‘ ,
‘
st
!
‘
By nih
,
, ae)
ver
,
Pn ee
Ca fe
et
at
a ae t
are
ee
ft
woke
he
ber iwe ad
ai weve b i bea
Hae we
sree oa
BN ES ee, ee ee ee ee ee
kb WO OF Oe ae ee Ie ee ee ALT HE HOI TD Nhe mye te the dg WE end ME: pA Cok eee Bed HE bate oR ta at oe
Cer ke Ce oe ere sri ryt war ier weer ie wot ee Mn ee ere ee ee
Ce a Pere) re eee ee Se eee i eee ee ie ee ee rn ee ete rarer Newt ae vie ire err Brit ty ety
A er ek eee ie Sr Cr er Te oO Oe ewer 6 er Moet ory Aan ias een le erik ie 9
Ce Le RARE Ae Ek OQ Ok Rh a Gee ae MPL ge ew Bik hrbesi) otee: Berg irh va meas
te ce
ewe eae ee eee woe Noe ene ere ie eC eee ene * HER ee we ge Op
ren a) reer. See eke ie ee ee eee ea see eae bese tg Re get ME a ARs OL oln Oana, Hs Hye Anh
errr rere mre era ee Pe ee ere Me Wer ee er We ee eh Pe ee
nie hi rere ee ern Si 0 ee earner Ser tir te ee ee ee oe ee ae
re) Pee Wet ee eee oe
SC ee ee ee eee ee eC | Aer eee Cre rs oe ee
pees Parent eee er ree ae ie Mer are Tee oe re Te ee ee ee edd eee sonny
Briere tein ir | eee NOL be Ae We Cee EO OO Od, AM. © Gb here 44. Oc AUR MELO eh we prereset er ict)
See ee re arate ‘ we Seen ar freee tere ene Se eer eee Wee re CEC end
Ce ree a 144 et ee on + , Loh bok mee age awe ee ms
Cree eer AEE Ag Reka ep nthe ee Pe Meee Sn eT ere BC oreo tae de a
earn arate evr iin erm ken mim CW te here Se Rr ee Pe i re err ee oy Maren ty teste ee Rew t Fak Mir ideks Weel
rarer me eC Me SC me ee Ce re eed ee nee te ty teeth nw @UNetLee NUMA. We ws Me kb MB yah Mele De A. wmatew a)
ra i ir ah a a re cern ir ck he ik che ee eee wee ee Ce ee CTE OPO e ee ea
ee er a ee eC Cas
owe Piha Preys
Serer: wire es he te ee eee ee en ete
Pere ae ree te See eC es eee ee
a few een oh SNA ter ies ee 8 gy ih GB M0 Ade. OL ei yo ow Ts be
Cee SC ie See ere eee ee cee CeCe oe eee
Ne A A te Reig eg tg erm ty meet
Ce ee ee ee ee a
sees
Ce. ee
eee ee ee yee ees
ree Pee een
wane Vn At owe Pt Ee nn P Memte tet .
pee oA Eas gif AO sey tanteine de gk Menesipt MA HM Aw, Farad ot mtn
teen (earn ere: fe Teeter ieee Metre Tne Sc ae ne acer re Were her)
Bet ae mM he tM
eo
eT eer Mr Ce eRe ec wre TO Mrer RCM Ct eke el alas
THA en eee ee er Sone he eo
ater aku 40,4 ANOS large ik hy dee Buk’ NOW A aed eres asia tats Bee Mee et
tree Cee ns ee te Oper win aint Marre Tome art hy er a
rere Cr wee oe OO ee ee Femi GA ny MMe
at avags . ee ae rete metbi tt Fw hee ee toon bot
4 sae re eC ec ee ee) eee rrr reenter a
- oa How bee Lae WE RR a a ue me eit Pet bathe ot ”
‘ vee Ppa ge See er Cn Mee Cees poreey Carrera rer in ey
ean a CEE EH Fe Rhee ey Ca eee) rere eee
ob LAE Sie MAC wR a ke Qui heese, ALAS 4 bee peed TANT OR ae ee ey nee ee ce ee Coen ee a
CE EE Be Or abe yt eyes
re meen ete eee eit
bee
++ rire remeewe oC nee, Wir irs Meee ete LS tre ea
teow Pes S SOe oe ee ee
vi * are Vem de ara
hohe my re ere eee ae emerge reine fete k
te ae He ene eo oe
Fk ele et rk mie eee We ork & te Msenet ahome phe ae me
ee ee Yt he ages WR Amato tee tem ap ati de
(ener x tien) pepe ween a kot ie we Se Oe eo
Cee or it fier qete oe
ees ee ted betel
Yet ge > Meds Aeyeorer
NEE Re Meme mn Be te ape et
ah ube at
ae
2 EEE Wee
wt ited PN tte tag he
Cerny aie are SU yo er
‘ Sonn ee SS eb ete eons ”
Lee ey ee ep ee geen rere eee mer eee econ
Pee ere ee rc Wr oe a) SS ehh ted em ST ie oir
yh bytes An
Dem ees
DUPE MERE ee ee Rode
SN Rm eee he
PE Mike tote
eer Cnr Macon tet Yet A le
ny teow Pagan ree ve eM aor
hee RR ATE thet ete
Pevrer Manni irr: nce Se i oo veer ei a 2
SNM aN IN A aT ante taety MIM Gee gta tbe tery
Sr Site Tie Pett en ee ee
ash te Saad eas
SP ey
ey th Motes &
Bertheibee!
Hee
4. oh
Vite betig eae mem gr eer git ets
St eb. bed” vo ee tothe tetewe eet tele a te
S Sere Roce ikelh Br itaie le te tetagrova fle tha fee
Rye be Lp St eae e Meme ttersigrt
Wie Pea hyge ty Boite bette Be foley serait be eg
Patino atria eee Ort ie eee ere
Seve ee rr ee
A oaeagte ee ote bets
FO Sete te de he te te
eros
raed:
FAs es he eS tes
Vv one
Caras one Pe eee
mat em
Sy eos
sete 8
ergrey irk)
ee ere ene
seme Sd
4g eet Sth
elBry
%
aoe #
vy
atsteee
Wats dees
VAR FRB Was D3 ee) osaay a
wee
VHB |
bd ee ph deat
It ae es a Fas
a inh Them
Peery ee rn en ae a
. aft Re
wee Od
sheet
atest
sabe tgler ie ab hin we Oe
ai Dae aw Odd eae ede Rb 38
Becks dg ded aie &
Pa ee ed
sare hain
Fo Ryd ase
ns rte
’
error etme t |
He ae ee anh den
G0 al weed Foot bet ia
eae deed
ee Lr een
ee ee ee ae
daw We me eet &
Dodie dO pbs 49 Mtns AG
Leyes o deat ;
ee ee weed ate
ee
oie
Bab abe ae mean tnt ary
aes
*
we er
ad mie ge ee
Pierre ee ec
Ce ed i te
oe
Brera eee a ort etd
Pareene trae iar rot td
Sale Scare ee
Ad wd ween
tay Gata tpl ee ate
Biro
bern i ways ‘ we
- re) oy eat wea vein Wee
ne Aeurrrcers werrr writ incr Tie ers
. a Ase BAAR a?
. ¥ ee ee ee
coc Wieoarc rere retiree liar ir
ne ere re ie ae ar a
Vague eenwrrrerarer es nv tr me rieaers Caen oie
lie ey fb soc sbaee ol otal wh AL oe ei e ey Piet en
Wom ae memati Me a
Tene) ee ema et ott
ae tek eg bates wet ae
STERN Ms OETA ee Aah AOR A Havas Laem res ee
BB) Be hen i tn aie ar Gie ae Rene eee
Pereror mr mrer rit matt er te ad i Meth
CR ae erie ee ne ee)
ro ee Ae hee Oe Tees
eine ges
ee wre eee wee Ane t
pera we eae ry Sees cL teaches: aha ea de Bae Prone semen ttre or arr ee
wn 2 Le jeoee x
Perera SNe Cnet en fe ee hi St pA haw ay felipe ede
Preston ron ae cee
eomrae oe ent @ whee bea arom em
ak ek ek ea
Per ee ne
Peer Re Se a ee
eee be Bb Da A dae anne ie! Bd etAle nt at
aca cect Perr
Pe pd *e
or
Pernt ret So ee
Pee Se
Pe ee
a aoe ODD)
Le pene eee se mens roe rete
Ly ee Rarer tener ewer een pert
yea Gee cea a TA EAs guna ado ataetieal aimed acuri sad @vihlr amine @ uti im @. hbo Oo ari
Pera ee ee oe Se Anbrere wren y rie 0 ere ert eto ee “4 Seer Bak aa anon oe mead abn at aba at)
Ree eae a ire 2 ts Nee ee ar Sd ot hh moras ane ae trie b yivede Ra bleamGecdd whsye Sie yi Qarelzre mat Lod aitante Gas ete dt Gate
Fine Aas Gaara tea ae hs Sts Latin tt telethon
: x aid ND ts aia aA Pre see rh | POULT n et LOM tC ik CCB hal ae aaah We
ae ree PYRE RRC n i ie ari | Rarer er fy bt fr) Ere IL’ Hae ent prnbaeiamrnenniien LYE ete,
ER ee ee ge LAV PEO a8 8 Og wae meas ds arvetecowile menssshgees road oh wh RG Us 1 prs Oa
Ser Se Or en ee 1a art Gre eye Wehr ar gere ws
‘ ka eb nal be eae Bera Perera eet foes Bateres trace we bed
24 Pay ey i ar ee eS sada fer erer Steen ee err wert ht ke
sad Vea eek aren rice ee ieee Se ee eC Cerri wr i rk neni s ‘i
sev bet ew ete tae ae Ss a2 ch Ay di oh cee Bat we seca we gtr ee
Saree erin PNT ar ee car ret ie ates he
. oF : + aa ded me te ee ee
tes ten ie Ek aed RH tt aed eeAhl ahi pI IE AS oe awed 48 he Pome 0 9 fe ie
Perce eer ea ore aa er ceak eae S08 GT Gat taal Uh eee LU SST MA GU nL ame oar og ee ME A A ot
4 awe e ae) Oyen erin ir eo Me eels Ue ERE. A Oe ig WAP LL Ha he Tent Wes A ft Ba hy
yd fe SNS og pete ee re ee Awe eee a As bee boy me Aas Fad Ut erted or uw creda nya g Medan nee Ne eC
Ferd eras cg BAe Aide Ald, ulep ame Hr waa ed a Be hc BEN Oe ne ote Oren
ror Pinar rae we ee ee ee ey ee é ‘bes oy Ay . wt * are Va wad aah oe eu eee 6 oh ee ipierlete- te Lae wpe ty
a ere rer eee Oe ere eR a) PPI reo) eet wma aed | uh weg ede dhe My! bate ah ge Gelahe WAM Wb) Hy she tis Utag She @ eet eae §
Sor Pee eOn Ay rier OL Bee a me Re te ae asta a de Wor lt Gas 6 thet Pe woth a
Onno eran LaG sah EER Pi A alee tor Whi as aesds ema or ops pee dono say wed 2b OW: aH Tne ¥8/ 8 Oy)
‘ ee a pagreraaaeeronrter inet ar Ment Wel Teer We COS Ded ae teed EES SA Dla bali car arent ane
a oat Aight eee eee ala bam 0's C050D seu ths a 5b AG Wife. presses ees U6 08h eth Bs ihe $20) Os He Olde ESOT OES He
OR ae : a Pe ene nar here or tewene eee Mrer ny tee Nor oe kA leer Karke:
P aM ate a tank hae fee & Peery mm ec ee ne ee ee Ue Padi Seriya sealer
Ad iret ini te Se a tae at aati athe wiety odie. gal hie Rees yy Pree tn anew vam
a ‘‘ i ith Gh tah i arta © ve: aici Gwe ei eee sain eM tar less oe date Ow alg etnew aa bene® Nome
i
Lr oe
ean
ae
PROCEEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF THE
ZOOLOGICAL SOCIETY
OF LONDON.
1922, pp. 1-481,
WITH 21 PLATES AND 214 Trxr-FrictREs.
pei ASONTARSS
of
a ee
SS LIBRARIES
PRINTED FOR THE SOCIETY,
SOLD AT ITS HOUSE IN REGENT’S PARK.
LONDON:
MESSRS. LONGMANS, GREEN, AND CO.
PATERNOSTER ROW
1 Meyish Je
OF THE
COUNCIL
Ohya
AND OFFICERS '
OF THE
ZOOLOGICAL SOCIETY OF LONDON.
1922.
Patron.
His Masusry Tue Kine.
COUNCIL.
His Grace Tan Duk or Beprorp, K.G., F.R.S., Presedent.
Tar Hon. Ceci Barine, M.A. |
Lr.-Con. 8. Monerron Copr-
MAN, M.D., F.R.S.
CHARLES DRUMMOND,
Treasurer.
AuFRED Ezra, Esq., O.B.E.,
Vice-President.
HiucH SS. GLADSTONE,
M.A., F.R.S.E.
Tne Rr. Hon. 'Tun Viscount
Grey, K.G., B.C.
Sim Sipnry F, Harmer, K.B.E.,
WigsNe, IDSs, Ilatsses Ge
President.
Masor Tur Lorp ALASTAIR
2L0BERY [nnES-KeEr, D.S.O.
Pror. Ernest W. MacBripn,
IDisies,, IbAb Day Wotton, Wace
President.
Ksq.,
Ksq.,
Cou. Sir A. Henry McManon,
G.C.M.G., K.C.1.E., Vice-
President.
E.G. B. Mreapr-Watpo, Hsq.
P. Guaumers MircHery, Esq.,
GRIER, Wiles IDES, Jbl, 1D).
F.R.S., Secretary.
Tue Hari or Onstow, O.B.E.
Masor Artzverr Pam, O.B.E.
Lue Lorp QUEENBOROUGH.
THe MarquEss oF SLIGO,
F.S.A., Vice-President.
Pror. G. Euiior Suire, M.A.,
MED baie Ss.
Ricwarp 8. Taytor, Esq.
