.
Brailsford Trap (all sizes).................. 687 209 33 |
GuineerayS) eee ee et aoe rers ne reese 973 173 ie
Do. (placed under wire tunnel) ... 135 36 26 |
Nip perinapy sh see teem ern eterna Ulan 1962 237 12 |
Do. (placed under wire tunnel). 126 24. 19 |
Round Wire Trap (7" high)............... 124 12 10 |
Do. (Oe lichens 61 0 Osa
Ordinary Caze Trap (with bait hook |
Only) ere 82 0) | (0) |
Do. (with, platform) . 166 16 10 |
Helpot or Wonder Trap................0006 195 17 9 |
Oiherstymestecestaseeestecse cree cree 528 12 2 |

METHODS OF RAT DESTRUCTION. 243

Hunting with Dogs and Ferrets.

Although this method is practically useless in most towns, it is
of the greatest assistance in keeping down the rat population in
the country. A good ratting dog seldom fails to locate the
presence of a rat, and will always differentiate between the in-
habited and uninhabited holes. The results obtained when ferrets
co-operate with a dog are usually very satisfactory, and such hunts
with sticks and shot-guns in the Society’s Gardens, in conjunc-
tion with trapping. has led in the past few years to a very
distinct reduction in the numbers of rats, in spite of the general
increase elsewhere. The time spent on rat-huntimg with the
help of dogs and ferrets has during the mouths of April to
September averaged eight hours a week and has resulted in
340 rats being killed. Like trapping, hunting with dogs and
ferrets should be pursued with special activity during the winter
months, and during the periods following poisoning with Squill
or Barium Carbonate.

Gassing.

This method has the distinct advantage over all others in that
it kills not only the adult rats but also the newly-born in their
nests. ‘The gases most frequently advocated for rat-destruction
are Carbon Bisulphide, Hydrocyanic Acid, and Sulphur Dioxide.
Although both Carbon Bisulphide and Hydrocyanic Acid are very
effective, the dangerous nature of these gases, the former being
highly inflammable and explosive, and the latter very poisonous,
odourless, and invisible, makes it impossible to recommend their
use. Sulphur Dioxide, however, is visible and non-inflammable,
and harmless to man when inhaled in small quantities.

In the course of this summer we were afforded opportunities of
being present at fumigations by means of sulphur candles, and
an apparatus manufactured by the Clayton Fire Extinguishing
and Disinfecting Company, ia which the sulphur is burned in a
furnace supplied with air induced by a draught, and the gas
driven off under pressure through a hose.

The use of the candles for killing rats in their burrows we
found ineffective, the gas penetrating too slowly; when the
sulphur dioxide is, however, driven into the holes under pressure
the whole network of runs is permeated in a few seconds, making
the existence of the rats underground impossible. Although
many of the rodents escape only to die in the open, some no doubt
recover, and the co-operation of a dog is therefore desirable. The
large Clayton machines which are used for fumigating ships and
large buildings are driven by petrol or electricity. I was shown,
however, the model of a new portable hand-driven apparatus
which appeared suitable for the treatment of hedges and embank-
ments. No opportunity was afforded us of witnessing gassing
on ships with these machines, but we were able to investigate
the system of rat destruction by means of sulphur candles as

Proc. Zoot. Soc.—1919, No. XVIII. 18

244 REPORT ON METHODS OF RAT DESTRUCTION.

employed by the Port of London Authority. The method gives
very satisfactory results and could not we think be improved
upon.

Preventive Measures.

The chief preventive measures, such as the protection of food
supplies, the destruction and prompt removal of garbage, the
repairing of defective drains, and the rat- proofing of buildings,
have been preached so consistently of late that it is unnecessary
to allude to them further. It must remain with the medical
officers of health and other sanitary authorities to see that these
conditions are enforced.

Protection of Ricks.

The building of ricks on rat-proof platforms has been officially
recommended. These platforms, however, besides being extremely
costly are not easily portable, and as the ricks are annually erected
on different sites, this method of protection can only be adopted
in rick yards. It has been found by expomment, which We have
confirmed, that rats will not dig to a depth of more than 22 feet,
and therefore all that is needed is to surround the rick with
galvanized iron sheets, 55 feet high, dug slightly over 2 feet into
the ground. The cost of such an erection would amount to less
than half that of a platform. The sheets are portable and would
last for many years.

Increase in the numbers of the Old English Black Rat.

We ascertained in the course of our investigations that not
only had the Common Brown Rat very greatly increased in
numbers in recent years, but that since 1910 the Old English
Black Rat had become much more abundant, and in London both
Species are now in some parts living in harmony, not only on the
same premises but in the same rooms. On the one floor of a
factory in Holborn we captured both species of rats, and also
specimens of the Alexandvine Rat—the brown variety of Rattus
rattus, and the black variety of Rattus norvegicus. A well-known
London rat-eatcher, who has kept records of his captures, informs
me that he is at the present time catching as many Old English
Black Rats as Common Rats in localities where, prior to the war,
the latter only were found.

This recent increase in the numbers of the Old English Black
Rat is disquieting, and can only be checked by the enforcement
of stricter measures for their destruction on incoming vessels.

ON THE OTIC REGION OF LEPIDOSTEUS. 245

15. On certain Features of the Otic Region of the Chon-
drocranium of Lepidosteus, and Comparison with
other Fishes and Higher Vertebrates. By Hpwarp
Pratps Anhisgdirs 2:8:

[Received May 6, 1919: Read May 27, 1919.]

In a figure of a 149 mm. specimen of Lepidostews osseus, Veit
(07) shows the larger part of the dorso-lateral edge of the otic
portion of the chondrocranium formed by a laterally projecting
vidge which he says is currently called the crista parotica, and
he ‘adopts this name for it. It will, however, be later shown
that the anterior portion of this ridge is not included in the
crista parotica of current descriptions of mammalian embryos,
and there is some doubt as to any part of it being the strict
homologue of that crista. The name is therefore inappropriate,
and as the ridge lies, in fishes, in part on the sphenotic and in
part on the pterotic portion of the chondrocranium, I shall call it
the spheno-pterotic ridge.

The sphenotic portion of this spheno-pterotic ridge is formed,
in Veit’s 149mm. specimen of Lepidosteus, by the ‘dorso- lateral
edge of the autosphenotic, and it begins, cater oly, at the
summit of the postorbital process. Its pterotic portion is w holly
of cartilage, but it supports, in this fish as in Amia, the lateral
edge of the dermo-pterotic. The anterior three-quarters,
approximately, of this cartilaginous portion of the ridge forms
the dorsal edge of the ar ticular facet for the hyomandibula, and,
-as shown in fine figures given, it apparently lies along the dorso-
lateral edge of the ridge of the lateral semicircular canal. At
the hind end of the facet for the hyomandibula the ridge is joined
by a much less pronounced one, which forms the ventral edge of
the facet for the hyomandibula, the two ridges, united, then con-
tinuing onward a short distance as a stout and rounded ridge,
which lies along the lateral, instead of the dorso-lateral, surface
-of the ridge of the lateral semicircular canal and ends somewhat
abruptly with a curved hind edge slightly posterior to the
anterior edge of the epiotic bone and some little distance anterior
to the transverse plane of the vagus foramen. The marked,

postero-laterally projecting corner 2 the hind end of this ridge
of Ama (Allis, 97, fig. 8) is thus wanting in Lepidosteus, car
the ridges of these anne fishes must, nevertheless, have approxi-
mately ‘the same posterior extent, for the hind end of the ridge
is traversed in each of them by a canal which transmits the
ramus dorsalis of the nervus glossopharyngeus.

Along the lateral portion of the dorsal surface of the chondro-
cranium there is a sulcus longitudinalis, which is bounded
laterally by the spheno-pterotic ridge and mesially by a ledge,
‘the rounded edge of which lies at a much higher level than the

246 MR. E. P. ALLIS ON THE OTIC REGION OF

spheno-pterotic ridge and in the line prolonged of the lateral
edge of the epiotic bone. The lateral portion of this sulcus.
lodges that thickened lateral portion of the dermopterotic that
is traversed by the main latero-sensory canal, and its deepest.
portion lies between the ridges of the anterior and posterior
semicircular canals. The sulcus runs posteriorly over the hind
edge of the ridge of the lateral semicircular canal, and is there:
in communication, across the ridge of the posterior semicircular
canal, with a slightly depressed region which lies directly beneath
the overhanging lateral edge of the epiotic and extends ventrally
to the dorsal edge of the vagus foramen. ‘This depressed region
doubtless lodges a part of the thymus, for, in an 80 mm. specimen
of this fish, I find a large anterior portion of that gland lying
close against the cranial wall, immediately postero-lateral to the
descending limb of the posterior semicircular canal, and having a
small dorso-anterior prolongation which passes upward postero-
mesial to the levatores arcuum branchialium and the adductor:
and levator operculi, at their imsertions, and then over the hind
edge of the spheno-pterotic ridge on to the dorsal surface of its.
hind end.

The depression for the thymus, above referred to, is separated
from the posterior surface of the chondrocranium by a sharp.
ridge, which Veit calls the crista occipitalis lateralis, this name
evidently having been adopted from Gaupp’s (93) descriptions
of Rana, where the so-designated ridge is said to be a band of
eartilage which forms the dorsal boundary of the vagus foramen,
and connects the otic capsule at the middle of its height with the
dorsal end of the occipital arch. The crista of Lepidosteus is thus.
not the exact homologue of the crista of Rana, and would seem
to be a ridge secondarily developed upon that crista. It forms.
the boundary between the dorsal and lateral surfaces of the occi-
pital portion of the chondocranium, and lies in Lepidostews as in.
Rana wholly upon the os occipitale laterale (exoccipital), but in
many of the Teleostei the corresponding ridge lies as much upon
the basioccipital, opisthotic, and pterotic as upon the exoccipital.
In Lepidosteus it is said by Veit to vanish, anteriorly, on the roof
of the labyrinth region of the cranium, but it is not so shown in
his figures, there running directly into the ventro-posterior edge.
of the epiotic bone, but separated from that bone by a narrow
space which has the appearance of being a notch in the edge of
the ridge. The epiotic forms the dorso-postero-lateral corner
of the chondrocranium, and apparently has no relatioas whatever-
to the ridge of the posterior semicircular canal, for it is shown
lying definitely postero-mesial to the prominentia canalis semi-
circularis posterioris. Veit says that he could find no indication
whatever of the epiotic being formed of two components, an
epiotic and opisthotic, as described by Parker (’82) in embryos
of this same fish. An opisthotic bone is, In any event, wholly
wanting in this fish, as is also, even in the adult, an autopterotie.

In a 204mm. embryo of this fish, Veit (11, fig. 18, pl. c.)-

THE CHONDROCRANIUM OF LEPIDOSTEUS. QAT

‘shows, on the dorsal surface of the otic capsule, a marked depres-
sion, which corresponds to that portion of the sulcus longi-
tudinalis of the 149 mm. specimen that les posterior to the ridge
of the anterior semicircular canal. This supraotic depression
opens posteriorly, across the posterior portion of the ridge of the
dateral semicircular canal, between that ridge and the ridge of the
posterior semicircular ¢ canal, on to the lateral (surface of the chondro-
cranium, and a slight depression leads from its dorso-mesial edge
across the ridge of the posterior semicircular canal into a depres-
sion which lies postero-mesial to the latter ridge, between that
ridge and the bounding edge of the foramen magnum. This
latter depression is called by Veit the fossa supratemporalis, and
it corresponds to the postero-ventral prolongation of the supra-
temporal groove of my descriptions of Scomber (Allis, 03).
There is in this embryo no epiotic process in any way com-
parable to that in the 149mm. specimen, but the basal portion
of that process must be represented in the summit of the ridge
of the posterior semicircular canal, for, in my 80 mm. specimen of
this fish, I there find the epiotic already represented by a thin
layer of perichondrial bone. The strongly developed epiotic
process of the 149mm. specimen must, therefore, have been
formed by the addition, to this perichondrial bone, of bone of
membrane origin.

he so-called crista parotica of this embryo is said by Veit
(11, p. 168) to be a ridge which forms an anterior prolongation
of the ridge of the lateral semicircular canal (‘‘ springt in ihrer
Fortsetzung eine scharfe Leiste lateralwarts vor’’), and it ends,
‘anteriorly, at the summit of the postorbital process. The anterior
portion of the ridge is said to be perforated by a canal which
lodges the recessus dorsalis spiracularis, this part of the ridge
thus having no relations whatever to the ridge of the lateral
semicircular canal. The posterior portion of the crista is said
to form the dorsal edge of a small oval depression which
apparently occupies the full width of the ventral surface of the
ridge of the lateral semicircular canal, and Veit calls this entire
fignrepsion the articular facet for the hyomandibula.

In a 20 mm. embryo of this fish, examined in serial transverse
‘sections, I find a longitudinal depression which has exactly the
position of the so- called facet for the hyomandibula of Veit’s
204mm. embryo, but the hyomandibula, which is thin, articu-
lates with the dorso-lateral quarter only of this depression, there
lying immediately beneath the spheno-pterotic ridge (Veit’s
crista parotica). The ventro-mesial edge of the depression is
formed by a slight ridge, which gives insertion to the adductor
hyomandibularis, this ridge forming the dorsal edge of a groove
which lies between it and the hulle) acus tica and lodges the vena
jugularis. Between this slight ridge and the hyomandibula
there is a large lymph vessel which, at the hind edge of the
adductor hyomandibularis and anterior to the levatores arcuum
‘branchialium, separates into two parts, one running ventrally

248 MR. FE. P. ALLIS ON THE OTIC REGION OF

along the lateral surface of the cranium to join and accompany
the vena jugularis and the other running outward and ventrally,
internal to the opercular bones, into the gill-cover. Posterior to
the adductor hyomandibularis, the ridge that forms its surface
of insertion gradually vanishes along the ventro-lateral surface of
the ridge of the lateral semicircular canal, and, posterior to the
ridge, the levatores arcuum branchialium and the adductor and
levator opereuli have their insertions at a higher level, imme-
diately ventral to the hind end of the spheno-pterotic ridge, or
even posterior to that ridge. In my 80mm. specimen the
hyomandibula is much stouter than in the 20mm. one, and a
definite ridge there forms the ventral edge of its articular facet,

this ridge lying at about the middle of the depression described
in the 20 mm. embryo, and corresponding to the ridge that forms
the ventral edge of the facet in Veit’s 149 mm. specimen.

Ina 14mm. embryo of Lepidostews the so-called crista parotica
is said by Veit (11) to be a short ridge which forms the ventral
border of the foramen by which the ramus oticus lateralis issues.
on the dorsal surface of the chondrocranium, and it is shown in
the figures given lying anterior to the ane ior edge of the
articular facet for the hyomandibula, and separated from it by
a depressed region which probably lodges the recessus dorsalis:
spiacularis. The ridge lies between the transverse planes of the
trigeminus and facialis openings of the trigemino-facialis chamber,
and, as the lateral wall of that chamber is formed by the otic
process of the palato-quadrate (pars ascendens quadrati), the
crista parotica is a ridge that lies approximately along the line
where that process fuses with the cranial wall. The so-called
erista parotica of this embryo and the ridge that forms the dorsal
edge of the articular facet for the: hyomandibula thus have similar
relations to structures derived, respectively, from the branchial-
ray bars of the mandibular and hyal arches (Allis, 18). The
articular facet fer the hyomandibula is said to Jie,in this embryo,
on the ventral surface of the prominentia ampulle lateralis, and
its dorsal edge is not referred to as a part of the crista parotica.
In 11-12 mm. embryos the hyomandibula is said to be here

fused with the wall of the otic capsule.

The spheno-pterotic ridge of Lepidosteus, the crista parotica of
Veit’s descriptions, is thus a ridge which apparently has two
distinctly different parts, which are related, respectively, to the
dorsal ends of the branchial-ray bars of the mandibular and hyal
arches. The ridge is of relatively late ontogenetic development,
and there is nothing in the descriptions of this fish to warrant
the assumption that either of its two parts has been developed
primarily in special pa to the related branchial-ray bar.
Veit’s conclusion (07, p. 179) that the entire ridge has been
developed in relation to the articulation of the hyomandibula
with this part of the cranial wall is, furthermore, evidently incor-
rect, for the hyomandibula of his 149 mm. specimen lies, approxi-
mately, beneath the two middle quarters only of the entire ridge.

THE CHONDROCRANIUM OF LEPIDOSTEUS. 249

The ridge forms the bounding edge between the dorsal and
lateral surfaces of the chondrocranium, and it may be that it has.
developed for this particular purpose, but there is the evident
suggestion that it has been developed in some relation to the
main latero-sensory canal, for that canal always lies parallel to
and slightly mesial to it, and the lateral edges of the dermal bones.
that later enclose that canal are always supported by it. The
relations of the ridge to the ridge of the lateral semicircular canal
are then simply fortuitous. The ventral edge of the so-called
facet for the hyomandibula of Veit’s 203 mm. embryo corresponds.
to the opisthotic ridge of my descriptions of Amia, Scomber, and
the Mail-cheeked Fishes, and has no relations whatever to the
articular facet for the hyomandibula.

In the prepared skull of a large specimen of Lepidosteus
platostemus, I find the slightly depressed region which, in Veit’s
149 mm. specimen, lies immediately posterior to the hind end of
the spheno-pterotic ridge, as an irregular and relatively deep
fossa which lies mostly on the lateral surface of the cranium but
partly also on its posterior surface, the ventral portion of the
fossa cutting across the dorso-anterior end of the lateral occipital
ridge and its dorsal portion, which has pit-like depressions in its
floor, extending upward mesial to the hind end of the spheno-
pterotic noses The lateral occipital ridge lies wholly on the

exoccipital, ending at the dorsal end of that bone , and mesial to
its dorso-anterior end an entirely independent ridge begins, and,
lying wholly on the epiotic, runs dorso- -anteriorly toward the
summit of that bone. This latter ridge is thus the homologue of
the epiotic ridge of my descriptions of Amia, Scomber, and the
Mail-cheeked Fishes, and the lateral occipital ridge is the homo-
logue of that part of the lateral occipital ridge of the latter fishes
that lies ventral to that process of the opisthotic that gives
articulation to the pedicel of the supraseapula.

In the prepared skull of a much smaller, but still adult spe-
cimen of Lepidosteus osseus, somewhat similar conditions were
found ; but in a second specimen of this fish, not previously dis-
sected, the lateral occipital ridge was found continving upward
on to the ventral surface of the overhanging epiotic process, but
there double, the postero-mesial portion of the ri idge corresponding
to the independent epiotic ridge of the other specimen. In this
second specimen there is a short but well-defined groove on the
lateral surface of the chondrocraninm, anterior to the dorsal end
of the lateral occipital ridge. This groove lies external to the hind
end of the lateral semicircular canal. in the hollow between that
ridge and the ridge of the posterior semicircular canal, is seen in
posteri lor as oar as lateral views of the cranium, iad leads.
dorsally into the hind end of the supraotic depression. It
evidently corresponds to the ventro-posterior continuation of the
temporal groove of Amia, Scomber, and the Mail-cheeked Fishes,
but in Lepidostews it lodges a part of the thymus, and it is un-
questionably this gland that is the cause of the marked differences

250 MR. E. P. ALLIS ON THE OTIC REGION OF

in the grooves of different specimens of this fish, and of the
irregularities in the epiotic and lateral occipital ridges. A stout
tendon arises from the hind end of the spheno-pterotic ridge,
and, running posteriorly, at first Jateral to the thymus but later
enveloped in that gland, gives insertion to an anterior extension
of the trunk-muscles; this tendon thus representing the stout
posterior process of the teleostean pterotic, which is wanting in
Lepidosteus. This tendon spreads posteriorly, and a part of it is
inserted on a ventral process of the suprascapula, and another
part on the large ee supraclavicular ligament; these attach-
ments gnenshne the ligament which, in certain of the Teleostei,
replaces the pedicel of the suprascapula of others of those fishes.
The occipito-supraclavicular Ligament is a large tough fascia
which arises from a process on the lateral surface of the basi-
occipital, lies upon the external surface of an anterior extension
of the trunk-muscles which has its insertion on the lateral surface
of the cranium ventral to the vagus foramen, and is inserted
mainly on a process along the internal surface of the lateral
edge of the supraclavicula, but partly also on the clavicle.
The levator and adductor operculi, and the levatores arcuum
branchiahum, arise from the cranial wall ventral to the ligament
that represents the posterior process of the pterotic, and anterior
to the groove for the thymus. <A stout aponeurotic formation,
which extends ventro-posteriorly into the trunk-muscles, arises
from the epiotic process, and that process is evidently developed
in relation to it. The epiotic ridge is strongly developed and
is certainly largely of membrane-bone, the line of the ridge not
even following the line of the posterior semicircular canal, as it
loes in many of the Teleostei.

There is thus in Lepidostews, as Veit stated, no functional
temporal fossa, but there is a supraotic depression which, if
invaded by the trunk-muscles through the interval between the
epiotic process and the hind end of the spheno-pterotie ridge,
would give rise to that functional fossa, and a small anterior
extension of tne trunk-muscles is in position so to invade it.
If the trunk-muscles were to invade it they would evidently tend
to deepen it by cutting into the massa angularis from behind
-and above, and this would give rise to a temporal ¢ eroove similar
to that in Amia, which would not, primarily, extend anterior ‘ly
beyond the anterior semicircular canal. The development of an
‘autopterotic, and the accompanying reduction of the dermo-
pterotic, would then give rise to the temporal groove of Scorpena
(Allis, ?09). this groove naturally lying mesial to the line of
fusion of the dermo- and auto-components of the pterotic, and
hence mesial to the latero-sensory canal that traverses that bone.
If the trunk-muscles were then to push forward dorsal to the
anterior semicircular canal, the groove of Scomber would arise,
the anterior prolongation of the groove so formed lying dorsal to
the parietal and frontal, being much less deep than its primitive
posterior portion, and being separated from that portion by

THE CHONDROCRANIUM OF LEPIDOSTEUS. 251

a marked ledge (Allis, 03). A perforation of the mesial wall
of this anterior prolongation of the groove would give rise to the
conditions described by Ridewood (’05) in certain of the Clupeide,
this perforation of that wall being called by him the temporal
foramen. The foramen is, however, probably simply a fenestra,
for, in a single specimen of Clupea alosa that I have examined, I
‘can find no nerves or vessels traversing it. It opens directly into
the dorsal end of the mid-brain recess of the er Panial cavity, and
it would seem as if it must be closed by membrane, but no such
membrane was evident in my specimen. The pre-epiotic fossa of
Ridewood’s descriptions of these fishes is simply a depression
in the mesial wall of the deeper posterior portion of the entire
temporal groove.

A modification of the temporal groove, as above described,
occurs when, as in certain of the Teleostei. the massa angularis
is excavated from its lateral surface, ventral to and in the hollow
of the semicircular canal, to form the subtemporal fossa of
Sagemehl’s (91) deseriptions. The axis of this fossa les trans-
versely to that of the temporal groove, and its arched roof forms
‘a transverse ridge across the floor of the latter groove, at about
the middle of its length ; and when the temporal groove is roofed
by dermal bones, and so becomes a fossa, and the subtemporal
fossa is extensive, as in certain of the Barbide, it practically
suppresses that part of the temporal fossa that les dorsal and
anterior to it. The temporal fossa then becomes a large but
shallow depression on the posterior surface of the cranium, from
which, according to Sagemehl (’91, p. 553), a simple cleft may
extend forward between external and internal plates of the
pterotic (Sqguamosum, Sagemehl) and represent the anterior
portion of the primitive fossa. This cleft is said by Sagemehl
to be in large part filled with fatty tissue which contains pigment-
‘cells and a few scattered muscle-fibres, the presence of the
muscle-fibres seeming to show that the groove of these fishes was
primarily more extensive, and that it has suffered reduction as a
result of the development of the subtemporal fossa. In Albula
(Ridewood, ’04) this type of fossa is also found, but the anterior
portion of the fossa has not been completely pinched off and
‘suppressed, and the fossa further differs from that in the Barbidee
in that its anterior portion lies beneath the frontal bone, and in
that its floor is in part formed by the prootie.

In Elops the temporal groove is similar to that in Albula, but
the trunk-muscles have here pushed forward in the direction of
the little recess described by me (’03, p. 51) at the antero-lateral
corner of the deeper posterior portion of the temporal groove of
Scomber. The resulting anterior extension of the temporal fossa
of this fish thus lies lateral to the anterior semicircular canal, in
the angle between that canal and the anterior portion of the
lateral semicircular canal, and ventral, instead of dorsal, to the
investing bones on the dorsal surface of the cranium. Its
anterior end reaches the alisphenoid region, and is there bounded

252, MR, E. P. ALLIS ON THE OTIC REGION OF

both laterally and mesially by portions of the alisphenoid bone,.
the groove thus splitting the dorsal edge of that bone and being
lodged between its two parts. In Albula the trunk-muscles fill
the! entire temporal groove. Whether or not this is also true of
Elops I cannot tell from the remnants that I have of the skull of
this fish, but a re-examination of these remnants shows that the
anterior semicircular canal is enclosed in the prodtic, and not, as
stated in my work on the Mail-cheeked Fishes, in the alisphenoid.

In Catostomus and Moxostoma, the temporal groove is said by

Sagemehl (91, p. 550) to be nearly transverse In position, and to
open on to the lateral instead of the posterior surface of the
chondrocranium. The epiotic is said to form the posterior
boundary of the groove, and the pterotic its anterior boundary ;
and the mesial portion of the groove is said to be roofed by the
epiotic, pterotic, and parietal. It is not said what muscle, or
muscles, occupy the groove in these particular fishes, but as no.
muscle is anywhere mentioned in relation to the groove, in any
of the several species of the Cyprinids deseribed, excepting only
an anterior extension of the trunk-muscles, it is natural to
assume that this muscle was considered by ‘Sagemeh] to have-
here invaded it. In Catostomus, Sagemehl found a greatly
reduced opisthotic, but in Iocostonea succetta he could find no
trace of this bone.

In a specimen of Moxostoma aureolum, I find the temporal
groove directed postero-mesially, practically as shown in Sage-
meh s figures of Catostomus teres, but its anterior wall is formed
by a tal aad Shiba ridge, instead of | oy a broad surface, as is-
apparently shown in Sagemehl’s figures of Catostomus. The
antero-mesial end of the groove is roofed by the postero-lateral
portion of the parietal, that bone here widely separating the:
superficial portions of the epiotic and pterotic. A deep V-shaped
incisure, which lies in the axis of the temporal groove, cuts into
this roof- forming part of the parietal, the two limbs of the V
forming the antero-mesial portions of the anterior and posterior
bounding edges of the groove. The remainder of the anterior
bounding edge of the groove is formed by a ridge on the pterotic,
the corresponding part of the posterior bounding edge being
formed, in its mesial portion, by a ridge on the epiotic, and, in
its lateral portion, by the dorsal edge of the opisthotic. The
epiotic portion of the edge corresponds to the mesial edge of the
groove of Scorpena, and to that edge of the deeper posterior
portion of the groove of Scomber, this! portion of the edge of the
groove of Moxostoma ending abruptly in a per pendicular portion
which corresponds to the mesial edge of the posterior opening of |
the groove of the other two fishes. The opisthotie portion of
the edge begins at the base of this nes teat portion, and
corresponds to the floor and the lateral edge of the posterior
opening of the groove of Scorpena and Scomber. There is
accordingly, in this edge of the groove of Moxestoma, a large
angular incisure, and it opens directly into the dorsal portion of

THE CHONDROCRANIUM OF LEPIDOSTEUS. Ua

a conical depression on the posterior surface of the cranium,,.
which gives insertion to a portion of the trunk-muscles.

The opisthotic is a relatively large, concave-convex bone, the
coneave surface presented posteriorly and the convex surface
antero-laterally, and it is difficult to comprehend how it could
have been wanting in Sagemehl’s specimens of J/oxostoma succetta.
The posterior portion of the posterior process of the pterotic rests
against the antero-lateral surface of the opisthotie, near its dorsal
edge, and has there anchylosed with it, but the base of the
process does not touch the opisthotic, an opening there being
left between the two bones and giving passage to the supra-
temporal branch of the nervus vagus. The opisthotic of this fish
thus corresponds to that part of the opisthotic of Scomber and
Scorpenw that lies mesial to the lateral occipital ridge, this.
accounting for the fact that it lies against the mesial, instead of
the lateral, surface of the posterior process of the pterotic. The:
posterior process of the pterotic forms the floor of the lateral
opening of the temporal groove, this floor of the opening lying
at a considerably higher level than the floor of the remainder of
the groove. The dorsal portion of the supraclavicula lies in a
somewhat vertical position, and its internal edge has a slight
antero-posterior sliding motion on the posterior portion of the
floor of the opening of the temporal groove, a curved process on
the dorsal edge of the supraclavicula hooking over the edge of
the epiotic, ine om its anterior to its posterior surface, between the
summit of the epiotic and a slight process mesial to it; the
supraclavicula thus completely closing the angular incisure in
the posterior edge of the temporal groove.

The suprasea pula of Sagemehl’s descriptions lies, as he has
stated, along the posterior surface of that part of the epiotic that
forms the posterior edge of the temporal groove, and is without
pedicel, and neither it nor the supraclavicula come into contact
with the opisthotie. The suprascapula has no pedicel, and,
furthermore, is not traversed by the main latero-sensory canal.
This bone of this fish thus cannot, as will be later further
explained, represent the whole of the similarly named bone of
Amia and most of the Teleostei, The absence of a pedicel to this
bone, together with the presence of a large opisthotic, is unusual,
for the two are usually associated ae each other, a ligament
first developing between the suprascapula and the cranium, as
above described in Lepidosteus, this ligament then undergoing
ossification at its suprascapular end, and this ossification later
inducing ossification at its point of articulation with the cranium.
In #sox, where the suprascapula has a well-developed pedicel,
the opisthotic is said to be wanting (Starks, 04), but in a single
specimen of this fish that I have examined I find it well
developed as a large membrane- bone loosely attached to the
cranium.

The temporal groove of Moxostoma lodges a muscle which,
after it issues from the groove, lies posterior to the levator

“254 MR. E. P. ALLIS ON THE OTIC REGION OF

opereuli, in contact with it, and to all appearance a part of it,
but it is inserted in part on the dorsal ‘end of the supraclavicula
and in part in the dermis that forms the dorsal corner of the gill-
opening. he distal portion of this muscle is quite certainly
innervated by the ramus opercularis facialis, and is hence a
derivative of the levator operculi, and it has acquired entrance
into the temporal groove by passing upward external te the
posterior process oP the pterotic. This suggests a secondary
rather than a primary arrangement, and leads one to Suppose
‘that the groove was primarily developed i in relation to an invasion
by the trunk-muscles, and that the dorso-mesial portion of the
muscle that actually occupies the groove has, possibly, been
derived from a portion of the trunk-muscles that was cut off
from the parent muscle by the peculiar attachment of the
ssupraclavicula. In favour of this assumption are the facts that
this dorso-mesial portion of the muscle seems to be in a somewhat
disintegrated condition, that the groove here contains considerable
fatty tissue, and that, in certain other fishes (Amia, Allis, ’97,
p. 567), the levator operculi does actually invade a temporal
groove already occupied by an anterior extension of the trunk-
muscles. Furthermore, the position of the epiotic, somewhat
removed from the dorso-postero-lateral corner of the cranium,
would seem to indicate that the trunk-muscles had, in the
ancestors of this fish, occupied the temporal groove. It is,
however, to be noted that, even in a 57 mm. specimen of
Catostomus occidentalis, which I have examined in serial sections,
this muscle has no slightest connection with the trunk muscles.
In Ameiurus, MeMurrich ('84) describes a cavity enclosed
between the pterotic, epiotic, and supraoccipital bones, which he
‘says 1s, in all probability, a rudiment of the temporal fossa of
Sagemehl’s descriptions of Amia. It is said to contain only fatty
tissue, and its opening is said to be almost closed by the supra-
clavicula. The supraclavicula is said to be ‘a T-shaped bone, of
which the upper portion of the transverse limb articulates with
the pterotic and epiotic, and almost occludes the opening of the
temporal fossa, while the extremity of the vertical limb articulates
with the side of the basioccipital.” I have shown, in an earlier
work (Allis, 04), that the superficial portion of the transverse
limb of this bone is traversed by the main latero-sensory canal,
‘and this led me to call the entire bone the suprascapula. The
bone has a mesial limb which was said by me to lie in a vertical
plane, its anterior surface resting against the hind edge of the
«lorsal surface of the cranium, and its ventral edge partly upon
the epiotic and partly upon a bone which I considered to be a
‘greatly reduced parietal. The relations, to the cranium, of this
mesial limb of this bone of Ameiwrus thus strongly resemble
those of the dorsal end of the supraclavicula of Jfoxostoma,
provided that the parietal bone of my descriptions of Ameiurus
is the so-called suprascapula of Movostoma, which seems highly
probable. It must then be that the suprascapula of dma and

THE CHONDROURANIUM OF LEPIDOSTEUS. 255»

most of the Teleostei is composed of two components, one of
latero-sensory and the other of membrane origin, that these two:
components are found separated from each other in both MJoxo-
stoma and Ameiwrus, and that the latero-sensory component
persists as an independent, superficial, dermal ossicle in Moxostoma,
but has fused with the supraclavicula in Ameiwrus. The temporal
fossa of Ameiurus would then be strictly similar to that of
Moxostoma, but greatly reduced in size, the muscle, or muscles,
that primarily invaded and occupied it having been entirely
excluded from it by the progressive utilization of the fossa as an
articular cavity for the dorsal end of the supraclavicula. In
Macrones, the temporal fossa is apparently much more strongly
developed than in Ameiurus, and Bridge and Haddon (793).
definitely say that it serves as an articular cavity for what they
call the ascending process of the post-temporal, but which is, in
fact, the dorsal end of the supraclavicula. In Silurus glanis, I
find conditions strictly similar to those in Macrones, and they
are indicated in Jaquet’s (98) figures of this fish.

In Ameturus the adductor mandibule has invaded the dorsal
surface of the cranium by passing upward over the ridge of the-
lateral semicircular canal and external to the main latero-sensory
canal, and it occupies what corresponds to the region of the
primitive supraotic depression. In Silwrus it is the trunk-muscles
that invade this region of the dorsal surface of the cranium, but
they here pass mesial to the summit of the epiotic bone, thus
traversing what corresponds to the mesial, instead of the lateral,
one of the two branch depressions at the hind end of the supra-—
otic depression of Lepidosteus.

In Conger conger there is a very definite supraotic depression,
but there is no vestige, even, of a branch depression leading
from it, either laterally or posteriorly, between the pterotic and
epiotic. The adductor mandibule has here invaded the dorsal
surface of the cranium, as in Ameiuwrus, by passing upward over
the ridge of the lateral semicircular canal and external to the
main latero-sensory canal.

In embryos of all of these teleosts there undoubtedly is a
supraotic depression similar to that im embryos of Lepidosteus,
for both Parker (73) and Gaupp (’05) show such a depression in
embryos of Salmo. In Parkev’s fifth stage of Salmo (fry of the
second week after hatching) there is as ‘yet no noticeable post--
orbital process, but there is said to be, on the ventral surface of
the ridge of the lateral semicircular canal, a slight ridge which
‘“‘ sets bounds externally to the facet for the extended head of the
hyomandibular.” The conditions in embryos of the Holostei and
Teleostei are thus probably strictly similar, but in the Holostei
the pterotic portion of the spheno-pterotic ridge persists as.
cartilage, while in the adults of the Teleostei above considered
the entire ridge is of bone, and this bone has so-called primary
‘relations to the chondrocranium. ‘The ridge of the adult teleost.
thus overlies the ridge of embryos, and is a ridge of the dermo--

“2.56 MR. E. P. ALLIS ON THE OTIC REGION OF

cranium instead of the chondrocranium, excepting, possibly, in
fishes such as Moxostoma, where the latero-sensory ossicles have
not, in this region, fused with the underlying bones. ‘There is,
in all of these fishes in which there is an opisthotic bone, a more
or less developed opisthotic ridge, which runs antero-ventrally
from the summit of the suprascapular process of the opisthotic
towards the facialis opening of the trigemino-facialis chamber,
and it must, primarily, have ended, anteriorly, in the dorsal edge
of that opening. The vena jugularis, after it issues from the
facialis opening of the trigemino-facialis chamber, runs posteriorly
ventral to the opisthotic ridge, thus lying considerably ventral to
the spheno-pterotic 1idge and in no relation whatever to it.
The subtemperal fossa, when present, lies dorsal to the opisthotic
ridge, and the adductor and levator opereuli, and the levatores
arcuum branchialium, have thei origins either on, or dorsal to
the latter ridge, the surface of origin lying, in the Teleostei,
in part on the secondarily developed posterior process of the
pterotic.

In the Selachii the conditions are somewhat different from
those above described, and this is unquestionably related to the
fact that the hyomandibula of these fishes has been developed
from the epihyal, instead of from the branchial-ray hars of the
hyal arch. Apparently related to this, the lateral semicircular
-eanal lies lateral to the auditory vesicle (sacculus ¢?) and in a
nearly horizontal position.

In Acanthias, both Gegenbaur (72) and Wells (17) show, on
‘the dorsal surface of the chondrocranium, ridges which are said
to mark the positions of the anterior and posterior semicircular
canals. Between these two ridges, mesially, and the dorso-lateral
-edge of the chondrocranium laterally, there is a slight depression
which, according to Wells’s figures, forms the surface of insertion
of a portion of the trunk-muscles, but | find it giving insertion
to those muscles in its posterior portion only, its anterior
portion being occupied by a tough ligamentous structure which
has the appearence of being a tendinous anterior extension
-of the trunk-muscles, but is not actually a tendon of those
muscles. This depression is thus a supraotic depression similar
to that in the Holostei and Teleostei, but it corresponds to the
mesial portion, only, of the depression of the latter fishes, for
the dorso-lateral edge of this portion of the chondrocranium lies,
in Acanthias, considerably dorsal to the ridge of the lateral
semicircular canal, separated from it by a large concave surface
which lies on the lateral surface of the chondrocranium and gives
insertion to the levator maxille superioris (Csd,) and the muscle
Csd,. The lateral edge of the supraotic depression forms, as above
stated, the dorso-lateral edge of the chondrocranium, it begins
anteriorly at the summit of the postorbital process, and the main
infraorbital latero-sensory canal lies superficial and slightly mesial
to it. This edge of the depression, which is called by Wells the
-supraotic crest, 1s therefore certainly the homologue of the spheno-

THE CHONDROCRANIUM OF LEPIDOSTEUS. 257

pterotic ridge of the Holostei and Teleostei, notwithstanding that
it does not lie along the ridge of the lateral semicircular canal.

The hyomandibula is said by Wells to articulate with the
““ postero-lateral surface of the cranium, in the region of the
horizontal semicircular canal,” and she shows it, in her figures,
lying much farther posteriorly than Gegenbaur shows it, and
than I find it in two specimens that I have examined. She
shows it apparently directed dorso-posteriorly, while I find it,
when the palato-quadrate 1s put in the position shown in her
figure 5, directed dorso-anteriorly, with its dorsal end partly
hidden internal to the palato-quadrate. Its articular facet lies
ventral to the ridge of the lateral semicircular canal, and external
to the ventral portion of the auditory vesicle (sacculus ?), a slight
ridge marking its dorsal edge. The vena jugularis lies between
this latter ridge and the summit of the ridge of the lateral
semicircular canal, and traverses a short canal through the
postorbital process. The’ posterior opening of this canal lies
immediately ventral to the anterior end of the ridge of the lateral
‘semicircular canal, and is apparently not shown in Wells's figures.
The ridge of the lateral semicircular canal thus forms the dorsal
edge of the groove that lodges the vena jugularis, and in this
corresponds, as will be seen later, to the opisthotic ridge of
Polypterus. The levator maxille superioris has its insertion, in
my specimen, on that part of the palato-quadrate that lies pos-
terior to the so-called palatine process of Wells's figure, instead
of anterior to that process, as shown by her, and no part of the
adductor mandibule has its msertion near the anterior end of
the palato-quadrate.

In Heptanchus, the conditions, as shown in Gegenbautr’s (’72)
figures, are sunilar to those in Acanthias, but the aa cemior por zo
of the spheno-pterotic ridge has undergone special development
to form an articular facet for the dorsal edge of the palato-
quadrate.

In a prepared skull of a 10cm. embryo of J/ustelus, I find the
dorso-lateral edge of the chondrccranium formed by the ridge of
the lateral semicircular canal, and between this ridge and the
ridges of the anterior and posterior semicircular canals there is
‘a marked supraotic depression. The postorbital process is but
slightly deveioped, and is hardly recognizable. In adult specimens
of this fish the postorbital process 1s strongly developed, as shown
in Gegenbauv’s (72) figures of this fish and of the closely related
Galeus, and a spheno-pterotic ridge runs postero-mesialiy from
the outer end of this process to the summit of the ridge of the
lateral semicircular canal, and there vanishes ; the spheno-pterotic
ridge of this fish thus forming the dorso-latera] edge of the larger
anterior portion of the otic portion of the chondrocranium, but
the posterior portion of that edge being formed by the ridge of
the lateral semicircular canal. The Hvoniantelily ui articulates
with the lateral surface of that part of the otic capsule that
lodges the ventral portion of the auditory vesicle, and there is a

258 MR. E. P. ALLIS ON THE OTIC REGION OF

sharp ridge forming the dorsal edge of its articular facet. The
vena jugularis lies dorsal to this latter ridge, in a groove which.
has a slight ridge along its dorsal edge. Dorsal to this groove,
between it and the dorso-lateral edge of the chondrocranium,.
the levator maxille# superioris and the muscle Csd, have their in-
sertions. The trunk-muscles have invaded the supraotic depres-
sion, and occupy the whole of it. The main infraorbital latero-
sensory canal lies superficial to the lateral portion of the supraotic
depression.

In the Selachii, there is thus, as in the Holostei and Teleostei,.
a spheno-pterotic ridge which is developed wholly independently
of the ridge of the lateral semicircular canal, but may lie, in part,.
along that ridge. The sphenotic portion of the ridge is developed
in relation to the postorbital process, and in Heptanchus gives.
articulation to the otic process of the quadrate. No portion of
the ridge has any relations whatever to the articular facet for
the hyomandibula.

In embryos of Polypterus, Budgett (02) shows a supraotic:
depression similar to that in Lepidosteus, and, as in that fish, it
does not become a functional temporal fossa. The lateral edge
of the depression is formed by the pronounced ridge of the
lateral semicircular canal, this ridge being called by Budgett the-
pterotic ridge. The anterior portion of this ridge is said to give:
off the ‘“ sphenotic wing,” which forms a pronounced postorbital.
process. In a 75mm. specimen of this fish I find the dorso-
lateral edge of the postorbital process forming a pronounced
sphenotie ridge, which vanishes posteriorly along the dorso—
lateral edge of the anterior portion of the ridge of the lateral
semicircular canal. Posterior to this ridge, another ridge begins.
along the same edge of the ridge of the lateral semicircular canal,
and there forms the dorsal edge of the articular facet for the
hyomandibula, this ridge evidently being developed in definite
relation to the facet. Posterior to the facet, this ridge vanishes,,
and the dorso-lateral edge of the chondrocranium is there formed,
at first by the ridge of the lateral semicircular canal, and then
by a ridge which lies at first along the lateral surface of the ridge
of that canal, but, farther posteriorly, rises gradually to the level
of the dorsal edge of that ridge. Ventral to this ridge, a low
and rounded ridge marks the position of the lateral semicircular
canal, and the vena jugularis lies immediately ventral to it,
between it and the swelling of the bulla acustica.

In the adult Polypterus there is a strongly developed opisthotie-
ridge, which begins, anteriorly, at the dorsal edge of the facialis
opening of the trigemino-facialis chamber and runs posteriorly to
the summit of the process-like, dorso-postero-lateral corner of the:
chondrocranium. It forms a projecting roof to the groove that
lodges the vena jugularis, and the articular facet for the
hyomandibula lies immediately dorsal to it and partly on its
dorsal surface, the ridge thus, in a measure, forming the ventral
edge of that facet. The dorsal edge of the latter facet is a sharp.

THE-CHONDROCRANIUM OF LEPIDOSYRUS. 959

and strongly defined ridge, the anterior portion of which lies on
the chondrocranium and the posterior portion along the lateral
edge of the dermal parieto-pterotic. When this latter bone is
removed, that part of the ridge that lies on the chondrocranium
is seen to vanish posteriorly along the dorsal surface of the
opisthotic process, the pterotic and opisthotic ridges of the
chondrocranium thus being there confluent. The sphenotic
portion of the spheno-pterotic ridge is not well defined in the
region of the postorbital process, but, anterior to that process, an
anterior continuation of it is formed by the lateral edge of that
Jong, anteriorly projecting, process-like portion of the postfronto-
sphenotic that is shown in Traquair’s (’71) figures of this fish.
Between this process-like portion of the postfronto-sphenotic and
the sphenoid bone, there is a large supraorbital fontanelle, roofed
by the frontal, and the musculus temporalis has its origin on the
ventral surface of this roofing portion of the frontal bone. The
postorbital bone is attached both to the anterior end of the
process-like porticn of the postfronto-sphenotic and to an ad-
jacent angle on the lateral edge of the frontal, the posterior
margin of the orbit thus lying considerably anterior to the
postorbital process of the chondrocranium.

The conditions in a few of the higher vertebrates may now be
considered, the supraotic depression, which is found more or less
developed in all of them, being left out of further consideration.

In a 29mm. embryo of Rana fusca, Gaupp (93) describes a
so-called processus oticus quadrati which is said to be a short rod
of cartilage which extends from the most postero-lateral portion
of the quadrate to the most antero-laterally projecting portion of
the ridge of the lateral semicircular canal. It is said to be, as its
name implies, a process of the quadrate, but it has exactly the
position of that posterior part of the sphenotic portion of the
spheno-pterotic ridge of Veit’s 203 mm. embryo of Lepidosteus
that lies lateral to the recessus dorsalis spiracularis. In an older
specimen of Rana, at a stage which represents the end of the
period of metamorphosis, a strong and overhanging ridge has
developed along the outer surface of the ridge of the lateral
semicircular canal, at the poimt where the processus oticus
quadrati had previously fused with the latter ridge. Gaupp calls
this overhanging ridge the crista parotica, and the so-called
processus oticus quadrati forms a connection between the outer
edge of this crista and the postero-lateral corner of the quadrate.
In a still older specimen, a young frog immediately after the
metamorphosis, the crista parotica is said to be much more
strongly developed, and to form a sort of roof to the tympanic
cavity (Paukenhohle); and, because of this, Parker is said to
have called this ridge the tegmen tympani. The processus
oticus quadrati is said to have now been absorbed by the crista
parotica, and it is the body itself of the quadrate that rests
against, and is fused with, the so-formed crista parotica. The
vena jugularis lies directly beneath this crista, and the nervus

Proc. Zoou. Soc.-—i919, No. XIX. 19

260 Mik. E, Ps ALLIS ON ‘tHE OTIC REGION OF

facialis directly beneath that vein. The pars ascendens of the
quadrate, called by Gaupp the pars metapterygoideus, lies in a
nearly vertical position, and its dorsal end lies opposite that part
of the cranial wall from which, in fishes, the postorbital process
projects laterally or ventro- laterally. The columella and the
annulus tympanicus are both connected, by cartilage, with the
crista parotica, the columella lying anterior to the ramus hyoman-
dibularis facialis and hence corresponding to the anterior articular
head of the teleostean hyomandibula. ‘The posterior portion of
the hammer-shaped paraquadratuin of Gaupp’s (05) later descrip-
tions, the ossa tympanica of one of his earlier works (’96), lies
upon the outer surface of the crista parotica, the anterior portion
of this limb projecting anteriorly as the processus zygomaticus ;
this bone thus strongly vecaliing but probably not representing
the’ dermal, postfrontal portion of the postfronto-sphenotic of
Polypterus. Parker (71) calls this bone the temporo-mastoid,
and says that it “would seem to combine the supratemporal
and preopercular of the Triton, or of the Siluroid or Ganoid
Fishes.”

In embryos of Lacerta, the crista parotica is said by Gaupp
(00) to be, as in Lana, a ridge which projects ventro-laterally
from the most projecting portion of the ridge of the lateral
semicircular canal, but this projecting portion of the latter ridge
lies in its posterior portion, instead of, as in Jana, near its
anterior end. On the crista parotica there is said (1. ¢. p. 451)
to be an independent, anteriorly directed process, the summit
of which lies internal to, and closely against, the hind edge of
the dorsal end of the quadrate. Gaupp calls this little process
the processus paroticus, and says that a thin strand of tissue,
composed of closely agglomerated cells, extends forward from its
summit and is continuous with a sharp ridge which lies dorsal to
the columella. his latter ridge is shown, in the figures given,
lying dorsal also to the foramen faciale, ina position that suggests
a remnant, or primordium, of the lateral wall of the pars ganeli-
onaris of the trigemino-facialis chamber of fishes, which wall is,
otherwise, wanting in this reptile. The thin strand of closely
agglomerated cells above referred to, then possibly represents a
part of the sphenotic ridge of fishes, and hence also the so-called
processus oticus quadeati. of Gaupp’s descriptions of Rana.

The processus paroticus of Lacerta is said by Gaupp (0. ¢. p. 463)
to have been primarily independent of the crista parotica and
continuous with the columella auris, and only in later stages of
development to have become independent of the columella and
completely fused with the crista parotica. In embryos of
Crocodilus biporcatus, the processus paroticus of Gaupp’s deserip-
tions is called by Shino (14, p. 325) the processus dorsalis of the
columella, and it is said by him to never become either detached
from the columella or fused either with the crista parotica or the
quadrate. In Crocodilus porosus this process is said to fuse with
the quidrate. he quadrate of Lacerta does not fuse with the

THE CHONDROCRANIUM OF LEPIDOSTEUS. 261

eranial wall. In Crocodilus biporcatus it fuses with the otic
capsule at a point which corresponds exactly to the point of
origin of the postorbital process of fishes, and it is separated
from the anterior end of the crista parotica by a considerable
interval, The process formed, in Lacerta, by the fusion of the
processus paroticus with the crista parotica is said by Gaupp
(J. c. p. 519) to quite certainly be the homologue of the processus
styloideus of Mammals; and the vena jugularis and the nervus
facialis both run posteriorly beneath this process and the erista.

In embryos of the rabbit, the crista parotica is said by Voit
(09, pp. 449-451) to be a tall and plate-like ridge which projects
ventro-anteriorly from the anterior edge of the ridge of the
lateral semicircular canal, the anterior edge of the latter ridge
of the rabbit corresponding to the ventral edge of the ridge of
fishes. Posterior to the crista parotica there is said to be a
_ depression (Grube), the lower part of which is particularly deep
and forms a fossa subarecuata lateralis which sinks into the massa
angularis. This fossa must accordingly lie between the crista
and the ridge of the lateral semicircular canal, for otherwise it
could not sink into the massa angularis. The figures given
favour this interpretation of it,and it would seem to be confirmed
by the conditions in embryos of Hehidna, where Gaupp (08)
shows, in figures of transverse sections, the crista parotica arising
from the lateral wall of the otic capsule considerably ventral to
the ridge of the lateral semicircular canal and separated from it
by a slight depression. The upper (anterior) end of the crista of
the rabbit is said by Voit to “encounter” the lower (posterior)
end of the tegmen tympani, and then to be continued onward in
a slight ridge, the crista facialis, which ends anteriorly in the
dorsal edge of the foramen by which the nervus facialis issues
from the cavum supracochieare. ‘fhe latter foramen is called
the foramen faciale externum s. secundarium, and its external
opening is called the apertura tympanica canalis facialis. The
dorsal end of the hyal cartilage (Reichert’s cartilage) fuses with
the internal surface of the anterior (morphologically ventral)
edge of the crista parotica to form the processus styloideus.
Posterior to the latter process, the processus mastoideus arises
from the outer edge of the crista parotica. Beneath the crista
facialis there is a groove, the sulcus facialis, which lodges the
nervus facialis after it issues from the apertura tympanica canalis
facialis, and this sulcus is continued posteriorly in the angle
between the crista parotica and the lateral wall of the otic
capsule. The crista parotica of this mammal thus has the
general position and relations of the opisthotic ridge of fishes,
vather than those of any part of the spheno-pterotic ridge, and
its anterior prolongation, the crista facialis, certainly does not
represent any part of the latter ridge.

The tegmen tympani of the rabbit is said by Voit (l.c. p. 448)
to arise from the lateral surface of the but slightly developed

prominentia utriculo-ampullaris superior, thus being related to
19%

962 MR. E. P, ALLIS ON titk OVIC REGION OF

the anterior (superior), instead of the lateral semicircular canal.
It is said to project laterally and upward as an arched umbrella-
shaped roof (“leicht gewolbtes schirmfrmiges Dach), and also to
extend forward a cer ain distance, there arching over the nervus
facialis and, beyond that nerve, fusing with the pars cochlearis
of the otic capsule, along the lateral edge of the planum
supracochleare ; thus fonmine the lateral wall of the cavum
supracochleare, the ventral portion of which is pertorated by the
foramen faciale externum s. secundarium. That part of the
tegmen that lies anterior to this foramen is said (d.c. p. 533)
to. correspond to the freely projecting processus perioticus
superior Gradenigos of certain other mammals, and to be
continued anterior fly a certain distance by procar tilaginous tissue
which later chondrifies. The commissura capsulo-parietalis fuses
with the dorsal surface of the wnbrella-shaped tegmen tympani,
and it is that part of the tegmen that lies ventro-mesial to the
line of this fusion that forms the lateral wall of the cavum
supracochleare, the part that lies lateral to the line of fusion
forming a tall ridge which projects ventro-laterally and extends
posteriorly slightly beyond the anterior end of the crista parotica,
where it curves ventrally to meet the basal portion of that crista.
This ridge, alone, is frequently referred to in the text as the
tegmen tympani, and it is it alone that is so designated, in
the figures, by the index letters. The fossa that les between
this ridge and the crista facialis is called the fovea epitympanica,
and its posterior portion is said to be deepened to form the fossa
incudis, this latter fossa lying, as shown in the figures, on the
external surface of the otic capsule, while the anterior portion of
the fovea epitympanica les upon the external surface of the
lateral wall of the cavum supracochleare. The tegmen tympani
is said to be a direct anterior prolongation of the crista parotica,
buf one is in doubt as to whether this applies to that part of the
tégmen that forms the mesial wall of the fossa epitympanica
or the part that forms its roof. The crista facialis is also said, as
above explained, to be a direct anterior prolongation of the crista
parotica, and yet this crista is simply a ridge on the external
surface of that part of the tegimen that forms the mesial wall of
the fovea epitympanica.

Van Kampen (05), in an earlier work, gives a somewhat
different description of the tegmen tympani of Mammals in
general. According to him (J. ¢. p. 344), Reichert’s cartilage
fuses with a cartilaginous ridge which arises from the outer
surface of the ridge of the lateral semicircular canal. Anterior
to the point of this fusion, the ridge separates into two parts,
one of which is called by Van Kampen the crista facialis, and
corresponds to the similarly named crista of Voit’s descriptions
of the rabbit. The other part of the ridge is said to lie at a
higher level than the crista facialis, and the two ridges enclose
between them a part of the wall of the pars vestibularis of the
otic capsule which Van Kampen calls the mesial wall of the

THE CHONDROCRANIUM OF LEPIDOSTEUS. 963

recessus epitympanicus, this recess, as thus described, thus lying
wholly on the externa! surface of the otic capsule, and not partly
on the lateral wall of the cavum supracochleare. The upper
one of the two enclosing ridges forms the voof of the recessus
epitympanicus, and corresponds exactly to the roof of that recess
of Voit’s descriptions of the rabbit, and it is said that it is this
roof alone that forms, in human anatomy, the tegmen tympani.
The tegmen tympani of human embryos, as thus defined, is thus
a ridge which forms a part of the external, instead of the internal,
wall of the recessus epitympanicus, and hence is in no way
comparable to the tegmen tympani of the adult Man, where it is
a thin plate of bone which lies between the tympanic and cerebral
cavities and separates them one from the other. In Mammals
other than Man,'the recessus epitympanicus is said to be less tall,
and as, in consequence of this, the roof of the recess and the
evista facialis are not distinctly separated from each other, they
are both considered to be included in the tegmen; the tegmen
of these Mammals, as thus defined, thus corresponding to the
tegmen of the rabbit as described by Voit, and forming
the mesial wall and part of the external wall of the recessus
epitympanicus. The crista facialis alone is said by Van
Kampen to probably be the homologue of the crista parotica of
Lacerta, and Gaupp (08, p. 687) accepts this as correct. Gaupp
also further says that the tegmen tympani is a new formation
which first appears in Mammals, but that it is wanting in Mchidna;
and yet he shows, in one of his figures (l. c. fig. 44, p. 628), a
portion of the erista parotica which has, to the incus, exactly the
relations of that part of the tegmen tympani of both Voit’s and
Van Kampen’s descriptions that forms the roof of the recessus
epitypanicus. Van Kampen, in the figure of a new-born Ovis,
shows the tegmen tympani as a right-angled ridge the anterior
portion of which forms the roof of the fossa muscularis major
and ends in the transverse plane of the hiatus facialis. The
anterior portion of the tegmen is said by Van Kampen to be
very variable, and to often be wholly wanting, and in Man it is
said to arise, not as a ridge, but as an anteriorly projecting
process, the processus perioticus superior Gradenigos. The
rocess lies, as shown in the figure given (/.c. fig. 3, p. 342),
definitely anterior to the recessus epitympanicus, and anterior
also to the column of cartilage that forms the anterior boundary
of the foramen faciale externum s. secundarium; and the
ossification of this process and the layer of connective tissue
that separates it from the pars cochlearis of the otic capsule is
said to form the tegmen tympani of the adult. The mesial wall
and the roof of the recessus epitympanicus of embryos thus,
neither of them, enters into the tegmen tympani of the adult.
There is thus some want of accord and precision in the
descriptions of the tegmen tympani by these authors, and this
has in a measure been perpetuated in later descriptions. Fawcett,
for example, says (717, pp. 319-321) that the tegmen tympani of

264 MR. BE, P. ALLIS ON THE OTIC REGION OF

Microtus is a curved shell-like cartilage which projects forward
from the base of the triangular lateral surface of the otic capsule,
‘‘over the incus and malleus cartilages”; that the recessus
epitympanicus has been formed by the hollowing out of the
under surface and root of the anterior prolongation of the
tegmen tympani; and that the posterior portion of the latter
recess has been deepened to form a fossa incudis which lies on
the ‘upper aspect” of the evista parotica. The tegmen tympani
of this mammal, as thus described, would thus seem to be
nothing more than the ridge that forms, in Voit’s descriptions of
the rabbit, the roof of the fovea epitympanica; and it would
seem as if it could not be an anterior prolongation of the crista
parotica, for its posterior portion, which forms the dorsal
boundary of the fossa incudis, must he either on the external
surface of the crista parotica, or on the external surface of the
otic capsule immediately dorsal to that erista. In Hrinaceus
the tegmen tympani, as described by Fawcett (18), also forms
the roof of the recessus epitympanicus, but it would here seem to
be a direct anterior prolongation of the crista parotica. Terry
(17) does not consider that part of the ridge of these descriptions
that forms the roof of the fossa incudis to be a part of the tegmen
tympani, for he says (/. ¢. p. 300) that the tegmen tympani is
not present in a 23:1 mm. embryo of the cat, and yet he shows,
in his figure 3, a well-developed fossa incudis which is bounded
dorsally by a strongly marked but not projecting ridge which is
evidently the homologue of the posterior portion of the roof of
the fovea epitympanica of Voit’s, Van Kampen’s, and Fawcett’s
descriptions. The mesial wall of the fossa incudis is apparently,
and correctly, not considered by either Fawcett or Terry to form
part of the tegmen tympani.

Comparing these conditions in Mammals with those in Fishes,
it is evident that that part of the tegmen tympani of Voit’s
descriptions of the rabbit that forms the lateral wall of the cavum
supracochleare, together with its anterior procartilaginous pro-
longation, is the post-trigeminus portion of the lateral wall of the
pars ganglionaris of the trigemino-facialis chamber of Fishes.
That so-called part of the tegmen that forms the roof of the
fovea epitympanica must then be the sphenotic portion of the
spheno-pterotic ridge of Fishes. The incus, which lies immediately
ventral to this roof, then has the relations to it that it normally
should, if,as I have lately endeavoured to show (Allis, in press), it
is the homologue of the otic process of the quadrate of Heptanchus,
of the corresponding part of the quadrate of the Amphibia and
Reptilia, and of the lateral wall of the pars jugularis of the
trigemino-facialis chamber of Fishes. The crista facialis certainly
corresponds to the anterior portion of the opisthotic ridge of
Polypterus, and the crista parotica is usually shown as a direct
posterior continuation of this crista, and not of the roof of the
fovea epitympanica. The crista parotica would thus seem to cor-
respond to the posterior portion of the opisthotic ridge of Fishes,

i la Nl tl

THE CHONDROCRANIUM OF LEPIDOSTEUS. 265

rather than to the pterotic portion of the spheno-pterotic ridge.
The relations of the processus zygomaticus of Man to the region
of the roof of the fovea epitympanica at is to the sphenotic
ridge—would therefore seem to indicate that it has been derived
from, or at least contains, the long, anteriorly projecting process
of the postfronto-sphenotic of Polypterus; and the temporal
fossa of Man would arise if that part of the frontal of Polypterus
that roofs the supraorbital fontanelle were to be suppressed by
the wearing action of the musculus temporalis. That angle of
the frontal of Polypterus that gives articulation to the post-
orbital bone would then correspond to the external angular
process of the frontal of Man, and the postorbital bone would
correspond to the malar bone of Man; this all being in accord
with my conclusion (Allis, 00) that the maxillary bone of
Polypter us is the homologue of the maxillary part of the superior
maxillary bone of Man, ‘and that neither of these bones is the
homologue of the teleostean maxillary.

Palais de Carnolés, Menton, France,
April 30, 1919.

Literature.

Autis, f. P., Jy. 1897. The Cranial Muscles, and Cranial and first Spinal Nerves
in Amia calva. Journ. Morphology, vol. xi. No. 3.
Boston.
1900. ‘The Premayillary and Maxillary Bones, and the Maxillary
and Mandibular Breathing Valves of Polypterus bichir.
Anatom. Anz. Bd. 18. Jena.
7 1908. The Skull, and the Cranial and first Spinal Muscles and
Nerves in Scomber scomber. Journ. Morphol. vol. xviii.
No. 2. Lancaster, la., U.S.A.

1904. The Latero-sensory Canals and related Bones in Fishes.
Intern. Monatsschr. f. Anatom. u. Physiol. Bd. 21.
Leipzig.

1909. The Cranial Anatomy of the Mail-cheeked Fishes. Zoo-
logica, Bd. 22.

1918. On the Origin of the Hyomandibula of the Teleostomi.
Anat. Ree. vol. xv.

(In Press). On the Homologies of the Antitiost Ossicles and
the Chorda tympani.

Brive, T. W., 1893. The Air-bladder and Weberian Ossicles in the Siluroid
& Happon, A.C. Fishes. Phil. Trans. Royal Soc. London, vol. elxxxiv.
Bepaerr, J.S. 1902, On the Structure of the Larval Polypterus. Trans. Zool.

Soc. vol. xvi. London.

Fawoertt, E. 1917. The Primordial Cranium of Wierotus amphibius (water-rat),
as determined by Sections and a Model of the 26 mm.
Stage, with Comparative Remarks. Journ. Anat. vol. li.
pt. 4. London.

1918. The Primordial Cranium of Hrinaceus europeus. Ibid.
vol. li. pt. 2.

Gaver, ©. 1893. Beitrage zur Morphologie des Schadels. 1. Primordial-Cranium
und Kieferbogen von Rana fusca. Morph. Arbeiten, Bd. 2
Heft 2. Jena.

1896, Anatomie des Frosches (Ecker & Wiedersheim), Lehre vom Skelet
und vom Muskelsystem. Aufl. 3. Abth.1. Braunschweig.

1990. Das Chondrocranium yon Lacerta agilis. Kin Beitrag zum
Verstiindnis des Amniotenschadels. Anat. Hefte, Bd. 14.
Tal, 8)

bP)

266 ON THE OTIC REGION OF LEPIDOSTEUS.

Gavpp, H. 1905. Die Entwickelung des Kopfskelets. Handbuch d. vergl. u.
experiment. Entwickelungslehre d. Wirbeltiere von Oskar
Hertwig, 1906, Bd. 3. Teil 2.
i 1908. Zur Entwickelungsgeschichte und vergleichenden Morphologie
des Schadels von Hehidna aculeata var. typica. Jenaische
Denkschriften, Bd. 6. T. 2. Jena.
GEGENBAUR, C. 1872. Das Kopfskelet der Selachier. Untersuchungen zur Ver-
gleichenden Anatomie der Wirbeltiere, H.3. Leipzig.
JAQuet, M. 1898. Recherches sur l’anatomie et Vhistologie du Silwrus
glanis, l. Arch. Sc. Méd. Bucarest, Tome 3.
Kampern, P.N. 1905. Die Tympanalgegend des Siugethierschidels. Morph.
VAN. Jahrb. Bd. 34.

MoMorricnu, 1884. The Osteology of Amiurws catus. Proc. Canad. Instit.
alee vol. il. fase. 3.

Parker, W.K. 1871. On the Structure and Development of the Skull of the
Common Frog (Rana temporaria, L.). Phil. Trans.
Royal Soc. London, vol. ¢elx1.

4 1873. Onthe Structure and Development of the Skull in Salmon
(Salmo salar, lu.) Phil. Trans. Royal Soc. London,
vol. elxiit.

P 1882. On the Development of the Skull in Lepidosteus osseus.
Phil. Trans. Royal Soc. London, vol. elxxiii.

Rrprwoop, W.G. 1904. On the Cranial Osteology of the Fishes of the Families
Flopidx and Albulide, with Remarks on the Morphology
of the Skull in the Lower Teleostean Fishes generally.
Proc. Zool. Soc. London, 1904, vol. 11. p. 35.

f 1904. On the Cranial Osteology of the Clupeoid Fishes. Proc.
Zool. Soc. London, 1904, vol. 11. p. 448.

Sacpment, M. 1891. Beitrige zur vergleichenden Anatomie der Fische. 1V.
Das Cranium der Cyprinoiden. Morph. Jahrb. Bd. 17.

Surro, Kk. 1914. Studien zur Kenntnis des Wirbeltierkopfes. I. Das
Chondrocranium von Crocodilus mit Berticksichtigune
der Gehimnerven und der Kopfegefiisse. Anat. Hefte,
Bd. 50, H. 2.

Stars, E.C. 1904. A Synopsis of Characters of some Fishes belonging to the
Order Haplomi. Biol. Bull. vol. vii.

TErry, R. J. 1917. The Primordial Cranium of the Cat. Journ. Morphol.
vol. xxix. No.2. Philadelphia.

Traquair, R. H. 1871. On the Cranial Osteology of Polypterus. Journ. Anat.
Phys. vol. vy. London.

Var, ©: 1907. Uber einige Besonderheiten am Primordialeranium von Lepi-
g é p
dosteus osseus. Anat. Hefte, Bd. 33.
*y 1911. Beitrige zur Kenntniss des Kopfes der Wirbeltiere. 1. Die

Entwickelung des Primordialeranium von Lepidosteus osseus.
Anat. Hefte, Bd. 44, H. 1.

Vor, M. 1909. Das Primordialeranium des Kaninchens unter Berticksicht-
sigung der Deeckknochen. Ein Beitrag zur Morphologie des
Saiugethierschadels. Anat. Hefte, Bd. 38.

Wetts, G. A. 1917. The Skull of Acanthias vulgaris. Journ. Morphol. vol. xxviii
No. 2.

ces

ON THE SNAKES OF WEST AFRICA, 267

16. A List of the Snakes of West Africa, from Mauritania
to the French Congo. By G. A. Bounencer, F.R.S.,
F.Z.8.

[Received May 20, 1919: Read June 17, 1919.}

(Published by permission of the Trustees of the British Museum.)
(Text-figures 1 & 2.)

After an interruption of four years, due to the restrictions
imposed on the Society’s publications, | am now able to continue
the series of lists of African Snakes, accompanied by artificial
keys and arranged according to districts, with the object of
facilitating identifications *.

The present instalment deals with the western parts of Africa,
from Mauritania to the French Congo inclusively. This division
is as artificial as the keys, for we know a great many Snakes,
until lately believed to be characteristic of West Africa, to
extend across the great forest region as far as Uganda and the
north-west of Lake Tanganyika. But it is convenient for
practical purposes, especially if this list be used in conjunction
with that dealing with the Belgian Congo, to which, however,
many additions have been made since its publication. Our
knowledge of the Snakes of the interior of Africa is still so im-
perfect that attempts at mapping out the distribution must be
regarded as very provisional.

The figures, in lieu of a glossary, which have already appeared
in the ‘ Proceedings,’ ave here reproduced (text-figs. 1 & 2).

Synopsis of the Families.

IT. Worm-like, with small inferior month, eyes hidden or visible under the head
shields, and body covered with uniform imbricate scales above and beneath.

Ocular shield not bordering the mouth; tail not or but little

_ longer than broad ; 18 scales or more round middle of body .... Typmnopip®.
Ocular shield bordering the mouth; 14 scales round middle
OT NNO CV dap aarss seen nye se sev to tee Ong RRS eT ne tote se eT cla eT aces ae ORT ANT IO INGTATID SAE

IT. Mouth lar ge, eyes distance an eee noe aaah shintecd shields beneath.

Ventral shields much narrower than the body; supraocular, if

distinet, broken up into two or more shields ...... .. Bor.
Ventral shields at least nearly as broad as the body ; supraocular

single; poison-fangs, if below the eye, pigeued by smaller

teeth.. : .. CoLuBRrip®.
Ventral shields. nearly : as broad as the ‘body ; : large poison- -fangs
in a very large sheath below the CY. Cas eestor ie aaa ote ete VPRARED ai

Family TyPHLOPIDA,
A single genus.
1, TyPHLops.

Schneid. Hist. Amph. ii. p. 339; Bouleng. Cat. Sn. i. p. 7

* P. Z.S. 1915, pp. 193, 369, 611, 641.
+ Cf. Boulenger, Ann, Zool, Afr, vii. 1919, p. 1.

68 MR. G. A. BOULENGER ON 'THE

Text-figure 1.
(From P. Z.S. 1915, p. 612.)
ocular rostral nasal

‘ preeocular

. upper
~Jabial

supraocular
nasal, *

preefrontal /, |
prasocular & S=
ocular--—

TYPHLOPS PUNCTATUS.

rostral —prazfrontal rostral _nasal

supraocular nasal ; x upper

nasal-... / ocular {3 Ns PA labial
rostral | SARS eaillen

ocular.

frontal --

meal i
upper labial pi

loreal postecular
post.nasal =; preeocular..-”
ehuaesl ee __.-. ant.temporal
rostral ns
upper labial ~~

subocular x
rostral ---..... ; leweniabial mee Postral

symphysial
"lower labial

ant.nasal
internasal
prefrontal A{§
_ supraocylar’ \ fower labial

parietal aca At--ventral

CAUSUS RHOMBEATUS. JGREEN OcL

SNAKES OF WEST AFRICA. 269

Text-figure 2.

(From P. Z. 8. 1915, p. 613.)

TT Oe
UL Vo)
TRS

TOR
Wy

\\ %
as uae
sh ne

‘ ’ ue
TANYA) SR
RN 1 = :

Mi KL

W528)

iN
TREE

ventrals scales (15 rows) ventrals
iN B.
Scaling of thickest part of body.
A. Gastropyxis smaragdina, with keeled seales and bicarinate ventral shields.
B. Dipsadomorphus blandingii, with oblique scales and enlarged vertebrals.

The present list deals with Division VI. of the following
TE :

Division of Africa into seven districts.

270 MR. G. A. BOULENGER ON THE

Synopsis of the Species.

I. No subocular; ocular in contact with two or three upper labials.
A. Rostral not more than one-third width of head; nasal completely divided
into two; diameter of body 35 to 50 times in total length.

20 scales round middle of body; eyes distinct ; anterior part of

nasal extending to upper surface of head ...ccccccccee eee. ZL. braminas.
18 scales round middle of body ; eyes hidden ........ even. Li vececatus:
22 scales round middle of body ; eyes just distinguishable Lapel ae _ T. leucostictus.

B. Rostral at least half width of head.

1. Nasal completely divided into two; przeocular much narrower than the
ocular; eyes distinct.

18 or 20 scales round middle of body, the diameter of which is

36 to 49 times in total length _..... wa. T. elegans.
24 scales round middle of body, the diameter of which is 66
HOTS WN HOLA WSREANH — Gopnevooocso cnaccacsvaccooanpadencbocascccenane toh MABOIROSHIS:

2, Nasal incompletely divided ; snout (seen from above) rounded.
a. HKiyes hidden.

28 scales round middle of body, the diameter of which is

39 to 40 times in total length; przocular much narrower

than ocular... .. 1. batesii.
98 scales round middle ‘of ody, ‘the diameter of which is 19

times in total length; praeocular much narrower ,than

ocular ..... veces ZT. hallowelli.
24 scales round middle of body, the diameter of which is 53

times in total length ; preocular nearly as broad as ocular. T. buchholzi.

b. Hyes distinguishable ; 24 to 80 scales round middle of body.

Diameter of body 41 to 45 times in total length ; ae oe a

little narrower than ocular . .. TT. steinhausi.
Diameter of body 21 to 32 times in total Jength ; g “preocular
much narrower than ocular ..........c:cceccecec tee eeseeeeerseeeee D, punctatus.

3. Nasal completely divided ; preocular as large as ocular; snout (seen from
above) pointed ; eyes hidden ; tail without terminal spine.

Snout obtusely pointed; 22 scales round middle of body, the

diameter of which is 54 times in total length . Meech erie ae CHOSSts
Snout obtusely pointed; 20 scales round middle of body, ‘the

diameter of which is 51 times in total length .................. 7. fee.
Snout acutely pointed ; 22 scales round middle of body, ‘the

diameter of which is 62 to 66 times in total length ......... T. principis.

If. A subocular separates the ocular from the upper labials; eyes hidden.

18 scales round middle of body, the diameter of which is 35 to

45 times in total length; snout rounded; rostral { width

of head; no caudal spine _..... T. zenkeri.
20 scales round middle of body, the diameter ‘of which is 70

times in total length; rostral large, with obtuse horizontal

edge ; a small cauidallspiniets i)... ..0)ova ieee eames Meee aceanen: au,
29 scales round middle of body, the diameter of which is 60 to

74 times in total length ; rostral large, with coe horizontal

edge; a small caudal spime ........ T. ce@cus.
28 se ales round middle of body, the diameter ‘of: w which i is 55 5 to

70 times in total length ; rostral large, with sharp horizontal

Ed@el) Moleandal spine. pees: ene sucuaniedcneereeeeesee et ateeen ==) Un VECUCOnate

rufescens.

1. TypuLors BRAMINUS Daud.
Bouleng. Cat. Sn. 1, p. 16.
Southern Asia; islands of the Indian Ocean; South Africa ;

SNAKES OF WEST AFRICA. 971

Lagos (fide Peters); Mexico.—Distribution probably ascribable
to transport by human agency.

2. Tyentors cHcatus Jan, Icon. Gén. p. 9, 1. 3, pls. iv. & v.
fig. 2; Bouleng. t. c. p. 32.

Gold Coast.

3. TypHiops LEucosrictus Bouleng. Ann. & Mag. N. H. (7)
i, 1898, p. 124.

Liberia.