Anvuony H. WINGFIELD, HsqQ.
A. Smrra Woopwarb, Esq.,
LL.D., F.R.S:, Vice-Pre-
sident.
PRINCIPAL OFFICERS.
P. Caatmers Mircueny, C.B.M., M.A., D.Sc., LL.D., F.B.S.,
Secretary.
R. I. Pococr, F.R.S., F.L8., Curator of Mammals and
Resident Superintendent of the Gardens.
D. Sers-Surru, Curator of Birds and Inspector of Works.
Epwarb G. BouLuncer, Curator of Reptiles.
Miss L, HK. Curusman, F.E.S., Curator of Insects.
C. F. Sonnrac, M.D., Anatomist.
N.S. Lucas, M.B., Ch.B., Pathologist.
R. J. Orrterp, M.A., Hon. Parasitologist.
F, Martin Duncan, F.R.M.S., Librarian.
F. W. Bonn, Accountant.
W. H. Conn, Chief Clerk.
LIST OF CONTENTS.
1922, pp. 1-481.
EXHIBITIONS AND NOTICKS.
The Srcrerary. Report on Additions to the Society’s
Menagerie during the months of November and
Mecem berimlODil ppaknAn ep Ai wanes 6 Lie aes cake
Mr. A. H. Evans. Exhibition of Cuckoos’ eggs taken near
Wanmilorialcerye pe. 3y8 . Sneha ee Nae hk Oy oe a eaie
Lord Ciirrorp or Cuupieien, F.Z.S. Exhibition of photo-
graphs of Wotothervum mvitchellt 2.2.0.6. .00.cccceeesetee es
Miss L. E. Cuznsmay, F.E.S. On the position and function
of the Siphon in the Amphibious Molluse, Ampullaria
BOM UORTUAS ps Serciascrcu ch otha seeuraa eee at hia! ater eh tid:
Mr. G. C. Rogson, F.Z.8. Exhibition of Models demon-
strating the respiratory mechanism of Ampullaria
QEIIE ONMUUSI: as SUAVE A thi ke he Saha doh ite hath Ae Sahin tas bees
The Sucrerary. Report on Additions to the Society’s
Menagerie during the month of January, 1922 ......
The Secrerary. Exhibition of a photograph of the
Society’s Gardens taken from an aeroplane ............
Miss L. KE. Cuzrsman, F.E.S. Exhibition of living speci-
mens of the Amphibious Molluse, Aimpeullaria vermi-
“OUTS, BIOS as SORE MATE SOLO a os See aE ee
Mr. G. C. Rosson, F.Z.8. On the respiratory mechanism
OLE (lave) Ja Uc OU VENICE en bake Aaah Or hares pedal aah seer Cees cee
Mr. F. Marvin Duncan, F.R.M.S., F.Z.S. Exhibition of
cinematograph films taken in the Society’s Gardens..
202
1V
Miss L. KE. Curzsman, F.E.S. On the habits, in captivity,
of the Fresh-water Crab, Cardisoma armatum .........
Mr. C. Tare Rucan, M.A., F.R.S. Exhibition of living
specimens of the Indian Tortoise-Beetle, Aspido-
WO NO SEHOCR AG FONENS © soc shnooens@anoegnads Wedeednaonscoads
The Secretary. Report on the Additions to the Society’s
Menagerie during the month of February, 1922 ......
Mr. C. Tare Reean, M.A., F.R.S. Exhibition of a series
of lantern-slides illustrating specimens of various
blimdsiresh=waiterpMishesy)..0 eken.-cecere see ert aseeeeee
Mr. R. H. Burne, M.A., F.Z.S. Exhibition of specimens
demonstrating the recessus orbitalis in Flat Fishes...
Dr. L. Hocpen, M.A., F.Z.S. Exhibition of, and remarks
upon, some small examples of metamorphosed Mexican
Salamanders (Amblystonva tigrinwm) .......60esece eee e es
The Secrerary. Report on Additions to the Society’s
Menagerie during the month of March, 1922 .........
The Secrerary. Exhibition of some dressed skins of
a Tree-Hyrax (Procavia valida) from Tanganyika
Reh eI 01 Bea toe GP RIO a RPR MAUR HUbcy AaaNododnddencic
Prot. P. T. Fuynn. Exhibition of, and remarks upon, a cast
of the skull of a Squalodont Whale ...................4.
The Secretary. An account of Mr. Loveridge’s experiences
while watching “ Lions at their Kill”
eee ter ee cee wee ees
Page
203
203
479
479
A79
479
480
481
481
bo
ite
12.
PAPERS.
. The Fauna of East Africa and its Future. By C. W.
ROBT EW. (OVI G OHM Zins. biarhs,srascelectne nae atete a aitretes
. Onthe Temporal Arches of the Reptilia. By R. Broom,
D.Se., F.R.S., C.M.Z.8. (Text-figures 1-4.) .........
. Animal Communities in the Southern North Sea. By
Tab; [sieitenyiw, Wye edailk, (uesxteabletingey Wy Goncoosacuocobe
. Evolution within the Genus Dendronephthya (Spon-
godes) (Aleyonaria), with descriptions of a number of
Species. By W. Rat Suerrirrs, M.A., D.Sc., F.L.S.
(PlatesMcS il) Nexto ures pl 30))i (ae seenuesseses. ee
. The Life-history of the Water-Beetle, Pelobius tardus
Herbst. By Frank Bartrour- Browne, M.A.,
IDS eehoel Dayle Ags loan (Leena) ta Le LIL) Wade ere ganwetann sous
. On the Vagus and Sympathetic Nerves of the Kdentata.
By Cuarves F. Sonnrac, M.D., F.Z.S. (Text-
JBSUORSYS ALEETS i ca La NP etna CE an cs
. Contributions to the Morphology, Classification, and
Zoogeography of Indian Oligocheta. By J. SrspHen-
son, M.B., D.Sc., F.Z.8. (Plate I.; Text-figures 1-19.)
. On the Vagus and Sympathetic Nerves of Hyrax
capensis. By CuHartes F. Sonnrac, M.D., F.Z.S.
(Hest eM MBO SOS) ea a anise tenes) « Maps tad aeatn ey
. The Cichlid Fishes of Lake Victovia. By C. Tare
Regan, M.A., F.R.S., F.Z.S. (Plates I.-IV.; Text-
ICSU Sl lho) ane APE PORE aR a one aici ana naran re
. Direct Development in a Dromiid Crab. By SrepHEen
K. Monveomery, B.A., B.Sc. (Text-figures 1-3.) ...
Notes on the Life-history of Cweulus canorus, with
exhibition of Eggs. By A. H. Evans, F.Z.8. .........
A Revision of the Nematode Parasites of Elephants,
with a description of four new species. By M.
Kuauit, M.D, D.P.H., D.T.M. & H. (Text-
FICHUIMOS Mel lene aise a eae es UNUht nid sulliitCid hte
Page
79
wy)
109
149
157
193
197
205
13.
14,
15.
16.
Wie
18.
AOR
bo
bo
23.
V1
Report on the Deaths which occurred in the Society’s
Gardens during 1921. By N.S. Lucas, M.B., F.Z.8.
Some Spiders from South Annam. By H. R. Hoee,
Wh Nog Ate ) UNepaserthernnes dkI\0)5) Saassgecstotoanss0853
New Reptiles from Tanganyika Territory. By Arruur
MovERTDGE, ©. MNZiSe (ee aeeey wen. od ee uence eee
Notes on the Zebras and some Antelopes of Angola.
By Gineert Buarse, ¥.Z.8. (Plates 1—VIIL.) ......
Notes on the Respiratory Mechanism of the Ampulla-
mde, 9 ByiGuy C. ROBSON, MAC REA /Z2>) anne ee eee
Monkeys and the Fear of Snakes. By C. P. Cuaumrrs
Whinerenoin. [DyStese DIL ED ES Walesiee, C)8)J91, ccoccosancoosse
Mendelian Experiments on Fowls. II. Production
of Dominant Pile Colour. By J. T. Cunnincuam,
MA B28; \(Blates 1-11.) oe. ayence en momar eres
. Observations on the Land Crab, Cardisoma armatum,
with especial regard to the Sense Organs. By
inser TORO (Oinaimee IMG pes gyoeeanocosoacaccccosesacaa04 3
. On the External Characters of some Hystricomorph
Rodents. By R. ft. Pococw, F.B.S., H:Z.S. (Wext-
ITV UU Gs lca) Eee Xo) Mae Re eR ee ARE PUBL AR PR Dis Noanondacdace
. On the Anatomy of the Drill (Zandrillus lewcopheus).
By Cuartes F. Sonntac, M.D., F.Z.S. (Text-
foes, OPA rss easiest MeN opt oo aac a eee ee
On the Persistence of the Mesopterygoid in certain
Reptilian Skulls. By R. Broom, F.R.S., C.M.ZS.
(Text-figures 5-7.)
Cee eee wer eee ew neers reece ne sere res ene eseree
. A New Species of Gisophagostomum (Wsophagostomwum
wert, sp. n.) from a Rodent (Xerws sefosus). By
R. J. Ortuepp, M.A. (Text-figures 1-6.)
eer eee ne ee
. Note on the Bat-Parasite, Cyclopodia greeffi Karsch,
and on a new Species of Hymenopterous (Chalcid)
Parasite bred from it. By F. W. Unica, Hueu
Scott, M.A., D.Se., and J. Warsrston, D.Sce., F.Z.S.
Wextetioumes 1). i860). PON Oe Ae EA ni a
Page
281
285
313
D7
341
B47
349
361
365
461
471
ALPHABETICAL LIST
CONTRIBUTORS,
With References to the several Articles contributed by each.
(1922, pp. 1-481.)
BLAINE, Gineerr, F.Z8. oe
Notes on the Zebras and some Antelopes of Angola.
(GR reS ils VAIPELS iret et ec ES eee tL) ENE aE Ce 317
BurevaD, H., Dr. Phil.
Animal Communities in the Southern North Sea.
(GV ce IST NY etsy cro. rera wr accls snpacse.c od maya Mea eee eae e ose rte 27
Broom, R., D.Sc., F.R.S., C.M.Z.S8.
On the Temporal Arches of the Reptilia. (Text-
Ue A rate taal Gacsta SU ter ene pare tame we bh 17
On the Persistence of the Mesopterygoid in certain
Reptilian Skulls. (Text-figures 5-7.)..................00008. ABD
Browne, FRANK Batrour-, M.A., F.R.S.E., F.Z.8.
The Life-history of the Water-Beetle, Pelobius tardus
Hest Ueeubese ll WI co 5 Sota quan gy seinen. arieiactate salen eee 79
Burne, R. H., M.A., F.Z.S.
Exhibition of specimens demonstrating the recessus
OUD Mal I Mend NVSINES Gry eda rinsuaunoncnnnn damon eemonasa a aseet 479
vill
Page
CuereEsman, Miss L. E., F.E.S.
On the position and function of the Siphon in the
Amphibious Molluse, Ampullaria vermiformis .........+4. 202
Exhibition of living specimens of the Amphibious
Mollusc, Ampullaria vermiforniis.............00ceesnene sas ore 203
On the habits, in captivity, of the Fresh-water Crab,
OLA RUOSOTOCMOTALOLLIGHIO: spticso aa shoe Roo npEpaoac OA sbobbmenasAgn nboooC 203
Observations on the Land Crab, Cardisoma armatum,
with especial regard to the Sense Organs .................. 361
CrupieIeH, The Lord Ciirrorp oF, F.Z.8.
Exhibition of photographs of Vototheriwm mitchelli ... 202
CunnineHamM, J. T., M.A., F.Z.8.
Mendelian Experiments on Fowls. III. Production
of Dominant PileiColour:) (Blabes) 1h) ete ee seaadee 349
Duncan, F. Martin, F.R.M.S., F.Z.S.
Exhibition of Cinematograph Films taken in the
Socretys Gardens | i..6 5.0 ae eee eee eee 203
Evans, A. H., F.Z.S.
Notes on the Life-history of Cweulus canorus, with
exiiloitonvotwhiogs:-.... cin tae teak. .a6b ke eee eee ee eae ach SOM
Exhibition of Cuckoos’ eges taken near Cambridge ... 202
Bova. Prot Pet.
Exhibition of, and remarks upon, a cast of the skull
Olansqualodonth Wihaleseiser cs. coss sce. 4s enna eee eee eeeee Gee Leolt
Hostey, C. W., C.M.G., C.M.Z.S.
The Fauna of Wast Africa and its Future ............... i
Hoesen, Dr. L., M.A., F.Z.8.
Exhibition of, and remarks upon, some small examples
of metamorphosed Mexican Salamanders ( dinblystoma
IGE UO CHT) SUSUR ep SEN eR PEN a RRO an dee DMEM SRG 5 479
Hoe, H. R., M.A., F.Z.8.
Some Spiders from South Annam. (Lext-figures 1-10.)
Jeet Niitivg, UG, WED Da ebiS ins DEOMI Leaccg dal,
A Revision of the Nematode Parasites of Elephants,
with a description of four new species. (Text-
HERTNE SUES abana ccc c tence ee con mDM ERE na oc pon SU MOeoer
LoveripGcE, Arruur, C.M.Z.S.
New Reptiles from Tanganyika Territory ...............
Lucas, N.8., M.B., F.Z.S. (Pathologist to the Society).
Report on the Deaths which occurred in the Society’s
Gardens durrimon lO 2 1.) iacccmam cep mcactyaae ites? as sacle Hone salsa
Mrroueny, Dr. P. Cuaumers, C.B.H., F.R.S8. (Secretary to
the Society).
Report on the Additions to the Society’s Menagerie
during the months of November and December, 1921 ...
Report on the Additions to the Society’s Menagerie
dlimime thepmonthvom January, LO Aeenasees essence nent ser
Exhibition of a photograph of the Society’s Gardens
falkcermatrccrmmelmae Op larlel ee eee ner nee eee. ae aeee =
Monkeys amd thie Mearof Snakes senses ee serere :
Report on the Additions to the Society’s Menagerie
during the month of February, 1922) .....5......5...cee00 +0
Report on the Additions to the Society’s Menagerie
dunimesthemmonth of Maxehy 1922) ies ceeseaeennene cca
Exhibition of some dressed skins of a Tree-Hyrax
(Procavia valida) from Tanganyika Verritory...............
An account of Mr. Loveridge’s experiences while
Watchmaceabionsrat ther Keil (Mh eae ian en lt cculye en:
MontcoMErRY, SrtepHEN K., B.A., B.Sc.
Direct Development in a Dromiid Crab. (Text-
BAUCUS LON) ug. cect tcncd oeyatetn ance eet gangaana rn va sla
Proc. Zoo. Soc.—1922. b
Page
205
315
281
480
481
x
Ortierp, R. J., M.A. (Hon. Pathologist to the Society).
A New Species of Gisophagostomum (Hsophagostomum
xeri, sp. n.) from a Rodent (Xerus setosus). (Text-
\
Ao uMes L265) 0 eco 5 5). Hees eee kt tele Sates oe See Ee ee
Pocock, R. I., F.R.S., F.Z.S. (Curator of Mammals).
On the External Characters of some Hystricomorph
Rodents... (lext-teures) Ie28.) sce." ...cchue waeseeee ee ane
Reean, C. Tarn, M.A., F.R.S., F.Z.S.
The Cichlid Fishes of Lake Victoria. (Plates I.-IV.;
Mextstioures WA) ee cites «ce cil oae eee ieee eRe eee
Exhibition of living specimens of the Indian Tortoise-
Beetle, Aspidomorpha sanctoe-Cructs ..........0..ecersseee econ
Exhibition of a series of lantern-slides illustrating
specimens of various blind fresh-water Fishes ............
Rogson, G. C., M.A., F.Z.8.
Exhibition of models demonstrating the respiratory
mechanism of Ampullaria vermiformis ........224- v.10.
On the respiratory mechanism of the Ainpullaride ...
Notes on the Respiratory Mechanism of the
rtp willl a eas". 8 i'. arinvaigaaln ounce ate semen cree Re ee aR eee
Scorr, Hucu, M.A., D.Sc. See Warerston, J., D.Sc., F.Z.S.
SHERRIFFS, W. Rag, M.A., D.Sc., F.L.S.,
Evolution within the Genus Dendronephthya (Spongodes )
(Alcyonaria), with descriptions of a number of Species.
(Blates) IID. Pext—-figureswl3.0);) 22a eee eee ease
Sonnrac, Cuaries F., M.D., F.Z.8. (Anatomist to the
Society).
On the Vagus and Sympathetic Nerves of the
Bidembatae u(dlext-fieuires 10.) aleasaopeere: bea er pees
On the Vagus and Sympathetic Nerves of Hyrax
Capensrs, exit eUres)O—O4) sneak tener are eee eee eeaeEe ee
On the Anatomy of the Drill (J/andrillus leucopheus).
(Ressbafionies O22 4) octane a eee eRe ee ea
Page
461
369
479
33
99
‘
x1
Page
STEPHENSON, J.. M.B., D.Sc., F.Z.S.
Contributions to the Morphology, Classification, and
Zoogeography of Indian Oligocheta.—IV. (Plate I.;
PEG TT OMT Ss Me DE oy ea 8 cara cie ware eaaneeeramEM Meenas ook 109
Unico, F. W. See Warersron, J., D.Sc., F.Z.8.
Wavrerston, J., D.Sc., F.Z.S., and others.
Note on the Bat-Parasite, Cyclopodia greeffi Karsch,
and on a new Species of Hymenopterous (Chalcid)
Earagiver bred iron ib. \(WMext=tieure ls): 1. .9st-eees sence: ATi
INDEX OF ILLUSTRATIONS,
Acanthion longicauda, Figs. 20, 26,
pp. 404, 414.
Adrastis lashbrooki, Fig. 5, p. 294.
Agalena doris, Fig. 8, p. 301.
tenuis, Hig. 8, p. 301.
Agama, Fig. 6, p. 457.
hispida var. aculeata, Fig. 5,
p. 456.
Amira pileata, Figs. 65-68, pp. 267-
269.
—— sameera, Figs. 69-71, pp. 270-272.
Antelope, Angolan Sable, Pl. V., p- 317.
S
Antidorcas angolensis, Pls. VI1., VIIL.. |
p. 317,
—— euchore, Pls. VII., VIII, p. 317.
Argyroelos micans, Fig. 2, p. 288.
Atherura africana, Figs. 1, 8, 26,
pp. 368, 384, 414.
Oy
Capromys pilorides, Figs, 2, 9, 21, 27,
pp. 370, 386, 406, 416.
Captorhinus agutt, Fig. 1, p. 18.
Cavia aperea, Fig. 6, p. 378.
—— porcellus, Figs. 4,
407, 418.
Chetogaster, Figs. 2-6, pp. 118, 114.
—— orientalis, Fig. 1, p. 111.
Chart showing Stations worked by the
23; 28, pp. 373,
‘George Bligh,’ 6th-Sth July, 1921, |
Fig. 1, p. 30.
Chinchilla lanigera, Figs. 4, 7, 14, 25,
pp. 378, 382, 594, 411.
Proc. Zoon, Soc,—1922.
Celogenys paca, Figs. 3, 5,7, 13, 23, 28,
pp. 872, 375, 382, 398, 409, 418.
Coendu nove-hispanie, Fig, 20, p. 404.
—— prehensilis, Figs, 2, 4, 10, 11, 26,
pp. 370, 378, 387, 388, 414.
Conodectes favosus, Fig. 7, p. 459.
Ctenodactylus gundi, Figs. 4, 6, 18, 28,
pp- 9738, 378, 400, 418.
Ctenomys imendocinus, Figs. 4, 6, 18, 22,
pp. 878, 378, 400, 407.
Cienus flavidus, Fig. 7, p. 299.
Dasyprocta
pp. 378, 393, 409.
spa’, Bigs. 16,
| Dendronephthya, Fig. 5, p. 40.
-——- annectens, Pl. IIT., p. 33, Figs. 27,
28, pp. 73, 74.
brevirama, Fig. 26, p.71.
cervicornis, Fig. 14, p. 55.
clavata, Pl, I., p. 38, Fig. 7, p. 45.
collaris, Fig. 20, p. 68.
coronata, Fig, 25, p. 69.
— disciformis, Fig. 22, p. 65.
—— ehrenbergi, Fig. 11, p. 51.
jlammea, Fig. 16, p. 57.
—— gigantea, Pl. I1., p. 33, Figs. 8, 9,
pp. 47, 48.
habereri, Fig. 23, p. 67.
—— hyalina, Fig. 30, p. 77.
—— hklunzingeri, Fig. 17, p. 59.
{ (eset longicaulis, Fig. 21, p. 64.
—— microspiculata, Fig. 18, p. 60.
c
X1V
Dendronephthya mirabilis, Fig. 15,
p. 96.
—— mollis, Hig. 18, p. 53.
—— orientalis, Fig. 19, p. 61.
pumilio, Big. 24, p. 68.
—— punicea, Wig. 10, p, 50.
— simplex, Fig. 29, p. 79.
—— suensoni, Fig. 12, p. 52.
Desmogaster, Figs. 11, 14,15, pp. 186, _
140, 142.
Diea shirleyi, Fig. 3, p. 291.
Dinomys branickii, Figs. 6, 19, pp. 378,
402.
Divaricate, a typical, Figs. 2, 6, pp. 56,
41.
Dolichotis patagonica, Figs. 5, 7, 16, 23,
28, pp. 372, 382, 398, 409, 418.
Drawida, Figs, 10, 13, 14, pp. 135, 187,
140.
= enjoomniaga’,, VPN Voy Fos WO),
Hchimys cajennensis, Fig. 6, p. 378.
Krethizon dorsatum, Figs. 2, 6, 10,11, |
26, pp. 370, 378, 887, 388, 414.
Hupelinus urichi, Fig. 1, p. 475.
Huphractus vitlosus, Sigs. 3, 4, pp. 104, |
106.
Hupotygaster, Figs. 12, 14, pp. 187,
140.
Husthenopteron foordi, Fig. 7, p. 459.
Galea littoralis, Figs. 6,17, 28, pp. 378, |
399, 409.
Glomerate, a typical, Figs. 1, 6, pp. 36, |
41.
Haplochromis acutirostris, Fig.7,p.181.
altigenis, Pl. 1., p. 157.
annectens, Vigs. 2,
190.
—~ argenteus, Fig. 12, p. 187.
—— dentex, Pl. 1ED., p. 157.
—— dichrourus, Fig. 6, p. 179.
— gracilicauda, Pl. IV., p. 157.
— longirostris, Pl. LV., p. 157.
— macrodon, Fig. 4, p. 176.
—-- macrognathus, Pl. IIT., p. 157.
14, pp. 168, |
INDEX OF ILLUSTRATIONS.
Haplochromis imaculipinna, Fig. 5
p. 178.
melanopus, Fig. 1, p. 165.
orthostoma, Fig. 9, p. 184.
pellegrini, Fig. 11, p. 186.
plagiostoma, Fig. 8, p. 181.
—-— sauvage, Fig. 13, p. 189.
serranoides, Pl. II,, p. 157.
—— teniatus, Fig. 3, p. 171.
xenostoma, Fig. 10, p. 185.
Hippotragus equinus, Pls. I11., IV.,
p. 17.
niger, Pls. I¥., 1V., p. 317.
variant, Pls. 1.,1V., p. 317.
Hoplotilapia retrodens, Fig. 18, p. 189.
Hydrocherus capybara, Figs. 7, 17, D4.
28, pp: 382, 399, 410, 418.
Hyrax capensis, Figs. 6-8, pp. 150, 152,
154.
Hystrix africe-australis, Figs. 1, 8, 20,
26, pp. 368, 384, 404, 414.
Ichthyosaurus communis, Fig. 5, p. 22.
Kerodon rupestris, Fig. 15, p. 396.
Lagidium pernanum, Pig. 15, p. 396.
Lagostomus trichodactylus, Figs. 7, 14,
25, pp. 382, 394, 411.
Langliana klossi, Big. 1, p. 286.
Leiperenia galebi, igs. 5-7, pp. 211—
213.
--—— leiperi, Pigs. 1-4, pp. 207-210.
Loncheres armatus, Wig. 6, p. 378.
Tumbricus, Fig. 8, p. 158.
Macropleurodus bicolor, Fig. 13, p. 189.
Mandrillus leucopheus, Wigs. 9-22,
pp- 480, 431, 433-435, 437, 458-442,
444,445, 447, 449-451.
Memphisia aziza, Figs. 59-64, pp. 261—
264.
memphisia, Figs, 52-58, pp. 256-
260.
Mendelian Experiments on
Pls We, py S49:
Menemerus dabanis, Fig. 10, p. 308,
—— felia, Fig. 9, ». 306,
Fowls,
INDEX OF ILLUSTRATIONS.
Mizosairus atavus, Fig. 3, p. 22.
Moniligaster, Bigs. 13, 14, pp. 187,
140.
Murshidia hadia, Bigs, 15-21, pp. 224—
227.
—— linstow?, Figs. 13, 14, p.
Myocastor coypus, Bigs. 3, 6, 12, 21, 27, |
pp. 372, 378, 390, 406, 416.
999
aa.
Ocnerodrilus occidentalis, Fig. 9, p. 134.
Octodon degus, Figs. 4, 6, 9, 22, 27,
pp. 373, 378, 586, 407, 416.
(Hsophagostomum wert, Figs.
pp. 462-466.
Oligocheta, Pig. 7, p. 138.
Oiigochzete, ancestral terrestrial, Fig. 19,
p. 144.
1-6,
Palystes ledleyi, Fig. 6, p. 297.
Pandercetes ochrea, Fig. 4, p. 292.
Parabronema smithii, Figs. 8-12,
pp. 216-219.
Pardosa dranensis, Fig. 7, p. 299.
Pareiasaurus sp., Fig. 1, p. 18.
Pelomedusa galeata, Fie. 2, p. 20.
Pelobius tardus, Pls. 1—I1., p.79.
Petalomera lateralis, Figs. 1-38, pp. 195, |
196. |
Placochelys placodonta, Fig. 2, p. 20.
Placodus sp., Fig. 2, p. 20.
Platyteniodus degeni, Rig. 14, p. 190.
Flesiosaurus macrocephalus, Fig. 2, p. 20.
XV
Polyp, Armature of lateral point of,
Fig. 4, p. 38.
Pteridopharyne
pp. 251-254.
anisa, Figs. 47-51,
Quilonia africana, Fig. 46, p. 248.
apiensis, Figs. 22-28, pp. 251-
234.
— brevicauda, Figs. 35-39, pp. 240-
243.
ethiopica, Bigs. 40-45, pp. 244—
247.
uganda, Figs, 29-34, pp. 236-289.
Springbuck, Angolan, Pl. V., p. 317.
Syngenodrilus, Figs. 16-18, pp. 142,
143.
Tamandua tetradactyla, Figs. 1, 2,
pp- 100, 102.
Tatusia novemcincta, Fig. 5, p. 107.
Thalattosaurus alexandre, big, 4, p. 23.
Thrynomys swinderianus, Rigs. 1, 7, 19,
pp. 368, 382, 402.
Umbellate, a typical, Figs. 5, 6, pp. 37,
ails
Youngina capensis, Fig. 4, p. 23.
Zebra, Hartmaun’s, Pl. VI., p. 317,
iat
i
i y
; c 1) OS
Bib, aa
igs ies =)
a | } :
a fs
4
~
ea
1
oy ONO nee SIE
Ba pipe
PND EX:
1922.—Pages 1-481.
[New names in clarendon type. Systematic references in italics.
(z.8.1..) indicates additions to the Society’s Menagerie. |
Abra prismatica, 81.
Acanthion, 371, 383, 405, 412, 413.
Acanthobdella, 117.
Adrastis lashbrooki, <p. n., 294.
Agabus abbreviatus, 88.
didymus, 88.
Agalena doris, sp. n., 301.
— tenuis, sp. n., 302.
Agama hispida, 455.
Aleyonaria, 33.