4, TypHLops ELEGANS Peters, Mon. Berl. Ac. 1868, p. 450,
pl. ui. fig. 1; Bouleng. Cat. Sn. 1. p. 37.

Prince’s Id., Gulf of Guinea.

5. Typunops pecorosus Buchh. & Peters, Mon. Berl. Ac. 1875,
(ds NOs

Cameroon.

6. TypHtors BArEsit Bouleng. Ann. & Mag. N. H. (8) vii.
1911, p. 371.

Cameroon.

7. TYPHLOPS HALLOWELLI Jan, Icon. Gen. p. 29, 1. 4, pls. iv. & v.
fig. 6; Bouleng. Cat. Sn. 1. p. 40.

Gold Coast.

8. TypuLors BucHHOLZL Peters, Sitzb. Ges. Nat. Fr. Berl. 1881,
p. 71; Bouleng. Cat. Sn. i. p. 41.

Cameroon.

9. TyPHLops sTEINHAUSI Werner, Jahrb. Hamb. Wiss, Anst.
xxvi. 2, 1909, p. 209.

Cameroon.

10. TypHLops punctatus Leach.

Bouleng. Cat. Sn. i. p. £2.

Typhlops bocagii Bethencourt Ferreira, Jorn. Se. Lisb. (2) vii.
1904, p. 114.

Typhlops adolphi Sternt. Mitt. Zool. Mus. Berl. v. 1910, p. 70.

¢ Typhlops dubius Chaban. Bull. Mus. Paris, 1916, p. 364, fig.

Tropical Africa, as far north as the Gambia and Uganda.

11. Typutors cross1 Bouleng. Cat. Sn. i. p. 52, pl. iii. fig. 5.

Southern Nigeria.

12. Typutors Fea bouleng. Ann. Mus. Genova, (3) 11. 1906,
p. 209, fig.

San Thome Id., Gulf of Guinea,

13. TypuLops principis Bouleng. |. c. fig.
Prince’s Id., Gulf of Guinea.

YTz MR. G@. A. BOULENGER ON THE

14. Typnnops zenkeri Sternf. Sitzb. Ges. Nat. Fr.

Ds o2.

Typhlops vermis Bouleng. Aun. & Mag. N. H. (8) xiv. 1914,

p. 482.

Cameroon.

15, TypHiops RUFESCENS Chaban, Bull. Mus. Paris, 1916, p. 375,

fig. (1917).

French Congo.

16. TyPHuors caicus.

Onychocephalus cecus A. Dum. Rev. et Mag. Zool, 1856, p. 462,

pl. xxi. fig. 4.
Lyphlops cecus Bouleng. Cat. Sn. 1. p. 55.

Sierra Leone to Congo.

17. 'yputors Newrontt Bocage, Jorn. Se. Lisb. (2) u. 1891,

p. 61; Bouleng. 1. c.

I. das Rolas, Gulf of Guinea.

Family GLAUCONIIDA.
A single genus.
1. GQLAUCONTA.

Gray, Cat. Liz. p. 139; Bouleng. Cat. Sn. 1. p. 59.

Synopsis of the Species.

I. A single upper labial between the nasal and the ocular.

First upper labial as large as or a little smaller than lower part of
nasal; diameter of body 38 to 50 times in total length; tail
5 to 10 times as long as broad ‘

Hirst upper labial much smaller than lower . part of nasal; : ‘dia-
meter of body 55 times in total length; tail about 5 times as
long as broad .. Ne

First upper labial much) smaller than ‘lower part of nasal ; ‘dia-
meter of body 57 times in total length ; tail hardly twice as
NGrHyee AVS JOHFORIGL a dens dun voontouse bot coveveweeedans

II. Two upper labials between the nasal and the ocular.

Second upper labial twice as Jarge as first but much smaller than
the ocular ; diameter of body 42 times in total length ........

Second upper labial very large, nearly as large as the ocular; dia-
meter of body 38 to 43 times in total length

First and second upper labials small ; diameter of body 50 to 65
times in total length x

First and second upper labials small ; diameter of body. 78 times
in total length

1. GLAUCONIA NARIROSTRIS.

Stenostoma narirostre Peters, Mon. Berl. Ac. 1867, p. 708,

pl. —. fig. 2. é
Glauconia narirostris Bouleng. Cat. Sn. 1. p. 65.
Lagos, 5. Nigeria, Cameroon.

Ber). 1908,

G. narirostris.

G. boueti.

G. brevicauda.

G. sundevalli.
G. gestri.
G. bicolor.

G. gruveli.

SNAKES OF WEST AFRICA. Die

2. GuAuCcoNTA BouvETT Chaban Bull. Mus. Paris 1917, p. 9,
figs.

fo)
French Soudan.
3), (GLAUGONIA BREVICAUDA,.

Stenostoma brevicauda Bocage, Jorn. Sc, Lisb. xi. 1887, p. 194.
Glauconia brevicauda Bouleng. t. c. p. 67.
Dahomey and Ashantee.

4, GLAUCONIA SUNDEVALLI.

Stenostoma sundevalli Jan, Avch. Zool. Anat. Phys. 1. 1862,
Delo le

Glauconia sundevalli Bouleng. t. c. p. 68.

Togoland.

5, GuAUCONIA GEstrI Bouleng. Ann. Mus. Genova, (3) 11. 1906,
p. 210, fig.

Fernando Po and Cameroon.

6. GLAUCONIA BICOLOR.

Stenostoma bicolor Jan, Icon. Gen. p. 40, 1. 1, pl. v. fig. 15.
Glauconia bicolor Bouleng. Cat. Sn. 1. Pp. 69.

Gold Coast, Togoland.
GLAUCONIA GRUVELI.

Glauconia bicolor ygruveli Chaban. Bull. Mus. Paris, 1916,
p. 367 (1917)

Dahomey.
Family Bop &.
Three genera :—

Head distinct from neck, with shields above; rostral and anterior

upper labials with deep pits; subeaudals paired... . Python.
Head not distinct from neck, with shields above ; tail short, rounded

at the end, with single subcaudals .. ; . Calabaria.
Head not distinct from neck, with sm: ll scales ‘above ; “tail ‘short,

pointed, with single SUDCUG tee oa Hrye.

1a BXmEON,.
Daud. Hist. Rept. v. p. 266; Bouleng. Cat. Sn. i. p. 85.
Two species :—

2 upper labials pitted ; scales in 81-93 vows ; subcaudals 63-77
4d upper labials pitted ; scales in 538-63 rows; subcaudals 30-37

i eae P. sebe.
Breast P. regius.

* In comparing this snake with G. bicolor, M. Chabanaud has, through an over-
sight, given the length of the tail as the diameter.

974 MR. G. 4. BOULENGER ON THE

1. Pyraon SEB.
Coluber sebe Gmel. 8. N. i. p. 1118.
Python sebe Bouleng. t. c. p. 86.
Tropical and South Africa.
. PyrHon REGIUS.
ee regia Shaw, Zool. 11 p. 347, pl. xevi.
Python regius Bouleng. ths Ge Dp: 88.
aes to Niger, eastwards to the eyiben Soudan.

2. CALABARIA.
Gray, P. Z.S. 1858, p. 154; Bouleng. Cat. Sn. i. p. 92.

1. CALABARIA REINHARDTI.

Hryx reinhardti Schleg. Bijdv. tot de Dierk. i. p. 2, pl. —.
Calabaria reinhardti Bouleng. |. ¢.

Liberia to Congo, eastwards to the Ituri.

3. Eryx.
Daud. Hist. Rept. vi. p. 251; Bouleng. Cat. Sn. i. p. 122.

1. ERYX MUELLERI
Gongylophis ees Bouleng. Ann. & Mag. N. El (6) ix.
1892) p. (4.

Lryx muellert Bouleng. Cat. Sn. i. p. 128, pl. v. fig. 2.
Mauritania, N. Nigeria, Togoland, Egyptian Soudan.
Family CoLUBRIDs,

Three parallel series :—

No poison- fanes!-palletlne Geet laysolicliseaen erty. reree eee cee terest tee eie Ne) eA ipo lies
Poison-fangs Hehindisces, cee UE a ea eet oA sala METS Opisthogly pha.
Poison- fangs in EPO MR MRO EUTS UMOATS . AARON TNR! Proteroglypha.

A. Aglypha.
I. Loreal present.
A. Rostral without angular horizontal edge.

1. Pupil round ; a single anterior temporal; not more than two upper Jabials
entering the eye ; body not very slender, with not more than 165 ventrals ;
subcaudals not more than 95; scales not at all oblique; anal usually
divided (if entire, scales smooth in 17 rows).

IMO STRIVES IS sk soon cdocso cso oce spodan von sacogudandosntencncocndadsccagotoan LARA DICOROMLUISS
A single internasal; nasal semidivided ; scales in 26 rows ...... Helicops.
A single internasal; nostril between two nasals; scales in

DIOTEZOUOWS) (desonee een cee RCCL Ce eee PEE REE eee eee nme EEO CIRCE LEO OSs

2. Pupil round; a deep concavity on the side of the snout, between the
nostril and the eye; anal entire.
Scales keeled), in 23 VOWS 2.00.0... 0.0 ccccstcetcetsecesestsveresrsesvernvsers Dothrophthatmus.
Scales smooth, Tal LEZ OOD AURORE GoncisasuoodosoNnncoBson ssn boo svedovc0 LEC DRAANENE,

SNAKES OF WEST AFRICA. Oi

3. Pupil vertically elliptic; anal entire.
a. Scales smooth, vertebral row not enlarged ; subcaudals less than 75.

SCalessin’ 25607 so LOWSaet eee ee eeetee ee eee tenoncc as wana LB oodons

Seales in - THOME! oon capod ese sesncasseooo cus guaeboscmobosooeocaoscsaensecasea lMearaondlann.
. Scales of ver ae = row oe

Scales a head very distinct from neck ..... Hormonotus.

Scales keeled, yertebrals bicarinate ; anterior teeth not enlar eed, Gonionotophis.
Scales keeled, vertebrals bicarinate ; anterior teeth strongly
in Eye eA oe cng soo cen 306 oc ode Lobe0 Goo aso UnD BdULEE bee Gan eagadd ane ehacaeaba” SCILORCIAGMITG.

4,, Pupil ound two Br esd anterior eae or, if anterior temporal
single, 3 upper labials entering the eye, or ventrals more than 165.

. Scales not oblique, in 31 rows ; ventrals 245-254 ... Zamenis.
: Scales more or less oblique, at least on the satetten part of the body, in
13 to 19 rows ; body usually very slender (Tree-Snakes).
. Scales in 13 or 15 rows, of vertebral row not enlarged, laterals as long
as dorsals.
* Seales smooth.

Subcaudal shields not keeled ...... .... Chlorophis.
Subcaudal shields keeled on each side and with a notch corres-
ponding to the keel, same as on the ventrals .................. Philothamnus.

** Scales keeled.
Subcaudal shields keeled and notched on each side; a single

anterior temporal .. gonbodsno doo on03b0 weg eeombUOs acadne acbane LERRRIRD RIN:
Subcaudal shields not keeled ; s usually two ge anterior
HEmypOua Sis) eee ore : ae .. Hapsidophrys.
B. Scales in 13 or 15 rows, very spiinas Nites
shorter than dorsals; eye very large ............... Thrasops.
y. Seales in 17 or 19 rows, very oblique ; eye very
Target cnc. Danogsoun dsangoogansescncdoon JMUCMDIDOLOS:

e. Seales not silfiens, & m 15 to. 21 rows.
A single anterior temporal; ventrals shania rostral not twice

as broad as deep ...... See OA OMELlar
Two superposed anterior temporals ; 5 ventrals 125-168 |... Grayia.

B. Rostral very large, with angular horizontal ae
Hye in contact with labials: scales in 15 rows ..... . Prosymna.
Suboculars separate the eye from the labials ; scales in 19 to 25

THONG) © nocooyeqanestes:oonan.o¢e 12d doa doe bouickin seindan ene eRe Se bpoDapsOnseca | | AAONMOTO Dob.

II. No loreal, nasal in contact with procular.
Hye small, with round pupil; no internasals, prefrontal single ;
SOULS) SHON OKOLEN AS WH WS) THONGS, <n bes ase ane dupeeo nen sagederooanocnestacne — LSCACHUOYANOLNIS
Hye moderate or rather large, with vertical pupil; a pair of
internasals and a pair of prefrontals ; scales strongly keeled,
IN ZO\GO27 LOWS Awe, coeeeen ce emaeassne meee. oaks. Serer em neem. eS MelGIss

1. TROPIDONOTUS.

Kuhl, Bull. Sc. Nat. u. 1824, p. 81; Bouleng. Cat. Sn. iii.
p. 192. ;
Synopsis of the Species.
I. Scales smooth; two upper labials entering the eye.
Scales in 17 rows; ventrals 117-135; anal usually entire; sub-

GAMIGRNIS FOAVS | dovsnansapecaosssusdaad Gdoaseds sesesnescuce shesepucane Ab I/THMMe IMCS.
Seales in ot rows; ventrals 125-143; anal divided; subcaudals
GB S77 rekese duce ghd gab us ReSuE eee oan en Oea eens Haines t-urpcion an SER aniegatis®
Scales in ‘9 ows ; ventrals 131-150; anal y diviged subcaudals
G)=G)5) | nsosoracenea ® : .. 1. olivaceus.
II. Scales strongly feel, in onl 27 rows ; subosnlnes sepa
the eye from the labials ........ Meee: ceepenous

Proc. Zoou. Soc.—1919, ‘No. XX. 20

276 MR. G, A. BOULENGER ON THE
1. TROPIDONOTUS FULIGINOIDES.
Coronella fuliginoides Giinth. Cat. Col. Sn. p. 39.
Tropidonotus fuliginoides Bouleng. t. c. p. 217.
Gold Coast to Congo.
co)
2. TROPIDONOTUS VARIEGATUS.
Mizodon variegatus Peters, Mon. Berl. Ac. 1861, p. 358.
Tropidonotus variegatus Bouleng. |. ¢.
Gold Coast to Cameroon.

3. TROPIDONOTUS OLIVACEUS.
Coronella olivacea Peters, Mon. Berl. Ac. 1854, p. 622.
Tropidonotus olivaceus Bouleng. t. ¢. p. 227.

Tropical Africa, from the Soudan to Namaqualand and
Southern Rhodesia.

4, Troprponorus FEROX Giinth. Ann. & Mag. N. H. (8) xii.
1863, p. 355, pl. vi. fig. F; Bouleng. t. c. p. 241.
Sierra Leone to Calabar.

2. HELICcOPS.

Wagler, Syst. Amph. p. 170; Bouleng. Cat. Sn. 1. p. 272.

1. Hexicors cenprir Bouleng. Ann. & Mag. N. H. (8) v. 1910,
Ns OA,

French Guinea.

3. HyDR&THIOPS.

Giinth. Ann. & Mag. N. H. (4) ix. 1872, p. 28; Bouleng. Cat.
Sn. 1. p. 280.

Two species :—
Scales keeled, in 25 rows; ventrals 1438-156 ; 5th and 6th or 6th

_ _and 7th upper labials entering eye... .......ceccsccecencee cer eee ees Hi. melanogaster.
Scales smooth, in 21 rows ; ventrals 154-165 ; 4th and 5th upper
labials entering eye, 6th and 7th in contact with parietal... HH. levis.

1. HyprR#TrHIoPs MELANOGASTER Giinth. Ann. & Mag. N. H. (4)
Ixe NS (2 ison. 28, pl. ie neGo. boulenes tye, paZcl
Cameroon to Congo, eastwards to the Ituri.
2. Hypre#ruiops L&avis Bouleng. Ann. & Mag. N. H. (7) xiii.
1904, p. 450.
Cameroon.
4, BoTHROPHTHALMUS.

Peters, Mon. Berl. Ac. 1863, p. 287; Bouleng. Cat. Sn. i. p. 324.

1. BorHRoPHTHALMUS LINEATUS.
Elaphis (Bothrophthalmus) lineatus Peters, |. c.
Bothrophthalmus lineatus Bouleng. 1.c¢.

West and Central Africa, from the Gold Coast and Uganda to
the Congo.

SNAKES OF WESY AFRICA. 277

5. BorHRoLycus.
Giinth. P. Z. 8. 1874, p. 444; Bouleng. Cat. Sn. i. p. 325.

1. Bornrotycus arpr Giinth. |. ¢. pl. lvii. fig. B; Bouleng.
is (Ga ]Op BIA).

eae cy albopunctatus Anderss. Bih. Svensk. Ak, xxvii.

. no. 5, 1901, p. 6, pl. i. figs. 2-4.

eee br evicandatus Anderss. t. c. p: 8.

Cameroon, Fernando Po, Ituri.

6. Boopon.

Dum. & Bibr. Mém. Ac. Se. xxiii. 1853, p. 460; Bouleng. Cat.
SVM | ane
Synopsis of the Species.
I. Subcaudals in two rows.

A. Seales in 23 (very rarely 25) rows; belly blackish brown
withithemnddleline yellowish) Sees oR virgatus.

I. Scales in 25 to 33 rows.
Parietal eae longer than distance between frontal and end of

snout; 2 (very rarely 3) upper labials entering the eye ......... B. lineatus.
Parietal gnats longer than distance between frontal and end of
snout; 3 upper ‘labials entering the eye......... .... DB. bedriage.
Parietal shields as long as distance between frontal and end of snout. B. fuliginosus.
II. Subcandals single; scales in 25 to 29 rows .........e0.00....... +B. olivaceus.

1. BooDON VIRGATUS.

Celopeltis virgata Hallow. Proc. Ac. Philad. 1854, p. 98.
Boodon virgatus Bouleng. t.¢. p. 331.

Gold Coast to Calabar; Hast Africa (°).

2. Boopon tingeatus Dum. & Bibr. Erp. Gén. vii. p. 363;
Bouleng. t.c. p. 332.
Tropical and South Africa and South Arabia.

3. Boopon BeDRIAGH Bouleng. Ann. Mus. Genova, (3) ii. 1906,
p. 211.
S. Thomé and Prinece’s Id., Gulf of Guinea.

4, BoobDon FULIGINOSUS.

Lycodon fuliginosus Boie, Isis, 1827, p. 551,

Boodon fuliginosus Bouleng. t. ¢. p. 334.

Mauritania to N, Nigeria, eastwards to the Egyptian Soudan.

5. BoOoDON OLIVACEUS.

Holuropholis olivaceus A. Dum. Rev. et Mag. Zool. 1856, p. 4€€
Boodon olivaceus Bouleng. t. ¢. p. 335.
West and Central Africa, from Nigeria and Uganda. to the

Congo.
20*

278 MR. G. A. BOULENGER ON THE

7. LYCOPHIDIUM.

Dum, & Bibr. Mém. Ac. Se. xxiti. 1853, p. 462; Bouleng. Cat.
Sn. 1. p. 336.
Synopsis of the Species.
I. Loreal separated from the eye by the praeocular. —
A. 8 upper labials.
1. Parietals considerably longer than distance between frontal and end of
snout.
2 labials entering the eye, the diameter of which equals its dis-
tance from the mouth; ventrals 174-199 ; subcaudals 32-44 ... ZL. laterale.
3 labials entering the eye, the diameter of which exceeds its dis-
tance from the mouth; ventrals 164-208 ; subcaudals 24-47... LL. capense.
2. Parietals not or but slightly longer than distance between frontal and end
of snout.
3 labials entering the eye, the diameter of which is greater than
its distance from the mouth; ventrals 164-193; subcaudals
sr Oe) Ges ooh doe doa er eBe epee pasneduu es aotataneacsqpbascuecoacas tno ccasccudy \ILy Cr ROMREUOID.
3 labials entering the eye, the diameter of which is but slightly
greater than its distance from the mouth; ventrals 188-219 ;

Rinlocemmclallsy SYA «25-5 oat naneas cbacsnunnaubacbiecasrndesnssncosseenace Ll QEMNACIMOIII
B. 7 upper labials; ventrals 178-198; subcandals 30-58 ...... LL. fasciatum.
- IT. Loreal entering the eye.
7 upper labials; ventrals 185; subcaudals 43 ........................... Ds. werneri.
8 upper labials; ventrals 225-253 ; subcaudals 66-74 ................ DL. elapoides.

1. LycopHIDIUM LATERALE Hallow. Proc. Ac. Philad. 1857,
p- 58; Bouleng. t. c. p. 338.

Gold Coast to Congo.

2. LYCOPHIDIUM IRRORATUM.

Coluber irroratus Leach, in Bowdich, Miss. Ashant. p. 494.
Lycophidium wroratum Bouleng. t. ec. p. 340.

Senegambia to Gold Coast and N. Nigeria.

3. LYCOPHIDIUM CAPENSE.

Lycodon capensis A. Smith, S. Afr. Quart. Journ. (1) no. 5, 1831,
De Tks,

Lycophidiun capense Bouleng. t.¢. p. 339.

Tropical and South Africa.

4. LycopHipium semicinctum Dum. & Bibr. Erp. Gén. vii.
p. 414; Bouleng. t. c. p. 341.

Senegambia, French Guinea, N. Nigeria; East Africa (2).

5. LiycoPHrDIUM FASCIATUM.

Alopecion fasciatum Giinth. Cat. Col. Sn. p. 196.

Lycophidium fasciatum Bouleng. t. ce. p. 342.

Sierra Leone to Gaboon, eastwards to the Congo Forest west
of Mt. Ruwenzori.

6. LycopHipiIuM WERNERI Mocquard, Bull. Mus. Paris, 1902,
p. 411.

Gaboon.

SNAKES OF WEST AFRICA, 279

7. LycopHiprumM ELApPOIDES Giinth. P. Z. 8. 1874, p. 444;
Bouleng. t. ¢. p. 343.
Cameroon.
8. HoRMONOTUS.

Hallow. Proc. Ac. Philad. 1857, p. 56; Bouleng. Cat. Sn. i.
p. 343.
1. Hormonorvus MoDESTUS.

Lamprophis modestus Dum. & Bibr. Erp. Gén. vil. p. 429.
Hormonotus modestus Bouleng. 1. c.

Goid Coast to Gaboon.
9. GONIONOTOPHIS.

Bouleng. Cat. Sn. 1. p. 323.

Synopsis of the Species.

I. Scales strongly keeled.
Loreal and prefrontal entering the eye; scales in 21 rows; ventrals

IBS Isis o ey vor NO HIS VSO). 5s Gob sac eceaodcodbos cdocoacscoenssdcowentonn (GH ISG ROREA
Loreal separated from the eye by a procular; scales in 19 rows;
ventrals 6/—Wjss sulpcaudals: 9O=040es me eeeneerecs.caceeseeee a eee G. klingii.
II. Scales rather feebly keeled, in 15 rows (19 on neck).
Loreal twice as long as deep; ventrals 167-173 ; subcaudals 67 ...... G. grantii.

Loreal as long as deep or a little longer; ventrals 210-211; sub-

CSNY ISAS ee 8 or OA Re dee OC EME do tome cate cere G. microps.

1. GONIONOTOPHIS BRUSSAUXI.

Gonionotus brussauxt Mocquard, Bull. Soc. Philom. (8) i. 1889,
p. 146, pl. 11.

Gontonotus vosst Boettg. Zool. Anz. 1892, p. 418.

Gonionotophis brussauxt & vossi Bouleng. t. ¢. p. 323.

¢ Semocephalus imsignis Coaban. Bull. Mus. Paris, 1916, p. 369,
fig.

Cameroon to Congo.

2. GONIONOTOPHIS KLINGIZ Matschie, Sitzb. Ges. Nat. Fr. Berl.
1893, p. 172; Bouleng. Cat. Sn. ii. p. 614.
Togoland.

3. GONIONOTOPHIS GRANTII.

Simocephalus grantit Gtinth. Ann. & Mag. N. H. (3) xii. 1863,

p. 361.
Gononotophis grantu Bouleng. Cat. Sn.1. p. 324, pl. xxiii. fig. 1.
N. Nigeria, Gold Coast, Togoland.

4, GoNntonotoPpHis Micrors Bouleng, Ann. & Mag. N. H. (8)
vA OL aps ot O..
Cameroon.

280 MR. G. A. BOULENGER ON THE

10. SIMOCEPHALUS.
Giinth. Cat. Col. Sn. p. 194; Bouleng. Cat. Sn. 1, p. 344.

Synopsis of the Species.

1. Scales in 15 rows on body; temporals 1+2 (varely 1+3).
A. Eye much larger than nostril.
1. Ventrals 203-255 ; subcaudals 45-70.
a. 2 labials (8rd and 4th) entering the eye.
Scales with parallel secondary keels or tubercles ; loreal present. S. capensis.
Scales with strong striation directed ogee towards the keel ;
loreal absent ......... cree S. phyllopholis.

6. 3 labials (3rd, Ath, “sth) veuteniie %the ¢ eye, Snnless 5th separated by a
subocular detached from it.

Scales strongly keeled, with strong striation directed obliquely

towards! theikeel sy cis.. sic: cseheecet cee meena Eee tise Ne MLenCL ts
SMES) TTDI SENET co casove scalcbosda coo voscovadesbesoosecensbacsocoasagens | (So UAMROGHOIO.
2. Ventrals 239-262; subcaudals 75-124; scales without
secondanvalkeelst se tree ceerereheeee ease en: oeeraceente se em Sse DOC IUSIS:
3. Ventrals 178; su Heaudale G2 eel Dau S. rostralis.
B. Kye scarcely larger than the mecca scales foxbly
keeled ; ventrals 205-228; subcaudals 49- 59 . ... WS. stenophthalmus.
If. Seales in 17 or 19 rows on oe ventrals 999-934, subcaudals 53-68.
Scales in 17 rows; temporals 2+3...........000000000 ee cts. SL Crossi.
Scales in 19 rows; temporals 1+2.....0.........c...:.c000e0.-s.. SS. riggenbacht.

1. SIMOCEPHALUS CAPENSIS.

Heterolepis capensis A. Smith, Ill. Zool. 8. Afr., Rept. pl. lv.
Simocephalus capensis Bouleng. t. ¢. p. 345.

Gaboon, E. Africa, Nyassaland, Natal.

2. SIMOCEPHALUS PHYLLOPHOLIS Werner, Zool. Anz. xxiv. 1901,
p. 301.

Cameroon.

3. SIMOCEPHALUS GUIRALI.

Heterolepis guirali Mocquard, Bull. Soc. Philom. (7) viii. 1884,
p. 145.

Simocephalus guirali Bouleng. t. ¢. p. 346.

Cameroon to Congo.

4. SIMOCEPHALUS BAUMANNI Sternf. Mitt. Zool. Mus. Berl. iv.

1908, p. 214.
Togoland.

5. SIMOCEPHALUS POENSIS.

Heterolepis poensis A. Smith, Il. Zool. S. Afr., Rept.
Simocephalus poensis Bouleng. t. ¢. p. 346.

Sierra Leone to Congo and Uganda.

6. SIMOCEPHALUS ROSTRALIS Sternf. Mitt. Zool. Mus. Berl. v.
ISIN, joe Gas

Cameroon.

SNAKES OF WEST AFRICA. 281

7. SIMOCEPHALUS STENOPHTHALMUS.
Heterolepis stenophthalmus Mocquard, Bull. Soc. Philom. (7)

xi. 1887, p. 16, pl. i. fig. 1.

Simocephalus stenophthalmus Bouleng. t. c. p. 347.
Gold Coast, Togoland, Gaboon.

8. SIMOCEPHALUS CROSSI Bouleng. Cat. Sn. iij. p. 618.
S. Nigeria.

9, SIMOCEPHALUS RIGGENBACHI Sternf. Mitt. Zool. Mus. Berl.

. 1910, p. 63.

Cameroon.
11. ZAMENIS.

Wagl. Syst. Amph. p. 188; Bouleng. Cat. Sn. 1. p. 379.
1. ZAMENIS DORRI.
Periops dorri Lataste, Le Natural. 1888, p. 227
Zamenis dori Bouleng. t. c. p. 410.
Senegal.

12. CHLOROPHIS.

Hallow. Proc. Ac. Philad. 1857, p. 52; Bouleng. Cat. Sn. ii.

dened

Synopsis of the Species.
I. No trace of ventral keels; ventrals 152-166; subeaudals
85-99... Beth Sanasonons (Oh OP DGKIS
Wt, Wi Pipeals in more or ieee eormcn ieee teen.
A. Anal divided ; scales in 15 rows.

Preocular separated from frontal ; body very slender anteriorly ;

ventrals 175-190; subcaudals WISN) oe C. heterolepidotus.
Preocular in contact with or narrowly seoaaiel from frontal ;

ventrals 150-182 ; subeaudals 90-133 .. Sogooacnnsaseseose On CRAHOUCHPOS:

B. Anal entire; is 141-162 ; subcaudals 15- 96,
SUES TH TUS TRONS ph saud edooakecasoedasdunnsasseodoacavedebonsvercanoantoneden CR IDGACIROMNIG OME
SLOOS TA, IIBTIRON Son Won acoder eoenlaccdanbodoodeadibar ncadunocawon Myooedsabencaca . Ob CEARIOGIAS,

1. CHLOROFHIS ORNATUS.

Philothamnus ornatus Bocage, Jorn. Se. Lisb. ii. 1872, p. 80.
Chlorophis ornatus Bouleng. t. ¢. p. 93.

Portuguese Guinea, Angola.

2. CHLOROPHIS HETEROLEPIDOTUS.

Ahetulla heterolepidota Giinth. Ann. & Mag. N. H. (3) xi. 1863
286.

Chlorophis heterolepidotus Bouleng, t. ¢. p. 95, pl. v. fig. 3.
Chlorophis gracilis Sternf. Mitt. Zool. Mus. Berl. v. 1910, p. 64.
Gold Coast to Angola, eastwards to the coast of Zanzibar.

282 MR. G. A. ROULENGER ON THE

3. CHLOROPHIS IRREGULARIS. .
Coluber irregularis Leach, in Bowdich, Miss. Ashant. p. 494.
Chlorophis irregularis Bouleng. t. ¢. p. 96.

Senegambia and Uganda to Damaraland and 5. Rhodesia.

4. GHLOROPHIS HETERODERMUS Hallow. Proc. Ac. Philad. 1857,
p. 54; Bouleng. t. c. p. 97.

Sierra Leone to Congo, eastwards to L. Tanganyika.

5. CHLOROPHIS CARINATUS Anderss. Bih. Sv. Ak. Hand]. xxvii.
Ts Ge Dy IMO, jou, Be

Cameroon to Congo, eastwards to the Ituri and the Stanley
Falls.

13. PHILOTHAMNUS.

A. Smith, Ill. Zool. 8. Afr., Rept.; Bouleng. Cat. Sn. 11. p. 98.

Synopsis of the Species.

J. Seales in 15 rows.
A. Subcaudals 110-155.

Temporals usually 2+2; ventrals 166-207................see
Temporals 1+2 or 2+2; ventrals 150-165 0...
Temporals 1+1; ventrals 167-190 .............

B. Subcaudals 160-175 ; ventrals 201-220 ; temporals 1+1

. Semivariegatus.
. nitidus.
. dorsalis.

me} tach gel !na} ne

Og Ae hepa edane tone ucts 2 camopuesmaticonsgn soebas ode uodbEccophed . thomensis.
II. Scales in 13 rows; ventrals 186-194; subcaudals. 143-153 ;
trerodyovorma Ws) JECT IL Gye Pr foe oe eho eaoecocdaean aon ure engeacnan acc bar .girardi.

1. PHinorHAMNUS SEMIVARIEGATUS A. Smith, op. cit. pls. lix.,
lx.; Bouleng. t.c. p. 99.
Tropical and South Africa.

2, PHILOTHAMNUS NITIDUS.

Ahetulla nitida Ginth. Ann. & Mag. N. H. (3) xi. 1863, p. 286.
Philothamnus nitidus Bouleng. t. c. p. 100, pl. v. fig. 4.

Lagos to Cameroon.

3. PHILoTHAMNUS DORSALIS Bocage, Jorn. Se. Lisb. 1. 1866,
p- 69; Bouleng. t. c. p. 101.

Gaboon to Angola.

4, PHILOrTHAMNUS THOMENSIS Bocage, Jorn. Se. Lisb. ix. 1882,
Dell sttes > (Boulenesitcegp koi

S. Thome Id., Gulf of Guinea.

5, PHILOTHAMNUS GIRARDI Bocage, Jorn. Se. Lisb. (2) 11. 1893,
p. 47; Bouleng. t. c. p. 102.

Anno Bom Id., Gulf of Guinea.

SNAKES OF WEST AFRICA. 283

14, GASTROPYXIS.
Cope, Proc. Ac. Philad. 1860, p. 556; Bouleng. Cat. Sn. 11.
p. 102.
Two species :—

Ventrals 150-174; subcaudals 129-172...............scecceeeeeeeteeeeeeere GE Smaragdina.
Wentralsslg5 lot sculicarrdalsaldO Udine eee aernaer erer sere: -eeneee ssi (Gre PRUILCUNES

1. GASTROPYXIS SMARAGDINA.

Dendrophis smaragdina Schleg. Phys. Serp. 1. p. 237.

Gastropyxis smaragdina Bouleng. t. c. p. 103.

Tropical Africa, from Sierra Leone and Uganda to the Congo
and Northern Angola. ;

2. GAsTROPYXIS PRINCIPIS Bouleng. Ann. Mus. Genova, (3) ii.
1906, p. 213, fig.

Prince’s Id., Gulf of Guinea.

15. HApsIDOPHRYs.
Fischer, Abh. Nat. Ver. Hamb. ili. 1856, p. 110; Bouleng.
Cat. Sn. ii. p. 103.
1. Hapstpopurys Lineata Fischer, t.c. p. 111, pl. i. fig. 5;
Bouleng. t. c. p. 104.
Gold Coast to Congo, eastwards to Uganda and L. Tanganyika.

16. THRASOPS.
Hallow. Proc. Ac. Philad. 1857, p. 67; Bouleng. Cat. Sn. 11.
p. 104.
Two species :—

Rostral little broader than deep; ventrals 179-215; subcaudals

ZS AVAG Wein. cece ter issele Bee meee as eee Uae centers LEM LAULGMALaRTS.
Rostral much broader than deep; ventrals 163-174; subcaudals
BIC) LS) i i eee aPeeeer eee eicis aie Le aes we eae T. batesit.

1. THRASOPS FLAVIGULARIS.

Dendrophis flavigularis Hallow. Proc. Ac. Philad. 1852, p. 205.
Thrasops flavigularis Bouleng. t. ¢. p. 105.

Sierra Leone to Congo.

2. Turasors BAtESII Bouleng. Ann. & Mag. N. H. (8) ii. 1908,
p. 92.
Cameroon.
17. RHAMNOPHIS.

Gunth. Ann. & Mag. N. H. (3) ix. 1862, p. 129; Bouleng.
Cat. Sn. fil. p. 632.

Two species :—

Scales smooth ; a pair of large shields behind the parietals ............. R. ethiops.
Scales more or less distinctly keeled on the middle of the back; no
lax serocenpiralesmield sss ste.nesneee ene eee ene ne een ery cea | rem nCeCleSOmets

284 MR. G. A. BOULENGER ON THE

1. RuAmyoputs aruiors Gunth. |. c. pl. x.; Bouleng. |. ¢.

Thrasops splendens Anderss. Bih. Sv. Ak. Handl. xxvii. iv.
UNO). iy KONE Fog le
Sierra on to Congo, eastwards to aks Tturi.

2. RHAMNOPHIS JACKSONII.

Thrasops jacksonit Giinth. Ann. & Mag. N. i. (6) xv. 1895

. 528.
4 Rhannophis jacksonii Bouleng. 1. c.

French Guinea, Gold Coast, eastwards to Uganda and
L. Tanganyika.

18. CoRONELLA.
Laur. Syn. Rept. p. 84; Bouleng. Cat. Sn. 1. p. 188.

Two species :—

Scales in 21 rows; rostral much broader than deep ..................... CO. semiornata-
Scales in 19 rows; rostral a little broader than deep .................. CG. coronata.

1. CORONELLA SEMIORNATA Peters, Mon. Berl. Ac. 1854, p. 622 ;
Bouleng. t. c. p. 195.

Zamenis tchadensis Chaban. Bull. Mus. Paris, 1917, p. 451,
fi

wie}

Soudan, Hast Africa, N. Rhodesia.

2. CORONELLA CORONATA.

Calamaria coronata Schleg. Phys. Serp. ii. p. 46.
Coronella coronata Bouleng. t. c. p. 196.
Senegal to Gold Coast.

19. GRAYIA.
Giinth. Cat. Col. Sn. p. 50; Bouleng. Cat. Sn. ii. p. 188.

Synopsis of the Species.

I. Scales in 17 or 19 rows; ventrals 143-168.
Lower anterior temporal longer than its distance from the loreal ;

7 upper labials (varely 8); subcaudals 89-102 ........ G. smythiz..
Lower anterior temporal not longer than its distance from the loreal ;
8 or 9 upper labials ; subcaudals 71-84 .....-ccsccesesecssecssceseen GE ornata.
IL. Seales in 15 rows ; ventrals 125-149.
Kye much shorter than snout; subeandals 100-128 ....................... G. tholloni.
Hye as long as snout; subcaudals 125-161 20.0.0... eee. GE cesar.

1. GRAYIA SMYTHII.

Coluber smythii Leach, in Tuckey’s Explor. R. Zaire, App..
p. 409.

Grayia smythii, part., Bouleng. 1. e. .

Gold Coast to Angola, eastwards to Uganda and L. Tanganyika.

GRAYIA ORNATA.
Macrophis ornatus Bocage, Jorn. Sc. Lisb. i. 1866, p. 67.

ie)
~t

SNAKES OF WEST AFRICA. 2

-Grayia ornata Bouleng. P. Z. 8. 1909, p. 944, fig.

Cameroon to Angola, eastwards to the Ituri and Lake
Tanganyika.

3. GRAYIA THOLLONI Mocquard, Bull. Soc. Philom. (8) ix. 1897,
p- 11; Bouleng. t. ¢. p. 951, fig.

French Congo, Katanga, Uganda, Egyptian Soudan.

4. GRAYIA CHSAR.

Xenurophis cesar Giinth. Ann. & Mag. N. H. (3) xii. 1863,
p. 357, pl. vi. fig. C; Bouleng. Cat. Sn. i. p. 288.
Cameroon to Congo, eastwards to the Ituri.

20. PROosYMNA.
Gray, Cat. Sn. p. 80; Bouleng. Cat. Sn. i. p. 246.

Two species :—

Two internasals, two prafrontals ..0.....0....0..ceseecccesteeeeesessesseres. -P: greigerti.
A single internasal, a single prefrontal SpOpUDEDE LOD ASS OobesUnameeRastedds fey HOCUEEL POS

1. PRosyMNA GREIGERTI Mocquard, Bull. Mus. Paris, 1906,
p- 466.

French Soudan.

2. PRoSYMNA MELEAGRIS.

Calamaria meleagris Reinh. Vid. Selsk. Skr. x. 1843, p. 238,
pl. 1. figs. 4-6.
Prosymna meleagris Bouleng. t. c. p. 249.

Sierra Leone to 8. Nigeria, eastwards to the Egyptian Soudan.

21. ScAPHIOPHIS.
Peters, Mon. Berl. Ac. 1870, p. 644; Bouleng. Cat. Sn. ii.
p 264.
. SCAPHIOPHIS ALBOPUNCYATUS Peters, t. c. p. 645, pl. i. fig. 4
oes l.e.
Tropical Africa, from the Soudan to the Congo.

22. PacrLorHo.is.
Bouleng. Ann. & Mag. N. H. (7) xii. 1903, p. 352.

1. PacILopHoLis CAMERONENSIS Bouleng. 1. c.
Cameroon.

23. DAsyPELris.
Wagl. Syst. Amph. p. 178; Bouleng. Cat. Sn. ii. p. 353.

Two species :—

Eye less than } length of head; scales in 23 to 27 rows................... D. scabra.
Eye g to 4 length of head ; scales in 20 to 24 rows ......................... D. macrops

286 MR. G. A. BOULENGER ON THE

1. DASYPELTIS SCABRA.
Coluber scaber Linn. 8. N.1. p. 384.
Dasypeltis scabra Bouleng. t. c. p. 354.
From Sierra Leone and Egypt to the Cape of Good Hope.
DasyPELtiIs MACROPS Bouleng. Ann. & Mag. N. H. (7) xix.
1907, p. 324.

Cameroon.
B. Opisthoglypha.

I. Eye moderate or large ; head more or less distinct from neck ; loreal present.
A. Pupil vertically clliptic ; head short, much broader than neck.
1. Subcaudals in two rows.

Vertebral scales not enlarged; scales in 19 to 23 rows; two

superposed anterior temporals... : Tarbophis.
Vertebral scales not enlarged ; scales in 17 or 19 rows; 3 a a single
anterior temporal ......... sondesbecdauusaan, ALAMO IRE
Vertebral scales enlarged ; scales in 19 to 25 YOWS woes. Dipsadomorphus.
2. Subcaudals single; scales in 17 or 19 rows............... _ _Dipsadoboa.

B. Pupil round, exceptionally vertically subelliptic.
1. Scales not oblique, in 17 or 19 rows; loreal not longer than deep.

Scales keeled ; anal entire; a single anterior temporal ............ Geodipsas.
Scales grooved in the adult; anal divided; two eee frontal

at least twice as long as broad ....... vu. Ceoelopeltis.
Scales smooth; anal divided; frontal about twice as ‘long as

JOHROP- V0 bse tanan cannes MaMa eaaGmce dod oneme Sroctrearaabbbar danse aadsnacad eT AO OOUG ADIOS.

2. Scales more or ibse Ae. in 15 or 7 (rarely 19) rows, smooth; loreal
at least 13 times as long as deep; rostral not or but little broader than

deep.
Frontal, in the middle, not or but slightly narrower than supra-
ocular ; ; a single anterior tempor ‘al. : svoonocesesnnasen — -DROMONOIS.
Frontal, in the middle, narrower than supraocular ; ‘usually two
superposed anterior (EUMDONAS cso noonovovacsacancoocanacsenave ana LESMMPUOOOOS.

3. Scales not oblique, in 19 to 25 rows, seen 3 ‘eerie at
least 13 times as long as deep: rostral at least twice

BS) lOO GIS) CIEE) Sco casconcovsy bec cascnsoue ean acerscssdonason JkeemRoyn Roan.
4. Scales very oblique, very narrow, in 19 or 21 rows,
more or less keeled ; nostril im an undivided nasal... Dispholidus.
C. Pupil horizontal; scales narrow, oblique, feebly keeled,
in 19 rows; nostril in an undivided nasal ........... .... Dhelotornis.

II. Eye small or very small; head not at all distinct from neck; noloreal ; scales
im 15 or 17 rows, not oblique.

A. Subcaudals in two rows.

Nasal in contact with rostral; 5th upper labial in contact with

__ parietal; scales in 17 rows; ventrals 173-208 ............... Calamelaps.
First labial in contact with internasal; no Jabial in contact
_ with parietal; scales im 15 rows ; ventrals 180-249 ......... Miodon.
First labial in contact with internasal; 5th upper labial in
contact with parietal ; scales in 15 rows; ventrals 296 .... Hlapocalamus.
B. Subcaudals single; one or two upper labials in contact with parietal.
First labial in contact with internasal ..... .... Polemon.
Nasal in contact with rostral ; posterior “maxillary. teeth. large
and strongly grooved ....... .. Aparallactus.

Nasal in contact with rostral ; posterior “maxillary teeth feebly
enlarged and feebly grooved ...........0..ccc0ccceeeeeeer eee Llapops.

SNAKES OP WEST AFRIGA, ase

1. TARBOPHIS.

Fleischm. Dalm. nov. Serp. Gen. p. 17; Bouleng. Cat. Sn. iii.
p. 47.

Two species :—
Scalesim 19 rows; 7 to 9iupper labials 10.0.4... eres. 2. variegatus-

Scales in 21 or 23 rows; 9 to 11 upper labials T. oblusus.

1. TARBOPHIS VARIEGATUS.

Dipsas variegata Reinh. Vid. Selsk. Skr. x. 1848, p. 249, pl. i.
figs. 15-17.

Tarbophis variegatus Bouleng. t. c. p- 51.

Leptodira probeguint Moequard, Bull. Mus. Pavis, 1902, p. 45.

French Guinea to Cameroon.

2. TARBOPHIS OBTUSUS.
Coluber obtusus Reuss, Mus. Senckenb. 1. 1834, p. 137.
Tarbophis obtusus Bouleng. t. ¢. p. 52.
Mauritania, Northern Nigeria, Egypt to Somaliland.
? =) 7 Syt

2. Lepropira.
Giinth. Cat. Col. Sn. p. 165; Bouleng. Cat. Sn. ili. p. 47.
Two species :—
Ventrals 144-180; anal entire; subcaudals 32-54 ..................... DL. hotambeia-

Ventrals 201-225; anal divided; subcaudals 94-121 ................... LE. duchesnis.

1. LepropIRA HOTAMBGIA.

Coronella hotambaia Laur. Syn. Rept. p. 85.
Leptodira hotambeia Bouleng. t. ¢. p. 89.
Tropical and South Africa.

2. LeproDIRA DUCHESNII Bouleng. Ann. Mus. Congo, Zool. 1.
EIOI, pe LOS pla tvesnse ole

Dipsadoimorphus viridis Sternf. Mitt. Zool. Mus. Berl. iii. 1908,
p. 411, fig.

Dipsadomorphus brevirostris Sternf. |. e.

Cameroon to Congo, eastwards to the Ituri.

3. DiIpsADOMORPHUS.

Fitzing. in Tschudi, Faun. Per., Herp. p. 55; Bouleng. Cat.
Sn. i. p. 99.

Two species :—

Scales in 19 rows; ventvals 236-276; anal entire; subcaudals

QOHUBY patos saoass sed pou nme eau spebeg Abi vas ouedonusoo4<saabs sateondonqeonea, | HDs YUNNAN AIS:
Scales in 21 to 25 rows; ventrals 240-289; anal divided ; sub-

caudals wat Sr. ee) cea woe eae Del Ce aura hae blandingii.

288 MR G. A. BOULENGER ON THE

1. DIPsADOMORPHUS PULVERULENTUS.

Dipsas pulverulenta Fisch. Abh. Nat. Ver. Hamb. iti. 1856,
Jo Cell joule mints tates, ale

Dipsadomorphus pulverulentus Bouleng. t. c. p. 68.

Dipsadomorphus boweti Chaban. Bull. Mus. Paris, 1916, p. 314,

he.
Sierra Leone to Congo, eastwards to the Ituri.

2. DiIPpSADOMORPHUS BLANDINGII.

Dipsas blandingu Hallow. Proc. Ac. Philad. 1844, p. 170.
Dipsadomorphus blandingit Bouleng. t. ¢. p. 77.
Senegambia to Congo, eastwards to Uganda.

4, DIPSADOBOA.
Giinth. Cat. Col. Sn. p. 182; Bouleng. Cat. Sn. 11. p. 81.

Two species :—

“Scales in 17 rows, vertebrals strongly enlarged .............................. D.wnicolor.
“Scales in 19 rows, vertebrals scarcely enlarged ............................. D. isolepis.

1. DresADoBoA UNICOLOR Giinth. op. cit. p. 183; Bouleng. 1. c.
Sierra Leone to Congo, eastwards to the Iturt.

2. DresaposoA 1soLePis Bouleng. Ann. & Mag. N. H. (7) xix.
1907, p. 329.
Cameroon.
5. GEODIPSAS.
Bouleng, Cat. Sn. i. p. 32.

1. GEODIPSAS DEPRESSICEPS.
Tropidonotus depressiceps Wern. Verh. zool.-bot. Ges. Wien,

xlvii. 1897, p. 139.
Geodipsas mapajensis Anderss. Bih. Sv. Ak. Handl. xxvii. iv.

INOS 5), WON jog UY).
Geodipsas depressiceps Sternf. Mitt. Zool. Mus. Berl. i. 1908,

p. 410.
Cameroon, Fernando Po, Ituri.

6. C@LOPELTIS.
Wagl. Syst. Amph. p. 189; Bouleng. Cat. Sn, in. p. 141.

1. C@LOPELTIS MONSPESSULANA.
Coluber monspessulanus Hermann, Obs. Zool. 1. p. 283.
Celopeltis monspessulanus Bouleng. |. c.
Borders of the Mediterranean, southwards to Mauritania,
-eastwards t) Persia.
7. RUAMPHIOPHIS.

Peters, Mon. Berl. Ac. 1854, p. 624; Bouleng. Cat. Sn. ii.
p. 144.

SNAKES OF WEST AFRICA. 289

Two species :—

Snout with angular horizontal edge, curved in profile; 2 or 3

superposed anterior temporals ; subcaudals 90-110 ............ BR. oryrhynchus.
‘Snout obtusely pointed; 1 or 2 anterior temporals; subcaudals
GAS SO i 7 ee ee eee mia tag CEPA ee, A ACO G OCNSISS

1. RHAMPHIOPHIS OXYRHYNCHUS.
Psammophis oxyrhynchus Reinh. Vid. Selsk. Skr. x. 1843,

p. 244.
Rhamphiophis oxnyrhynchus Bouleng. t. ¢. p. 146.

Tropical Africa, as far north as the Gold Coast.

2, RHAMPHIOPHIS TOGOENSIS.
Psammophis togoensis Matschie, Mitth. Deutsch. Schutzgeb.

WAG oempeeik2
Rhamphiophis togoensis Bouleng. t. c. p. 147.

Togoland, N. Nigeria.

8. Dromopuis.
Peters, Mon. Berl. Ac, 1869, p. 447; Bouleng. Cat. Sn. iii. p. 149.

Two species :—

Scales in 17 rows; ventrals 140-159: subcaudals 78-105 ......... D., lineatus.
Scales in 15 rows; ventrals 161-180; subcaudals 110-122 ......... D. preornatus.

1. DROMOPHIS LINEATUS.

Dryophylax lineatus Dum. & Bibr. Exp. Gén. vil. p. 1124.
Dromophis lineatus Bouleng. |. c.
Coast of Guinea to Egyptian Soudan and Zanzibar Coast.

DROMOPHIS PRHEORNATUS.

Dendrophis preornatus Schleg. Phys. Serp. 11. p. 236.
Dromophis preornatus Bouleng. t. ¢. p. 150.
Gold Coast to Niger.

9, PSAMMOPHIS.

Boie, Isis, 1827, p. 521; Bouleng. Cat. Sn. i. p. 152.

Synopsis of the Species. ‘

I. Rostral well visible from above; snout 14 to 2 times as long as eye; sub-
caudals 64-149.
A. Frontal narrower than the supraocular ; anal divided. °

Loreal 3 to 4 times as long as cee 5 me 9 upper labials, 5th and 6th

entering the eye. : : P. schokari.
Loreal 13 to 25 times as s long as “deep ; usually g upper - labials
4th and 5th entering the eye ............ P. sibilans.

B. Frontal as broad as the supraocular ; i orl 25 times as le as
deep ; 8 upper labials, 4th and 5th entering the eye ; anal
GIIHRS: Tare Aan Preece Per oGoc bn bretco rcnatie 38a ict acu cua aah Sane ace Warm adem APC KS

If. Rostral scarcely oss from above; snout 2 to 24 times as
long as eye; loreal 3 to 4 times as long as deep : ; 9 upper
labials, 5th and 6th entering the eye; subcandals 144-172... P. elegans.

290 MR. G. A. BOULENGER ON THE

1. PSAMMOPHIS SCHOKARI.

Coluber schokari Forsk. Descr. Anim. p. 14.

Psammophis schokari Bouleng. t. ¢. p. 157.

North Africa to Mauritania and Somaliland; Arabia and
Syria to Afghanistan and Sind.

9. PSAMMOPHIS SIBILANS.

Coluber sibilans Linn. 8S. N. 1. p. 383.

Psammophis sibilans Bouleng. t. c. p. 161.

Tropical and South Africa, Egypt.

3. PsAMMOPHIS REGULARIS Sternf. Mitt. Zool. Mus. Berl. ui.
1908, p. 412.

Cameroon and Togoland.

4, PSAMMOPHIS ELEGANS.

Ooluber elegans Shaw, Zool. iii. p. 536.
Psammophis elegans Bouleng. t. c. p. 167.
Mauritania to Niger.

10. MAcRoPROTODON.
Guichen. Explor. Sc. Alg., Rept. p. 22; Bouleng. Cat. Sn. ii.
p. 175.
1. MAacroproroDoN CUCULLATUS.
Coluber cucullatus 1. Geoff. Descr. Egypte, Rept. pp. 148, 151,
pl. viii. fig. 3.
Macroprotodon cucullatus Bouleng. 1. ce.
Mauritania, North Africa, S. Spain, Baleares, Lampedusa,
S. Palestine.
1i. DisPpHOLIDUs.
Duvernoy, Ann. Sc. Nat. xxvi. 1832, p. 150; Bouleng. Cat.
Sn. il. p. 186.
1. DisPHOLIDUS TYPUS.
Bucephalus typus A. Smith, Zool. Journ. iv. 1829, p. 441.
Dispholidus typus Bouleng. t. ¢. p. 187.
Tropical and South Africa, northwards to Portuguese Guinea
and Abyssinia.
12. THELOTORNIS.
A. Smith, Ill. Zool. 8. Afr., Rept.; Bouleng. Cat. Sn. in. p. 184

1, THELOTORNIS KIRTLANDII.
Leptophis kirtlandti Hallow. Proc. Ac. Philad. 1844, p. 62.
Thelotoriis kirtlandii Bouleng. t. e. p. 185.

Tropical and South Africa, northwards to Sierra Leone and
Uganda.

SNAKES OF WEST AFRICA. 29}.

13. CALAMELAPS.
Giinth. Ann. & Mag. N. H. (3) xvii. 1866, p. 26; Bouleng.
Cat. Sn. il. p. 245.
Two species :—

Scales in 17 rows; 6 upper labials, 3rd and 4th entering the eye ...... C. unicolor.
Scales in 15 rows; 5 upper labials, 2nd and 3rd entering the eye ...... C. fea.

1. CALAMELAPS UNICOLOR.

Calamaria unicolor Reinh. Vid. Selsk. Skr. x. 1843, p. 236,
pl. i. figs. 1-3.

Calamelaps unicolor Bouleng. |. c.

Sierra Leone to Niger; East Africa.

2. CALAMELAPS FE& Bouleng. Ann. Mus. Genova, (3) 11. 1906,
p- 214, fig.

Portuguese Guinea.

14. Mtopon.

A. Dum. Arch. Mus. x. 1859, p. 206; Bouleng. Cat. Sn. iii.
p- 249.
Synopsis of the Species.
I. Anal entire; ventrals 190-216 ...................c.:eeseeeees J, aeanthias.

IT. Anal divided.

Internasals considerably shorter than the prefrontals; nasal divided ;

VET AS 22 ON O28 warren ty ak deere eer eeeene EMCEE a na aac sere tngponen uleveoulanis:
Internasals as long as or slightly shorter than the prefrontals ; nasal

entire or incompletely divided; ventrals 214-249..... 0.000.022... M. gabonensis.
Internasals as long as or slightly shorter than the jrefrontals; nasal

dividediiventrals 18) —2)4tesseee ee er enn te meno ciecrse

1. M1IoDON ACANTHIAS.

Urobelus acanthias Reinh. Vid. Meddel. 1860, p. 229, pl. iii.
Miodon acanthias Bouleng. t. ¢. p. 250.

Gold Coast, Ashantee, Nigeria.

2. M1o0DON COLLARIS.

Microsoma collare Peters, Sitzb. Ges. Naturf. Fr. Ber]. 1881,
p. 148.

Miodon collaris Bouleng. t. c. p. 251.
Old Calabar to Angola,

3. MIoDON GAPONENSIS.

Elapomorphus gabonensis ‘. Dum. Rev. et Mag. Zool. (2)
vill. 1856, p. 468.

Miodon gabonensis Bouleng. t. c. p. 252.

Old Calabar to Congo, eastwards to the Ituri.
Proc. Zoot. Soc.—1919, No. XXJ. 21

292, MR. G, A. BOULENGER ON THE

4, MiopoN NOTATUS.

Microsoma notatum Peters, Sitzb. Ges. Nat. Fr. Berl. 1882,
p. 127

Miodon notatus Bouleng. t. ¢, p. 252

Cameroon, Congo.

15. HLAPOCALAMUS.
Bouleng. Ann.& Mag. N. H. (8) viii. 1911, p. 371.

1. ELAPocALAMUS GRACILIS Bouleng. |. ec.
Cameroon.
16. PoLEmon.
Jan, Rev. et Mag. Zool. (2) x. 1858, p.520; Bouleng. Cat. Sn.
lil. p. 253.
Two species :—

A single postocular; ventrals 221-226................sccceteette eee Be bart.
Two postoculars; ventrals 174-206 ........0......seeceeeeeeeeee reese BP. bocourti.

1. Potemon sarrut Jan, |. c.; Bouleng. t. c. p. 254.
Gold Coast, Ashantee.

2. Potemon nocourtt Moequard, Bull. Soc. Philom. (8) ix.
USD, JOo ls}

Aparallactus hagmann Gough, Zool. Anz. xxv. 1902, p. 646.
Cameroon, I'rench Congo.

17. APARALLACTUS.

A. Smith, Ill. Zool. S. Afr., Rept., App. p. 15; Bouleng. Cat.
Sn. iii, p. 255.
Synopsis of the Species.
I. Two prefrontals ; a single labial in contact with the parietal ; ventrals 139-156.
A. 6th upper labial in contact with the parietal; 2 postoculars.

Frontal not 14 times as long as broad .............:..::eeeeeeeeeeeeeeee. A dollot.
Frontal 13 times as long as ‘proad.. aetna UL Saeed wee bahesin.

B. 5th upper labial im contact qth the parietal 5 fl Aeinele postocular.
Frontal at least twice as long as broad | .........000cce eee ceeeee teste eee A. nigrocollaris.
Frontal hardly 13 times as long as broad ..............eeee eee A. roucheti.

IL. A single preefrontal; 5th and 6th labials in contact with the parietal; ventrals
152-170.

Scales smooth; frontal 14 times as long as broad.. wth eA ineatus.
Scales smooth; frontal 13 times as long as broad . oes eee, A anomalus:
Scales keeled on posterior part of body and on tail; “frontal

TEE THINGS Gis MOTD AIS |OHORNCL sas ccoscoccoven ceo ascsoouncossesceeeccnoceons ko (POGKELN

1. APARALLACT’S DoLLor Werner, Verh. zool.-bot. Ges. Wien,
lii. 1902, p. 346.

French Congo.

2, APARALLACTUS BATESIT Bouleng. Ann. & Mag. N. H. (7)
sab IO Os Oa

Cameroon.

SNAKES OF WEST AFRICA. 293

3. APARALLACTUS NIGROCOLLARIS Chaban. Bull. Mus. Paris,
1916, p. 377, fig.

French Congo.

4, APARALLACTUS ROUCHETI.

Aparallactus nigrocollaris, var. roucheti Chaban. t.c. p. 378, fig.

French Congo.

5. APARALLACTUS LINEATUS.

Uriechis lineatus Peters, Mon. Berl. Ac. 1870, p. 643, pl. i. fig. 3.

Aparallactus lineatus Bouleng. Cat. Sn. iii. p. 261.

Keta, Guinea.

6. APARALLACTUS ANOMALUS.

Uriechis anomala Bouleng. Ann. & Mag. N. H. (6) xii. 1893,
p. 273.

Aparallactus anomalus Bouleng. Cat. Sn. iii. p. 262, pl. xi. fig. 3.

Gold Coast.

7. APARALLACTUS NIGER Bouleng. Ann. & Mag. N. H. (6) xix.
1897, p. 154.

Rouleophis chevalieri Chaban. Bull. Mus. Paris, 1916, p. 379, fig.

Sierra Leone, French Guinea.

18. Kvapops.
Giinth. Ann. & Mag. N. H. (3) iv. 1859, p. 161; Bouleng.
Cat. Sn. i. p. 262.
1. Exarops mopsstus Giinth. |. e.; Bouleng. 1. c.

Aparallactus boulengeri Werner, Verh. zool.-bot. Ges. Wien,
xlvi. 1896, p. 363.

Aparallactus perafinis Werner, op. cit. xlvii. 1897, p. 404.
Liberia to Congo, eastwards to the Ituri.

C. Proteroglypha.
(Loreal absent in all the genera.)

I. Head short; subcaudals less than 95.

Scales not at all oblique; ventrals 192-230; subcandals 67-80...... Boulengerina.
Scales more or less oblique; ventrals 141-172; subcaudals 13-36... Hlapechis.
Scales oblique; ventrals 189-228; subcaudals 50-92 .................. Naia.
II. Head long, narrow, snout not broader than long ; scales very
oblique; ventrals 202-279; subcaudals 97-121 ................... Dendraspis.

1. BoULENGERINA.

Dollo, Bull. Mus. Belg. iv. 1886, p. 159; Bouleng. Cat. Sn. iii.
p. 357.

Two species :—

Rostral nearly as deep as broad; temporals 1+2; 3 upper labials in

contact with lower subocular.. a .. B. annulata.
Rostral much broader than deep ; temporals 2+2 or r2+3; 2 upper
labials in contact with lower subocular ........ yotanoaponces Jato CWROOREIEO

21*

294 MR. G. A. BOULENGER ON THE

1. BouLENGERINA ANNULATA.
Naia annulata Buchh. & Peters, Mon. Berl. Ac. 1876, p. 119.
Boulengerina annulata Bouleng. P. Z.S. 1900, p. 455, pl. xxxi.

Cameroon to Congo, eastwards to the Uellé and the Stanley
Falls.

2. BouLENGERINA DYBOWSKII Mocquard, Bull. Soc. Philom. (8)
1x, ISS 5 ee Ie
French Congo.
2. HLAPECHIS.

Bouleng. Cat. Sn. 111. p. 358.

1. ELAPECHIS GUENTHERI.

Hlapsoidea guentheri Bocage, Jorn. Se. Lisb. i. 1866, p. 70,
pl. i. fig. 3.
Elapechis guentheri Bouleng. t. c. p. 359.

Elapechis moebiusi Werner, Verh.. zool.-bot. Ges. Wien, xlvy1i.
1897, p. 400.

Togoland, Northern Nigeria, and Uganda to Angola and
Nyassaland.
3. Nata,

Laur. Syn. Rept. p. 90; Bouleng. Cat. Sn. ii. p. 372.

Synopsis of the Species.
I. 19 to 29 scales across the neck, which is dilatable, 17 to 23 across the body;
53-70 subcaudals.

6th or 7th upper labial largest and deepest, in contact with post-

oculars ; eye separated from the labials by suboculars ......... N. haie.
6th upper labial largest and deepest, in contact with postoculars ;

3rd and 4th upper labials entering the eye .....................65 NV. melanoleuca..
3rd or 3rd and 4th upper labials deepest and entering the Bie 6th

and 7th not in contact with postoculars ........... . WN. nigricollis.

II. 15 scales across the neck, which is not damianle! 13 or 15
across the body; 4th or 8rd and 4th upper labials enter ing
WINE Gye VASE] SOE FAIS “Go ,coonsdoun assude 506050 s9p b45eG0008 Sas n08 NV. goldii.
1. NATA HATE.
Coluber haie Linn. in Hasselq. Reise Palest. p. 366.
Naia haie Bouleng. t. ¢. p. 374.

Borders of the Sahara, East Africa southwards to the Tyrans-
vaal and Zululand, Palestine, Arabia.

2. NAIA MELANOLEUCA.

Naia haie, var. melanoleuca Hallow. Proc, Ac. Philad. 1857
pp. 61, 72.

Naia melanoleuca Bouleng. t. c. p. 376,

Tropical Africa, from the Gold Coast and Uganda to Angola
and Nyassaland.

3. Nara nigRicoiuis Reinh. Vidensk. Selsk. Skr. x. 1843,"p. 269,
pl. i. figs. 5-7; Bouleng. t. c. p. 378.

Senegambia and Upper Egypt to Bechuanaland and Natal.

SNAKES OF WEST AFRICA. 295

4. Nata cotpit Bouleng. Ann. & Mag. N.H. (6) xvi. 1895,
p. 34, and Cat. t. e. p. 387, pl. xx. fig. 2

Naia guentheri Bouleng. Cat. t. c. p. 388, pl. xx.

Sierra Leone to Congo, eastwards to the Ituri and the Kasai.

4, DENDRASPIS.

oe Versl. Zool. Gen. Amsterd. 1848; Bouleng. Cat. Sn.
5 0 434,

.
Two species :—

Scales in 15 to 19 rows, outer not shorter than dorsals ................ D. gamesonii.
Scales in 13 rows, outer half as long as dorsals ........................ D. angusticeps.

1. DENDRASPIS JAMESONII.

Hlaps jamesonti Traill, in Schleg. Phys. Serp., Engl. Transl.
p. 179 ple nies O20:

Dendraspis jamesonit Bouleng. t. c. p. 436.

Tropical Africa, from Nigeria and Uganda to the Congo and
Angola.

2. DENDRASPIS VIRIDIS.

Leptophis viridis Hallow. Proc. Ac. Philad. 1844, p. 172.
Dendraspis viridis Bouleng. t. c. p. 435.

Senegal to Niger and 8. Thomé Id., Gulf of Guinea.

Family VIPERIDz.

I. Eye moderate or large, usually separated from the upper labials by suboculars.

A. Upper surface of head with large symmetrical shields ; ape
TROWDUNG LS Bosconsonasosad : Saccseleaeen, OCUSUSE

B. Upper surface of head) a scales ; sprpill secteall.
1. Subcaudals in two rows

Lateral scales not smaller than dos sals, without serrated keels ......... Bitis.
Lateral scales smalier than ae disposed obliauelse with serrated
keels! gsceceeceare : 5 joseoddaccel, \OARARIOS.

Y. Subcaudals Sinaia: ‘eet ecules ties ie toa and more or less
oblique or inregul ar.

Lateral scales with serrated keels.. ss Doe umaconconarwou eos
Lateral scales without serrated keels ; p ‘tail prehensile .. caonseabocbaon, lQIPES.

II. Eye minute, with round pupil; upper surface of herd with
large =ymmnetrical shields; no loreal ; ; a small preocular nsually

pr esent . Atractaspis.

1. Causus.
Wagl. Syst. Amph. p. 172; Bouleng. Cat. Sn. ui. p. 465.

Two species :—

Scales in 15 to 21 rows; subcaudals in two rows ...........-...5.... OC. rhombeatus.
Scales in 13 rows; subcaudals single ...........2:::c cece a CO. Lichtensteinii.

1. CAUSUS RHOMBEATUS.

Sepedon rhombeatus Licht. Verz. Doubl, Mus. Berl. p. 106.
Causus rhombeatus Bouleng. t. c. p. 467.

Tropical and South Africa.

296 MR. G. A. BOULENGER ON THE

2. CAUSUS LICHTENSTEINII.

Aspidelaps lichtensteinti Jan, Rev. et Mag. Zool. 1859, p. o11.
Causus lichtensteinii Bouleng. t. c. p. 470.

From the Gold Coast and Uganda to the Congo.

2. Brris.
Gray, Zool. Miscell. p. 69; Bouleng. Cat. Sn. iit. p. 492.

Synopsis of the Species.

One or two series of scales between the nasal and the rostral; 8 to
11 scales across the head, from eye te eye.. eee LBA anietamns.
4 or 5 series of scales between the nasal and the ‘rostral ; 13 to 16
scales across the head, from eye to eye; a singlé enlarged, some-
times horn-like scale above the internasal, in contact with its
TSN VON rica ake ie Sari oa Seen ae ocd dona ace, “cram BRncbmte shogun dauiepuoahedn Saas oood B. gabonica.
4. to 6 series of scales between the nasal and the rostral; 14 to 16
scales across the head, from eye to eye; 2 or 3 enlarged, horn-
like scales above the internasal, usually with small scales
LOYETANIGLESTI (r] VSTITLL GOS dlsr.ntn eho doe aocude bes onadbeconoss cop oa gHeRnoB arene cssnadaes © WEI OMSKCOM TKS
1. Birts ARIETANS.
Vipera arietans Merr. Tent. p. 152.
Bitis arietans Bouleng. t. ¢. p. 493.
Tropical and South Africa, northwards to 8. Morocco; Southern
Arabia.
2. Brvris GABONICA
Echidna gabonica Dum. & Bibr. Erp. Gén. vu. p. 1428,
Dig ibe Op :
Bitis gabonica Bouleng. t. e. p. 499.

Tropical Africa.

3. BIrIs NASICORNIS.
Coluber nasicornis Shaw, Nat. Miscell. ii. pl. xciv.
Bitis nasicoriis Bouleng. tees pa00:
Tropical Africa.
3. CERASTES,
Wael. Syst. Amph. p. 178; Bouleng. Cat. Sn. ii. p. 501.
1. CerasrEes cornutus Forsk. Deser. Anim. p. ix; Bouleng.
t. c. p. o02.
Borders of the Sahara and Soudan; Arabia and Palestine.

4. Kcuis.
Merr. Tent. p. 149; Bouleng. Cat. Sn. i. p. 504.

1. HcHis CARINATUS.
Pseudoboa carinata Schneid. Hist. Amph. 11. p. 285.
Echis carinatus Bouleng. t. ¢. p. 505,

Desert and sandy districts of Africa north of the Equator ;
Southern Asia from Transcaspia and Arabia to India.

SNAKES OF WEST AFRICA. 297

5, ATHERIS.

Cope, Proc. Ac. Philad. 1862, p. 337; Bouleng. Cat. Sn. 11.
p. 508.
Synopsis of the Species.

9-11 scales across head, from eye to eye, 15-17 round eye; 25-36

SCMES DGOSIS WONT — go0co0 aoc 0da000 0n0e edonab eueonnobeogecds aebaso vaaaed A. chlorechis.
7-9 scales across head, from eye to eye, 10-16 round eye; 15-24
scales across body ...... .. A. squaniger.

8-10 scales across head, from eye to eye, ‘16-17 round eye; 25
scales across body ; several erect, horn-like superciliary scales, A. ceratophorus.

1. ATHERIS CHLORECHIS.

Vipera chloroechis Schleg. Versl. Ak. Amsterd. 111. 1855, p. 317.
Atheris chlorechis Bouleng. I. c.
Liberia to Gaboon.

2. ATHERIS SQUAMIGER.

Echis squamigera Hallow. Proc. Ac. Philad. 1854, p. 1938.
Atheris squamiger Bouleng. t. c. p. 509.

Calabar to Angola, eastwards to Uganda and the Ituri.

3. ATHERIS CERATOPHORUS Werner, Verh. zool.-bot. Ges. Wien,
xlv. 1895, p. 194, pl. v. fig. 1; Bouleng. t. c. p. 510.
Togoland, East Africa.

6. ATRACTASPIS.
A. Smith, Ill. Zool. 8. Afr., Rept.; Bouleng. Cat. Sn. ii. p. 510).

Synopsis of the Species.

I. Anal divided ; all or most of the subcaudals paired.
2nd lower labial separated from its fellow by the chin-shields ;

scales in 23-27 rows; ventrals 217-207 ............-...00 tee eee A. irregularis.
2nd lower labial forming a suture with its fellow ; scales in 19-21

rows; ventrals 3038-331 .. .... A, reticulata.
2nd lower labial forming a suture with its ‘fellow ; “scales in 1 21-23

rows; ventrals 336-359 . ete isserseeee A. heterochilus.

II. Anal entire; all or ribet of fhe saieauilalecr pair red

Scales in 21 rows; ventrals 198-222 .8..........000cecseceeeteteeesess. A. matschiensis.
Scales in 29 rows; ventrals 226 ............. La. Ae caudalis.

III. Anal entire; all or most of the eabeaudals nels
A. Postocular in contact with a large temporal.
1. Snout rounded, feebly projecting.
Qnd lower labial very large, forming a suture with its fellow;

scales in 23-27 rows; ventrals 178-193 . : i.e. A. corpulenta.
3rd lower labial longest ; ‘scales in 21 rows ; ; ventrals 195 ......... A. boulengeri.
3rd lower labial longest; scales in 19-21 rows; ventrals 251-300 A. aterrima.
2. Snout cuneiform ; scales in 31 rows; ventrals 240 ...... A. dahomeyensis.

B. Temporals small, 2 or 3 superposed in front.

Snout cuneiform; upper part of rostral as long as its distance from

the frontal; ‘scales in 25 rows ; ; ventrals 210-223 .. A. micropholis.
Snout rounded, upper part of rostral much shorter than its dis-

tance from the frontal; scales in 25-29 rows; ventrals 214-

225 A, watsonii.

298 ON THE SNAKES OF WEST AFRICA.

1. ATRACTASPIS IRREGULARIS.

Elaps irregularis Reinh. Vid. Selsk. Skr. x. 1843, p. 264, pl. 11.
figs. 1-3.
Atractaspis irregularis Bouleng. t. c. p. 513.

West and Central Africa, from the Gold Coast and Uganda to
the Congo.

2. ATRACTASPIS RETICULATA Sjostedt, Zool. Anz. 1896, p. 516.
Cameroon.

3. ATRACTASPIS HETEROCHILUS Bouleng. Ann. Mus. Congo,
FAO)! itl Jy) UBS Toll aN ities! Ihe

Cameroon, Ituri, Tanganyika.
4, ATRACTASPIS MATSCHTENSIS Werner, Verh. zool.-bot. Ges.
Wien, xlvii. 1897, p. 404.

Cameroon.

5. ATRACTASPIS CAUDALIS Sternf. Sitzb. Ges. Nat. Fr. Berl.
1908, p. 94.

Gold Coast.

6. ATRACTASPIS CORPULENTA.

Brachycranium corpulentum Hallow. Proc. Ac. Philad. 1854,
p. 99.

Atractaspis corpulenta Bouleng. Cat. Sn. i. p. 514.
Liberia to Congo, eastwards to the Ituri.

7. ATRACTASPIS BOULENGERI Mocquard, Bull. Soc. Philom. (8)
1.5 IUSO5 je MSs

Ogowe.

8. ATRACTASPIS ATERRIMA Giinth. Ann. & Mag. N. H. (8) xii.
1863, p. 363; Bouleng. t. c. p. 515.

Gold Coast to Niger, eastwards to Uganda.

9. ATRACTASPIS DAHOMEYENSIS Bocage, Jorn. Sc. Lisb. x1. 1887,
p. 196; Bouleng. t. ec. p. 516.

Dahomey.

10. ArracrasPIs MICROPHOLIS Giinth. Ann. & Mag. N. H. (4)
ix. 1872, p. 36, pl. i. fig. EH; Bouleng. t. c. p. 516.
Senegambia, Northern Nigeria.

11. Arracraspis warsonir Bouleng, Ann. & Mag. N. H. (8) ii.
1908, p. 93.
Atractaspis nigra Pellegr. Bull. Mus. Paris, 1909, p. 414.

Mauritania, French Soudan, Northern Nigeria.

ON THE SNAKES OF NORTH AFRICA. _ 299

17. A List of the Snakes of North Africa.
By G. A. BouLencer, U.RS., F.ZS8.
[Received May 20, 1919: Read June 17, 1919.]
(Published by permission of the Trustees of the British Museum.)
This list, dealing with the comparatively few species known

from North Africa (Section VII. of map on p. 269), concludes the

series I have prepared for the easy identification of the Snakes
of Africa *.
Only four families are represented in this area :—

Worm-like, covered with uniform scales above and beneath ; mouth

small, inferior ; eyes rudimentary, under the head-shields....... GLAUCONIID®.
Mouth large; eyes exposed; head covered with smail scales ;

ventral shields much narrower than the body ...................... Borpm.
Mouth large; eyes exposed; head with large shields; ventral

slivel dsibnoddlte stares terre h ncn esas neptes/oecercdcepebeemaad | LCOLUMBRTDIAT
Mouth large; eyes exposed; head covered with small scales;

TOMA SOG ATEN | as psnnasnmdeoase sonuonosenepnnc soseBeacebeeccsunceme NV TTDI IYO

Family GLAUCONIIDE.
A single genus.
1. GuaucontA.