Amblystoma tigrinum, 479.
Amira, 266.
pileata, 2613.
sameera, sp. n., 269.
Amphiura filiformis, 21.
Ampullaria glauca, 344,
tnsularum, 344.
Ampullartide, 341.
Anomapsida, 26.
Antidorcas, 333.
——- angolensis, sp. n., 335.
Argiope etherea, 290.
Argyroelos, gen. n., 288.
— micans, sp. n., 288.
Ascaridia rodhaini, 279.
Aspidomorpha sancte-crucis, 203.
Proc. Zoou. Soc.—1922.
vermiformis, 202, 203, 344.
Atherura africana, 367, 371, 375, 376,
383, 413, 415.
Axinus flexuosus, 31,
Bathmestomum, 275,
sangeri, 275.
Belascaris, 214.
lonchoptera, 205.
Bidessus minutissimus, 83.
Birgus latro, 345,
Bradypus tridactylus, 99,
Bubo bubo (2.8 1.), 481.
Bunostomum, 275,
—— foliatum, 275.
Caiman sclerops (2. 8. u.), 201.
Canis lupus (2. 8. 1.), 202,
Cannabateomys, 371, 379, 389, 417.
Capromyide, fam. n., 423.
Capromys, 369, 372, 875, 377, 387, 406
413, 417, 420, 423.
Captorhinas, 19.
Cardisoma armatum, 205, 361,
Cardium edule, 27.
Carterodon, 371, 879, 385.
’
Cavia, 369, 371, 875, 380, 695, 397, 403,
415, 418, 421, 425, 427,
d
XVill
Caviide, fam. n., 425.
Caviine, 426.
Cebus fatwellus (z. 8. u.), 201.
Cephalophus melanorheus, 399.
Ceratodus forstert (4.8. u.), 201.
Cerchneis tinnunculus satwratus (4. 8. L.),
481.
Cervus hortulorum (4.8. u.), 203,
Chetogaster, 109.
diaphanus, 109, 111.
limnei, 111.
—— opientaiis, 109, 111,
pellucidus, 109.
Chetomyine, 422.
Chinchilla, 370, 371, 875
417, 421, 425.
Chinchillide, 363, 425.
Chinchilline, 425,
Chirononus, 83.
992
weOe
115-118.
, 381, 392, 412,
Choniangium,
epistomunr, 229.
Ccelogenyidas, fam. n., 424.
Celogenys, 368, 369, 371, 375, 879, 384,
392, 403, 414, 421, 424.
Coendine, +22
Coendu, 369, 371, 375, 377
413, 416, 420, 422.
Conodectes favosus, 457.
Coracias caudatus (4. 8. L.), 201.
Ctenodactylide, 425.
Ctenodactylus, 371, 375, 382
419.
Ctenomys, 369, 371, 375, 379, 390, 401,
405, 417, 425
Ctenus flavidus, sp. n.,
Cuculus canorus, 197.
Cyclopodia greeffi, 471.
Cymbospondylus, 24
Cynonycteris stramined, 471.
, 089, 405,
, 403, 415
299.
Damatlis senegalensis, 8.
Dasyprocta, 369, 371, 380, 391,
408, 414, 419, 421, 424.
prymnolopha (4.8. u.), 201.
Dasyproctide, fam. n., 365, 424.
Dasypus villosus, 468.
Decrusia, 220.
—— additictr1, 220.
403,
INDEX.
Dendronephthya, 33.
annectens, sp. n.,
—— brevirama, 44, 70.
cervicornis, 44, 54.
clavata, 44.
collaris, 44, 61.
coronata, 44, 69.
disciformis, 44, 64.
- ehrenbergi, 44, 50.
flammea, sp. n., 44, 56.
—— gigantea, 44, 46.
habereri, 44, 66.
—— hyalina, 44, 76.
Klunzingeri, 44, 58.
longicaulis, 44, 63.
microspieulata, 44, 59.
mirabilis, 44, 55.
mollis, 44, 52.
orientalis, 44, 60.
—— pumilio, 44, 68.
—— punicea, 44, 49.
—-— simplex, sp. n.,
——- swensoni, 44, 51.
136, 137-189, 140, 145.
Diza shirleyi, sp. n., 200.
44, 72,
44, 78.
Desmogaster,
Dimetrodon, i9, 458.
Dinomyide, 365, +24.
Dinomus, 367, 371, 3877, 383, 424.
Diphyllodes hunsteini (4.8. 1.), 201.
Dolichotine, 426.
Dolichotis. 309, 371, 5
415, 419, 421, 425.
Donax vittatus, 30.
Drawida affinis, 128, 138, 141, 144.
-—- hbahamensis, 119.
124. *
—— ——— yar. imputusus, 128.
75, 881, 397, 403,
—— harwelli,
hrunnea, 128.
—— chulakundiana, 128.
decourcy?, 128.
Jakir, 128.
-— ghatensis, 128.
——- grandis, 123.
—— hodgarti, 128.
indicus, 128.
japonica, 119.
— kanarensis, 128.
—— kempi, 128.
INDEX.
Drawida nepalensis, 128.
papillifer, 128.
—— parambikulamana, 128.
pellucidus vay. stewarti, 128.
—— rangamatiana, 128.
robusta, 125, 128, 141.
rotunga, 128.
—— schunkarai, 141,
travancorensis, 128.
-—— willsi, 128, 138.
Dromia vulgaris, 194.
Dytiscus lapponicus, 86, 87, 02.
Echinocardium cordatum, 80.
flavescens, 31.
Echinocyamus pusillus, 30.
Eidolon helvum, 471.
Elephas maximus (2.8. u.), 479.
Elseya, 21.
Entomavtus fasciatus (4. 8. u.), 481.
EHquinurbia, 278.
stpunculiformis, 273.
Equus hartmanne, 330.
Evethizon, 369, 371, 375, 377, 388, 405,
413, 416, 421, 422.
—— dorsatus (4. s. u.), 480.
Erethizontide, 422.
Lrethizontine, 422.
Hupelnus, 471.
—— trichi, sp. n., 474.
Huphractus villosus, 99, 103.
Eupolygaster, 186, 137, 138, 140, 141.
Husthenopteron foordi, 459.
Hutypheus, 134.
Felis concolor (% .8. 1..), 20).
—— leo (4. 8. u.), 201, 202, 479.
—— lynx isabellinus (4. s. u.), 202.
Galea, 427.
Galepus, 19.
Gasteracantha pretextata, 290.
Geodipsas proctere, sp. n., 313.
Gorgonops, 19.
Granmocephalus, 274.
-—— clathratus, 205, 274.
varedatus, 274.
Grison allamandi (4%. 8. u.), 479.
Xx1x
Hatiaétus leucogaster (4. 8. u.), 201.
Haplochromis, 160.
—— acutirostris, sp. n., 162, 180,
—— altigenis, sp. n., 162, 175.
—— annectens, sp. n., 161, 168.
-—— argenteus, sp. n., 162, 186.
—— hayoni, 162, 176.
-—— cavifrons, 162, 183.
——- chilotes, 161, 170. _
—— cinereus, 161, 166.
—— crassilabris, 161, 167.
= ACItex sp. n., 162162:
—— dichrourus, sp. n., 162, 178.
—— flavipinnis, 161, 172.
—— gestri, 161, 164.
—— gracilicauda, sp. n., 161, 188.
—— gwiarti, 162, 174.
—— humilior, 161, 169.
— ishmaelt, 161, 169.
—— longirostris, 162, 187.
——— macrodon, sp. n., 162, 176.
—— Mmacrognathus, sp. n., 162,
182.
—— jmacrops, 161, 166.
—— maculipinna, 162, 177.
—— martini, 161, 171.
—— melanopus, sp. n., 161, 165.
——- mento, sp. n., 162, 183.
—— microdon, 162, 173.
—- nigrescens, 161, 172.
-—— nigricans, 161, 165.
—— nubilus, 161, 164.
-— nuchisquamulatus, 161, 1638.
—— obesus, 161, 170.
—— obliquidens, 162, 188.
—— orthostoma, sp. n., 162, 184.
—— pellegrini, sp. n., 162, 185.
—- plagiostoma, sp. n., 162, 181.
-—~ prognathus, 162, 177.
—— sawvager, 161, 167.
——-- serranoides, sp. n., 162, 180.
—— serranus, 162, 174.
—— spekit, 162, 179.
—— squamulatus,
Wey,
—— teniatus, sp. n., 161, 170.
—— wenodon, 162, 188.
—— xXenostoma, sp. n., 162, 185.
nom. n., 162,
xx INDEX.
Haplotaxis, 145.
Henicophaps albifrons (2. 8, u.), 201.
Hippotragus equinus, 318.
niger, 318.
variant, 318, 319.
Hoplotilapia, 190.
retrodens, 190.
Hydrobius fuscipes, 83, 94.
Hydrochoeride, fam. n., 425.
Hydrocherus, 371, 397, 408, 410, 412,
415, 419, 421, 425.
Hydromedusa, 22.
Hygropoda longitarsis, 304.
Hyrax capensis, 149.
Hystricide, 365, 522.
Hystricomorpha, 365.
Hystrix africe-custralis, 367, 369, 371,
375,
421.
Ichthyosaurus, 19, 23, 24.
Jaculus, 371.
Kerodon, 397, 400, 421, 427.
Labidosaurus, 18.
Lacerta, 458.
Lagidium, 370, 3874, 392, 421, 425.
Lagostomine, 425,
Lagostomus, 370, 3871, 875, 381, 394, |
412, 417, 421, 425.
Lamprodrilus satyriscus, 118, 147.
Langbiana, gen. n., 286.
klossi, sp. n., 286.
Lanistes affinis, 344.
olivieri, 344.
Leiperenia, 206.
—— galehi, 205, 210.
leipert, 206.
Lumobricilius, 134.
Lumbriculus variegatus, 147.
Lumbricus, 134.
-—— rubellus, 146.
Macoma baltica, 27.
Macropleurodus, gen. n., 189.
— bicolor, 189.
noE s |
376, 379, 388, 405, 413, 415,
Macropus giganteus (2.8. u.), 201.
—— ind (%.8.L.), 481.
Macrothele, 285.
maculata var. annamensis,
var. n, 285.
Macetra clliptica, 51.
—— subtruncata, 30.
Mandrillus leucopheus, 429.
Marisa cornu-arietis, 344.
Marmota marmota (2.8. .), 480.
Megaloprepia magnifica (2.8. u.), 201.
Memphisia, 25.
—— aziza, 261.
memphisia, 256.
Menemerus crassus, sp. n., 307.
— dabanis, sp. n., 307.
—— felix, sp. n., 305.
Mixosaurus, 24.
Monitigaster, 135, 187, 141, 144.
—-— deshayesi, 127,
—— indicus (= Drawida robusta), 125, °
138.
Murshidia, 220.
—— falcifera, 205, 222
—— hadia, 223, 228.
—— linstowi, sp. n., 205; 222; 228)
—— murshida, 221, 228.
Mya arenaria, 27.
Myocastor, 869, 372, 375, 377, 389, 405,
413, 420, 423.
228,
d
Natica alderi, 27, 30.
Naxioides serpulifera, 198.
Nematodrilus griseus, 125.
Nephila maculata, 299.
Nototherium mitchelli, 202.
Nucras kilose, sp. n., 314.
Ocnerodrilus, 134.
Octodon, 369, 371, 379, 385, 405, 407,
Ar), Gall7j, e240)
Octodontide, 423.
(Hsophagostomum tuberculatum, 468.
—- X€r?i, sp. n., 461.
Ophelia limacina, 30.
Ophioglypha albida, 27.
Oryx gazella, 336.
Ourebia rutilus, sp. n., 325.
INDEX.
Pachylabra sp., 344.
gordoni, 344.
Palystes ledleyi, sp. u., 296.
Pandercetes ochrea, sp. n., 2')2.
Pantylus, 18, 19.
Parabronema africana, 216.
indicum, 215.
—— smithii, 216.
Pardosa dranensis, sp. n., 303.
Parotia lawesi (%. 8. u.), 201.
Pectinaria koreni, 30.
Pelobius tardus, 79.
Perameles nasuta (z. 8. L.), 201.
Petalomera lateralis, 193.
Petaurus australis (4.8. 1.), 201.
sciureus (4.8. L.), 201.
Petromyide, 423.
Phascolomys mitchelli (a. 8.1.), 201.
Pheretima, 134.
Philhydrus maritinus, 94,
Placochelys, 22.
Platambus maculatus, 88.
Platyteniodus, 190.
—— degeni, 191.
Podargus strigoides (2. 8. U-), 201.
Pontoscolex corethrurus, 118.
Pristina, 109.
Procavia valida, 481.
Pteridopharynx, 250.
——- africana, 250.
anisa, 250.
Quilonia, 229.
—- africana, 235.
—— apiensis, 230.
__— brevicauda, sp. n., 238.
—— ethiopica, sp. n., 248.
—— renniet, 230.
tranvacra, 230.
— uganda, 239.
ji We 5
Ranatra linearis, 85.
Rangifer tarandus (4. 8. 1.); 480.
Redunca arundinum, 324.
Xx1
Sabella alveolata, 30.
Saimiris sciureus (u%.8.%.), 201.
Sarcophilus harrisé (2. 8.t.), 201.
Scops bakkamena lettia (z.8.u.), 481.
Scrobicularia plana, 27.
Seramba picta ?, 298.
Seymouria baylorensis, 457.
Spirurine, 215.
Sphenodon, 17, 21, 25.
—— punctatus, 495.
Spongodes, 33.
Storenomorpha, 285.
comottot, 285,
Stronglocentrotus drobachiensis, 30.
Sylvicapra grimmi leucoprosopus, 326.
Syngenodrilus, 135, 142, 144, 147.
Tachyskarthmos, gen. n., 310.
annamensis, sp. n., 310.
Tamandua tetradactyla, 99, 149.
Tatusia novemcincta, 99, 107.
Tellina fabula, 30.
Terrapene, 21.
Testudo calecarata (2.8. t.), 202.
Thalattosaurus, 24.
Thrynomyide, nom. n., 423.