Gray, Cat. Liz. p. 139; Bouleng. Cat. Sn. i. p. 59.

Two species :—

Snout hooked, the preoral portion flat or concave inferiorly ;

diameter of body at least 100 times im total length......... G. macrorhynchus.
Snout rounded ; diameter of body less than 100 times in total
Ten bere. bs52i2) kee er a en RU va Meee. SEGRE G a Ts

1. GLAUCONIA MACRORHYNCHUS.

Stenostoma macrorhynchus Jan, Arch. Zool. Anat. Phys. i.
1862, p. 190.

Glauconia macrorhynchus Bouleng. Cat. Sn. 1. p. 61.

Glauconia algeriensis Jacquet, Bibl. Anat. iv. 1896, p. 79, figs.

Algeria, Nubia, Mesopotamia, Persia.

2. GLAUCONIA CAIRI.

Stenostoma cairt Dum. & Bibr. Erp. Gen. vi. p. 323.
Glauconia cairt Bouleng. Cat. Sn. i. p. 66.

Egypt, Nubia, Abyssinia, Somaliland, Mauritania.

Family Borp #.
A single genus.
Lise Dunner

Daud. Hist. Rept. vil. p. 251; Bouleng. Cat. Sn. i. p. 122.

* P. Z.S. 1915, pp. 193, 369, 611, 641, and 1919, p..267.—According to my latest
estimate, the number of African species of Snakes amounts to 475.

300 MR. G. A. BOULENGER ON THE

T'wo species :—

12 to 15 scales from eye to eye across forehead ; tail pointed, ending
ANY AY COMMCAI SO UGS fe Be... cberaaier tases Me acc aOR a AE Hi, thebaicus.
5 to 7 scales from eye to eye across forehead ; tail obtuse ............... E. jaculus.

1. Eryx THeparcus Reuss, Mus Senckenb. i. 1834, p. 134 ;
Bouleng. Cat. Sn. i. p. 125.

Upper Egypt to East Africa.

2. Eryx FACULUS.

Anguis jyaculus Linn. 8. N. i. p. 391,
Eryx jgaculus Bouleng. Cat. Sn. 1. p. 124.

Algeria, Egypt, Syria, Asia Minor, Greece.
Family COLUBRID.
Three parallel series :—

No poison-fangs; all the teeth solid .................0.:.0:00:.... A. Aglypha.

TRUITT TAVERS WSTMUAGL os one cose Aonces seqnco cos ens ccoseedoe sop see seaooces Jey Ojonstilnoadhyyalag.
JEUSOSMHENOERS TN TRO — SoncvocosesdcvocscoosseonoeHn esnecourcocanooawa Op JPROR@KoglhyalNel

A. Aglypha.

I. Loreal present; scales in 19 rows or more.
A. Scales strongly keeled, in 19 to 23 rows; suey round; a
single anterior temporal ................ cesses. Dropidonotus.

B. Scales smooth or feebly keeled ; 2 to re superposed anterior temporals ; rostral
not twice as broad as deep.

Head elongate; snout rounc:»? at the end; pupil round; one or

more suboculars ; ventra.s 195-278 ; subeaudals 65- 154 ee: Zamenis.
Snout cuneiform, rostral four-sided; pupil vertically elliptic or
subelliptic ; ventrals 160-188; subcaudals 35-46 ............... Lytoriynchus.
Snout rounded or obtusely pointed ; ee round ; no suboculars ;
ventrals 170-200; subcaudals 49-72 . Beer ee eee Cononellas
If. Loreal present or absent; scales in br rows, amootl: ; Enont
strongly projecting ........ , : .. Oligodon.
Ill. Loreal absent; pupil anleells Pe o 23 40, 271 rows,
strong lyr keeled’ ee. eects eine saucuencecton eect ene eeeiny ce eee LD SUE LOIS

1. TROPIDONOTUS.

Kuhl, Bull. Sc. Nat. ii. 1824, p. 81; Bouleng. Cat. Sn. i.
pelo 2F

Synopsis of the Species.

Scales in 19 rows ; third and fourth upper labials entering the eye;

upper postocular not in contact with the temporal ....... oa SL AGIOS?
Scales in 19 rows; fourth or fourth and fifth upper labials enteri ing

the oye ; upper postocular not in contact with the temporal . T. tessellatus.
Scales in 21 or 23 rows; third and fourth upper labials entering

the eye; upper postocular in contact with the temporal ...... TZ. viperinus.

1. TROPIDONOTUS NATRIX.
Coluber natriw Linn. S. N. i. p. 380.

SNAKES OF NORTH AFRICA. 301

Tropidonotus natrix Bouleng. t.c. p. 219.

Algeria and Tunisia north of the Atlas, Europe, Western
Asia.

2. TROPIDONOTUS TESSELLATUS.

Coronella tessellata Laur. Syn. Rept. p. 87.
Tropidonotus tessellatus Bouleng. t.c. p. 233.

N.E. Egypt, Europe, Western Asia.

3. TROPIDONOTUS VIPERINUS.

Coluber viperinus Latr. Hist. Rept. iv. p. 49.
Tropidonotus viperinus Bouleng. t.¢. p. 235.

Morocco, Algeria, Tunisia, Spanish Peninsula, France, Switzer-
bd 7 ? ? 7)
land, Italy.

ZAMENIS.

Wagl. Syst. Amph. p. 188; Bouleng. Cat. Sn. 1. p. 379.

Synopsis of the Species.

T. A pair of internasals and a pair of preefrontals.
A. Scales in 19 rows ; two labials entering the eye.
Frontal anteriorly not or but little broader than the supraocular ;

ventrals 205-218; subcaudals 98-132 ...... Z. dahlii.
Frontal anteriorly considerably broader than the supraocular ;

ventrals 212-262; subcaudals 113-154 . Z. rhodorhachis-
Frontal anteriorly considerably broader than the supraocular ;

ventrals 195-201; subecaudals 95-105 ......................... Z. rogersi.

B. Scales in 21 (rarely 23) rows, smooth ; two labials enter-

ing the eye; ventrals 198-228; subcaudals 79-111 ...... Z. florulentus.
C. Scales in 23 or 25 rows, obtusely or faintly keeled ; usually

a single labial entering the eye; ventrals 197-216; sub-

CRG RUS OSTOMY ojo ois sce dasons eb8 con ccubbacoodgeeessadeeusnoncdeoe Z. nummifer.
D. Scales in 25 to 29 (rarely 23) rows, smooth.

Scales usually in 25 rows; usually one labial entering the eye ;

ventrals 214-232; snbcaudals 87-104 ..... Z. algirus.
Scales in 25 to 29 rows; eye usually separ: ated: from the labials

by a series of suboculars; ventrals 222-258; subcaudals

MG NOT a eeihceetcmas on tu Co eee Cee EAE eee esecaewesaniial, , Laehappocnepiss

If. Prefrontals broken up into 3 or more shields; eye separated
from the lJabials by suboculars; scales in 25 to 31 rows,
usually more or less obtusely keeled; ventrals 210-278 ;
Sule CU SMI) So. ccdone- concn seacooncs cosoro den seo osonesastonceas 4 CKIMIGTATAG In

1. ZAMENIS DAHLII.

Tyria dahlii Fitzing. N. Class. Rept. p. 60.
Zamenis dahlii Bouleng. t. ec. p. 397.

N. Egypt, 8.W. Asia, S.E. Europe.

2. ZAMENIS RHODORHACHIS Jan, in De Filippi, Viagg. Pers.
p. 356 ; Bouleng. t.c. p. 398.

Egypt to Somaliland, $.W. Asia.

302 MR. G. A. BOULENGER ON THE

3. Zamenis RocERst Anders. Ann. & Mag. N. H. (6) xi. 1893,
p. 439; Bouleng. Cat. Sn. iii. p. 623.
N. Egypt.

4, ZAMENIS FLORULENTUS.

Coluber florulentus Geoffr. Descr. Egypte, Rept. p. 146, pl. vin.
fig. 2.

Zamenis florulentus Bouleng. Cat. Sn. 1. p. 402.

Egypt to Somaliland.

5. ZAMENIS NUMMIFER.

Coluber nummifer Reuss, Mus. Senckenb., 1. p. 135.
Zamenis nummifer Bouleng. t.c. p. 407.

N. Egypt, Syria, Cyprus, Asia Minor.

6. ZAMENIS ALGIRUS.

Periops algira Jan, Elenco, p. 60.
Zamenis algirus Bouleng. t.c. p. 408.
Algeria and Tunisia.

7. ZAMENIS HIPPOCREPIS.

Coluber hippocrepis Linn. 8. N. i. p. 388.

Zamenis hippocrepis Bouleng. t.c. p. 409.

Morocco, Algeria, Tunisia, Spain and Portugal, Sardinia.

8. ZAMENIS DIADEMA.
Coluber diadema Schleg. Phys. Serp. i. p. 148.
Zamenis diadema Bouleng. t.c. p. 411.
Algeria, Tunisia, Tripoli, Egypt, Arabia to Turkestan and
N.W. India.
3. LYTORHYNCHUS.

Peters, Mon. Berl. Ac. 1862, p. 272; Bouleng. Cat. Sn. i.
p- 414.

1. LyToRHYNCHUS DIADEMA.

Heterodon diadema Dum. & Bibr. Erp. Gén. vii. p. 779.

Lytorhynchus diadema Bouleng. t.¢. p. 415.

Algerian and Tunisian Sahara, Tripoli, Egypt, Nubia, Arabia,
Syria, Mesopotamia, Persia.

4, CORONELLA.
Laur. Syn. Rept. p. 84; Bouleng. Cat. Sn. u. p. 188.

Two species :—
Rostral as deep as broad, produced between the mternasals ............ ©. amalie.
Rostral much broader than deep, just visible from above ............ C. girondica.

SNAKES OF NORTH AFRICA. 303:

1. CoRONELLA AMALI.

Rhinechis amalie Boettg. Zool. Anz. 1881, p. 570.
Coronella amalice Bouleng. t.c. p. 193.

Morocco and Algeria.

2. CORONELLA GIRONDICA.

Coluber girondicus Daud. Hist. Rept. vi. p. 432.

Coronella girondica Bouleng. t.c. p. 194.

Morocco and Algeria, Spain and Portugal, 8S. France, Italy.

5. OLiGopon.
Boie, Isis, 1827, p. 519; Bouleng. Cat. Sn. ii. p. 233.

1. OLIGODON MELANOCEPHALUS.

Homalosoma melanocephalum Jan, Arch. Zool. Anat. Phys. ii.
1862, p. 34.
Oligodon melanocephalus Bouleng. t. ce. p. 246.

N. Egypt, Sinai, Syria.

6. DASYPELTIS.
Wagl. Syst. Amph. p. 178; Bouleng. Cat. Sn. 11. p. 353.

1. DASYPELTIS SCABRA.

Coluber scaber Linn. Mus. Ad. Frid. p. 36, pl. x. fig. 1.
Dasypeltis scabra Bouleng. t.c. p. 354.

Egypt, Tropical and South Africa, 8. Arabia.

B. Opisthoglypha.

J. Head short, very distinct from neck ; pupil vertically elliptic; scales in 19 to
23 rows; subcaudals 60-86.

Hyeunoderateswanallldivid eden peers aera Labo piase
Pyevlarcespanalkentinel’c-a ce peeeeeeee re eeoece ae ee eee Anes Le epeodena
II. Head elongate; eye large, with round pupil; scales in 17 or 19 rows.

Scales not oblique; loreal not longer than deep; subcaudals

CAST Bemrietee weaee a Somber i sarcpscd saab See Hoscos gas HORGEREE Der OeER IBSEN eC ey EAL HOS
Scales more or less oblique; loreal at least 13 times as long as
deep; subcaudals 90-149 2.02.0... eee eee eeeeeeeeeeeeeeses Psammophis.

ILI. Head short, not very distinct from neck ; eye rather small,
with round or vertically subelliptic pupil; rostral at least
twice as broad as deep; scales in 19 to 25 rows; sub-
Caudal’ 4LO=B4i 00. ysis yea idem aateen eee Mane inbicneniie sn conan eiee Macroprotodon.

1. TARBOPHIS.

Fleischm. Dalm. noy. Serp. Gen. p. 17; Bouleng. Cat, Sn. iti.
p. 47.

304 MR. G. A. BOULENGER ON THE

Two species :—
Scales in 19 rows; ventrals 174-190 ......0......ccccccceeesseeseeeeeeeeeeeee LL savignyi.
Scales in 23 (rarely 21) rows; ventrals 218-272 .....0..0.00........ ZL. obtusus.
1. TArBorHis sAviGNyI Bouleng. t.c. p. 48.
N. Egypt, Syria.

2. TARBOPHIS OBTUSUS.

Coluber obtusus Reuss, Mus. Senckenb. i. 1834, p. 137.
Tarbophis obtusus Bouleng. t.c¢. p. 52.

Egypt to Somaliland, Mauritania and N. Nigeria.

2. LEPTODIRA.

Giinth. Cat. Col. Sn. p. 165; Bouleng. Cat. Sn. 11 p. 88.

1. Lepropira TRrIPoLiTANA Werner, Zool. Jahrb., Syst. xxvii.
ISOS), J. OHS
Tripoli.
3. C@LOPELTIS.

Wagl. Syst. Amph. p. 189; Bouleng. Cat. Sn. 1. p. 141.

Two species : —

‘Two loreals; frontal very narrow in the middle; subcaudals

GSENO2 aissctaestee odtcceaie sue tuowanime nea ua i meteeaany cae brbioctnsee C. monspessulana.
A single loreal: frontal as broad as the supraocular; sub-
GMUGENIS BATS -soccaccsces sgsoso cor taascuvan esacdscencavcarcoonacceces Cy MAOWEMSIS.

1. C@LOPELTIS MONSPESSULANA.

Coluber monspessulanus Herm. Obs. Zool. i. p. 283.

Celopeltis monspessulana Bouleng. |. e.

Borders of the Mediterranean, southwards to Mauritania, east-
wards to Persia.

2. C@LOPELTIS MOILENSIS.
Coluber moilensis Reuss, Mus. Senckenb. i. 1834, p. 142, pl. vii.
fiona
Ceelopeltis moilensis Bouleng. t. c. p. 143.
Northern Sahara, from Algeria to Egypt, Nubia, Arabia,
Western Persia.
4, PSAMMOPHIS.

Boie, Isis, 1827, p. 521; Bouleng. Cat. Sn. ii. p. 152.

Two species :—

Loreal 3 to 4 times as long as deep ; usually 9 upper labials, 5th and

Ghhventerimg phe eye =...0.4.6 iihecateee) ene etc kotet ne dee ne tee an RISC DIGG.
Loreal 13 to 25 times as long as deep; usually 8 upper labials, 4th

RUNG LSE oY HMMS TPA HOI OYE ENVE) cou son ostelcaonosd ace -scsne saccdsededessoobeace P. sibilans.

SNAKES OF NORTH AFRICA. 30a

1. PSAMMOPHIS SCHOKARI.
Coluber schokari Forsk. Descr. Anim. p. 14.
Psammophis schokari Bouleng. t.c. p. 157

North Africa to Mauritania and Somaliland; Arabia and Syria
to Afghanistan and Sind.

2. PSAMMOPHIS SIBILANS.

Coluber sibilans Linn. 8. N. i. p. 383.
Psammophis sibilans Bouleng. t.c. p. 161.
Egypt, Tropical and South Africa.

5. Macroproropon,

Guichen. Expl. Sc. Alg., Rept. p. 22; Bouleng. Cat. Sn. iii.
p. 175.

1. MacroPROTODON CUCULLATUS.

Coluber cucullatus I. Geoftr. Descr. Egypte, Rept. pp. 148, 151,
pl. viii. fig. 3.

Macroprotodon cucullatus Bouleng. 1. e.

North Africa from Morocco to N. Egypt, Mauritania, 8. Spain,
Baleares, Lampedusa, 5. Palestine.

C. Proteroglypha.

Two genera :—

Scales oblique; neck dilatable ; subcaudals 53-68 ..................... Naia.
Scales not oblique; subcaudals 45-48 000... ceeeeecceseeeeevsseseese Wealterinnesia.

1. Nata.

Laur. Syn. Rept. p. 90; Bouleng. Cat. Sn. iii. p. 372.

Two species :—

‘6th or 7th upper Jabial largest and deepest, in contact with post-
oculars; eye separated from the lalials by suboculars ; 21-23
scales across neck, 19-21 across middle of body ...... . WN. haie.
3rd upper labial deepest and entering the eye, 6th and 7th not in
contact with postoculars ; 25-29 scales across neck, 21-25
PUTIN THAN CT KO OIE OWI? sag nona osban8 easwunanseenenecadoadcsesosseonseee bvy dawRnGaLUTS.

1. NAIA HAIE.

Coluber haie Linn. in Hasselq. Reise Paleest. p. 366.
Naia hate Bouleng. t.c. p. 374.

Borders of the Sahara, Kast Africa southwards to the Transvaal
and Zululand, Palestine, Arabia.

2. Nata NIGRICOLLIS Reinh. Vid. Selsk. Skr. x. 1843, p. 269,
pl. i. figs. 5-7; Bouleng. t.c. p. 378.
Upper Egypt and Senegambia to Bechuanaland and Natal.

306 MR. G. A. BOULENGER ON THE

2, WALTERINNESIA.

Lataste, Le Natural. 1887, p. 411; Bouleng. Cat. Sn. i.
p. 392.

1. WALTERINNESIA HGYPTIA Lataste, ].c.; Bouleng. 1. c.
Eeypt?, Nubia ?
Family VIPERID2.

J. Lateral scales not smaller than dorsals, without serrated keels.
Nasal in contact with rostral or separated from it a a naso-rostral

shield; scales in 19 to 27 rows... ... Vipera.
Nasal separated from rostral by small scales ; ‘scales in 29 to 41 rows... Bitis.

II. Lateral scales smaller than dorsals, disposed obliquely, with ea keels.
Ventrals angulate laterally; subcaudals in two rows ............ ........... Cerastes.
Ventrals rounded; subcaudals single .................. cece eee Hehis.

1. VIPERA.

Laur. Syn. Rept. p. 99; Bouleng. Cat. Sn. i. p. 471.

Two species :—

Snout turned up at the end, produced into a small ray: Saas
scales in 21 rows ...... BAS es ea ee ... V. latastii.
Snout rounded; scales in 25 or 27 rows SRG ee SS oe UCD EGU Ge

1. Vipera tatastit Bosca, Bull. Soc. Zool. France, 1878, p. 116,
pl. iv.; Bouleng. t.c. p. 484.

Morocco and Algeria north of the Atlas, Spain and Sperines

2. VIPERA LEBETINA.

Coluber lebetinus Linn. S. N. i. p. 378.
Vipera lebetina Bouleng. t.c. p. 487.

Morocco, Algeria, Tunisia, §.W. Asia, Greek Archipelago.

2. BIris.
Gray, Zool. Miscell. p. 69; Bouleng. Cat. Sn. iil. p. 492.

1. Brivis ARIETANS.
Morocco (Valley of Sous), Tropical and South Africa, S, Arabia.

- 3. CERASTES.
Wagl. Syst. Amph, p. 178; Bouleng. Cat. Sn. 11. p. 501.

Two species :—

15 to 21 scales across the head, from eye to eye, 14 to 18 round the

eye; fr equently an erect horn- like scale above the eye; scales in

DpBiS TONGS ema aUks) NEYO SAUGS 5 onc ccc soa cnacuc nto cn con ohn caeceosed 200 ULE C. cornutus.
9 to 13 scales across the head, from eye to eye, 9 to 14 round the eye ;

scales in 23-27 rows; ventrals 102-122... 12.20. .:.... 0.0... sec een ceeee C. vipera.

SNAKES OF NORTH AFRICA. 307

1. Cerastes cornutus Forsk. Descr. Anim. p.ix; Bouleng. t. c.
p. 502,
Borders of the Sahara and Soudan, Arabia and Palestine.

2, CERASTES VIPERA.

Colubra vipera Linn, in Hasselq. Reise Palest. p. 314.
Cerastes vipera Bouleng. t.¢. p. 503.

Northern border of the Sahara, from Algeria to Egypt, Sinai.

4. Kents.
Merr, Tent. p. 149; Bouleng. Cat. Sn. i. p. 504.

Two species :—

Scales on snout and vertex more or less strongly keeled; 2 (rarely 1
or 3) series of scales between eye and upper labials; ventrals

TEP STE ce:can danas Bop SCO a0 GOR GHEE ER OOE ACES SoA Gen Cae Oar aa Bc OCR Aras comieR RMAC nO XARA TIS
Scales on snout and vertex smooth or obtusely keeled; 3 or 4 series
of scales between eye and upper labials; ventrals 174-205 ...... FE. coloratus.

1. EcHIS CARINATUS.

Pseudoboa carinata Schneid, Hist. Amph. ui. p. 285.

Kchis carinatus Bouleng. t.c. p. 505.

Desert and sandy districts of Africa north of the Equator ;
Southern Asia from Transeaspia and Axabia to India.

2. Kcuis cotoratus Giinth. P. Z. 8. 1878, p. 978: Bouleng.
t.c. p. 905.

Kgypt, Socotra, Arabia, Palestine.

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ANE Wet SUNNEREEGUND (SIGN 1S) 4 al

ON A NEW SPECIES OF ZEUGLODONT. 309

A Description of New Species of Zeuglodont and of
Leathery Turtle from the Hocene of Southern Nigeria.
By C. W. Anprews, D.Sc., F.R.S. (British Museum,
Natural History).

(Published by permission of the Trustees of the British Museum.)
[Received April 29, 1919: Read June 17, 1919. |
(Plates I. & II.*)

Two small collections of vertebrate remains from the Ombialla
District of Southern Nigeria have recently been received by
the British Museum, one having been sent by Sir Frederick
Lugard, G.C.M.G., the other by Sir John Eaglesome, K.C.M.G.
The sternum of a large carinate bird included in the latter
collection has already been described 7, and it is now proposed to.
give a short account of some remains of a Zeuglodont Whale and
of a Turtle belonging to the so-called Athecate group.

PAPPOCETUS LUGARDI, gen. et sp. nov.

Portions of the lower jaws of a Zeuglodont are included in
both collections, and, although in both cases incomplete, the
specimens to some extent supplement one another, so that the
structure is fairly clear. The most complete specimen (M 11414,
referred to as specimen A) consists of the imperfect left ramus
wanting the articular and angular regions, but united with a
considerable portion of the anterior region of the right ramus
including the hinder part of the symphysis (Biot. fics) wor
tions of five teeth are preserved on the left side and of two on the
right. The bones were embedded in an intensely hard pyritous
clay, including many fragments of molluscan shells. This may
be regarded as the type-specimen.

The other specimen (M 11086, specimen B) is a left ramus of a
mandible broken into three pieces (text-fig. 1), The anterior of
these bears the sockets for the incisors, canine, and front half of
the single-rooted pm,. Behind this a length of about 2-5 em.,
which must have carried the posterior half of pm, and anterior
half of pm,,is wanting. The length missing is estimated by com-
parison with specimen A which belonged to an individual of nearly
the same size. The two other fragments unite below. but, unfor-
tunately, the portion of the alveolar border bearing pm, is lost.
The hinder piece bears the basal portion of the very large pm,, and
following this without interval are the three molars, of which the
first is nearly complete, the second is represented by the roots
only, while the third, which had not yet emerged, has been
exposed by cutting away the bone; the articular and angular

regions are missing.

* For explanation of the Plates see p. 319.
+ Proc. Zool. Soc, 1916, p. 519.

bo
bo
*

310 DR. C. W. ANDREWS ON A NEW SPECIES OF

The outer face of the mandibular ramus is convex from above
downwards, the convexity being most marked in the symphysial
region, which extends back to the level of the anterior root of
pm,; in Prozeuglodon it does not seem to extend beyond the
hinder end of pm,. he symphysial surface itself bears a
strongly rugose surface for union with the opposite ramus, the
rugosities being most strongly developed posteriorly : the union
between the two rami must have been much stronger than in
Zeuglodon or Prozeuglodon, where the symphysial surfaces bear
only a few straight and slightly developed longitudinal rugosities.
Behind the symphysis the inner face of the ramus is convex from
above downwards immediately beneath the alveolar border, but
towards the ventral edge becomes gently concave in the same
direction. The ventral border is nearly straight from before
backwards as far as beneath the hinder end of m,, where it turns
somewhat upwards for a short distance and then continues in the
original direction, so that at this point a slight step-like prominence
is formed (text-fig. 1, 2). The posterior portion of the ramus is
lost in both specimens. In its general form the mandibular
ramus is very similar in form to those of Pr -ozeuglodon and
Zeuglodon, but at the same time is distinguished from them by
being more massively constructed and by the presence of the
slight step in the ventral border referred to above.

Text-figure 1.

Left ramus of mandible of Pappocetus luyardi, gen. et sp. nov.

n, step-like notch on lower border (M 11086). About % nat. size.

Teeth.—So far as can be ascertained the dental formula of the
mandible was I. 3, C. 1, Pm. 4, M. 3—the full Eutherian dentition.
‘Of these, I, is represented in specimen B by the socket only ;
this is situated at the extreme anterior end of the jaw and was
separated from its fellow of the opposite side by a very thin
wall of bone only. The tooth must have been directed forwards
and somewhat upwards. I, and I, are represented in the same
specimen by their broken bases, atin are somewhat wider, from
before backwards than from side to side, and appear to have
possessed a slight keel on the anterior border. Of the incisors
I, is much the iat gest, the Jongitudinal diameter of its base beine,
about 27 mim., while in I, this measurement is only 13 mm.
I, was about the same size at I,. This relatively large size if

1
I, seems to be characteristic of this genus, not occurring in
2 Oo ? (a)

ZEUGLODONT AND OF LEATHERY TURTLE. SIL

Prozeuglodon or Zeuglodon. 1, followsimmediately behind I,, but
between it and I, there is interval of about 23mm. The canine
is separated from I, by a diastema of 30 mm.; it is represented
by its broken base, which shows that it was about the same size
as I,, and was probably somewhat compressed from side to side
with an anterior angle ; like the incisors it was directed forwards.
Behind and a little to the outer side of the lower canine there
is a slight depression in the outer surface of the jaw, presumably
for the reception of the point of the upper canine. Pm, is
represented by the anterior half of its broken base in specimen B,
and by its socket only in specimen A; it is separated from the
canine by an interval of about 45 mm., and was considerably
compressed laterally, its long diameter being about 30 mm., the
transverse only 14. The remaining teeth are all represented in
one or other of the specimens by t their more or less broken crowns.
Pm, is preserved in specimen A only, where it is present on both
sides, following pm, at an interval of 19 mm. It is a two-rooted
tooth, the greatest length of which is 37 min., while its greatest
width above the front of the posterior root is 13mm. So far as
can be made out, the compressed crown formed a single cusp
without accessory serrations. Both the anterior and posterior
borders of the crown are blunt and rounded ; at the base of the
crown there is a well-marked constriction and on the inner side
there is a slightly developed cingulum. The enamel is much
roughened, being raised into knotted ridges, which for the most.
part run vertically. On the posterior lobe of the tooth the
enamel ridges of the outer and inner side meet, forming a keel
which is situated rather more on the inner than on the outer
side of the crown.

Pm, is represented in specimen B by its roots only, but in
A is present on both sides, that on the left being nearly
complete. It is a long, laterally compressed two-rooted tooth,
the length of the crown being 53 mm., its greatest breadth only
15mm. The crown forms one large cusp, the anterior slope of
which is shorter and steeper than the posterior. On the anterior
border there seem to have been no accessory cusps, but on the
posterior there are two with perhaps a rudimentary basal cusp
just above the cingulum, which is well developed on the postero-
internal side; it is also distinetly marked on the outer and less
clearly on the inner face of the tooth. The summit of the main
cusp has undergone considerable wear, which also extended down
the anterior edge; the top of the upper accessory cusp is also
worn.

Pm, is unfortunately represented in specimen B by its two
roots only, while in A the crown is very imperfect. It was-even
more strongly compressed laterally than pm,, and from before
backwards was considerably the longest tooth in the series,
measuring 59 mm. in this direction, while from side to side the
greatest width (above the posterior root) is only 18mm. It
consisted of a main anterior cusp, which may or may not have
borne small accessory cusps, and a posterior heel-like cusp which

2311 DR. GC. W. ANDREWS ON A NEW SPECIES OF

is broken but which seems to have had a cutting-edge. There is
a distinct cingulum round the whole crown, but it is much more
strongly developed at the posterior end. The enamel, except
near the base of the crown, is smoother than in pm,, perhaps
partly through wear. The posterior root is considerably the
larger.

The first molar (PI. I. fig. 2) is present in both specimens, though
in each case the summit of the main cusp is wanting; 1t is much
smaller than pm, with which it is in contact; its antero-posterior
length is 44mm. and its width (above the anterior root) 18°5 mm.
The crown formed a cutting-blade consisting of a large anterior
and a smaller posterior lobe, both inclined a little backwards.
On the outer face of the posterior lobe there is a well-marked
surface of wear produced by shearing against the upper tooth and
extending down to the cingulum. This latter is well marked
and is most prominent at the hinder border of the tooth.
Immediately above the cingulum on the antero-internal side of
the tooth the enamel is raised into a group of small rounded
tubercles, a line of which is continued up on the antero-internal
face of the main cusp towards its summit. The general surface
of the enamel is raised into faint irregular ridges, which,
increasing towards the unworn edges of the hind cusp, give them
the appearance of being ae erenulated.

In specimen A m, is onl y just emerging, while in B, though
more advanced, the tooth i is not fully in place ; in both specimens
ib 1s Imper fectly preserved, but, so far as can be seen, was quite
similar tom,. In specimen A it can be seen that the posterior
edge of the main cusp is raised into a number of crenulations,
which pass on the sides into irregular ridges of enamel.

M, was not cut in either specimen, hat in B it has been partly
exposed and its form seems to be like that of the other molars.
The molars and the Jast premolar form a closed series.

The collection also includes an isolated single-rooted tooth
(M 11087) with a conical crown, probably one of the incisors of
this species (PI. I. fig. 3). The inner face is somewhat flattened,
and is limited anteriorly by a well-defined ridge. The posterior
surface of the tooth has the enamel raised into numerous irregular
more or less vertical ridges of enamel, the outer one being the
best defined and running up to the summit of the crown. There
is a slightly developed cingulum, most marked on the inner and
posterior faces.

Another specimen probably belonging to this species is an imper-
fect axis vertebra (M 11089), cea the neural arch and lateral
processes (PI. I. fig. 4). The odontoid is a blunt prominence, oval
in section; it has a longitudinal ridge on its upper surface and a
shght dimple-lke depression on its anterior end. From the base
of the odontoid the lateral surfaces for the atlas slope strongly
backwards, widening as they go; their upper edge is marked by
a strong ridge, behind which the stout pedicle of the neural arch
arises, inclining to its fellow of the opposite side. The posterior

ZEUGLODONT AND OF LEATHERY TURTLE. 313

surface of the centrum is nearly flat, and between its lateral
edge and the broken base of the transverse process is a well-
marked coneayity. The ventral surface has a strong prominence
in the middle line, deepening towards its posterior border.
In the middle line on the dorsal surface, on its posterior half, is a
deep circular pit —probably the nutritive foramina opened into it.
A similar depression has been noticed on the axes of a Cave-Bear
and of the Creodont Apterodon; probably it is found elsewhere,
but it is not seen in two axis ver tebree of Prozeu glodon. Compared
with the axis of Protocetus as figured by Fraas*, this specimen
has a blunter odontoid, the fateral surfaces for the atlas slope
more strongly backwards, and the posterior surface is much wider
in proportion to its depth. In Protocetus the centrum is about
as long and has a similar ventral prominence. In Prozeuglodon 7
the whole vertebra is shorter, the ventral ridge is indicated only
by a slight posterior prominence, and the surfaces for the atlas
do not slope so much backwards. The length of the centrum
and the backward slope of the atlantal surface, the width of
the posterior surface and the ventral prominence, seem to be
primitive characters, approximating to what is seen in the
Carnivore-Creodont group.

This new Zeuglodont, for which the name Pappocetus lugardi
is suggested, is especially interesting on account of the Car-
nivore-like characters of the teeth, which seem to point to a
Creodont-Carnivore ancestry as was suggested ly Fraas from his
study of the Middle Hocene genus Pr olocetus t. Unfortunately
in this animal only the upper dentition is known, but pro-
bably the lower teeth were not very unlike those now described.
To this relationship with the Creodonts some ebjections have
recently been raised by Matthew and Gregory $, who point to
a number of characters which seem to indicate that the
Zeuglodonts may have branched off from the primitive Insecti-
yora, perhaps from some such form as the Kocene Pantolestes, a
type which, at any rate, shows that some of the early members of
the order attained a considerable size and became adapted to
an aquatic life ||. Attention is also drawn to the remarkable
superficial similarity in the form of the Aeuglodont skull to
those of some of the Centetidee, notably Memicentetes. This
similarity is also particularly noticeable in the form of the
mandible, in which even so small a point as the step-like notch
on the ventral border is present, though farther back. In the
structure and arrangement of the teeth there seems to be
nothing that altogether excludes the possibility of such a

* Braas, “ Neue Zeuglodonten aus dem unteren Mitteleocan vom Mokattain bei
Cairo,” Palwxont. Abhandl. Bd. x. (1902-5), p. 211, pl. il. fies, 2-5

+ Andrews, Catal. Tertiary Vertebrata of the Fayum (1906), p. 254, text-fig. 82
B-D.

+ Palwont. Abhandl. n.s. Bd. vi. (1904), p. 199.

§ “The Orders of Mammals,” Bull. Amer. Mus. Nat. Hist. vol. xxvii. (1910),
p. 414. ;

|| Memoirs of the Amer. Mus. Nat. Hist. vol. ix. (1909), p. 523.

314 DR. C. W. ANDREWS ON A NEW SPECIES OF

relationship, and it is interesting to note that probably the -
ancestors of the Centetidz lived in Africa in early Tertiary times,
and it is there also that the Zeuglodonts probably origina ted,

CosMOCHELYS DOLLOT, gen. et sp. nov.

The second specimen (R 4338) now described consists of
portions of the central region of a carapace belonging to a
Chelonian referable to the so-called Athece, to which belong the
recent Leathery Turtle, Dermochelys coriacea, and the Tertiary
genus Psephophorus to which the present form seems to be most
nearly allied. ‘This specimen was presented to the Museum by
Sir John Eaglesome, K.C.M.G. Unfortunately, the fragments
seem to have been picked up at random, so that much has been
lost; but, nevertheless, it has been possible to join a certain
number of pieces, which together make up a portion of the
carapace measuring about 37 cm. long by 27 em. wide. On the
right side of this the upper portions of five ribs are preserved
and on the left, three, corresponding to the posterior three on
tke right side. In the case of these three posterior ribs, the
upper portion of their articular ends are preserved, articulating
with the neural arches. These bore neural short spines, to the
upper ends of which the remnants of the disappearing neurals are
joined. ‘The whole outer surface of the carapace is covered by a
thick armour of epithecal plates, corresponding to the shell of
Dermochelys.

This epithecal shell may be described first. As in Dermochelys
it consists of several rows of longitudinally keeled plates separated
by a mosaic of smaller plates without ridges. The keeled plates
forming the median dorsal ridge are roughly hexagonal and elon-
gated in a longitudinal direction ; they present much the appear-
ance of the neurals ef a thecal carapace. The keeled plates of the
two upper lateral rows are seen also to be irregularly hexagonal,
but shorter than those of the median row. ‘T'wo or three isolated
keeled plates (Pl. Il. fig. 3), to which the outer points of ribs
are adherent, are much larger than the rest and are nearly
quadrate in outline; they were probably near the margin of the
carapace and are thinner than the central plates. The exact
number of keeled ridges cannot be determined, but there were at
least seven. The central part of the carapace preserved shows a
median and two lateral ridges, while the large (@ marginal) plates
above referred to bear another , and others may have been present,
since it is unknown how much of the carapace is missing
between the outer part of the main fragment and the margin;
the distance must have been considerable to allow for the
narrowing of the broad ribs towards their outer pointed ends.
In Dermochelys a median and three lateral ridges are present,
seven in all, but Volker * thinks that probably there were
originally nine.

* Volker, “ Dermochelys coriacea;’ Zool. Jalybiicher (Anatcmie), vol. xxviii.
(1913), p. 477.

ZEUGLODONT AND OF LEATHERY TURULE. 315

The plates filling the intervals between the ridges are smaller
than the keeled plates and of more irregular forms (PI. IT. fig. 2).
So far as ean be seen, between each pair of ridges there are only
two rows of these plates, except that here and there a small irre-
gular plate may be intercalated. This comparatively small number
of intermediate plates seems to be a primitive character, since in
Psephophorus and Dermochelys there is a progressive increase 1n
their numbers: in Psephophorus there were at least six rows and
in Dermochelys they are very numerous. The epithecal plates
are very thick and massive; in the median row the thickness of
the lateral portions is about 10mm., while at the ridge it may be
as much as 16 mm. In the lateral plates above noticed, the
general thickness has decreased to 7mm. or, at the ridge, 9 mm.
According to Seeley *, the plates of Psephophorus may attain a
thickness of nearly 10 mm., but are usually thinner. The outer
surface of the plates is beautifully sculptured, with a, series of
irregular tuberosities in the middle bordered by ridges more or
less radially arranged, and running to the margin which is usually
bevelled off, so that the suture between neighbouri ing plates runs.
at the bottom of a groove. In Psephophorus also the surface of
the plates 1s sculptured, but the ornament appears to consist
of radial ridges only and to be less pronounced than here.
Probably in life the outer surface was covered by a leathery skin,

possibly by horny plates, though this does not seem likely-
The presence of this strongly marked ornament seems to show
that this Turtle was not adapted for rapid motion through the
water, but was probably a littoral or even a swamp-living form
hike Trionyx. It is unfortunate that nothing is known of the
plastron.

Text-figure 2
=,

Transverse section through epithecal and thecal shell of Cosmochelys.

cost., costal plate; ep7., epithecal shell; 7.7., lateral ridge of ditto; m.2., mediam
ridge of ditto ; ., neural plate; ».a., neural arch; x.s., neural spine; 7. rib,
+ nat. size,

The thecal skeleton, so fav as preserved, consists of the remnants
of the costals and neurals. The costal plates are confined to the
upper end of the ribs, where they are fairly well developed,

* Seeley, “ Note on Psephophorus polygonus v. Meyer,’ Quart. Journ. Geol. Soc
vol. xxxvi. (1880), p. 406.

316 DR. C. W. ANDREWS ON A NEW SPECIES OF

projecting forwards and backwards so as te unite with one
another in a short suture; external to this they narrow rapidly
to the edge of the broad and strongly developed rib. Towards
the middle line of the carapace the costals are free from the ribs
for a short distance, forming flanges which project towards the
vestigial neurals, but do not reach them. In this respect the
degree of reduction is intermediate between what is seen in
Protostega and Archelon, in which the greatly reduced costals still
meet the neurals and unite with one another, and in Dermochelys,
where the upper borders of* the costals, though marked by a
distinct ridge, do not form flanges projecting towards the
neurals, which are, in fact, absent or perhaps in part represented
by the bilobate upper ends of the neural spines; moreover, in
Dermochelys the costals do not unite with one another. ‘The
upper ends of the ribs are thick and no doubt had a considerable
articulation with the vertebral centra, but this region 1s abraded
and only the portion articulating with the neural arch remains.
The distance between the heads of the ribs of opposite sides is
about 33 mm. The most anterior of the ribs preserved appears
to be curved backwards towards the rib behind and to have had
the costal plate on the hinder side only: this was, probably, the
first rib. The outer free portion of the other ribs is very broad
and strongly developed.

The neural arch bears a short stout neural spine, to the upper
end of which is attached a thin, flat, table-like neural, which
projects on either side for about 7 mm., but remains separated
from the upper edge of the costal by an interval of 10 mm.

In the figure (Pl. IL. fig. 4, text-fig. 2) the relations of the
eostals, neurals, ribs, and neural arches to one another and to the
epithecal shell is well shown. It will be noticed that the median
ridge of the epithecal shell is not immediately above the middle
line of the underlying neural, but in the crushing that has been
undergone has been displaced to one side. This probably indi-
cates that in life the epithecal and vestigial thecal shells were
separated by a considerable layer of soft tissue now represented
by a thin film of matrix only.

Of the remainder of the skeleton nothing is yet known. It is
greatly to be desired that engineers and others who are on
the spot where excavations are opened, would collect all the
fragments that are exposed, especially in tropical localities like
Nigeria, where fossils once uncovered are rapidly disintegrated
and cuttings very soon overgrown and rendered inaccessible. Of
course, 1t would be still better if, when such works are in progress,
a skilled collector could be present to see that nothing was lost.

The Chelonian above described approaches most nearly to
Psephophorus, but, as pointed out, differs from this in several
respects, notably in the arrangement of the plates on the
epithecal shell, and it is therefore referred to as a new genus
Cosmochelys, the specific name being Cosmochelys dollot in honour

ZBUGLODONT AND OF LEATHERY TURTLE. Bhs

of Professor L. Dollo, who has done so much to elucidate the
history of this group of Chelonians.
This specimen is of especial interest, because it appears to help

Text-figure 3.

Inner surface of part of carapace of Cosmoenelys (R. 4338). +} nat. size.

c.p., costal plate; z.p., median epithecal plate; 7.s., neural arch ; 7, rib.
A-B, line of section shown in text-fig. 2 and PI. II. fig. 4.

318 DR. C. W. ANDREWS ON A NEW SPECIES OF

to fill up one of the gaps in the series of forms Which end in
Dermochelys. According to Dollo’s* view, the Cheloniide and
related forms, which no doubt were derived from littoral and,
more remotely, from terrestrial types, underwent a reduction of
their thecal armour in consequence of their becoming more and
more adapted to pelagic life; this reduction seems to have
culminated in the later Cretaceous, in such forms as Alloplewron,
Protostega, Archelon. Dollo then supposes that some such pelagic
form re-adopted a littoral or perhaps a swamp life and that these
new conditions rendered a protective armour again necessary.
This was not formed by a re-development of the thecal skeleton,
but by the formation of an epithecal armour external to it.
Whether, as Dollo seems to believe, this was an entirely new
formation or resulted from the increased development of epithecal
elements already present, as Volker, Hay, and others think, is not
certain. The presence of epitheeal elements in the neural and
probably in the supramarginal regions of the shell of Archelon
gives some support to the latter view, especially as it is generally
agreed that the marginals of all Chelonians are of epithecal
origin. However this may be, it appears that in forms like that
now described a very strong epithecal skeleton with a strongly
sculptured outer surface was developed outside the reduced thecal
shell, and this could only have been of use to a littoral or even
partly terrestrial animal. This stage in the series should theoreti-
cally occur in the early Tertiary period, and it is precisely from
this horizon (Lower or Middle Kocene) that Cosmechelys comes.
It is supposed that subsequently this or some similar form
returned to a pelagic mode of life, which in turn resulted in the
reduction of the epithecal skeleton. In the Oligocene-Miocene
genus Psephophorus, this has not advanced very far, and is chiefly
manifested inthe multiplication of the number of plates between
the ridges and in the less strongly developed sculpture of the
surface. In the culminating form, Dermochelys, the reduction
has advanced so far that the plates of the carapace are very thin,
smooth, and very numerous between the ridges: in the plastron
they have almost entirely disappeared. Unfortunately, in
Cosmochelys nothing is known of the plastron, either thecal or
epithecal, but pr obably the epithecal was well developed.

The discovery of further remains of this interesting Chelonian
will be awaited with great interest.

* Dollo, “ Premiére Note sur les Chéloniens oligocénes et néogénes de la
Belgique,’ Bull. Mus. roy. Hist. Nat. de Belgique, tom. 5 (1888), p.- 59. Also.
“Sur V Origine de la Tortue Luth (Dermochelys coriacea),” Bull. Soc. roy. des
Sciences médicales et naturelles, 1901. Also “ Hochelone brabantica .... et
PEvolution des Chéloniens marins,” Bull. Acad. roy. des Belgique, 1903, p. 792, and
other papers.

Lists of papers relating to the origin of the Athece are given by Versluys, Report
Brit. Assoc. 1913 (Birmingham), p. 806; and by Volker, on “Ueber das ‘Stamm,
Gliedmassen-, und Hautskelet von Dermochelys coriacea,” Loologische Jahrbiicher
(Anatomie), vol, Xxxiii. (1912-13), p. 543.

ZEUGLODONT AND OF LEATHERY TURTLE.

EXPLANATION OF THE PLATES.

PLATE I,
Pappocetus lugardi, gen. et sp. nov.

Fig. 1. Mandible, outer side. (Type-specimen M 11414.) 4 nat. size.

2, 2a. Outer and inner aspects of first molar of specimen M 11086.

3. ? Incisor (M 11087). Nat. size.

4, Anterior face of imperfect axis vertebra (M 11089). 4 nat. size.

Prins JOE

Cosmochelys dolloi, gen. et sp.nov. (Type-specimen R 4338.)

Fig. 1. Outer surface of middle portion of epithecal shell. 3 nat. size.
2. Outer face of intermediate plates of epithecal shell. Nat. size.
3. Outer face of two lateral plates of epithecal shell. 4% nat. size.
4. Vertical transverse section through thecal and epithecal shell.

c.p., costal plate.
er, central ridge.
ép., epithecal shell.
L7v., lateral ridge.
Ny neural plate.
1.Sp., neural spine.
Res Toop

319

3 nat. size.

Nat. size.

le 4S, IG, CARVER, Pl it

XIPHIAS GLADIUS.

a
t

Ee er boe Oat gh mett

2, 2,8, i910, CARTER, Pl, im,

ISIS IT KOPN HORMOS.

oy
;
es
a
4
?
i
'

I 4 8; (Ol@, CARTIER, IP, Wut

1 BEENNIUS, 21ce) EAC EIS:

DENTICLES ON THE SNOUT OF XIPHIAS GLADIUS. 321

On the Occurrence of Denticles on the Snout of Xiphias
gladius. By J. Taornton Carter (Hon. Research
Assistant, Department of Zoology, University of
London, University College).

{Received June 16, 1919: Read June 17, 1919.!
(Plates I.-III.*)

The purpose of this communication is to describe the appearances
seen in the examination of a portion of the rostrum of a young
eghias gladins which was kindly given by Dr. Smith Woodward,
F.R.S., to Professor J. P. Hill, ith FR S., who handed it to me for
repor a since [am engaged, miner his direction, in an investigation |
of the histogenesis of dental tissues.

Sections of the material disclose the presence of denticles,
hitherto described amongst Teleosteans only in various Siluroid
fishes.

These little conical denticles (Pl. I. fig. 4, 2) are seen to rest on
pediments () to which they are attached by a substance having a
translucent appearance (2), and this area of attachment invariably
coincides with the point of contact of epithelium (e) and the
underlying connective tissues (7). These pediments are connected
with their neighbours by trabecule (Pl. I. fig. 4, tr), and so a
continuous bony layer is formed overlying the premaxilla (Pl. I.
fig. 4, pv), with which however it enters, at certain places, into a
Goapernie so intimate that no line of demarcation is visible
(Pl. i. fig. 1). Where the section passes transversely through
the ae of such a pediment it is seen to have an annular shape
(Pl ig. 2. ) the lumen of which is occupied by a continuation
of the dental papilla. '

The denticles develop (PI. I. fig. 3) as is usual, from a
mesoblastic papilla which is invested by an ingrowth of the
deeper layer of the oral epithelium to form an enamel organ
consisting of a double layer of cells, the innermost of which is
made up of columnar cells, the ameloblasts, whilst the outer
layer or external epithelium consists of cuboidal cells.

I have not been able to procure a portion of the restrum of an
adult Xiphias for comparison with my sections of the young, but
Mr. Tate Regan, F.R.S., has kindly given to me a piece of the
rostrum from a skeleton of an adult Histiophorus and also a
fragment of the skin taken from the area of junction of the
rostrum with the rest of the skull. This portion of skin
contains, lying in its substance, the small tips of denticles, which
would seem to show that they are developed in the same manner
as those seen in the sections of Yiphias.

But, as the rostrum elongates, a change occurs in the site of
development, for now the forming denticles are seen to lie in

* For explanation of the Plates see p. 325.

Se MR, J. i. CARTER: DENTICLES ON

crypts at the sides of the bases of denticles already erupted and
attached (PI. IT. fig. 1, d). .

As these successional denticles grow their predecessors become
detached and shed, their removal being brought about by absorp-
tion at the base. In sections of the rostrum there is no trace of
the superficial denticles, which consist of typical fine-tubed dentine
(Pl. IT. fig. 2, d) becoming embedded in the substance of the bone.
The denticles are larger along the lateral margins than on the
upper and lower surfaces.

‘The presence of denticles in any Teleostean fish is a point of
interest, for the question arises “‘ What part do the denticles
play in the development of the bony layer to which they are
attached”? Goodrich (Proc. Zool. Soc. 1907, pp. 763-764)
states :—‘‘ Nor is there any evidence that denticles do ever
really contribute to form dermal bones. Even in the case of the
palatal bones of fish and amphibians, the teeth do not actually
combine to build up the supporting bone, but become fused
sooner or later to bony substance independently developed at
their base ” (the italics are mine).

On this important point I am unable to agree with Goodrich
for reasons advanced later in this paper.

Denticles develop in the soft tissues and usually are supposed
to obtain a secondary attachment to the bone, but Iam not aware
of any detailed account of how such attachment takes place.
In the absence of fixed material of older Swordfishes I have
had to turn to other fishes to trace out the development of the
denticle and its pediment and how attachment between them is
effected. I realize the objection can be raised that the conditions
obtaining in the case of teeth in the mouth may be adaptations
to special conditions, but the manner of development of denticles
in Xiphias and Histiophorus, and their mode of attachment,
appears to be identical with that of teeth in the various areas of
the mouth in other Teleosts.

As in the case of the denticle of Xiphias, the point of attach-
ment of the tooth to its pediment is always at the point of
junction of epithelium with mesoderm. Whether development
takes place in a bony crypt or deep in the soft tissues, far from
the surface, it is invariably found that the epithelial inflection,
the enamel organ, extends down the whole Jength of the tooth
to the transparent area, where it ends.

In Pl. III. fig. 1, which is a photomicrograph of a section
through the pharyngeal plate of a Blenny, a developing tooth is
shown lying beneath the functional teeth. It is seen to consist
of a tooth (d), a translucent zone (¢z), and a pediment (p), all
forming on the surface of one papilla (dp). The epithelium
(c) is seen to extend down to the limits of the tooth.

On examining the functional teeth it is apparent that the
relationship between the various parts still exists: the teeth are
still connected, by means of the translucent area (¢z), to the
pediments but now these latter are joined one to another by

THE SNOUT OF XIPHTAS GLADIUS. BAe

trabecule of bone (é). When the teeth first erupt they are
slightly movable and easily detached, breaking’ away at the
point of junction of tooth and pediment, but rapidly they become
so firmly anchylosed that considerable force is required to detach
a tooth. The line of fracture, however, is still the same.

In Pl. IIL. fig. 2, a developing tooth of a Sea-Bream is shown in
which the relationship of the various parts is seen to be similar
to that described in the Blenny. The continuity of the cells on
the surface of the dentinal papilla (d.p) is well shown. I have
followed out a complete cytomorphosis of these cells in several
Teleostean fishes and find that the changes through which they
pass are identical whether they go to form the denticle, the
pediment, or the connecting area.

Pl. IIL. fig. 3, exhibits a transverse section through the pedi-
ment of a recently erupted jaw-tooth of a Sea-Bream. It is seen
that trabeculze (¢7) pass outward from the surface of the pediment
(p) and, traversing the connective tissues, blend with those of
adjoining teeth already fixed in place. These trabecule are at
first thin, flexible, and transparent, but rapidly thicken until
eventually a section through this area presents the appearance of
an almost solid plate of bone.

Once firm attachment is effected the pediment in its growth
downwards no longer retains its annular form but divides into
separate trabeculae, the bone cells on the outer surface being
continuous with and indistinguishable from those on the inner
surface and derived from the dental papilla. The puip chamber
of the erupted tooth becomes almest completely obliterated, the
typical dentinal structure being maintained, whilst coarse tra-
becule grow inwards and fill the Jumen of the pediment.

Briefly put, the tooth with its pediment develops deep in
the tissues as a single entity, and not until it has assumed its
functional position does its pediment enter into connection with
those of adjacent teeth to form a continuous supporting plate ;
the tooth invariably retains its characteristic dentinal structure
but the pediment in its growth undergoes a gradual and pro-
gressive transition, both on its outer and inner surface, from
dentine to bone and becomes incorporated in the substance of
the bone. f

The translucent zone (¢z) constituting the junction of tooth and
pediment, though formed before the eruption of the tooth, calcifies
latest and usually appears to have a glassy structure.

The mode of development described above for the teeth holds
good for the denticles in the young Xiphias, for though the
soft tissues are somewhat macerated, my material being from
a specimen preserved in spirit, yet the developing denticles
which have not yet entered into any connection with the bone
are found to consist of a cap of dentine and a pediment, connected
by a transparent area, and, later, trabeculze may, be seen forming
to join adjacent pediments. These pediments are seen to stand up
above the level of the surrounding bone, but this bone continues

Proc. Zoot. Soc.—1919, No XXIII, 23

324 MR. J. T. CARTER: DENTICLES ON

to thicken and grow until its surface lies almost at the level of
the transparent areas.

The denticles in the adult Histiophorus, developing as they do
in the substance of the bone, may be said to have become teeth,
and the appearances seen in sections from the rostrum conform
with the observations described in the Blenny and the Bream.

Mr. Tate Regan has most generously given to me the post-
temporal bone of one of the Tha icariide., an adult Pseudacanthus
seriatus, the examination of which confirms the account I have
given as to the mode of development and attachment of the
denticles and their part in the formation of the bone.

The surface of the bone is studded with denticles, some fully
erupted and so firmly attached as not to become displaced in the
preparation of ground sections of the whole bone, others but
partly erupted and freely movable in their sede which are
widely open like the denticles in the rostrum of Histiophorus
(Pl. If fig. 1,d). Where denticles have become detached the
surface presents an appearance of circular areas with slightly
raised edges, connected one with another by radiating trabecule
and, but for the soft tissues being completely removed, affording
a picture identical with that seen in Pl. UT. fig. 3.

A ground section reveals the denticle, composed of hard tubed
dentine with a tip of enamel, resting on a pediment which on its
outer surface merges into the con oundins bone, whilst towards
the pulp piamrbes: the structure aagern bles coarse dentine; the
denticle is continued downwards a little way into the cavity of
the pediment, or in other werds, is slightly socketed. In the
anchylosed denticles attachment to the pediment is effected by a
layer of almost structureless calcified tissue which occupies the
area of Junction.

The condition is identical with that seen in certain of the
Gadidee where the tooth, the pediment, and the connecting
substance develop on the surface of the one dentinal papilla in
the manner described earlier in this paper. At first the line of
termination of the base of the tooth is straight as in Pl. ITI.
fig. 1, but gradually the inner surface of the connecting trans-
parent zone becomes converted into dentine to form the little
downward extension seen in sections. Eventually, in certain
areas of the mouth, the remainder of the connecting zone calcifies
and the tooth is firmly anchylosed into place.

It is interesting to compare this description with that given by
Williamson (Phil. Trans. 1851, Part II. p. 659) of the “scale of
Loricaria cataphracta. We wrote : —

“On the surface of the scale there are numerous small circular
cavities which communicate inferiorly with branches from the
network of Haversian canals. Hach cavity is surrounded by a
narrow projecting rim, upon which the flanging shoulders of the
tooth rest, whilst its eonstvieted base is hited into the enelosed
hole, thus producing an arrangement which closely resembles a
ball-and-socket joint, and which must allow of a considerable

THE SNOUT OF XIPHIAS GLADIUS. 325

degree of motion in every direction. The tooth is apparently
held in its place by a capsular expansion of the membrane which
covers the surface of the scale........ It only requires the
tooth to be fixed instead of movable, and depressed instead ot
acuminate, in order to render it the exact homologue of one of
the areolz in the cosmine of Jegalichthys.”

Since teoth or denticle, pediment, and connecting area are
formed on the surface of the same papilla, it seems certain that
the difference of structure seen in the fully developed hard
tissue is due to some influence exercised by the investing tissues.
Over the area invested by epithelium the typical dentinal
structure obtains; where the investment is mesodermal there is
an approximation to bone, whilst at the point of junction of
epithelium with mesoderm a layer is formed which, when
calcified, presents a glassy appearance.

Though I have advanced evidence to show that the bone to
which the teeth or denticles are attached is not independently
developed, but is ‘‘an extension of the denticle cone,” so that the
sharp line of demarcation drawn by Goodrich between the tooth-
bearing bones in Teleostei and the bases of Placoid scales does not
exist, yet I have refrained, purposely, from discussing the very
interesting theoretical problems which arise from a comparison of
the conditions described in this paper with those existing in other
orders of fishes. Until we possess a satisfactory account of the
process of histogenesis of bone and dentine in fishes, based on
properly fixed material, we cannot answer certain essential
questions. Only the possession of fresh facts can advance our
knowledge beyond the position so admirably presented in the
classical papers of Williamson and of Goodrich.

My cordial ‘thanks are due to Mr. F. J. Pittock, of the
Zoological Department of University College, for the photographs
used to illustrate this communication.

EXPLANATION OF THE PLATES.

The letters used have the following significance :—

d, denticle. Pp, pediment.
d.p, dentinal papilla. pme, premaxilla.
e, epithelium. tr, trabecule.
m, mesoderm. tz, translucent zone.
Prats I.

Fig. 1. Xiphias gladius. The bone developed by the junction of trabecule (¢7)
connecting the pediments (p) is seen blending with the surface of the
premaxilla (pma). X90.

2. The same. The circular pediments () are seen connected one with another
by means of trabecule (¢) which also attach them to the neighbouring
bone. X 40.

Fig. 3. The same. On the right-hand side of the figure denticles (d) are seen
developing in the soft tissues; denticle (@) and pediment ( p) are seen
connecting by a translucent zone (tz). > 40.

Uys

le>|
on
S

DENTICLES ON THE SNOUT OF XIPHIAS GLADIUS.

Fig. 4. Thesame. A number of denticles (¢) are seen resting on their pediments (p).
The connecting translucent zone (¢z) is seen to correspond ‘with the
junction of epithelium (e) with mesoderm (m). ‘he continuous
scaffolding of bone which supports the denticles is not connected at

any point in the section with the premaxilla. > 40.

Fig. 5. The same. A section showing the relationship of the premaxilla to the
secondary bone. X 40.

Prats II.

Vig. 1. Histiophorus, Photograph of the surface of the rostrum showimg
developing denticles (d) lying im widely open crypts. ‘The erupted teeth
are anchylosed in place. X 7d.

Fig. 2. The same. Photomicrograph of a portion of a section of the rostrum
showing two denticles (d) attached to the surface of the rostrum. X 40.

Prare ITI.

Fig. 1. Blennius. Photomicrograph of a section of the pharyngeal! plate showing a
developing tooth (d), its pediment (p), and connecting substance (¢z).
The epithelium (e) extends down to the limits of the tooth.

The pediments of the erupted teeth are joined one to another by means
of trabecule of bone (¢7). > 40.

Fig. 2. Pagellus centrodontus. Photomicrograph of a developing tooth showing
the continuity of the cells of the dentinal papilla (dp). X 90.

Fig. 3. The same. Photomicrograph of a transverse section through the bone

underlying the teeth, showing the pediment (py) of a newly erupted tooth
becoming attached to its neighbours by trabeculae (é). > 45.

Pe ZoS5 Ie, SIMS ING, Ik i

Del., T. R. R. Stebbing.

PELTARION SPINOSULUS (White) juv.

1 Ze Ss IGG, SIMASISUNG, Pk, We

Del., T. R. R. Stebbing.

ZOEA of a BRACHYURAN.

1ES 25 Se HONG) SIMEIBISMING. IB, Wit

Del. T. R. R. Stebbing.

MEGALOPA of an OXYRRHYNCH.

PZ. Ss. 1919; SHEBBING, Pl ive

S » ot eee

Del., T. R. R. Stebbing.

TANAIS NIERSTRASZI, n. sp.

Del., 7. R. R. Stebbine.

lea 4otSs IDI, SMEZSITING, I, VW.

MUNNA ANTARCTICUS (Pfeffer).

a re

ON CRUSTACEA FROM THE FALKLAND ISLANDS. BAL

20. Crustacea from the Falkland Islands collected by
Mr. Rupert Vallentin, F.L.8S.—Part III. By the
Rev. THomas R. R. Srespine, M.A., F.R.S., F.L.S.,
Ose

[Received June 13, 1919: Read June 17, 1919.]

(Plates I—-V.* and Text-figures 1-8.)

In re-examining Mr. Vallentin’s Malacostraca I have observed
some forms, chiefly specimens of very small size, which seem
worthy of notice in this concluding report.

Brachyura. Isopoda Genuina.

Tribe Cy cLOMETOPA, Tribe FLABELLIFERA.
Family ATELECYCLID®. Family erp

4 Pavey
Genus Peltarion Jacquinot.

Peltarion spinosulus (White) juv. Genus Alga Leach.

Aiga semicarinatus Miers.
Family PortTUNID®. ‘

aN. ilw > ,
Larval genus Zoea Bosc. Family Spo mRomipa.

= \ a aa An G “i
TihSins OUR SeiRIRIN ROTEL. Genus Dynamenella Hansen.

Larval genus Megatopa Leach. Dynamenella eatoni (Miers).

Macrura Anomala. Tribe ASULLOTA.

Tribe GALATHEIDEA. Family Munnrp2.

Family GALATHEID2. Genus Munna Kroyer.
Genus Munida Leach. | Munna antarcticus (Pfeffer).
Munida gregarius (Fabricius). ‘
Amphipoda.
Schizopoda.
Family Lystanassip a.
Tribe THYSANOPODACEA. é
Maye Genus Tryphosites Sars.
Hamily Tvs NOFODEDZ. Tryphosites chevreuxi Stebbing.
Genus Nematoscelis Sars.
Nematoscelis rostratus Sars. Family Mrerorip®.
(Cyrtopia stage.) :
Genus Metopoides Della Valle.

Isopoda Anomala. Metopoides parallelocheir (Stebbing).
(or Apseudacea).

Rael yall a Family PonroGENEIID®.
Genus Tanais Audouin & M. Edwards. Genus Paramera Miers.

Tanais nierstraszi, sp. 0. Pauramera austrinus (Bate).

* Kor explanation of the Plates see p. 339.

328 REV. 'T. R. R. SLEBBING ON CRUSTACEA

MALACOSTRACA.
BRACHYURA.
Tribe CYCLOMETOPA.
Family ATELECYCLIDS.
Genus PELrartion Jacquinot.

The systematic position of this genus and its nomenclature
have been already discussed in Proc. Zool. Soc. for 1900, pp. 518—
519, where also bibliographical details are supplied for the
following species.

PELTARION sPInosuLus (White) juv. (PI. I.)

Writing as to his collecting of Crustacea between November
1901 and March 1902 Mr. Vallentin says :—‘“ I have dredged one
specimen of this species in Stanley Harbour in 3 fins. in the black
mud. It is common in certain protected bays fringing the ocean,
being easily procured during low-water spring tides. Its presence
can at once be detected by a slight blister in the sand. About
an inch deep in the sand under one of these mounds a crab can
always be secured. Gulls, Larus dominicanus, are splendid
fellows at finding these crabs. With one dig with their bill and
a twist they turn them out from their hiding places, and directly
tear them in pieces and devour them.”

Though the adult form of the species has long been well known,
IT have not been able to find any description of the juvenile
stages, one of which seems to me to be represented by the minute
specimen which I have figured.

The carapace measures about 3mm. in length by 2 mm. in
breadth, while the adult may have a breadth of 50 mm. and a
length somewhat greater. The eyes of the small specimen are in
the Megalopa stage, and the five spinulose teeth on each side of
the carapace to the rear of the eyes and the spinulose eminences
along its medio-dorsal line must undergo considerable modifi-
cation in the later development. On the other hand, the micro-
scopically denticulate rostrum and many other details are strongly
in favour of the proposed identification. Many points of agree-
ment may be observed by comparing the account which Miers
gives of the genus (‘ Challenger ’ Reports, vol. xvii. p. 210, 1886)
with various details here figured. Attention may be called to
the third maxillipeds ; to the chelipeds (prp. 1) with the “‘ fingers
robust, scarcely as long asthe palm, and rather obscurely dentated
on the inner margins, distally acute; the dactylus spinuliferous
on the superior margin,” as described by Miers for the adult
male, and here only differing by the greater length of the fingers
in relation to the palm, the other perzopods also agreeing with
Miers’s description, “‘ dactyli styliform, slender, and much longer
than the penultimate joints.”

co
Lo
io}

FROM THE FALKLAND ISLANDS.

Family PornTUNIDS.
Larval Genus Zona Bose. (PI. IT.)

1769. Monoculus Slabber, Natuurkundige Verlustigingen, part 5,
paso; pl. o. wes! 12.

1802. Zoeaw Bosc, Hist. Nat. Crust. vol. 11. p. 135.

1813. Zoe Leach, Edinb. Eucyel. vol. vii. p. 389.

1818. Zoéa Leach, in Tuckey’s River Zaire Exp., Appendix 4,

p. 414.
1830. Zoeq Thompson, Zoological Researches, vol. 1. [Milne
Edwards}.

1837. ,, Milne Edwards, Hist. Nat. Crust. vol. 1. pp. 431-438.
1878. Zoéa Claus, Untersuch. des Crustaceen-Systems, pp. 1, 31,
63, ete.
1903 ,, Williamson, Fishery Board Scotland, Rep. xix. pt. 3,
p- 136.
1911. ,, Williamson, Fisheries, Scotland, Sci. Invest. 1901,
Now:
1918. Zoea Meek & O. Jorgensen, Rep. Dove Marine Lab.
pp. 23, 62.

Slabber’s description and figure of his Monoculus tawrus seem
to give him priority in the observation of this form of crustacean
life. By his laudable anxiety not needlessly to increase the
number of genera he has lost the credit, such as it is, of giving
it its first generic title. Yet he recognised the absurdity of
including in the definition of JMJonoculus ‘‘oculi duo,” whether
expressed or implied in the plural “ oculi approximati.” It may
be noticed that Leach gives a very uninstructive figure of his
Zota clavatu. The account by Milne Edwards of fluctuating
opinion down to 1837 is of great interest, as is that by Claus
later on. Professor Meek proposes that the term Zoea should
be limited to the larve which have “more than eight but not
more than thirteen pairs of appendages.’ The specimen which
I have figured from the Falklands shows much likeness to that
repr esented by Claus (Loe. cit. pl. x1.) as the Zoea of some member
of the family Portunide.

Tribe OXYRRHY NCHA
Larval Genus Mecatora Leach, (PI. IIT.)
1813. JJegalopa Leach, Edinb. Encyel. vol. vii. pp. 394, 431.

1816. 33 » Encycl. Brit., Suppl., Ed. 5, p. 417.

1818. * ie in Tuckey’s Riv er Zaire Exp., Appendix 4,
p. 414.

1825. BA Desmarest, Consid. gén. Crust. p. 200.

1837. Megalops Milne Edw: rds, Hist. Nat. Crust. vol. ii. p. 260.

1874. 4 S.J. Smith, Inv ert. Vineyard Sound, p. 237 (531),

pl. 8. fig. 38.

330 REV. T. R. R. STERBING ON CRUSTACEA

1876. Megalopa Claus, Untersuch. des Crustaceen-Systems, pp. 66
etc.

1911. Megalops Williamson, Fisheries, Scotland, Sci, Invest.
1909 pp. 42°65 Sho:

1918. dMZegalopa Meek, Rep. Dove Marine Lab. p. 30.

1918. Megalops Olga Jorgensen, Rep. Dove Marine Lab. p. 61.

Various other references will be found indicated in the works
above cited. In 1769 or 1770 Slabber, in his ‘ Natuurkundige
Verlustigingen, Part 18, p. 159, pl. 18. fig. 1, deseribes and
figures ‘‘an oblong-quadrate sea-crab,” the size of a grain of
wheat, which is no doubt a J/egalopa, but Slabber supplies no
Latin name. In 1783 Herbst in allusion to its size named it
Cancer granarius (Naturg. Krabben und Krebse, Parts 2—5,
p. 107, pl. 2. figs. 28a, A.). His reproduction of Slabber’s figure
is not specially accurate. Later on, in the third volume of O. F.
Miiller’s ‘ Zoologia Danica,’ edited by Abildgaard (p. 56, pl. 114.
figs. 1-3 ; 1789) appears Cancer faeroensis, also with a tridentate
front, and recognised by Milne Edwards (loc. cit. p. 262) as a
Megalopa. In 1804 Montagu described and figured his Cancer
rhomboidalis (Tr. Linn. Soc. vol. vii. p. 64, pl. 6. fig. 1), a species
apparently belonging to the Cyclometopa, and on this Leach in
1813 founded his genus Jlegalopa, renaming Montagu’s species
as Megalopa montagui (Malac. Pod. Brit. pt. 14, pl. 16. figs. 1-6;
1817).

The Jegalopa of Cancer irroratus Say has been carefully
ascertained by 8. [. Smith, and as the adult is clearly allied with
Cancer pagurus, presumably the Jegalopa stage will be nearly
the same in the two species. The MWegalopa of Carcinus menas
figured by Spence Bate is reproduced in Huxley’s ‘The Cray-
fish’ (p. 282, figs. 74, C, D. ed. 3; 1881) by a slip under the
name of C. pagurus. Williamson, who uses Jlegalopa as the
plural of Megalops, supplies figures of this stage for Portunis
holsatus, Portunus puber, and a species which he believes to be
Ilyas araneus. As our Falkland Island specimen shows good
agreement with the last-named form it may reasonably be allotted
along with it to the Oxyrrhyneha, leaving open the question of
its genus and species.

MACRURA ANOMALA.

Tribe GALATHEIDEA.

Family GALATHEID®.
Genus Munrpa Leach, 1820.

MunipA Grecarivs (Fabricius), 1793.

The adult form has been already mentioned in these Proceediv es
for 1914, p. 846. The figures here given refer to a very early

FROM THE FALKLAND ISLANDS. Sal

larval stage, in which the carapace has only a length of 2 mm.
The generic identification may, I think, be relied on by a com-
parison with the description and figures which Professor G. O.
Sars supplies for a similar stage of Munida rugosus (Fabricius)
in his * Bidrag til Kundskaben om Decapodernes Forvandlinger,”
ii. p. 178, tab. 6 (Arch. Naturv., 1889).

The figures give a dorsal view of the specimen in two divisions,
the line v.s. indicating the actual length of the carapace.

Text-figures 1 & 2.

Munida gregarius, early larval stage.

SCHIZOPODA.
Tribe THYSANOPODACEA.
Family Tuysanopopip®.
For the classification see Ann. S. African Mus. vol. vi. pp. 395,
396: 1910. .
Genus Nematroscenurs G. O. Sars.

1883. Nematoscelis Sars, Vid. Selsk. Forhand]. Chrvistian., No. 7,
Dey ile

NEMATOSCELIS ROSTRATUS Sars.
1885, Nematoscelis rostrata Sars, Rep. Voy. ‘Challenger,’ vol.
xii. Schiz., pp. 135, 169, pl. 25. figs. 8-10, pl. 31.
figs. 23-29.
Among numerous specimens of larval forms belonging to other
groups there occurred a single slender form 4°5 mm. in length,

B32 REY. T. R. R. STEBBING ON CRUSTACEA

having a telson in minute agreement with that figured by Sars
for the Cyrtopia larva of his species above-named. His descrip-
tion of the telson says, ‘‘The middle projection of its extremity
(fig. 29) is considerably produced, but narrowly truncate at the
tip; and of the seven original spines, three only remain. Of the
three outer spines, the fer nnast on either side is much larger
than the others, and has assumed the character of the subapical
spines.” The outermost, as shown in the figure 29 is microscopic,
and in the upper part of the telson but below the middle (not
included in fig. 29) there is another microscopic pair. ‘The cara-
pace has a denticle on each side below the middle. The first legs
in the Falkland specimen, however, have not attained the same
relative length as that shown in fig. 25 of the ‘ Challenger ’
report. In various papers H. J. Hansen makes JV. rostratus a
synonym of WV. microps Sars. On this I am not presuming to
pass an opinion, but retain the name rostratus for the better
identification of the Falkland Island specimen with the ‘ Chal-
lenger’? Cyrtopia form.

ISOPODA ANOMALA.
(or Apseudacea).

Family TANAID&.

See Proc. Zool. Soc. 1914, p. 348, and add for the present
purpose :—

1884. Tanaide Studer, ‘ Gazelle’ Isopoden, p. 24.

1886. Rs Beddard, Rep. Voy. ‘Challenger,’ vol. xvii.
Part 48, p. 119

1914. As Barnard, Ammo A SaqeAir. Mus. volvxt ata lle
p. 331 a.

Genus Tanais Audouin & M. Edwards, 1829.

TANAIS NIERSTRASZI, sp. n. (PI. IV.)

The present species belongs to that division of the genus in
which the pleon has six segments. In having the last three
abruptly narrowed it agrees with 7. normant Richardson,
differing from it by having the ramus of the uropods 10-joited.
In this respect it stands between the large blind 7. willemoesii
Studer, which has 8 joints, and 7’. hirsutws Beddard, which
has, including peduncle, “‘about 12.” From the latter, taken
“* off Prince Edward Island; depth 50 to 150 fathoms,” it appears
to be distinguished by the very different proportions of many of
the hody segments.

The eyes are dark, piriform, at the rounded angles of the
cephalothorax, which has a broad front with short rostrum, and
gradually attains a breadth at least equal to the length. The

FROM THE FALKLAND ISLANDS. 333

first antenne are as in 7’. hirsutus, with crowded sete on joints
of the peduncle, but only a minute one-jointed flagellum tipped
with long sete.

The mandible ends in four crowded teeth or short sete from
which a narrow strip of the trunk leads to the strong molar.

Text-figures 3-8.

gi. Te

Tanais nierstraszi.

a.s. First anteana. m. Mandible. ma. 1. First maxilla. map. Mavxillipeds.

gn. 1. First gnathopod. wrp. One of the uropods in attachment to part of
pleon ; the ramus should be 10-jointed.

The first maxilla has its oblique apical margin spinose, with a
group of subapical sete on the outer margin; the long two-
jointed palp ends in several sete. The maxillipeds have the

304 REY. T. R. R. STEBBING ON CRUSTACEA

apex of the palp’s fourth joint, like the two preceding joints,
provided with a crowd of sete. The broad third joint is apically
narrowed. The first gnathopod is normal, the fingers closing
without a gap, and the apical teeth overlapping. The second
gnathopods are very slender. In the fifth pereeopods the penulti-
mate joint has the lower half of its front margin fringed with
small spines. The rami of the pleopods have very long fringes.

A specimen nearly 7 mm. in length was taken by Mr. Vallentin
at Roy Cove from a depth of 3-4 fathoms. The smaller specimen,
5 mm. long, he took from the surface. To this the text-figures
reter.

The specific name is given in recognition of Professor H. F.
Nierstiasz’s valued studies of the Isopoda.

ISOPODA GENUINA.

Tribe FLABELLIFERA.
Family AiGip &.
Genus Alga Leach, 1815.

AGA SEMICARINATUS Miers.

1875. ga semicarinata Miers, Ann. Nat. Hist. ser. 4, vol. xvi.

Deals

We fae a, $5 Miers, Phil. Trans., Zool. Kerguelen,
Grewtsines Jos 2p Jol stale tives, I,

Woh, sag 5 Dollfus in Crust. Miss. Cap Horn (A.

M.-Edw.) p. 57, pl. 8. figs. 2, 2 a.
1914. ,, wrotoma Barnard, Ann. 8. Afr. Mus. vol. x. p. 367,
pl. 32 A.

In a manuscript note Mr. Barnard identifies his wrotoma with
the present species. His figure of the telsonic segment, however,
does not show nor does his description mention the slight medio-
dorsal carina which is recorded and figured by Dollfus and is
present in the Falkland specimen. This was found by Mr. Val-
lentin on drift Macrocystis near West Point Island. It measures
49 mm. in length, with a breadth rather over 21mm. In the
first antenna the flagellum is 10-jointed. The difference of four-
teen joints in that of the Cape specimen cannot be considered
important, as the total length of the Cape example was also
larger, being 53 mm.

Our specimen has the whole dorsal surface of the pleon and the
last side-plates of the person strongly pitted. In the first
gnathopods the fourth and fifth joints are very short, the sixth
has a minute process on the inner margin, and the seventh is
strongly bent with the apex acute and black.

FROM THE FALKLAND ISLANDS. 335

Family SPH HROMIDS.

Genus DYNAMENELLA Hansen.

1905. Dynamenella Hansen, Q. J. Microsc. Sci. vol. xlix. pp. 96,
Oe ly, 125).

1905. a H. Richardson, Mon. Isop. N. Amer. p. x.

1906. - H. Richardson, Pr. U.S. Nat. Mus. vol. xxxi.
jon tes

INES 5 Barnard, Ann. 8. Afr. Mus. vol. x. p. 410.

DYNAMENELLA EATONI (Miers).
1875. Dynamene eatonit Miers, Ann. Nat. Hist. ser. 4, vol. xvi.

4 (hase
1891. t » Dollfus, Crust. Miss. Cap Horn, p. 66.
1905. Dynamenella eatoni Hansen, Q. J. Microse. Sci. vol. xlix.
p- 125.

Mr. Vallentin’s specimens, taken on the shore at Stanley
Harbour and from a depth of 3 to 4 fathoms in Roy Cove, were
all females.

Tribe ASELLOTA.

Family MuUNNID 4.

1899. Mannide Sars, Crust. Norway, vol. 11. p. 105.
1916. AMunniné (group) Hansen, ‘ Ingolf’ Malacostraca, iii. p. 33.

Genus Munna Kroyer.

1839. Munna Kroyer, Naturhistorisk Tidsskrift, vol. i. p. 612.
1882. », Chilton, Ann. Nat. Hist. ser. 5, vol. ix. p. 1.

1887. Haliacris Pfeffer, Krebse von Siid-Georgien, Part 1, p. 97.
1899. Munna Sars, Crust. Norway, vol. 11. p. 106.

1902. Haliacris Hodgson, ‘Southern Cross’ Crustacea, p. 253.
1905, Munna H. Richardson, Isop. N. Amer. p. 480.

1906. Haliacris H. Richardson, Exp. Antarct. frangaise, Isop.,

p. 16.
1909. ce Chilton, Subantarctic Is. N. Zealand, Crust.,
p. 6950.
1910. x Hodgson, Nat. Antarctic Exp., Isopoda, p. 58.
IO > H. Richardson, Deuxsieme Exp. Antarct. fran-

caise, p. 19.
1916. Junna Hansen, ‘ Ingolf’ Malacostraca, iii. p. 34.

The species of this genus have caused no little ditticulty by the
smallness and transparency of some parts and the great length
and fragility of others. Some curious slips of the pen may also be
noticed. Thus Sars attributes the genus to Boeck, just after
writing of it as Kroyer’s. Pfeffer in defining Haliacris states

336 REY. T. R. R. SLEBBING ON CRUSTACEA

that the second to the fourth pairs of walking-legs are longer
and stronger than the fifth to the seventh pairs, though his
specific description shows that he means just the reverse. Hodg-
sen in describing the mandible says of the palp, “ first and third
joints subequal, third the longest,” his figure showing correctly
the second joint as the longest. Chilton and Hodgson, with a
lingering retention of the name Haliacris, agree that the name
must be regarded as a synonym of M/wnna. Hansen points out
that the character “eyes distinct” must be withdrawn from the
definition given by Sars, if the genus is to include such species as.
Munna ceca Richardson, J. truncata Richardson, and MW. acanthi-
jfera Hansen. But he does not notice Miss Richardson’s proposal in
1908 (Pr. U.S. Nat. Mus. vol. xxxv. p. 79) to substitute the generic
name Cecimunna for the species truncatus and Haplomunna for
the species cecus. Should these proposals be adopted, Hansen’s.
acanthifer would probably be allotted to Cecimunna, thus with-
drawing all the blind species from JZunna. In 1913 Miss
Richardson advocates the retention of Haliacris on the ground
of the special structure of the first gnathopods in the male and
their great size, This distinction would require the inclusion,
along with Pfeffer’s species, of J/unna palmatus Lilljeborg, 1851,
and Munna neozelanicus Chilton, 1892. But it is at least highly
inconvenient to have the adult male in one genus, while the
females and young males can be appropriately placed in another.
In J. kréyert Goodsir the carpal joint of the male’s first gnatho-
pod is large, while in J/. palmatus it is very much larger, but
surely this by itself should not count for generic difference. In
instituting his genus Pfeffer was himself unacquainted with the
full development of the flrst gnathopod in the adult male.

Monwa Antarcricus (Pfeffer). (Pl. V.)
1887. /aliacris antarctica Pfeffer, Krebse Siid-Georgien, Pt. 1,
p. 97, pl. 6. figs. 28-47.

1902. ir australis Hodgson, ‘Southern Cross’ Crust.,
p. 293, pl. 34. figs. 1 a-d, pl. 37.

1906. ¥, a H. Richardson, Exp. Antarct. fran-
caise, p. 16, fig. 20.

1909. se antarctica Chilton, Subant. Is. N. Zealand, Crust.,
p. 650, fig. 14 6.

1910. Bf a Hodgson, Nat. Antarct. Exp., Isop.,
pp. 58-61.

1G) 15), H. Richardson, Deuxiéme Exp. Ant.

frangaise, p. 19.

Mr. Hodgson says of the specimens obtained by the ‘ Dis-
covery’ that some of the old males “attain a length of seven
millimetres.’’ None of the Falkland Island specimens exceeded
3mm. Yet the single example of an adult male first gnathopod
is very characteristic of the advanced development. It differs
slightly from the only other available figure, given by Miss.

FROM THE FALKLAND ISLANDS, 337

Richardson in 1906, as there theinner margin of the large carpal
joint’s process is serrate, in place of the well-marked inner tooth
of our specimen.

In the first antennze I found two stout joints, the second
longer than the first ; to the second succeeds a minute joint which
L suppose to be the third joint of the peduncle. It is followed
by a similar joint which should, I think, be considered the first
of the slender flagellum.

In the male the second antenne may attain a great length,
fully twice that of the body, the transparent flagellum slightly
exceeding that of the peduncle. Pfeffer’s figure gives this fla-
gellum without any divisions, and those which I have marked
are very uncertain, notwithstanding the high magnification.
As shown on the Plate the specimen carrying this long antenna
on the right had on the left one very much shorter, and the
peropods on the left are rather shorter than those on the right.
Mere size has to be carefully considered before it can be used in
clussification.

The curved third joint of the mandibular palp seems naturally
to bend away from the cutting-edge rather than towards it. In
the maxillipeds the broad plate of the second joint has three or
four minute hooks on the inner margin and four little teeth on
the truncate distal border.

The first pleopods of the male are described by Hansen as the
“median lamella of the abdominal operculum of that sex,” and
for specific distinction he says ‘“‘in reality the shape of this
lamella, especially its terminal part, affords, perhaps, the sharpest
and most reliable character.” Unfortunately in small specimens
its details are excessively difficult to determine. Even for the
larger divisions of the perzeon my figures cannot claim exactitude.

The specimens were obtained by Mr. Vallentin from a hulk
at low water.

AMPHIPODA.
Family LystaAnassip®.
Genus TryPHostres Sars.
TRYPHOSITES CHEVREUXI Stebbing.
1914. Tryphosites chevreuxi Stebbing, Proc. Zool. Soc., p. 355, pl. 3.

The oviginal description states that in this species the third
pleon segment “has the lower half of the postero-lateral margin
convex and cut into a serration of nine little teeth.” An exam-
ination of additional specimens shows the variability of this
character, a small example having only three such teeth, and
one somewhat larger having four on one side of the pleon and
six on the other side.

In J. Linn. Soc. vol. xxix. p.58; 1903, Mr. A. O. Walker

338 REY. T. R. R. SCEBBING ON CRUSTACEA

describes specimens of Atyloides serraticouda Stebbing with
seven teeth on the margin above discussed, instead of only two
in the specimen first deseribed. The moral which Mr. Walker
dvaws as to the untrustworthiness of small characters for specific
distinction is enforced by the additional example in Zryphosites,
a genus remote from Atyloides. But it is difficult to profit by
the warning when a single specimen with apparently novel
characters has to be classified.

Family Mreropips.

Genus Merroporpes Della Valle.
MesroporDES PARALLELOCHEIR (Stebbing).

1888. Metopa parallelocheir Stebbing, Rep. Voy. ‘ Challenger,’
vol, xxix. p. 762, pl. 43.

1893. J/etopoides 4 Della Valle, F. Fl, Neapel, vol. xxi.
p. 907.

1906. se a Stebbing, Das Tierreich, vol. xx1.
p. 186.

The specimens obtained by Mr. Vallentin at the Falkland
Islands had unfortunately become too dry for satisfactory ex-
amination in detail before I attempted dissection. Beyond
identifying the species I can add nothing to the description and
figures supplied in the ‘Challenger’ report and ‘ Das Tierreich.’
The depth of 100 metres from which the ‘ Challenger’ specimen
purports to come loses such authority as it had by comparison
with Mr. Vallentin’s taking of the species at very small depths.
They were found by him “in the branchial sac of a simple
ascidian.”

Family PoNtroGENEIID.
Genus ParaAMa@ra Miers.

1875. Paramera Miers, Ann. Nat. Hist. ser. 4, vol. xvi. p. 79
(see also Rep. Voy. ‘Challenger,’ vol. xxix. p. 447).

ParaAM@RA AUSTRINUS (Bate).
1914. Paramera austrinus (Bate), Proc. Zool. Soc., p. 364.

Specimens which I aim inclined to include in this seemingly
variable species were taken by Mr. Vallentin some nine years
ago at Crooked Inlet. In regard to the first of them he writes :
“Tt was found under the mantle of the common limpet Patella
cenea. Colour, body ivory-white with a dark red line running
down directly in the median line from head to tail. | Eyes fiery-
red.” It is remarkable that the body colouring was retained till

FROM THE FALKLAND ISLANDS. 339

examination in the year 1918, though the eyes had become orange
iather than red.

Chevreux in 1913 deseribes his Stebbingia gracilis as having the
hody “teinte de blanc et de rose” and the eyes “ d’un rouge vif.”
It also agrees with our Falkland specimens in having no accessory
flagellum to the first antenna, and the telson slit for half its
length, but the smoothly rounded apex of each lobe is devoid of
the spiule which our specimens have, and the slender spinose
perzeopods cannot be reconciled with the comparatively stout and
smooth lower joints of the Falkland species, of which ‘‘ six more
specimens removed fiom as many different limpets were found
later.” One of these smaller examples, however, proved to belong
to a different genus. As to the Paramara specimens, so far as
IT have been able to verify the details, they agree with those
which I have figured in the ‘Challenger’ Amphipoda, pl. 76, the
species being there named Atyloides australis (Miers).

EXPLANATION OF THE PLATES.

Prare Td.
Peltarion spinosulus (White) juy.

v.s. Lines indicating size of carapace in the adjoining dorsal view.

r., a.8., a.i. The rostrum ; first and second antenna magnified to the same scale as
the rostrum and mouth-organs.

m., mx. 1, mv. 2, map. 1, 2,3. Mandible, first and second maxille, first, second, and
third maxillipeds.

prp.1, prp. 2. Four terminal jomts of the first and second perzeopods, with fingers
of first to scale of the mouth-organs.

Prare IT.
Zoea of a Brachyuran.

a.s. Line showing length from apex of frontal to apex of dorsal spine in specimen
figured below im lateral view.

car. Carapace more cularged, dorsal view, frontal spine omitted.

T. 'Telson to the same scale as the antennee and anterior mouth-organs.

1.8., @.S., @.i., m., mx. 1, mx. 2. Upper lip, first and second antenne, mandible, first
and second maxillee.

mep. 1, 2, prp. 1,3, 5. First and second maxillipeds, first, third, and fifth perzeopods,
to a lower scale than preceding details.

Prarn III.
Megalopa of an Oxyrrhynch.

».s. Line showing length of specimen figured helow in dorsal aspect.
car. Profile view of carapace and base of pleon.
PI. Vast three segments of pleon im dorsal view, more magnified.

a.s., a.i. First and second antenne.

m., mx.1, mx. 2, mep.1, 2, 3. Mandible (part), first maxilla (palp incomplete), second
maxilla, first, second, and third maxillipeds.

prp- 1, prp. 4, plp. First and fourth persopods (less magnified than the other
details), a pleopod, and terminal joints of prp. 1 on higher scale.

Proc. Zoon. Soc.—1919, No. XXIV. Q4

340 ON CRUSTACEA FROM THE FALKLAND ISLANDS.

Prace LV.
Tanais nierstraszi, sp. Vi.

a.s. Line showing length of specimen figured below in two aspects.

oc., a.8., at. Eye; first and second antenne more highly magnified.

l.s., m., map. Upper lip, mandible, maxillipeds ; these, the end of prp. 5 and the
urp. on higher scale than the other details.

gn. 1, gn. 2, prp. 5. First and second gnathopods and fifth perzeopod.

plp., wrp. Pleopod and uropod.

Puate V.
Munna antarcticus (Pfeffer).

n.s. 6. Line showing length of male specimen, roughly sketched, incomplete.

a.s., @.t., 9. Dorsal view of female; first antenna more highly magnified.

Pl. 2, ov. Pleon of female highly magnified ; ovum to the same scale.

m.3,m. 2, mxp., op. 6. Mandibles of male and female; a maxilliped ; opereular
lamella of male; all more magnified than other details.

ai. 6,gn.1, 6, gn.1, 2, prp.1, prp. 5. Second antenna; first gnathopod of a
male (from separate specimen), first gnathopod of female, part of first
pereopod, fifth perzeopod; all to one scale of magnification.

FIELD-NOTES ON MAMMALS IN THE SOUTHERN SUDAN. 341

21. Field-Notes on some Mammals in the Bahr-el-Gebel,
Southern Sudan. By Major J. Stevenson HAmMILron,
O.M.Z.8.

"Received October 18, 1919: Read November 18, 1919.]
(With Chart.)

It should be noted that my intimate personal experience
extends only to the area between Bor and Shambe, and from
these points eastwards to a distance of 80 miles, N. and N.K. of
Shambe 60 miles, and west of the river about 20 miles. It is
only in this area that I have been resident. As regards the
region between Mongalla and Bor, I have spent only two months
of the dry season there, and south of Mongalla I know the
country only from occasional short visits.

Tatane (Damaliscus tiang Heugl.).

West bank.—Through the flat country from Lado northwards
to Shambe.

Kast bank.—From a little south of Bor northwards to the limit
of the area considered.

Found east of the Nile in much greater numbers than to the
west. Country probably more suitable. ‘The great open plains
in the Nuer, and in the west of the Dinka country seem especially
fitted to their habits. The type found west of the river is of
much deeper coloration and contrasted markings than that occur-
ring west of it, forming perhaps a different local variety.

During the dry months Thiang may be seen on these plains in
countless numbers. In the vicinity of water-pools herds of from
1000 to 2000 are common, and two or three such herds may be in
sight at one time.

The habits of the Thiang appear more akin to those of the
Blesbuck than to those of its other cousin the Tsessobe, which
latter is never found in droves and seldom far from forest or

those great plains by the water, which covers the country during
several months of the year, and at such times they break up into
small family-parties of from half a dozen to a score of individuals,
something after the fashion of the Tsessobe, and then roam
through the bush-country or wherever there is dry ground, while
large numbers doubtless migrate far to the east or south-east to
higher country.

The calving season is regular, lasting from about the middle of
February to about the middle of March—that is to say, it takes
place just previous to the advent of the first rains, and in this
assimilates closely to the habit of the Tsessobe.

While in large herds in the open, Thiang are very wary and
will seldom allow approach nearer than 250 yards; probably at

24*

31°E. Long. a2?

CHART OF STATION OF CERTAIN
ANIMALS IN BAHR-EL-JEBEL AREA
SUDAN

MALEK eee

|
3°

Miles 10 5 O 10 20 39 40 50 60 76 Mites

Bubalis telwel a) Gazella albonotata

Equus bureneni

Damaliscus corrigum Riinoceros simus

i, I 1 Kobus leucotis

x xX

ee Kobus maria

Kobus vaughani

rae Kobus kab thomasi

FIELD-NOTES ON MAMMALS IN THE SOULHERN SUDAN. 343

such a distance considering themselves safe from the lions, which
follow the great herds everywhere, as also from the natives who
oceasionally hunt the full-grown animals. In the bush, and when
in small parties, on the other hand, they appear easier to approach
than most game.

The species doubtless tends to be of larger size and to bear
longer horns as it trends north: but to divide it into two sub-
races, with a dividing-line about the latitude of Lado, is with-
out justification. Although it is unusual to meet with Thiang
south of Bor along the river, they of course exist on the great
inland plains, which are a continuation of those of the Nuer and
Dinka countries, and doubtless the herds make their seasonal
migrations south-eastwards towards the Abyssinian and East
African borders, when the winter grazing-grounds become water-
logged.

During the dry season of 1918-19 the Thiang, which had
migrated westwards in immense herds, were collected in thou-
sands over the Duk and “Toich” country between lats. 7° 15’
and 7° 45’ in the north of Bor District (E. of Bahv-el-Gebel).
A virulent form of dysentery broke out among these great herds
about the middle of March 1919. This disease: atch in its
symptoms appeared indistinguishable from the dysentery which
affects human beings, was undoubtedly specific in nature and was
spread by the droppings of the sick animals fouling the grass.
That this was the case was proved by cattle grazing over the
same ground becoming affected, while herds kept on clean eround
ren aimed healthy. The stomachs of dead animals were usually
erammed with food—the lungs, liver, and heart healthy, but the
gall-bladder unnaturally distended. Animals rapidly lost con-
dition, but the disease seemed too rapid in its effects to give time
for more than a staring coat and a generally pinched appearance.
Animals turned away from the herd usually sought the shelter
of a tree or long grass, stood for a few hours, then lay down, and
presently died.

The disease was first noticed in the herds grazing in the swampy
land bordering the river, which had been under water for two
years and was covered with long rich thick grass, but it quickly
spread to the other herds which were grazing on the short sweet
grass of the Duk country 30 miles inland, and affected them to
an extent quite as great as the others. I did not happen to see
the remains of any animals except Thiang which had died,
although there were large numbers of can Giraffe, Gazelle,
Reedbuck, Oribis, and a few Waterbuck and Zebra in the same
country. I calculate that perhaps rather more than 10 and less
than 15 per cent. of the total Thiang died in about a month.
With the advent of the rains the big herds broke up, and the
dry germs of disease could no longer be distributed by the wind,
which probably accounted for the cessation of the sickness.

I believe that the disease, which T do not consider to have been
rinderpest, although it closely resembled the latter in some of its

344 MAJOR J. 8S. HAMILTON: FTELD-NOTES ON

symptoms, was originally brought to the river by herds migrating
from far inland, probably from the Abyssinian boundary. I feel
the more confident of this because natives stated that it often
made its appearance in years of extreme drought, and that some-
times all species of game, including elephants, died from it.

Harreseest (Lelwel) (Bubalis leliwel).

West bank.—To about 6° North (roughly the latitude of Tombe).

East bank.—Extends about as far north as Bor, and this is
about its limit northwards. It is never met with in either the
Lau Nuer or the Twi Dinka countries (N. and N.E. of Bor). Its
northern limit is therefore practically identical on both the west
and the east bank, and it slightly overlaps the southern limit of
the Thiang along the river on both banks, more so on the west
than on the east.

Nite Lecuwe. Mrs. Gray’s Waterbuck (Aobus maria).

The most southerly limit of this species undoubtedly is about
halfway between Tombe and Mongalla at about 5° 30’ N., where
in the dry season there is always a herd on an island.

North of this pomt they may be encountered in suitable
localities on both banks of the river. A little south of the Bahr-
el-Zaraf and east of the Bahr-el-Gebel they exist in the Jonglei
swamps, where I have seen them. They are, however, more
numerous on the west bank, not continuously, but in isolated
herds considerable distances apart. An extraordinarily tame herd,.
which can never have had any experience of firearms, exists at one
point. The Dinkas hunt the young ones with dogs, but if a man
is alone, unaccompanied by a dog, and strolling along very slowly,
the animals after a time will disr egard his presence and may be
approached within 50 yards in the open. This herd consists of
about 120 full-grown animals, of which a dozen or so are dark-
coloured males.

The part of the country in which this herd is found is a big
reed-swamp bordering the river, opening out into a great, flat,
treeless plain, which is cut up into sections by a number of
swampy streams averaging 20 feet wide by 3 feet deep. Of
course, during the rains this is all inundated. If the herd finds
itself separated from a man by one of these swampy streams it
seems to consider itself perfectly safe. When chased by natives
the animals make at once for the nearest of these channels, and,
dashing across, halt on the other side. Should the natives follow,
the petiormmnice is repeated to the next channel. The herd
spends most of its time on its open grazing-ground, where it
can be seen from a distance of several “miles - away, and appears
only to use the river reed-swamp as a place of refuge. I think,
however, that the females with younger calves spend their
whole time therein, until the calves are able to run and swim
well. Although the usual calving-time is seemingly January and
February, I have never seen any small calves with the herds, and

SOME MAMMALS IN THE SOUTHERN SUDAN. 345

those caught by the natives are usually, they say, found in the
swamp. Of course, the dogs sometimes get a half-grown one in
the open, but the natives say they never catch a full-grown one.

The habits of these animals are almost exactly those of the
Zambesi Lechwe, as is their general appearance, method of hold-
ing their heads when running, and the manner in which the heid
splashes through shallow water. When the plain is water-logged,
I have seen the whole herd standing in water halfway up to
their hocks, just as I have seen the Lechwe herds in the Kwando
swamps of the South and elsewhere.

The young ones seem rather delicate and difficult to rear in
captivity.

I have never seen in this herd the old dark rams without the
white wither-patch spoken of by Col. Roosevelt. The younger
rams have an indistinct patch, but the older the animal the more
defined the patch, in my experience. There are aberrant types in
all species, or possibly in some localities there may be variety of
type, but I do not think anyone else has ever had so unique
opportunities of studying a herd at close quarters as I have
had, and in a large one, such as this, there cannot fail to be
animals of all ages; besides which I have been able to watch each
individual male separately, and am sure none of them display
this peculiarity.

Cos (Xobus).

Cob extend all the way along the west bank of the Bahr-el-
Gebel from south of Rejaf to beyond Shambe in suitable localities.
There is a very gradual but well-marked variation in type from
the quite brown-eared specimens seen near Rejaf through the
buff-eared types from between Mongalla and Tombe to the
perfectly white-eared but rufous-bodied animals found opposite
Bor. The latter agree with what is known as Vaughan’s Cob in
all characteristics ; but they shade off gradually southwards into
the typical Uganda Cob found on the same side of the river. On
the east pane of the Nile “station” is much more irregular.
However, Uganda Cobs are found more or less sporadically from
Mongalla northwards, but, so far as I know, the intermediate type
is not found, and after a gap wherein no Cobs oceur, the typical
Kobus lencotis begins to occur in the plains near Jonglei.

vine Vaughan ” Cobs opposite Bor were seen in open country
near forest, where a herd of some sixty grazed with several parties
of Reedbuck and Thiang. ‘They had the Cob-like habit of
mounting ant-heaps to scrutinize intruders. This herd had never
been molested, and I found it very tame and easy to approach so
long as concealment was not attempted.

Probably the extreme permanent southern limit of the true
leucotis is at Jonglei on the east bank of the Bahr-el-Gebel, lat.
6° 45' N.; here they are tolerably numerous over an area of about
20 square miles in a strip along the river. They spend the nights
and mornings during the dry season, as a rule, in the open plains,

346 MAJOR J. §. ILAMILYON : FIELD-NOTES ON

inland from the forest strip, where they graze and eat the lolob
fruit. About 1 p.m. they begin to saunter back to the forest near
the river, where they stand in the shade, and, having about 4 p.m.
drunk at one or other of the forest-pools, they graze about in the
forest until sunset, when they return to the open country 2 or 3
miles away.

Northwards from Jongiei no more Cob are met with south of
the Bahr-el-Zaraf, where, as is well known, they are very
numerous.

There are no lewcotis on the western bank of the river south
of Shambe, lat. 7° 15’, and no Uganda nor Vaughan’s Cob on the
east bank north of, at most, lat. 5° 15’. Thus, whereas. the
Uganda Cob shaded by perceptible gradients into the rufous
white-eared animal found near Kenisa on the west bank, there is
no form intermediate between the true White-eared Cob found
oa the east bank, and the true Uganda Cob found very much
farther south on the same bank. As regards breeding-season, I
saw no young animals during the dry season, and, from the size ‘of
the immature animals, am of the opinion they lamb about May
or perhaps late April. In the wet season the White-eared Cob
disappear from Jonglei and migrate long distances to the east,
probably following the course of the iehores, towards the Abyssinian
foothills in common with other game.

ZEBRA (Hquus quagga).

Two opportunities occurred—one in early July and the other
In late February—of inspecting herds at very close quarters. No
animals possessed what, by any stretch of imagination, could be
termed a mane, though a small minority of both sexes had a very
slight ridge of hair about an inch long and very thin and ragged.
This appeared about equally the case in young and in old animals—
in fact, some of the colts and fillies looked like newly-hogged polo
ponies. It seemed a matter of individual variation. On the
other hand, a very young foal brought to me from another herd,
which I did not see, had a long fully developed hog-mane exactly
like that seen among the southern types of Zebra. This was
from a place 20 miles from where I saw the herd in February,
and was in the month of April.

The herd which £ saw in July was in lat. 6° 10’, and that in
February was in lat. 7° 15’.

I do not think the ie of mane can be rightly attributed to

_seasonalism, for one of the herds was seen well on in the cool wet
season and the other in the middle of the hot dry season. In any
case, I do not think there is so much as 40° Fahr. difference
between the hottest day and the coldest night here all the year
through. ,

Shadow-stripes seemed entirely a matter of individual variation.
Some members of the herd, apparently without special reference
to sex or age, had very strongly developed shadow-stripes on the
quarters, while others showed absolutely no trace of them. Of

a ee

SOME MAMMALS IN THE SOUTHERN SUDAN, 347

the two stallions shot, both over 6 years old, one had strongly
marked shadow-stripes, while in the other any traces of such
marks were entirely lacking. The newly-born foal had indica-
tions of shadow-stripes on the quarters.

All members of both herds without exception had white ears.

The extreme northern range of the Zebra in Bor District east
of Bahr-el-Gebel is at 7° 30',and the small troops occasionally
seen so far north in the height of the dry season may be regarded
as stragglers.

Reeppuck (Redunca).

The type of Reedbuck, formerly very common in the neigh-
bourhood of Mongalla, was distinguished by the peculiarly wide
spread of the horns. The rinderpest epidemic in 1912 almost
exterminated these animals. Nevertheless, the peculiarity of the
branching does not seem to have been confined to Mougalia, nor
does it seem to be typical of any variety so much as of individual
specialisation. I have seen several male Reedbucks as far north
as Duk Fadiat, which is nearly 3 degrees of latitude north of
Mongalla, possessing these same wide-spread horns, but far the
larger number of the males in the locality have the ordinary type
of horns well hooked forward at the tips, which one associates with
Behar Reedbuck.

These Reedbuck are not only much smaller in size than, but
differ considerably in their habits from, the common Reedbuck of
the south. They are usually found grazing in parties with or
near herds of other game in the open, which is foreign to the
custom of Hleotraqgus arundinum. ‘Chey also run with their necks
outstretched, unlike the latter, which always caries the head
high.

GAZELLE (Glazella).

The local Gazelle in Mongalla Province though smaller have
the same general appearance and display the same habits as the
Thomson Gazelle of Hast Africa.

The females bear weak malformed, decadent horns, and both
sexes continually wag their black tails in the familiar manner of
the “Tommy.” The herds and single rams act when approached
just as do the herds of the latter, and they scatter about when
grazing among the crowds of Thiang, which here take the place
of Kongoni Hartebeest,

The external markings do not vary more from those of thomson
than one would expect from a local variety, while the buffy band
which is spoken of by Roosevelt as existing between the black
side-stripe and the white underpait is, I think, purely an indi-
vidual eccentricity, and far from being universally present.

Towards the middle of very dry seasons these Gazelle migrate
northwards and spread along what is known as the Duk country—
where there is always sweet grass and water—as far as lat. 7° 30"

348 FIELD-NOTES ON MAMMALS 1N THE SOUTHERN SUDAN.

but no further. With the first rains they disappear to the south,
and though a few may be found lingering well on into the
summer in the low country east of Mongalla (lat. 4° 30’), IT am
of the opinion that the great majority migrate very long distances
towards the higher and drier country lying in the extreme S.E.
of the Sudan near the Kast African border, and that somewhere
among these plateaux he their summer quarters. Here they
come not very far from the northern range of the true thomsoni,
and are not separated from this animal by any natural boundary
so formidable as that formed by the Sobat River and its swamps,
which cut the species off from G. rujifrons in the north. More-
over, the Mongalla Gazelle never ranges within 100 miles of the
Sobat River, the intermediate country being, in fact, unsuited to
the habits of Gazelles to a considerable degree. Probably the form
is almost exactly intermediate between thomsone and rufifrons,
with tendencies towards the former.
It is not found west of the Bahr-el-Gebel.

Wuaire Ruinoceros (2/inoceros sinus).

This animal exists all along the west bank of the Nile in the dry
season, but is always very rare in the area under consideration.
The natives say they drink at the river at night, and retire great
distances into the forest during the day.

The above notes, which only deal with quite a small area of
country (approximatery 200 miles by 150) and have no preten-
sions to discuss the distribution of the animals mentioned outside
thereof, tend to show how peculiarly patchy even within the same
small area the distribution of any given species tends to be, and
is an example on a small scale of what we often find and with as
little apparent reason throughout the whole continent of Africa.

ON A NEW SNAKE FROM THE TRANSVAAL. 349

22. Descriptions of a new Snake from the Transvaal, together
with a new Diagnosis and Key of the Genus Xeno-
calamus, and of some Batrachia from Madagasexr. By

The Hon. Paut A. Meruven, M.A., F.Z.8.
‘Received October 9, 1919: Read November 4, 1919. |
(Text-figure 1.)

Genus XENOGALAMUS Gthy.

The discovery of a new form of Yenocalamus, described below,
and the examination of the skull of Y. bicolor Gthr. and of
X. transvaalensis, sp. n., calls for a revision of the characters
of this genus, for which I propose the following diagnosis :-—

Maxillary short, with 4 or 5 solid teeth of moderate size
gradually increasing in length posteriorly, followed, after a short
interspace, by a pair of somewhat enlarged grooved fangs situated
below the posterior half of the eye ; palatines toothless or bearing
a few small teeth ; lower jaw with 7 to 9 rather small teeth on
each side, those in the middle of the row the largest. Postfrontal
bone fused with parietal. Quadvate showing tendency to enlarge-
ment and to having direct attachment to the skull, at the expense
of the squamosal which may be very much veduced. Basioccipitals
reniform in profile. Posterior vertebrze without heemal processes.

Head small and elongate, not distinct from neck; snout much
depressed, very prominent; rostral large, with obtuse horizontal
edge, flat or excavate below; eye minute, with round pupil ;
nostril between a large posterior and a small anterior scale, or in
a single shield which may show incipient signs of similar division.
A large preocular ; no loreal; no prefrontals (fused with frontal) ;
no anterior temporal.

Body cylindrical ; tail short ; scales smooth without pits, in 17
or 21 vows; ventrals rounded ; subecaudals in 2 rows.

Tropical Africa, as far south as the Transvaal.

Synopsis of the Species of Xenocalamus.

I. Palatine without teeth ; rostral flat below.
A. Seales in 17 rows; nasal divided ; 6 upper labials, third and fourth entering

the eye.
1. A narrow supraocular ; ventrals 218, subcaudals
OH TON GAOL ame eee aes ban tac setibor.be o padeuneeesendeecseey ite Aucorol Eien
2. No supraocular; ventrals 229 to 239; sub-
GANG EME GIL TO) XO gasasbosn coe a V. mechovii Pet.

B. Seales in 21 rows; nasal entire; 5 upper labials,
second and third entering the eye ; ventrals 257,

STULL NCAT 27 le een don scene qagedsuE ese X. michelli Mull.

* The Transvaal Museum has an adult female specimen of XY. bicolor (‘T. M. Cat.
Rept., No. 1151), with exactly the same number of ventrals and subcaudals as the
type; the specimen is, however, remarkable in that the third and fourth upper
labials are fused into one large scale which alone enters the eye ; in colour it is slate-
blue above, head and neck lighter ; from Rechtuit, Waterberg District, Transvaal.

350 THE HON. P. A. METTLUEN ON A NEW SNAKE,

If. Palatine bearing 3 or 4: small teeth; rostral excavate
below; nasal entire with incipient signs of divi-
sion; 5 upper labials, second and third entering
eye; scales in17 rows; ventrals 195, subcaudals 81. XY. ¢ransvacalensis, sp. n.

XENOCALAMUS TRANSVAALENSIS, Sp. 11.

Description.— Maxillary bone short, with 4 solid teeth of mode-
rate size, the posterior ones the largest, and followed after a short
interspace by a pair of enlarged grooved fangs situated below the
posterior corner of the orbit. Palatine bearing 3 or 4 small teeth.
Lower jaw with about 9 rather small teeth on each side, those in
the middle of the row the largest. Quadrate lnrge, attached
directly to the skuil in its anterior half; squamosal very much
reduced, that part which is visible being only two-thirds the
length of the quadrate (vide text-fig. 1).*

Text-figure 1.

Posterior part of skulls seen in profile :
(A) Xenocalamus bicolor. (B) X. transvaalensis.

Head as broad as neck; snout depressed, prominent but not as
much so as in 1. bicolor ; rostral large, rather acutely rounded,
with rounded horizontal edge, excavate below, in contact with the’
nasal. Nostril pierced in a single scute which abuts on the
rostral (the division of the rostral shield into'a very small and a
large scale as in LY. bicolor is suggested by incipient sutures; in
X. bicolor the sutures are distinct). Internasals large, forming a
median suture, separated from the first upper labial by the
rostral. The large preocular forms a suture with the rostral and
the internasal in front. Supraocular and postocular scales
minute. Frontal very large, rounded in front, more or less heart-
shaped, a little more than half as long as the distance from the tip
of the rostral to the posterior limit of the frontal (actual measure-
ments being 52 mm.:9°4 mm.). Parietals elongate, forming a
long suture, not quite so long as the frontal. Five upper labials ;
the first and fifth very small, the fourth enormous, the second
and third entering the eye; the third forms a short suture with
the postocular, Six lower Jabials, third very large, first and

* In X. bicolor the quadrate differs somewhat in shape and is not as large as
in XV. transvaalensis ; further, the visible part of the squamosal is seen as a curved

horn-shaped process extending a considerable distance across the supratemporal
region (vide text-fig. 1).

AND BATRACHIA FROM MADAGASCAR. 351

second very small; the first pair of lower labials forms a median
suture. <A single ‘pair of rather small chin-shields, which form
sutures with abe first, second, and third lower ipl:

Ventral scales 195, rounded; anal divided; 31 pairs of
subeaudals.

Bo ly cylindrical, slightly depressed.

Blue-black above, below white with dark brown transverse
markings on the ventral scales; throat, lower jaw, and upper lip
nearly entir ely white.

Total length 414 mm., of which the tail measures 44 mm.

A burrowing suake found in sandy soil, north of the Zoutpans-
bergen, Northern Transvaal, near the Ingelel River, within 25
miles of the Limpopo: collected by Messrs. Noomé and Roberts,
in September, 1913: it was observed to be sluggish in its
movements.

Type in the Transvaal Museum, Cat. Rept. No. 1689.

BATRACHIA.

In 1913 Mr. Hewitt and I published an account of a collection
of Batrachia made in Madagascar (2): I have since been able to
compare many of the specimens referred to in this paper with
material in Kuropean Museums, and especially with that in the
British Museum. I have also been able to profit by the criticisms
of Mr. G. A. Boulenger on several specimens I submitted to him.
T gladly avail myself of this opportunity for thanking him for his
advice on many occasions, and among other things fo suggesting
the names of the two new genera described below, and for
pointing out their affinities to the genus M/antidactylus.

GEPHYROMANTIS, gen. nov.

Vomerine teeth present ; digits with supernumerary phalanx ;
terminal phalanges dilated at end, the dilatation with shallow
notch distally, reniform ; lower surface of digits with ring-shaped
groove; outer metatarsals united; style of sternum and omo-
sternum long, slender, and bony ; pupil horizontal; tongue well
developed, bifid behind.

GEPHYROMANTIS BOULENGERIT, Sp. n.

Description.—-Head longer than broad; snout subacuminate ;
nostril a little nearer tip of nostril than eye ; loreal region deeply
concave ; canthus rostralis curved ; diameter of eye very nearly
equals the distance from eye to tip of snout. Interorbital space
equals breadth of upper eyelid. Tympanum distinct, 4 the dia-
meter of eye. Fold over tympanum swollen anteriorly and
posteriorly. Vomerine teeth as in Mantidactylus granulatus.
Fingers moderate, first a little shorter than second, considerably
shorter than fourth, their tips expanded into dises which are
large on the third and fourth fingers, being about double the
breadth of the penultimate joint ; subarticul: ur tubercles of digits
and metacarpus prominent. ‘Toes moderate, 7 webbed, their tips

52 THE HON. P. A. METILUEN ON A NEW SNAKE,

expanded into discs of moderate size, not quite as large as those
on third and fourth fingers; a sliced pear-shaped inner meta-
tarsal tubercle, barely 4 length of first toe; a small outer
metatarsal tubercle.

The tibio-tarsal joint of the adpressed leg reaches a point
between the eye and the nostril. Heels strongly overlapping.
Length of tibia 4 the distance from snout to vent, about 33 times
as long as broad.

Upper surfaces of head, body, and limbs very granular with
numerous warts and elongated tubercles, which on the head and
back form an irregular pattern. Belly, sides, and inner parts of
thighs granular. .

Habit like the JZ. granulatus section of the genus d/fanti-
dactylus (1).

Colour: Above dark bluish-brown with irregular lighter mark-
ings more in evidence on the head than elsewhere ; upper lip,
loreal region, throat, and chest mottled with dirty white and dark
brown ; an irregular median dirty white line on throat and chest.
Upper surfaces of limbs same colour as back, the lighter brown
colour of the inner parts being carried across the upper surfaces
as thin irregular transverse bars.

Length from snout to vent 27°50 mm.; length of outstretched
hind limb from vent to tip of fourth toe 44 mm.

Size of eggs in oviduct 2°75 mm. in diameter.

Type a female, T. M. Cat. Rept. No. 1013, and cotype No. 1012,
in the Transvaal Museum ; origin, Folohy, Hast Madagasear, 1911.

In 1913 Mr. John Hewitt and I placed these two speci-
mens—for lack of comparative material—pro tem. in the genus
Mantidactylus, the genus to which this new form is most nearly
related (2).

TRACHYMANTIS, gen. nov.

Hemimantis: H. horridu Bttgr. Zool. Anz. 1880, p. 282;
Abhand. d. Senck. Gesell., B. xii. p. 492, Taf. 11. fig. 14, 1881.

Arthroleptis: A. horridus Bouleng. Cat. Bat. p. 118.

Microphryne Methn. & Hwtt. (2) p. 55, preoccupied.

The only character I have to add to our original diagnosis
concerns the discs of the digits, as in Mantidactylus and Gephy-
romantis the lower surface of the digits has the characteristic
ring-shaped groove. This character in Mantidactylus was pointed
out by Mr. Boulenger (1) when he made the genus Aglypto-
dactylus for Mantidactylus madagascariensis. J must also correct
what was our impression at the time, that Zrachymantis was
related to Rhacophorus; Gephyromantis, which is very closely
related to Mantidactylus and may even be a comparatively recent
offshoot from a M. granulatus-like form, leads naturally to
Trachymantis ; the relationship can be shown thus :—

Mantidactylus : vomerine teeth ; outer metatarsals separated.
Gephyromantis: vomerine teeth ; outer metatarsals united.
Trachymantis: novomerine teeth; outer metatarsals united.

AND BATRACHIA FROM MADAGASCAR. aoe

In 1913 (2) we pointed out that we strongly suspected Rana
labrosa to be the only truly endemic Ranid in Madagascar which
was not supplied with the supernumerary phalanx to the digits ;
and that Bottger’s ‘* Arthroleptis (Hemimantis) horridus” belonged
to the same genus as our ‘ Zrachymantis (Mier ophryne) mala-
gasia.” In 1914, 1 had. an opportunity of examining Bottger’ S
type at Frankfort with Dr. Sternfeld: we were both “of opinion
that Arthroleptis horridus should be referred to our genus, and
that the two species horrida and malagasia were very closely
allied. This fully confirmed our presumption as to Madagascar
Ranids *.

Besides the differences cited in the two original descriptions,
the following distinctions were noticed : in 7’, malagasia the snout
is slightly longer as compared with the diameter of the eye and
more pointed ‘than i in 7. horrida ; in the latter species the dises
of the digits are larger than in 7. malagasia. The tympanum in
T. horrid is more visible and a shade larger than in the other
species. The femurs in 7. horrida ave glandular, but lack the
huge glands of 7. malagasia which we suggested might be an
abnormal development. In Bottger’s species ae eranules on the
ventral surface end in a sharp point; they might almost be
described as small pointed tubercles: in our species they are
replaced by swollen granules of larger size.

MANTIDACTYLUS ARGENTEUS, Sp. 0.

This species falls into the group with large discs to the fingers,
and with granular belly (1): nearest ally seems to be M/. granu-
lutus : she. upper surfaces have however no asperities,

Description Head flat and depressed, longer than broad ;
snout subacuminate, practically pointed, strongly projecting
beyond the mouth ; nostril considerably nearer tip of snout than
eye; distance from eye to nostril $ diameter of eye. Tympanum
distinct, ? diameter of eye. Interorbital space a “fraction greater
than breadth of upper eyelid, equal to diameter of tympanum.
Loreal region concave; canthus rostralis lightly curved; fold
over tympanum feebly developed. Exposed part of vomers
bearing the teeth not so prominent as in JZ. granulatus, with
large median space between them. Fingers well developed, their
tips expanded into discs about, or a fraction more than, double
the breadth of the penultimate joint; on the first finger, which
is much shorter than the second, the disc is smaller than this.
Toes moderate, their tips expanded into discs which are much
smaller than those of the hand, being barely 4 their diameter.
Toes 2 webbed. Inner metatarsal tubercle rather small, not
prominent.

Tibio-tarsal joint of adpressed leg reaches the nostril or just
short of it. Heels strongly overlapping. Distance from snout to
vent 1? times as long as tibia, which is 5 times as long as broad.

* Vide also (8).

304 THE HON. P. A. METHUEN ON A NEW SNAKE,

Dorsal surfaces without asperities, but with very fine reticu-
lations; some glandular granulations in coccygeal region. Belly,
sides, inside of thighs, and round vent, granular.

Colour: Above brownish-purple or vinous flecked with dirty
white; limbs same colour with hght narrow transverse bars.
Sides marbled with brown and silver; tympanum dirty white
flecked with silver; lower half of eye, loreal region, snout and
lips, dirty white; throat and chest silver white, yellower on chest,
becoming dirty htie on belly and thighs.

Length from snout to vent 32 mum. ; length of leg from vent
to tip of fourth toe 52 mm.

Size of egg in oviduct 2-5 mm. in diameter. :

Type a female, T. M. Cat. Rept. No. 1009; cotype a juvenile
specimen, toes } webbed, tympanum ? diameter of eye, tibio-
tarsal joint of adpressed leg reaching tip of snout, No. 1035;
both in the Transvaal Museum : : origin, Folohy, Hist Madag? scar,

Oval

PLETHODONTOHYLA TUBERIFERA, Sp. 0.

P. notosticta Gthr., Mthn. & Hwtt. (2) p. 60.

Under the name P. notosticta, Mr. Hewitt and I provisionally
identified seven examples of this genus, indicating at the same time
that our specimens were by no means typical (/.c.). I have since

compared ours with the examples of this genus im the British
Museum and find that our specimens repr esent a new form. Its
distinguishing characters are as follows:—Head flat and de-
pressed ; loreal region very oblique; no canthus rostralis; a rounded
or more or less pointed snout projecting only slightly beyond the
mouth ; discs of digits larger than in any other known species of
the genus ; on each: side in the sacral region a small prominent *
tubercle. In the natural arrangement of the genus, P. inguinalis
appears to be intermediate between the new species and
P. notosticta.

Description.—Uead # as long as broad, flat and depressed ;
snout rounded or more or less pointed, extending only slightly
beyond the mouth; loreal region very oblique; no canthus
rostralis ; interorbital region twice the breadth of the upper
eyelid ; tympanum distinct, 4 or nearly equal to the diameter of
eye. ‘l'ongue typical for the genus. Vomero-palatine teeth in a
jong cheyrone shaped transverse series, interrupted in the middle,
and extending just beyond the choanze. Fingers moderate, dilated
into large triangular dises, that on the third finger being # the
diameter of the eye; first finger considerably shoiter than
second, with only slightly expanded disc; hand with flat inner
and outer metacarpal tubercles coalescing medianly. ‘Toes rather
short, expanded into triangular dises, that on the fourth toe being
about 3 the diameter of the eye, that on the fifth toe small. Toes
free*. A flat elongated inner metatarsal tubercle, rather poorly
developed.

* Tn 1913 (7. c.) we stated that the toes had trace of webbing at the base. We

wrongly interpreted the integument which is usually present m a more or less
developed form at the base of webless toes in many frogs as a rudimentary webbing.

AND BATRACHIA FROM MADAGASCAR. 355

Tibio-tarsal joint of adpressed leg reaches as far as the eye.

Upper parts smooth with a prominent little tubercle on each
side overlying the expanded ends of the diapophyses of the
sacral vertebra. Posterior parts of belly, sides, and of thighs
near the vent, granulate.

Colour: Ground-colour dirty white ; dorsal surface blotched or
spotted with dull magenta, darkest on head and snout; the
tubercle on sacral region surrounded by dark horseshoe-shaped
blotch ; a dark line from eye, through tympanum, along side
nearly to a point reached by tibio-femoral joint of adpressed leg.

Length from vent to snout 30 mm.

Types in the Transvaal Museum, Cat. Rept. Nos. 1265 to 1271;
No. 1269 presented to the British Museum. Origin, Ambato-
haranana, forest of Central Hast Madagascar, 1911.

The terminal phalanx is Y-shaped ; in P, notosticta it is broadly
Y-shaped ; and in P. inguinalis it is also Y-shaped. So I think,
in the diagnosis of the genus, this bone should be described as
Y-shaped or broadly Y-shaped, rather than as T-shaped as in the
Brit. Mus. Cat. The internal structure of this species was
examined ; it is typical for the genus.

Libliography.

(1) “On the Madagascar Frogs of the genus JMJantidactylus.”

By G74 Boulenger, Ws. P2729. 1918) pp. 2ov—261),

(2) “On a Collection of Batvachia from Madagascar, made during
the year 1911.” By Paul A. Methuen & John Hewitt,
Beate cA Wransvaale Muss Nosy 2. vols ava 1913)
pp. 49-64.

(3) *‘A comparative review of the Amphibian Faunas of South
Africa and Madagascar, with some suggestions regarding
their former lines of dispersal.” By John Hewitt, B.A.
Ann. Transvaal Mus., April 1911, pp. 1-11.

he
Or

Proc. Zoon. Soc.—1919, No. X XV.

Dey 2

rng os
cave a

ON THE DORSAL SCALE-ROWS IN BRITISH SNAKES, 357

23. On the Variation in the Number of Dorsal Seale-rows
in our British Snakes. By Miss Joan B. Procrsr,
F.ZS.

[Received August 14, 1919: Read November 4, 1919.]
(Text-figures 1-3.)

In a valuable paper* Dr. Ruthven has drawn attention to
certain points in the lepidosis of the Garter-snakes or North
American 7ropidonotus of the group Zhamnophis, and has for-
mulated a series of laws regarding the loss of certain rows of
dorsal scales, which throw much light on phylogenetic questions.
The principal of these are as follows :—

“ That the decrease in the wumber of scale-rows posteriorly is
brought about in all of the forms of garter-snakes by the loss of
certain definite rows.

“That the order in which these rows are lost posteriorly in the
different forms is the same as in the form having the maximum
number of rows for the genus.

“The individual, geographic, and racial variations in the number
of dorsal scale-rows in the garter-snakes is brought about by the
shortening and loss of the same scale-rows as are ordinarily dropped
posteriorly in conformity with the taper of the body, and there is
evidence that this decrease is due to a dwarfing of the body.”

A short time ago Mr. Boulenger, who guides and encourages
me in my work, gave me a copy of this most interesting paper,
and suggested that I should find how far these laws apply to other
snakes. J have therefore made a careful study of our three
British snakes, as representing widely differing types subjected to
the same climate.

In all I have found that there is this abridgment of certain
longitudinal rows of dorsal scales, and that in each of these species
the manner in which this takes place is highly characteristic.

On examining any snake with abridged series of scales, one at
first concludes that these have been formed by the regression of
rows which were complete in the primitive state. All the implicated
Species, however, are not being subjected to dwarfing influences,
as is evidently the case with the Garter-snakes, and in some
groups, notably the Vipers, evolution seems to tend towards
enlargement and increased number of scales. ‘To determine the
manner in which any given abridgment has been arrived at, it is
necessary to consider the scale-formula of the primitive form of the
genus, and, in some cases, to make comparisons with individuals
having the next highest and next lowest formula.

1. Tropidonotus natriw has 21 scale-rows upon what may be
called the neck, the reduction to 19, brought about by the termi-

* “Variations and Genetic Relationships of the Garter-snakes.”’ Bull. 61, Smith-
sonian Inst. U.S. National Museum, 1908.

358 MISS JOAN B. PROCTER ON THE NUMBER OF

nation of the 10th series, usually takes place at about the level of
the 6th ventral plate—a range bearing relation to the scaling at
the back of the head, rather than that of the body. Since the
primitive number of series for the body is probably 19*, it is
this pair of series which would develop in species having 21.
Exclusive of this, the seale-formula is 19-17. The reduction to
17 is brought about by the dropping out of the [Vth series in
accordance with the laws of sequence formulated by Dr. Ruthven
for the Garter-snakes ; which is exactly what one weuld expect, as
these snakes also belong to the genus 7ropidonotus in the wide
sense.
19-17.

The abridged series responsible for the drop from 19 to 17 rows
ave unquestionably in course of reduction. The termination of
the [Vth series (see text-fig. 1) occurs at a point 3/5ths down the
length of the body, or sometimes 5/Sths. Occasionally this series

Text-figure 1.

Tropidonotus natriv, lateral view, to show termination of series IV.,
2ths down the body.

fuses with the Vth, this point being marked by a large scale
bearing two converging keels. The exact range of variation where
the IVth series may be dropped, is °55 to ‘60 of the length of the
body 7, which shows that there is very little individual variation,
and proves that the scutellation of the Grass-snake is in a stable
condition.

2. Coronella austriaca.—On the neck this snake has 21 rows of
scales, the reduction to 19 usually taking place at about the level
of the 8th ventral by the termination of series V. or its fusion
with LV.

The primitive number of scale-rows for this genus is 19.
Series V., therefore, which is only present on the neck of
('. austriaca, becomes progressively developed in species having
21, or more, rows of scales.

The complete scale-formula for C. austriace exclusive of the
neck is 19-17-15, these two pairs of abridged scale-rows being
formed by reduction.

* Tn view of the fact that it is the most frequent number in the genus, as well as

in Colubrids generally, irrespective of the affinities of the species.
+ “Body” in this paper signifies the head and trunk exclusive of the tail.

DORSAL SCALE-ROWS IN BRITISH SNAKES. 359

19-17.

The drop from 19 to 17 rows takes place by the fusion of
series I[V.* with III. (see text-fig. 2) or occasionally with V. (the
Viths of the original 21). The final loss of this row usually takes
place at a point 3/5ths down the body, although the complete
range of variation is from *55 to ‘75,

Text-figure 2.

4
4
es

+t
S

6
3

Coronella austriaca, lateral view, to show fusion of series IV. and III.,
2this down the body.

The fusion of series LV. with IIT. (or V.) is in active progress,
for one finds (@) individuals on which the reduction from 19 to
17 rows takes place at a definite point on the third quarter of
the body, anterior to which the number is invariably 19, and
(6) individuals on which series LV. fuses, or tends to fuse, inter-
ruptedly with IIT. on the entire anterior half of the body, and
continues in this way until that point is reached, on the third
quarter of the body, where the loss is final. Between these two
extremes every intermediate form may be found, the degree even
varying on the two sides of the body. This instability shows a
tendency for this loss to become complete.

17-15.

The reduction from 17 to 15 rows takes place by the suppression
of series IIT. or its fusion with IV. (tke VIth of the original 21),
the final loss being at a point on the posterior quarter of the body,
but subject within this limit to considerable variation, which
suggests that this reduction is comparatively recent, and still in
active progress.

3. Vipera berus.—The apparently most primitive form among
living Vipers, V. ursinii, also has 19 rows of scales, and there-
fore V7. berus, with scale formule varying from 19-17-16 to
23-21-19-17-16, must have four of its abridged series in course
of production, aud three in course of reduction.

23-21.
In the average individual there are 23 rows of scales imme-

* Although in many cases the fusion 1s completed in such a manner as to make it
impossible to say which is the supernumerary row, I judge it to be 1V., as im some
cases the final scales of this row are rudimentary, and alse the occasional union with
the Vth instead of the IIIrd points to this conclusion,

360 MISS JOAN B. PROCTER ON THE NUMBER OF

diately behind the head, the drop to 21 rows taking place by the
termination of series V. at about the level of the 15th ventral.
This series is in course of development, and in several specimens
it crops up on the middle third of the body to a degree varying
from a single scale to a complete row, which, though it commences
by cropping up in an intermittent manner on this region of the |
body of some individuals, develops and progresses both forwards
and backwards in others, and in one specimen unites with its
originated portion on the neck giving this snake 23 uninterrupted
rows for more than half its length. This progressive development
of series V., which in J’. berus does not ever continue beyond the
anterior two-thirds of the body, evidently continues in species
having 25 or more scale-rows.
21-19.

The second pair of series in course of development is the [Vth.
In normal specimens which have 21 rows of scales this series
terminates about two-thirds down the body, usually by fusing
with the IIIvd. In specimens which have 19 rows instead of 21
it is only present on the neck; its development from this point
taking place in precisely the same manner as that of series V

19-17.

The first pair of series in course of reduction is that next to the

vertebral ; its termination takes place on the third quarter of the

Text-figure 3.

ne
RY

tll

are

A NIK

wi (
cae eer

= =u — oo

,
ANN i een
CA ny aud
= =o i ss

body usually two-thirds down. In the normal Viper, which has
21 rows of scales on its anterior two-thirds, this point almost

DORSAL SCALE-ROWS IN BRITISH SNAKES. 361

invariably coincides with the termination of series 1V.; and as the
corresponding points vary on the two sides of the snake, a variety
of combinations result, all of which have the effect of reducing the
number of scale-rows from 21 to 17 in an abrupt and often highly
irregular manner. One such combination of the ending series
was as follows :—21-20, IVth left side: 20-19, Xth right side;
19-18, TXth left side ; 18-17, Vth and I1Ivd united, right side
(see text-fig. 3). In some other specimens the penultimate series
ended over an inch anterior to the [Vth, the regression of the
one pair thus overlapping the progression of the other. This
overlap is always present in specimens having 23 rows, as in
these the amount of regression remains the same, whilst the
progression of series V. and TV. has advanced reciprocally.

17-16.

Just before the commencement of the tail there is yet another
reduction caused by the vertebral row, which drops out usually
within about four ventrals of the end of the body. Occasionally
the loss is less advanced, and constitutes the first reduction on the
tail-proper ; conversely regression may be more advanced, and in
one extreme instance this series ended 22 ventrals before the end
of the body.

Anatomical reasons for the halting-points in the reduction
and production of scale-rows.

It is clear, since the size of the dorsal scale is more or less fixed,
that the number of rows is correlated with the calibre of the body,
and that this number must decrease or increase, if smaller or
larger forms are to be produced. Also, the series must vary in
number according to the taper of the body. The posterior reduc-
tions are amply accounted for by this, but the general taper of
the body seems quite insufficient to account for the constant
halting-point in reduction or production of two, in the case of
V. berus four vows, about two-thirds or three-fifths down the
body. :

Now the number of rows must be regulated by the normal
girth of the body; or by the girth attained by necessary expan-
sion at certain points. It also stands to reason, if scale-rows are
to be dropped, in the case of reduction, that the abridgment will
commence where the loss in girth will be least missed, and pro-
eress in this manner, so that the portion of body where these
rows finally persist will be that part where the skin has the
greatest strain imposed upon it. Conversely this will be the
point where the development of new rows will commence *.

In 7. natrix this portion is from the neck to a point just
beyond the middle of the body. In the Garter-snakes when
further reduction is necessary after the abridged rows have

* In Naia, for instance, the more developed the hood, the more numerous the
scales,

362 MISS JOAN B. PROCTER ON THE NUMBER OF

reached this point, 7¢ takes place from the neck backwards until
complete.

In C. austriaca the TVth, which is in process of medaenens
halts at the same point, somewhere on the third fourth of the
body, and when further reduction is required the whole series
commence to break up by intermittent fusion with its neighbour.

It is thus obvious that the portion of the trunk-proper where
the skin receives the most strain is that anterior to this halting-
point, the greatest strain of all being on the middle third. The
reason for this is the presence of the stomach, a fact which I have
verified by the dissection of 15 spirit-specimens*, confirmed by
the dissection and special study of three fresh specimens, taking
particular notice of the relative positions of the pyloric sphincter,
and the point where the final loss occurs in the involved scale-
rows. In every case the halting-point is on a level with the
pyloric sphincter, or some way posterior to it; in no case does it
occur anterior to the pylorus. ‘To test this I took a small viper
enormously distended with food (spirit specimen) and stuck a pin
into it where it first regained its normal girth. On examination
this pin proved to have been lodged w ithin a ventral of the point
where the IVth series fused w ith the Illvd, on the left side;
dissection proved this point to coincide also with the pyloric
sphincter ; the stomach-proper in this case was distended by a
large mouse. Most cases are of course less exact, as there is much
individual variation, the given points usually varying a little on
the two sides of the body.

If one considers a snake’s manner of feeding, and the length of
time that the stomach remains distended, it is not surprising
that the reduction of scale-rows should be held up where the
strain on the skin is so great, or that new rows should commence
by out-cropping at this region.

Dr. J. C. Thomson, in his paper on ophidian anatomy, has
studied this question of the variation in the number of scale-
rows, together with the external landmarks of the principal
organs, the relative positions being given in term of ventrals.
Of Thamnophis ordinoides he says :—‘‘ he relation existing be-
tween the position of the viscera and the added and suppressed
seale-rows has been studied in fifty specimens of this species.”
“19-21-19-17, that is 19 rows forward, increasing to 21 at about
the level of the heart and continuing to the end of the liver,
then decreasing to 19, and further on to 17.” He does not,
however, mention the oma and its physiotogical bearing on
the question, nor does he record its position in the tables of
“The external landmarks of the principal viscera.’

The high number of rows behind the head can also be accounted
for, this area of skin being subjected to the greatest strain of all,
by the enormous distension of the j jaws in the str ugele to swallow

* 4 T. natrix, 5 C. austriaca, and 6 V. berus.
+ “Further Contributions on the Anatomy of the Ophidia.”” Proc. Zool. Soc, 1914,

—

DORSAL SCALE-ROWS IN BRITISH SNAKES. 363

the prey whole. What would be the use of the highly elastic
ligamentous attachment of the mandible, and the loose attach-
ment of the bones of the skull generally, if the skin over the
back of the head and neck were not also capable of the highest
distension ¢ The scale-rows appear first and last upon this small
area, but the development of fresh rows almost invariably halts
on the neck, and recommences over the region of the stomach,
afterwards continuing forwards and backwards until the scale-
series is complete from the head to the level of the pyloric
sphincter—the second halting-point. It is probable that this
procedure in the production and reduction of scale-rows will be
found to hold good for all snakes.

Sexual dimorphism.

In accordance with these principles, it follows that the female
snake should have, or tend to have, a higher number of scale-
rows than the male, on account of her greater girth. Dr. Ruthven
has already. made this supposition*, but finds that it is not
realised in the Garter-snakes, except in the case of 7. radix, in
which the greater number of males have the formula 19-21—19-17,
whilst the greater number of females have that of 21-19-17.

In 7. natrix both sexes have the formula 19-17, and I can
detect no sexual variation in the amount of abridgment of
series IV. in the 25 British specimens which I have studied.

All the British specimens of C. qaustriaca in the British
Museum have the formula 19-17-15. It may be that there is a
sexual variation in the position of the posterior reduction, as in
8 out of 9 females this occurred from *85 to °96 in the length
of the body, whilst in 3 out of 4 males it occurred *75 to °85 in
the length of the body; one cannot base conclusions, however,
on such small material as this, or the preceding.

In I’. berus there is undoubtedly a tendency for the females to
possess a higher formula than the males. The number of rows
on the middle third of the body varies from 19 to 23. In
170 British specimens the 4 having 23 rows were all females.
Mr. Boulenger, who has statistics relating to 460 European speci-
mens, finds that 3:74 per cent. males and 2°81 per cent. females
have less than 21 rows, and 2°79 per cent. males and 6°32 per
cent. females have more than 21 rows. This tendency is still
more marked in the Green Viper, Atheris squamiger + Hallow., in
which the male has from 19 to 21, and the female 21 to 23 rows.

At present there is only one species known, in which this
differentiation between the sexes is complete and constant. Mr.
Boulenger = has recently discovered that in the Cameroon Bothro-

* Op. cit.

+ Boulenger, “ Batraciens et Reptiles recueillis par le Dr. Christy au Congo
Belge 1912-1914.” Revue Zool. Afric. vol. vii. fase. 1, 1919.

£ “Un cas de dimorphism sexuel chez un Serpent Africain (Bothrolycus ater
Giinth.).” Inst, de France, C. R. Ac. Sci. 1919,

364 ON THE DORSAL SCALE-ROWS IN BRITISH SNAKES.

lycus ater Giinth. it is normal for the male to have 17, and the
female 19 scale-rows on the anterior two-thirds of the body,
caudad to which the number of rows is reduced to 15 in both
sexes. The sexual dimorphism in this case, which so amply fulfils
Dr. Ruthven’s supposition, led to the sexes being described as
two species, Psewdoboodon albopunctatus and P. brevicaudatus
Andersson. In the female of this snake the reduction from 19
to 17 scale-rows is effected by the convergence of series VIII.
and IX., two-thirds down the body, and that from 17 to 15 by
the convergence of series VILI. and VIT. about nine-tenths down.
Tn the male the reduction to 17 rows is accomplished on the neck,
and that from 17 to 15 takes place three-quarters of the way
down the body, also by the convergence of the VITth and VIIIth
series; even this caudad reduction is therefore further advanced
in the male than in the female, thus completing the sexual
dimorphism of this unique species.

Pein

PZ. I9l9, WANRSISVAILIL, JP. It

BORNEAN BALANINUS.,

Bs 4S 1G INMVAIRESISVAUL JE, 1, UI.

BORNEAN BALANINUS.

ON SPECIES OF BALANINUS OCCURRING IN BORNEO. 365

24. On the Species of Balaninus occurring in Borneo (Coleo-
ptera, Curculionide). By Guy A. K. MarsHatt,
Db. Ses keZs:
[Received August 12, 1919: Read November 4, 1919. |
(Plates I. & I1.*)

>. G. E. Bryant having succeeded during a visit to Sarawak
in ae a number of weevils of the genus eaten us asked me
to work them out for him. On examining the literature up to
1914, and sueh publications since that Anes as are available in
this country, it came as a surprise to find that not a single species
of the genus had been described from Borneo. It is true that
Faust (Ann. Mus. Civ. Genova, xxxiv. 1894, p. 234) has recorded
the oceurrence in both Borneo and Burma of Lalaninus inter-
ruptus Kirsch (1875), but there seems to be some reason for
doubting the accuracy of his identification, at least so far as his
single Bornean 2 was concerned.

Kirsch’s insect was described from Malacca, and unfortunately
his type is not available (being in Dresden), nor have I been able
to find any species from the Malay Peninsula that can be attri-
buted to it. On the other hand, his description applies adequately
to one Indian and two Bornean forms, which represent three
undoubtedly distinet species. It is more probable that B. inter-
ruptus will be the same as the continental species, though it
would not be surprising if the Malacca insect should prove to be
yet a fourth species of the group. In the circumstances it has
seemed wiser to treat the Bornean species as distinct, and they
are described below as B. analis and B. imitator, spp. n.

In all, Mr. Bryant took 32 different species of Balaninus, of
which 3 were represented by single specimens in too poor con-
dition for description. Among the Curculionide sent from
Kuching by the late Mr. R. Shelford to the Oxford and Cambridge
Museums, which I have been able to examine through the kind-
ness of Prof. E. B. Poulton and Dr. Hugh Scott respectively,
there were only 7 species, 38 of which were not met with by
Mr. Bryant. Although Dr. A. R. Wallace paid special attention
to the collection of Coleoptera during more than a year’s stay in
Sarawak, on examining his material at the British Museum, I
could find only a single specimen of the genus (B. unifasciatus,
Siam.)

The species of Balaninus ave certain to prove far more
numerous in the tropics than would appear at present; for they
would be met with only occasionally, unless specially searched for,
owing to the fact that they breed in arboreal fruits and nuts,
and probably therefore they will be found mainly at the tops of
the forest trees. Mr. Bryant informs me that he obtained a large
proportion of his specimens from the flowers of a single felled
forest tree (Vernonia arborea) on Mt. Matang.

* For explanation of the Plates see p. 397.

366 DR. G. A. K. MARSHALL ON THE SPECIES

In spite of certain obvious diversities I have preferred to retain
all the species here described within the genus Balaninus in its
broad sense. The ventral structure in &. bryanti and the sepa-
ration of the front coxee in 4. discreticowis are notable departures
from the normal; but in a group of which we admittedly know
extremely little, undue haste in erecting new genera on single
species is by no means desirable.

The strong sinuation of the lateral margin of the elytra, which
has been used in the key for the primary division of the genus,
will no doubt be treated by some authors as of generic value.
But in the writer’s opinion it does not present the features of a
satisfactory generic character ; it is not correlated with any other
structural difference, and the. two divisions created by it are no
more homogeneous among themselves than is the whole undivided
series. It seems not unlikely that this sinuation is in the nature
of a late adaptation ; for it will probably be found to be correlated
with a greater activity in flight, the cutting away of the elytral
margin allowing greater freedom in the use of .the membranous
wings.

In the following descriptions no reference is made to the
sculpturing of the elytra, except in B. glabricollis, because it 1s
practically similar in all the other species. In these the punctate
striae are deep and broad throughout, but narrower than the
intervals, which bear transverse granular rugosities, these being
coarsest near the hase and becoming gradually finer behind.

I have pleasure in expressing my thanks to Miss O. F. Tassart
for the great care and trouble she has taken in the preparation of
her excellent drawings.

Mr. Bryant bas now generously presented the whole of his fine
collection of Curculionide to the British Museum, in which will
be found the types of all the following species, unless otherwise
indicated.

A Key to the Bornean Species of Balaninus.

1. (24.) Margo elytrorum lateralis profunde sinuatus supra metepi-
sternum lJatissimum.

2. (3.) Segmentum ventris secundum angulo laterali posteriore valde
=) . 5 . . . .
producte fere usquead basin segmenti quarti, segmentis primo
et secundo pone coxas brevissimis, simul sumptis ibi tertio non
longioribus; elytra nigra, tertia parte apicali testacea; ¢ tibiis
posticis intus et femoribus anticis subtus longe fimbriatis,
tibiis anticis intus dente lato armatis. 1. bryanti, sp.n., o 2.

3. (2.) Segmentum ventris secundum angulo laterali posteriore non aut
perparum producto; elytrorum integumentum concolor;
pedibus simplicibus.

4, (5.) Pronotum leve nitidum, punctis parvis sejunctis; articulus
funiculi primus quatuor sequentibus simul sumptis fere
wequalis. 2. glabricollis, sp. n., .

(4.) Pronotum opacum, punctis magnis reticulatim approximatis,
nonnumauam confluentibus ; articulus funiculi primus tribus
sequentibus simul sumptis brevior.

6. (21.) Segmentum ventris primum pone coxas secundo non longius.

7. (20.) Articulus funiculi primus secundo longior; pronotum regu-

lariter reticulato-punctatum.

OF BALANINUS OCCURRING IN BORNEO.

8. (11.) Elytra nigra, macula transversa alba singulatim pone medium
ornata; stria 9 ad basin elytrvorum percurrens; femora postica
ultra elytrorum apicem evidenter extensa; prothorax lobis
postocularibus angustis preditus.

9. (10.) Prothoracis longitudo latitudini baseos eequalis; macula alba in
elytro ab interstitio tertio ad octavum extensa; mesepimerum
undique dense albo-squamosum, metepisternum area triangu-
lari nuda in angulo laterali anteriore preditum; ¢ setis in
pygidio auvantiis, lacuna basali ventris minus profunda; 2
rastro quam femore antico plus duplo Jongiore.

3. bispilotus, sp. n

10. (9.) Prothorax latitudine baseos breyior; macula alba in elytro ab
interstitio quarto ad sextum aut septimum extensa; mesepi-

merum squamulis albis nullis aut perpaucis, metepisternum

ad basin omnino albo-squamosum; ¢ setis in pygidio nigro-

brunneis, lacuna basalr ventris profunda; 2 rostro quam

femore antico sesqui-longiore. 1. excavatus, sp. n., 6 2.

11. (8.) Hlytra fasciis albis nullis; stria 9 elytrorum basin non attin-
geus; femora postica ultra elytrorum apicem non aut perpa-
rum extensa; prothorax lobis postocularibus nullis.

12. (17.) Elytra interstitio secundo conspicue pallido-squamoso, excayva-
tione suturali ineequali in elytro dextro profundiore quam in
levo.

13. (16.) Pronotum vitta media pallida ornatum.

14. (15.) Vitta media prothoracis e squamulis transversis composita ;
elytra vitta pallida in interstitio sexto perfecta; ¢ tibiis pos-
ticis interne supra corbulam profunde excisis.

5. eveisipes, Sp. 1.

15. (14.) Vitta media prothoracis e squamulis longitudinalibus compo-
sita; elytra vitta pallida interstiti1 sexti in tertia parte basali
obsoleta; ¢ incognitus. 6. sesquilineatus,

16. (13.) Pronotum vitta media nulla; ¢@ tibiis posticis interne non

od FS.

, 3%.

SPeailes Gre

OCICS i120, eer (@
eXCISIS. 7. bilineatus, sp. n., 62.

17. (12) Elytra interstitio secundo non conspicue pallido-squamoso, ex-
cayatione suturali equali.

18. (19.) Integumentum nigrum, supra squamulis fuscis inconspieuis
maximam partem indutum; pronotum utrimque in dimidio
basali vitta lata laterali miniato-squamosa; elytra interstitio
tertio in dimidio basali sparsim, sutura in dimidio apicali late
et dense miniato-squamosa, 8. semisuturellus,

19. (18.) Integumentum rufo-brunneum, supra sat dense ferrugineo-squa-
mosum, linea media pronoti et elytris versus apicem pallido-

squamosis. 9. rufulus, sp. 0.,
20. (7.) Articuli duo basales funiculi equales; pronotum crebre gyroso-
carinulatum. 10. gyrosicollis, sp.

21. (6.) Segmentum ventris primum pone coxas secundo longius ; sutura
elytrorum in dimidio posteriore setis spiniformibus oblique
elevatis armata.

22. (23.) Corpus et pedes nigri; fimbrie tibiarum apicales rufo-flavide ;

pronotum sine vitta media pallida; scutellum subquadratum

sp.n., 3.

Qe
me, SP.

elevatum. 11. nigrocinereus, sp. n.. 2.

23. (22.) Caput et prothorax et pedes ferruginei, elytra nigra; fimbrie
tibiarum apicales fuscxe ; pronotum vitta media pallida orna-
tum; scutellum depressum, longius quam latins.

12. nigrorufus, sp. u., &.

24, (1.) Margo elytrorum lateralis non sinuatus; metepisternum non
dilatatum : segmentum ventris primum pone coxas secundo
longius.

25. (62.) Coxe antic contigue:.

26. (59.) Antenne articulo clave secundo multo latiore quam longiove.

27. (56.) Sutura elytrorum in dimidio basah evidenter depressa.

28. (53.) Serobes antennarum prope basin infra rostrum producti; margo

inferior rostri (a latere inspectus) caput attingens longe supra

marginem inferiorem oculi.