Thrynonys, 401, 415, 420, 425.
swinderianus, 367, 3871, 380, 401,
433.
Tilapia, 160.
variabilis, 160.
—— zillii, 160.
Trichomys (Nelomys), 371, 417.
Trichys, 871, 421.
Trionyx, 21.
Turritella terebra, 31.
Venus gallina, 80.
NXerus setosus, 461.
Youngina, 18, 24-26.
Zootoca, 495.
PRINTED BY TAYLOR AND FRANCIS,
RED LION COURT, FLEET STREET.
uy vo oe
2 OMS RG
i
VIN CATON RARE (aR
yy
Bey
PROCEEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF THK
ZOOLOGICAL SOCLETY
OF LONDON.
1922.
PART I.
conTAINING Paces 1 ro 208, wita 11 Puarres anp
APRIL 1922.
PRINTED FOR THs SOCIETY, —
SOLD AT ITS HOUSE IN REGENT’S PARK.
LONDON :
MESSRS. LONGMANS, GREEN, AND CO,,
79 ‘Vex-FIGURES. Z
PATERNOSTER ROW, *
y& : [Price Twelve Shillings.)
WEN
eZ
LIST OF S003 TEA Es.
1922, Parr L (pp. 1-203).
EXHIBITIONS AND NOTICES.
* Page
The Secrerary. Report on Additions to the Society's Menagerie during the months of
November and December, 1921
Mr. A. H. Evans, F.Z.8. Exhibition of Cuckoos’ eggs taken near Cambridge
Lord Currrorp or Cuvupinien, F.Z.8. Exhibition of photographs of Nototherium mitchelli.
Miss L. E. Creusmay, F.E.S. On the position and function of the Siphon in the
Awphibious Molluse, Ampudlaria vermiforits 12. or 00 connec eee ewes es ae ete
Mr. G. C. Rozsoy, F'.Z.S. Exhibition of Models demonstrating the respiratory mechanism
Ole Ay CHEE OORMOLOP UO MGA AQHOCE SAG down; rabdnodaconmansoebsofhoongoe ls
The Secrurary. Report on Additions to the Socicty’s Menagerie during the month of
UETTIE NS al MS Pee eA ME Mmmm ogqocaden och duadiouabeuesdon wo dca o
The Secretary. Exhibition of a photograph of the Society’s Gardens taken from an
BELOPlANe ceigeice es cess regs
Miss L. E. Cunrsman, F.E.S. Exhibition of living specimens of the Amphibious Molluse,
Ampullaria vernifornis
ee oeeer ee oe ee ee ee ee ee ee ese ese teow BF ee seeece ts eeeegsee
Mr, G. C. Rosson, F.Z.8. On the respiratory mechanism of the Ampullariide
Mr. F. Martin Duncan, F.R.MS., F.Z.S. Exhibition of Cinematograph Films taken in
the:Soctetyis Gamdlens 2c. c's) 5 «ove oct Rem RRase Melee tey ener sta leae | eer eie tame ene gegen
Miss L. E. Cnzssmay, F.E.S. On the habits, in captivity, of the Freshwater Crab,
CHMSCLE CHUMUINTD ‘sd ocabacdneaddoocn6 herp cteraiet allen eerereaeieus NCR ER anisoe on
Mr. C. Tarr Recay, F.R.S. Exhibition of living specimens of the Indian Tortoise-Beetle,
Aspidomorpha sancte-crucis
208
203
203
Oontents continued on page 3 of Wrapper.
PROCEEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OV THE
ZOOLOGICAL SOCIETY OF LONDON.
PAPERS.
1. The Fauna of Hast Africa and its Future.
By C. W. Hosiery, C.M.G., C.M.Z.S.
[Received October 17, 1921: Read February 7, 1922.
I have been asked by the Secretary to give some of my
impressions of British Hast Africa, which is now known as
Kenya Colony.
fam naturally diffident in addressing a learned society like
this, and my only claim to attention is that there are probably
few present this evening who have had such a long experience of
the country of which I propose to speak. My connection with
East Africa began in 1890, and since that time I have served
continuously in the Uganda Protectorate and British East Africa,
and have during the whole of that period been a keen observer
of the fauna of the area, so have naturally chosen that subject for
my remarks this evening.
The country has been so often described in books of travel and
sportsmen’s records of slaughter, that any general description is
unnecessary. I therefore propose to confine myself to a few
remarks on the fauna based on personal observation.
I was fortunate enough to be able to travel in the country
when it was in what may be termed the exploration stage; that
is to say, before the advent of any European settlement and the
consequent disturbance of natural conditions. Needless to say,
game was far more numerous over the whole country, but
particularly in the favoured areas (7. ¢., the great plains) in those
days than it is to-day; and on looking back and trying to compare
Proc. Zoou. Soc.— 1922, No. I. 1
M MR. C. W. HOBLEY ON THE
the stock of game in the country as a whole to-day with that in
say 1895, probably the present stock would only represent 15 per
cent. to 20 per cent. of the former: this is, however, only to be
expected when one considers that upwards of 6000 square miles
of land, most of it game country, has been alienated to colonists.
I am not complaining, for it cannot be expected that fertile
lands which will provide homes for people of our race and grow
products essential to civilization, will remain for ever in the
ossession of wild game.
T will, however, revert to the question of the future of the game
later on, and now propose to recall the conditions which prevailed
20 years ago before man took a hand in the matter, 7. e. to any
great extent.
At that time the areas noted for great profusion of game were
the Athi and Kapiti Plains, the Yatta Plateau, the Serengeti
Plains, the Loita Plains, the Rift Valley, and the Uasingishu
Plateau, A portion of the Kapiti Plains and the Loita Plains
are included in the game reserves, so presumably carry a good
stock in places, but all the other areas are decimated, mostly
owing to the effect of settlement.
The main factors which determined the distribution of game
in the early days were yearly variations in rainfall which resulted
in a sufficiency or insufficiency of grazing in particular areas ;
epidemic diseases also periodically affected certain species, and
the number of carnivores also counted. All these factors operate
at present, but the first mentioned, viz., the variation in rainfall,
operates more harshly than formerly, for nowadays the area over
which the game can migrate in search of grazing is restricted.
Tf, for instance, the Southern Game Reserve is seriously affected
by drought, large numbers of game are doomed; some may
attempt to migrate into the farm lands, but many are shot down,
and the survivors retreat to the reserve where the grazing is
finished and many of the water-holes dry. A good example of
this occurred in 1910, when the plains were so dry that zebra and
hartebeest came up in force into the town of Nairobi, regardless
of man, in their search for water; the lions followed them and
killed game nightly in the open land in the centre of the town.
Natives and others killed many of the invaders, and the emaciated
remnant was driven back to the plains.
Speaking of epidemics, one of my earliest recollections in East
Africa was the great rinderpest visitation of 1891. JI was then
exploring the course of the Tana River, and the buffalo were
coming down to that river literally in thousands to die. The
bush country fringing the Tana between Hameye and Mumoni is
not ideal buffalo country, for the grazing is not too plentiful at
any time, but once they contracted the disease they appeared to
be impelled to seek water, doubtless coming from great distances
to the river, and I estimate that im the stretch of country above
mentioned, a distance of about 80 miles, we saw several thousand
buffalo in all stages of disease and death, attended by vultures and
FAUNA OF EAST AFRICA. 3
marabou storks in myriads, all gorged to repletion. Occasionally
dead giraffe, waterbuck, and bush buck were seen; the eland also
suffered, and [ am told that kudu and roan were also attacked.
From the buffalo the disease spread to the cattle, and at
Ngomeni in North-East Kitui, which was a great cattle centre
betore the epidemic, we were shown some 20 odd beasts, the
sole survivors of many thousand head, and the desiccated carcases
of the victims were piled up as a wall outside the villages.
It was during that journey that we happened to follow au day
behind a large. impt of Laikipia Masai which was engaged in
raiding the Embu and Mbe tribes. I will not dwell on the
evidences of ruthless slaughter we witnessed, but some fifteen year's
later I was recalling the incident to the chief of the Laikipia
Masai, and he laconically told me that he was a “moran” or
warrior at the time and had participated in the raid; he paren-
thetically added that it was an unfortunate venture, for some of
the captured cattle were infected with rinderpest, and so they
took the disease back to their own herds on Laikipia and practi-
-eally all perished. Nemesis indeed !
This outbreak was the worst epidemic known in the recent
history of Africa; it spread rapidly south through what was then
German East Africa, crossed the Zambezi, reached Buluwayo
about 1895, and by the end of 1896 it had reached the Cape.
During the last year of the visitation its progress was remarkably
rapid, viz., about 1000 miles ; it was probably spread to a great
extent by the transport riders. Since that devastating attack we
have had minor epidemics of the disease; in 1904 I saw eland
«lying of it near Naivasha, and the Masai then lost over 600 head
of stock. Although sporadic outbreaks still occur, this disease is
now well in hand.
In very dry years, when grazing is scarce and the plains are
very dusty, outbreaks of anthrax occur among the game, the
principal species to suffer being Coke’s hartebeest; the last
‘serious outbreak which was identified was in 1905, when several
thousand head of game died on the Athi Plains.
Pleuro-pneumonia 1s rarely absent from the herds of Masai
cattle grazing in the South Reserve; but, curiously enough, there
is no record of the disease ¢ attacking the game, and the Game
Warden has stated that cases are inewn where eland and buffalo
graze over the same land as cattle infected with this disease and
‘are apparently unaffected.
About 1906 an epidemic of what is believed to be distemper
broke out among the jackals on the Athi Plains, and large numbers
died; a year or ‘two later the same disease was recorded from the
Rift Valley, and then from Laikipia. It is not, however, known
whether the disease is endemic, or whether it has been introduced
‘by civilization.
Wild game is in some areas infected to a considerable extent
with intestinal parasites. I have frequently observed the intra-
muscular cysts of what is commonly called ‘“‘*measles” in mpala,
4 MR. C. W. HOBLEY ON THE
Grant’s gazelle, and hartebeest. The health of the host appears.
to be unimpaired.
Taking it all round disease is rare among game and serious ep1-
demics are uncommon ; as Mr. Percival has pointed out, the reason
is that directly an animal becomes sick it either leaves the herd
or is driven out, and being alone and dull with sickness it more
readily falls a prey to one of the killers or one of the scavengers ;
next day the vultures complete the story, and very little is left to.
form a breeding ground for the flies which are the great dis-
seminators of disease.
It would take far too long to attempt to deal with each species.
of the various game animals in the part of Africa with which I
am acquainted, and so much has already been written by others.
I will, however, venture to refer to one or two of particular
interest.
Everyanr.—First comes the elephant. The numbers of
elephant have greatly decreased during the last thirty years, for
apart from the European sportsmen from overseas, the residents.
in the country have accounted for a large number; further, the
improvement in communications has indirectly led to a greater
amount of illicit slaughter. It is true that game laws have been
in force in British Kast Africa and Uganda for about twenty years.
and administered by a keen staff, and the ports have been care-
fully watched; but a great deal of ivory has nevertheless been
smuggled out over our and frontiers both to the north and south,
and without heavy expenditure this was impossible to check.
There are certain small tribes like the Dorobo and Sania who are:
professional hunters, and many of the Hastern Akamba are great
offenders. A native with a few tusks has up to now had no great
difficulty in disposing of them to some itinerant Arab, Baluchi,
or Somali trader who was wandering about ostensibly trading in
hides, and Indian traders could always be found to finance such:
undertakings.
The elephant endangers itself by conservatism, for it is a
beast of very regular habits ; ; at certain seasons 1t invariably
seeks certain loznltsies. and it is interesting to map out the lines
of migration.
Generally speaking, in the hot dry season it seeks the shelter
of the high forests, and in the rains treks to more open countr Wee
native hunters say that the perpetual drip from the trees in thie
forest is distasteful to it. Hlephants in the Kilimanjaro region,
for instance, move down the northern flanks of the leanne
Range about April and spread out through the Nyika Plains
almost to the coast, one of the attractions shan being the ripe
fruit of great groves of Hyphene palms.
Similarly, another herd migrates each year from the same
region and marches east, crossing the Uganda Railway near
Kinani and on eastward across the Athi into Southern Kitui.
For many years past, about June, a herd of elephants has come:
FAUNA OF EAST AFRICA, 5
down to the sea north of Kilifi and has spent a month or so
wandering about and feeding in the thorn bush near Malindi.
The herd may be the one referred to from Kilimanjaro, or it may
come from the Middle ‘Tana or even from the Lorian.
very year a herd of elephants was wont to descend from the
Aberdare range to the Rift Valley between Naivasha and Longonot
Mt. ;. this herd has, I think, been obliterated. Other “Tags
of migration were across South Laikipia from the Aberdares
to Kenya and then north-east towards Lorian*. Still other herds
migrated from the Aberdaves northwards, passing east of Baringo
towards Mount Nyiro.
Elephants from the Mau forests also yearly come down to the
high grass country in the Lower Nyando Valley, and others
annually frequented a swampy valley at the foot of Gwasi Mount.
The main routes of these migrants were so well marked, and
they followed practically the same roads year after year and
about the same month, that a native hunter’s task was easy.
These elephant roads are in certain places striking features; on
the east side of Ol Bolossat Lake they can be seen climbing
«hagonally up the flank of the escarpment, and form mnitoetaly
graded roads about 8 feet wide. On the summit of Mau there
ave other roads originally elephant tracks but later on widened
out and deepened by mobs of Masai cattle, and about which some
imaginative people have tried to force the opinion that they are
the relics of an ancient highway from Egypt to Zimbabwe.
Hearing that considerable stocks of old ivory were in possession
of the natives, some years ago the local government instituted the
practice of buying in ivory at half the market value; this resulted
in the production of a large amount of old ivory but also of a
large number of tusks mostly derived from immature beasts.
Recently most of the ivory brought in has been obviously obtained
from animals Killed fairly recently, but often camouflaged to
resemble old ivory.