368 DR. G. A. K. MARSHALL ON THE SPECIES

29. (32.) Elytra nee fasciis nee maculis albis ornata; scutellum dense
albo-squamosum. :
30. (31.) Tibiz postice margine dorsali sinuate et squamis latis indute ;
venter sezmentis duobus basalibus squamis latis albis dense
indutis. 13. subpartitus, sp. u., 2.
31. (380.) Tibie postices recte et squamis angustis setiformibus indutee ;
venter totus squamis angustis sparse 1dutus, macula parva
laterali e squamis latioribus in sezmento secundo excepta.
14. mestus, sp. n., 2.
32. (29.) Elytra nigra, fasciis aut maculis albis ornata.
33. (34.) Rostrum in parte basali late dilatatum; antennarum scapus
brevissimus, artrculis duobus basalibus funiculi brevior.
15. tumidirostris, sp. n., 6.
34, (33.) Rostrum versus basin non dilatatum; antennarum scapus
longus, articulis 4 aut 5 basalibus funiculi zequalis.
35. (38.) Pronotum maculis basalibus albis nullis ; scutellum dense albo-
squamosum.
36. (37.) Elytra macula postimediana albo-squamosa in interstitio secundo,
et ibidem squamis et setis suturalibus:albis in interstitio
primo; ¢ segmento ventrali quinto fasciculis duobus apicali-
bus e setis longioribus compositis. 16. trinotatus, sp... od 9.
37. (36.) Hlytra maculis postmedianis albis in interstitiis 2 et 3, nec
squamis nec setis albis in interstitio primo; ¢ fasciculis ven-
tralibus e setis brevibus cum uno solo longiore compositis.
17. consocius, sp. n., ¢.
38. (35.) Pronotum in utroque latere macula basali alba.
39. (40.) Dorsum squamis angustis albis et flavo-brunneis mixtis indu-
tum ; pronotum maculis basalibus albis parvis et inconspicuis ;
scutellum dense albo-squamosum ; elytra pone medium albo-
fasciata. 18. wnifasciatus, sp. n., 3d &.
40. (39.) Dorsum (maculis albis exceptis) squamis tantum fuscis indu-
tum; pronotum maculis basalibus albis conspicuis.
41. (48.) Elytra linea brevi apicali alba in interstitio primo aut secundo ;
scutellum nigrum.
42, (43.) Elytra linea apicali alba in interstitio secundo ; femora postica
ultra elytrorum apicem longe extensa. 19. decemnotatus, sp.n., 9.
43. (42.) Elytra linea apicali alba in interstitio primo; femora postica
elytrorum apicem tantum breviter excedentia.
44, (45.) Elytra ad basin tantum in interstitio primo albo-vittata.
20. quincunx, sp.n., oY.
45. (44.) Elytra ad basin in interstitiis 1-4 fascia curvata alba utrimque
cum maculis pronoti conjuncta, et fascia secunda alba pone
medium.
46. (47.) Funiculus articulo primo quam secundo paulum bzeviore; elytra
ad apicem linea marginali albo-squamosa; ¢ segmento ventris
secundo ad Jatera pestice evidenter angulato, sezmento quinto
ad apicem profunde exciso et utrinque triangulariter producto ;
® segmento quinto in medio profunde foveato.
21. analis, sp.n., 6°.
47. (46.) Funiculus articulo primo quam secundo paulum longiore; elytra
in margine apicali non albo-squamosa; g segmento ventris
secundo postice truncato, sezmento quinto ad apicem vix
sinuato nec producto; 2 seemento quinto non impresso.
22. imitator, sp.n., ¢ 2.
48. (41.) Klytva linea alba juxta-suturali nulla.
49. (50.) Scutelluam nigrum, antice profunde excisum; statura major
(5-5'5 mm.). 23. deceptor, sp. u., 3d.
50. (49.) Scutellum integrum, squamulis albis plus minus dense indutum;
statura minor (2°5-3°S mm.).
51. (52.) Prothorax macula basali alba utrimque mesepimerum non attin-
gente; elytra ad basin squamis albis tantum in interstitio
secundo et maculis postmedianis albis in interstitiis 2, 3, 7, 8.
24. commodus, sp. n., 6.

wcAZr

OF BALANINUS OCCURRING IN BORNEO. 369

52. (41.) Prothorax macula basali utrimque mesepimero contigua; elytra
ad basin squamisalbis in interstitus 1-4, ct fascia postmediana
in interstitus 1-8. 25. grypus. sp.n., 3g.

53. (28.) Scrobes antennarum omnino laterales; margo inferior rostri
caput attingeus ad marginem inferiorem oculi; rostrum ad
basin fronte non Jatius.

54, (55.) Sutura elytrorum eqyualiter nmpressa; prothorax carina media
tenui instructus ; funiculus articulo primo quam secundo

paulum longiore. 26. shelfordi, sp.u., 2.
55. (64.) Sutura in elytro recto magis impressa ; prothoray non carinatus ;
funiculus articulo 1=24+3+4, 27. pusio, sp. n., 7.

56. (27.) Sutura elytrorum ad basin non impressa.

57. (58.) Dorsum nigrum, maculis albis definitis notatum. :
28. delicatulus, sp. u., d 2.

58. (57.) Pronctum nigrum; elytra picea, squamulis fulvis irrorata.

29. eugeni@,sp.u.. d&.

59. (26.) Antennze articulo claves secundo fere duplo longiore quam
latiore.

60. (61.) Elytrorum sutura ad basin non impressa; funiculus articulo
1=2+3, articulo clave secundo ad basin valde constricto ;
elytra vitta suturali lata alba in tertio basali et linea suturali
angusta in dimidio apicali ornata. 30. longiclavis, sp. u., 9.

61. (60.) Elytrorum sutura ad basin profunde et inwqualiter impressa ;
funiculus articulo 1=2+3+4+45, articulo clave secundo ad
basin vix constricto; elytra area magna communi cervino-
squamosa. 31. sellautus, sp.u., &.

62. (25.) Coxw antic evidenter separate. 32. disereticowis, sp. n., 2.

1. BALANINUS BRYANT, sp.n. (PI. I. fig. 8.)

3 2. Integument biack, except part of the elytra, rather
more than the apical third being testaceous; the lower surface
of the head, the sides of the prosternum and the black portion of
the elytra thinly clothed with small hair-like scales, the basal
margin of the pronotum with a single row of rather broader
scales ; the testaceous portion of the elytra densely clothed with
broad yellow scales (except interval 1), which mostly become
narrower towards the apex; the front portion of the prosternum,
the meso- and metasternum with similar and equally dense scales,
those on the venter being paler and narrower.

Head with the rostrum arising from about the middle of the
eye (in lateral view), the forehead nearly parallel-sided and about
two-thirds the width of the base of the rostrum. LRostruwm— d,a
little shorter than the body (7:9), rather coarsely punctate near
the base, with a broad lateral furrow ending shortly before the
end of the scrobe, which latter passes beneath the rostrum at the
base and does not extend beyond the antenne; 9, as long as the
body, the basal punctures. less coarse, the lateral furrow shorter
and shallower. Antenne inserted at (¢) or behind (2) the
middle of the rostrum; joint 1 of the funicle equal to 2+3,
2 longer than 3, 3 equal to 4, 5 to 7 shorter and subequal, but 7
a little broader ; the club fusiform, joint | slightly longer than
2or 3. Prothorax a little broader than long. broadest near the
base, slightly constricted at the apex. the sides gently rounded ;
the basal margin very slightly produced in the middle, the apical

370 DR. G. A. K. MARSHALL ON THE SPECIES

margin almost vertical at the sides, without any trace of post-
ocular lobes ; the upper surface with reticulate and partly con-
fluent punctures, with a mere trace of a median smooth line, and
with short recumbent brown setee. Sewtellwm almost circular,
with a few minute pale sete. Hlyira more elongate than usual,
the shoulders rather accentuated by a shallow sinuation just
behind them, the apices separately rounded, the lateral margins
very deeply sinuate above the metepisternum, the suture deeply
and quite symmetrically impressed in the basal half, and striz
9 and 10 not quite reaching the base. Legs thinly clothed with
recumbent white sete, all the femora with a small sharp tooth ;
‘$ with a short truncate process on the front coxe, the front
femora with a fringe of long hairs beneath, the front tibie with a
broad internal tooth behind the middle, the hind femora with
a long sharp tooth midway between the usual tooth and the base,
the hind tibie strongly flattened internally and there clothed
with a fringe of long hairs; 2 with the Jegs simple, the inner
apical face of the front tibie rather deeply excavated. Venter
with segments 1 and 2 extremely short behind the coxe, 1 being
shorter than 2, and the two together about as long as 3 along
the same line; the outer angle of 2 strongly produced, almost
reaching the base of 4 and thus separating 3 from the elytra; the
pygidium of 3 broadly exposed and very convex, with short
recumbent sete.

Length, 3°8-4mm.; breadth, 19-2 mm.

Sarawak: 1 ¢, Mt. Matang, 1000ft., 13.11.14, and 1 9,
Mt. Merinjak, 2200 ft., 27. v. 14 (G. #. Bryant).

A very distinct and aberrant species, not only in its colouring,
but also in the structure of the basal ventral segments and the
secondary sexual characters on the legs of the ¢, the additional
lower tooth on the hind femora being a very unusual feature
among the Curculionide.

2. Black or dark piceous, shiny; the whole of the lower sur-
face and the elytra (except the shoulders) densely covered with
elongate and apically truncate fawn-coloured scales, and a row of
similar scales along the basal margin of the pronotum.

Head with the rostrum arising from about the middle of the
eye (in lateral view), the forehead with a central fovea, nearly
parallel-sided and almost as broad as the base of the rostrum.
Rostrum extremely long (14 mm., without allowing for the curve),
porrect for two-thirds of its length, densely punctate close to the
base, except for a rather broad smooth central line, without any
lateral furrow or carina, the scrobe entirely lateral and not
extending beyond the antenne. Antenne inserted far behind
the middle of the rostrum; joint 1 of the funicle very long, as
long as the succeeding 34 joints, 3 nearly as long as 2, 4 a little

OF BALANINUS OCCURRING IN BORNEO. Bil

shorter and equal to but thinner than 7, 5 and 6 still shorter and
equal; joints 3-7 bearing recumbent setiform white scales, their
density increasing outwardly; the club fusiform, clothed with
fulvous pubescence, joints 1 and 2 equal and a little longer than
broad, 3 distinctly shorter. Prothorax about as long as broad,
broadest at the base, the sides scarcely rounded except just before
the broad shallow apical constriction; the base marginate and
produced in the middle, the apical margin rounded dorsally and
without postocular lobes; the upper surface smooth and shiny,
with numerous small separated punctures and no smooth central
line, the sides of the prosternum with larger and closer punctures.
Scutellum relatively small, longer than broad and with a few
minute pale scales. Hlytra elongate cordiform, the shoulders not
very prominent, the sides nearly straight, the apices almost
jointly rounded, the lateral margins deeply sinuate near the base ;
the upper surface almost flat, with comparatively narrow and
shallow strize, which become much shallower behind, strie 9
and 10 practically reaching the base, the suture deeply and
symmetrically impressed in the basal half; the intervals broad
and plane, with fairly close and slightly wrinkled punctures.
Legs very long and slender, with sparse setiform pale scales; the
hind femora projecting shortly beyond the apex of the elytra and
the tooth on them but little larger than that on the mid femora.
Sternum with a deep impression along the upper edge of the
metepisternum, the projecting margin bearing a fringe of over-
hanging scales. Venter with segment 1 ssa y longer than 2
behind the coxee, the external angles of 2 scarcely produced.

Length, 8° 4 mom. ; breadth, 4 mm.

SaRAwAK: 1 2, Mt. Mevinjak, 2200 ft., 27.v.14(G. #. Bryant).

Differs from all the knowa Bornean species owing to its
smooth pronotum and the impression on the metepisternum,

3. BALANINUS BISPILOTUS, sp. n. (PI. I. fig. 2.)

3 Q. Rather dull black, with patches of pure white scales ;
the elytra with a transverse white band behind the middle
extending from interval 3 to 8 or 9, and an elliptical marginal
patch at the apex which does not quite reach the suture; in the
2 only there are also a few white scales near the ‘base of
interval 2; the front of the prosternum, the mesepimeron, the
mesosternal process, the whole side of the metasternum (except
the apex of the metepisternum and a small triangular patch at
its base) and the entire venter, densely covered with broad white
scales; the coxe, the rest of the mesosternum (except the
episterna, which are bare) and the centre of the metasternum
with narrower, greyish white scales. MHewd with the rostrum
arising from samt (2) or above (3) the middle of the eyes, the
forehead about two-thirds the width of the rostrum at its base
and with a shallow fovea. ostrwm—d, only a little more than

Proc. Zoou. Soc.—1919, No. XX VI. 26

372 DR. G. A. K. MARSHALL ON THE SPECIES

half the length of the body (6:11), rather strongly and closely
punctate in the basal half and finely and sparsely so beyond, with
a smooth central line in the basal half, a shallow punctate lateral
furrow (nearly as long as the scrobe) bounded below by a low
eavina, below which is an impunctate and longitudinally striolate
area, the scrobe passing partly beneath the rostrum at the base
and extending slightly beyond the antenne; 9, much longer
than the body (3: 2), less strongly punctate at the base than im
the ¢ and extremely minutely punctate beyond the antenne, the
lateral furrow only one-third the length of the scrobe, and the
striolate area beneath it lacking. Antenne inserted at (¢ )
or far behind (2) the middle of the rostrum, the seape not
reaching the eye; joint 1 of the funicle longer than 2, joints
3 and 4 shorter and subequal, 5 and 6 again a little shorter and
equal, 7 equal to 3; the club with brown pubescence, joint 1 dis-
tinctly longer than 2, and 2 than 3. Prothorasz as long as broad,
broadest at the base, the sides gently rounded, the apical constric-
tion very slight; the base not marginate and but little produced
in the middle, the apical margin feebly arcuate, the postocular
lobes feeble; the upper surface with regular reticulate punctures
and a very short central smooth line, the punctures on the sides
of the prosternum with rather broader intervals. Sewtellwm
small, bare, usually with a transverse shallow impression. Hlytra
longer than broad (13:10), the lateral margins deeply sinuate
near the base, the apices separately rounded; the basal sutural
impression deep and not quite symmetrical so that interval | on
the right elytron is narrower than that on the left, strie 9 and
10 practically reaching the base; the intervals transversely
rugose and on the black areas with short dark recumbent sete,
which are longer and slightly raised on the apical third of the
suture. Legs very long and slender, especially the front femora
in the 2, the hind femora exceeding the apex of the elytra and
bearing a tooth that is no larger than that on the mid femora.
Venter with segment | distinctly shorter than 2 behind the coxe,
the apical angles of the latter moderately produced; segment 5
of ¢ broadly sinuate at the apex and with «a broad central
depression, flanked on each side at the apex by a dense erect tuft
of reddish-yellow hairs, the pygidium being densely clothed with
similar erect hairs; segment 1 of d very broadly but shallowly
impressed.

Length, 8:4-8°8 mm. ; breadth, 4-4-4 mm.

Sarawak: 1 9, Mt. Matang, 2000ft., 24. xii. 13(G. #. Bryant);
1S, Quop; ie VOI (GB) AS oe Ono Mite leremnyeilce
2200 ft., 23. v.14 (G. #. B.—types) ; 1 9, Banting, 12.iv.1915
(G. D. Allen). S.H. Borneo: 1 3, Martapura, 1891 (W. Doherty).

The Martapura ¢ differs from the others in having a white
spot at the base of interval 2, and the transverse band and the
apical patch on the elytra linked together along intervals 8
and 9.

aoe

OF BALANINUS OCCURRING IN BORNEO. BD

4, BALANINUS EXCAVATUS, Sp. n.

Colouring as in B. bispilotus, sp. n., except that the transverse
white patch on the elytra extends only from interval 4 to 6 or 7,
and the white scales at the base of interval 2 in the 9 are
lacking; beneath, the scales in the centre of the prosternal white
patch are much narrower than the outer ones, exposing the
integument, the mesepimeron bears at most only a few white
scales, the mesosternal process is clothed with narrow scales, the
metepisternum has no bare basal patch, and the scales on the
venter are much narrower.

The following are the principal structural differences from
B. bispilotus :— Head with the rostrum arising from about (do) or
well below (?) the middle of the eye. ostrwm— ¢, much less
strongly or densely punctate, without a smooth central line, the
lateral furrow almost impunctate; 9, much shorter, being only
of the same length as the body. Antenne with the scape reach-
ing the eye in the 9 but notin the ¢.. Prothorax shorter than
its basal width, its sides more strongly rounded. lytra jointly
and rather shallowly sinuate at the apex. Venter with segment I
of g deeply excavated and containing a short median furrow at
the base; segment 5 of ¢ hardly sinuate at the apex, not
depressed in the middle, and with a small curved tuft of brown
hairs on each side of the hind margin, the hairs on the pygidium
being blackish brown.

Length, 52-64 mm.; breadth, 3-3:2 mm.

Sarawak: 1d, 19, Mt. Merinjak. 600-2200ft., v. 1914
(types), and 1 ¢, Mt. Matang, 3200 ft., 18. x11.13 (@. #. Bryant).

The ¢ from Mt. Matang has a few white scales near the base
of interval 2 on the elytra.

5. BALANINUS EXCISIPES, sp. n.. (Pl. I.-fig. 3.)

3 2. Dull black, with yellowish-grey stripes; the head with
a few white setiform scales above and bélow ; the prothorax with
a central yellowish stripe and scattered isolated white scales at
the sides; the elytra with two complete yellowish stripes on
intervals 2 and 6 (the scales in the former being larger),
intervals 8-10 covered with pale scales, those on the basal third
being narrow and whitish, the remainder broad and yellowish,
intervals 3, 4 and 7 with some pale scales at the apex, and a few
at the base of 4, the rest of the surface rather thinly clothed with
narrow brown scales; the prosternum with only a few larger
whitish scales on the intercoxal process, the sides of the meso-
and metasternum densely clothed with broad yellow scales, the
centre of the sternum and the whole venter fairly closely covered
with narrower whitish scales,

Head with the rostrum arising from about (9) or well above
(3) the middle of the eye (in side view). Rostrwm— 3, much
shorter than the body (3:5), with a smooth median line in the
basal half and three rows of punctures on each side of it, without

26*

374 DR. G. A. K. MARSHALL ON THE SPECIES

any definite lateral furrow or carina, the scrobe not quite passing
beneath the base of the rostrum; 9, rather shorter than the
body (4:5), the punctures finer, the scrobe quite lateral.
Antenne inserted at (¢) or behind (9) the middle of the
rostrum, the funicle with joint 1 distinctly longer than 2,
3 longer than 4, and 4-7 almost equal; the club with joint 1
very slightly longer than 2, and 2 than 3. Prothorax distinetly
broader than long, the sides subparallel for a short distance from
the base, then roundly narrowing to the apex, the apical con-
striction obsolete; the apical margin feebly arcuate dorsally, the
postocular lobes absent; the upper surface reticulately punctate
throughout. Scwtellum small, rounded, with a few minute sete.
Elytra a little longer than broad (6:5), the lateral margins
deeply sinuate, the apices jointly sinuate; the basal sutural
depression deep and asymmetrical, the edge of the right elytron
being more depressed than that of the left, the apical portion of
the suture without erect spine-like setee; strie 9 and 10 not
quite reaching the base. Legs comparatively short, the hind
femora not exceeding the apex of the elytra; the femoral teeth
sharp and spine-like, the hind tibize of the ¢ with a deep notch
on the inner edge just at the top of the corbel. Venter with the
two basal segments of equal length behind the coxe ; segment 5
of g sinuate on its posterior edge and with a shallow apical
impression ; the pygidium with a dense pointed tuft of brown
hairs. ;
Length, 4mm.; breadth, 2mm.

Sarawak: 1¢, on flowers of Vernonia arborea, Mt. Matang,
1000 ft., 10. 11.14 (type), and 1 ¢ 1 9, Quop, iv. 1914
(G. E. Bryant).

6. BALANINUS SESQUILINEATUS, sp.n. (PI. I. fig. 6.)

2. Colour dull black, with yellowish-white stripes; the head
with a few sparse scales beneath; the pronotum with a pale
central stripe, which is broadest at the base and gradually nar-
rows to the apex (the scales themselves being broad near the base
and diminishing towards the apex), and a small basal pale spot on
each side; the elytra with a complete stripe of broad scales on
interval 2, that on 6 being obsolete in the basal third, the apical
half of intervals 7 and 8 and almost the whole of 9 and 10
thinly clothed with narrow whitish scales, and intervals 3 and 4
with a few pale scales at the apex ; the prosternum with a dense
patch of broad white scales in front of the coxee, and isolated
narrow seales on the sides; the meso- and metasterna with
separated broad scales, but these are densely packed on the mes-
epimeron and on the sides of the metasternum, except on a patch
at the base of the metepistérnum where the scales are minute;
the veater with fairly dense whitish scales of intermediate
width :

Head with the rostrum arising from well below the middle of

eee SOIT

OF BALANINUS OCCURRING IN BORNEO. BD

the eye, the lower surface being about on a level with the lower
edge of the eye. Rostrum shorter than the body (5:7), finely
and sparsely punctate at the base, without any lateral furrow or
carina, the scrobe entirely lateral. _Anéernnce inserted well behind
the middle of the rostrum, the scape reaching the eye; the
funicle with joint 1 slightly longer than 2, and 3 than 4, 4-7
almost equal in length; the club elliptical, joint 1 slightly longer
than 2, and 2 than 3. Prothorax very slightly broader than
long, the sides subparallel near the base, then roundly narrowed
to the apex, the apical constriction faint: the apical margin
feebly arcuate dorsally, and with no postocular lobes; the upper
surface with regularly reticulate punctures. Hlytra as in
B. excisipes, but more rapidly narrowed behind, so that the
shoulders appear more prominent; the scales on interval 2 are
distinctly broader, being 3-4 deep in the broadest part and 2 deep
near the apex (5 and 2 respectively in B. ewcisipes). Legs asx in
B. excisipes 9, except that the internal basal sinuation of the
tibize is distinctly deeper, and the scales on the upper surface of
the hind tibiz are rather broader than any others on the legs.

Length, 3mm.; breadth, 1°6 mm.

Sarawak: 1 9, on flowers of Vernonia arborea, Mt. Matang,
1000 ft., 12.11.14 (G. H. Bryant, type); 1 9, on flowering tree.
Kuching, vii. 1900 (2. Shelford).

In B. excisipes the central stripe on the pronotum is much
broader and composed entirely of transverse scales, whereas in
the present species all the scales le longitudinally, except those
at the extreme base.

7. BALANINUS BILINEATUS, sp.n. (PI. I. fig. 4.)

3 %. Colour dull black, the pronotum without pale markings;
the elytra with a stripe of broad, pale yellow scales on interval 2
from the base to beyond the middle (the scales 4-5 deep at its
broadest), an apical patch of similar scales between stria 2 and
the margin and extending forwards on intervals 5—7 as far as the
termination of the stripe on 2, intervals 8-10 thinly clothed with
minute setiform white scales; the prosternum with a dense ante-
coxal patch of broad yellowish scales, the sides with narrower
isolated whitish scales; the meso- and metasternum with broad
yellowish scales which are fairly dense throughout except in the
middle of the mesosternum; the abdomen densely clothed with
narrower whitish scales.

Head with the rostrum arising at about ( ¢ ) or well below ( ¢ )
the middle of the eye. Rostrwm—d, two-thirds the length of
the body, rather irregularly punctate in the basal half, with
a very narrow smooth central line and a punctate lateral furrow
extending for about four-fifths of the scrobe, which passes
beneath the base of the rostrum and not beyond the antenna ; 9°,
shorter than the body (7 : 9), the punctures finer, and the lateral
furrow shorter, Antenne inserted about (3) or well behind (¢ )

376 DR. G. A. K. MARSHALL ON THE SPECIES

the middle of the rostrum; the funicle with the outer joints very
short, 1 much larger than 2, joints 3 and 4 equal, 5-7 a little
shorter and subequal, 7 as broad as long; the club elliptical and
comparatively large, being about as long as the five preceding
joints, joint 1 longer than 2, and 2 than 3. Prothorax and
seutellum as in B. eacisipes. LHlytra also as in B. excisipes,
but slightly narrower (length : breadth as 5:4). Legs com-
paratively short, the hind femora not exceeding the apex
of the elytra, the femoral teeth sharp and spine-like; the
basal sinuation of the tibie as deep as in B. sesquilineatus,
the hind pair clothed above with small setiferm scales lke
the rest of the legs and not notched in the ¢. Venter with
segment 1 shorter than 2 behind the coxe; segment 5 of 3
with a small bunch of hairs on each side, the pygidium rather
broadly exposed and covered with yellowish-white hairs.

Length, 3-4mm.; breadth, 1°-6-2mm.

Sarawak: 1 3d 1 Q, Quop, iv. 1914 (G. H. Bryant).

8. BALANINUS SEMISUTURELLUS, Sp. n.

3. Colour dull black, the pronotum on each side with a broad
stripe from the base to the middle composed of dark pink
separated transverse scales, and a few pale setiform scales on the
disk and at the sides; the elytra with a diffuse stripe of pale
narrow scales from the base to the middle, a broad dense sutural
stripe on the apical half formed of broader pale pink scales,
interval 7 with a diffuse stripe of similar whitish scales, and the
remaining intervals with a few white setiform scales scattered
among the dark ones; the prosternum with sparse narrow white
scales, a small dense ante-coxal patch of broader pink scales and
a few pink scales on the sides ; the meso- and metasternum with
sparse white scales and a broad lateral stripe of dense pink scales ;
the venter with fairly dense whitish scales.

Head with the rostrum arising from about the middle of the
eye. Rostrwm comparatively short and stout, much shorter than
the body (4:7), rather coarsely and confluently punctate on the
basal section, with a very narrow smooth central line and a broad
punctate lateral furrow, nearly reaching the antenna and bounded
below by a distinct carina, the scrobe passing beneath the base
of the rostrum and not beyond the antenna. Antenne inserted
just behind the middle of the rostrum, the scape not quite
reaching the eye; the funicle with joint 1 longer than 2, 3 and 4
subequal, 5—7 shorter, subequal and about as broad as long; the
club as long as the four preceding joints, joint 1 much longer
than 2, which is but little longer than 3. Prothorax nearly as
long as broad, broadest at the base, the sides gently rounded,
the apical constriction very feeble, the apical margin truncate
dorsally and without postocular lobes; the upper surface deeply
and reticulately punctate. Scutellwm almost circular, bare.
filytra comparatively elongate (5: 4), the margins deeply sinuate,

OF BALANINUS OCCURRING IN BORNEO. BY i

the apices almost jointly rounded; the sutural impression deep
and almost symmetrical, but slightly inclined to the right, the
apical area without raised sete; striw 9 and 10 almost reaching
the base. Legs comparatively short, the hind femora not exceed-
ing the apex of the elytra, the femoral teeth sharp and spine-like;
the basal sinuation of the tibiw rather shallow, the hind pair
clothed above with broader pink and white scales. Venter with
segment | shorter than 2 behind the cox, segment 5 simple.

Length, 2°8mm.; breadth, 1°6 mm.

Sarawak: 1G, on flowers of Vernonia arborea, 8.11. 1914
(G. H. Bryant).

9. BALANINUS RUFULUS, Sp. 0.

3. Colour brown, the upper side fairly densely clothed with
rather narrow, reddish brown scales; the pronotum with a
narrow central line of longitudinally placed yellowish scales; the
elytra with a large apical area clothed with broader yellowish
scales and a short stripe of similar scales at the base of interval 2;
the whole sternum clothed with separated broad yellowish scales
which are denser on the mesepimeron and the margin of the
mesepisternum, the scales on the venter being denser and
narrower.

Head with the rostrum arising from well below the middle of
the eye; the vertex thinly, the forehead densely covered with
sealing. Rostrum (broken at the apex) distinctly punctate
laterally at the base, but without any lateral furrow or carina,
the serobe not passing beneath the rostrum nor exceeding the
antenna. Antenne inserted far behind the middle, the scape
almost touching the eye; the funicle with joint 1 distinctly
longer than 2, and 8 than 4, 5 the shortest, 4, 6 and 7 subequal
and a little longer than broad ; the club about as long as the four
preceding joints, joint 1 a little longer than 2, and 2 than 3.
Prothorax a little broader than long (8:7), the sides subparallel
for a short distance from the base, then roundly narrowed, with-
out any apical constriction, the apical margins feebly arcuate
dorsally and without any postocular lobes; the upper surface
with regular reticulate punctures. Scutellum nearly circular,
with a few minute scales. Hlytra with the lateral margins
deeply sinuate and the apices jointly emarginate; the basal
sutural impression deep and asymmetrical, the edge of the right
elytron being depressed below that of the left, the suture without
raised sete at the apex, strie 9 and 10 not nearly reaching the
base. Legs comparatively short, the hind femora not exceeding the
elytra, the femoral teeth sharp and spine-like, the basal sinuation
on the tibiz well marked, the hind pair with fine setiform scales
above like the rest of the legs. Venter with the two basal
segments equal behind the coxe.

Length, 2°6 mm.; breadth, 1-4 mm.

Sarawak: 1 @, on flowers of Vernonia arborea, Mt. Matang,
1000 ft., 8. 11,1914 (G. #, Bryant).

378 DR. G. A. K. MARSHALL ON THE SPECIES

10.. BALANINUS GYROSICOLLIS, Sp. n. (PI. II. fig. 2.)

é¢ 2. Dull black, clothed above with narrow dark brown scales
mottled with whitish ones, and with a faint central paler stripe
on the pronotum; beneath, with separated narrow whitish scales
throughout, and without any of the usual dense patches of
broader scales.

Head with the rostrum arising from well above the middle of
the eye in both sexes, its upper surface being on a level with the
top of the eye (d) or a little below it (2). Rostrum— ¢ , two-
thirds the length of the body, rather coarsely and confluently
punctate in the basal section, without any smooth central line,
but with a very shallow broad lateral furrow reaching the
antenna and bounded below by a distinct carina, the scrobe
passing beneath the base of the rostrum and extending shortly
beyond the antenna; 9, seven-eighths the length of the body,
the lateral furrow scarcely shorter, and the punctures rather
finer. Antenne inserted behind the middle of the rostrum in
both sexes, the scape not reaching the eye; the funicle with the
two basal joints equal, 3 slightly longer than 4, 5-7 a little
shorter and equal in length; the club short, ovate, as long as the
two preceding joints, joint | slightly longer than 2, and 2 than 3.
Prothorax slightly broader than long (6: 5), broadest at the base,
the sides rather strongly rounded, with a distinct apical constric-
tion, the apical margin truncate dorsally and without postocular
lobes ; on the upper surface the reticulate punctures have coalesced
so that the interspaces form numerous fine sinuous ridges.
Scutellum longer than broad, with fairly dense pale setiform
scales. lytra elongate (4x3), the margins distinctly sinuate,
the apices jointly rounded; the basal sutural impression com-
paratively shallow and asymmetrical, being almost entirely con-
fined to interval 1 on the right elytron, the apical portion of the
suture without raised setee; striz 9 and 10 not reaching the base.
Legs moderately long, the hind femora slightly exceeding the end
of the elytra, the femoral teeth sharp and spine-like, the tibie
with the basal sinuation feeble and clothed only with setiform
scales. Venter with segment | shorter than 2 behind the coxe.

Length, 5:8-6:4 mm.; breadth, 2-4-3 mm.

Sarawak: 3 dod, 2 2 2, Quop, iv. 1914 (types), and 2 3 6,
Puak, v.1914(G@. #. Bryant).

The sculpture of the pronotum and the short antennal club are
very distinctive characters.

11. BALANINUS NIGROCINEREUS, sp. n. (PI. I. fig. 5.)

3 @. Colour dull black, the upper surface covered with rather
sparse short hair-like white scales (producing a dull grey effect)
and with the following black patches on which the scales are
dark: the forehead and vertex of the head, a large triangular
patch in the middle of the base of the pronotum, a transverse

OF BALANINUS OCCURRING IN BORNEO. 3n9

band on the elytra before the middle extending from stria 2 to
the margin and including the whole shoulder and the base from
stria 4 to the margin, and a compiete subapical transverse band
about 4mm. wide; on the lower surface the scales are rather
wider, except on the sides of the prosternum, the middle of the
metasternum, and on the whole of the last three ventral seg-
ments; on the mesepimeron and at the sides of the first two
ventral segments the scales are larger and closer together, and on
the front part of the prosternum they are almost round and
separated.

Head with the rostrum arising above the middle of the eye (in
side view), its upper edge being on a level with (2) or above(¢ )
the top of the eye. ostruwm— ¢, shorter than the body (10: 13),
rather more distinctly punctate than usual in the apical half
(especially at the sides), in the basal half with a smooth central
line, and two rows of punctures on each side of it, below these a
lateral furrow as long as the scrobe and containing a single
regular row of punctures, its lower edge being distinctly carinate,
the scrobe passing beneath the rostrum at the base and not
extending in front of the antenne; 2, also shorter than the
body (10:12) and rather more slender than that of the do,
the punctures finer (except close to the base) and the lateral
furrow slightly shorter. Antenne inserted slightly behind the
middle of the rostrum in both sexes, the joints of the funicle
with pale recumbent sete, the scape not nearly reaching the eye;
joints | and 2 of the funicle equal, 3 longer than 4, 4—5 subequal,
7 equal to 3; the club with yellowish-grey pubescence, joint 1
longer than 2, and 2 than 3. Prothorax broader than long
(5:4), the sides almost parallel for a short distance from the
base, then strongly rounded, the apical constriction well marked ;
the apical margin truncate dorsally and with feeble ocular lobes ;
the upper surface with regularly reticulate punctures and no
distinct central line. Scutellwm subquadrate, covered with dense
white scales, except a narrow strip on each side. Elytra slightly
longer than broad (8:7), the apices almost jointly rounded, the
lateral margins deeply sinuate; the basal sutural impression rather
shallow and asymmetrical, interval | of the right elytron being
lower than that of the left, the apical third of the suture with
erect black spine-like bristles, the strize 9 and 10 not reaching
the base. Legs elongate, the hind femora distinctly exceeding
the elytra; the femoral teeth long and sharp, especially that on
the hind pair; the second joint of the hind tarsi longer than
broad. Venter with segment | longer than 2 behind the coxe;
segment 5 of 3 not impressed and with a stout erect seta on each
side at the hind margin.

Length, 4°8-5:2mm.; breadth, 2°4—2°8 mm.

Sarawak: 1 ¢ 1 9, on flowers of Vernonia arborea, Mt.
Matang, 1000 ft., 11.1914 (G@. #. Bryant).

380 DR. G. A. K. MARSHALL ON THE SPECINS

12. BALANINUS NIGRORUFUS, sp.n. (PI. I. fig. 1.)

3. General colour ferruginous, the elytra black; the forehead
and a narrow central stripe on the pronotum with dense yellow
scales, the sides of the latter with scattered and narrower yellow
scales; the elytra with the following yellow scaling: a dense
stripe on interval 2 from the base to the middle, a transverse
postmedian band on intervals 3-8, and. widely scattered isolated
scales elsewhere, interspersed with hair-like white scales; the
underside for the most part rather thinly clothed with narrow
yellowish scales, a compact patch of broad creamy scales before
the front coxe, and a similar patch on the mesepimeron and on
the whole side of the metasternum.

Head with the rostrum arising from above the middle of the
eye. ostrum about half the length of the body, with five
narrow carine at the base, the intervening strie being shallowly
punctate, the scrobes passing beneath the rostrum at the base
and scarcely extending beyond the antenne. Antenne inserted
distinctly beyond the middle of the rostrum, the scape not
reaching the eye; the funicle with joint 1 a little longer than 2,
joints 3 and 4 equal, 5 and 6 a trifle shorter ue equal, 7 equal
to 3; the club with joint 1 longer than 2, and 2 than 3.
Prothorax a trifle broader than long, the sides shightly rounded,
the apical constriction almost obsolete ; the apical margin slightly
arcuate dorsally, with very feeble postocular lobes; the upper
surface with rather shallow and barely confluent reticulate punc-
tures. Scutellun elongate and clothed with dense yellow scales.
Elytra slightly longer ‘than bro ad, the lateral margins sinuate,
the apices almost jointly rounded; the basal sutural depre ssion
broad and deep, and almost sy mmetrical, the apical third of the
suture with short, obliquely raised scales, strie 9 and 10 not
reaching the base. Legs comparatively short, the: hind femora
not exceeding the apex of the elytra, the femoral teeth short and
sharp: the front cox with a conical process; the hind tibie
with three-fourths of the upper surface clothed with broad yellow
scales, all the remaining scales on all the legs being setiform and
white. Venter with segment 1 much longer than 2 behind the
coxe.

Length, 446 mm.; breadth, 2°6 mm.

SARAWAK: | ¢, Kuching, 1897 (R. Shelford. type);
Quop, 7. iv. 1914 (G. E, Bryant).

13. BALANINUS SUBPARTITUs, sp. n. (PI. II. fig. 6.)

2. Colour black, variegated with dark brown, bright fawn and
white scales ; the head beneath with small separated white scales ;
the pr onotum with scattered fawn and white scales at the sides
and with a broad median stripe of elongate fawn scales, which is
broadest near the base and gradually narrows in front, the scales
converging towards a point on the median line behind the middle ;
the scutellamn with dense silvery white scales; the elytra thinly

OF BALANINUS OCCURRING IN BORNEO. 381

clothed for the most part with fawn or brown scales and a few
seattered white ones, with a very indefinite transverse band
behind the middle formed principally of white scales; the proster-
num with large separated white scales on the antecoxal area and
fine white and brown scales at the sides; the rest of the sternum
with narrow white and fawn scales, except for a dense patch of
white ones on the mesepimeron; the venter with dense white
scaling on the two basal segments, and fawn scales on the others.

Head with the upper surface of the rostrum on a line with
the top of the eye. Rostrum strongly bent downwards beyond
the middle, about two-thirds the length of the body, strongly
punctate in the basal section, with a low central carina, as well
as a lateral one, but no lateral furrow, the scrobe passing beneath
the base of the rostrum but not exceeding the antenna. Antenne
inserted far behind the middle of the rostrum ; the funicle with
the two basal joints equal, 3 longer than 4, 4-7 subequal and
with appressed white sete; the club ovate, as long as the two
preceding joints and clothed with golden pubescence, joint |
rather longer than 2, and 2 than 3. Prothorax much broader
than long (10:7), broadest at the base, the sides strongly rounded,
the apical constriction deep, the apical margin truncate dorsally
and with distinct postocular lobes; the upper surface with deep
reticulate punctures. Scztellwm strongly elevated, longer than
broad. lytra with the lateral margins not sinuate, the apices
jointly rounded, the sutural impression rather shallow and asym-
metrical, interval 1 of the right elytron being lower and broader
than that of the left, the apical half of the suture with stout
raised spine-like bristles, striz 9 and 10 not nearly reaching the
base. Legs moderately long and stout; the hind femera some-
what exceeding the end of the elytra, the femoral teeth broad
and sharp, that of the hind pair broadly laminate and ending in
a sharp point; all the tibiz clothed above with broader scales,
the basal sinuation obsolete on the anterior pairs, but well marked
on the hind one, the upper edge of which is distinetly sinuate.

Length, 4-4-4:°6 mm.; breadth, 2°5-2°6 mm.

SaRAwAk: 1 9, on flowers of Vernonia arborea, Mt. Matang,
1000 ft., 12.11.1914, and 1 9, Mt. Merinjak, 1500 ft., 25.v. 1914
(G. E. Bryant).

14, BALANINUS MG@STUS, Sp. n.

@. Black, thinly clothed above with recumbent white setz
and without any markings formed of scales ; the seutellum with
dense white scales; the lower surface thinly clothed with white
sete or narrow scales, with the following broader white scales :
a few on the prosternum in front of the coxe, a dense patch on
the mesepimeron, a small patch at the apex of the metepisternum,
and a small lateral patch on the 2nd ventral segment.

Head with the rostrum arising from above the middle of the
eye. ostrwm shorter than the body (5:7), punctate at the

382 ‘DR. G. A. K, MARSHALL ON THE SPECIES

base, with a smooth central line and a very shallow punctate
lateral furrow, bounded below by a fine carina ; the scrobe passing
beneath the rostrum at the base and not exceeding the antenna.
Antenne inserted a little behind the middle of the rostrum, the
scape not reaching the eye; the funicle with joint 1 very slightly
longer than 2, 3 a little longer than 4, and 4-7 subequal; all the
joints with a few recumbent white sete; the club about as long
as the two preceding joints and with joint 1 distinctly longer
than 2. Prothorax broader than long (11:9), broadest at the
base, the sides rather strongly rounded, the apical margin slightly
arcuate dorsally and without any postocular lobes; the upper
surface with reticulate punctures and a low median carina.
Scutellum slightly raised and a little longer than broad. Hlytra
agreeing with the description of those of B. subpartitus, but more
pomted behind. JZegs rather long and stout, the hind femora
exceeding the apex of the elytra; the femoral teeth compara-
tively small; the tibize clothed only with setiform scales, the dorsal
edge straight, and the basal sinuation very shallow.
Length, 55 mm.; breadth, 3 mm.

Sarawak: 1 2, Quop, 6.i11.1914 (@, #. Bryant).

15. BALANINUS TUMIDIROSTRIS, sp. n. (Pl. IJ. fig. 10.)

3. Black, with blackish recumbent sete, and the following
markings formed of rather broad white scales: the prothorax
with a large basal patch on each side, linked together by a narrow
band along the base and continued outwards so as to join up with
the mesosternal patch ; the elytra with a common square median
patch extending on each side to stria 2, a broad transverse band
on each side about the middle extending from stria 6 to the
margin, a short stripe near the apex of interval 3, a small spot
near the apex of interval 2, a fringe of single white scales along
the apical margin and a few narrow white scales sparsely scattered
on the black areas; the scutellum with dense white scales; the
lower surface fairly densely clothed with white scales, but more
thinly so on the sides of the prosternum and on the basal half of
the metepisternum.

Head with the upper edge of the rostrum above the top of the
eye. ostrwm less than half the length of the body (5:11),
strongly curved downwards from the base, thick at the base and
eradually widening to the insertion of the antenne, then abruptly
narrowed and becoming cylindrical and shiny, ath rows of fine
punctures ; the basal area opaque and rugose, with three distinct
caring ; the scrobe passing immediately beneath the rostrum, so
that the scape in repose is quite invisible from above. Antenne
inserted well behind the middle of the rostrum; the scape very
short (shorter than the two basal joints of the funicle together).
the club forming nearly half its length, and not quite reaching
the eye; the funicle thinly clothed with recumbent light brown
sete, joint | a little longer than 2, and 3 than 4, 5-7 shorter and

OF BALANINUS OCCURRING IN BORNEO. 383

equal to one another ; the club nearly as long as the three pre-
ceding joints, the apex obtuse and joint 1 distinctly longer than 2.
Prothorax somewhat broader than long (5:4), broadest at the
base, the sides almost straight and slightly converging from the
base to the middle, then rounded, and strongly constricted at the
apex ; the apical margin truncate dorsally and with feeble post-
ocular lobes; the upper surface with deep reticulate punctures.
Scutellum small and narrow, twice as longasbroad. Hlytra with
the shoulders very sloping, the conjoint apices shallowly sinuate,
the sutural impression shallow and asymmetrical, the sutural
margin of the right elytron being distinctly sinuate, and the
apical half of the suture with raised interlocking sete ; striz 9
and 10 not nearly reaching the base. Zegs not long, the hind
femora not extending beyond the elytra; the femoral teeth long
and sharp, the tibiz with the basal sinuaticn shallow. Venter
with segment 1 flattened in the middle; segment 5 with the
apical margin broadly sinuate, the two angles bearing a tuft of
brown hairs; the pygidium with erect brown hairs shorter than
the ventral tufts.

Length, 45 mm.; breadth, 2°2 mm.

Sarawak: 2 ¢ d, Kuching (Rf. Shelford).

The structure of the rostrum distinguishes this species from
all the others here described, but it is probable that the character
may prove to be less pronounced in the female sex ; the conceal-
ment of the scrobe and tie shortness of the scape are also peculiar
characters.

16. BALANINUS TRINOTATUS, sp. n. (PI. II. fig. 8.)

$ 2. Colour dull black, thinly clothed above with narrow
brown scales interspersed with white ones, the latter frequently
deciduous but always present along the base of the elytra, which
also bear an elongate dense patch of broader white scales on
interval 2 behind the middle and a few white scales and bristles
on interval 1 at the same place; the scutellum with dense white
scaling ; the lower surface clothed with fairly broad but isolated
white scales, and with much denser patches on the mesepimeron,
at the posterior angles of metasternum, and along the sides of the
abdomen.

Head with the rostrum arising from well above the middle of
the eye, its upper surface being a little above the level of the top
of the eye in both sexes. ostrwm about three-fifths the length
of the body in both sexes, stout in the basal half and distinctly
narrower beyond the antenne ; the basal area with two irregular
rows of punctures on each side of the smooth median line, then
a broad lateral furrow, edged below by a sharp carina; the scrobe
passing beneath the rostrum at the base. Antenne inserted at
about the middle of the rostrum in both sexes, the scape not
uearly reaching the eye; the funicle with jot 1 as long as 2
and half of 3, joints 3-5 gradually diminishing, and joints 4, 6

384 DR. G. A. K. MARSHALL ON THE SPECIES

and 7 subequal in length; the club a little longer than the three
preceding joints, and with joint 1 distinctly longer than 2. Pro-
thoraw nearly as long as broad (5:6), broadest at the base, the
sides gently rounded, the apical constriction shallow, the apical
margin truncate dorsally and with feeble postocular lobes; the
upper surface with shallow reticulate punctation and occasionally
with a faint central carina. Scwtellum alittle longer than broad.
Hlytra with the lateral margins not sinuate, the apices jointly
rounded, the sutural impression moderately deep and quite asym-
metrical, being almost entirely confined to the right elytron and
extending backwards to the top of the declivity, and the apical
third of the suture set with raised interlocking black bristles.
Legs moderately long, thinly clothed with white setiform scales ;
the hind femora shortly exceeding the apex of the elytra, the
femoral teeth rather long and sharp; the tibize with the dorsal
margin straight and the basal sinuation very shallow. Venter of
3 with two tufts of fairiy long yellowish sete on the apical
margin of segment 5; the pygidium with dense erect blackish
sete.

Length, 3°5-4°5 mm.; breadth, 2--2°5 mm.

Sarawak: 2 ¢ 5 Q, Kuching, vii. 1900 (2. Shelford); 1 3
1 2, Mt. Matang, 1000 ft., xii. 1913 (@. #. Bryant, type):

17. BALANINUS CONSOCIUS, sp. n

3. Very closely allied to B. trinotatus, sp. n., and apart from
its larger size, differing only in the following characters :—The
club of the antenne proportionately smaller, being a little shorter
than the three preceding joints ; the prothorax distinctly more
transverse; the elytra with a post-median patch of white scales
on interval 3 similar to that on 2, but a little shorter, without
any white scales on interval 1 and with the sutural bristles
entirely black; the fifth ventral segment of the $ with the
apical tufts composed of very short sete and a single long one.

Length, 5°2 mm.; breadth, 3 mm.

Sarawak: 1 ¢, Mt. Matang, 1000 ft., 26.1.1914 (@. #. Bryant).

18. BALANTWUS UNIFAsScIATUS, sp.n. (PI. IT. fig. 5.)

3 9. Black, subnitid, thinly clothed above with narrow white
and light brown scales, and with markings formed of broader
white scales; the pronotum with about a dozen broad white
scales at each basal angle; the scutellum with dense white scales;
the elytra with a postmedian macular wh?te band lying between
strix 1 and 8 or 9; the sternum clothed with separated white
scales and the usual denser patches, and the abdomen with denser
sealing throughout.

Head with the rostrum arising from about the middle of the
eye in both sexes. Rostrum— 6, shorter than the body (3:5),
the basal portion with a single row of punctures on each side,
followed by a distinct lateral furrow which is bounded below by

OF BALANINUS OCCURRING IN BORNEO. 385

a sharp carina, the scrobe passing beneath the base of the rostrum
and not exceeding the antenna; 2, longer than the body (13: 11),
the basal portion with two rows of punctures on each side and
the lateral furrow less distinct. Antenne inserted at (¢) or far
behind (@) the middle of the rostrum, and the scape not nearly
reaching the eye; the funicle with joint 1 a trifle longer than 2,
3 longer than 4, and joints 4-7 subequal; the club equal to the
three preceding joints, its first joint distinctly longer than the
second. Prothorax broader than long (11:8), broadest at the
base, with the sides gently rounded, and shallowly constricted at
the apex, the apical margin truncate and with very feeble post-
ocular lobes ; the upper surface with closely reticulate punctures.
Scutellum longer than broad. Hlytra with the lateral margins
not sinuate, the apices jointly rounded, the sutural impression
very shallow and confined to the right elytron, the raised sutural
bristles, varying from yeilowish or light brown to blackish. Legs
thinly clothed with white setiform scales; the hind femora
distinctly exceeding the elytra, the tooth on the femora dilated
at the base; the tibie straight and without any basal sinuation.
Venter of ¢ with the basal segment merely flattened in the
middle and not impressed, segment 5 also somewhat flattened
and with two isolated erect hairs on the apical margin; the
pygidium with a dense tuft of erect white sete.

Length, 2°75-4:25 mm.; breadth, 1°5-2°25 mm.

Sarawak: 1 3, Simunjon, 1855 (A. &. Wallace, type); 1 9,
Kuching, 16. vil. 1900 (2. Shelford).

19, BALANINUS DECEMNOTATUS, sp.n. (PI. II. fig. 1.)

2. Dull black, thinly clothed above with recumbent blackish
narrow scales or sete and with markings formed of broad white
scales; the pronotum with a basal trapezoidal white patch on
each side, the base of which lies between strie 4 and 8 of the
elytra; the elytra with a few white scales at the base of intervals -
2 and 3, behind the middle a transverse white patch between
strie 1 and 3 and another on a line with it between striz 5and 9,
and finally a short white line at the apex of interval 2; the lower
surface for the most part thinly clothed with widely separated
narrow white scales, with the following parts covered with dense
broader scales: the antecoxal area of the prosternuim, the meso-
sternal process, the mesepimeron and a part of the mesepisternum
adjoining it, the basal external angle of the metasternum, and a
large lateral patch on the two basal ventral segments.

Head with the rostrum arising from about the middle of the
eye. ostrum slender, distinctly longer than the body (17:14),
with two rows of very fine punctures on each side in the basal
portion and beneath them an obsolescent punctate lateral furrow,
bounded below by a carina; the serobe passing beneath the
rostrum at the base and not exceeding the antenna. Antenne
inserted far behind the middle of the rostrum, the scape not

386 DR. G. A. K. MARSHALL ON THE SPECIES

reaching the eye; the funicle with joint 1 very slightly longer
than 2, and 3 longer than 4, joints 4, 5 and 7 subequal and 6 a
little shorter ; the club as Jong as the preceding 23 joints, its first
joint nearly twice as long as its second. Prothorax broader than
long (6:5), broadest at the base, with the sides almost straight
and gradually narrowing from the base to the middle, then rounded
and with a distinct but shallow apical constriction, the apical
margin truncate dorsally and without any postocular lobes; the
upper surface regularly and reticulately punctate throughout.
Scutellum subquadrate, with a few minute brownish scales.
Elytra with the lateral margins not sinuate, jointly rounded at
the apex, the sutural impression very broad, deep and almost
symmetrical, the apical third of the suture with raised inter-
locking black bristles. Legs long and slender, thinly clothed with
white setiform scales; the hind femora with almost the whole
clavate portion extending beyond the elytra, the tooth’ on the
femora triangular at the base; the tibie straight, without any
basal sinuation.
Length, 55 mm.; breadth, 3 mm.
Sarawak: | 9, Lundu, 6.1.1914 (G. #. Bryant).

20 BALANINUS QUINCUNX, sp. n. (PI. LI. fig. 11.)

3 @. Black, dull or subnitid, thinly clothed above with
blackish narrow scales or sete, and with markings formed of
broad white scales; the pronotum with a transverse elliptical
basal white patch on each side extending from stria 3 almost to
the mesepimeron; the elytra with a short white stripe at the
base of interval 1, behind the middle a transverse macular patch
on intervals 2 and 3, on the same line with it a band on intervals
7-10 (or 6-10), and a short apical white stripe on interval 1 ;
the lower surface fairly closely covered with separated broad
white scales, with the usual denser lateral patches, that on the
prosternum extending upwards along the apical constriction to
the level of the middle of the eye.

Head with the rostrum arising well above the middle of the
eye, its upper surface being on a level with the top of the eye in
both sexes. ostrwum— 3, two-thirds the length of the body,
the basal portion coarsely punctate, with a smooth median line
and a broad lateral furrow bounded below by a sharp carina, the
scrobe passing beneath the rostrum at some distance from the
base and distinctly extending beyond the antenne; in the 9,
nearly as long as the body and more strongly curved, the puncta-
tion much finer and the lateral furrow less distinct. Antenne
inserted at (¢) or somewhat behind (9) the middle of the
rostrum, the scape not nearly reaching the eye ; the funicle with
the two basal joints equal, 3 longer than 4, and joints 4—7 sub-
equal; the club about equal to the three preceding joints, and its
first joint distinctly longer than the second. Prothorax (¢ )
distinctly transverse, the sides subparallel from the base to the

|
|
:

OF BALANINUS OCCURRING IN BORNEO. 387

middle, then strongly rounded and with a deep apical constriction,
the apical margin truncate dorsally and with no trace of post-
ocular lobes; in the 9 thesides are gently and regularly rounded,
and with hardly any apical constriction; the upper surface
coarsely and reticulately punctate throughout. Scutellwm slightly
longer than broad, with a few dark scales. Hlytraw with the
lateral margins straight, the apices jointly sinuate, the sutural
impression confined to the right elytron, shallow and extending
from the apex of the basal stripe to the top of the declivity, and
the apical third of the suture with raised interlocking black
bristles. Zegs rather thinly clothed with white setiform scales,
the hind femora extending shortly beyond the elytra, and the
tibie straight and with a shallow basal sinuation. Venter of 3
with the anal segment simple and with an apical fringe of
yellowish sete; the pygidium with long dense erect brown set.

Length, 4°5 mm.; breadth, 2°5 mm.

Sarawak: 1 ¢, Mt. Matang, 11.1914 (type), and 1 2, Quop,
iv. 1914 (@. #. Bryant).

Like a small B. imitator, sp. n., with reduced white markings,
except that the basal stripe on interval 1 of the elytra is at least
twice as long, the rostrum is more coarsely punctate, the last
ventral segment of the ¢ lacks the erect hairs, and the sete on

fo)
the pygidium are much longer.

21, BALANINUS ANALIS, sp.n. (PI. IT. fig. 9.)

3 2. Dull black, thinly clothed above with recumbent blackish
scale-like setee, with markings formed of dense broad white scales;
the prothorax with a large transverse basal white patch on each
side extending from the mesepimeron inwardly as far as stria 5
of the elytron, and a vertical lateral band on each side of the
apex extending from the prosternum up to the level of the top
of the eye; the elytra with a white patch surrounding the
scutellum on the bases of intervals 1-4, a broad white band just
behind the middle between strize 1-10, with its anterior edge
deeply sinuate and the portion on interval 4 reduced and some-
times absent, a short white stripe on the apical fourth of interval 1,
a few scales at the apex of interval 2, a row of single white scales
on the apical fourth of interval 10, and a similar row of much
narrower scales on the extreme margin ; the lower surface fairly
densely clothed with white scales, which are mostly narrow in
the median area and broad laterally, being densest on the front
portion of the prosternum, on the mesepimeron, mesosternal
process, external apical angle of the metasternum, and on the
extreme lateral margins of the abdomen; in the ¢ the scales in
the centre of the metasternum are very long, narrowly lanceolate
and obliquely raised, and those in the centre of the first ventral
segment are setiform.

Head with the rostrum arising in the ¢ far above the middle
of the eye, its upper surface being on a level with the top of the

Proc. Zoou. Soc.—1919, No. XX VII. 27

388 DR. G. A. K. MARSHALL ON THE SPECIES

eye; in the @ it arises only slightly above the middle of the eye,
its upper surface lying much below the top of theeye. Rostruwm—
3, only a little longer than half the body, the basal half with
a smooth central line and two irregular rows of punctures on
each side, then a broad lateral furrow, bounded below by a distinct
carina, the scrobe passing beneath the rostrum at the base and
extending shortly beyond the antenna; in the 9, slightly longer
than the body, with the punctures and lateral furrow much less
distinct. Antenne inserted near (d) or far behind (@) the
middle of the rostrum, the scape not reaching the eye; the funicle
with joint 1 slightly shorter than 2, joint 3 distinctly longer
than 4, and 4-7 subequai; the club slightly shorter than the three
pr eceding joints, its first joint longer than the second. Prothorax
distinctly broader than long (3: 2), broadest at the base, the sides
straight and gradually narrowed to the middle, then Homnited and
with a well-marked apical constriction, the apical margin truncate
dorsally and sloping obliquely forwards behind the eye; the
upper surface reticulately punctate throughout, sometimes with
a trace of a median line. Scutellwm subquadrate, with a few
dark scales. Hlytra with the lateral margins not sinuate, the
apices separately rounded, the sutural impression deep and
asymmetrical, the apical third of the suture with raised inter-
locking black bristles. Legs rather densely clothed with white
setiform scales; the hind femora extending well beyond the
elytra, and the mid femora with a fringe of hairs on the lower
edge in both sexes; the tibie straight and with the basal sinua-
tion very shallow. Venter of the 3 with a large deep depression
on segment 1, segment 2 with the posterior margin strongly
angulate laterally, 3 and 4 flattened in the middle, and 5 excavate
and bare in the middle, the posterior edge with a broad deep
emargination and fringed with hairs, the outer angles being
visible from above in the form of two conical prominences ; in
the 9 segment 5 bears an oval median impression at the apex.

Length, 4—5°25 mm.; breadth, 2-2°5 mm.

Sarawak: 4 ¢ 6,10 2 92, Kuching, v.1909 (J. H. A. Lewis,
type); 1 ¢, Puak, 30. iv. 1914 (@. #. Bryant).

22. BALANINUS IMITATOR, Sp. Nn.

3 9. Extremely similar to B. analis, sp. n.. but differing as
follows :—The apical white band on the prothorax does not extend
above the middle of the eye, and the lateral basal patch hardly
exceeds the fourth stria; on the elytra there is no apical row of
scales on interval 10 or on the margin; in the middle of the
metasternum of the ¢ the scales are small, flat and normal, and
those on the first ventral segment are not setiform. The rostrum
of the ? is shorter than the body (7:9 or 10), and the scrobe
extends much further beyond the antenna in both sexes. The
antenne have joint 1 of the funicle slightly longer than 2, and
the club is nearly as long as the four preceding joints. The

OF BALANINUS OCCURRING IN BORNEO. 389

sutural impression is rather shallower and the sutural bristles
less raised. The hind femora only slightly exceed the apex of
the elytra. The basal ventral, impression in the ¢ is much
shallower, the apical margin of segment 2 is quite straight and
not angulate externally, segments 3 and 4 are transversely
convex, and segment 5 is truncate at the apex, with a very
shallow median impression and a few erect hairs on each side of
it; the last ventral segment of the Q is simple and without any
impression.

Length, 3-4 mm.; breadth, 1°6-2°4 mm.

Sarawak: 73 5,32 9 (1Q on LHleocarpus), Quop, ii. 1914
(G. #. Bryant); 13,1 92, Puak, v.1914(G@. £. Bryant, type).

23. BALANINUS DECEPTOR, sp.n. (PI. IT. fig. 3.)

6. Dull black, thinly clothed above with small black narrow
scales or setze, and with markings formed of broad white scales ;
the pronotum with a narrow transverse basal white patch on
each side, almost reaching the mesepimeron externally and stria 5
internally ; the elytra with a few white scales at the base of
interval 1, a narrow transverse band at the base of intervals 2—4
(or 5), the portion on 2 being nearly twice as long as the rest,
and a narrow sinuate macular white band behind the middle
between striz 1 and 9, the spot on interval 4 being absent and
those on 2 and 9 a little longer than the others; the sternum
with separated narrow white scales and with dense patches of
broader scales on the precoxal area of the prosternum (emitting
a band up the apical constriction to the level of the middle of the
eye), on the mesepimeron, the mesosternal process, and the apical
external angle of the metasternuin; the venter more evenly
clothed with broader white scales.

Very similar structurally to P. analis, sp. n., and apart from
its larger size differing only in the following characters :—The
scutellum has a broad and deep V-shaped incision on its anterior
margin ; the tooth on the femora is triangularly dilated at the
base, and the mid femora entirely lack the fringe of hairs on the
lower edge; the narrow scales on the metasternum are not hair-
like and raised, but short, recumbent, and truncately fringed at
the apex; the second ventral segment is not angulate externally,
and the fifth is not excised at the apex, but bears a median
depression almost concealed by a dense tuft of incurved hairs
arising from a low elevation on each side of it; and the pygidium
is much more exposed.

B. imitator, sp. n., differs in having the scutellum not or ver
faintly indented in front; the first joint of the funicle is slightly
longer than the second, and the club is relatively longer; the
hind femora are shorter and the femoral tooth is less dilated at
the base; the basal ventral impression is much shallower and
the structure of the fifth ventral segment is much more simple.

Length, 55 mm.; breadth, 3 mm.

SaRawak: 2 5 3, Puak, iv-y, 1914 (G. #, Bryant).

27%

390 DR. G. A. K. MARSHALL ON THE SPECIES

24, BALANINUS COMMODUS, sp. n.

¢@. Black, subnitid, thinly clothed above with regu beat
blackish scales or sete, and with markings formed of broader
white scales; the prothorax with a subquadrate basal white patch
6n each side opposite intervals 4-7 of the elytra; the scutellum
with a few minute white scales; the elytra with a few white
scales at the base of interval 2, behind the middle a macular
white patch on intervals 2 and 3, and another in line with it on
intervals 7 and 8; the lower surface fairly closely covered with
separated white scales and with the usual denser lateral patches
of broader scales.

The structural characters-as in 5. guincuna, sp. n., except the
following :— Head longitudinally convex, with the rostrum arising
about the middle of the eye. Rostrum a little longer than half
the body (5:9), the serobe not exceeding the antenna. © Pro-
thorax less abruptly constricted in front and with feeble post-
ocular lobes. Seutellam with a few white scales. Venter of J
with a shallow median impression on the anal segment, without
any tufts of hair or apical fringe; the pygidium with short
white setze.

Length, 3°75 mm.; breadth, 2 mm.

Sarawak: | ¢, Quop, iv. 1914 (@. H. Bryant).

25, BALANINUS GRYPUS, sp.n. (PI. II. fig. 12.)

3. Just like a very small specimen of B. analis, sp. n., but
the white markings differ as follows: there is no vertical apical
band on the prothorax and no apical lines on the elytra; the
basal patches on the prothorax are broadly truncate internally
(instead of tapering to a point), and the postmedian band on the
elytra only reaches stria 8.

The structural characters agree also, except in the following
respects : the rostrum is fon geh in proportion to the body (5:7);
the funicle has joint 1 equal to 2; the prothorax is more elongate
(94-8), with the sides gently and more evenly rounded and
scarcely constricted at the apex, the sculpturing being propor-
tionately coarser; the mid femora have no fringe “of hairs
beneath ; the second ventral segment is not angulate externally,
and the fifth 1s quite simple and without any tufts of hairs; the
pygidium is clothed with dense erect white sete (in 4. analis and
imitator they are own

Length, 2°75 mm.; breadth, 1°5 mm.

SARAWAK: | GC, Quop, 24. i11. 1914 (G@. #. Bryant).

26. BALANINUS SHELFORDI, sp.n. (PI. II. fig. 15.)

2. Colour piceous brown, with dark brown scaling and mark-
ings of fawn and white scales; the prothorax with a narrow
central stripe of transverse fawn scales, and on each side of the
base a quadrate patch (occupying one-fourth the width and extend-
ing longitudinally nearly to the middle) formed of. fawn scales

OF BALANINUS OCCURRING IN. BORNEO. 391

which gradually turn to white at the base of the thorax, the inner
anterior angle of the patch being produced as a narrow longi-
tudinal stripe almost to the apex and with a few fawn scales just
below its apical termination; the elytra with a complete fawn
stripe on interval 1, and the following white markings: an
oblong patch at the base of intervals 2 and 3 3, just behind the
posterior line of this patch a small spot on intervals 3 and 5,
another at the base of 7, and just behind the middle a band on
intervals 2-4, which is broadest internally and narrows out-
wardly ; the front portion of the prosternum, the sides of the
metasternum, and the sides of segments 2-4 of the abdomen with
fairly dense creamy white scales, those on the mesepimeron and
metepisternum still denser.

Head with the front broader than the base of the restrum,
with a smooth central ridge and some fawn scaling on each side ;
the rostrum arising far below the middle of the eye, its lower
surface being on a level with the lower margin of the eye.
Rostrum shorter than the body (5:7), very smooth and shiny,
almost impunctate even at the base, and without any lateral
furrow or carina; the scrobe entirely lateral in position.
Antenne inserted far behind the middle of the rostrum, with the
scape reaching the eye; the funicle with joint 1 only shghtly
longer than 2, joints 4-7 subequal; the club nearly as long as the
four preceding joints and with joint 1 slightly longer than 2.
Prothorax slightly broader than long (5:4), the sides parallel
from the base to the middle, then rapidly narrowing, but scarcely
constricted at the apex ; the apical margin truncate dorsally and
without postocular lobes ; the upper surface with reticulate
punctures and a low central carma. Scutellum slightly trans-
verse, with a few pale fawn scales. Hlytra with the apices
separately rounded, the sutural. impression rather shallow
and quite symmetrical, and the apical half of the suture
with raised interlocking bristles. Legs moderate, the hind
femora slightly exceeding the apex of the elytra; the femoral
teeth stout and sharp; the tibie with the basal sinuation shallow,
the hind pair with the dorsal edge distinctly sinuate.

Length, 2°75 mm.; breadth, 1°5 mm.

SARAWAK: 1 2, on flowering tree, Kuching, 17. vii. 1900 (A.
Shelford).

Type in the Oxford University Museum.

27. BALANINUS PUSIO, sp. 0. (Etre yl)

2. Colour rather shiny black, thinly clothed above with
narrow recumbent dark brown scales and with markings formed
of broad white scales; the rostrum reddish brown ; the pronotum
with a transverse basal white patch on each side, almost touching
the mesepimeron externally and extending as far as stria 3
internally ; the elytra with a circum-sentellar white patch at
the base of intervals 1 and 2, and a rather smaller patch slightly

392 DR. G. A. K. MARSHALL ON THE SPECIES

behind the middle on the same two intervals but not actually
reaching the suture; the prosternum with dense white scales in
front of the coxe, but none at the sides; the rest of the lower
surface with separated white scales, except on the mesepimeron
and along the sides of the metasternum and abdomen, where the
scales are dense.

Head with the forehead slightly broader than the base of the
rostrum, the lower surface of which is on a level with the lower
margin of the eye. Rostrum as long as the body, only slightly
curved downwards in the apical third, very shiny and impunctate,
even at the base where it is slightly compressed, and without any
lateral furrow or carina; the scrobe entirely lateral and not
exceeding the antenna. Antenne inserted far behind the middle
of the rostrum, with the scape reaching the eye; the funicle
with joint 1 equal to 2 + 3 4 4, joint 2 half as long again as 3,
joints 3-5 gradually diminishing in length, 6 and 7 each about
equal to 4; the club a little longer than the three preceding
joints, and with joint 1 slightly longer than 2. Prothoraw rather
broader than long (16:13), broadest at the base, with the sides
moderately rounded, the apical constriction shallow, the apical
margin slightly arcuate dorsally and without postocular lobes ;
the dorsal surface rather coarsely reticulate throughout. Sceutellam
transverse, and with only a few narrow brown scales. Hlytra
with the lateral margins straight, the apices jointly rounded, the
basal sutural depression deep and asymmetrical, lying mainly on
the right elytron, and the apical fourth of the suture with raised
interlocked whitish bristles. Zegs mederately long, thinly
clothed with white setiform scales; the hind femora slightly
exceeding the elytra, the tooth on the femora narrow ; the tibiz
straight, with the basal sinuation shallow.

Length, 2 mm.; breadth, 1-2 mm.

SARAWAK: 1 9, Mt. Matang, 1000 ft., 11.01.1914 (@. Z.
Bryant).

28. BALANINUS DELICATULUS, sp. n. (PI. II. fig. 13.)

3 2. Colour rather shiny black, thinly clothed above with
small dark brown scales and with patches of broad white
seales ; the rostrum blackish near the base and reddish brown
externally ; the pronotum with a transverse basal white patch on
each side linking up with the mesosternal patch externally and
reaching stria 3 internally; the scutellum with dense white
sealing; the elytra with a broad white stripe on interval 1
reaching from the base nearly to the middle, and behind the
middle a transverse white band extending outwardly to interval 4
or 5 and not quite reaching the suture; the lower surface clothed
with separated small white scales and with dense patches of
broader scales on the following areas: the antecoxal area of the
prosternum, the whole of the mesepimeron, the posterior half of

the metepisternum, and a stripe along the extreme lateral edge
of the abdomen.

OF BALANINUS OCCURRING IN BORNEO. 393

Head with the rostrum arising from the middle (¢) or below
the middle (9) of the eye; the forehead narrower than (¢) or
about as broad as the base of the rostrum(@). ostrum shorter
than the body in both sexes (¢, 8:13; 9, 10:13), rather
strongly curved downwards in the apical third; ¢, strongly
striato-punctate in the basal half, with a smooth central line and
a distinct furrow above the scrobe, which latter passes beneath
the rostrum at the base; 9, very finely punctate in the basal
area, without any lateral furrow, and with the scrobe entirely
lateral in position. Antenne inserted at (¢) or far behind (¢ )
the middle of the rostrum; the funicle with joint 1 distinctly
longer than 2, joints 3, 4, 6 and 7 subequal in length, 5 shorter ;
the club nearly as long as the four preceding joints, its first joint
slightly longer than the second. Prothorax almost as long as
broad, the sides nearly parallel from the base to the middle,
thence roundly narrowed and shallowly constricted near the apex,
the apical margin truncate dorsally and without postocular lobes ;
the upper surface distinctly more convex than usual, reticulately
punetate and with a low median carina. Scutellwm nearly circular.
Elytra with the lateral margins not sinuate, the apices jointly
rounded, the suture not impressed and with a row of slightly
raised bristles on its apical third. Zegs moderately long, thinly
clothed with white setiform scales; the hind femora shortly
exceeding the apex of the elytra, the tooth narrow; the tibie
straight and with the basal sinuation distinet. Venter of 3 with
a few erect sete near the apex of segment 5.

Length, 2°5-2°38 mm.; breadth, 1°25 mm.

Sarawak: 1 3, Mt. Matang, 1000 ft. 13.11.1914 (G@. &. Bryant
—type); | ¢ 1 @, Kuching, 16. vi. 1900 (2. Shelford).

29. BALANINUS EUGENIA, sp. nb.

3 2. Ground-colour piceous black, the elytra and tibie dark
red-brown; the pronotum with the usual narrow brown scales,
mostly replaced by slightly broader pale scales on the basal third
and isolated pale scales scattered over the rest of the surtace; the
elytra fairly densely clothed with broader fawn-coloured scales,
variegated with darker patches formed of narrower brown scales ;
the sternum with separated whitish scales, but with dense fawn
or orange-coloured scales at the sides of the imeso- and meta-
sternmin, and a small patch of similar scales in the middle and on
each side of the prosternum ; the venter with fairly dense whitish
scales.

Head with the rostrum arising from about the middle of the
eye in both sexes. ostrwm about three-fourths the length of
the body in both sexes, but much more strongly curved in the d,
fairly strongly punctate in the basal section, with a smooth
central line and a lateral furrow reaching the antenna, the scrobe
passing beneath the base of the rostrum and not exceeding the
antenna. Antenne inserted well behind the middle of the

394 DR, G. A. K. MARSHALL ON THE SPECIES

rostrum in both sexes, the scape not quite reaching the eye;
the funicle with joint 1 not or very slightly longer than 2, joiut 3
a little longer than 4, 4-7 almost equal in length and about

3+ times as long as broad; the club about as long as the four
preceding joints, joint 1 rather longer than 2, which is equal
to 3. Prothoraw rather broader than long, broadest at the base,
the sides fairly strongly rounded, the apical constriction distinct,
the apical margin truncate dorsally and with feeble postocular
lobes; the upper surface with close reticulate punctation.
Scutellum quadrate, covered with dense fawn scaling. Hlytra
with the lateral margins straight, the apices forming a joint
shallow sinuation, the suture not impressed and without any
raised sete near the apex. Legs rather short, the hind femora
only slightly exceeding the end of the elytra, the femoral teeth
stout and sharp, the tibise evenly clothed with setiform scales.
Venter with segment 1 longer than 2 behind the coxe, segment 5
of ¢ sinuate at the apex and with a short tuft of whitish hairs
on each side.

Length, 3°2-3°6 mm.; breadth, 1°6-1°8 mm.

Sarawak: 1 ¢, Mt. Matang, ii. 1914, and 2 2 9, on Hugenia,
Quop, 26. 11. 1914 (@. H. Bryant).

30. BALANINUS LONGICLAVIS, sp. n. (PI. IT. fig. 4.)

3 2. Colour brownish black, rather thinly clothed with small
narrow brown scales, sparsely intermingled with white ones, and
the following markings formed of dense broad white scales: a
small basal lateral patch on each side of the pronotum, the whole
of the scutellum, a broad stripe on the basal third of interval 1
on the elytra, and a narrow line on the apical half of the suture ;
the lower surface also with brown and white scales, but broader
than those above and the white ones greatly predominating,
especially on the abdomen, and with the following patches of
dense larger white scales : a small antecoxal one on the prosternum,
a large vertical one on the side of the mesosternum, a smaller one
at the external apical angle of the metasternum, and a small
one at the side of the second ventral segment.

Head with the rostrum arising from above the middle of the
eye, but its upper surface below the level of the top of the eye.
Rostrum a little shorter than the body (9:10 or 7:8) in both
sexes; d, moderately punctate in the basal area, with a smooth
central line and a well-marked lateral furrow; 2, with the
punctures and furrow much less distinct; the scrobe not extend-
ing beyond the antenne and passing beneath the base of the
rostrum in the ¢, but not in the 2. Antenne inserted far
behind the middle of the rostrum in both sexes, with the scape
almost touching the eye; the funicle with joint 1 equal to 2 + 3,
3 shghtly longer than 4, 4-6 short and subequal, 7 as long as
5 + 6 and more pubescent than the others: the club as long
as the six preceding joints, with its two basal joints equal and

OF BALANINUS OCCURRING IN BORNEO. 395

unusually elongate, being distinetly longer than broad (4: 3),
and joint 2 markedly constricted in the basal half. Prothorax
broader than long (13:10), broadest at the base, the sides
moderately rounded in the @, less so in the ¢, the apicai con-
striction being fairly distinct in the former and almost evanescent
in the latter, the apical margin truncate dorsally and without
any postocular lobes; the wpper surface with shallow reticulate
punctation throughout. Sewtedlum subquadrate. Hlytra with
the lateral margins not sinuate, the apices almost jointly rounded,
the suture not impressed at the base and without any raised
bristles near the apex. Zegs moderately long. the hind femora
scarcely exceeding the apex of the elytra; the tooth on the
femora long and rather narrow, the basal sinuation of the tibie
shallow and the dorsal edge straight. Venter of 3 with an erect
bristle on each side at the apex of the fifth segment.

Length, 3-5-4 mm. ; breadth, 2-2°25 mm.

Sarawak: 1 ¢ 192, Quop, i. 1914, and 1 9, Puak, 4.v. 1914
(G. EB. Bryant).

Distinguished by the remarkable elongation of the club, in
which it resembles the otherwise dissimilar B. sellatus, sp. 0.
This character also occurs in a few African species.

31. BALANINUS SELLATUS, sp.n. (PI. II. fig. 16.)

2. Colour black, with a large discal patch on the elytra red-
brown, the black areas mostly with small narrow, dark scales, the
brown area with broader fawn-coloured scales: the prothorax
with a pentagonal patch of fawn seales in the middle of the base,
not extending outwardly further than stria 2 of the elytra: the
fawn patch on the elytra somewhat ill-defined, reaching from
the base to beyond the middle and outwardly to about stria 4,
though there are a few fawn scales just beyond this, and an
indefinite elongate lateral patch extending from the margin to
stria 8; the lower surface rather densely and uniformly clothed
with yellowish and whitish scales.

Head with the forehead only slightly narrower than the base
of the rostrum, which arises well below the iniddle of the eye, its
lower surface being on a level with the lower margin of the eye.
Rostrum as long as the body, slender, smooth, and almost im-
punctate at the base, and without any lateral furrow or carina;
the scrobe entirely lateral and not exceeding the antenna.
Antenne situated far behind the middle of the rostrum, with the
scape reaching the eye; the funicle with joint 1 as long as 2-5,
joints 4-7 subequal in length, each a little shorter than 3, and 3
shorter than 2; the club very elongate, as long as the six pre-
ceding joints, with its first joint a trifle longer than the second,
which is not constricted at the base. Prothorav broader than
long (3:2), broadest at the base, rapidly narrowing thence to the
apex, the sides being almost straight and the apical constriction

5
obsolescent, the apical margin truncate dorsally and without any

396 DR. G. A. K. MARSHALL ON THE SPECIES

postocular lobes; the upper surface with shallow reticulate punc-
tation and a mere trace of a raised central line. Sewtellum
punetiform. lytra with the lateral margins not sinuate, the
apices jointly rounded, the sutural impression not very deep and
asymmetrical, being almost confined to the right elytron, and the
suture with a row of raised bristles near the apex. Legs mode-
rately long, but the hind femora scarcely exceeding the apex of
the elytra; the tooth on the femora narrow and sharp, the basal
sinuation of the tibiz shallow, and only the front tibiz with the
dorsal edge slightly sinuate.

Length, 3 vm. ; breadth, 15 mm.

Sarawak: 1 9, Puak, 3.v. 1914 (@. #. Bryant).

32, BALANINUS DISCRETICOXIS, sp. n. (PI. II. fig. 7.)

2. Colour brownish black, subnitid, and apparently bare
above, but really with minute dark recumbent sete and markings
of straw-yellow scales; the pronotum with a narrow median
stripe (the scales longitudinal, broad at the base and narrow in
front), a broader lateral stripe on each side extending from the
base to the middle (scales oblique), and a spot on each side near
the apex ; the scutellum densely scaled ; the elytra with a broad
stripe on the basal third of interval 1 and extending partly on to
2, and about half-way between the end of this stripe and the
apex a spot reaching from the middle of interval 2 to the middle
of interval 3; the prosternum with an antecoxal patch that is
whitish in the middle and. yellow latevally; the mesosternum,
metasternum, and the two basal abdominal segments each with a
large patch of very dense straw-yellow scales.

Head coarsely punctate, bare, with the forehead distinetly
broader than the base of the rostrum, which arises slightly below
the middle of the eye. Lostrwm as long as the body, very slender,
almost of equal width throughout, very smooth and shiny, scarcely
punctate even at the base, and without any lateral furrow or
carina; the scrobe entirely lateral and not exceeding the antenna.
Antenne inserted far behind the middle of the rostrum, with the
scape reaching the eye; the funicle with joint 1 equal to 2 4 3,
and joints 3-7 very slightly and progressively diminishing in
length ; the club equal to be three preceding joints, with joint 1
slightly longer than 2. Prothorav almost as long as broad, the
sides gently rounded, broadest about the middle, the apical con-
striction slight, the apical margin truncate dor sally and without
postocular lobes ; the upper surface with shallow reticulate punc-
tures throughout. Seuwtellum almost circular. Elytra with the
lateral margins straight, the apices jointly sinuate, the basal
sutural impression short, shallow, and quite symmetrical, the
apical third of the suture with raised dark bristles, and stria 9
remote from the base. Legs long, slender, and sparsely clothed
with small recumbent white sete; the hind femora shortly

exceeding the apex of the elytra, the femoral teeth small and

sie

————

OF BALANINUS OCCURRING IN BORNEO. 397

sharp; the front and hind tibie with the dorsal edge shallowly
sinuate ; the front coxe distinctly separated.

Length, 3-3°5 mm.; breadth, 1-4-1°75 mm.

Sarawak: 1 2 on flowers of Vernonia arborea, Mt. Matang,
1000 ft., 13.11 1914 (Go B. Bryant); 3 2, Quop, 21. im. 1914
(GE. B.).

A very distinct little species which differs from all other
members of the genus known to me in having the front coxee
separated.

WXPLANATION OF THE PLATES,

Prac Ti

Fig. Page Fig. Page
1. Balaninus nigrorufus, sp.n., g. 380 5. Balaninus nigrocinereus, sp.n.,
9. bispilotus, sp. n., 2. 371 i pean OUD
3. 2 excisipes, sp... 6, eee 3 sesyuilineatus, Sp. l.,
and apex of hind tibia. 373 ‘ 374

glabricollis, sp.n., @ . 370
bryanti, sp.u., 6 ... 869

A, » » bilineatus, sp.n., 9 . 375

lo a |

Prats II.

Fig. Page | Vig. Page
1. Balaninusdecemnotatus,sp.u., 2 385 9. Balaninus analis, sp.u., d...... 387
2. 33 gyvosicollis, sp. n.,g 3878 | 10. ss tumidirostris,sp.n.,¢ 382

3. Fe deceptor, sp.n., § ... 389 | 11. a quincune, sp. n., d... 386

4. 33 longiclavis, sp.n., 2. 394 12. 5 GVUPUS, Sp. N., O...... 390
5. FF unifasciatus,sp.n.,2. 3884 , 13. “ delicatulus, sp.n., 6. 392
6. Py subpartitus, sp. 0.,2. 380 14. is pusio, sp.n., 92 ...... 391
He a discreticowis,sp.u.,?. 396 | 15. 5 shelfordi, sp.u., 2... 390

3. - trinotatus, sp.u., 2 . 383 | 16. m sellatus, sp.u., 2... 895

ON NEW FISHES FROM LAKE TANGANYIKA. 399

25. On some new Fishes from near the West Coast of Lake
Tanganyika. By G. A. BouLenecer, F.R.S., F.ZS.

(Published by permission of the Trustees of the British Museum.)

Received September 5, 1919: Read November 4, 1919.'
(Text-figures 6-10.)

A few months ago I reported in these Proceedings* on a
collection of Fishes made at Albertville by M. Dhont-De Bie, of
the Belgian East African Expeditionary Force. On returning
home in May last, the same gentleman has brought me further
specimens from localities near but outside the great lake, among
which were representatives of six undescribed species.

These Fishes are from three localities :—

(1) A ditch (marigot) along the Lukuga River: Protopterus
ethiopicus Heck., Clarias hilgendorfi Bler. (pr eviously known from
Lake Rukwa), Auchenoglanis occidentalis C.& V., Anabas ctenotis,
sp. n., and Mastacembelus teniatus Blgr.

(2) Tumbwe, a village S.W. of the Kalemie River, on a small
stream flowing into ihe Niemba River, a tributary of the Lukuga :
Labeo dhonti, sp. n., Barbus euchilus, sp. n., B. miochilus, sp. n.,
BL. holotenia Bier.

(3), Kabeke, a village 30 miles south of Tumbwe, on the Niemba
River: Allabenchelys dhonti, sp. n., Amphilius platychir Gthr.,
Phractura lukuge, sp. n., an ‘association ver y suggestive, so far a as
genera are concerned, ae the rivers of South Cameroon, whence
the two first known representatives of the genus Allabenchelys
Blgr. were described, and where several Amphilius Gthr., and
Phractura Blgr., are at home.

It is remarkable that no members of the family Cichlide, which
form the great majority in the Tanganyika Fish-fauna, should
have been collected in these three localities. Characinide are
also absent.

Descriptions of the new Species.

LABEO DHONTI.

Body feebly compressed, its depth 5 to 52 times in total length.
Head 4 to 41 times in total length, its width 3 its length; snout
rounded or very obtusely pointed, strongly projecting, with a
curved transverse groove above, its length less than half that of
head; eye small, supero- -lateral, 5 to 52 times in length of head ;
interorbital w idth 2 2 length of head : width of mouth (with lips)
2.to 4 length of head; lps strongly developed, upper straight-
edged, lower more or less expanded and bordered in front by a
fringe of papille, the posterior border strongly festooned; inner
surface of lips with small papille forming numerous transverse

* Of. P.Z,S. 1919, p. 17.

400 MR. G. A. BOULENGER ON NEW FISHES FROM

plice ; rostral flap large, completely detached on the sides, its
edge feebly festooned ; nuptial tubercles or their scars on ‘the
upper surface of the snout. Dorsal III 10-11, equally distant
from nostrils and from root of caudal, upper edge concave, last
simple ray as long or a little longer than head, Anal III 5,
reaching root of caudal or not. Pectoral as long as or a little
shorter than head, not reaching ventral, the first ray of which

Text-figure 6.

Labeo dhonti.

falls below fourth branched ray of dorsal. Caudal deeply emar-
einate, crescentic when fully spre out. Caudal peduncle 14

times as long as deep. Scales 35—36

Be , 33 between lateral line
and ventral, 12 (exceptionally 14) round caudal peduncle. Dark
olive above, white beneath; a more or less distinct dark laterai
band; sometimes ending in a black spot.

Total length 115 mm.

Six specimens from Tumbwe.

Distinguished from L. cylindricus Peters by a more elongate
body and a lower number of scales round the caudal peduncle.

BARBUS EUCHILUS.

Depth of oe 4 times in total length, length of head 34 times.
Snout rounded, 3 length of head; eye 4 times in length of head,
interorbital width ai times ; north inferior, its sada os ae:
in length of head; both lips much developed, the lower with a
rounded median lobe; two barbels on each side, anterior 2
diameter of eye, posterior 2 2, Dorsal TV 10, equally distant from

eye and from root of caudal: border concave ; ae simple ray not
enlarged, articulate in its distal half, smooth, + length of head.
Anal ITI 5, not reaching caudal. Pectoral 2 length of head, not
reaching ventral; base of latter below middle of dorsal. Caudal
peduncle 13 times as longas deep. Scales longitudinally striated,

45 : P
262, 2 between lateral line and ventral, 12 round caudal

a age ee ee

TILE WEST COAST OF LAKE TANGANYIKA. 40]

peduncle. Brown above, whitish beneath; a large dark brown
spot on the gill-cover.

Text-figure 7.

Barbus euchilus, with lower view of anterior part of head in same (a)
and in B. miochilus (b).

Total length 80 mm.

A single specimen from Tumbwe.

Very closely allied to B. caudovittatus Blgr., from the Ubanghi,
but eye smaller and lips more developed.

BARBUS MIOCHILUS.

Depth of body equal to length of head, 4 times in total length.
Snout rounded, 3 length of head; eye 4 times in length of head,
interorbital width 3 times; mouth inferior, with sharpish edge,
its ‘width 33 times in length of head; lips moderate, lower
restricted to the sides; two barbels on each side, anterior 4
diameter of eye, posterior $ to 3. Dorsal IV 10, equally distant
from centre of eye and from root of caudal; border concave; last
simple ray not enlarged, articulated nearly to the base, smooth,
a little shorter than head. Anal ITI 5, not reaching caudal.
Pectoral ? length of head, not reaching ventral; base of latter
below middle of dorsal. Caudal peduncle 13 to 1} times as long

as deep. Scales longitudinally striated, 25-26 =. 2 between
lateral line and ventral, 12 round caudal pedunele. “Brown above,
whitish beneath; a large dark-brown spot on the gill-cover.
Male with very small round nuptial tubercles on the snout and
larger ones on the lower surface of the head.

Total length 85 mm.

Four specimens from Tumbwe.

Closely resembles the preceding, but distinguished by the
character of the lips and the shape of the mouth,

402 MR. G. A. BOULENGER ON NEW FISHES FROM

ALLABENCHELYS DHONTT.

Dept of body 7 to 8 times in total length, length of head 44
to 5 times. Head 1; to 13 times as long as broad, smooth above,
the bony casque, in Ae middle of the head, 2 to 4 width of head ;
occipital process acutely pointed ; a rather large frontal fontanelle ;
occipital fontanelle small, in advance of Gceipital process ; eye
very small, 3 times in length of snout, 6 to 7 times in interorbital
width, which is 1? to 27 times in length of head; band of pre-
maxillary teeth ri to 5 times as lone as broad; vomerine teeth

Text-figure 8.

Ailabenchelys dhonti.

eranular, forming a crescentic band which is a little narrower
than the premaxillary band: nasal barbel 2 to # length of head ;
maxillary barbel as long as or a little shorter chan head, “be
reaching middle of pectoral fin; outer mandibular barbel 3 to #?
length of head, inner 4 to 2. Gil rakers moderately long, 10 to
12 on anterior arch. ible hidden ae the skin, Dorsal
55-60, its distance from occipital process 4 to 2 length of head.
Anal. 50-53. Both dorsal and anal ee ed separated from
eaudal. Pectoral 2 to 4 length of head, its spine strongly serrated
on outer side and Ceeeils on inner, and 3 to § the length of the
fin.» Ventral 1% to 14 times as ‘distant from root of caudal as
from end of snout. Caudal 2 to 2 length of head. Dark brown
above, whitish beneath.

Total length 165 mm.

10 specimens from Kabeke.

PHRACTURA LUKUG&.
Depth of body 63 times in total length, length of head 5

times. Head 14 times as long as broad, feebly rugose oe er

snout half lemat of head, obtusely pointed, projecting beyond
mouth ; space “between nostrils a little nearer eye than end of
snout; eye very small, on upper surface of head, 8 ‘diameters in
length of head, 2 in interorbital width ; lips and barbels covered
with large round papille beneath; maxillary barbel 3 length of
head, outer mandibular a little shorter, inner mandibular +.

THE WEST COAST OF LAKE TANGANYIKA. 403

Occipital process long and very narrow, not reaching interneural
shield. Dorsal I 6, twice as distant from base of caudal as from
end of snout; second dorsal a little nearer first than root of
eaudal. Anal II7. Pectoral as long as head, longer than ventral ;
latter not reaching anal. Caudal with crescentic notch. 24
dorsal and 19 ventral scutes, the last 5 united on caudal peduncle,
which is 3 of the total length. Pale yellowish above, white
beneath ; three ill-defined darker bars across the body and dark

Text-figure 9.

Phractura lukuge.

annuli on the caudal peduncle; pectorals, ventrals, and anal with
rather indistinct dusky cross-bars; black variegations on each
lobe of the caudal.

Total length 90 mm.

A single specimen from Kabeke.

This species is intermediate between P. bovii Perugia and
P. lindica Blgr., from the Congo.

ANABAS CTENOTIS.

Depth of body equal to length of head, 3 times in total length.
Snout rounded, a little shorter than eye, which is 4 times in

Text-figure 10.

Anabas ctenotis.

length of head and 11 times in interorbital width; maxillary
extending to below anterior fourth of eye; no palatine teeth ; pree-

orbital, preeoperculum, and interoperculum entire ; suboperculum
Proc. Zoot. Soc.—1919, No. XXVIII. 28

404 ON NEW FISHES FROM LAKE TANGANYIKA.

strongly denticulate; 8 to 10 strong, subequal serra above and
5 or 6 below opercular notch. Dorsal XV—XVI8; last spine
longest, 4 length of head; longest soft rays 3 length of head.
Anal X 7-8. Pectoral 3 length of head. Ventral reaching or

nearly reaching anal. Caudal peduncle very short, nearly as long
Ce yWea . om ns WEI
as eye. Scales rugose, strongly ctenoid, 27 ,-; lateral lines 7.

Brown, with small darker spots; fins dark brown, caudal blackish
at the base.

Total length 70 mm.

Two specimens from a ditch near the Lukuga River.

This species is very near A. nanus Gthr., from which it is dis-
tinguished by the denticulation of the gill-cover, the different
anal fin-formula, aud the shorter ventral fins,

ON THE RADULA OF THE MITRID&. 405

26. The Radula of the Mitridee.
By the Rev. A. H. Cooke, Se.D., F.Z.8.

[| Received October 16, 1919: Read November 18, 1919. }

(Text-figures 1-18.)

Fifty years have passed since F. H. Troschel published his great
work on the radule of Mollusca (11), and, during the interval,
the number of Mitride whose radule have been investigated may
be counted on the fingers of one hand. From one cause or another,
Troschel’s work on the group contained some serious errors of
misstatement, which have been copied into the subsequent
Manuals dealing with the Mollusca, e.g., those of P. Fischer (1)
and Tryon (12). Troschel figured, as belonging to Zmbricaria, a
radula closely akin to that of Vaswm, and placed the subgenus
close to the Turbinellide ; he held that Strigatella had a radula
whose form differed widely from that of the typical Mitride, and
was closely allied to that of Turricula (now Vextllum Bolt.), while
the true form of the Vewillum radula was imperfectly described
by him.

The recent acquisition, by the British Museum (Natural
History), of the vast collection of radule formed by the late
Prof. H. M. Gwatkin, enables fresh light to be thrown on this
and other groups, while the results can be checked, in part, by
the collections which he gave, during his lifetime, to Cambridge
and other Universities.

Troschel figured and described 13 species of Mitridee (Cylindra
1, Mitra proper 5, Imbricaria 1, Strigatella 2, Turricula (Veail-
dum) 4: Volutomitra is now more correctly placed with the
Volutide. The Gwatkin Collection contains 14 Veaillam and 37
belonging to other groups, of which 51 only 4 were known to
‘Troschel, and nearly all are new to Science.

In Mitra proper the radula is remarkably broad in proportion
to its length, a feature due to the extreme width of the laterals.
For instance, in adult specimens, we have :—

adusta Lam. Length in mm. 5:0; breadth in mm, 1°5,
cardinalis Gmel. __,, a 5:0 Li * TG.
digitalis Chem. a os 5:0 4 i ete
episcopalis L. a se 70 ss a 2-0.
glabra Swains, 45 - 6:5 st Ks iG;
papalis L. A a 5:0 a 9 1-0;

The number of rows of teeth varies :—

adusta (full-grown) has 76+ naszent rows.

99 (young) a” 64 9? 9)
cardinalis woo és ”
,, (another spec.) ,, 67 2} 4

406 DR. A. H. COOKE ON THE

cucumeriiud has 47-+ nascent rows.
digitalis sy bale 8 3
episcopalis Foes will eS Eee
ferruginea Day est A ay
fulva eee: a =
glabra = LOG . .
papalis ee A ci -

In many genera of rhachiglossate Mollusca, and particularly in
the Thaide and allied groups, the form of the laterals is nearly
constant, and affords little help in subdivision and classification ;
the shape of the rhachidian is the determining factor. In the
Mitride the reverse is the case, for while the rhachidian tooth
varies considerably in shape, the modifications of the laterals are
of great importance, and appear to be of equal or even superior
value in estimating the relationships of allied groups and species.
Thus, Vewxillwm is at once separated from all other forms of Mitra
by its single-hooked laterals.

Similar, though not so wide divergencies in the laterals form a
basis for classification within the various groups. In this respect,
the Mitvride fall into line with the allied genera Musinus, Lascio-
laria, Latirus, and Peristernia, in all of which the laterals, rather
than the rhachidian tooth, supply the best evidence for classifica-
tion. In these four genera, the rhachidian is so much reduced
in size that it has practically lost those characteristics which can
be employed to distinguish one group from another. The
Mitride have not quite reached this point of development, or, as
it might be called, of degradation, but the rhachidian is small in
proportion to the rest of the lingual apparatus.

Group 1. Mirra Martyn.

Rhachidian 6-11-cusped on a squarish* or oblong framework :
laterals long rake-shaped, multicuspid, the interior cusps inclining
slightly inward, cusps 2, 3, or 4 slightly the largest.

Text-figure 1.

Lateral and rhachidian tcoth of Mitra adusta Lam.

Section (a). Mitra proper.—Rhachidian with 8-11 deeply rooted
sharp cusps, the outermost of which lie obscurely behind the
others.

1. I. adusta Lam. Rotuma. Rhachidian 10-cusped, the 6
Inner cusps nearly equal, the 2 outer on each side smaller, and lying
somewhat behind and below the others; base nearly straight,

RADULA OF THE MITRID#. 407

framework of the tooth rather narrow : laterals with 30-32 cusps,
deeply rooted, gradually diminishing to the outside.

Troschel’s figure of the rhachidian is quite wrong.

2. M. cardinalis Gmel. Samoa. Rhachidian 8-cusped, cusps
rather short, the 6 interior about equal, the 2 exterior smaller
and withdrawn ; base very slightly curved: laterals with 13-15
cusps, diminishing to mere serrations outside, inside 6 or 7 deeply
rooted. Another specimen from Torres Strait agrees in essentials.

3. VW. cucumerina Lam. Samoa. Rhachidian 8-cusped, with two
curious projecting side-pieces, cusps nearly equal, deeply rooted;
framework of tooth narrow, base slightly rounded below : laterals
about 15, diminishing outwardly, inside 2 smaller, inclining
inwards.

4. M. digitalis Chem. Samoa. Lhachidian 10-cusped, cusps
rather long, deeply rooted, the 6 interior cusps more or less equal,
4 exterior smaller and projecting sideways; base straight, frame-
work of tooth deeper than in cucwmerina: laterals with 28-30
denticles, gradually diminishing to the outward end of the tooth,
all deeply rooted ; interior 3 or 4 inclining inwards.

Tn another specimen (also from Samoa) the forward end of the
radula tapers away almost to nothing, so that both rhachidian
and laterals are gradually much reduced in size.

5. MW. fulva Swains. Fiji. Rhachidian 8-cusped, cusps rather
long and narrow, deeply rooted, the 4 central the largest, outer
2 on each side more stumpy ; framework of tooth rather narrow,
base straight: laterals with 20-21 cusps, gradually diminishing
right to the end, interior 3 or 4 inclined inwards. 3

6. UW. glabra Swains. Tasmania. Rhachidian 8-cusped, mode-
rately long, but shorter than in fulva, central 6 nearly equal,
the 2 outside shorter and withdrawn; framework narrow, base
straight: laterals with 30-32 cusps, all deeply rooted, gradually
diminishing to the end; framework rather narrow.

7. M. rhodia Reeve. Port Jackson. Rhachidian 5-cusped on
a peculiarly shaped squarish framework, cusps very deeply rooted,
the 3 interior the largest, numbers 2 and 4 set at a shght angle
to the central cusp; base straight, upper margin curved : laterals
with 15-16 cusps, very deeply rooted, not very sharp, gradually
diminishing almost to the end, extreme end bare, inside 3 or 4
cusps inclining inwards ; framework of tooth nearly straight.

8. UW. scutulata Chem. E. Indies. Rhachidian 8-cusped on a
broadish-oblong framework, cusps deeply rooted, somewhat broad
and blunt, projecting well above the upper margin, the 4 central
the longest ; base straight: laterals with 26-28 cusps, gradually
diminishing to mere points, which continue nearly to the end,
about half deeply rooted, cusps 3, 4, 5 (from inside) the largest.

9. M. stephanucha Melv. Karachi. The specimen is badly
mounted and can scarcely be seen, but the radula apparently
belongs to this group. Rhachidian with 3 fairly long cusps, deeply
rooted, these are flanked by 2 or 3 smaller cusps on each side :
laterals many-cusped, cusps rapidly diminishing in size.

408 : DR. A. H. COOKE ON THE

In all the above species the characteristic orange colour of
the rhachidian is distributed all over the framework.

Section (6). Papalaria Dall.—Rhachidian with 7-9 cusps ; cusps
short, triangular, the external set at an angle to those in the
centre.

Text-figure 2.

re Bap

Lateral and rhachidian tooth of Mitra (Papalaria) papalis Lu.

1. I. episcopalis L. Funafuti. Lhachidian 8-9-cusped, cusps
stout and broad, set on an arching curve of the upper margin,
deeply rooted, base nearly straight : laterals with 20-22 cusps,
gradually diminishing to mere serrations at the end, inside 11
deeply rooted, the 3 innermost inclining inwards.

In an immature specimen from S. Pacific (Cambridge) there
are only 5 cusps on the rhachidian, the central much the thickest
and longest. ‘Treschel’s figure is very poor.

2, M. papalis L. S. Pacific. Rhachidian 7-cusped, the external
2 being obscure and withdrawn ; interior 5 nearly equal, central
slightly the largest, all set on a curved thickening of the
upper half of the framework, base slightly curved : laterals with
28-30 cusps, the interior 10-11 the largest, then diminishing to
mere serrations, which continue to the end.

In both these species the 4 central cusps seem to be mounted
on a superposed plate, which is coloured deep orange, the rest of
the framework is colourless or very light yellow.

Section (c) of IW. ferruginea.—Rhachidian cusps of equal size,
narrow, none projecting at the sides or withdrawn behind the
others.

Text-figure 3.

eth BUH

Lateral and rhachidian tooth of Mitra ferruginea Lam.

M. ferruginea Lam. Samoa. Ihachidian 6-cusped, cusps
rather thick and stumpy, mass of the tooth rather deep, the Ist
and 6th cusps descend perpendicularly at the sides, and there are
no obscure side cusps or pieces; base very slightly arched :
laterals 15-17, crowded together, the 5 inside much the largest ;
about half are deeply rooted, the rest diminish into serrations
which continue to the end.

RADULA OF THE MITRID®™. 409

The complete absence of any small cusps on the side of the
rhachidian, which often makes it difficult to count their actual
number, seems to justify the creation of a separate section for
this species.

Group 2. Subgenus Dipapnus Philippi.
p g pp

Rhachidian closely allied to that of Mitra proper, 8—-9-cusped :
laterals multicuspid as in Mitra.

D. edentulus Reeve. Polynesia. Rhachidian with 8-9 deeply
rooted sharpish cusps, projecting well beyond the upper margin,
the 5 central the longest; upper margin slightly curved, base
slightly arched: laterals with 19-21 deeply rooted sharp cusps,
set well apart, gradually diminishing to the outer end of the
framework.

Text-figure 4.

wuesvoooo aaa agp

Lateral and rhachidian tooth of Dibaphus edentulus Reeve.

A specimen from Mauritius has 9 cusps on the rhachidian
and 18-19 on the laterals, another (Cambridge) from Mauritius
has 43+ nascent rows.

Group (3) of coriacea.

M. coriacea Reeve. 8. Pacific. Rhachidian 3-cusped, cusps
thin and sharp, the central 3 times as long as the others, all
projecting well beyond the upper margin, which is much curved,
sloping sharply away from the central cusp ; on the outer sides of
the central mass ave two lateral projections, like broad denticles ;
base slightly curved : daéerals with rather a deep framework, set

Text-figure 5.

Const

Lateral and rhachidian tooth of Mitra coriacea Reeve.

with 10-12 sharp denticles along a straight upper margin, about
half of these are deeply rooted, they continue to the end, leaving
no bare space ; a small prow-like projection stands out from the
interior edge of the lateral, pointing to the similar projection on
the rhachidian.

In this group, of which coriacea is the single known repre-
sentative, the laterals are fairly typical, but the singular form of
the rhachidian separates it off.

410 DR. A. H. COOKE ON THE

Group 4. Subgenus SrricaATELLA Swainson.

Nha a ~ Neen
This group shows :

acuminata 8. 60+ nase. rows of teeth, 6 cusps on rhachidian.
astricta 5O - ei ae *
columbelleformis 75 i 45 Deb evades ss
limbifera 86 Fr a a as 5A
litterata 76--S802865870 5 ue uh i e
luctwosa 81 si ‘3 ‘i ae Me
pellis serpentis 52 Bs a a as iy
planilirata 95 9 3 6 a ne
tristis 74 ‘ss ss 8 m ;

Rhachidian tooth shaped more or less as in Mitra proper,
with 5-8 narrow cusps, sharp or blunt, short or prolonged, the
two external somewhat withdrawn and obscure: lateruls comb-
shaped, the inner half of the framework more or less arched, the
outer half straight, with a marked “break” or division between
the two halves; inner half set with well-marked smallish cusps
(the inner 3 or 4 incline slightly inwards), which become mere
points or fade away altogether on the onter half; extreme end
usually quite bare.

Text-figure 6.

Ca gt Sn

Lateral and rhachidian tooth of Strigatella litterata Lam.

1. S. acuminata Swains. Rotuma. Rhachidian 6-cusped, the
4 central cusps deeply rooted, the 2 outer small, and lying at the
side and somewhat behind; cusps rather thick, projecting well
above the upper margin : laterals with 7-8 well formed, deeply
rooted cusps, these rapidly diminish to mere serrations or points ;
there are about 10 of these, reaching almost to the end; the
“break” not very prominent.

An immature specimen at Cambridge, also from Rotuma,
has 5 cusps on the rhachidian, the central the longest, and fewer
cusps on the laterals.

2. S. astricta Reeve. Maui. Rhachidian 7-cusped, the 2 outer
obscure, lying as in acuminata, central the largest, all deeply
rooted, not projecting far beyond the margin; base scarcely
arched : laterals with 6 inner deuticles, conspicuous, deeply rooted,
then suddenly mere serrations, or the rest quite bare; 4th cusp
from inside the largest ; “break” conspicuous.

3. S. columbelleformis Reeve. Rotuma. LRhachidian squarish,
d-cusped, with an additional obscure cusp or projection at the
sides, cusps very deeply cut and chiselled out of the mass, tips
blunt, scarcely projecting above the margin: laterals with 6-7

aaa

RADULA OF THE MITRIDA. All

strong deeply rooted denticles, then a few serrations or points,
and finally blank, denticles 3 and 4 much the strongest ; “ break”
very pronounced.

4. S. limbifera Lam. Durban. Ihachidian 7-cusped on a
squarish-oblong base, cusps deeply rooted, short and stumpy,
scarcely projecting above the upper margin ; base straight or very
slightly rounded : laterals with 9-10 short ‘thick denticles which
rapidly diminish to mere points, then a long bare space, almost
ole to that occupied by denticles ; ‘‘ break” well marked.

5. S. litterata Lam. Andamans. Ihachidian 7-cusped, cusps
deeply rooted, the 2 exterior very small, and lying behind,
interior 3 equal, projecting considerably beyond the upper margin,
next two rather smaller, all rather thick and stumpy ; base quite
straight : laterals with 8-9 short, stout, deeply rooted denticles,
rapidly diminishing to points or serrations, end bare; ‘“ break ”’
sharply marked.

Specimens from Hilo, Hawaii, and from Scottburgh, Natal,
correspond in essentials, but have rather more (11— 13) denticles
on the laterals. One from Isipingo exhibits the 2 obscure exterior
cusps on the rhachidian much more plainly, while the 5 interior
are of about equal length.

6. S. luctwosa A. Ad. Durban. This might well be a specimen
of litterata, except that the cusps, both in rhachidian and laterals,
are sharper, and further apart; there are 7-8 cusps only in the
laterals, and beyond them the bare space is very long.

Se pellis serpentis Reeve. Mauritius. Radula small; rha-
chidian 7-cusped, cusps short, rather blunt, deeply rooted,
projecting beyond the upper margin, central the thickest and
longest ; base straight: laterals wily large cusps, rather sharply
meinen ve some distance a part, then a Weae space to the end, the
first two only incline slightly inwards ; ‘‘ break” not very strongly
marked.

This species, in the shape of the lateral cusps, stands rather
apart from the others.

8. 8. planilirata Reeve. Suez. Rhachidiain 6-cusped, cusps
somewhat sharp and narrow, deeply rooted, tips not projecting
far beyond the margin; base straight : laterals with 7—8 deeply
rooted cusps, rather near together, gradually diminishing to
ae bare space quite considerable; “break” not very marked.

9. S. tristis Sowb. Panama. Rhachidian with the upper
margin produced at the angles into two curious horns, cusps 8,
deeply rooted, the 2 exterior somal visible, central 4 the largest,
tips scarcely ‘projecting beyond the upper margin; base wavy-
straight : laterals with 6-7 strong blunt cusps, very deeply rooted
on the arched portion, then rapidly diminishing to points, which
continue right to the end, no bare part.

One would expect this species, the only Neotropical repre-
sentative of the subgenus whose radula is known, to exhibit
difterences in structure from the rest, and it does so.

412 DR. A. H. COOKE ON THE

Group 5. Subgenus Impricaria Schumacher.

- Rhachidian with 2 large and 2-3 small cusps: laterals with
one or more cusps considerably developed, remainder either
numerous or few and degraded.

Text-figure 7,

Lateral and rhachidian tooth of Imbricaria marmorata Swains.

1. J. marmorata Swains. Upolu. Rhachidian 5-cusped, all
sharp and projecting beyond the upper margin, the central small,
flanked on each side by a very large cusp, outside these one small
cusp, with sometimes a trace of a second ; all these are set upon
a thick plate, the base of which is somewhat angled in the middle ;
laterals with 12-14 cusps, the third from the inside much the
largest, inside this are 1 or 2 small denticles, all these incline
slightly inwards ; outside the big cusps are 9-11 upright, sharp,
narrow denticles, diminishing to mere points which continue
to the end of the framework; in the central portion of the
lower side of the framework is a roughly triangular “ false keel” ;
the framework itself narrows towards both ends, becoming
almost pointed on the inner side.

Text-figure 8.

Lateral and rhachidian tooth of Imbricaria ossea Yease.

Text-figure 9.

Lateral and rhachidian tooth of Imbricaria oliveformis Sowb.

2. J. ossea Pease. Samoa. Lhachidian with a roughly tri-
angular framework, nearly all of which is occupied by 2 long
prominent sharp cusps, whose roots are carried through to the
base, these are flanked on each side by 2-3 wrinkles rather than
cusps, their points not projecting above the slope of the upper
margin: laterals with the second cusp enormously developed,

RADULA OF THE MITRID®. ALI 3

pointing inwards, at its inner base is one sharp almost adherent
denticle, outside 4-5 very small denticles, which diminish to
mere points, all inclining sharply inward, but for the big denticle,
the appearance of the tooth is degraded; ‘false keel” well
marked and deep.

In a more mature specimen from Society Is., the flanking
cusps on the rhachidian are better marked ; in the laterals there
are only 2-3 denticles outside the big one, close together on one
side, further apart on the other; the framework is sharply
pointed towards the rhachidian, bluntly pointed on the outside.

3. I. oliveformis Sowb. 8. Pacific. Rhachidian with two
large, sharply pointed cusps, projecting well beyond a curving
upper margin, flanked on each side by a single small denticle
standing out against the mass of the tooth, its apex not reaching
the curve of the upper margin, sides of the tooth continue this
curve, base nearly straight : laterals quite bare, save for one large
cusp, inclining inwards, no trace of further denticulations ; upper
margin straight, lower curved, framework pointed towards the
vhachidian ; general appearance very degraded.

Text-figure 10.

an Hay

Lateral and rhachidian tooth of Imbricaria punctata Swains.

4. I. punctata Swains. W. Maui. Lhachidian 4-cusped, flanked
by wrinkles, the 2 large inner cusps deeply rooted, with no
intervening central cusp; the 2 side cusps larger than in mar-
morata ; the superposed plate from which the cusps spring 1s
extended angularly at the sides: laterals 5—6-cusped, the inside
2 small, then 2 large ones, then one or two much smaller, di-
minishing rapidly, all cusps inclining sharply inwards; a rather
shallow but well marked ‘false keel”; general appearance
degraded.

This species is often regarded as identical with ossea Reeve,
and the radula is not against this view.

Group (6) of fammigera Reeve.

Rhachidian 7—8-cusped on a narrow framework: laterals multi-
cuspid, cusps about 11-15, one or two of exaggerated size; frame-
work narrowed prow-like inside, produced below into a “false
keel.”

1. Jf. flammigera Reeve. Durban. Rhachidian 8-9-cusped,
cusps narrow and sharp, set on the margin of a very narrow frame-
work, the 5 interior alternately larger and shorter, externals mere

414 DR. A. H. COOKE ON THE

serrations; base straight: laterals with one of the denticles (in
place of the 3rd or 4th) immensely exaggerated in size, and in-
clining inward, forming a separation between the two parts of the
tooth; the inner part is narrow, produced like the prow of a
boat, and carries 3-4 fine needle-like denticles or serve close to

Text-figure 11.

een

Lateral and rhachidian tooth of Mitra flammigera Reeve.

the big denticle, while the tip is bare; on the outer part small
denticles (about 9) mount up towards the big denticle; they are
set apart from one another, gradually diminishing and leaving
the external portion of the margin quite bare.

In another specimen (also from Durban), the rhachidian is
smaller and narrower, and has a rather thicker framework; the
cusps ave very irregular, 2, 3, or 4 in number, of various sizes
and dispositions in different rows: the laterals are very faintly
serrated just inside the big denticle, which, on one side only of
the whole radula, is bifurcated (and in some cases trifurcated)
like a lobster’s claw.

2. M. interlirata Reeve. Durban. Rhachidian -cusped,
cusps shaped as in fammigera, the 3 interior much the largest, the
central the shortest of the three; base narrow, nearly straight ;
upper margin slightly curved: in the laterals the big denticle is
rather longer than in flammigera; there are 4-5 minute denticles
or serre on the interior part of the margin ; on the exterior side
of the big denticle are 7-8 small denticles, end of framework
bare*.

Text-figure 12.

Conan ni Yan

Lateral and rhachidian tooth of Mitra pretiosa Reeve, Durban.

3. JJ. pretiosa Reeve. Suez. LRhachidian 7-8-cusped, cusps
sharp and projecting far beyond the upper margin, the 4 interior
much the largest, all deeply rooted; base straight: laterals with

* Since this description was written, Mr. Burnup, of Durban, who sent the speci-
mens to Prof. Gwatkin, has assured me that the two species flammigera and
interlivata are conchologically identical.

RADULA OF THE MITRID. 415

one cusp, the 4th from inside, much the largest, next inside one
rather large, 11-12 in all, rapidly diminishing on both sides of
the big cusps, the exterior mere points.

Another more mature specimen from Durban corresponds,
except that both the 4th and dth lateral cusps are very large.

Group (7) of spherulata.

rhachidian thick, deeply coloured with orange and brown,
bicuspid; laterals degraded, cusps few and obscure.

AL.“ cireulata Kien. var.” Durban (Mr. H. C. Burnup*),.
Rhachidian thick, square, deep orange colour, opaque, with 2
prominent deeply-rooted cusps, the tips of which are blunt, and
do not project far beyond the upper margin, these are flanked
on each side by two obscure denticles or wrinkles: daterals thin,
framework rather deep, with 4-5 deeply rooted denticles, 2-3 of
which incline inward, half the margin quite bare. There are
73+ nascent rows.

Text-figure 13.

Lateral and rhachidian tooth of Wtra spherulata Mart.

There is evidently some confusion in the specimens forwarded
by Mr. Burnup, since those of “ ctrcewlata” exhibit quite a different
type of radula. Another specimen of ‘ cirewlata var.” in my own
collection corresponds exactly with that here described.

2. WM. crenifera Lam. Mauritius. Rhachidian squavish-oblong,
colour deep orange-brown, almost black, 6-cusped, the 2 central
strong, blunt and projecting, the side cusps smaller but distinct ;
base somewhat arched: laterals with 4-5 stumpy denticles, all
inclining inward, rest of margin blank.

3. M. spherulata Mart. §. Pacific. Rhachidian lozenge-
shaped, with 2 large deeply rooted cusps mounted on a flat plate,
which is superposed on the main framework, tips broad, project-
ing well above the upper margin; these are flanked on each side
by two small obscure cusps, set in the side of the tooth and some-
what behind; sides of tooth slightly produced with small blunt
wings; base slightly arched: laterals consisting of an obscure,
rather ‘deep- -oblong plate, 3-cusped (some trace of a 4th) on inside
part of the inner margin, rest blank. Rows 63+ nascent.

In a specimen com Upolu (Coll. Cambridge) and in one from
Uvea (coll. A. H. C.) there are 4 lateral cusps.

* This form, hitherto confused with MW. circula (not eirculata) Kien., will be
described as JL. burnwpiana. Mr. Burnup agrees that it is distinci.

416 DR. A. H. COOKE ON THE

Group (8) of variegata.

Rhachidian single-cusped on a triangular plate: laterals de-
graded, with one large cusp only.

M. variegata Reeve. Mauritius. This unique and most in-
teresting form shows a rhachidian formed of a simple triangular
plate, with 2 angles at the extremes of the upper margin, and the
3rd in the centre of the base; in the middle of the upper margin
is a single sharp, deeply rooted cusp, the tip of which projects well
beyond the margin, no trace of any other cusp: Jaterals in general
shape like znterlirata, with a single strong denticle of exaggerated
size, inclining inwards, no trace of any further denticles; the
projecting “‘ keel” much larger than in znéerlirata.

Text-figure 14.

Lateral and rhachidian tooth of Mitra variegata Reeve.

*“ The whole radula is rather thin and lacking in substance ;
while the merging of all the cusps in the laterals in a single
strong denticle is very suggestive of the lines on which radula

<levelopment may proceed.

Group (9) of erumnosa.

Radula small, rhachidian and laterals set with numerous small
sharp cusps; framework of teeth narrow, nearly straight.

1. WZ, erumnosa Melv. 8S. Africa. Khachidian 8-cusped, the
6 interior equal in length, the 2 exterior much smaller; cusps
sharp, small, none deeply rooted; base slightly arched: laterals
with 12-14 cusps, the 2 or 3 interior inclining slightly, but all in
the upper margin gradually diminishing in size.

2. M. bovei Kien. Suez. Rhachidian 8-9-cusped, cusps nearly
equal, not deeply rooted ; base very slightly arched: laterals with
12 nearly equal cusps, with 2 or 3 serrations at the outer end.

Text-figure 15.
Lateral and rhachidian tooth of Mitra bovei Kien.
3. M. filosa Born. Mauritius. Rhachidian 8-10-cusped, cusps

very small, framework narrow, upper margin slightly curved; the
cusps are inclined to become irregular in different teeth, and

RADULA OF THE MITRIDA. 417

sometimes two coalesce into one: laterals set with 20-24 tiny
sharp denticles on a slightly curved framework, not greatly differ-
Ing in size except at the extreme end, where they become very
small.

4. M. proscissa Reeve. Karachi. Lhachidian 11—-12-cusped,
cusps tiny, sharp, close together, equal in size, upper margin
slightly curved, framework very narrow: laterals with 13-14
cusps, the 2 or 3 innermost rather short and stumpy, the rest
larger and sharp, none inclining inward ; they gradually diminish
to the extreme end.

5. M. pura A. Ad. China. Rhachidian 6-cusped, the 2 ex-
terior below the plane of the rest; margin straight, base slightly
rounded: laterals with 12-13 sharp denticles, set rather apart
from one another, gradually diminishing to the end.

This and the following group probably represent separate
branches from the parent stem.

Group (10) of scabriuscula.

Radula with rhachidian and laterals set with numerous small
sharp cusps; framework of rhachidian narrow, deeply arched,
tips curving upward ; framework of laterals narrow, arched, then
straight.

1. MW. circula Kien. Durban. hachidian and laterals set
with very numerous tiny denticles, all about the same size. The
specimen is in bad condition, preventing the denticles from being

counted,
Text-figure 16,

ee oe

Lateral and rhachidian tooth of Mitra scabriuscula Li.

2. M. scabriuscula L. Upolu. Rhachidian with 30-32 tiny
denticles, not deeply rooted, gradually diminishing towards the
ends; framework narrow at tips, deeply curved, tips bare and turn-
ing slightly upward : laterals with about 32 similar denticles set
closely together, interior end bare of denticles, they continue to
the exterior end, gradually diminishing in both directions.

The form of the rhachidian in this group, corresponding closely
as it does with that of Zurricula, forms an interesting link
between the radula of that genus and that of Mi itra, While the
laterals, on the other hand, resemble in miniature those of Mitra

proper,

Genus Vexinium Bolten (olim Turricula [Klein] auctt.).

As pointed out long ago by Gray (3, 4), the laterals in Vexillum,
instead of being broad rand comb- shaped, with many sharp denticles,

418 _ DR. A. H. COOKE ON THE

consist of a single curved blade, as in Oliva, Murex, Trophon,
Thais, and the allied groups. As a rule the base is narrower,
and the blade thinner than in these genera. The rhachedian
tooth is bow-shaped, framework narrow, pointed at the ends, set
either with many small sharp cusps, which cover nearly the whole
of the upper margin, or with three only, closely grouped together
in the centre. The whole radula is as a rule extremely small.

Group (1). Rhachidian set with many small sharp cusps, tips
of framework curving slightly up, and pointed at the ends, which
are bare of cusps.

1. V. alauda Quoy. Mauritius. Framework of rhachidian
moderately curved, set with 16-17 minute denticles, diminishing
outwardly, ends bare, scarcely turned up: laterals simply hooked,
not deeply curved.

2. V. arenosum Lam. Philippines. Framework of rhachidian
deeply arched in bow shape, set with 20-22 sharp denticles, not
deeply set, which diminish gradually im size to mere points, and
finally leave the ends bare, central denticle the largest; ends
sharply turned up: laterals simple hooks, not deeply curved.

3. V. cruentatum Chem, Philippines. Rhachidian not very
deeply waved; denticles 19-20, closely packed and narrow,
central the largest; a small blank space at the ends of the frame-
work: laterals with very narrow blades, slightly hooked at the
tip..

Text-figure 17.

Lateral and rhachidian tooth of Vexillwm exasperatwm Gmel.

4. V.exasperatum Gmel. Mauritius. Rhachidian deeply curved
in bow shape, ends of framework narrow and bare; set with
15-16 sharp narrow cusps, equal in size, except those at the
extreme ends, which become suddenly smaller: laterals long and
narrow, hooked at tip.

5. V. lyratum Lam. Mauritius. Framework of rhachidian
deeply bow-shaped, ends scarcely turning up, tips bare; denticles
13-15, sharp, thin, rather distant from one another, central
rather the largest: laterals long, narrow, very sharply pointed.

6. V. nodosum Swains. Hilo. Rhachidian bow-curved, but
not very strongly, tips bare; denticles 7-8, sharp, triangular,
close together: blade of laterals much curved.

7. V. tasmanicum T.-Woods. N. Tasmania. Radula very
small; rhachidian bow-shaped, well curved in centre, tips of
framework bare and turned upward; denticles 10-11, sharp,
close together, equal-sized, but diminishing towards the ends:
laterals narrow-bladed, curving sharply in the centre.

8. V. teresie T.-Woods. Tasmania. Framework of rhachidian

RADULA OF THE MITRID. 419

not deeply bowed, tips of framework bare; denticles 10-11,
central slightly the largest: blade of laterals narrow, somewhat
angled in the middle.

Group (2). Rhachidian tricuspid, cusps small, placed close
together in the centre of the curve, all the rest of the framework
bare of cusps.

Text-figure 18.

Lateral and rhachidian tooth of Vewillwm australe Swains.

1. V. australe Swains. N. Tasmania. The specimen is in bad
condition, but enough can be seen to testify that the rhachidian
is tricuspid, cusps somewhat large, central slightly the smallest :
laterals thickish below, narrowing sharply at the ends.

2. V. ebenus Lam. Malta. Framework of rhachidian narrow,
deeply arched, but scarcely waved in bow shape; cusps very close
together, thin, sharp, equal in length, central slightly the
narrowest: blade of laterals narrow, slightly hooked at tip.

3. -hizenense Pils. Japan. Framework of rhachidian deeply
arched in bow shape; denticles smaller and somewhat further
apart than usual im this group, equal in size: blade of laterals
long and rather narrow.

4. V. porphyreticum Reeve. Tonga. Lhachidian with frame-
work deeply waved, cusps small, equal-sized, close together :
blade of laterals deeply narrow, angularly curved in centre. Rows
76+ nascent.

A general survey of the Mitridan radule suggests several con-
siderations.

In the first place, the rhachidian tooth exhibits wide differences
of structure, ranging from the lozenge-shaped 8—9-cusped form in
Mitra, to the unicuspid triangular form of the variegata group.
It is probable that the investigation of further material may
discover links between forms of Mitridan rhachidians at present
very dissimilar. But there appears to be warrant for the sug-
gestion that the Mitride represent an ancient group of Mollusea*,
so that many links in the chain of development may have become
extinct.

Secondly, these divergencies in the structure of the rhachidian
are accompanied by a general similarity of plan in the laterals,
subject however to a progressive modification in their form.
We can arrange the groups and sometimes the species in a

* Mr. R. B. Newton informs me that Bellardi enumerates nearly 200 species of
Mitridee from the Upper Tertiaries of Italy alone, and that the genus dates back to
Cretaceous rocks.

Proc. Zoou, Soc.—1919, No. XXTX. 29

420 DR. A. H. COOKE ON THE

group (cf. Jmbricaria), in a sevies indicating a transition from a
more complex to a simpler form vf tooth. The laterals, in fact,
exhibit every symptom of regress towards a gradual degradation.
At the head of the series we have the multicuspid lateral
of iitra proper, Papalaria, and Dibaphus, in which the 2nd,
ord, and 4th cusps are shghtly larger than the rest, and incline
slightly inwards. The first important modification occurs in
Strigatella, where, on the outer half of the lateral, the cusps
either diminish to mere dots, or are absent altogether, leaving the
framework bare. In Jmbricaria (a parallel development) one or
more of the interior cusps increases considerably in size, at the
expense of the others, which generally become smaller, or are
reduced in numbers and become degraded.

In the fammuigera and spherulata groups further modification
occurs. As a rule, one of the lateral cusps is enormously
developed, with a corresponding decrease in size, or with the
total disappearance, of the remainder of the cusps, the whole
tooth assuming a degraded appearance. In the variegata group
the big lateral cusp has ousted all the others, and the tooth is
mucno degraded.

According to Troschel, Cylindromitra (formerly Cylindra)
nucea Meusch. has no laterals, while specimens of Cy. dactylus L.
have been examined both by himself and Dr. Gray without any
trace of a radula being discovered. It is possible therefore that
Cylindromitra forms the last term in the series of degraded forms
of Mitridan radula; on the other hand, it may be wiser to wait
for further evidence before a final decision ean be reached.
Troschel’s figures can seldom be accepted without confirmation.
He places the Ziervogelia group of Strigatella close to Vexillum.
If the radule he figures are correct, Ziervogelia cannot be regarded
as akin to Strigatella; but here again further evidence is needed
before the true position of Ziervogelia can be settled.

It is tempting to pursue this branch of the subject a little
further, and to enquire whether, in the progressive degradation
of the Mitridan lateral, we can obtain a clue to the genesis of
the familiar bicuspid or unicuspid lateral of many of the Rhachi-
glossa. It seems within the bounds of possibility that the
coalescing, or gradual disappearance, of the cusps, in a multicuspid
lateral, produced, in more cases than that of the Mitride, a lateral
with one or two large cusps instead of many small ones*. In
Buccinum, Neptunea, "Obra a. Huthria, and the allied genera,
small cusps occur, with some degree of irregularity, between the
two, or three, great cusps. It is conceivable that these smaller
cusps may be vestiges of an armature, in which the cusps were
more numerous and more on a level as regards size. Even a
genus like Alectrion, which has settled down, in most species, to
a steady bicuspid lateral, occasionally exhibits more than traces

* Compare B. B. Woodward, Proc. Malac. Soc. London, vil. p. 2 “In the

Radula . . . there is consistent progress in the shape of the RET of nume-
rous, weak, little teeth by few strong ones, especially in the carnivorous groups.”

RADULA OF THE MITRIDA. 421

of extra intervening cusps. In most cases of bicuspid laterals,
one or other of the cusps, usually but not always the interior, is
decidedly the smaller of the two. This smaller cusp may show
signs of incipient disappearance, a stage which receives illustra-
tion from Troschel, pl. vil. figs. 15, 19, where the inner eas) of
a Hemifusus and a Iyr istica is very small, and pl. viii. figs. 5, 6,
where the outer cusp of a Vaswm is similarly reduced, Tt is no
-great step from laterals of this type to forms in which the second
eusp has vanished altogether. The framework of the tooth
would diminish in breadth, the fewer cusps it had to carry.

This process of redaction in the rhachiglossate lateral seems to
receive illustration from the case of the Volutide. Most species
of Voluta have lost their laterals altogether. Voluta concinna
and Volutilithes abyssicolt alone retain “them in a very degraded
form, unicuspid on an oblong base. A further stage is shown in
Neptuneopsis gilchristt, where the lateral cusp has ‘vanished, and
the framework is reduced to a small thin plate, of undefined shape,
“probably quite functionless” (Pace 7, Woodward, M. F. 14).

In other genera the process of degradation takes another
line. The laterals remain unimpaired, while the rhachidian
suffers degradation, or even disappears altogether. Fasciolaria,
Latirus, Perister nia, and Fusinus form a group of genera in which
the rhachidian i is reduced to extreme smallness, w ale the rhachi-
dian cusps have in many species almost vanished. Tt cannot be
doubted that the rhachidians of Columbella and of Liomesus
(= Buccinopsis), which are now.mere narrow blocks, are descendants
of teeth which once were furnished with cusps.

Bibliography.

1. Fiscuer, P.—Manuel de Conchyliologie. Paris, 1887.

2. Garretrr, A.—‘‘ Catalogue of the Polynesian Mitride, with
remarks on their .... station.” Journ. of Conchol. iii.
pp. 1-73, 1880.

3. Gray, J. E.—‘“ On the division of Ctenobranchous Gastero-
podous Mollusea into larger Groups and Families.” Ann.
Mag. Nat. Hist. ser. 2, x1. pp. 124-133, 1853.

4. Gray, J. E.—“‘ On the Teeth of the genus Aftra, Lamarck.”
Ann. Mag. Nat. Hist. ser. 2, xu. pp. 129-130, 1853.

5. Hoge, J.—‘:The Lingual Membrane in Mollusca, and its
value in classification.” Trans. Roy. Micros. Soc. xvi.
pp. 93-104, pl. x. f. 37, 1868 htra fusca Swains.).

6. Macponaup, Dr.—Letter in Trans. Roy. Soc. Mauritius, n. s.
v. p. 109, 1871 (Muuritia barclayi H. Ad.).

7. Pace, S.—‘‘On the Anatomy and Relationships of Volta
musica Linn., with Notes upon certain other supposed
members of the Volutide.” Proc. Malac. Soc. London,
v. pp. 21-31, 1902.

8. Sars, G. O.—Mollusea Regionis Arcticee Norvegice. Christi-
ania, 1878.

zoe

422 ON THE RADULA OF THE MITRID2.

9. ScHacko, G.—“ Radula Untersuchungen.” Conchol. Mittheil.
i. (ii.) pp. 122-128, pl. xxiv. f. 5, 1881.

10. Turin, J.—‘‘ Die beschalten Gastropoden der deutschen
Tiefsee-Expedition 1898-99. B. Anatomisch-systema-
tische Untersuchungen..... ” Wissen. Ergebn. deutsch.
Tiefsee-Hxped..... Chun, vil. pl. ix. f. 63.

11. TroscHet, F. H.—Das Gebiss der Schnecken. Berlin:
Vol. ii. part 2 (1868) contains Cylindra, Mitra proper, and
Imbricaria: Vol.i1. part 3 (1869), Strigatella and Turricula.

12. Tryon, G. W.—Manual of Conchology. Vol. iv., 1882.
Philadelphia.

13. Vayssthre, A.—‘ Etude zoologique et anatomique de la
Mitra zonata Marryatt.” Journ. Conchyl. xlix. pp. 77-95,
1901.

14. Woopwarp, M. F.—*“‘ Note on the Anatomy of Voluta ancilla
(Sol.), Veptuneopsis gilchristi Sby.and Volutilithes abyssicola
(Ad. & Rve.).” Proc. Malac. Soc. London, iv. pp. 117—
1255=plisc LIOL:

THE RIGHTING REACTION IN ASTERINA GIBBOSA. 423

27. Note on the Righting Reaction in Asterina gibbosa Penn.
By HE. 8. Russevy, M:A., B.Se., F.Z.8.

Received October 22, 1919 : Read November 18, 1919. ]

Introductory.—The experiments to be described in this paper
were carried out in June 1919 on the beach of Porthmear, a
little rocky bay in North Cornwall between Trevose Head and
Newquay. <A small colony of Asterina lived in the rock-pools
of this bay about halfway up the beach. They did not extend
further to seaward, and the reason for this appeared to be that
their habitat midway up the narrow iand-locked cove was less
exposed to wave-action than the lower section of the beach,
which at low water ran continuous with the coastal line of cliffs.

The method of experimentation was simple—the starfish were
collected in twos and threes, generally from the underside of large
stones, and transferred at once to shallow rock-pools, in which
they could be closely observed. No specimen was used for more
than ten experiments, so as to eliminate the factor of fatigue.

I attach some importance to the fact that the observations
were made under natural conditions, for it is obvious that if one
removes animals from their natural surroundings to the artificial
conditions of a laboratory experiment their behaviour is apt to
be upset by the change. Few animals seem to take kindly, for
instance, to life in a smooth glass basin exposed on all sides to
the light, and it is improbable that their behaviour remains
unaffected by such strange surroundings. Furthermore, there is
always the risk of aquarium specimens “losing vigour and getting
out of condition. Plessner (1913) says of Asterius rubens and
Solaster papposus weakened by aquarium life that they right
pcmsrres only with difficulty or even fail to turn.

It is best, then, when studying the behaviour of any animal to
study it in its native haunts or in surroundings approximating as
nearly as possible to those natural to it. This was comparatively
easy to manage with the Asterinas of Porthmear, and the obser-
vations recorded below, scanty though they are, have at least
the merit of referring to animals see normally in a normal
environment.

The Righting Reaction. —Records were kept of the righting
reaction in eleven individuals of various sizes, the exact way in
which the righting movements were performed being noted, as
well as the time taken, from the moment the starfish was placed
on its back to the moment when it recovered its normal position
and started to crawl away. The rapidity of the reaction in many
cases was striking, nearly 40 per cent. of the turnings observed
being completed in 20-30 seconds. These times compare favour-
ably with the times taken by Ophiura brevispina, which accord-
ing to Glaser (1907) averages 45 seconds for the turn. On the
other hand, if things went wrong and the starfish got tied up in

494 MR. E. 8S. RUSSELL ON THE RIGHTING

the peculiar way to be described later, the complete turn might
take as long as 10 minutes.

The detailed records are as follows :—
Specimen A: ca. 25mm. 13/6/19.
Method of turning.

f
Time taken.
40 secs. Turned on two adjacent arms.
3d 39 39 39 2? ’
24. 3 ” ” 29 |
30; Four arms attached, two outer gave up, turned on two. .
Bone 5 Three attached, one retracted tube-feet, turned on two adjacent arms. :
5) gy Turned on two. Interfered with by a shrimp. .
WHO op Turned mainly on one arm which bent back. Later on an adjacent .
arm rolled longitudinally and attached.
40 ,, Turned on adjacent pair, but first tried to fix with arm opposite the

pair on which it finally turned.

Specimen B: ca. 30mm. 13/6/19.

20 secs. Turned on adjacent pair.

30, Four attached, outermost gave up, turned on two.

20a Three took part, turned on adjacent pair.

BO op Turned on adjacent pair.

25s; ” 22

30 ” 29 op)

BE 55 Three attached, one loosed hold, turned on adjacent two.

BE} Sy Three most of the time, towards the end an outer arm gave up.
30 ,, Turned on two adjacent.

32, ” ”

Average time 27:4 secs.—a clever and vigorous specimen.

Specimen C: ca. 30mm. 14/6/19.

67 secs. First on three, then on adjacent pair.
BO) 35 Turned on adjacent two.
40 ,, First on three, then on adjacent pair.
35 3) ” y)
42 3” 4 2”? " 29 9 5
25 5; Four attached slightly, turned on adjacent pair.
2D yy Three ” 9 oy)
23 3) Four 39 29 3)
32) 33 Three 3 3 c
28 Turned on adjacent pair.
39 5}

Average time 34°7 sees.

Specimen D: ca. 25mm. 14/6/19.

30 secs. Turned on adjacent pair.

35 Three attached, turned on adjacent pair.

27 ” y) oy)
30 2 9 5B)

310) ep Four 4 3

SON ss Three ,, 45

BD ap Turned on adjacent pair, but second choice of pair.
58, First four, then three, turned on adjacent pair. :
32s, Turned on adjacent pair.

25) 5 y) ”

Average time 33°2 secs. This specimen seemed slow at bringing
over fifth arm.

REACTION IN ASTERINA GIBBOSA. 425

Specomen Hi: ca.25mm. Rather short raysand big body. 14/6/19.

secs.

Turned on adjacent pair.

Three at first, one gave up early, turned on adjacent pair.
Three, then four, turned on adjacent pair.

Five attached,
Three attached, a5 "

Turned on adjacent pair.

Three at first, then turned on adjacent pair.

39 bP]

33 33 39
Principally on one of an adjacent pair.
Three for a very short time, turned on adjacent pair.

Average time 31-4 secs.

Specumen BF:

5 secs.

ca. 32 mm. 16/6/19.

Three attached and transitory deadlock occurred, then a fourth
attached and the opposite outermost let go. ‘Turned on three
adjacent.

Three, then five, turned on two adjacent.

Turned on two, but second choice. Interfered with by surface-film
which caused exploratory movements of tube-feet.

Transferred to deeper part of pool :—

5 secs.

Turned on two adjacent.

Three attached, turned on two adjacent.

Four attached, then turned on two adjacent.
Four, then three, then turned on two adjacent.

39 3” 3)
Two, then three, then four, turned on adjacent pair.

39 33 33 23

This specimen showed a distinct tendency to attach all arms.

Specimen G: ca. 14mm. 16/6/19.

5 secs.

Two, then three, turned on adjacent pair.
Turned on adjacent pair.

29 3
a9 by)
32 39

25 ”
Three, then turned on adjacent pair.
Turned on adjacent pair.
One attached, then turned on adjacent pair.

23 92 22

Average time 28°3 sees. A small and very active specimen.

Specimen H: ca. 22mm. 16/6/19.

Specimen J :

72 secs.

60
52

Secs.

Three, then four, then turned on adjacent pair.
Four, then turned on adjacent pair.

Deadlock J. (see below, p. 429).

Two, three, turned on adjacent pair.

Two, three, four, then turned on adjacent pair.
Deadlock II. (see below, p. 429).

Deadlock IIT. (see below, p. 429).

Two, four, three, then turned on adjacent pair.
Three for very short time, turned on adjacent pair.
Two, three, turned on adjacent pair.

ca. 31 mm. 18/6/19.

Two, three, four, three, finally turned on adjacent pair.
Four, three, turned on adjacent pair.
Three for very short time, turned on adjacent pair.

426 MR. E. 8S. RUSSELL ON THE RIGHTING
67 secs. One took the lead, other two attached, one of these gave up, turned
on adjacent pair.
225, Deadlock IV. (see below, p. 429).
80 ., Two adjacent attached, tips curling in same direction, joined by a
' third, one of original pair gave up, turved on adjacent pair (second
choice).
80 ,, Turned on two adjacent, delayed by one opposite.
OQ Turned on one, delayed by an opposite arm holding on.
120) 5 delayed by an opposite pair holding on.
S20. Four attached, turned on adjacent pair.

Specimen K: ca. 28mm. 18/6/19.

55 secs. One, three, turned on adjacent pair, hindered by opposite arm.

A One, two, turned on adjacent pair.

25s, Turned on adjacent pair.

25, One, two, turned on adjacent pair.

50) One, tivo, three, four, turned on second pair (adjacent).

42 ,, Turned on adjacent pair, impeded a little by opposite pair.

BA 5, First on one, then superseded by opposite pair, on which it turned.

BO) on At first one arm turned right back, then opposite pair took hold and
superseded first, which retracted tube-feet.

60 ,, One arm, superseded by opposite pair. Unifying impulse noticeable.

Specimen L: ca. 13 mm. 18/6/19.

125 secs. One, two, turned on adjacent pair, long period of incoordination.
SOM Turned on adjacent pair.
5d; ) >
GOmee es es impeded by opposite arm.
30 Turned on adjacent pair.
SB» gp First one attached, then turned on adjacent pair.
52 22 2 2) ?
52 bed 32 39 bP]
GD op Delayed reaction. Turned on two, impeded by opposite arm.
Slee Turned on adjacent pair.

Norr.—Specimen J seemed to be atfected by the hot sun, so after the first four
trials it was kept in the shade of a stone, as were also specimens K and L.

From these records it is apparent that while there is great
variation in the way the reaction is commenced, there is great
uniformity in the way in which the turning is finally accom-
plished—almost all turn on an adjacent pair of arms, the actual
percentage in the 107 trials recorded being 92°5. In four trials
the turning took place on one arm which bent back from the tip
progressively : turning took place on three adjacent arms in four
cases, including three of the ‘‘deadiock” turns, and in two other
cases (counted as turnings on an adjacent pair) three arms were
engaged until a late stage in the righting, when an outer one
gave up.

When placed on its back Asterina immediately flattens out,
then curls the tips of the arms downwards, raising itself a little.
The tube-feet are extended and feel about actively. Any number
of the arms may attach themselves to the bottom by thei ter-
minal tube-feet. Of these arms two or three generally get a lead
over the others. The simplest case is when two adjacent arms
get ahead and keep ahead of the rest. Provided the ventral
surfaces of these two arms both turn inwards towards one another

REACTION IN ASTERINA GIBBOSA. 427

the righting is carried out with great precision and rapidity,
often in 30 secs. or less. The two arms on which the starfish
rotates are at the end of the turn disposed at a wide angle with
one another, a condition recalling the usual method of righting in
Ophiuroids, where the animal pushes itself over on two arms
stretched out nearly in a straight line. The wide angle is
immediately compensated for when the starfish begins to crawl
away.

If, however, the two arms turn their ventral surfaces in one
direction, so that the tips get a twist in either a clockwise or a
counter-clockwise direction, the turning is delayed and other arms
must be brought into play. In general, a disposition of the arms
with the tips pointing in the same circular direction is a very
unfavourable condition for turning, and much adjustment is
required before turning can be carried out.

The turning on coo. arms is very often delayed by the holding
on of one or two other arms, which later detach themselves.
The “unified impulse” of which Jennings (1907) speaks in his
elaborate analysis of the righting reaction in Asterias forrert 1s
certainly shown in Asterina, but appears to arise rather late in
the reaction. I have often seen the third arm detached rather
forcibly by the pull of the turning pair. In other cases, how-
ever, one can observe a voluntary retraction of the tube-feet of
the impeding arm or arms. That the unified impulse does play a
great part is shown by the fact that, in general, a bad start is
always or almost always compensated and a good solution arrived
at, for whatever number of arms is employed at the beginning
the turn is almost invariably made on two adjacent arms. If the
unified impulse did not at some stage arise the righting reaction
would be a long drawn-out and confused affair, coming only by
chance to a successful issue.

A second neat solution of the righting problem is afforded
when the righting is carried out mainly by one arm, helped
towards the ena of the reaction by the arms on either side. This
solution is rare, and the method is not a particularly speedy one,
the times taken ranging from 37 secs. to 120 secs.

A third method sometimes adopted is to turn on three adjacent
arms. On condition that the middle arm of the three takes the
lead this method is quite successful: it is, in fact, hardly to be
distinguished from the method of turning mainly upon one arm.
If, however, the middle arm Jags behind and the outer arms get
ahead of it, trouble ensues and a deadlock occurs

Turning on four arms is really not met with. Four arms may
firmly attach and an attempt be made to turn on all four, but the
actual turning must take place on the middle pair, the outer arms
being forced to let go. An attempt to turn on four resolves
itself into a turn on a single pair.

Tf all five arms attach, no progress can be made until at least
two relax their hold.

Comparison with the account given by Jennings of the methods

428 MR. E. S. RUSSELL ON THE RIGHTING

used by Asterias forrert shows that Asterina attempts practically
all the methods employed by Asterias, but is successful only in
the measure that its organisation permits. Asterina is a short-
rayed, comparatively rigid form, and cannot twist its rays (espe-
cially round a longitudinal axis) to anything like the extent
possible to the long-rayed Asiterias. Its righting reaction is
accordingly less varied than in dAsterias, for the simple reason that
certain solutions open to Astertas are mechanically extremely
ditticult or quite impossible for Asterina. Organisation, in fact,
must be regarded not only as a means towards action, but also
as a limit upon activity—a limit which the animal in its active
striving tries always to overcome.

The followi ing is a summary in Jennings’ own words of the
various modes ‘of righting which he has observed in Asterias
forreri :-

The sunplest and neatest method of turning is the follow-
ing: Two adjacent rays twist their tips in such a way that the
ventral surfaces of the two face each other; then the tube-feet of
these rays attach themselves and throw the starfish over in a neat
somersault......

“2. The tips of the two adjacent rays may so twist that the
ventral surfaces do not face each other, but both face in the same
direction. The tube-feet then take hold and throw the starfish
over,—twisting it about an axis which passes lengthwise through
one of the attached rays. This method of turning is extremely
dificult and awkward, but is seen at times. Usually when two
rays become attached in the way described, a third ray takes hold
and aids the turning, the method then forming a transition to
eat given next,

3. Three adjacent rays twist, attach themselves, and remain
ea all pulling throughout the reaction. Usual y the animal
turns pranany by the aid of the two outer rays, while the
middie one is relatively passive, and is compelled to double back
under as the animal turns. Often this middle ray walks back-
ward beneath one of the other rays, or the other walks actively
over its surface or there is a combination of these two movements,
till the normal position is reached......

“4, Four of the rays take hold, two extending to the right,
two to the left....:.. (all remain attached during the turn J.

All of the rays attach themselves. Now the turning can
be accomplished only by the release of certain rays ......

“6. An unusual method is that in which but one vay twists
and attaches itself, and by its unaided eftorts turns the starfish
about an axis passing through this vay. :

“7, A still more unusual type 1s seen in “the performance of
the righting reaction without attachment of the tube-feet of any
of the rays....” (1907, pp. 125 & 128).

This last method, in which the starfish raises its dise high by
standing on the tips of all five rays and then swings one or more
rays over or under until it topples over right side up, I have

—

REACTION IN ASTERINA GIBBOSA. 429

not observed in Asterina, and Jennings has seen it in Asterias
only when the tube-feet were prevented from taking hold.

Method 6 is used by Asterina, but the turning is round an axis
transverse to the ray, not longitudinal as Jennings’ account seems
to imply. (In general, the inverted Asterina tends to roll its rays
backwards from the tip rather than twist them to the side round
a longitudinal axis, as is customary 1n Asterias.)

Method | is in Asterina as in Asterias the neatest and most
successful solution. Method 2 also occurs, but seems in Asterina
always to require the assistance of a third ray, which generally
super’ sedes one of the original pair. Method 4, as already pointed
out, is not mechanically possible i in Asterina.

It is in Method 3 that the difference between the two species
comes out most clearly. If Asterina tries to turn on three rays
and the two outer get ahead, the middle one becomes sharply bent
back upen itself and is unable to straighten out owing to the rays
on either side pinning it down hard against the bottom. A dead-
lock follows, which lasts for several minutes, the two unattached
rays standing up almost vertically in the water the whole turning
movement being suspended half-way.

One gets the impression that the starfish is thoroughly puzzled
by the situation, and it certainly takes a long time to arrive ata
solution. Various solutions are found. In Case I. (supra, p. 425)
the starfish finally bent down one of the free rays and took hold,
then relaxed the two outer attached rays enough to push the
middle ray through. In Case II. both of the free arms were bent
down and attached, then one of the original outer pair let go and
so released the prisoned ray. In Case IU. the free arms bent
down and attached, pulling the body flat by main force, the
prisoned ray being subsequently freed by adjustment of the
neighbouring rays. In another case, not included in the general
series, in which the deadlock lasted about 4 minutes, an outer
ray let go, and the starfish turned on the other two attached rays,
the free arms taking no part in the solution. Case LV. in the
general series was of another character. Three rays had attached,
the tips of two rays twisting towards one another: then a fourth
ray took hold; finally the starfish turned on a pair of rays at the
side, having taken over three and a half minutes to work out this
solution.

In Asterias, when three rays attach and the outer rays get
ahead, as is apparently usually the case, turning is carried out
with comparative ease, the length and flexibility of the rays
permitting the middle vay to era wl over or under the ones at the
side, instead of being pinned between them as happens in Asterina
with its short and flat rays.

T did not investigate the question as to whether the individual
Asterina shows any preference for using a particular ray or
combination of rays, as Jennings has demonstrated for dsterias
forrert (1907, p. 144), but it is clear from the detailed records
given above that each individual tends to stereotype the method

430 MR. E. §. RUSSELL ON THE RIGHTING

used in the first few trials. Some specimens, notably B and G,
turned unhesitatingly on an adjacent pair, F showed a tendency
to attach all its rays, H got itself tied up in a deadlock three
times in ten trials, K showed a propensity to use one ray for
turning. It is difficult to say whether these tendencies are due
to individual idiosynerasy or to the formation of transitory habits.

Perceptions involved in the Righting Reaction —When Asterina
is turned on its back two obvious changes take place in its
relation to environment—the tube-feet lose contact with the
bottom, and the action of gravity upon the animal is reversed in
direction. The irritating effect of light upon the ventral surface
need not be considered as an essential factor, for, according to
Preyer (1886-7, p. 99), the righting reaction may take place in
the dark. The stimulation of the dorsal surface by contact with
the bottom may also be eliminated from the essential conditions,
since Asterina is often found under stones with its back in contact
with the bottom. And Preyer (2bid. p. 107) has shown that
removal of large pieces of the dorsal integument does not hinder
the righting reaction.

In order to study the relative importance of the two main
conditioning factors—contact and gravity—I carried out the
following experiments on the beach with a couple of large and
active specimens :—

I. The starfish was turned on its back, and a small piece of
slate was held 1-2 mm. above the ventral surface. The starfish
attached completely and could be lifted away adhering to the
slate—the righting reaction was not carried out, the starfish
remaining back downwards. If, however, the slate was not lifted
away and the starfish remained in contact with the bottom, it
left the slate and crawled on to the bottom, thus completing the
righting reaction. (10 trials.)

II. The slate was presented half-way through the turning
movement. The starfish attached some of its tube-feet tem-
porarily but completed the turn towards the bottom. (10 trials.)

III. A piece of Mucus of about the same area as the starfish
was placed on the ventral surface of an inverted Asterina. The
animal attached its tube-feet and carried out “ walking” move-
ments with them, by means of which the weed was carried right
off the ventral surface. The starfish then proceeded to turn.

ITV. The starfish was held lightly in position by means of
angled pins on the under side of a floating block of wood, the
ventral surface facing earthwards. It turned so as to bring its
ventral surface into contact with the wood. In spite of the fact
that the pins interfered somewhat with the righting movements,
less than one minute sufficed in two cases for the turn, less than
two minutes in three other cases.

V. Propped up with its back against a vertical side of rock,
the starfish invariably turned towards the earth, pivoting rapidly
on the pair of arms whose tips were in contact with the horizontal
surface.

REACTION IN ASTERKINA GIBBOSA. 431

From these experiments it seems clear that the primary aim
of the righting reaction is to bring the tube-feet into contact
with a solid and resistant surface. The first thing Asterina does
when detached from its hold is to stretch out its tube-feet and
feel round in all directions. Mere tactile contact is, however, not
sufficient, as Experiment III. shows—the surface must resist a
pull. The perception involved in the contact reaction is not so
much tactile as kinesthetic. It does not seem to matter very
much whether the surface of adhesion lies below the starfish, as
in the normal righting reaction, or above it, as in Experiment IV.
There is no pronounced tendency to take up a definite position
relative to gravity. Under natural conditions Asterina may be
found either adhering back downwards on the underside of a
stone or back upwards on the bottom underneath the stone, or
again wedged along the lateral edges of the stone. Experiment I.,
however, indicates that Asterina prefers to turn its ventral surface
to the bottom rather than crawl on a small inverted stone, and
this fact might be adduced, together with the similar results
obtained by Preyer (¢bid. p. 116), in favour of graviperception
taking a part in the reaction. On the other hand, Asterina
may crawl off the piece of slate simply because of its small area.
(The result of Experiment II. is best explained as due to that
“persistence of an established impulse,” which is a characteristic
feature of the behaviour of many lower animals (Jennings, ibid,
pp. 115-6, 145-6).