It is very noticeable, however, that during the last ten years, in
consignments of confiscated or purchased ivory, the average weight
per tusk has decreased, and old big tuskers are now, I believe, very
scarce. The future of the elephant may therefore be said to be
precarious—he will not stay in any reserve; the inherited instinct
with which he makes his annual migrations will, I fear, cause his
disappearance, although there may for many years to come be
vast areas wherein he could live in reasonable safety.
Lron.—A few words regarding the lion. This beast is in no
danger of extinction. As is well known, cats are very prolific, and
* Fyrom the Lorian swamp the migrant herds pass N.W. to Marsabit and then
southwards back to the Aberdares, a round trip of well-nigh 400 miles. ‘The
well-known hunter naturalist Mr. R. J. Cuninghame informs me that this journey
from the Aberdares back to the Aberdares is believed to take three years, for herds
have been identified leaving the forests on the above mountains without calves and
returning to the same place three years later with calves of about a year old. The
tavourite breeding place is said to be Marsabit forest.
6 MR. C. W. HOBLEY ON THE
as long as the reserves are maintained and contain any game, so
long will there be lions ; in fact, a reasonable supply of lions is, I
consider, a sine qua non in a reserve, for they clear off the weak-
lings and the sick, forming part of nature’s scheme in a natural
assemblage of animals mainly composed of herbivores. Curiously
enough, however, during the war they increased out of proportion,
for although an enormous amount of game was shot by the
troops, the lions were not hunted to any extent.
The result of this was that game being scarce in the area of
military operations, numbers of lions became cattle eaters, and
the Masai herds which tenant the Southern Game Reserve have
suffered heavily ; I believe that the Game Department has had to
take steps to reduce the number of lions there during the last
year or two. It is not easy to understand what normally limits.
the number of lions; one never sees a lion which has died of
starvation or disease, and lions which are shot always appear to.
be in a fair condition, but some cause undoubtedly does operate
in a natural ass semblage to preserve the proportion of carnivores
to herbivores.
Considerable troops of lions are sometimes seen, but it 1s rare.
Sir Frederick Jackson recorded a troop of 23 near Lukenya; the
Leases troop I have seen myself was 11, made up of 1 male,
4 females, and 6 half-grown cubs, near Neong, where I watched
the adults opened out in echelon on a hillside, hunting a herd of
zebra, the cubs being assembled under a tree watching the
performance, and doubtless learning the art.
Lions have now been well-nigh abolished in the older settled
areas and necessarily so, for no one ean successfully farm stock
in a country where lions are common. ‘There are still, however,
but few places where domestic stock could safely be allowed to
stay out in a pasture all night.
Ruinoceros.—This curious beast has suffered severely of recent
years, and great numbers have been killed. The greater pro-
portion inhabited the open plains and were thus conspicuous to
sportsmen. They have irritable natures and have a stupid habit
of charging down on anything that annoys their sense of smell.
Tn hundreds of cases this has induced their death, for many aman
has had to shoot a rhino in self-defence. I have myself had
several narrow shaves owing to thei sudden attacks.
Many were killed during the war in the southern portion of
the Game Reserve by the troops of both forces.
The high price which rhino horns have commanded in the Far
East during the last few years has, moreover, proved an induce-
ment to the native hunters to slaughter these animals; they
were instigated by the Indian traders, who surreptitiously shipped
them out in considerable quantities.
This beast does not make long migratory journeys, and, pro-
viding that the reserves are of ample size and contain an adequate
FAUNA OF BAST AFRICA, 7
supply of the acacia scrub which forms its diet, and water, there
is no reason to fear its extinction.
Burrato.—After the big rinderpest epidemic it was feared that
these fine beasts were practically extinct, and for some 20 years
or so they were undoubtedly rare.
They have, however, gradually increased in certain areas, and no
anxiety need now be exercised as to their extinction. There are,
it is believed, considerable numbers in the Southern Reserve, on
the northern stretches of Laikipia, in Southern Kitui, and various
other places.
A considerable number also frequent the coastal area where the
bush is very thick, and upto recently a few even annually visited
the mainland opposite Mombasa town, but left the spot when the
bush was cleared to make a plantation. In the spring of 1919
about half a dozen visited the place they knew, but finding no
cover they retreated into the interior; one, however, swam across
the mouths of the two harbours and landed on the mainland
south of the island, a distance of over half a mile.
In the old days buffalo could be seen in the early part of the
day and in the late afternoon grazing out in the open like herds
of cattle, but since the epidemic this is very rarely the case.
They now live in dense bush, only coming to graze in the open at
night, retiring to the bush again at dawn. Possibly now there
are fewer of them they have lost the confidence they formerly
possessed.
The geographical range of the buffalo is remarkable, for, as I
have remarked, they frequent the hot coast-lands, and then, again,
a desiccated buffalo carcase is recorded from the foot of a glacier
on Mount Kenya. As might be expected, the animals inhabiting
the higher altitudes have thicker coats than those at the coast.
Hrerororamus.—This beast is decreasing at a great rate; in
the old days the middle and upper Tana swarmed with them, but
now not more than 5 per cent. survive. The same applies to the
Athi River ; they have also greatly decreased in Lake Victoria
and in other smaller lakes.
Unfortunately the Game Reserves do not contain much water
suitable for them, but it will bea long time before they disappear
from the swampy bays on Lake Victoria, and from some of the
more remote rivers, such as the lower course of the Nyiro which
flows into the Lorian Swamp.
GrraFFre.—The area occupied by this fine beast 1s becoming
more restricted every year, and the settlement of the Uasingishu
plateau will probably settle the fate of the beautiful subspecies
which is found there, for the Dutch colonists find it hard to refrain
from slaughtering them, as giraffe hide has a peculiar quality
8 MR. C. W. HOBLEY ON THE
which makes it valuable for reims. There are, however, a good
number of the common species in the Southern Reserve, and one
can frequently see a herd from the mail train on the Uganda
Railway near Sultan Hamud Station.
WILDEBEEST.—In former times there were vast numbers of
these beasts to be seen, and in the spring of 1896 I saw an
enormous herd, which I estimated at nearly 10,000 head, migrating
northwards along the Athi Valley on the east side of where Athi
River Station now stands. The species is, however, in no danger
of extinction, for there are still a considerable number in the
Southern Reserve and on the Loita Plains.
I will now add a few words regarding the vagaries of
geographical distribution of some of our game species, and the
difficulty of ascertaining the laws that govern it.
T will give the following examples :—
1. Sastre AntELoPE.—This species is found from the border of
Tanganyika Territory near the coast and inland for about 20
miles, then very sparsely northwards to the Sabaki River, where
it ceases. In fact, it is extremely rare north of the Uganda
Railway. It is not found anywhere else in Kenya Colony or
Uganda.
2. Witpesesst.—In the plains near Kilimanjaro through the
Southern Reserve up to the neighbourhood of Donyo Sabuk this
abounds ; it stops abruptly at the Thika River. Farther west it
is found toa small extent in the southern portion of the Rift
Valley, but never extended northwards along that valley farther
than the Lower Kedong Valley, although the plains immediately
to the north abounded in grass and water. West of the railway
the wildebeest favours the Loita Plains; in that area, however, it
was shut off from extension to the north by the Mau Forest.
3. Roan.—the distribution of this antelope is very capricious.
It is said to be sporadically found near the coast, but is very rare,
and I have never seen it there. It is found on the big range of
hills west of Sultan Hamud, and on the foot-hills of the Ukamba
Range to the east of that place. Then there is apparently a gap,
for if have not heard of its oceurrence again until we come to the
Nyando Valley about 30 miles east of Kisumu. Sir Frederick
Jackson also mentions its occurrence near the Turkwell River.
It is, however, nowhere a common beast. Odd specimens may
yet turn up in unexpected places. a
4, Tort. Damaliscus senegalensis.—The distribution of this
antelope is worthy of attention. On the coast it is fairly common
between Malindi and Lamu, and some say that it is found farther
south near Vanga. Jackson states that it does not occur south of
the Sabaki, but I saw a few N.W. of Rabai. Proceeding inland
FAUNA OF EAST AFRICA. 9
there is an enormous gap, for we do not find it again until we
come to the Uasingishu Plateau, which is about 6000 ft. above
sea-level. This is an unexplicable hiatus. I have not, however,
had an opportunity of comparing Uasingishu skins with speci-
mens from the coast, so cannot testify to their absolute identity.*
5, Wauier’s GazeLie.—This species is another example of
intermittent distribution. It is found on the steppes on the east
or left bank of the Tana River below Hameye and extends
northwards to Somaliland. Then there is a great gap, for no
trace of it is seen until we reach the neighbourhood of Lake
Magadi, a distance of 200 miles; a few are also found on the
Serengeti Plains, principally to the north of the Voi—Taveta
Railway line. It is also believed to exist in Usambara district in
Tanganyika Territory, but this is its southerly limit.
6. Harreseest.—The common hartebeest of the country is
Coke’s, and, as a rule, whenever a square mile or two of open
country occurs we get this ubiquitous beast. It is locally called
the ‘‘ Kongoni.” A curious thing happens in the Rift Valley; at
the south end of the Rift Valley in Kenya Colony all the harte-
beest are cokei, but about the north end of Naivasha Lake
nearly every hartebeest seen belongs to the Neumann's variety,
and as we go farther north and west we find this gives way to the
curious Jackson’s variety, and on the Uasingishu Plateau there
is no species of the hartebeest other than Jackson’s. ‘This, of
course, is not so much a question of the distribution of one species
as a change of one species, and the variations of the Grant's
Gazelle and the Oryx in different areas could similarly be traced.
These facts of erratic distribution are difficult to understand,
and to find an adequate explanation J am inclined to believe that
we shall have to go back to the time when the progressive desic-
cation of this portion of Africa set in.
This question of desiccation cannot be discussed here at any
length. I have collected a considerable amount of evidence re-
garding it which was published in the ‘ Geographical Journal,’
November 1914. The causes are by no means clear, but’ certain
meteorologists have advanced the opinion that during the glacial
period of Europe there was a much greater rainfall in the areas
to which the glaciers did not extend. At any rate, there is clear
evidence that in Pleistocene times or thereabouts the glaciers of
Kenya extended some 5000 ft. below their present terininations, .
viz., to about 9000 ft. above sea-level, and on Ruwenzori glacial
deposits are, it is stated, to be found at the surprisingly low alti-
tude of 5000 ft. above sea-level.
Now the greater rainfall during that period, which only de-
creased gradually through a long term of years, must have resulted
in a marvellous expansion of the forest area; the high-level flora
* The topi is also common on the Loita Plains and southwards on the same
meridian ot Longitude into Tanganyika Territory.
10 MR. C. W. HOBLEY ON THE
would extend over a much larger area, and the low-level forest.
assemblages would extend over thousands of square miles now
only tenanted by dwarfed trees of species which can survive the
long droughts. Co nsequently the areas of open grass lands so-
necessary to many species would be restricted.
All this must have had a profound effect on the wild game of
those times and on its distribution. The low forest, for instarice,
now only survives in a few favoured spots such as the delta of the
Lumi River near Taveta, the banks of the Tana, and on a few
isolated avens near the coast, but traces of its existence are to be
found over a great extent of country, and ,these remnants are
evidence of its greater extension.
Piecing together the few facts I have quoted regarding distri-
bution, T premise that Kenya Colony may be looked upon as the
junction or the termination of three zoological areas :— The West.
Coast area, the Somaliland area, and South-East African area.
The West Coast area ofticiaily terminates at Ruwenzori Mountain,
but traces of its fauna are still found far east of that mountain,
in country which they invaded during the period of maximum
afforestation before referred to, a variety of the West African
potto ( Perodicticus) being found as far east as Kakumega Forest,
which is 50 miles east of Lake Victoria. ‘The bongo is also a
West African type, the butterflies of Kavirondo and West Nandi
also contain many West African types, and the grey parrot of
W. Africa comes as far as the Nandi Forest, and is very occasion-
ally seen as far east as Kikuyu.
As regards the Somaliland fauna, I look upon Waller’s gazelle
as the type-species illustrating the invasion from the north, it is
quite common on the north and east side of the Tana, but much
rarer at its southern limit; Grévy’s zebra is another example, and
possibly the lesser kudu. The greater kudu is a puzzle because
it is fairly common in parts of Some aliland, and common in parts.
of South Africa, but very rare in Kenya.
With regard to the South-East Afri ica area, the sable antelope
undoubtedly worked its way up the East coast from Portuguese
East Africa, and I am inelined to believe that the wildebeest
also came north from South Africa, developing new characteristics
on the journey.
The original focus of distribution of the bubalines, which in-
clude the various forms of hartebeest, is very difficult to settle,
but they possibly originated and split up into varieties in this.
avea, for more hartebeest probably exist in it than in any other,
and the forms are more numerous.
GAME AND Disnase.— Although, of course, there are many keen
sportsmen in the Colony, there is a strong local feeling against
wild game among a section of farmers, who believe that it
spreads the tsetse- flies and thus increases the area affected by
trypanasomiasis. They again argue that game fosters the supply
FAUNA OF EAST AFRICA. 1}
of ticks and forms a reservoir for Hast Coast fever, ete. They
thus urge the extermination of all the game.
The tsetse-fly has probably restricted the settlement area, but
up to the present there has been no dearth of land free from this:
insect. To what extent the tsetse depends for its perpetuation
on wild game is not quite clear, J, personally, saw more tsetses.
than anywhere else in « bush area on the coast where hardly any
game existed; then, again, there are tsetse-haunted areas on
the Kerio River where the Turkana tribe herd large numbers.
of donkeys and camels with impunity. The bionomics of this
insect need still more investigation, and it does not necessarily
follow that all the representatives of the genus Glossina carry
the disease.
The tick question in Africa is really one of more importance,
and why some areas literally swarm with these Arachnids, while
others are but sparsely supplied, is a mystery. I do not know
if it is anything move than a coincidence, but I have noticed
that the areas covered with black cotton soil contain far more
ticks than the sandy soils or red soil areas. The black soil cracks
a good deal in the dry season, and it is possible that these
cracks may form shelters for some of the ticks when the grass.
is fired, whereas in other areas the greater portion of the ticks
are annually destroyed by fire.