The probability is that graviperception in the starfish, if really
existent, is purely kinesthetic. It is well known that when a
starfish is walking undisturbed over a horizontal surface the
tube-feet do not attach firmly to the bottom but act very much
like legs which push the animal forward. This has been clearly
shown by Jennings for Asterina forreri, and is true also of
Asterina. If, however, Asterina is irritated it huddles down and
grips the bottom firmly with its suckers. Also when at rest it
holds on fairly tightly. Similarly, if it is mounting a steep slope,
ov adhering to the under surface of a rock, it must hold on tighter,
bringing its suckers into play. Graviperception in Asterina, if
one can properly use the term, must be little more than a dim
perception of the difference between a surface on which it may
walk and a surface to which it must cling. This gives the
possibility of a choice between a back downwards and a back
upwards position. Space to the starfish must be a tactile or
better a kineesthetic space.

To sum up, the two factors provisionally distinguished above—
contact and gravity—probably resolve themselves into one, for
both appear to depend upon kinesthetic impressions. The aim
of the reaction is to get a firm foothold on something solid ; once
this is found the starfish is probably able to distinguish by
muscle sense, in a rough and ready way, whether the surface is
horizontal, vertical, or inverted,

It is possible that a third factor plays a part in the orientation

432 THE RIGHTING REACTION IN ASTERINA GIBBOSA.

of the body with respect to gravity—the drag or pull of the
internal organs. The existence of this factor was recognized by
Preyer (ibid. p. 121), and its importance in Crustacea has recently
been emphasized by W. von Buddenbrock (1914). As the star-
fish shifts its position with respect to gravity the internal organs,
or at least such of them as are slung by mesenteries, must shift
their position within the body. It is conceivable that such
alterations in stress are perceived by means of an interoceptive
nervous mechanism. In Experiment V., for instance, the stimulus
to turn to the horizontal may have been supplied by the down-
ward drag of the internal organs.

In conclusion, a remark upon method may perhaps be per-
mitted. If the whole reaction is considered as behaviour im the
real sense, that is to say, considered strictly from the psychological
point of view—as has in effect been done in the above discussion—
the problem ought not be formulated in terms of physiological
reaction to single environmental stimuli, but in terms of response
to a situation presented to the animal as a whole, or, in the case
of an experimental dissociation of the normal combined action of
the factors involved (as in Experiment IV.), in terms of response
to one factor as representative or suggestive of the whole normal
presented situation.

References to Literature.

BuppEenprock, W. von. 1914. ‘“ Ueber die Orientirung der
Krebse im Raum.” Zool. Jahrb. (Allg.) xxxiv. pp. 479-
514.

Guaser, O. C. 1907. ‘Movement and Problem Solving in
Ophiura brevispina.” Journ. Exper. Zool. iv. pp. 203-20.

Jennines, H.S. 1907-8. ‘ Behaviour of the Starfish Asterias
forreri de Loriol.” Univ. California Public. Zool. iv.
pp. 93-185.

Pressner, H. 1913. ‘‘ Unters. ti. d. Physiologie d. Seesterne.”
Zool. Jahrb. (Allg.) xxxiul. pp. 361-86.

Preyer, W. 1886-7. ‘ Ueber die Bewegung der Seesterne.”
Mitteil. Zool. Stat. Neapel, vii. pp. 27-127 and 191-233.

papel ae.

WXPERIMENTS ON SEX DETERMINATION. 433

28. lixperiments on Sex Determination.
By. Lt.-Col. 8. Moncxron Copeman, M.D., F.R.S., F.Z.3.

[Received October 21, 1919: Read November 18, 1919.)

The experimental work forming the basis of the present
communication was carried out at intervals, as opportunity
served, during the years 1902-1905. But although the main
outcome of the work was demonstrated (with the aid of lantern-
slides) to the Physiological Society in May 1908, and at the
Dublin meeting of the British Association in September 1908,
no account of it has previously been published. ‘The reason for
this omission is to be explained by the unfortunate fact that the
collection of diagrams and other records, which was handed over
by the writer, for purposes of criticism, shortly after the research
came to an end, has only recently been recovered.

From the earliest ages various writers have suggested the
possibility that the reproductive glands, male or female, or both,
were concerned in the production of offspring of one or the other
sex. The most prevalent idea would seem to have been that the
glands of one side of the body gave rise to male-producing
spermatozoa or ova respectively, and vice versdé. Some observers,
among the most recent of whom Dr. Rumley Dawson should be
mentioned, have favoured the supposition that the male or the
female gland only was concerned; while others have elaborated
the necessity for inter-action of spermatozoa and ova produced
by the reproductive glands of the same, or of opposite sides of
the body.

In the hope of definitely determining whether or not any of
these suggestions could be shown to possess basis of scientilic
fact, a series of experiments were devised which were carried out
on a somewhat extensive scale. ‘The animals employed for the
purpose of these experiments were, for the most part, rabbits,
although use was also made, to a smaller extent, of pigs, cats,
guinea-pigs, and mice. Of these animals the greater number
were semi-castrated, or semi-spayed (according to sex), and
subsequently kept under observation until such time as they
had completely recovered from the operation. An additional
number of animals which had already been operated on in this
fashion were obtained from dealers and breeders. In a later
series of experiments, with the object of avoiding possible fallacy
from the loss of an internal secretion, ligature in two places of
the uterine cornu, or of the cord (subsequently of the vas
deferens only), on one side of the body, with partial or complete
section of the tissues between the ligatures in order to prevent
possible regeneration, was substituted for ablation of the ovai y
or testicle respectively.

Ultimately the experimental animals were bred :-—

(4) with entire animals of the opposite sex,
(6) with other semi-castrated, or semi-spayed individuals,

A34 LT.-COL. 8S. MONCKTON COPEMAN :

in such manner as was intended to comprise all possible permu-
tations. Owing, however, to loss of animals from inter-current
disease, more particulary tuberculosis, and from accidents of one
and another kind, it did not prove possible to carry out the
scheme in its entirety.

The intended scheme of the breeding experiments may perhaps
be best shown in tabular form as follows :—

3 Entire. Bred with 2 R.
6 Entire, be Peel Dp
Ge Met - jo Qe yates,
3 R. 60 Shak teu
3 t. 99 8) 2 Ibi
SG Ie A » ¢ Entire.
3 LL. 5 Re Onl te
Gy IVE =n Pai Oe bas
(R. = possessing gland on Right side only.
L. = ” ” ” Left ” ” )

Without entering into details concerning the number and sex of
the offspring resulting from interbreeding on this plan, it may be
mentioned that although, so far as possible, each particular mating
was repeated on one or more occasions, in only four instances
were families of one sex, either all males or all females, obtained.
But even so, when the same animals, or animals in similar
condition, were again mated, a uni-sexual family was not again
obtained, definite indication being thus afforded that the result
obtained in the first instance was quite fortuitous. As an
instance of completely contrary results, in respect of offspring,
obtained on mating two sets of animals in similiar condition the
following experiments may be cited :—

March \ith, 1902.

Bernau buck: Castrated on Lerr side + Bernau NORMAL doe.
Litter = 5 M, 0 F.

June 5th, 1902.

Blue buck: Castrated on Lery side + Wakehurst NoRMAL doe.
Litter =0™M, 2 F.

while the mating of yet another similar couple was as follows :—
September 12th, 1902. Litter =2M,1F.

Equally divergent results were obtained as the result of crossing
semi-spayed does with normal bucks; or when bucks and does, in
both of which the reproductive gland of one or the other side had
been removed, or put out of action as the result of a vasectomy,
or section of a tube, were mated.

In the later series of experiments the female was killed shortly
before her litter was expected, in order that the young might be
examined in sitw in the uterine tube or tubes. as it seemed not

EXPERIMENTS ON SEX DETERMINATION, 435

improbable that useful information might be obtained from deter-
mining the relative position in the tube of foetuses of one or the
other sex. Records of the results obtained were indicated in the
form of a simple diagram, devised for the purpose. (Lantern-
slides of these were shown on the screen.) One point of interest
emerging from study of these diagrams is the frequent occurrence
of male couples. Whether these, as in the case of identical twins
in the human subject, are to be regarded as the outcome of
division of a single ovum could not be certainly determined.

In a lecture on “ Internal Secretion and the Ductless Glands”
by Dr. Swale Vincent (Lancet, Aug. 1, 1906, footnote) will be
found a reference to a further outcome of this investigation,
personally communicated to him, “ pointing distinctly to an in-
ternal secretion on the part of the interstitial cells” of the
testicle. Dr. Vincent notes that the “results are generally in
agreement with those of Ancel & Bouin (Recueil de Médicine
Vétérinaire, Jan. 15, 1904).” It became apparent also that in
the case of those rabbits in which the vas only was tied (with
careful exclusion of blood-vessels and nerves), the animals, as
contrasted with those in which semi-castration had been per-
formed, showed no diminution of desire, or capacity, for sexual
intercourse; while, on the average, they obtained larger families.

The communication also included an experimental and critical
review (illustrated by lantern-slides) of previous work on sex
determination by various observers whose investigations had
dealt with the study of this problem in plants, insects, and certain
of the lower vertebrates.

Proc, Zoou. Soc.—1919, No. XXX. 30

ON THE DIGASTRIC MUSCLE OF THE MACAQUES. A37

29. The Variations in the Digastrie Muscle of the Rhesus
Macaque and the Common Macaque. By Cuas. F.
Sonnac, M.D., Ch.B., F.Z.8., Anatomist to the Society.

| Received October 24, 1919: Read November 18, 1919. ]
(Text-figures 1-5.)

In the course of several dissections to explore the extrinsic
muscles of the tongue of the Rhesus Macaque, I have been struck
with the various appearances of the digastric muscle. I decided,
therefore, to examine the muscle of every monkey brought to
the Prosectorium, and the present communication is based on the
dissections of fifteen animals.

The muscle belongs to Parson’s first type*. In it the posterior
bellies of the two digastric muscles terminate in long slender
tendons which, passing forwards and inwards, unite to form an
arch anterior and superficial to the hyoid bone. From thearch the
fibres of the united anterior bellies pass to the lower edge of the
inferior maxilla (text-fig. 1), This arch is closely applied to the
mylo-hyoid muscle and compels it to have a dome-like appearance.
In some cases a probe can be passed under the arch, but in most
of them they are fused and can only be torn apart. Moreover,
in about fifty per cent. of cases there is no connection between
the arch and the hyoid bone.

The extent and nature of the attachment of the anterior
bellies to the inferior maxilla vary. In some cases the muscle
extends right back to the angles on both sides, but more commonly
the extreme posterior limit is about half an inch anterior to the
angle, and lymphatic or submaxillary glands are lodged in the
small recess. In the majority of cases the attachment to the
maxilla is muscular, but in one case there were a number of
tendinous fibres interspersed among the muscles, In no case did
I find any exception to the rule that the two anterior bellies are
joined in the middle line.

In a little more than fifty per cent. of cases, the space between
the tendinous arch and the hyoid bone is occupied by a membrane,
by fibrous bands, or by both together, In one case, a membrane
alone filled the space (text-fig. 2), and there was no fusion be-
tween it and the mylo-hyoid muscle underneath. The anterior
bellies did not reach the posterior angles of the jaw.

Tn one case (text-fig. 3a) the space was filled up by a membrane,
and the tendons of the posterior bellies were connected to it by
short fibrous bands. This specimen was interesting in another
way, for the membrane between the arch and the hyoid bone was
common to the digastric and mylo-hyoid muscles, so that the two
could not be separated. The mylo-hyoid muscle was defective in
front, there being a V-shaped gap which was filled by membrane
(text-fig. 3).

* F. G. Parsons, Journ. Anat. Physiol. 1898, p. 436.
30*

438 ; DR. C. F. SONNTAG ON THE

Text-figures 1--5.

WS j
Fig. 4. NS Vif Fig. 5. WS
Text-figs. 1-4.—Variations in the digastric muscle of the Rhesus Macaque
(Macacus rhesus).

Text-fig. 5.—The digastric muscle of the Common Macaque (Macacus
fascicularis).

A.B.D.: Anterior belly of the digastric muscle. P.B.D.: Posterior bellies of the
digastric. M.H.: Mylo-hyoid muscle. MEM.: Membrane. F.B.: Fibrous bands.
S.M.G.: Submaxillary gland.

DIGASTRIC MUSCLE OF THE MACAQUES. 439

In six cases the space between the arch and the hyoid bone
was occupied by a triradiate fibrous band, the vertical limb of
the Y being attached to the bone, and the two lateral limbs to
the posterior edge of the arch anterior to the posterior extreme
of the anterior bellies. In the fresh specimen the contrast
between the white band and the subjacent red mylo-hyoid muscle
was very marked (text-fig. 4).

In the single example of the Common Macaque (J/acacus
Jascicularis) which I examined (text-fig. 5), the arch was connected
to the hyoid bone by a broad band of connective tissue into which
two slender bands, from the tendons of the posteriar bellies, were
attached.

The condition present in the Common Macaque (text-fig. 5)
differs only in degree from the variety of the Rhesus Macaque
present in text-fig. 3a.

As there is no gap between the two anterior bellies of the
digastric, and no solution in the continuity of the tendinous
arch, it is not easy, at first sight, to give a rational explanation
of the above appearances. Iwas enabled, fortunately, to examine
a series of monkeys which exhibited transitions between Parson’s
first and third types of muscie. Each of these had one or more
of the structural elements shown in text-figs. 2-5.

The variable degree of adhesion between the digastric anterior
bellies and the subjacent mylo-hyoid muscle indicates the origin
of the former from the latter, and is the beginning of the meta-
morphosis. This is followed by the appearances in text-figs. 2-5,
and the condensation of the lateral fibres of the anterior bellies,
constituting the nearest approach to a true digastric muscle, is
the last stage.

In my opinion : —

(1) The anterior belly arises from the mylo-hyoid muscle by
splitting. There then ensues :—

Two muscles joined to the hyoid bone by membrane (text-
figs. 2 and 36).

(3) The pull of the anterior and posterior bellies separates the
latter into bands.

(4) These bands disappear leaving a muscle of Parson’s first
type.

(5) The tendinous arch splits and the anterior bellies separate
leaving a true digastric muscle. These I have not seen in
Macaques, but they are present in Cercopithecus wthiops, Cerco-
cebus ethiopicus, and Cebus altifrons.

(6) The muscle in the Macaques has stopped short of being a
true digastric condition in which there are two muscles, each
with an anterior and posterior belly (Parson’s third type).

(7) There is no variation in the posterior bellies.

ie Se’
*

‘ } ’ Ae
‘ :
’ * ris
.
i : ; Asn eae |
: ene i f AT \ ‘

NEMATODE PARASITES OF CHAPMAN’S ZEBRA. 441

30. On the Nematode Parasites of a Chapman’s Zebra.
By M. Turner, B.Se.

| Received October 8, 1919; Read November 18, 1919. }
(‘Text-figures 1-6.)

In September of this year a Chapman’s Zebra, which came
from Africa to the Zoological Gardens, London, ten years ago,
died. The post-mortem showed that its large intestine contained
many Nematode parasites, A collection of them was made by
Professor Leiper, and, through his kindness, I was enabled to
examine the material in detail.

Altogether seven species of Nematodes were represented in the
Zebra. An enumeration of them is as follows :—

Oxyuris curcula Rad, 1803.
Strongylus vulgaris (Looss, 1901),
Strongylus edentatus (Looss, 1901).
Triodontophorus intermedius Sweet, 1909.
“Lsophagodontus robustus Giles, 1892.

(?) Cylichnostonum goldi Boulenger, 1916.
Cylichnostomum zebree, sp. n.

The real object of these investigations was to ascertain whether
any of the parasites had lived in the Zebra in Africa, and had
persisted throughout the ten years in England, or whether they
had only been acquired by the Zebra since its arrival in England.
This point, however, was not definitely cleared up, as none of the
Nematodes found can be said to have a solely English or African
distribution. Thus Oxyuris curvula, Strongylus vulgaris, and
Strongylus edentatus are found all over the world. Z%iodonto-
phorus intermedius has been recorded from Australia and England,
and C@sophagodontus robustus from India and England. The
localities are so far apart that it would seem probable that these
species are present in intervening countries. Cylichnostomum
goldi has been recorded only once, and that so lately as 1916,
when it was found in England. More records of its occurrence
and their locality are necessary before it is possible to state its
distribution definitely. Cylichnostomum xzebree is an hitherto
undescribed species.

The impression obtained from the above facts is that the
Zebra acquired the majority, at least, of its parasites in England,
though no definite proof of this is given.

Oxyuris cuRVULA Rudolphi, 1803.

Two males were found in the Zebra. This discovery is inter-
esting, as the males of this species of Oxyuris ave rather rare. It
was on account of this rarity that the male was only described.
and figured for the first time by Railliet, in 1895, although the

442 MISS M, TURNER ON THE NEMATODE

female, under the name 7richocephalus equi, had been described
by Schrank so long before as 1788.

Oxyuris curvula has been recorded in the Horse, Ass, and
Mule.

SrronGyLus vouteanris (Looss, 1901).
SLRONGYLUS EDENTATUS (Looss, 1901).

In 1901 Looss recognised the three species of Strongylus,
2. €., Strongylus vulgaris, Strongylus edentatus, Strongylus equinus,
which are known to occur in the Horse, Ass,and Mule. Of them
the first two were represented in the Zebra.

TRIODONTOPHORUS INTERMEDIUS Sweet, 1909.

This species of 7’riodontophorus was first described in 1909 by
Sweet, whose material consisted of three female specimens
obtained from a Horse in Victoria. In 1916 Boulenger found
Triodontophorus intermedius abundantly represented in Horses
in Worcestershire, England.

The female of Triodontophorus intermedius from the Zebra
measures 18°D-21°5 mm. in length, and the male 15-16 mm.
These measurements are slightly larger than those given by
Sweet or Boulenger. The spicule of the male is long and rather
thick. It measures 3°8 mm. in length by about 0:03 mm. in
breadth at its proximal end.

(EsopHacopontus RoBustuS Giles, 1892.

Hsophagodontus robustus was first discovered in Horses and
Mules in India by Giles in 1892. Apparently this Nematode is
rare in England, as there is only one previous record of its
occurrence here. This was in 1916, when Boulenger obtained
about twelve specimens from the colon of a Mare. Its numbers
in the Zebra were sparse.

In the female Wsophagodontus robustus from the Zebra the
vulva is 2°8 mm. in front of the anus. This measurement differs
from that of Giles, who gives ‘‘akout 1 mm.,” but confirms those
of Boulenger, who gives 2°3-3 mm.

The anus is 0-7-0°75 mm. from the posterior end of the body
in the Wsophagodontus robustus from the Zebra.

(2) Cyticunosromum Goupt Boulenger, 1916.

The only previous record of this species was made in 1916
by Boulenger, who obtained it from the intestine and caecum of
several Horses in Worcestershire, England. :

The material obtained from the Zebra and diagnosed as
(2) Cylichnostomum goldi, consists of a single female specimen.
So far as could be ascertained from this female it appears to
belong to the species goldi, although it is somewhat longer, being
8°35 mm., than the females recorded by Boulenger. In the
absence of more material it is impossible to be certain of the
species at present. The diagnosis of this female was rendered

See ae, ant «

PARASITES OF A CHAPMAN’S ZEBRA. 443

more difficult by the similarity presented by the two species
Cylichnostomum goldi and Cylichnostomum pseudo-catinatum
Yorke & Macfie, 1919, seen in the following comparison.

Both C. goldi and C. pseudo-catinatum are described as having
a small and delicate body. Boulenger gives the length of the
male OC’. goldi as from 5°2-6 mm. and its maximum breadth
2304-280 (average 255). Yorke and Macfie found that the
length in ten males of C. pseudo-catinatwm ranged from 5:2-
66 mm., and that the maximum breadth averaged 260. The
females of C. goldi were 6-6-7 mm. in length and 280-300
(average 290m) in breadth. Ten females of C. psewdo-catinatum
measured from 6°1--7-7 mm. in length and averaged 320 in
breadth. These measurements show that the size of the body
of the worms of both species is practically identical. The slight
difference between the two species present here is not greater
than that between the measurements of individuals of the same
species.

A neck is reported to be absent in C. goldi, but present in
C’. pseudo-catinatum. In the figure of the former species, how-
ever, a very slight constriction in the neck region is seen.

The mouth collar in both species is marked off from the rest of
the skin by a definite constriction. In the figures given, the
mouth collar, in both, is seen to be of a similar shape, being
rather high and almost ellipsoidal in lateral view.

The submedian head papiliz in C. goldi are small and do not
project beyond the middle of the external leaf crown. In
C. pseudo-catinatum these papille are larger and project beyond
the anterior edge of the external leaf crown. In both species
the submedian head papillz are conical and their extremities are
not separated off by lateral notches.

Boulenger says that the lateral head papille of C’. goldi are
very inconspicuous, while Yorke and Macfie say that those of
C. ‘pseudo-catinatum are prominent. In the figures, however,
the lateral head papille of both species appear very similar as
regards size, in relative proportion to the rest of the head, and in
shape.

Boulenger gives “ about 20” as the number of elements in the
_ external leaf crown of C, goldi, while Yorke and Macfie give 20
“as the number of these in C. pseudo-catinatum. These leaves
are large and pointed im both species.

The number of leaves in the internal leaf crown of C. goldi is
given as 30-32, and Boulenger says that these leaves are shorter
than those from the external corona. Yorke and Macfie only
say that the internal leaf crown of C. pseudo-catinatum consists
of numerous long narrow elements. Their figure shows these to
be shorter than the leaves of the external corona. In both
C. goldi and C. pseudo-catinatum the internal leaf crown arises in
more than one plane. This arrangement is present in the other
members of this group of Cylichnostomes, 7.e., C. catinatum and
C. alveatum.

444 MISS M. TURNER ON THE NEMATODE

The mouth capsules of both C. goldi and C. pseudo-catinatwm
are like those of C. catinatum and C’. alveatun in being ellipsoidal
in transverse section, with the longer axis dorso-ventral. ‘The
mouth capsule of C. pseudo-catinatum is described as having an
oblique floor. ‘This character is not mentioned in C. goldi. The
optical section of the mouth capsule differs in the two species.
In ©. goldi the walls in ventral view converge anteriorly, while
in C’. pseudo-catinatum they converge posteriorly. The height of
the mouth capsule of C. goldi is given as “about 20m.” In the
males of C. pseudo-catinatum this height varies from 22 54-25
and in the females from 24u—-29y, so that from these measure-
ments the mouth capsule of C, psewdo-catinatwny appears to be
slightly larger than that of C. goldi. ‘The measurement of the
breadth of the mouth capsule has been undertaken 1 such
different ways in the two species that it is not possible to com-
pare the measurements given. From the figures it appears that
the proportion of the breadth of the mouth eapsule to its height
is greater in C. pseudo-catinatwn than in C. goldi.

The dorsal cesophageal gutter does not project into the mouth
capsule in either species.

The esophageal funnel is well developed in both C. goldi
and C. pseudo-catinatum. In the latter, however, cuticular
thickenings lining this funnel are not figur ed.

The cesophagus. in eight males of C. psewdo-catinatum varies in
length from 314-349 and in breadth from 70u~—824. In eight
females the cesophagus varies in length from 3224-363 ane in
breadth from 724-83. ‘Taking the artile of these measurements
the cesophagus of C. pseudo- catinatum varies from 314y-363p In
length and from 7Cp—83 in breadth, and these measurements
are not very different from the 300p- 850m in length and the
70-75 in breadth that Boulenger gives for the cesophagus
of C. goldi. In the cesophagus of both species the posterior bulb
is not markedly swollen.

The cervical papille of C. goldi are situated about 2700-300
from the anterior end of the body. In (. psewdo-catinatwm the
cervical papille are at about the same level as the excretory
vesicle, that is from 67-152 from the posterior end of the
cesophagus. It is not possible to compare these two sets of
measurements. In the figure of C. goldi that part of the
cesophagus behind the excretory vesicle is about one-fourth of
the length of the whole cesophagus, while in the figure of C. pseudo-
catinatum the proportion of the same two parts is one-third. It
may be that this figure of C. pseudo-catinatwm is that of a
specimen where the excretory vesicle is about 152 from the
posterior end of the cesophagus. In the figures the cervical
papillae of C. goldi are slightly more posterior in relation to the
excretory vesicle than those of CO. pseudo-catinatum.

Female. In both C. goldi and C. pseudo-catinatum the posterior
region of the female is bent dorsally to make almost a right angle
with the axis of the rest of the body. Jn both, the tail behind

PARASITES OF A CHAPMAN'S ZEBRA. 445

the anus is narrowed suddenly to form a point. The figures
show that the relative proportion of the ovi-projector to the
vagina is about the same in both species, The ventral promi-
nence and other curves in this region of C’. goldi appear less than
those of O. pseudo-catinatum.

Boulenger gives the distance between the anus and vulva of
CO. goldi as about 90-100, and for the same in C. pseudo-
catinatum Yorke and Macfie give 454-854. Thus the anus
and vulva are almost the same distance apart in, at least, some
members of the two species.

Male. The two descriptions of the posterior regions of the
males of C'. goldi and C. pseudo-catinatum are identical in many
important points.

The median dorsal lobe of the bursa is short and almost semi-
circular in both species. In both, the dermal collar is well
developed on the anterior and posterior (ventral and dorsal,
Yorke & Macfie) surfaces. The pre-bursal papille are well deve-
loped in both.

The genital cone of both species is similar in having its
appendages in the form of very thin delicate plates (slight
elevations, Yorke & Macfie) each provided with two slender
finger-shaped processes. Of these two processes the inner one
is the larger in both species. The figure of the genital cone of
C. goldi is indeed very similar to that of C. pseudo-catinatum.

In C. pseudo-catinatuwm the main trunk of the posterior ray of
the bursa and its second lateral branch are each provided with a
small accessory branch. Boulenger makes no mention of any
accessory branches in his description of C. goldi, but his figure
of a lateral view of the bursa shows an indication of one on the
second lateral branch of the posterior ray, though nothing is seen
on the main posterior ray. In the lateral view of the bursa of
C. pseudo-catinatum the accessory branch of the main posterior
ray is not shown.

Boulenger makes no mention of the spicules of C’. goldi, so it

“it is not known whether they resemble or differ from those of
C. pseudo-catinatum which are figured by Yorke and Macfie.

This comparison of the two species, made only from descrip-
tions and figures, was made more difficult by the different
methods of treatment employed by the different writers. Never-
theless enough has been shown to prove that there are some
features of very great similarity, if not of identity, in C. golda
and CC. pseudo-catinatum.

CYLICHNOSTOMUM ZEBR4#, Sp. D.

Specific diagnosis :—

Cylichnostomum zebre is one of the largest of the Cylichno-
stomes, being slightly larger than C. elongatum, but smaller than
O. auriculatwm, which is the largest of all members of the family.
The male of C. zebree measures 13-13°5 mm. in length and has a

446 MISS M. TURNER ON THE NEMATODE

maximum thickness of 0:75 mm. The female is 17—20°5 mm. in
length with a maximum thickness of 1 mm. Two males and ten
females were measured. The body is thickest in the middle, but
is quite thick throughout most of itslength. About 3 mm. from
each end it tapers to the extremities.

‘The wall of the intestine, especially the anterior end, is dark
with a brownish pigment.

Text-figure 1.

Cylichnostomum zebre, sp. 0.

Anterior end of body. View from dorsal side.

The cuticle is finely striated, the striations or the middle of
the body being 15y apart. v

The head is marked off from the body by a very slight con-
striction. It has a diameter of 280u-300u in the five specimens
measured, The mouth collar is well developed, with a height of
3 the breadth. It is almost ellipsoidal in shape, being flattened
anteriorly. The lateral head papille are broad and do not project
beyond the mouth collar. The submedian head papille are
rather large and conical and project almost to the anterior edge
of the external leaf crown. -

The external leaf crown consists of 85-93 narrow and pointed
leaves, which project well in front of the anterior margin of the
mouth collar. The internai leaf crown has 48-59 leaves. Each
leaf is rather blunt and is about twice as broad as one from the
external leaf crown. Amongst the ordinary cuticular leaves of

PARASITES OF A CHAPMAN’S ZEBRA. 447

this internal corona are six longer sharply-pointed ones. They
are also more refractive.

Text-figure 2.

Cylichnostomum zebre, sp. 1.
3

Anterior end of body. View from above.

Text-figure 3.

Cylichnostomum zebre, sp. n.

Anterior end of body. View from dorsal side.

The condition of the leaf crowns in C. zebre recalls that in
C. bicoronatum and C. ewproctus, for in these three the internal
leaf crown consists of fewer leaves than the external crown. In
other Cylichnostomes the reverse of this is the case.

448 MISS M. TURNER ON THE NEMATODE

The mouth capsule has a height (from the anterior end of the
cesophagus to the base of the leaves of the internal corona) of
854-90 and a breadth of 210u-250u. In optical section its

Text-figure 4.

Cylichnostomum zebre, sp. n.

Posterior end of Female. Lateral view.

walls seem to be composed of two parts jointed together. The
anterior part is pointed and curved inwards, bearing on its

ine

PARASITES OF A CHAPMAN'S ZEBRA. 449

posterior outer side a short spur, The posterior part of the
mouth eapsule wall is thick and becomes broader towards the
base, which is notched to form two parts, one narrow and the
other broad and rounded.

There is no dorsal gutter.

The cesophagus is 675-800 in length and 300p—-350p wide
at its broadest part. It is flask-shaped, with the nerve ring at
the base of the neck portion. The cesophageal funnel is well
developed, The anterior énd of the cesophagus is covered with
a cap of cuticle, which forms the floor of the mouth capsules and
is oblique from side to side.

The cervical papille are 6604—700pu from the anterior end of
the body and are situated halfway between the nerve ring and
the posterior end of the cesophagus.

The excretory pore is slightly in front of the cervical papille.

Female. The poster‘or region is usually in a straight line with
the rest of the body, but may be bent shghtly dorsally. It
tapers to the tip, which is rounded and rather knob-like.

The vulva is 1°6-2 mm. from the posterior extremity and
0-9-1 mm. in front of the anus.

The vagina is about 0°6 mm. in length.

The eges are 70-80 long by 504-55 broad.

Male. The genital cone is small. More detailed examination
of it was not possible, as it was somewhat damaged in both males.

Text-figure 5.

Cylichnostomum zebra, sp. n.

Accessory piece of Male.

An accessory piece 1s present. It is saddle-shaped with inturned
lateral margins. Its proximal end is bifid, each part ending in
a point. The spicules are thick and about 0-95 mm. in length.
Their distal ends seem to be simple points.

One of the most striking characters of the male is the finely
serrate edge of the free margin of the bursa. This is a character
only described in U. poculatwm amongst the Cylichnostomes.

5
The pre-bursal ray is short and the dermal collar small,

A450 MISS M. TURNER ON THE NEMATODE

The median lobe of the bursa is very short and notched. The :
dorsal vay, which narrows abruptly to form a finger-like distal
extremity, with its two branches arising together and equal in
length, difters from the dorsal ray of the bursa of any Cylich-
nostome hitherto described. So that the dorsal ray forms one of
the chief distinguishing features of Cylichnostomum zebre.

Text-figure 6.

Cylichnostomum zebre@, sp.n.

Bursa of Male. (a) Lateral view. (6) Dorsal view.

Another character peculiar to C. zebrw is the possession of the
six elongated and highly refractive leaves interspersed between

PARASITES OF A CHAPMAN’S ZEBRA, 451

the ordinary leaves of the internal leaf crown. These specialised
leaves have not been recorded in a Cylichnostome before.

In passing the final proofs of the above paper for press on
January 27th, 1920, I note that the Nematode described therein
as Cylichnostomum zebre n. sp. has been named Hexodonto-
stomum markusi n. gen. n. sp. by Ihle in Cent. f. Bakt., Abt. i.,
Orig. Bd. 84, 1920, Heft 1.

References.

Bouuencer, C. L. (1916). ‘Sclerostome Parasites of the Horse in
England. 1. The Genera 7'riodontophorus and @sophago-
dontus.” Parasitology, vill. (4).

—— (1917). ‘“Sclerostome Parasites of the Horse in England.
2. New Species of the Genus Cylichnostomum.” Para-
sitology, ix. (2).

Gites, G. M. (1892). ‘On a New Sclerostome from the Large
Intestine of Mules.” Sci. Mem. of the Medical Officers
of the Army of India, part 7, article 2.

Looss, A. (1901). ‘The Sclerostomide of Horses and Donkeys
in Egypt.” Ree. Egypt. Govt. School of Med. i.
pp. 25-139.

Rarer, A. (1895). Traité de Zoologie Médicale et Agricole,
p. 415.

ed F. v. P. (1788). “ Verzeichniss der bisher hinlang-
lich bekannten Hingeweidewurmer, etc.” Kgl. Svenska
Vetensk. Akad. Stockholm.

Sweer, G. (1909). “The Endoparasites of Australian Stock and
Native Fauna Part 2. New and Unrecorded Species.”
Proc. R. Soc. Vict. xxi. p. 454.

Yorke, W., & Macriz, J. W.S8. (1919). ‘*Strongylide in Horses.
vi. Cylicostomum pseudo-catinatum, sp.n.” Annals of
Trop. Med. and Parasitology, vol. xii. Nos. 3 & 4,
February, 1919.

* Not seen.

Proc. Zoou. Soc.—1919, No. XX XI. al

ON MESENTERIES IN URTICINA CRASSICORNIS. 453

31. The Development of the Mesenteries in the Actinian
Urticina crassicornis*., By JAmus F. Gemminy, M.A.,
MoD DSc

[Received August 18, 1919: Read November 18, 1919.]
(Text-figures 1-5.)

The adult Urticina is remarkable for having its mesenteries
and tentacles apparently arranged in 10-cycled symmetry (Dixon 4,
Faurot 5, Haddon 7), and on that account has been placed -by
various authors among the Paractinee. It is, however, a Hexac-
tinian, the arrangement of whose mesenteries has become modified
during early growth.

The species investigated was the large fleshy one with few
warts, occurring beyond tide-mark down to a depth of at least 35
fathoms, variously and often brightly coloured, attaining a size,
when fully grown, of over six inches in expanded disc diameter,
and having 160 tentacles of which 80 occur in the outermost
circle. ‘The other circles from within outwards consist respec-
tively of 10, 10, 20, and 40 tentacles. Gosse (6. p. 211) thinks
that this is just the shore species (his Vealia crassicornis) modified
for living under water, though he also describes an apparently
identical form as a separate species under the name Bolocera
eques (6. p. 351). I find that in the Firth of Clyde both the
shore and the submerged forms shed their eggs prior to fertiliza-
tion (a distinction from Lhodactinia crassicornis (3. pp. 39, 41)),
that development is the same in both, and that cross fertilization
can occur between them. On general grounds I would have
judged that the shore and the submerged forms were varieties of
the same species had my account (see below) of the development
of the mesenteries in the latter agreed with that of Faurot
(5. p. 172) for the former. It appears to have been the submerged
species whose development was investigated by Appelléf (1).
T. A. Stephenson, reviewing the nomenclature’in a note to me,
concludes that the shore form should be called Urticina (Ehren-
berg) coriacea (Cuvier), and the submerged form Urticina
(Ehrenberg) crassicornis (O. F. Miller), and that the two are
probably distinct species.

My material consisted of (@) young specimens reared to the
12 mesenteried stage from eggs shed in the tanks at the Millport
Biological Station, and (6) growth stages dredged from c. 20 fms.
near the Station, the youngest of which had only 12 tentacles.
All were studied by the method of serial sections. Of the 8
Edwardsia mesenteries, the ventro-laterals are the earliest to

* T have to express indebtedness to the Carnegie Trust for a grant towards
expenses of research work, part results of which are given in this and the following

paper.
ou

A454 DR. J. F. GEMMILL ON THE DEVELOPMENT OF

develop and the laterals the next, while the ventrals (sulears) and
dorsals (sulculars) appear almost simultaneously a short time after-
wards. Throughout the text-figs. these mesenteries are numbered
1, 2, 3, 4 respectively. Attachment of the larva then occurs,
and in a week or a fortnight a new mesentery (No. 5) forms in
each lateral Hdwardsia space, and shortly thereafter a new
mesentery (No. 6) in each ventro-lateral Hdwardsia space. The

Text-figure 1.

Fig. 1

=|
ge
or

THE MESENTERIES IN URTICINA GRASSICORNIS. 455

Text-figure 5.

4 13

EXPLANATION OF TEX?-PIGURES 1-5.

. Transverse section (diagrammatic) of late larva of Urticiia crassicornis
to illustrate the arrangement of the 8 Hdzardsia mesenteries.

. Similar transverse section, fourteen days after fixation, to illustrate the early
12 mesenteried stage.

. Diagrammatic transverse section of early growth stage in which an addi-

tional pair of mesenteries (Nos. 7,8) has appeared in each lateral and
ventral primary exoccele.

. Similar section of later growth stage in which the additional mesenteries
seen in fig. 3 have become mesenteries of the Ist Order, and a further pair
(No. 9) belonging to the same (i. ¢., the 2nd) cycle has appeared in each
dorsal primary exocele. The latter will ultimately take rank as a
mesenteric pair of the IInd Order.

. Similar section of still later stage in which (a) mesenteric pairs 10, 11
(8rd cycle mesenteries) have appeared and are growing forward to take rank
with 9 as mesenteries of the IInd Order, while (6) new mesenteric pairs of
the IIIrd Order (12-16) are being formed (see explanation in text) in all
the exoceeles.

arrangement of the muscle banners on the 12 mesenteries now
present is such as to group them into the six primary pairs
characteristic of hexactinian symmetry. This agrees with the

resul

(1. p

ts furnished by the latest stages investigated by Appellof
. 82).

456 DR. J. F. GEMMILL ON THE DEVELOPMENT OF

Next appears a pair of mesenteries in each lateral exocele (No.7
in text-figs.), and a little later another pair (No. 8 in text-figs.) in
each ventral exocele. These four new pairs, though they belong
to what may be called the 2nd developmental cycle, grow rapidly
and take rank with the six primary pairs, the result being that
ten pairs of mesenteries of the Ist Order are produced. Mean-
time a pair of additional mesenteries has begun to form in each
dorsal exoccele (No. 9 in text-figs.). These are really the last of
the 2nd developmental cycle, and their growth is slow. 3rd cycle
mesenteric pairs (Nos.10,11 in text-fig. 5) next form in the remain-
ing 8 exoceeles and growing relatively quickly, take rank with the
two pairs last named as mesenteries of the IInd Order. There
are now 20 exocceles, and in each of these a new mesenteric pair
(Nos. 12-16 in text-fig. 5) appears producing the 20 pairs of the
Iltrd Order. It will be seen that, properly speaking, four of
these pairs (Nos. 12-13) belong to the 3rd cycle, and the
remainder to the 4th cycle. There are now 40 exoceles and in
each of these a new mesenteric pair appears, 8 of these pairs
belonging to the 4th and the remainder to the 5th develop-
mental cycle, the whole forming the mesenteries of the LVth
Order.

In the text-figs. appended, the first twelve mesenteries and the
succeeding mesenteric pairs are numbered in the order of their
formation. It will be seen that the mesenteries of the IInd,
IIIrd, and [Vth Orders tend to appear earliest in the dorsally
placed exoceles. This is natural since the mesenteries of these
exoceeles belong to earlier cycles than those which apparently
correspond with them in the other exocceles.

Boveri (2. p. 496) conjectured that the decamerous symmetry
of Tealia arose by the formation of an additional pair of Ist Order
mesenteries in each dorsal and ventral primary exocele. He
based this conjecture on sections of an undetermined larva
obtained from the Naples Station. Faurot (5) states definitely
that in Teaha felina Linn. (= Tealia crassicornis Gosse, i. ¢., the
shore Urticina) the four additional Ist Order mesenteric pairs
arise in the dorsal and lateral primary exoceles. His description
and figures bear out this statement fully, but it is to be noted that
the youngest specimen he examined had already 28 tentacles.
Assuming him to be right, then, in spite of superficial resem-
blances, there is a deep-seated developmental difference between
Urticina coriacea and Urticina crassicornis which will certainly
make them take rank as distinct species. Material is being
collected for a revisal of his work on this point.

References.

1. AppELLOr, A.—‘“‘Studien iiber Aktinien Entwickelung.”
Bergens Museum, Aarbog, 1901, pp. 1-99,

are

THE MESENTERIES IN URTICINA CRASSICORNIS. 457

. Bovert, Ta.—‘‘ Ueber Entwickelung u. Verwandschaftsbezie-
hungen der Aktinien.” Zs. Wiss. Zool. xlix. 1889-90,
pp. 461-502 (see specially p. 496).

. Carueren, O.—“ Die Aktiniarien der Olga Expedition.” Wiss.
Meeresuntersuch. Bd. 11. Abth. Helgoland, 1912, pp. 33-35.
. Drxon, G. Y. & A. F.—“* Notes on Bunodes thallia, Bunodes
verrucosa, and Tealia crassicornis.” Scient. Proc. Roy.
Trish Acad. vi. 1889, p. 310.

. Fauror, L,—“ Etudes sur Jes Actinies.” Arch. Zool. Expér.
et Génér. i11. 1895, pp. 43-262 (see specially pp. 172-191).
. Gossz, P. H—A History of the British Sea-Anemones and
Corals. London, 1860.

. Happon, A. C.—‘*A Revision of the British Anemones.”
Trans. R. Dublin Soe. ser. 2, vol. iv. 1889, pp. 299-360.

ON THE CILIATION OF MELICERTIDIUM OCTOCOSTATUM. 459

32. The Ciliation of the Leptomedusan Melicertidium octo-
costatum (Sars). By James I’. Gemmitu, M.A., M.D.,
Dise

[Received August 18, 1919: Read November 18, 1919. ]
(Text-figure 6.)

Few hydromedusan gonophores have radial and ring canals
wide enough to allow the action of the ciliated lining of these
channels to be adequately studied. Melicertidium is an exception,
and now that the importance of primary functions in relation
to form is more and more recognised, the following details cannot
fail to be of interest. (The adult medusa has eight radial canals
along which the gonads are developed as eight somewhat promi-
nent and sinuous ridges projecting into the sub-umbrellar cavity.
See H. T. Browne, Proc. Roy. Soc. Edinb., 1905, p. 72, and
also a forthcoming paper by the author on the Life History of
Melicertidium.)

INTERNAL SURFACES.

. Stomach lining.—Currents ... . outwards or centrifugal from
middle point of roof, and inwards or centripetal (7. e.,
towards manubrial junction) along floor.

Lining of manubrial canal.—Currents .... weakly upwards
(from the mouth-opening) all round, even in the radial
grooves. This is different from what obtains in Aurelia
where the radial grooves serve as channels exhalant
from the stomach.

. Lining of radial canals.—Currents .... strong, centrifugal

(7. e., from stomach towards ring canal along roof (exum-
brellar surface) of each canal), and centripetal (7. e.,
towards stomach) along the floor of each canal.

4, Lining of ring canal.—Currents .... confused, causing mixing,

but on the whole there appears to be a flow clockwise

(as viewed from the aboral side) along the floor of the

canal over the openings into the tentacle cavities, and a

conyerse flow along the opposite wall of the canal. An

inward and an outward current may be noted on
opposite sides of the opening of each ipuiae's cavity
into the ring canal.

EXTERNAL SURFACES.

1. Hxumbrellar surface.—Ciliation absent.

2. Sub-wmbrellar surface.—(a) Along the projecting edges of the
gonads there is strong ciliation upwards (7. ¢., towards
the manubrial region) ; (6) along the sides of the gonads
the currents are towards the projecting edges but

460 DR. J. F. GEMMILL ON THE CILIATION OF THE

slanting upwards and thus feeding the eurrents noted
under (a); (¢) in the spaces between the gonads ciliation
is absent except towards the centre of the bell where it
is feebly upwards (7.e., towards the manubrial junc-
tion); (d) below the floor of the stomach the ciliation
is strong and inwards towards the manubrium, the
surface of which is likewise strongly ciliated in the
direction of the mouth-opening.

3. Tentacles. — The tentacles show weak ciliation from their
attached to their free ends, except on their inner sides
near their bases where they are ciliated more strongly
and in the opposite direction.

Text-figure 6.

Diagram (vertical section) to illustrate ciliation of Melicertidiwmn. The arrows
indicate the direction of the ciliary currents. See explanation in text.

1, Mouth-opening; 2, cavity of manubrium; 3, stomach ; 4, radial canal ;
5, ring canal; 6, tentacle.

We may judge that the ciliation of the gastrovascular lining
subserves in the first place the mixing and transportation of
food, and that it is also capable of aiding the ingestion of
small food particles and the evacuation of the sexual products
through the mouth. The ciliation of the sub-umbrellar surface
will gather small food particles towards the mouth-opening.

LEPTOMEDUSAN MELICERTIDIUM OCTOCOSTATUM. 461

Since the tentacles are often found inturned into the umbrellar
cavity, and even with their ends projecting into the mouth, we
may infer that their ciliation will also assist, however slightly
and intermittently, the important business of food gathering.

So far as I know, the only author who has investigated the
direction of the ciliary currents in hydroid meduse is Boehm
(Jen. Zs. Natw. xii. 1878, p. 108), and what he states in this
connection is that the currents are always from stomach to ring
canal, and that he has not been able to make out ciliary. activity
in the roof of the latter.

Reference may be made to the following recent papers on
ciliation in other mayine animals :—

CARLGREN, O.—Biol. Centralbl. xxv. 1905, pp. 308-322 (Acti-
nians, Madreporarians).

Orton, J. H.—Journ. Mar. Biol. Assoc. U.K. ix. 1912, pp. 144-
478 (Ascidians, Molluscs).

Orton, J. H.—J/bid. x. 1913, pp. 19-49 (Amphioxus, Ascidians,
Molluses).

GeEmMILL, J. F.—Proc. Zool. Soc. Lond. 1915, pp. 1-19 (Starfish).

Wipmark, H. M. P.—Zs. Allg. Phys. Jena, xv, 1913, pp. 33-48
(Aurelia aurita).

GemMILL, J. F.— Proc. Zool. Soc. Lond. 1919, pp. 263-265
(Ctenophore).

THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 463

EXHIBITIONS AND NOTICES.

October 21st, 1919.

Prof. EK. W. MacBripn, F.R.S., F.Z.8., Vice-President,

in the Chair.

The Secrerary read the following Reports on the Additions
made to the Society's Menagerie during the months of June, July,
August, and September, 1919 :—

JUNE.

The registered additions to the Society’s Menagerie during
the month of June were 355 in number. Of these 267 were
acquired by presentation, 11 were deposited, 7 were received in
exchange, 66 were purchased, and 4 were born in the Menagerie.

The following may be specially mentioned :—

2 Cheetahs (Cynelurus jubatus), from Berbera, Somaliland,
deposited on June 26th.

2 Pandas (Hlurus fulgens), born in the Menagerie on June
26th.

A collection of birds from Gambia, West Africa, consisting of
Fire- Finches, Quail-Finches, Weavers, and two Black-shouldered
Kites, presented by Dr. HE. Hopkinson, D.S.O., on June 19th.

2 Bateleur Kagles from Berbera, Somaliland, presented by
G. F. Archer, C.M.G., on June 26th.

2 Glaucous Gulls, hatched in the Menagerie on June 27th.

JULY.

The registered additions to the Society’s Menagerie during the
month of July were 343 in number. Of these 37 were acquired
by presentation, 32 were deposited, 5 were received in exchange,
238 were purchased, and 33 were born in the Menagerie.

The following may be specially mentioned :—

1 Hippopotamus (Hippopotamus amphibius), g, born in
Amsterdam, purchased on July 16th.

1 Yak (Bos grunmiens), 2 (Tibet), deposited on July Ist.

1 White-bearded Gnu (Connochetes albojubatus), born in the
Menagerie on July 30th.

1 Pere David’s Babbler (Pterorhinus davidi), from North China,
new to the Collection, deposited on July 31st.

2 Greenish Hangnests (Pseudoleistes virescens), and 1 Yellow
Troupial (Ageleus flavus), from Argentina, presented by M. Jean
Delacour on July 18th.

1 Ross’s Plantain-eater (/usophaga rosse), | Bare-faced Fruit-
Pigeon (Vinago calva), and 2 Grant’s Francolins (/rancolinus

464 THE SECRETARY ON ADDITIONS TO THE MENAGERIE.

granti), from British East Africa, presented by Dr. Van Someren
on July 22nd.

A collection of North-American Reptiles, including 2 Horned
Lizards (Phrynosoma cornutum), 2 Bull Snakes (Pitwophis say),
2 Testaceous Snakes (Zamenis flagelliformis), and 3 Western
Diamond Rattlesnakes (Crotalus atroa), purchased on July 28th.

AUGUST.

The registered additions to the Society’s Menagerie during the
month of August were 68 in number. Of these 29 were acquired
by presentation, 9 were deposited, 3 were purchased, and 27 were
born in the Menagerie.

The following may be specially mentioned :—

1 Leopard (Felis pardus), 2 , from Kismayu, British Somaliland,
presented by H.M.S. ‘Hyacinth’ on August 6th.

2 Young Indian Hlephants (Hlephas maximus), from Assam,
deposited on August 23rd and 25th.

1 Kilima-Njaro Sunbird (Wectarinia kilimensis), from Uganda,
new to the Collection, deposited on August 9th.

2 Hlephantine Tortoises (Zestudo elephantina), from the Sey-
chelles, presented by H.E. Lt.-Col. Sir Eustace Fiennes, Bt.

SEPTEMBER.

The registered additions to the Society’s Menagerie during the
month of September were 153 in number. Of these 33 were
acquired by presentation, 22 were deposited, 91 were purchased,
and 7 were bred in the Menagerie.

The following may be specially mentioned :—

1 Red-bellied Cercopitheque (Cercopithecus erythrogaster)
(Lagos), purchased on September 17th.

1 Black Mangabey (Cercocebus aterrimus) (Congo), purchased
on September 9th.

2 Red-cheeked Ibises (Comatibis eremita), from Birijik, Upper
Euphrates, new to the Collection, presented by Capt. E. H.
Buxton, on September 6th.

Mr. Oxupristp Tuomas, F.R.S., exhibited three interesting
Mammals obtained by Dr. Aders, F.Z.S., in Zanzibar: namely,
an example of Cephalophus adersi, » recently described new
species; an example of Colobus kirki, which until lately was
supposed to be almost extinct, and a specimen ofa rare Insectivore
belonging to the genus Petrodromus.

POSITION AND AFFINITIES OF 'TARSIUS. 465

Discussion on The Zoological Position and Affinities of Tarsius.

Dr. A.Smiru Woopwarp, F.R.S., in opening a “ Discussion on
the Zoological Position and Affinities of Tursius,” said :—

The small mammal Tarsius, though technically a lemur, ex-
hibits somany anatomical resemblances to the higher anthropoids
that it proves to be one of those solitary links between groups
which paleontologists welcome as “ living fossils.” It survives
only in the Philippines and in the Indo- “Malay an region, which
furnishes some other primitive mammals, among which may be
specially mentioned the ancestral ruminant Zragulus. Apart
from its enlarged eyes and its highly specialised jumping feet,
it might well be regarded as belonging to the earliest Tertiary
period, the Hocene.

Among Hocene fossils already known bork from Europe and
North America, there are numerous small jaws with a dentition
much resembling that of Zarsius. Some of these may belong to
prinitive ecceu on es, carnivores, and ungulates, which were not
well differentiated at the beginning of the Tertiary epoch; but
complete skulls of Anaptomorphus and Votharetus from the Lower
and Middle Eocene of North America, and others of Vecrolemur
from the Upper Hocene or Oligocene of France, are essentially
identical with the skull of Zarsius. The greater part of the
skeleton of Votharctus has also been discovered, and the hind foot
differs from that of Zarsias only in the shortness of the caleaneum
and navicular. <A survey of ali the known fragments of Eocene
lemurs suggests that they were generalised forms, from which
both the medeon lemurs and the anthropoids may have arisen.

For the latest information reference may be made to papers
by Dr. W. K. Gregory in Bull. Geol. Soc. America, vol. xxvi.
pp. 419-446 (1915), and Bull. Amer. Mus. Nat. Hist. vol. xxxv.

pp. 258-271 (1916).

Prof. G. Exiior Sarru, F.R.S.:—T'welve years agoin a letter to
‘Nature’ (May 2, 1907, Tol. 76, p. 7) I explained my reasons for
thinking that the analysis of the Primates which gave most
natural expression to the known facts involved the splitting up
of the Order into three subdivisions of subordinal rank, of which
the central Suborder includes Z'arsius alone of the living members,
in addition to the extinct Eocene families Anaptomorphide and
Microcheride. This mede of subdivision had been suggested by
Dr. Gadow in 1898.

My presidential address to Section H of the British Association
in 1912 was devoted, in the main, to the discussion of the part
played by the Tarsioidea in the Evolution of Man, and the dis-
cussion of the significance of the persistence in Man and Tarsius
of so many generalised features, which had been modified in
most other Primates, and even more profoundly in most mammals
of other Orders. Im that address I attempted to explain the
preservation of a generalised strueture, in association with excep-
tional efficiency, as a token of the fact hey Man’s ancestors were

466 ON THE ZOOLOGICAL POSITION

able to survive and maintain the plasticity of a generalised
structure and the functional adaptability that goes with it, in
virtue of the fact that they were cultivating their intellectual
powers rather than specialising for one particular kind of life.
Their nimbleness of mind and agility of action enabled them to
adapt themselves to a great variety of changing circumstances
without sacrificing the generalised structure of their limbs; so
that, when their opportunity came, they still had the mental and
bodily plasticity to take advantage of it, and acquire the dominant
position in the animal kingdom. The Lemurs and most Monkeys
sacrificed their chances of attaining such pre-eminence when they
adopted specialisations of structure and of habits to avert the
risk of extinction. Z'ursiws represents the phylum whose progress
was brought to a sudden stop in Hocene times by an over-develop-
ment of, and an exaggerated reliance upon, those specialisations
of vision and its cerebral instruments which were responsible for
the differentiation of the Tarsioidea from the Lemuroidea, and
gave impetus to those developments which produced the Anthro-
poidea from one of the Tarsioid families. But Zarsius was able
to escape extinction only by adopting the safe nocturnal habits
which also played a part in sparing the Lemuroidea.

The evidence upon which my own views were based was
primarily a detailed examination of the brain in the Primates,
the first results of which were submitted to the Linnean Society
on March 6, 1902*.

Inthat memoir I explained howconclusively the structure of the
brain demonstrates (@) that the Lemuroidea differ from mammals
of all other Orders in presenting cerebral features that are dis-
tinctive of the Primates; (6) that Zarsiws has a brain which, in
most respects, closely resembles, and is no better developed than,
that of such Lemuroids as the Galagine ; but (c) that it reveals
a reduction of the olfactory areas and an expansion and precocious
development of the visual cortex, which represent the commence-
ment of specialisations foreshadowing the emergence of distine-
tively Simian features.

This evidence is so precise and conclusive and its significance
so unmistakable that those who have attempted to exclude the
Lemuroidea from the Primates have perforce been driven to
repress all reference to the brain, the organ which above all
others has been responsible for raising the Primates to the rank
expressed in their Ordinal name, and most plainly gives expression
to the distinctive features of the Order. For the outstanding
characteristic of Man is the range of his intellectual abilities, and
of the Order to which he belongs the nature of the instrument—
the cerebral structure—which made possible the emergence of
such extensive powers of discrimination in one of its members.

In each of the mammalian Orders there is a distinctive mode
of folding of the cerebral cortex that is due to the relative

* “On the Morphology of the Brain in the Mammalia, with Special Reference to
that of the Lemurs, Recent and Extinct,” Trans. Linn. Soc. of London, 2nd Series,
Zoology, vol. viii. p. 319.

AND AFFINITIES OF TARSIUS. 467

development of the different physiological areas, and especially to
the rate and order of their growth during feetal life. The ground
plan of this pattern of the sulci was already laid down in the
._Hocene ancestors of most of the Orders; and it is so peculiar to
each of them as to afford a sure criterion of the right of any
creature to be regarded as a true member of an Order. The fact
that the Lemuroidea strictly conform in every respect to this
distinctive test of membership of the Primates is conclusive proof
of their right to be included in the Order.

Text-figure 1.

Perodicticus. Pithecia.
A comparison of the cerebral hemispheres of a Lemuroid (Perodicticus) anda
New-World Monkey (Pithecia)—natural size.

}’.— Inferior frontal sulcus. ‘’.—Superior temporal sulcus,
C.— Central sulcus. P.—Postecentral (intraparietal) sulcus.
S.—Sylvian fissure. L.—Lunate sulcus.

In the Primates it was the precocious expansion of the brain
and the simultaneous cultivation of the visual, auditory, tactile,
and motor areas of the cerebral cortex that first differentiated
the earliest Primates from all other mammals, and provided them
with the germs of the capabilities and the means of attaining the
supreme position expressed in the name of the Order.

The pattern formed by the cerebral sulci, equally in the
Lemuroidea and the Anthropoidea (see text-figure), is the direct
expression of the factors I have already mentioned. The pre-
cocious expansion of the visual cortex causes this area to become
folded along its axis, so that the major portion of the area striata
is bent in to form the retrocalcarine furrow in a way that is
peculiarly distinctive of the Primates, and is found equally in all
three Suborders*, but in no other mammal. The simultaneous
expansion of the tactile and auditory areas leads to the develop-
ment of the equally distinctive Sylvian fissure (text-figure, 8.),

* ] have discussed this at some length in my Linnean memoir (op. cit, supra).

Proc. Zoou. Soc.—1919, No. XXXIT. 32

468 ON THE ZOOLOGICAL POSITION

which is found in no other Order, except in a somewhat modified
form in certain Chiroptera and Edentata. This characteristic
fissure is found in all the Lemuroidea and Anthropoidea, but in
Tarsius the enormous size of the orbits stretches the brain and
tends to erase this characteristic fold of the cortex. The great
expansion of the sensory and motor areas is responsible for the
development of the central sulcus in Perodicticus (see text-fig., C)
and certain of the Indrisine; but in most of the Lemuroidea it
is represented only by two slight puckers or is absent altogether,
as in Z'arsius, the Hapalide, and some of the Cebidee (Aotus and
Callithriv). But this sulcus is so peculiarly distinctive of the
Primates that its occurrence in some of the Lemuroidea is of
the utmost value as an indication of the affinities of the Suborder.
The infevior frontal (text-figure, F.), superior temporal (T.),
lunate (L.), and orbital sulci complete the picture of the Primate
plan; and they not only reproduce in the Lemuroidea the same
form and grouping as in the Cebide (see text-figure), but are
subject to the same variations. Specially significant is the fact
that the postcentral (so-called “intraparietal”) suleus in Vycti-
cebus and some of the Indrisine shows a tendency to fuse with
the Sylvian fissure, which produces the same peculiar pattern
that is found among the Cebide in Chrysothrix, Aotus, Alouatta,
and at times in Lagothrix.

Everyone who is acquainted with the wide divergence between
the fissural pattern of the Primates and those of other Orders
cannot fail to be impressed with the completeness of the identity
between the Lemuroidea and the Anthropoidea in respect of
these features, and of its significance as positive evidence of
kinship.

But these superficial resemblances are merely the outward
expression of a deep-seated structural and functional identity
which demonstrates beyond any possibility of mistake that the
Lemurs are primitive members of the Order Primates. Working
in conjunction with the late Dr. Page May and Professor W. H.
Wilson, | made an experimental examination of the reactions of
the cerebral cortex in Loris and Lemur *, and not only confirmed
the conclusions suggested by the study of the morphology of the
cortex, but also obtained results which showed a differentiation
of cortically-controlled movements much superior to that of any
other mammals with the exception of the Anthropoidea. The
researches of Vogt subsequently (1906) confirmed our results.
But the evidence afforded by the minute structure of the brain is
even more decisive. In the ‘Transactions’ of this Society some
years ago (May 1908, Vol. xviii. p. 175) I referred to my histo-
logical studies of the Prosimian cerebral cortex and their general
significance. Later in the same year the account was published
of amuch more detailed examination of the Lemur’s cortex than
JT had made.

* See Report of the British Association, 1904, p. 760.

+ K. Brodmann, “Die cytoarchitektonische Cortexgliederung der Halbaffen,”
Journal fiir Psychologie und Neurologie, Bd. x. 1908, p. 287.

AND AFFINITIES OF TARSIUS. 469

Brodmann not only examined (in Vogt’s laboratory) the
microscopic structure of the Lemus cortex, but also made an
elaborate comparison between it and the cortex of the Apes and
a considerable series of other mammals*. He arrived at the
conclusion that in the arrangement of its cells and fibres the
Liemur’s cortex reveals ‘‘weitgehende Uebereinstimmung mit
den Primaten [2.e. Simi], in einzelnen jedoch Abweichungen
mancher cytoarchitektonischer Typen teils im Sinne einer
niederen, teils aber auch im Sinne einer héheren Differenzierung
aufweist.”

Brodmann did not examine the brain of Tarsius ; but my own
studies enable me to say that its visual cortex is not only more
extensive than that of the Lemuroidea, but also more highly
differentiated and more like that of the Apes. The other regions
of the cortex, however, conform much more closely to the
corresponding areas in the Lemuroidea, both in extent and in
structure, than to those of the Apes.

The Tarsioidea seem to have become differentiated from the
Lemuroidea by a reduction of the face, which permitted the much
fuller development of stereoscopic vision. This in turn stimulated
the higher specialisation of the visual cortex and provided the
guidance for the performance of movements of a much greater
skill and precision. It was the cultivation of these powers that
brought one branch of the Tarsioidea to Simian rank.

Not content with the recognition of the fact that the Apes
were derived from the Su Inoaselast Tarsioidea, some of those who
are taking part in this discussion want to promote Z'arsius to the
full status of an Ape. This is an unwarrantable claim. The
anatomy of every part of the body, and more particularly of the
brain, reveals the extent of the profound difference between
the Anthropoidea and the Tarsioidea. On the present occasion I
shall not attempt to enumerate all the criteria of a true Ape, and
shall refer only to one point (which, however, is of fundamental
importance as a distinctive feature between the Anthropoidea
and the other two Suborders of the Primates). In the true Apes
all the great cortical areas to which I have already referred
(visual, auditory, tactile, and motor) are very much larger and
more highly difter entiated than they are either in the Tarsioidea
or the Lemuroidea. But, in ae tieta, the relatively insignificant
parietal, temporal, and fr ontal ‘association areas” of the Pro-
simiew have undergone so much expansion that the brain of a real
monkey is, at lenge about three or four times the bulk of that of
a Tarsioid or Lemuroid of the same size; and these overgrown
territories have also become highly differ entiated and specialised.
In my Linnean memoir (1902), to which I have already referred,
the data for comparing the size of the brain in different mammalian
Orders has been set forth, so that I need refer here only to one
relevant point.

* Vergleich. Lokalisationslehre der Grosshirnrinde,’ 1909.
oe

470 ON THE ZOOLOGICAL POSITION

Comparing WVycticebus and Pithecia as representatives of the
Lemuroidea and Cebide of approximately the same size (each
weighing about 500 grm.), it is found that the brain of the
former varies in weight from 7:72 to 818 grm., whereas the
monkey’s brain ranges from 22 to 36:2 grm. In other words,
the lowly Platyrrhine has from three to four times as big a
brain as the Lemuroid. ‘T’o one who studies the meaning of the
size of the cerebral cortex in the different mammalian Orders,
and realises the significant rdle such expansion of the brain has
played, ever since “Eocene times, in the evolution of the higher
mammals, and especially of the Primates, it will be evident that
a vast chasm separates the monkeys from the Lemurs. Now
Eugen Dubois states * that the proportion of brain to body in
Tarsius i is not appreciably different from that of Nyeticebus. In
other words, judged by this fundamental test, Zarsius is sharply
differentiated from the Apes and occupies a rank not unlike that
of a Lemur. .As Cope pointed out more than thirty years ago, the
brain of Tarsius is not appreciably bigger than that of the Eocene
Anaptomorphus 7.

The ancestors of V'arsius, in fact, fell out of the race for
intellectual supremacy in Early Eocene times and ceased
cultivating their cerebral organs. Hventually, like the Lemurs,
they had to adopt nocturnal habits to avoid the risk of extine-
tion. At the same time another branch of the Tarsioids was
cultivating more highly skilled movements, and acquiring greatly
enhanced powers of discrimination and ability to profit from
experience. In course of time—and probably long before the
close of the Hocene--this particular family of big-brained
-Tarsioids was transformed into real monkeys and became the
ancestors of the Anthropoidea.

Before leaving this aspect of the problem, and while em phasising
the fact that so far as size of brain is concerned Jarsins is on the
same level as the Lemuroids, it 1s important to remember that at
the commencement of the Eocene period the representatives of
both the Lemuroidea and the Tarsioidea (and no doubt their
common Prosimian ancestors) were equally distinguished from
all other Orders by their relatively large brain. Both, in fact,
shared alike in the fundamental structural change that brought
the Primates into being. Even the modern Lemuroids, poorly
equipped as they are in brain-substance as compared with the
Apes, are better off in this respect than members of similar size
of any other Order. Thus in the Carnivora, which come next to
the Primates in respect of size of brain, it is found that in

+ E. D. Cope, “The Lemuroidea and the Insectivora of the Eocene Period of
North America,” American Naturalist, May 1885, p. 467.

Discussing the size of the cranial cavity of Anaptomorphus, he wrote :—<The
brain and its hemispher es are not at all smaller than those of Tarsius. . . . This is
important in view of the very small brains of the Hesh-eating and ungulate Mammalia
of the Eocene period as yet known. In conclusion, there is no doubt but that the
genus Anaptomorphus is the most Simian lemur yet discovered, and probably repre-
sents the family from which the anthropoid monkeys and men were derived.”

AND AFFINITIES OF TARSIUS, A471

animals of the same size as Lemurs the brain is about 1/100th
(or less) of the body-weight, whereas in Lemurs it is usually
about 1/70th, but may rise to 1/60th or more. But in a monkey
of similar size it is 1/20th or even 1/15th of the body-weight.
But we cannot on these grounds exclude the Lemuroidea from
the Primates, because the Tarsioidea are no better off than the
Lemurs so far as the quantity of brain is concerned.

The remarkable claim has been made that all the resemblances
between the Lemuroidea and the other Primates were due to the
fact that the former are primitive mammals and the Tarsioidea
and Anthropoidea generalised creatures that have preserved
many primitive features; or that they were the result of con-
vergence in animals leading similar modes of life. The former
statement can be ruled out of the argument at once, because the
brain of the Lemuroidea is definitely specialised in the manner I
have already described. Nor can the mode of life be regarded as
the explanation of the likenesses, because such arboreal animals
as the Tree-Shrews, Squirrels, Galeopithecus, et cetera, present
none of the numerous Primate features seen in the Lemur’s brain.
The fashionable and seriously overworked doctrine of convergence
is alsoa mere evasion of the real issue. There is abundant evidence
of convergence in the three Suborders of the Primates, but it is
clearly the expression of the tendency of similar traits to develop
in the various descendants of the same common ancestor, and
therefore can hardly help those who refuse to admit the close
connexion of the Lemuroidea with the other Primates. Henry
Fairfield Osborn has emphasised the fact that ‘‘the same results
appear independently in descendants of the same ancestors” *.
No more admirable illustrations of this principle could be found
than those elicited in the comparison of the Lemurs and Apes,
in the various individuals and species of which peculiar confor-
mations of brain, arrangements of muscles and arteries, form of
bones, structure of viscera and genital organs, which occur in no
other mammals, tend to reveal sahgranael nes in both the Lemurs
and the Apes. As illustrations of these striking demonstrations
I might refer to the centrai suleus of Perodicticus, the tendency
of the Sylvian fissure to fuse with the intraparietal, which is
found in Vycticebus and among the Apes in many of the Cebide ft.
Note also the striking likenesses in the temporal bone and the
course of the internal carotid in Zarsius and the Lorisiformes,
the tarsus of the Galagine and Tarsiws, and the similar variations
in the lacrymal region of the skull in Lemurs and Apes which
Dr. Forsyth Major has described in the ‘ Proceedings’ of this
Society. These are, for the most part, illustrations of similar
peculiarities developed independently in divergent descendants
of the same common ancestor.

* “The Four Inseparable Factors of Evolution.” Science, N.S., vol. xxvii.
January 24, 1908, p. 150.

+ See my account in the ‘Catalogue of the Museum of the Royal College of
Surgeons,’

472 ON THE ZOOLOGICAL POSITION

Ever since the time of Burmeister (1846) everyone who has
studied Tarsius has admitted its peculiarly distinctive position,
which Burmeister himself described so clearly when he claimed
it as a connecting-link between the Lemurs and the Apes
(‘‘Uebergangslied, wenn auch gerade nicht eins der auffallendsten
und merkwiirdigsten ”). He clearly recognised its generalised
character and its resemblances both to the Insectivora and to the
higher Primates, and epigrammatically summarised his con-
clusions in these words :—“ Aber Tarsius ist nicht mal ein Affe,
er ist vie'mehr nur ein Halbaffe”*. It is generally admitted
that Tarsius is a remarkably generalised creature which in many
respects is akin to the Menotyphlous Insectivora, and has per-
sisted with extraordinarily little change from the beginning of
the Eocene period. But it is also recognised that every part
of its anatomy, brain and skull (including the developmental
history of the skull +), face and rhinarium, muscles and viscera,
genitalia and mode of placentation, reveals its affinity to the
higher Primates and affords evidence of its differentiation from
the Lemuroidea.

Hence we are concerned in this discussion, not so much with
the facts of the case, which are admitted, as with the right
perspective in which they should be viewed, and their significance
estimated and expressed in classification. This cannot be done
merely by enumerating lists of differences between Tarsius and
the Lemurs or resemblances of the former to the Apes, especially
if all the evidence that lends support to the reality of the kinship
of the Lemurs and the other Primates, and that indicates the
lowly rank of Zarsius, is suppressed. By means of such methods
of special pleading a case might be stated for the view that the
whale was a fish, if care were taken to omit all reference to
the mammalian characters of the Cetacea. Yet the facts that
establish the right of the Lemurs to be regarded as Primates are
no less definite than those that make the whalea mammal. It
reveals a singular lack of logic to exclude the Lemurs from the
Primates because they are not Tarsii. One does not deny the
rank of Carnivora to dogs because they are not bears !

In 1830 Wagler claimed that the Lemurs should be put into
an Order (Lemures) distinct from the Apes (Simiz). Then in
1846 Burmeister with his wider knowledge and clearer insight
restored the true perspective, as I have already explained. But
since then a vast literature has grown up as the result of the
repeated reopening of these old controversies. Wagler has had
many followers, such as Gratiolet, Gervais, Milne-Edwards,

** H. Burmeister, ‘ Beitrage zur Kenntniss der Gattung Tarsius, 1846, p. v1.

+ Eugen Fischer, “On the Primordial Cranium of Tarsius spectrum,’ Konink-
lijke Akademie van Wetenschappen te Amsterdam, Proceedings of November 22,
1905, p. 400, “ exceedingly close relationship of the developing cranium of Tarsius
and that of the ape and man” and “ the striking resemblance between this type of
skull and that of reptiles,”

AND AFFINITIES OF TARSIUS. 473

Hubrecht, and Max Weber, in his attempt to remove the Lemurs
into an. Order apart from the other Primates: but the fallacy of
the arguments brought forward in support of such views has
been repeatedly exposed *. In view of the much more extensive
and precise knowledge of comparative anatomy, embryology, and
paleontology that is now available, these claims to exclude the
Lemuroidea from the Primates have even less justification than
in the past, and can only be given a plausible appearance by
special pleading of a desperately biassed kind.

The zoological rank and affinities of no mammal are more
precisely determined than are those of Zarsius. The evidence
presented in Filhol’s memoir‘, published forty-five years ago,
clearly demonstrated that the Hocene ancestors of the Lemurs
were closely akin to the contemporary forerunners of the
Tarsioids, and mention has already been made of Cope’s recog-
nition (in 1885), not only of the derivation of the Apes from
the Anaptomorphide, but also of the human likenesses which
he expressed by giving the specific name homunculus to a
Tetonius. But any lingering doubts on this subject have been
dispelled by the recent papers published by members of the staff
of the American Museum of Natural History =.

The evidence of anatomy and paleontology is thus unanimous
in support of the right of the Lemuroidea to be included in the
Primates. Nevertheless, the cleavage between Zarsius and the
Lemuroidea is so great as to establish the right to a separate
Subordinal rank for the former. The fact that its ancestors parted
company with those of the Lemuroidea so early as Paleocene or
even perhaps Cretaceous times affords strong corroboration of
the claim to put it into a distinct Suborder Tarsioidea. I have
already referred to the unjustifiable claim that the Tarsioids are
already monkeys and ought to be put into the Suborder Anthro-
poidea. Those who argue in this illogical way might perhaps
appreciate the fallacy anderlyine their claims by studying an
analogous case. The Cynodonts are a group of very primitive

* As, for example, by
Sir William Turner, “On the Placentation of the Lemurs,’ Philosophical
Transactions of the Royal Society, vol. 166, pt. 2, 1876, p. 569 (who gives
the earlier bibliography),
and

Charles Earle, “The Lemurs as Ancestors of the Apes,” Natural Science, vol. x.,
May 1897, p. 309:

_also “On the Affinities of Tarsius: a Contribution to the Phylogeny of the
Primates,” American Naturalist, vol. xxxi., July 1897, p. 569, and August
1897, p. 680.

+ H. Filhol, “Nouvelles observations sur les Mammiféres des Gisements de
Phosphates de Chaux,” Annales des Sciences Géologiques, T. v. Pl. 7 (1874).

* W. D. Matthew and Walter Granger, “A Revision of the Lower Kocene
Wasateh and Wind River Faunas,” Bull. American Museum of Natural History,
vol. xxxiv. (1915). William K. Gregory, “On the Relationship of the Kocene Lemur
Notharctus to the Adapide and the other Primates” and “On the Classification
and Phylogeny of the Lemuroidea,” Bulletin of the Geological Society of America,
vol. xxvi., Nov. 1915, p. 419; and “Studies on the Evolution of the Primates,”
Bulletin of the American Museum of Natural History, vol, xxxv., June 1916, p. 289.

474 ON THE ZOOLOGICAL POSITION
fossil reptiles which are distinguished from all other reptiles
(including all the surviving members of their Class) by a large
number of features which they share with Mammals. Moreover,
it is now generally admitted that the Mammalia were derived from
one of the Cynodont families. Yet no paleontologist, so far as
IT am aware, has been reckless enough to suggest that the
Cynodonts should be removed from the Class Reptilia and pro-
moted to the Class Mammalia. But those who want to remove
Tarsius from it slowly station and elevate it from Prosimian
to Simian rank are making a claim that is as preposterous as
the hypothetical parallel just suggested. In some respects the
analogy may seem inconsistent with the course I have adopted
here. It may perhaps be argued that, just as the Cynodonts are
still retained in the Class Reptilia, so the Tarsioids ought to be
retained in the Suborder Lemuroidea. Matthew and Gregory
have, in fact, recently (op. cit. swpra) reaftirmed their belief in
this traditional method of subdividing the Primates; and there
is unquestionably a goed deal to be said for preserving the term
Prosimie. But the ‘problem differs from that of the Cynodonts
in that we are now discussing the grouping of the constituent
parts of a single Mammalian Order, whereas the Cynodont
problem involves the interrelationships of two Vertebrate Classes.
The depth of the cleavage between the Tarsioids and the
Lemuroids and the remarkable affinities of the former to the
Anthropoidea signifies that the Tarsioidea occupy a position
definitely intermediate between the other two groups of Primates,
which finds most natural and most convenient expression if
a separate Suborder is made to include the Tarsioids. This is no
mere compromise between the extremists on the two sides, but
the arrangement which the whole trend of research since the time
of Burmeister has made more and more insistent and necessary.
If the exact status and affinities of the Tarsioidea as Kocene
Prosimie and their little-altered survivors has been definitely
established, the recent discoveries * of the Early Oligocene genera
Par. apithecus and Propliopithecus in the Egyptian Faytim have
no less definitely settled the relationships of the Suborder to the
Anthropoidea. Parapithecus retains sufficient of the primitive
traits to establish the truth of the Tarsioid ancestry of the Apes,
but it also provides evidence which can only be adequately
explained on the supposition that the transformation of a
Tarsioid into a primitive Ape must have occurred before the
close of the Eocene. For in the lowest Oligocene the primitive
ape Parapithecus 1s found in association with Propliopithecus,
which had developed far beyond the stage represented by the
former and become a real Anthropoid Ape. Parapithecus, then,
at the beginning of the Oligocene, must have been a survival from
* M. Schlosser, “ Beitriige zur Kenntniss der Oligozéinen Landsiugetiere aus dem

Fayum (Agypten),” Beitrage zur Pal, u, Geol. Osterreich-Ungarns u. d. Orients,
Bd, xxiv, 1911, p. 52, ;

AND AFFINITIES OF TARSIUS. 475

a still earlier period, 7. e. there were real monkeys in the Eocene.
T do not suppose that anyone will refuse to admit the certainty
of the derivation of the tailless Anthropomorpha from tailed
Catarhines; but the early development of true Simide shows
that the differentiation of the Old-World Apes into Cerco-
pithecide and Simiide must have occurred almost immediately
after the Catarhines themselves came into being.