The Athi and Kapiti Plains swarm with ticks. and the soil
there is a black clay formed by the weathering of the underlying
sheet of phonolite, and which opens out into deep cracks during
drought. The Rift Valley soil is a light powdery volcanic dust,
and ticks are by no means plentiful. In the light soil of the
coastal region ticks are also not very numerous.
There is no doubt that wild game is tolerant of the parasites.
which cause diseases such as Hast Coast fever and Redwater
fever in cattle, and there is little doubt that the ticks become
infected by biting these hosts ; further, it is alleged that a tick can
engorge itself once in each of its three stages of existence, also
that in most cases when a tick engorges on an animal it cleans
itself-—that is to say, its internal pavasites, if any, are discharged
into the mammal upon which it feeds. Presumably if this
animal is already infected the tick absorbs a new crop of parasites
with its blood.
It is also alleged that it has been proved that in some cases
ticks hand on their parasites to their progeny. It will therefore
ke seen that the problem is fairly complex.
The veterinary authorities in East Africa and South Africa
have done a good deal of work on the subject, but it appears to
me that further investigation is necessary before we can be
dogmatic on these dificult questions.
It must, moreover, also be borne in mind that if the big game
is exterminated in an area, there will always remain a supply
of ground game also presumably tolerant, which may keep the
disease parasites alive.
2 MR. C. W. HOBLEY ON THE
The abolition of the tick is, I fear, impossible; on a fenced
farm frequent dipping of the cattle apparently does greatly
reduce the population of ticks within the fenced area, but of
course dipping 1s impossible with wild game.
At any rate, the wild game is incriminated as regards the
spread of disease to domestic animals, and there is strong evi-
dence as to its guilt, particularly as regards Hast Coast fever and
Redwater disease of cattle; canine tick fever is another case in
point.
Game Reserves.—The Game Reserves in Kenya are not
tenanted only by the wild game, for they are also occupied by
certain native tribes.
The Southern Reserve contains the Masai tribe with its huge
flocks and herds; the Northern Reserve contains a sparse popu-
lation of Suk and Samburu people.
The Masai do not hunt or kill wild game to any great extent ;
they probably kill a few buffalo in order to get hides for their
shields, and they also kill a few hons. In this way they are not
detrimental to the game; their young men are useful, too, as game
scouts. During normal years matters adjust themselves, but in
periods of drought, when grazing is scarce and water scarcer still,
the game has to take second place to the Masai stock, and there
is little doubt that the game suffers.
It is not usually so much shortage of grass as shortage of
water, for wild game can graze farther from water than cattle,
and the question of increasing the water storage of the reserves
should be carefully gone into, and works carried out. The Masai
would probably contribute to the cost, as the area has been dedi-
cated to their use, and they are looked upon as having the first
claim on it; presumably if they decided to kill all the game there
as being detrimental to their grazing rights, it might “be argued
that they were legally entitled to do SO 5 ‘this is, however, an
academic point at present.
The Northern Reserve is mostly arid desert, and it receives
little attention from any one; it is rarely patrolled by the Game
Department, for a large portion of it is not too safe at present.
A few favoured spots such as Marsabit and Mt. Nyiro are the
haunt of elephants, buffalo, and probably kudu; Grevy’s zebra
is also found in this part of the Colony.
GENERAL.—The wealth of wild game with which nature has
endowed Hast Africa should, I think, be viewed as a national trust
to a greater extent than it is at present. In Kenya itself efforts
as regards its preservation are looked upon as being reactionary,
and the Game Department is too apt to Become k a cockshy for
the local politicians, and only tolerated because it brings in a
little more revenue than is spent upon it.
Before the war big game sportsmen spent many thousands
annually in the country, and afforded employment to a con-
FAUNA OF EAST AFRICA, 13
siderable number of Europeans and natives; further, the income:
from licenses averaged over £10,000 per annum.
It is, however, parochial to assess the value of the game on
these lines, and I feel sure that our American friends do not
measure the value of their fine reserves in terms of dollars.
In my view the wonderful fauna of our Hast African posses-
sions does not solely belong to the people who happen to settle
there, and who somewhat reluctantly vote a minimum amount
per annum for its preservation—it belongs to the Empire—and
should not be entirely at the mercy of local interests.
Of course, as I said betore, settlement must go on and full con-.
sideration must be given to the economic interests of the colonists,
and sympathetic attention to complaints regarding the depre-
dations of game, as well as the question of game and cattle
diseases. Generally the relations between the Game Department
and the colonists require great tact and judgment.
I believe that a proposal to reduce the game reserves materially
would have to obtain the Secretary of State’s approval, but pre-
sumably he would follow the recommendations made by the
Governor at the time, and that officer’s views might be:
coloured by the limited view of his Legislative Council, and
therein is the danger.
1 should like to see a Committee selected from the Council
of this Society and the Council of the Society for the preservation
of the fauna of the Empire duly recognised by the Home Govern-
ment as the official advisers on the question of game preservation
in East Africa, and on the policy of the Game Departments in the:
various Crown Colonies and the mandatory territory of
Tanganyika.
If this could be brought about I feel sure that a more scientific
treatment of the question would ensue.
IT am loth to introduce any personal note, but will venture to.
record that the present Game Warden of Kenya is devoted to the
game in his care, and has an unrivalled field naturalist’s know-
ledge of the fauna of the country; he, moreover, does ali that
is possible with the limited staff at his disposal. He would
probably be extremely thankful to receive support from a body
of great weight outside the country; such support could not.
fail to stre nsthen his hands, and to give him greater confidence
in his efforts.
J, however, strongly desire to urge that the whole outlook should
be reviewed on a scientific basis, and that the bionomics of the
fauna be given greater prominence. Up to now what may be
termed a hand to mouth policy has been the vogue. It is true:
that the various Governors have been more or less interested in
sport, and have generally supported game preservation, but no:
one knows whether a Governor might appear whose policy would
render nugatory any systematic game preservaticn.
Much, moreover, requires to be done if the Reserves are to be
14 MR. C. W. HOBLEY ON THE
made permanent sanctuaries for the big game. An improvement
of the water-supply will, for instance, do a great deal to anchor
the game to their Reserves.
The fencing of the portion of the Game Reserves which abuts
on the settled areas will eventually have to be undertaken, and
this is a serious matter, for it will cost a considerable sum to
erect, and annual sums for maintenance.
The relation of the tsetse-fly and ticks to game needs further
investigation by trained observers, and it Is, I think, important
that a biological station should be founded in the main Reserve
with a research zoologist in charge, and this should be the centre
for detailed investig gation into the life- -history of the game
animals, the problems of distribution, the diseases of game, and
the bionomics of insect life, with particular attention to the
tsetses and ticks. Attention should also be given to locust ex-
termination, for the destruction of large areas of grazing by these
insects profoundly affects the game, and when the locusts spread
to the farms, they lead to losses and consequent grievances. For
a, similar reason attention should be directed to the destruction
of the myriads of larve of a butterfly, of the genus Belenois,
which periodically destroys large areas of grazing in the Reserves
and elsewhere.
The Game Department should officially photograph the game
in their natural haunts, both by telephoto apparatus and film
cameras ; the rangers have opportunities of seeing game under
conditions denied to most. A series of photographs ‘would form
a very valuable record, and also prove of service for educational
purposes. The Department should also collect from time to time
series of heads of the various species for our national collections,
and a complete reference collection for the local museum. The
lesser fauna and the avifauna should also be collected for refer-
ence and study-——the field is very wide.
With proper arrangements and funds the Department could
also supply the Society with live specimens to fill its gaps. ‘The
Department at present is so limited in personnel that it almost
entirely confines its energies to the game-keeping side, 7. ¢., the
prevention of poaching, doubtless an important duty ; but more
stress requires to be laid on observation, and the collection of
accurate scientific data regarding the fauna.
Im the past, vacancies in the Game Department have been
filled in somewhat haphazard fashion from local applicants. ‘This
practice is, | believe, a mistake, and it is impossible to obtain
scientific data if it is continued. Here, again, this Society might
ask to be allowed to act as a selection board to see that only men
with the necessary zoological and biological qualifications were
appointed to the Game Departments in this group of territories.
Tt will, think, be allowed that the whole question of the future
of the game in East Central Africa is one of great interest,
FAUNA OF EAS! AFRICA. 19
whether viewed from the point of view of the zoologist and
naturalist, or from that of a great national asset.
‘The problem is one of some complexity. On one side we have
the interests of an increasing number of settlers who have in
many cases invested their all in their farms, and who cannot be
ignored ; for as time goes on they are steadily acquiring the right
to decide the future fate of the country in which they have
settled. On the other hand, we still have in our trust a wealth
of wild fauna such as exists nowhere else in the world to-day ;
similar conditions ruled in other parts of Africa, but in most
areas the game has been well-nigh annihilated by the advance of
settlement. In Kenya Colony the conditions are not quite ana-
logous, for here we have large areas not suited for white settle-
ment, and if game can be definitely confined in those areas by
the conservation of adequate water and grazing, and later by the
addition of fencing, the solution of the problem is in sight.
The question is how this state of affairs can be brought about,
and that is where I venture to hope that this Society and also
the Society for the preservation of the fauna of the Kmpire may
be able to play a great part.
I do not propose to pose as an alarmist, but having had the
opportunity of watching the growth of the country for many
years, I consider that the interests of the wild-game fauna are in
a precarious condition, and therefore crave no excuse for urging
that more interest should be taken in its future by the zoologists
of this country.
, Leda eg
pine enna
saps 4 re ern a itt i Prana, act page
hi
; perenne a rs Bees is
Mute ss a sik ten tation ha naead
iS ra stg tha aid ‘gale
ina
arn 2
Liha nigneeans #) canter ty an Ht
ry itis ish. Se ats ee ee
Ce sh GA Tag eae Be
, hab edes® cit ised
bei » ites, ect: ORR
Nudes 5 ay ues cre fa 7 eaaaie’ iy
éi,.\\spi ela fins ie Dre Oran
SR i a cade: Rael Gees
on Pee eet
. une daay ‘i goat a
CY 9 eee VS
Maeh Whe CNIS AN ee ony NSS ie Aa
‘aa aban BAUD FRReN a} se
a &., SS 3 ' hg es iy, +i ¢ i i
‘ Wi oh Valery ie A! et wen * ‘alas Diy
cae Bagi Bi Ee iueahes Pa
ON THE TEMPORAL ARCHES OF THE REPTILIA, We
2, On the Temporal Arches of the Reptilia.
By R. Broom, D.Sc., F.R.S., 0.M.Z.S.
[Received November 15, 1921: Read February 21, 1922. |
(Text-figures 1-4.)
In the last thirty years most of the attempted classifications of
the Reptilia have been based on the structure of the temporal
arches. It is generally agreed that all the later reptiles
have descended from a Lower Permian ancestor or ancestors
that had a roofed temporal region, and the arches found in
later forms have manifestly resulted from a fenestration of the
ancestral roof.
Many reptiles have a temporal region which has resulted
from one opening in the ancestral roof: others have a temporal
region in which two openings have been formed. If all later
reptiles could be classed into those with two temporal openings,
those with only one, and those with none, and if we could be
quite certain that the pair of openings in those reptiles with two
and the single opening in those with one were homologous the
classification would be a very easy matter. Unfortunately, we
have a number of groups where the nature of the temporal region
is not quite clear, and at present there is no general agreement
as to the origin or affinity of these groups.
There is almost universal agreement that at least all the later
reptiles with two temporal openings are closely related and form
a natural group to which the name Diapsida has been given by
Osborn. In this group we have the Dinosaurs, Crocodiles,
Phytosaurs, Pterosaurs, Rhynchosaurs, Rhynchocephalians, and
the primitive order I added a few years ago, the Eosuchia.
There is also general agreement that the Mammal-like reptiles
with a single temporal opening form a second natural group.
But we are left with the following important orders, concerning
which there is doubt—Ichthyosauria, Plesiosauria, Placodontia,
Chelonia, and Lacertilia.
The Ichthyosaurs, Plesiosaurs, and Placodonts agree in having
a single temporal opening. The doubt is whether this single
opening is homologous with the single opening in the temporal
region of the Mammal-like reptiles. The very large majority of
morphologists ave of opinion that it is—I think it is not.
The Chelonians have a temporal region which looks as if it
had never been fenestrated at all, but merely in many types
encroached on from behind or below. Some hold that the
temporal region is still a simple roofed one. But there are
at least two other possibilities. The Chelonians may be derived
from an ancestor which had one temporal opening like the
Plesiosaur, or two as in Sphenodon, where the openings had
become secondarily closed, or they may have originated from an
Proc. Zoo. Soc.—-1922, No. If. 2
18 DR. R. BROOM ON THE
ancestor which had a single temporal opening and whose imme-
diate descendants lost the posterior squamoso-parietal bar.
The lizards, though held by most to be derived from a two-
arched ancestor like Youngina by the loss of the lower bar, are
held by others to be the modified descendants of ancestors which
never had more than one temporal opening.
Some discussion of these debatable points may not be un-
profitable.
Text-figure 1.
A. Skull of Captorhinus aguti (Cope).
B. Skull of Pareiasawrus sp. After Watson.
Cope, Smith Woodward, and all later writers have taken as the
primitive reptilian condition of the temporal region a roofed
temporal region such as is seen in the Labyrinthodonts or in one
a 1 r na a = .
of the early Cotylosaurs, such as Pantylus or Labidosaurus or the
TEMPORAL ARCHES OF THE REPTILIA. 19
allied Captorhinus. But whether or not the reptiles have been
derived froma Labyrinthodont or even a Seymouria-like ancestor
need not at present concern us, as we have a temporal roof in
Pantylus primitive enough to have been the origin of all the
later types of temporal regions seen in higher reptiles. Those
who believe that in many later reptiles, such as /chthyosaurus,
Pleurosaurus, and others, we still have a well-developed supra-
temporal element, must necessarily derive them from an ancestral
type such as we have in Pantylus, where the supratemporal is
still preserved. Personally, I am of opinion that all the so-called
supratemporal bones in later reptiles are really tabulars, that the
supratemporal was very early lost never to reappear, and that a
temporal region such as we have in Captorhinus gives us a better
idea of the ancestral reptilian temporal roof than any other
well-known animal.