Anyone who conscientiously investigates the anatomy of the
Platyrrhine Apes, and attempts to interpret the vastly com-
plicated series of cerebral transformations that were necessary to
convert a Tarsioid into a monkey, must be forced to admit that
this did not happen twice, but that the Platyrrhines and the
Catarhines were derived from a common stock, some archaic Ape
more primitive and more Tarsioid even than Parapithecus, and at
a time long before the close of the Eocene period. The history
of the ancestors of the, Anthropoidea thus becomes clear.

In North America (which was clearly the home of the Order
Primates) the Lemuroidea and the Tarsioidea were differentiated
from the ancestral Primate probably at the close of the Creta-
ceous period. At some time during the Eocene (and somewhere
in the neighbourhood of America) true monkeys were differ-
entiated from one of the Tarsioid groups. Some of these found
an asylum in South America and became specialised as the
Platyrrhines. But others (in Eocene times) made their way
to the Old World along with the Adapid and Anaptomorphid
ancestors of the Lemuroidea and Tarsioidea respectively. During
this migration these primitive monkeys became transformed into
Catarhines, and the remains of Parapithecus provide the evidence
of their reality and give a hint as to their size and distinctive
features.

There is one other aspect of the problem under discussion
which has been fruitful of much misunderstanding, The
Lemuroidea represent a lower stratum of Primate evolution
than the Tarsioidea, just as the latter are on a very much lower
plane than the Anthropoidea. But, while the Lemuroidea retain
many features of brain, skull, face, placentation, et cetera, which
are survivals from their Paleocene or Cretaceous ancestry—the
earliest Primates,—during the long span of time that has elapsed
since the Cretaceous period they have acquired a host of minor
specialisations of structure which have modified or masked much
of their original likeness to the other Primates. T'arsius,
however, although on a distinctly higher plane of Primate
development, has managed to escape extinction with fewer and
slighter specialisations than the Lemurs. Hence it has retained
a mueh more generalised and obviously primitive structure along
with the germs of the features that are distinctive of monkeys.

475 ON THE ZOOLOGICAL POSITION

Prof. J. P. Hitt, FVR.S.

The Affinities of Zarsiws from the Embryological Aspect.
(With Table, Plate I., and Text-figures 1-5.)

My task in this discussion * is to consider what Jight the facts
of development throw on the question of the affinities of Z'arsius.
In furtherance of that object, [ have thought it might be both
useful and time-saving if I presented you with a brief summary,
in the form of the accompanying comparative table (v. p. 490), of
what I take to be the most important of the known facts relating
to the development, foetal membranes, and placentation of the
main groups of the Primates. For the purposes of this discussion,
I have set forth the facts relating to Tarsius in column 2 of the
table, in order that you may the more readily compare them with
those appertaining on the one hand to the Lemuroids (Lemuri-
formes and Lorisiformes) in column 1, and on the other to the
Anthropoids in column 3. Jt remains to be seen in how far
this tripartite mode of grouping the Primates is justifiable on
embryological grounds.

The first question which arises is that of the systematic position
and affinities of the Lemuroids. It is generally agreed that the
Lemuroids are a lowly and in many respects primitive group, and
even Hubrecht admitted that they ‘are in no respect a very
specialised order of Mammals.” The prevailing view, widely
held both by comparative anatomists and paleontologists, is that
they lie at the base of the Primate series; but certain authorities,
notably Hubrecht, deny that they are in any way related to the
other Primates. That is a view which, on embryological grounds
alone, | am unable to accept.

Unfortunately our knowledge of the development of the
Lemuroids is very fragmentary, but what we do know shows,
in my opinion, perfectly clearly that the existing forms are
no such forlorn and degenerate creatures as some would have us
believe, but, on the contrary, are to be regarded as the repre-
sentatives of a very old and primitive group of Mammals from
which the other and higher Primates may very well have taken
their origin. In their simple central type of development (the
blastocyst developing free in the uterine lumen), in their mode
of amnion formation (the amnion developing from folds in the
presence of a proamnion), and in the mode of development and
the relations generally of their fcetal membranes (in particular,
in the presence of a vesicular allantois, which grows out as a free

* Since the date of the discussion, I have had the opportunity, thanks to the
great kindness of Dr. Dan. de Lange, Junr., Director of the International Institute
ot Embryology at Utrecht, of examining a uterus of Tarsius, containing a nearly
full-term foetus with its placenta and of preparing sections of the latter. I wish
here to express my most cordial thanks to Dr. de Lange for his generosity in enabling
me to examine for myself this rare and valuable material. Its study has led to some
modification of the views I expressed at the meeting.

185 4, So WO; ISN0LIL, Pil I.

~ak cf.
% ees
p) KS ays

ye F
Se

Sea

Bea

ext

PLACENTATION OF TARSIUS.

AND AFFINITIES OF TARSIUS. AGT

vesicle and subsequently unites with and vascularises the entire
chorion), we see features all of which we are familiar with in the
development of the lower members of the Mammalian series, and
all of which are, in my opinion, primitive.
Moreover, the primary relations of their foetal membranes
are precisely those which we are justified in postulating for
the primitive ancestral stock from which presumably both the
Didelphia and Monodelphia diverged; whilst their simple,
diffuse, non-deciduate placenta (involvi ing the loose interlocking

Text-figure 1.

all. ch.

~omph.

Nycticebus tardigradus. Diagram (after Hubrecht) to show the arrangement of
the foetal membranes of the early embryo. Note especially the allantois (a/7.)
already fused with the chorion (ch.) to form a discoidal area of allanto-chorion
(all.ch.). The arrows indicate the direction of extension of the extra-embryonal
ceelom (ex.cc.) into the mesoderm of the omphalopleure (omph.) so as to
separate that into yolk-sac wall and chorion. amn. amnion. y-s.c. yolk-sac
cavity.

BEetoderm (including trophoblast) is represented by a thin line, entoderm by
a thick line, and mesoderm by a dotted line.

of short villous outgrowths of the allanto-chorion with corre-
sponding crypts of the uterine mucosa, the persistence of the
uterine epithelium and of the uterine glands) is, In my view,
essentially primitive, presenting us with a simple little spe-
cialised type of placenta from which the much more advanced,
and presumably more efficient, arrangements in the other

47 ,
18 ON THE ZOOLOGICAL POSITION

Primates may quite easily have originated as the result of
adaptive specialisation in the course of evolution. Hubrecht,
however, took an entirely opposite view and held that the
placentation of these forms is not genuinely primitive, but had
arisen from some hypothetical, more complicated form of
placenta as the result of secondary reduction and degeneration ;
and Assheton, inclining to the same point of view, suggested
that it might have been derived from a primitive Carnivore
type, also by reduction. For these views, expressed in con-
nection with particular theories of placental evolution, there is,
so far as i am aware, not the slightest direct evidence.

On the other hand, the evidence derivable from a study of
the development of the feetal membranes seems to me to provide
us with a perfectly definite lead. Hubrecht’s observations on
Nycticebus show that the allantois grows out into the extra-
embryonal ccelom as a small stalked vesicle, just as it does in all
primitive Mammals. It rapidly increases in size and already in
the embryo of 4:2 mm. (G. L.) has extended over and fused with
the discoidal area of chorion, which is thus transformed into
vascular allanto-chorion (text-fig. 1, all.ch.). In later stages,
following the splitting of the extra-embryonal mesoderm, it
rapidly spreads and the chorion, over its entire extent, is
converted into vascular allanto-chorion. With this rapid and
very marked growth of the allantois are to be correlated
two other occurrences to which I would direct attention:
(1) the extra-embryonal ceelom rapidly extends throughout the
entire extent of the mesoderm of the blastocyst wall or omphalo-
pleure, which thus becomes split into chorion and yolk-sae wall ;
(2) the yolk-sac becomes established as an independent vesicle,
and the yolk-sac placenta, if such temporarily exists in con-
nection with the early embryo, becomes completely replaced by
the allantoic. .

These development features in the Lemuroids—viz., the
establishment of a complete chorion and its early vascularisation
by the rapidly growing allantois, the early formation of an
extensive extra-embryonal coelom and the separation of the
yolk-sac as an independent vesicle—seem to me to foreshadow
in the most unmistakable way corresponding and_ highly
characteristic events in the early development of both Zarsius
and the Anthropoidea. We have only to suppose these onto-
genetic happenings in the Lemuroids telescoped into still earlier
stages as the result of developmental adaptation in order to
reach the structural conditions characteristic of the early
blastocysts of the other Primates, and for such adaptations
to become perfected, ample time has been available, seeing
that the Lemuroids and the Tarsioids were already well
differentiated from each other in the Lower Kocene.

From the standpoint of embryology, I would accordingly
range myself with those who, on comparative anatomical and
paleontological grounds, see in the existing Lemuroids the

AND AFFINITIES OF TARSIUS, AT9

representatives of the stock from which the higher Primates
originated.

Turning now to Zarsius itself, our knowledge of its deve-
lopment may be said to be fairly complete, thanks to the
untiring labours of that illustrious embryologist, the late
Professor Hubrecht. In a series of papers he has provided us
with richly illustrated accounts of its early development, its
foetal membranes and placentation, and he has discussed its
attinities at some length. As the result of his investigations,
Hubrecht came to the conclusion that Varsius is no Lemur,
but is more nearly related to the Anthropoidea and should be
classified with them.

And there can be no doubt at all that in many features
of its development Tarsiws does appear to be more closely
related to the Anthropoids than to the Lemuroids. That much
is obvious from the data set forth in the table, but there are
also certain differences to be noted, and our task is to try and
evaluate these resemblances and differences in terms of aftinity.

The principal developmental features in which Zarsius differs
from the Lemuroids and agrees with the Anthropoids may be
vecapitulated as follows:—(1) the early attachment of the
blastocyst to the uterine lining through the proliferative activity
of the ‘trophoblast ; (2) the precocious formation of the extra-
embryonal mesoderm and celom and the correlated early
separation of the yolk-sac as a small reduced structure ; (3) the
functional replacement of the vesicular allantois by the so-called
connecting or ventral stalk : and (4) the formation of a massive
cdeciduate ‘placenta i in which the maternal blood circulates through
lacunar spaces in the syneytial trophoblast.

Set down in this bald fashion, these striking resemblances
between Zarsius and the Anthropoids would seem to settle
the question of its affinities without more ado; but let us
examine them a little more closely.

The attachment of the Varsius blastocyst to the mien wall,
at a time when it measures only 0-3 mm. in diameter, is effected
by the activity of the trophoblast over a localised patch situated
immediately behind the posterior margin of the embryonal area
(text-fig. 2, pl.tr.). All that we can say in regard to this attach-
ment is that it represents a definite advance of an adaptive nature
on the Lemuroid condition. It is the necessary preliminary to the
formation of the discoidal deciduate placenta, and it is a point
of interest that the attachment is effected immediately behind
the embryonal area and so in proximity to the region where
the allantois normally develops, although later on the definitive
placenta, as the result of differential growth in the fetal
membranes, comes to lie opposite the embryo. And it may
also be regarded as the first step in the evolution of the relations
of the Anthropoid blastocyst to the uterus, which range from the
same primitive central type of development that is seen in
Tarsius, with in some cases an additional second attachment

480 ON THE ZOOLOGICAL POSITION

to the uterine wall (as in the Old- and New-World Monkeys), to
the interstitial mode of development in which the blastocyst,
whilst still quite minute, burrows its way, through the agency
of its trophoblastic covering, into the uterine decidua and so
becomes completely imbedded, its trophoblast proliferating over
its entire extent (as in Man and the Anthropoid Apes). Thus
the relatively simple attachment in Zarsius points the way
to the much more highly specialised Anthropoid condition.

Then, in respect of the very early formation of the extra-
embryonal mesoderm and celom, Zarsius exhibits marked
adaptive specialisation as compared with the Lemuroids,
and this same specialisation again reaches its acme in the
Anthropoids.

Text-figure 2.

Tursius spectrum. TWiagram (after Hubrecht) to show the structure of the early
blastocyst, shortly after attachment to the uterine wall. Note the connecting
stalk (¢.s¢.) already present in the form of a band of mesoderm extending from
the posterior end of the ectoderm of the embryonal shield (embr.ecto.) to the
margin of the attached area of placental trophoblast (pl.é7.), the yolk-sac
(y-s.c.) not yet free and the extensive extra-embryonal cwlom (¢2.cw.).
ch. chorion.

Whereas in the Lemuroids, the extra-embryonal mesoderm
would appear to be formed, like that of the lower Mammals,
simply by the gradual peripheral extension of the embryonal
into the bilaminar wall of the blastocyst, here, in Varsius,
it is formed precociously, long before the embryonal mesoderm
has made its appearance, as a cellular mass proliferated, in

AND AFFINITIES OF TARSIUS. 481

the middle line, from the hinder margin of the embryonal
ectoderm. Hubrecht compared it with the ventral mesoderm
of the Amphibia, but, without entering into that question,
what I want to suggest now is that it is none other than a
precociously formed part of the primitive streak mesoderm *—

Text-figure 3.

embr. eclo

Tarsius spectrum. Diagram (after Hubrecht) to show the arrangement of the fcetal
membranes, prior to the closure of the amnion. Note in particular, the yolk-
sac (y-s.c.) now established as an independent vesicle; the connecting stalk
(c.sé.) ito the proximal half of which extends the allantoic duct (a/l.d.}; the
placenta (P/.); the thick layer of loose mesoderm (mes.) into which the extra-
embryonal coclom does not extend with the result that the distal portion of the
connecting stalk does not become separated from the chorion; and the head
and tail-folds of the amnion (hd-fl.amn. and tl-fl.amn.). The short oblique
canal perforating the embryonal area (em67".ecfo.) is the neurenteric canal, and
behind it is the primitive streak region, the mesoderm of which is directly
continued into the connecting stalk (c.sé.).

a point of considerable interest, to which I refer again later.
The cellular mass, so formed, extends downwards and_back-
wards in contact with the inner surface of the. attaching area
of trophoblast, and in all but its proximal attached part it

* Prof, T. H. Bryce, in 1908, was, I find, the first to put forward the view that this
mesoderm in Tarsius is to be considered as the equivalent of the mesoderm which
is proliferated from the posterior end of the primitive streak in lower Mammals.

482 ON THE ZOOLOGICAL POSITION

becomes hollowed out by the appearance in its interior of a
cavity, the extra-embryonal celom. The entodermal yolk-sac
earlier established only partially fills the space enclosed by the
trophoblast, being in contact with the latter only in front (text-
fig. 2, y-s.c.). The ceelom now rapidly expands so as to fill this
pace. and it also extends forwards into the front wall of the
blastocyst, separating the yolk-sac entoderm from the trophoblast,
with the result that the yolk-sac becomes provided with an
independent wall of its own and projects into the celom as a
small free vesicle, whilst the chorion is completed as a continuous
membrane, which forms the outer wall of the embryonal for-
mation (text-figs. 3 & 4, y-s.c. & ch.).

Thus in the mode of development of the extra-embryonal
mesoderm and cceelom, in the precocious formation of the yolk-
sac, and in the early differentiation of the chorion, Varsius is
much more specialised than the Lemuroid—nevertheless, it still
retains in its ontogeny evident traces of the ancestral mode of
development of these structures.

The proximal part of the extra-embryonal mesoderm in Zarsius,
referred to above, into which the extra-embryonal ccelom does not
extend, persists in the form of a solid, short, axial strand which
directly connects the hinder margin of the embryonal ectoderm
with the region of the chorion over which the placental tro-
phoblastic attachment has already been effected. This strand,
Hubrecht regards as the primordium of the ventral or connecting
stalk, the significance of which we shall presently discuss.

Coming now to the Anthropoids, although we know com-
paratively little of the details of their early development, it is
quite clear from our knowledge of the structure of the early
blastocyst that their early ontogeny is much more specialised
than that of Zarsius.

'The earliest-known blastocysts are already either attached to,
or actually embedded in, the uterine decidua, the trophoblast has
proliferated to form a syncytial network, into ‘the meshes of which
maternal blood has penetrated. Inside the trophoblastic wall
there is already present a layer of extra-embiryonal mesoderm
(i. e., the chorion is established). This layer thickens at the upper
pole. to enclose the embryonal primordium proper, in the form
of two closed vesicles—an upper, the amnio-embryonal vesicle,
and a lower, the entodermal yolk-sac. Here the embryonal
ectoderm which forms the floor of the amnio-embryonal vesicle
never becomes exposed on the surface as it does in Varsius, and
the cavity of the vesicle, the primitive amniotic cavity, persists
to form the cavity of the definitive amnion, the amnion arising
by the closed method and not by the closing i in of folds as in
Tarsius and the Lemuroids. The entodermal yolk-sac is most
preeociously differentiated as a small closed vesicle, and appa-
rently from the first lies remote from the trophoblast. The
extra-embryonal mesoderm is also most precociously developed,
but as to its mode of origin we have no knowledge. It is present

AND AFFINITIES OF TARSIUS. 483

long before there is any trace of a connecting stalk, from which
we may conclude that it does not arise like that of Varsius. It
is possible that it takes origin as a diffuse proliferation from the
marginal ectoderm of the amnio-embryonal vesicle, as is said to
be the case in Galeopithecus and T'atusia, a mode of origin which
assuredly is purely secondary and adaptive.

In respect, then, of these developmental occurrences, it is
evident, I think, that Yarsiws provides the intermediate link
between the primitive Lemuroids and the highly specialised
Anthropoids.

We may pass on now to the consideration of the two features
in the development of Zarsiws to which Hubrecht attached most

Text-figure 4.

amn.cl

Tarsius spectrum. Diagram (after Hubrecht) to show the relations of the foetal
membranes after closure of the amnion (amn.). Note the temporary con-
nection between the amnion and the chorion, marking the last point of closure
(amn.cl.). Other reference-letters as in text-fig. 3.

importance as guides to its affinity, viz., the occurrence of a
connecting or ventral stalk and the presence of a massive
placenta of the hemochorial deciduate type. When Hubrecht
first put forward his views, the existence of a connecting stalk
outside Zarsius and the Anthropoids was unknown, and so he
naturally attached great importance to it as a token of affinity,
but we now know, through the researches of Newman and

Proc. Zoou. Soc.—1919, No. XX XIIL 33

484 ON THE ZOOLOGICAL POSITION

Patterson, that a genuine connecting stalk, presenting a re-
markable similarity to that of Yarsiws, is also present in the
Armadillo (Vatusia novemcincta), in which obviously it must have
been evolved quite independently of that of the Primates.
Nevertheless, the common occurrence of this structure in these
two groups of Primates is a feature of very great interest, and
the first question we have to consider is its functional and
morphological significance. When fully established, it consists
of a strand of mesoderm, into which there extends a tubular
diverticulum (known as the allantoic duct) from the hind-gut of
the embryo, and it serves to connect the posterior end of the latter
directly with the chorion (text-figs. 4 & 5, e.st.). It is simply
a mesodermal short-cut between the embryo and the enclosing
chorion, and its function is to facilitate the early vascularisation
of that membrane by furnishing a direct path for the umbilical
(allantoic) vessels which form the essential feetal constituent of
the definitive or allantoic placenta. In the lower Mammals the
chorion is vascularised as the result of the secondary union with
it of the vesicular allantois, which likewise carries the umbilical
vessels. Prior, however, to the establishment of the functional
allantoic placenta, the nutrition and respiration of the embryo in
the lower Mammals are provided for by means of a temporary
omphalopleural or so-called yolk-sac placenta, involving the
vitelline or yolk-sac vessels of the omphalopleure or primitive
blastocyst-wall.

Now, in the Lemuvoids, as we have seen, the allantois unites
with the chorion relatively early, and the entire omphalo-
pleure is rapidly resolved, through the extension of the extra-
embryonal ccelom, into chorion and yolk-sac wall, with the
result that a yolk-sac placenta, if it exists at all, is of quite
transitory duration. In Yarsiws and the Anthropoids, owing to
the much earlier differentiation of the entodermal yolk-sac and
the chorion, a yolk-sac placenta cannot be formed at all, and so as
a compensation what appears to have happened in these forms is
that the allantois, and more particularly the vessel-carrying
allantoic mesoderm, became precociously developed in the form
of a solid cord, running directly from the hinder end of the
embryo to the attached area of chorion, marking the site of the
future placenta.

The entodermal lining of the allantois at the same time under-
went reduction, and now appears in the form of a diverticulum,
usually slender and tubular, which runs from the hind-gut for a
longer or shorter distance into the mesoderm of the cord.

This entire structure, then, is the connecting stalk, and what
I want to insist on is that it is not something new nor is it a
primitive formation (as Hubrecht maintained). It is none other
than a precociously formed and adaptively specialised allantois,
the object of which is to provide for the early and direct vaseu-
larisation of the chorion—or, in other words, for the nutrition and
respiration of the embryo at the earliest possible moment. That

AND AFFINITIES OF TARSIUS. 485

this view of its significance 1s correct is clearly shown by its mode
of development in Tutusia, and by its occurrence in the Marmoset
(Hapale) in a condition which can best be described as semi-
vesicular. In Yarsius, its primordium is constituted at first,
according to Hubrecht, simply by the unspht proximal portion
of the axial mesoderm proliferated from the hinder margin of
the embryonal ectoderm; if we are justified in regarding this
mesoderm as precociously formed primitive streak mesoderm,
then the connecting stalk of Z’arsiws can also be brought into line
with what we know of the development of the allantois in lower
Mammals. The predisposing factor in the evolution of the
connecting stalk in Zatusia and Tarsius is doubtless to be sought
in the placental attachment which is early established by the
trophoblast situated just behind the embryonal area and in
immediate proximity to the normal seat of origin of the allantois.

As to the development of the connecting stalk in the Anthro-
poids, we have no certain knowledge. It seems probable, however,

Text-figure 5.

Diagram to show the structure of the early blastocyst in the Anthropoidea. Note
the embryonal ectoderm (cross-lined) forming the floor of the amnio-embryonal
vesicle (amm. amnion), and the underlying yolk-sac vesicle (y-s.c.); the eon-
necting stalk (c.sé.) with the allantoic duct (al/.d.); and the extensive extra-
embryonal ccelom (ex.cw.) bounded by the chorion (ch.).

that it does not arise in the same way as in Jarsius, but that, to
begin with, it 1s simply formed by the persistent hinder portion
of the layer of extra-embryonal mesoderm which in the earl

blastocyst lies between the amnio-embryonal vesicle and the
trophoblast. Through the extension of the cclom, this layer
would seem to become split into chorionic mesoderm on the
outside and amniotic mesoderm on the inside, except posteriorly,

3a”

486 ON THE ZOOLOGICAL POSITION

where it apparently persists as a solid strand, connecting the
embryonal formation directly with the chorion (text-fig. 5, c.st.).
Even if this strand is reinforced later by mesoderm of primitive
streak origin, it is clear, if the connecting stalk arises in the way
outlined, that it is more ceenogenetically modified than is that of
Tarsius, the latter providing the link betwee: the Lemuroids and
the Anthropoids.

In both Varsiuvs and the Anthropoids, the connecting stalk
later on becomes enclosed with the yolk-stalk in a tubular
prolongation of the margin of the umbilical opening, which
earries the amnion with it. The cord-like structure so formed is
the umbilical cord, which connects the embryo with the placenta.
In examining the feetal Varstws which Dr. de Lange so kindly
sent me, I noticed that the cord is related to the placenta some-
what differently to that of the Anthropoids. Whereas in the
latter, the cord takes the form of an elongated, more or less
twisted, rope-like structure which runs as a free cord to become
connected with the chorion covering the free surface of the
placenta, usually near its centre, In Tarsiws the cord is quite
short, and instead of passing directiy to the placenta, it joins the
chorion soon after it becomes clear of the body of the foetus and
runs down in that to join the distal margin of the placenta
(text-fig. 1). This difference confirms the conclusion we reached
above that the development of the cord does not follow identical
lines in the two groups.

Lastly, let us see what conclusions as to the affinities of Tarsius
may be drawn from the study of the placenta itself. The
development and structure of this remarkable organ were de-
scribed by Hubrecht in a lengthy paper published in 1899, but in
that paper Hubrecht was more concerned with demonstrating the
occurrence of a supposed blood-forming or hemopoietic process
in the placenta than with the description of the structure of the
ripe organ itself, and his figures fail to convey an adequate idea
of its characteristic structural features. That defect I have
attempted to remedy by the provision of the microphotographs
of sections of the nearly full-term placenta shown in fig. 1, and
herewith reproduced as figs. 2, 3, and 4, PI. I.

These figures supplement Hubrecht’s account of the placenta
very materially, and illustrate quite adequately, I think, the more
important features in its structure. rom these figures, anyone
who is familiar with the sectional appearance of the human
placenta, will recognise at once that he is dealing with a placenta
of the Anthropoid type. Hubrecht, with reference to his own
fig. 66, remarks ‘‘eine entfernte Aehnlichkeit mit der mensch-
jichen Placenta, wie sie Sedgwick Minot abbildet, ist nicht
zu verkennen,” and that is certainly the conclusion that any
competent embryologist would come to from an inspection of
my fig. 4. The general resemblance is indeed extraordinarily
striking, but, whilst that is so, it must be emphasized that in its
detailed structure, the Zarsius placenta shows peculiarities of its

AND AFFINITIES OF 'TARSIUS. 487

own which clearly mark it off from the Anthropoid placenta.
Although it foreshadows the latter in the most unmistakeable
fashion, it has failed to attain the same level of structural and
functional differentiation.

The mature placenta of Varsius, viewed as a whole (cf. fig. 1,
Pl. I., where it is seen in section) appears as a massive knob-
shaped or rather cone-shaped structure, measuring, in my specimen,
10x11 mm. in diameter, which projects freely into the uterine
lumen and is attached to the very thin uterine wall in the apical
region of the uterine horn by a very short staik through which
the maternal vessels pass. Its distal, slightly concave surface is
clothed by the chorion which marginally is, on the one hand,
reflected down to invest the remainder of the free surface, right
down to the stalk, and on the other, is continued on as non-
placental chorion, a very thin non-vascular membrane, with whose
inner surface the amnion lies in close apposition. In its cone-
shaped form and freely projecting character, it contrasts with the
sessile, cake-like, discoidal placenta characteristic of the Anthro-
peids. ‘These features are dependent on the fact that the placenta
develops in relation to a localised knob-like thickening of the
subepithelial tissue of the uterus, the decidual swelling or tropho-
spongia. ‘To this, attachment is effected in the first instance, and
it later on projects and serves as an axis round which the cone-
shaped placenta develops. In the process it undergoes progressive
degeneration, and only a remnant of it is preserved in the stalk-
region of the completed placenta. Its presence in the developing
placenta no doubt conditions the appearance of the extensive
blood-extravasation which is such a marked feature in the ripe
organ (fig. 2). No such conspicuous, localised, decidual swelling
has so far been described in the placental development of any
Anthropoid.

The Tarsius placenta agrees with that of the Anthropoids im
that it is deciduate and of the hemochorial type, 7.e., the fune-
tional placenta consists, except for the maternal blood present in
it, exclusively of foetal tissue and the maternal blood circulates
through lacunar spaces bounded solely by the foetal trophoblast.
This same type of placenta, however, occurs also in such diverse
orders as the Rodentia, Cheiroptera, Insectivora, and Xenarthra,
so that this similarity does not carry us very far. A more impor-
tant agreement, from our present point of view, lies in the fact
that the functional placenta comes to be established as the result
of the outgrowth from the mesoderm of the chorion of more or
less massive sprouts which grow into the syncytial layer formed
by the trophoblast and which branch abundantly to form charac-
teristic dendritic villi, in which the fcetal vessels are situated.
At the same time, the trophoblast provides, apparently in a some-
what different fashion in the two groups, an enclosing layer
vound each villus (including all its branches), whilst its blood-
filled lacunze extend so as to form a system of intervillous blood-
spaces. But there is an important difference in the villi in the

488 ON THE ZOOLOGICAL POSITION

two groups, for, whereas in the Anthropoid placenta the villi
are strongly marked and individualised structures which project
freely into what is practically a continuous blood-sinus formed by
the confluence of the intervillous spaces, in Zarsius they are less
prominent structures and except at their origin from the chorion
cannot be said to be individualised at all, since thei enclosing
layers of syncytial trophoblast are not individually distinct but
are connected with each other by anastomoses, the trophoblast
persisting in the form of a syncytial network, in the walls of
which the mesodermal villi are enclosed (Pl. I. figs. 3 & 4).

Furthermore, the intervillous spaces, except round the peri-
phery of the placenta, immediately below the chorion, where they
have coalesced to form definite blood-sinuses of some size (Pl. I.
figs. 2 & 3), take the form elsewhere of narrow and tortuous
channels, which, deed, in the deeper parts of the organ are
frequently incompletely hollowed out, many of them being more
or less blocked by a light-staining reticular material (PI. J. fig. 4),
no doubt derived from the breaking down of the trophoblast
during the formation of its lacune, but which has not been com-
pletely removed, owing perhaps to the slowness of the circulation
in the central region of the organ. Compared with the Anthro-
poid placenta, that of Zarsius strikes one as being on a much
lower plane of functional efficiency.

Hubrecht in one of his more recent papers (1908) has himself
emphasised the above-described difference in the relations of the
villi. He writes :—‘‘ The freedom with which they float about
in the maternal blood is another characteristic of Man and
the Monkeys. In Yarsius and in the Hedgehog their arrange-
ment is more that of a suspension in a very delicate and at the
same time most intricate trellis-work formed by the trophoblast
cells that have become spun out into this. When the connecting
trabecule of this trellis-work are suppressed, as we see it in the
higher Primates, the surface available for osmotic interchange
_is naturally increased and the free movements of the villi may
also be considered as an advantageous circumstance.” Leaving
aside consideration of certain other differences in detail in the
placentation of the two groups, e.g. in the early development, m
the constitution of the investing trophoblastic layer of the villi,
and in the occurrence of placental blood-extravasations, we may
deduce from the above quotation that Hubrecht regarded the
difference between the Varsvws and Anthropoid placenta as one
of degree only and that he looked upon the former as a much
less perfect organ functionally than the latter.

With these deductions, | am in agreement with the reservation
that I hold the degree of difference to be such as to justify us in
definitely excluding Tarsius from the Anthropoid group. In its
placentation, Tarsius is clearly on the line which leads to the
Anthropoids, but it has failed to attain their status, and im this
respect, as in so many others, is a true ‘ Halbaffe,” intermediate

AND AFFINITIES OF TARSIUS. 489

between the Lemuroids and the Monkeys, but approaching much
more closely to the latter than to the former.

The conclusions we have arrived at from the consideration
of the data of development may be summarised as follows:
(1) The Lemuroids represent the basal stock from which the
higher Primates evolved. They have retained in their develop-
ment many primitive Mammalian features, including a primitive
form of diffuse non-deciduate placenta, Developmentally they are
free from marked specialisation, and they present us with a
developmental ground-plan of such a generalised type as to be
easily susceptible of such adaptive modifications as have occurred
in the higher types in the course of evolution. (2) The Tarsioids,
early separating from the primitive Lemuroid stock, were more
progressive. They show in their development the beginnings of
those adaptive changes which reach their culmination in the
Anthropoids, and by acquiring an early attachment to the uterine .
wall they developed a localised deciduate placenta of the heemo-
chorial type; but for some reason, perhaps owing to a too active
participation on the part of the maternal decidua, they failed to
exhaust its possibilities and to evolve an organ of the highest
possible efficiency. In many features of their development, they
are transitional between the Lemuroids and the Anthropoids, but
they are plainly on the Anthropoid line, and from them the
Anthropoids undoubtedly took their origin. (3) Starting from
the Tarsioid stock, already provided with the beginnings at least
of a hemochorial placenta, the Anthropoids went on to make the
most of their inheritance, and evolved a highly efficient type of
nutritive organ in which the individualised villi are directly
bathed by the maternal blood—an efficiency which is reflected in
the advanced grade of organisation exhibited by the new-born
young. In them and more particularly in the highest forms, the
Anthropoid Apes and Man, developmental adaption has reached
its acme, as witness the complete implantation of the early
blastocyst and the correlated development of a complete decidual
capsule ; and here I may be permitted to add, of the close genetic
affinity of Man and the higher Apes there can be no question on
embryological grounds.

- Finally, as regards the systematic position of Tarsiws, Hu-
brecht’s contention that it must be removed from the Lemurs
I fully accept, but I am unable to agree that its true position is
with the Anthropoids. The remarkable annectant characters
which Yarsius exhibits justify us in placing it, along with its
extinct allies, in a subdivision of its own, and I am, therefore, in
agreement with those who, like Gadow and Elliot Smith, have
advocated, on quite other grounds, the division of the Primates
into three great radiations—call them what you may,—viz. a basal
or Lemuroid group, a Tarsioid group, and an Anthropoid group.
This tripartite arrangement seems to me most in accord with the
embryological data and best expresses the phylogenetic importance

490

ON THE ZOOLOGICAL POSITION

of this extraordinary creature, outside Man—-perhaps the most
interesting of all the Monodelphian Mammals.

I desire to express a thanks to my Laboratory Assistant,

My. F. J. Pittoek, for the skill and care he
preparation of the Plate.

has expended on the

Comparative Table, Development of Primates.

(2)

Tarsius.

(1)

LEMUROIDS.

(3)

ANTHROPOIDS.

as 1 (1).

1. pWescloument central.

as In (1) or sraceuctiels,

2. No attachment to ie Kemene very guilty ef.
rine wall. ‘Trophoblast tected by localised thick-|
relatively inactive. ening of trophoblast.

Trophoblastic activity
more marked than in (2).’
A single or double at-|
tachment or interstitial
inbedding very early ef. ;
fected.

|
:

3. onan fonued by fold- as in (1), but without pro-|

(loved aimnion- Funnton
the primitive amniotic
cavity persisting. !

formation with proam-| amnion.
nion. |

4. Allantois free and vesi- Allantois never free and.
cular; unites fairly early, vesicular, transformed)
yah chorion. / into ‘connecting stalk.” |

as in (2), but connecting,

stalk semivesicular in|
Hapale.

5. Yolk-sac Foenede in nor-| Yolk-sac precociously
mal fashion, but is early) formed and reduced from)
separated and soon be-| the first, never completely
comes reduced. filling space enclosed by
the tr rophoblast.

as in (2), but even more|
precocious and more re-
duced.

; :
6. Formation of extra-em-) Extra-embryonal mesoderm

as in

(2) but distinetly

throughout by allantois., and non-vascular, except
over placental area, which.
is vascularised through)
connecting stalk.

bryonal mesoderm and} and ccelom precociously, more specialised.
celom normal, but they) tormed. |
extend relatively early |
through entire omphalo-
pleure. |
7. Giecoa deculamenal Chorion extremely thin Chorion vascularised

throughout by way of,
connecting stalk.
|

18. Yolk-sac placenta, 1f|
DEESe BRAUN

No yolk-sac placenta.

|
| 1!

as in (2).

=| at |
non-) Placenta conical, deciduate,
hamochorial in type,!
but villi not individu-
| alised and clothed by syn-
cytio-trophoblast only.

ae
9. arena Ganeedl
| deciduate, epithelio-cho-
rial in type.

Placenta Sines or double!
discoidal and hamocho-|
rial, villi distinct and
projecting into maternal
bleod-sinus. Tropho-
blast of villi distinguish-
able into syneytio- and
cyto-trophoblast.

AND AFFINITIES OF TARSIUS. 49]

EXPLANATION OF PLATE TI.

Fig. 1. Photograph (X 1°5) of foetus of Tarsius spectrum (measuring 2°9 cm. in
G. L.), atter removal from the uterine hom and with the cone-shaped
placeuta attached. The short umbilical cord is seen to join the chorion
shortly after becoming clear of the body of the foetus and to run down in
that membrane to join the placenta marginally. ‘The placenta has been
divided by a vertical cut, the cut-surface showing (cf. fig. 2 and explanation
thereof).

[Hubrecht Coll., Tarsinus 76. Coll. Dr. Fock, Muntok Bankaleg, 17th
March, 1893. |

Fig. 2. Photomicrograph (xX 6) of vertical section of the entire placenta, well to one
side of the area of attachment to the uterine wall. Note the investing
chorion with the main stems of the villi arising from it (best seen on upper
side of the section), the peripheral blood-sinuses, and the large blood-
extravasation centrally.

Fig. 3. Photomicrograph (X 34) of a small portion of the superficial region of the
placenta (including the large villus visible in the upper left-hand sector of
fiz. 2). Note the chorion forming the boundary of the section on the
upper side, the peripiieral blood-sinus, and a large and a small villus-stem
arising from the chorion, the former showing well the characteristic
method of branching.

Fig. 4. Photomicrograph (x 80) of a portion of the placenta, situated a little deeper
than the base of fig. 3, to show the vil under higher magnification. The
villi are mainly cut transversely, and in the region photographed are, on the
whole, of greater tnan average diameter. Kach villus cousists of a core
of choriomie mesenchyme carrying the wmbilical vessels and invested by a
more deeply stained sheath composed of a thin nucleated layer of syncytial
trophoblast. An underlying layer of cyto-trophoblast such as is found in
the villi of Anthropoids is at vo time present in Tarsius. Adjoining villi
are connected by bridges of syncytio-trophoblast, the latter thus forming
an irregular network. Between the villi ave the intervillous spaces, many
of them being more or less completely occupied by the reticular light-
staining material referred to im the text (p. 488).

Prof. F. Woov-Jones, D.Sc., M.B., F.Z.S.:—Although the
general anatomy o7 reine may be said to be fairly well known,
there is still great need for complete accounts of dissections of
special systems and for the general examination of a larger
number of specimens. Apart fronr the gaps in our knowledge of
several important details, there is as yet no basis for forming an
estimate of the range of individual variation.

Asa result of comparing published accounts of the anatomy of
Tarsius and from dissecting an adult female specimen, one can
only conelude that either the range of individual variation is
considerable or the interpretations of different investigators
show a rather unusual Jack of agreement. Hxternal characters
have been studied in two specimens, both adult females, and the
details of bodily structure in one of them; in addition, I have
had X-ray photographs of various portions of both specimens.
[For my material I am indebted to Profs. G. Elliot Smith and
J. P. Hill, and for the X ray piates to Dr. Stanley Meiv ille. ]
For the purpose of this discussion it is impossible to furnish more
than a sammary of the anatomical details, and I have thought it
best to mention only the outstanding features which are likely to
throw light on the affinities of Zarsiws, dealing especially with
those points which link it to, or separate it from, the typical
members of the Lemuroidea and Anthropoidea respectively.

492, ON THE ZOOLOGICAL POSITION

It cannot be doubted, from an examination of its entire anatomy,
that Tarsius is a remarkably primitive mammal which has early
acquired an over-specialisation in vision and in powers of frog-
like arboreal leaping. In these two features it roughly resembles
Galago, and it may be doubted if, had this likeness not been so
apparently exact, zoologists would have classed Zarsius in the
Lemuroidea with so much confidence. One primitive feature
retained by the specimen dissected is the presence in the carpus
of an element which I can only homologise as a second os centrale.
This small bone has not previously been described, and its de-
tection without the use of X-rays would be rather a matter
of chance. The true os centrale (os accessorium, Lurmeister)
articulates with carpale I., II., and 111., as well as with the
scaphoid and semilunar (thus differing from the condition as
described and figured by Burmeister).

The second os centrale exists on the radial side of the carpus
in the gap between carpale I. and the scaphoid ; it does not
actually articulate with any bone, being held in place by fibrous
tissue. It is not situated in tendon, “but is enveloped in the
capsule of the carpal articulation. Should this small bone prove
to bea constant element in the carpus of Tarsius, it would furnish
very interesting evidence to which it is hoped attention will be
given by other “observers having command of material. Tarsius
differs from all members of the Lemuroidea in certain of its
external characters:—(1.) In having the nostril completely
ringed by the meeting of the lateral and medial nasal processes.
In “uhis encircling of the nares it resembles all members, and
surpasses some, of the Anthropoidea. (11.) In having the upper
lip simple in the mid-line. There is no lemmrine incisura
between maxillary processes, but a truly Simian philtrum com-
posed of medial nasal processes. (r1r.) The hair-tracts resemble
those of the highest of the Anthropoidea far more nearly than
they do those of the Lemuroidea, or even the lower Anthro-
poidea. (1v.) The digital formula differs from that typical of the
Lemurs, the middle line digit being the longest. (v.) The meta-
carpal formula shows the same primitive and non-lemurine
features. (vr.) The external genitalia, especially those of the
female, are formed upon a plan altogether unlike that seen in
the Lemurs and resembling that typieal of the highest
Anthropoidea.

Tn its internal anatomy Varsius differs from all members of the
Lemuroidea in (vit.) its dental characters ; (vuir.) in the forma-
tion of the auditory bulla; (1x.) in the form and fate of the
tympanic bone, and in the inception of an external auditory
meatus. Combined with these last two characters is (x.) the
non-lemurine cranial course of the internal carotid artery. In
the formation of the orbit two characters, typical of the Anthro-
poidea and differing widely from anything seen in the Lemurs,
are conspicuous : (xt.) with the inner wall of the orbit the
os planum of the ethmoid enters, and (x1.) the orbit is furnished

AND AFFINITIES OF TARSIUS. 493

with a posterior wall in the formation of which the alisphenoid
takes part. (xit.) The hyoid, with its relatively short ‘ lesser”
cornua, differs from the typical hyoid as seen in the Lemurs, and
resembles that of the monkeys and apes (Burmeister’s figure
and account of the hyoid does not accord with the condition
present in my specimen). (xiv.) The trachea is not formed upon
the peculiar lemurine fashion, for the rings are incomplete behind,
as they are in the Anthropoidea. (xv.) The two halves of the
mandible are synostosed in the middle line. (xv1.) The ventral
pelvic symphysis is shallow, and is limited to the pubis. In the
gastro-intestinal tract the very widest differences exist between
Tarsius and any members of the Lemuroidea, and for the
purpose of summary we will only note (xvit.) the simplicity of
the gut pattern of Zarsius, and the absence of any coiling which
is so characteristic of the Lemurs. In many features the
myology of Zarsius differs from that of the Lemurs and resembles
that of the Monkeys and Apes; as an example, (xvu1.) the very
human disposition of the digastric muscle may be instanced. In
the vascular system (xIx.) the arrangement of the vessels on the
arch of the aorta provides a striking contrast to the condition
seen in the Lemurs.

In summing up the entire anatomy of the Lemurs and of the
Monkeys and Apes it is impossible to avoid appreciating “the
differences in structure that indicate the wide separation of
the Lemuroidea and the Anthropoidea” (Elliot Smith, ‘ Nature,’
May 2, 1907, p.7). Judged by such standards as are commonly
employed in mammalian classification, such basal features as
those comprised in the structure of the nose and lips, in the
tympanic and orbital regions of the skull, and in the genitalia
appear to justify the separation of Zarsiws from the Lemuroidea,
and warrant its inclusion in the Anthropoidea. But in several
very striking details Z’arsius differs from all members of the
Anthropoidea except the very highest. Im many characters
Tarsius resembles Homo and differs from the Monkeys, in some
it resembles Homo and certsin of the Aunthropoid Apes. These
features are only to be termed primitive mammalian characters,
and as a result of summarising the anatomy of the Anthropoidea
it appears to me to be legitimate to conclude that Zarsiws and
Homo retain a remarkable series of primitive mammalian
characters, some of which are retained in part in the Anthropoid
Apes, but which are departed from increasingly widely as the
zoological seale of the Anthropoidea is descended. (1.) Zarsius
differs from all Anthropoidea except Man and the Orang in the
arrangement of the elements in the basis erani; here it re-
sembles all primitive mammals. (11.) It differs from all Anthro-
poidea, with the same two exceptions, in the normal retention of
the alisphenoid-parietal pterion. (1.) In the digital and meta-
carpal formulas it shows a primitive (and human) character in
avoiding the relative reduction of the 2nd digit of the manus
typical of the remaining members of the Anthropoidea. (1y.) In

494 ON THE ZOOLOGICAL POSITION

the external genitalia the affinities of Tarsius are far nearer
to the highest members of the series than they are to the lowest.
(v.) In the absence of a penile ossicle or penile cartilage, the
likeness is with Homo alone. (vy1.) In the arrangement of the
aortic trunks Varsiuws differs from all monkeys, and finds an
absolute parallel only in Man normally and certain Anthropoids
occasionally. Various points in myology are of interest as
showing retention of muscles in Zarsiws and Homo in a primitive
condition not preserved in other members of the Anthropoidea.
(vit.) Zarsius retains, like Homo, a well-developed pulmaris
brevis. (vitt.) The flexor accessorius is confined to Zarsius and
Man alone among the Primates. (1x.) The supinator brevis 1s
pierced by the posterior interosseus nerve exactly as it is in Man,
and the condition differs widely from that which is occasionally
_ present in the Chimpanzee, and which is otherwise the most
human manifestation of this muscle within the limits of the
Anthropoidea. (x.) The palmaris longus and plantaris are both
well developed. (x1.) The levator ang tli scapulee is in its human
condition. (xir.) The flexor pollicis longus, though judging from
published accounts displaying a vari iability i in difterent specimens,
approaches the human form and differs from the Simian in a very
remarkable manner.

In conclusion, Varsius appears from a summation of its
anatomical characters to belong to the Anthropoidea, of which
group it constitutes one of the most primitive members. In the
retention of primitive features of bodily architecture it finds its
paraliel in omo; and it differs from the rest of the Anthro-
poidea in which Simian specialisations have effected definite
alterations.

Tarsius appears to be a very primitive member of the Anthro-
poidea in which early specialisation of vision and arboreal leaping
activities absorbed the phylogenetic development of the species.
HIomo appears also as an extremely primitive form in which
cerebral advances, and lack of unequal physical specialisation,
save that of bipedal progression, is the phylogenetic keynote.
The Anthropoid Apes have departed more from the primitive
mammalian type in definite “Simian” specialisations—which
specialisations become increasingly conspicuous in the ‘lower ”
members of the Order.

R. 1. Pococx, Ksq., F.R.S., F.Z.8.:—In the ‘ Proceedings’ of this
Society, Aug. 1918, I pointed out that Varsius differs from all
the Lemurs and resembles the higher Primates in the structure
of the nose and muzzle and in the mobility of the lips; and that
the external genitalia of the female in the concealment of the
small clitoris and the orifices of the urethra and vagina by a
pair of labia are unlike those of the Lemurs, especially of the
Oviental and African forms (Lorisiformes), and recall rather those
of the Old-World Pithecoid Primates.

These differences between Z'arsius and the Lemurs, added to

AND AFFINITIES OF TARSIUS. 495

those previously established in connection with the skull and
teeth, the placenta and the digital formula of the hand, out-
weighed, in my opinion, the likeness between Varsius and the
Lemurs, and enforced the removal of the former from the latter
and its segocietien with the Monkeys, Apes, and Man in a group
of the Primates for which the term Haplorhini was proposed, the
true Lemurs, the Lorises, and Chiromys being graded in contra-
distinction as Strepsirhini. The Haplorhini were divided into
two suborders, the Tarsioidea for Varsiws and the Pithecoidea
for the Platyrhini (American Monkeys) and the Catarhini (Old-
World Monkeys, Apes, and Man).

This classification appeared, and still appears, to me to express
the known facts more accurately than its predecessors, the
nearest to it being that of Gadow who in 1898 definitely dis-
sociated Zarsius from the Lemurs, dividing the Primates into
tne three suborders Lemures, Tarsii, and Simiz. Possibly he
would have anticipated my systematic arrangement had he been
acquainted with the structures connected with the muzzle and
vulva in Zersius and other Primates and attached to them
the importance that Ido.

J. T. Cunnincuam, M.A., F.Z.S.:—In the development of
Mammals generally segmentation of the ovum produces a small
internal mass of cells covered by a single layer of external cells.
Accumulation of watery liquid between these two parts produces
the blastocyst, a vesicle of epiblast cells with the inner cell-mass
adhering to the inner surface of the vesicle at one small area. The
wall of the vesicle is the trophoblast, which corresponds to the
epiblast layer of the serous membrane or false amnion of birds
and reptiles. The next step in the mammal is the differentiation
of the hypoblast from the lower surface of the inner cell-mass :
this hypoblast grows round the inner surface of the blastodermic
vesicle, and so forms the inner lining of the yolk-sac. The
amnion is formed either by coalescence of external folds above
the inner cell-mass or as a closed cavity within the latter. The
mesoderm is formed next by differentiation between the epiblast
and hypoblast within the inner cell-mass, which may be now
called the embryo. The mesoblast extends between trophoblast
and hypoblast, and between trophoblast and amnion, splitting as
it goes to form a cavity called the extra-embryonic celom. By
folding in of the sides of the embryo the connection of yolk-
sac and embryonic gut is narrowed, and from the hinder end
of the gui grows out the allantois as a hollow sac into the extra-
embryonic ccelom.

We have thus a membrane called the chorion, consisting of an
external layer of epibiast and an internal layer of mesoblast,
entirely enclosing the embryo with its three membranes, the
amnion surrounding it dorsally, and the yolk-sac and allantois
extending from the gut ventrally. These two sacs oceupy varying
proportions of the inner surface of the chorion in different

496 ON THE ZOOLOGICAL POSITION

mammals. In Ungulates the cavity of the allantois is very large
and its outer wall is in contact with the inner surface of the
chorion over very nearly its whole extent, the yolk-sac becoming
very small. The chorion grows out into villi over its whole
surface, and these villi penetrate into pits or erypts of the inner
surface of the uterus, without becoming united with uterine
tissue. The mesoblast of the allantois with its blood-vessels
grows into these villi and thus is formed the diffused placenta.

In Man the amnion appears to be formed as a closed cavity,
and the hypoblast develops as described above, but instead of
extending round the inner surface of the chorionic vesicle it
forms a small closed sac, and mesoblast develops on the inner
surface of the chorion and the outer surface of the yolk-sae with
a large cavity between, which is the extra-embryonic ccelom.
There is at first no mesoblast within the embryo, but it extends
outside the amnion and attaches the embryo te the mesoblastic
lining of the chorion. As the embryo develops, this connection
lengthens somewhat and becomes first posterior then ventral.
This connecting mesoblast is the umbilical cord, and represents
a solid alllamioie. into which a very rudimentary cavity extends
from the hinder end of the gut. The mesoblast at the outer
end of the stalk forms villi over a dise-shaped area of the
chorion and so forms a discoid placenta.

With regard to these peculiarities of development, the ordinary
Lemurs agree with the Ungulata and Zarsiws agrees with Man.
It may be said, therefore, that there is as much difference between
the development of Varsius and that of a Lemur as there is
between that of Man and that of an Unguiate. Moreover, in
Man the trophoblast destroys and absorbs the uterme mucous
membrane and the allantoic villi extend into embryonic tro-
phoblast-tissue. In this respect also Tarsius agrees with Man
and differs from Lemurs. ‘To suppose that the mode of dev elop-
ment of Tuarsius and Man has been independently evolved
without any close relationship between the two seems to me
unreasonable.

The development of 7'arsiws is more similar to that of Man in
some respects than that of Monkeys and Apes. In many
Monkeys there are two placentas, a dorsal and a ventral; in Man
only one, which is dorsal. In Hylobates and Sina there is only
one placenta, but it is ventral. In Zarsius the placenta is
single and dorsal. The Apes, however, are nearer to Man in
having the chorion entirely imbedded in uterine tissue, which
is not the case in Monkeys or Varsius.

Dr. P. Caaumers MircHenpt, F.R.S.:—Characters have to be
judged as well as counted, if it be intended to use them for
estimating the relative degree of affinity between animal types.
No anatomist doubts but that Man retains many primitive
characters; Anthropoid Apes, Old-World Monkeys, American
Monkeys, ZYarsius, and Lemurs also retain many primitive

AND AFFINITIES OF TARSIUS. 497

characters. It is reasonable to assume that the common ances-
tors of all these animals possessed all the primitive characters
retained by any of them. And so it is not surprising to find
any primitive character in any descendant of a common stock,
but there is no reason to suppose that, because any two have
retained the same primitive character, they should for that reason
be judged more nearly related than either may be with some other
descendant of the common stock. Primitive characters may be
useful for the description or definition of a group—they have
no value for assigning degrees of affinity. These considerations
ought to be commonplaces in zoological argument, but they are
often forgotten, and I think they have been entirely forgotten by
Professor Wood-Jones in the imposing list of common characters
that he has drawn up for Man aind Lursius. Fortunately
they have been remembered by Mr. Pocock, and Professors Hill
and Elhot Smith, and the considerations they adduce have dis-
posed of Professor Wood-Jones’s argument that Varsius has a
special relation to the ancestry of Man. It may not be a Lemur,
but it is no nearer to Man than to other Primates.

Professor Wood-Jones, to whom we are indebted for originating
the interesting discussion, was more reticent to-night than in his
addresses to more popular audiences. He attacked Darwinian
evolutionists, and Huxley in particular, on the supposition that
they believed genealogical trees to be linear, that a higher group
took origin from the highest members of a lowe group. It was
a strange misreading of familiar evidence. Were he to consult
Huxley’ S Hssay on “Man's Place in Nature,” or any general state-
ment of the case for Evolution, as, for instance, the Article under
that heading in the ‘ Encyclopedia Britannica,’ he would see that
he was attacking a bogey that does not exist. Writer after
writer, with increasing insistence in recent ye ars, has dwelt on
the obvious fact that existing groups are at most in the relation
of collateral descendants of a common ancestor, and the tendency
has been to place the common ancestor ever lower and lower on
the tree of life.

Prof. MacBripg, F.R.8.:—In summing up the discussion the
Chairman said that there was one point on which all the speakers
were agreed, viz., that Zarsiws was much more nearly allied to
the higher Primates than it was to the Lemurs. But that was a
point which under any circumstances few would have disputed,
especially since Prof. Hubrecht’s researches on the placentation
of this form which had been so ably summarised for us by Prof.
Hill. The whole interest of the qvestion lay in Prof. Wood-Jones’s
attempt to prove that Zarsius and Man agreed in retaining
important primitive characters which the other Monkeys had
lost—and in the obvious inference from this position, from which
Prof. Wood-Jones had rather shrunk during the discussion, but
to which he had liberally committed himself in recent books
published by him. This inference was briefly that the human

A98 POSITION AND AFFINITIES OF TARSIUS.

stock had separated from the stock of Primates at the J'arsius
level and that all resemblances between the two stocks were due
to convergent evolution.

In Prof. MacBride’s opinion the points of similarity between
Tarsius and Man adduced by Prof. Wood-Jones were superficial
peculiarities, and the assertion that these structural charac-
teristics were absent from Monkeys seemed to rest largely on our
imperfect knowledge of the anatomy of Primates, as the ad-
mission of Prof. Wood-Jones that some of them appeared as
“exceptions” in the Chimpanzee and the Gorilla amply proved.
Prof. Elliot Smith had rightly emphasised the deep and funda-
mental resemblances between the Higher Apes and Man, and his
exposition would leave no doubt in the minds of most of us that
the older view that Man was sprung from a Simian stock was
true.

The great interest in the question of Man’s origin was to discover
the cause of his evolution, for we might rest assured that Man did
not evolve in response to an innate tendency lodged in a Monkey’s
constitution, but in response to needs created by a change in the
environment. The Apes were arboreal animals; and Man was,
anatomically, a Ground-Ape. When towards the end of the warm
Tertiary period, the forests began to shrink and to be replaced by
steppes swarming with swift-footed grazing animals, the restriction
of food on the trees and its obvious abundance in the plains led
to the most enterprising Apes venturing on to the ground and
assembling in troops to run down their prey, and thus a begining
in the evelution of Paleolithic Man the primitive hunter was
made. .

The race of squirrels, essentially tree-loving animals, had, in like
manner, spread into steppes and prairies, and given rise to short-
tailed short-legged forms like the Prairie-Dog and the Marmot,
which were essentially Ground-Squirrels.

If these considerations were justified, it followed that the
phrase ‘* Arboreal Man” was a contradiction in terms.

November 4th, 1919.

A. Smirn Woopwarb, Hsq., LL.D., F.R.S., Vice-President,
in the Chair.

Mr. F. Marvin Duncan, F.R.M.S., F.Z.8., exhibited a series
of photographs showing the actinic quality of the light from a
living Pyrophorus Beetle, and described the method he had
employed to obtain his records. He stated that the results
obtained appeared to show that photo-spectroscopically the
ereatest intensity of light action was in the yellow-green region.
Unfortunately the single specimen at his disposal was already in
a somewhat exhausted condition on coming into his hands, so

THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 499)

that he was unable to investigate the matter as thoroughly as he
would have desired.

Mr. EK. Heron-Auten, F.R.S., F.Z.S., exhibited a series of
Skiagraphs of the Foraminiferan genus Vernewilina from examples
grown in a hypertonic tank, and described some of the further
results that he and Mr. A. Earland, F.R.M.S., had obtained in
the course of their investigations, drawing particular attention
to the remarkable modifications produced in the morphology of
the “‘ test” of the specimens exhibited.

November 18th, 1919.

Prof. E. W. MacBripsz, F.R.S., F.Z.S., Vice-President,
in the Chair.

The Secretary read the following Report on the Additions
made to the Society’s Menagerie during the month of October
L919; —

The registered additions to the Society’s Menagerie during the
month of October were 138innumber. Of these 64 were acquired
by presentation, 55 were deposited, 15 were purchased, and 4
were born in the Menagerie.

The following may be specially mentioned :—

2 Musk-Oxen (Ovibos moschatus), new to the Collection, from
Greenland, purchased on October 9th.

2 Lions (Felis leo), $ 9, from Africa; 3 Leopards (felis
pardus), 3,1 Nylghaie (Boselaphus tragocamelus), 2 , from India :
presented by The Rajah of Payagpur on October 4th.

1 Leopard (Felis pardus), 2 Striped Hyenas (Hyena hyena),
from India, presented by The Rani of Bansi Basti on October 4th.

1 Blackbuck (Antilope cervicapra), 2 , 1 Sambur (Rusa unicolor),
2, from India, presented by W. B. Cotton, Esq., I.C.8., on
October 4th.

1 Blackbuck (Antilope cervicapra), S$, from India, presented by
Dr. Gulzeri Lal on October 4th.

A collection of Birds from New Guinea and the Malay Archi-
pelago, including four Lesser Birds of Paradise (Paradisea minor),
and containing several species new to the Collection, deposited on
October 27th.

Sir Epmunp Gites Loprr, Bt., F.Z.8., exhibited and made
remarks on a series of skulls of the Beaver, showing a separate
ossicle between the parietals.

Proc. Zoou. Soc.—1919, No. XXXIV. 34

Sg ai
Hig eae

No. 196.

ABSTRAOT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

October 21st, 1919.

Prof. H. W. MacBrins, F.R.S., F.Z.S., Vice-President,
in the Chair.

The Secretary read a Report on the Additions made to the
Society’s Menagerie during the months of June, July, August,
and September.

Mr. OnprietpD Tuomas, F.R.S., exhibited three interesting
Mammals obtained by Dr. Aders, F.Z.S., in Zanzibar: namely,
an example of Cephalophus adersi, a recently described new
species; an example of Colobus kirki, which until lately was
supposed to be almost extinct, and a specimen of a rare Insectivore
belonging to the genus Petrodromus.

Mr. E.G. Bounenesr, F.Z.8., read a “ Report on the Research
Experiments on Methods of Rat Destruction carried out at the
Society’s Gardens,” and exhibited some of the traps that had
proved most successful.

* 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 Stapence, or, if desired, sent post-free for
the sum of Six Shillings per annum, payable in advance,

Dr. A. Surra Woopwarp opened a “ Discussion on the Zoological
Position and Affinities of Tarsius,” and the discussion was con-
tinued by Prof. F. Woop Jonss, D.Sc., F.Z.8., Prof. J. P. Hint,
E.R.S., F.Z.S., and Dr. G. Extiorr Surrs, M.A., F.R.S., F.Z.S.
Contributions by Mr. R. I. Pocock, the Secretary, and the Chair-
man had to be taken as read.

The next Meeting of the Society for Scientific Business will
be held on Tuesday) November 4th, 1919, at 5.30 p.m., when the
following communications will be made :—

Exhibition of Skiagraphs of Vermiculina from examples
grown in a hypertonic tank.

Guy MarsHALL, D. Se., i Z. 8.

on the Species of Bulaninas occurring in Borneo (Coleoptera,
Curculionide ).

On the Variation in the Number of Dorsal Scale-rows in
our British Snakes.
G. A. Boutencsr, F.R.S., F.Z.8.

On some new Fishes from near the West Coast of Lake
Tanganyika.

The Hon. _PauL METHUEN, ) Le A. 5.

Description of a new Snake from the Transvaal, together
with a new Diagnosis and Key of the Genus Xenocalmus, and
of some Batrachia from Madagascar.

The following Papers have been received :—
M. Turner, B.Sc.

On the Nematode Parasites of a Chapman’s Zebra.

Rev. A. H. Cooker, E.Z.S.

The Radula of the Mitride.
eo SuenoN Pamiucon C28:

Field-notes on some Mammals in the Bahr El Gebel,
Southern Sudan.

Lt.-Col. 8S. Monckton Copeman, F.R.S8., F.Z.8.

Experiments on Sex Determination.

The Publication Committee desire to call the attention of
those who propose to offer Papers to the Society, to the great
increase in the cost of paper and printing. This will render it
necessary for the present that papers should be condensed and
be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should
be addressed to

P. CHALMERS MITCHELL,
Secretary.

ZOOLOGICAL Socrery or Lonpon,
ReGeENt’s Park, Lonpon, N.W. 8.
October 27th, 1919.

ve Yi a é
a ee
Pa dd

No. 197.

ABSTRACT OF THE PROCEEDINGS

ZOOLOGICAL SOCIETY OF LONDON.*
November 4th, 1919.

A. SuirH Woopwarb, Esq., LL.D., F.R.S., Vice-President,
in the Chair.

Mr. F. Martin Duncan, F.R.M.S., F.Z.S., exhibited a series
of photographs showing the actinic quality of the light from a
living Pyrophorus Beetle, and, in describing the method employed
to obtain the records, stated that photo-spectroscopically the
greatest intensity of light action appeared to be in the yellow-
green region.

Mr. EK. Heron-Apuen, F.R.S., F.Z.8S., exhibited a series of
Skiagraphs of the Foraminiferan genus Verneuilina from examples
grown in a hypertonic tank, and described some further results
that he and Mr. A. Harland, F.R.M.S., had obtained in the course
of their investigations.

Miss Joan B. Proctor, F.Z.S., communicated her paper on
“The Variation in the Number of Dorsal Scale-rows in our
British Snakes.”

Dr. G. A. Boutenerr, F.R.S., F.Z.8., gave a résumé of his
paper on “‘ Some new Fishes from near the West Coast of Lake
Tanganyika.”

* 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 subseribe to the Publications; but it may be obtained on the
day of publication at the price of Siapence, or, if desired, sent post-free for
the sum of Six Shillings per annum, payable in advance.

32 ‘

In the absence of the Authors, the following communications
were taken as read:—Mr. Guy Marshall, D.Sc., F.Z.8., ‘‘ On
the Species of the Balaninus occurring in Borneo (Coleoptera,
Curculionide)”; The Hon. Paul Methuen, F.Z.8., ‘ Description
of a new Snake from the Transvaal, together with a new Diagnosis
and Key of the Genus Xenocalamus, and of some Batrachia from
Madagascar.

Prof. J. P. Hitt, F.R.S., F.Z.S., exhibited and described a
series of lantern-slides illustrating the placentation of Zarsius.

Mr. R. I. Pocock, F.R.S., F.Z.S., exhibited a series of -
lantern-slides illustrating some of the external characters of
Tarsius.

The next Meeting of the Society for Scientific Business will
be held on Tuesday, November 18th, 1919, at 5.30 p.m., when the
following communications will be made :—

pip Bowen Guus Lover, Bt Eas:
Exhibition of the skull of a Beaver.

Major J. Stevenson Hamitron, C.M.Z.S.

Field-Notes on some Mammals in the Bahr el Gebel, Southern
Sudan.

(1) The Development of the Mesenteries in Urticina crassi-
cornis (Actinozoa).

(2) The Leptomedusan Melicertadiwm octocosiatum.

M. Turner, B.Sc.

On the Nematode Parasites of a Chapman’s Zebra.

Rev. A. H. Cooks, F.Z.S.
The Radula of the Mitride.

Lt.-Col. S. Moncxron Copeman, F.R.S., F.Z.S.

Experiments on Sex Determination.

Cuas. F. Sonntac, M.D., Ch.B., F.Z.S.

The Variations in the Digastric Muscle of the Rhesus
Macaque and the Common Macaque.

_E.S. Russenn, M.A., B.Sc., F.Z.8.
Note on the Righting Reaction in Asteria gibbosa Penn.

The Publication Committee desire to call the attention of
those who propose to offer Papers to the Society, to the great
increase in the cost of paper and printing. This will render it
necessary for the present that papers should be condensed and
be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should
be addressed to

P. CHALMERS MITCHELL,

Secretary.
ZOOLOGICAL Society OF LonpDon,
Recent’s Park, Lonpon, N.W. 8.
November 10th, 1919.

a a

k i Hae

i
Valen

ve
ai.

No. 198.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON”

November 18th, 1919.

Prof. E. W. MacBrins, F.R.S., F.Z.8., Vice-President,
in the Chair.

The SecRETARY read a Report on the Additions made to the
Society’s Menagerie during the month of October, 1919.

The Secrerary exhibited and made remarks on a photograph
of a White Tiger.

Sir Epmunp Gites Lopsr, Bt., F.Z.8., exhibited and made
remarks on a series of skulls of the Beaver, exhibiting a separate
ossicle between the parietals.

Major J. Stevenson Hamiron, C.M.Z.8., gave a résumé of his
paper “ Field-Notes on some Mammals in the Bahr el Gebel,
Southern Sudan,” and illustrated his remarks by means of a
fine series of skins.

In the absence of the Author, Prof. J. F. Gemminy, M.A., M.D.,
D.Se., his papers (1) ‘The Development of the Mesenteries in
Urticina crassicornis (Actinozoa),” and (2) “The Leptomedusan
Melicertidium octocostatwm,” were taken as read.

The Rev. A. H. Cooxes, Sc.D., F.Z.S., gave a réswmé of his
paper on “ The Radula of the Mitride.”

Dr. Cuas. F. Sonnrac, F.Z.S., communicated his paper on
“The Variations in the Digastric Muscle of the Rhesus Macaque
and the Common Macaque.”

Mr. E. S. Russert, M.A., B.Se., F.Z.S., communicated his
paper on “The Righting Reaction in Asterina gibbosa Penn,”
illustrating his remarks with a model showing the ‘* deadlock”
position in the righting reaction, caused by the middle ray
lagging behind.

In the absence of the Authors, the following papers were taken
as read :—Lt.-Col. S. Moncxron Copeman, F.R.S., F.Z.S.,
“«¢ Experiments on Sex Determination”; M. Turner, B.5Sc., ‘“‘ On
the Nematode Parasites of a Chapman’s Zebra.”

The next Meeting of the Society for Scientific Business will be
held on Tuesday, February 10th, 1920.

A Notice stating the Agenda for that Meeting will be circu-
lated early in February.

The following Papers have been received :—

R. E. Turner and J. WATERSTON.

A Revision of the Ichneumonid Genera Labiwm and Pecilo-
cryptus.

F. D. Wetcu, M.R.C.S.

Remarks on Forster’s Milvago or Carrion-Hawk.

EK. Heron-ALLEN and A, HARLAND

An Experimental Study of the Foraminiferal Species
Vernewilina polystropha (Reuss), and some others, being a
Contribution to a Discussion ‘* On the Origin, Evolution, anil
Transmission of Biological Characters.”

The Publication Committee desire to call the attention of
those who propose to offer Papers to the Society, to the great
increase in the cost of paper and printing. This will vender it
necessary for the present that papers should be condensed, and
be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should -
be addressed to

P. CHALMERS MITCHELL,
Secretary).
ZOOLOGICAL Socrery or Lonpon,
ReEceENt’s Park, Lonpon, N.W.8.
November 29th, 1919.

Exhibitions and Notices (continued).

16. A List of the Snakes of West Africa, from Mauritania to the French Congo, By G. A.
Bovieneer, F.R.S., F.Z.S. (Text-figures 1&2.) ..... SUG UD Ite Eat ponboce dc

17. A List of the Snakes of North Africa. By G. A. Boutencer, F.R.S., F.Z.S. ........

18, A Description of New Species of Zeuglodont and of Leathery Turtle from the Eocene
of Southern Nigeria. By C. W. Axprews, D.Sc., F.R.S. (British Museum, Natural
History.) (Plates I. & II. and Text-figures 1-3.)...

See ere ee se ee ee oe es OH OE HH eo Oe

19, On the Occurrence of Denticles on the Snout of AX%phias gladius. By J. Tuornton
Carter (Hon. Research Assistant, Department of Zoology, University of London,
University College). (Plates I.-III.)

20. Crustacea from the Falkland Islands collected by Mr. Rupert Vallentin, F.Z.S.—
: Part III. By the Rev. Tuomas R. R. Srespine, M.A., F.RS., F.LS., F.Z58.
(Plates T-V. and Dext-fioures 1-8.) 5... 00 ee ce cee e wee nace ene ccceoncseteruses

ee cere FF ee GEOFF e ee BH se ee FF oe ee ee sets oe

21. Field-Notes on some Mamuinals in the Bahr-el-Gebel, Southern Sudan. By Major J.
Srrvenson Hamivron, C.M.Z.S. (With Chart.)

22. Descriptions of a new Snake from the Transvaal, together with a new Diagnosis and
Key of the Genus Xenocalamus, and of some Batrachia from Madagascar. By The
Hon, Paut A. Metnven, M.A., F.Z.S. (Text-figure 1.)

ee ee ee Oe ee POOH oe OH ee HH Oe

23, On the Variation in the Number of Dorsal Scale-rows in our British Snakes. By Miss
Joan B. Procrser, F.Z.S. . (Text-figures 1-3.).. 2... cscceccecccrccsecerescerenee

24, On the Species of Balaninus occurring in Borneo (Coleoptera, Curculionidx). By
Cryo ee kGrViansmanre DLSc., HeZis. CPlates Wl, QUE an eccrsye siepeiciele selec © see elelloe

25. On some new Fishes from near the West Coast of Lake Tanganyika. By G, A.
Bouxencer, F.R.S., F.Z.S. (Text-figures 6-10,)

ee sre ce PF eoee Fee oe eese et se ee oe

26. The Radula of the Mitride. By the Rev. A. H. Cooxs, S8e.D., F.Z.S. (Text-
Merares JUS) sce i ee coer e canes ns eee eneee cere cece f

27, Note on the Righting Reaction in Asteriva gibbosa Penn. By E. 8, Russrxt, M.A.,
RSPR EY Lite nite occ tele wince ns 3" os

28, Experiments on Sex Determination. By Lt.-Col. 8. Moncrton Corzmay, M.D., F.RS.,
RIES eee aaa nist oe ieee aU Sila pmtarvlellos, sfeicie Rewiele'e

29, The Variations in the Digastric Muscle of the Rhesus Macaque and the Common
Macaque. By Cuas. F. Sonnrac, M.D., Ch.B., F.Z.S., Anatomist to the Society.
(Text-figures 1-5.) 2... cece cece cece cere cece erersereeere bogac

ese 088 ee ere ce oe

30. On the Nematode Parasites of a Chapman: s Zebra. By Miss M. Turnzr, B.Sc. (Text-
figures 1-6.) ... ®Coecrseecensre ere eet eee ee ee ee ee sere ee ee se oe @82ese ee 2982 see 8 ee oe oO

31, The Development of the Mesenteries in the Actinian Urticina erassicornis. By James
F. Gewuiiy, M.A., M.D., D.Se. (Text-figures 1-5.) ....... Soishatavetens acts wale ui teleebers

32. The Ciliatiow of the Leptomedusan Melicertidiwm octocostatum (Sars). By Janus F.
Guuint, M.A., M.D., D.Sc... (Lext-figure 6.) 65. seeseee cere rete teen erences

Mitlepage ......+-. efepeetet se no en obra se oO dle ceouandas Uulmod seldouotdion oGod gobi sheets
Listiof Council and Officers 2... 252.6... 00c5 eee Seine Siadeisjerte eset SPU CAM ACME de oo
AEE IMO COMES) e (ory stature igi sls tip aleve oyece, 6 ow! eiesie vue cloln) «\e\ nic 4 An)

Alphabetical List of Contributers co cece cece cece eee cree cet n ee tee teeeeene ee Src
Tides RM ercreh gel suse k natary oes en. 6! si mares sass TOS CA GS HID omiGdic cutee sciocrce ccia bir abn

Page
267

309

433

437

441

Vil

PLATES.

1919, Parts III. & IV. (pp. 227-499).

Plate Page
ANDREWS: dag alee TOE AAHIS IAN OAc ener melee Ma coh ce Ne eran Gar Als 2 209.
Ts Cosinocheltys: COllot iin thease teed ce ee eae eee J 4
Carter: Tee Denticles of Mi pnias Gla iis vate ak.s ree. ae ee
Il. Denticies of Histiophorus ..... ........0 fas
III. Teeth of Blennius and Pagellus ........++.+..00..
STEBBING : I, Pelterion spinosulus (White), juv. ....-......... ft)
LV. Zoea ofia Brachvoran. ome oon. <b ee oe lee oe
III. Megalopa of an Oxyrrhynch ....-.-.-2.....00.e008 327
IV. Tanais nierstrasei, sp.M. .... +2120 ee eee tae |
V. Munna antarcticus (Pfeffer) .... 2... .cccee se eee ee J
Peo ALe 7 \ New species: of Balamiittse sot.en a. 0cs ae 1 tn ene 365
Hi: I. Fetus and placenta of Tarstus ..............-.2:.. 476
NOTICE,

The ‘ Proceedings’ for the year are issued in four parts, paged consecutively,
30 that the complete reference is now P. Z. 8.1917, p.... The Distribution
is usually as follows, but on account of war conditions Parts I. & IT. are
issued together :—

Part I. issued in March,
te dlls June.

| Evo lB fh < September.
er als paling a, December,

‘Proceedings,’ 1919, Parts I. & IT. (pp. 1-225), were published
together in September, 1919. .

The Abstracts of the ‘ Proceedings,’ Nos. 196-198 are
contained in this Part. j

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