It is, of course, a point which might be debated at considerable
length whether it is the tabular which is lost and the supra-
temporal retained in the primitive Diapsids or the supratemporal
lost. The evidence is not as convincing as one would desire;
but, as I am showing elsewhere, there seems good reason to believe
that the upper bone in the temporal region of the lizard is the
tabular, and in the most lizard-like of the Cotylosaurs such as
Procolophon it is certainly the tabular that is retained and the
supratemporal lost, so we are probably justified in concluding that
except in a few primitive Cotylosaurs such as Pantylus and
Diadectes a supratemporal is never found in reptilian skulls.
Captorhinus, though it has all the ancestral elements of the
temporal region and only those that are met with in most later
reptiles, is considerably specialised. The squamosal is probably
considerably larger than in the more generalised ancestor, and
the tabular is certainly more reduced than it must have been.
Still the type as seen in Captorhinus is our best starting-point for
tracing the evolution of the temporal region.
In all the Mammal-like reptiles there seems to me no reason-
able doubt that the temporal condition has arisen by an opening
forming between the postorbital and the squamosal, exactly as
figured by Ver sluys 1 in his ““Synapsider Typus I.” *. This leaves,
when the opening is well formed, a temporal fossa ‘bomendedl above
by the postorbital and-squamosal, and below by the jugal and
squamosal. In Diametrodon and Gorgonops and Galepus we find
this type of fossa, and there is no doubt that, even where as in
Cynodonts and some others, the parietal forms part of the temporal
border, the condition is a secondary one.
The Mammal-like reptiles with this single lower temporal fossa
form a very natural group, to which the name Synapsida has been
given by Osborn.
There is another group of reptiles which, having also a single
* Versluys, I. “ Ueber den Phylogenie der Schlifenruben und Jochbogen bei den
Reptilien,” Sitzungsber, der Heidelberger Akad. der Wissenschaften. Heidelb erg,
1919.
20 DR. R. BROOM ON THE
temporal fossa, have so much superficial resemblance to the
Synapsid type that the larger majority of zoologists have
concluded that they are really allied to the Mammal-like reptiles.
These are the Plesiosaurs. In 1904 I first pointed out that the
Text-figure 2.
Tllustrations of temporal regions in a Plesiosaur, Placodonts, and a Chelonian.
A. Skull of Plesiosaurus macrocephalus Owen. From skull in Brit. Mus.
B. Skull of Placodus sp. From skull in Brit. Mus.
C. Skull of Placochelys placodonta Jaekel. After Jaekel.
D. Skull of Pelomedusa galeata.
fossa in the Plesiosaurs is really formed between the parietal
above and the postorbital and squamosal below, and that it is
thus the homologue of the upper fossa in Sphenodon and not of
TEMPORAL ARCHES OF THE REPTILIA. 21
the fossa in the Mammal-like reptiles which really corresponds to
the lower fossa in Sphenodon. Though this early view of mine
has been opposed by Andrews, Williston, Watson, and others,
I have never seen reason to alter it, as apart from the superficial
resemblance of the upperside of the skull to that of a Therapsid,
the fundamental structure of the Plesiosaur is so different from
that of the Mammal-like reptiles that I never could agree that
the two groups could be at all nearly related. It is with some
satisfaction, therefore, that I find Versluys supporting my view,
that the fossa in the Plesiosaur has had quite a different origin
from that in the Mammal-like-reptiles. Versluys makes it his
“‘ Synapsider Typus IT.”
Hlsewhere * I am dealing at some length with the question of
the aftinities of the Plesiosaurs, and the conclusion to which I
come is that they are a specialised offshoot from land-reptiles on
the line of descent which is leading to the Diapsids, but which
have not yet developed a lower temporal fossa.
In near association with the Plesiosaurs must be placed the
Placodonts. They are a different specialisation sprung from
very similar and closely allied ancestors. There is no essential
difference in the temporal regions of the two groups.
The Chelonians are by far the most aberrant and puzzling
group among the reptiles. Unfortunately, we know nothing of
the ancestry. The earliest-known fossil forms are already typical
Chelonians, and help us very little. Among living forms we have
great variations in the temporal regions. Some have the region
well roofed. Some have practically no roof at all, like the
American Yerrapene. Some have a small roof very deeply
encroached on from below, like the Zvustioalen Water-Tortoise
Lilseya.
For many years I have adhered to the view, which was also
held by Baur, that there is some close relationship between the
Chelonia and the Plesiosaurs. Unfortunately, each group is so
extremely specialised that hardly any apparent resemblance
remains. It is like comparing a Whale and a Bat. The
Chelonian and the Plesiosaur has each a remarkable type of
shoulder-girdle, nowhere else to be found, and each is merely a
modification of the same common type. Itis difficult to believe
that this remarkable type could have been twice independently
evolved. The pelvis, the tarsus, and many points in the skull-
structure confirm the affinity.
If we assume that the Chelonian is a second remarkable
specialisation from a land-type, suchas gave rise to the Plesiosaurs
by another line, we must conclude that the ancestor of the
Chelonian had, like the Plesiosaur, a single temporal fossa, and
that the condition now met with in Chelonians has resulted from
the loss of the posterior arch. A skull like that of Trionya with
its greatly elongated supraoccipital and the squamosal perched on
* Williston Memorial Volume.
22, DR. R. BROOM ON THE
the top of the quadrate readily suggests that the posterior
squamoso-parietal arch has been pushed out, as it were, by the
unusual development of the temporal muscle. And _ this is
confirmed by the fact that occasionally, as in Hydromedusa, the
posterior arch is still met with.
If we are right in this conclusion, we can hardly resist the
further possibility that the Chelonian is quite a near ally of the
Text-figure 3.
A. Skull of Mivosaurus atavus Quenst. After v. Huene. 4% nat. size.
B. Skull of Ichthyosaurus communis Conyb. After the reconstruction by
Sollas, and with the missing squamosal added.
Placodont. Jaekel has shown that Placochelys has a turtle-like
carapace. Perhaps it may yet turn out that Chelonians are
extremely degenerate cousins of the Placodonts.
Another group of aquatic reptiles that have caused some
trouble are the Ichthyosaurs. By some they have been held to
be aquatic specialisations from a Sphenodon-like ancestor. Others
TEMPORAL ARCHES OF THE REPTILIA. 23
have derived them directly from Cotylosaurs with no near
affinity with later forms.
Fortunately, as the result of the work of Sollas and others, we
now know the skull fairly well. In the temporal region there is
a single supratemporal fossa. Behind and below the fossa are
two large membrane-bones, which have given rise to much
difference of opinion. By some, the upper bone has been looked
upon as the squamosal and the lower called the supratemporal.
Most recent writers, however, have identified the lower bone
as the squamosal and the upper the supratemporal. J am also
Text-figure 4.
A. Skull of Foungina capensis Broom. From the type-skull, further restored
from a second specimen. 1% nat. size.
BL. Restoration of greater part of the skull of Thalattosaurus alexandre
Merriam. Further restored from Merriam’s figure in the light of the
later researches of v. Huene.
satisfied that it is the lower bone that is the squamosal. In an
extremely specialised skull like that of Jehthyosaurus the relations
of the bones are manifestly somewhat modified from what was the
primitive condition, but it is agreed by most that the two bones
are undoubtedly the homologues of the two bones usually found
behind the temporal fossa in lizards. These I have elsewhere
agreed are the squamosal and tabular. The supratemporal in the
Stegocephalians and those Cotylosaurs in which it occurs is
apparently merely a roofing-bone with no other important
24 DR. R. BROOM ON THE
functions. ‘The tabular is a much more important element.
Besides being a bone of the top of the skull, it is also an occipital
element with important bony and muscular attachments. The
supratemporal is apparently very rarely, if ever, an occipital
element at all. For these and other reasons which [ am else-
where giving, I regard the upper element in all the later reptiles
in which it occurs as the tabular.
In Mivosaurus, Cymbospondylus, and Ichthyosaurus the back
of the temporal fossa and much of the occipital margin is formed
by the tabular. The squamosal is situated below it. The
relations of the two bones are strikingly similar te those in
lizards.
For the discussion of the temporal region in the two-arched
reptiles we are very fortunate in having two good skuils of an
animal that must be pretty near to the ancestral type. Youngina
capensis, which I described a few years ago, is too specialised in
having a long snout to have itself been the Diapsid ancestor,
but it is sufficiently near for all morphological purposes. The
temporal region is remarkably interesting. The squamosal is
large and lies almost entirely behind the infratemporal fossa.
Above the squamosal lies a bone which I think there can be
little doubt is the tabular. It is situated between the parietal
and the squamosal. The relations of the bones will be best
understood from the figure given. Whatever doubt there might
have been about the identification of the squamosal in the type-
skull is removed by the second skull which I was fortunate in
discovering. This second skull, though it lacks the occiput, has
the snout well preserved.
The Thalattosauria forma group of very remarkable marine
reptiles, which seem to have some aflinity with the lizards and
Sphenodon. Unfortunately, they are very imperfectly known.
Merriam, who first described them, gave good figures of the
specimens and attempted restorations of the skull. Huene, who
has since examined the material, differs from Merriam in the
interpretation of some of the parts of the temporal region. The
most important difference of opinion is that Huene considers
that the squamosal of Merriam is really the supratemporal, and
finds the true squamosal in a much smaller bone situated below
the larger bone. I am inclined to think that in this Huene is
probably right. I give a restoration of the skull, modified from
Merriam in the light of Huene’s later observations, and with
the. occiput restored as I think it may have been. If this
figure be compared with that of Youngina (text-fig. 4, A & B), it
will be seen that Thalattosawrus is strikingly similar in type,
but specialised in a manner curiously parallel to that seen in
the primitive Ichthyosaurs Cymbospondylus and Mixosaurus.
The Lacertilia have given rise to much difference of opinion,
not only as to which of the two post-temporal bones is the
squamosal, but also as to whether they are really descended from
ancestors with two temporal fosse. It has long seemed to me
TEMPORAL ARCHES OF TILK REPTILIA. 25
that it is impossible to get over the fact that the pterygo-quadrate
arch in the lizard is developed exactly as in Sphenodon, as 1 showed
many years ago, and that the immediate ancestor of the lizard must
have had a fixed quadrate. The lizard still has a lower temporal
fossa, which only differs from that of Sphenodon or Youngina
in having become ligamentous when the quadrato-jugal was
lost and the quadrate became movable. The two post-temporal
bones are manifestly homologous with the two in Youngina, and
are thus the squamosal and the tabular. Elsewhere * I am giving
further evidence in confirmation of this view from the consider-
ation of the facts revealed by the study of the skeletogenesis.
The lizards are interesting, as showing in many cases how the
upper temporal fossa of the ancestor may disappear by the
approximation of the bones, but outside the Lacertilia I know of
no forms where this has manifestly happened.
If the interpretations of the temporal regions given above be
agreed to, we can very satisfactorily divide the Reptilia into
four subclasses :—
1. We have the primitive types with a roofed temporal region
which are usually grouped as Cotylosaurs. The Cotylosauria can
hardly be considered as an order. For convenience, every reptile
with a roofed skull is at present placed in it, but such a type as
Procolophon has really very little in common with, say, Diadectes
or Pantylus, and still less with Seymowria. If we still wish to
keep the roofed forms together we must regard them as forming
a subelass and not an order, and Williston’s term ANAPSIDA
seems a ver'y appropriate one, excluding from it the Chelonia.
II. The Drarsipa of Osborn is certainly a very natural group.
Starting in the Upper Permian we have the primitive Hosuchia,
from which we can easily derive all the later two-arched forms,
and also the Lacertilia and the Thalattosauria.
TIl. The Synarsrpa of Osborn, restricted to the Mammal-like
veptiles with a lower temporal fossa, also forms a thoroughly
natural third subclass.
TV. Weare then left with the Mesosauria, the Ichthyosauria,
the Plesiosauria, the Placodontia, and the Chelonia. Can these
be placed in a fourth natural subclass? Very early the veptiles
divided into two distinct lines—those that by a lizard-like habit
early lost the true coracoid and became possessed of a supra-
temporal fossa, and those that by a more Mammal-like habit
retained both coracoid and precoracoid and developed an infra-
temporal fossa. Until the Upper Permian times, when the
two-arched reptiles make their appearance, all reptiles belonged
to the Synapsida, the Anapsida, or to this other proposed sub-
class. Unfortunately, very few of the land-types are known.
Doubtless they became early extinct in the struggle with the
* Williston Memorial Volume.
26 ON THE TEMPORAL ARCHES OF THE REPTILIA.
better-equipped two-arched forms, and we can at present only
examine the very specialised aquatic modifications that survived.
Broonvia is pr obably the only well-known land-form. <Arwosceles
is probably also a member of the group, but so very early that it
still retains the true coracoid. In no other is a true coracoid
known. The Mesosaurs are very early aquatic modifications.
The Ichthyosaurs are much later aquatic forms. The Plesiosaurs
and Placodonts and Chelonians are, I believe, all descended from
land forms closely similar to Youngina, but with only the upper
temporal fossa developed.
This group of primitive lizard-like forms, which had only the
upper temporal fossa developed, and their aquatic descendants
the Mesosaurs, the Ichthyosaurs, the Plesiosaurs, and Placodonts,
together with the very highly specialised Chelonians, which I
consider to be all related, I proposed to unite in a fourth
subclass, which I suggest may be called the ANOMAPSIDA.
ON ANIMAL COMMUNITIES IN THE SOUTHERN NORTH SEA. 27
3. Animal Communities in the Southern North Sea.
By H. Burevap, Dr.Phil., Assistant at the Danish
Biological Station *.
[Received December 24, 1921: Read February 21,:1922.]
(Text-figure 1.)
Having determined the quantity and charted the distribution
of the bottom animals in the Danish Seas (‘‘ Valuation of the
Sea,” I. and II.: Reports XX. and XXI., from the Danish
Biological Station, 1911 and 1913), Dr. C. G. Joh. Petersen in
1914 published an appendix to Report XXI. (in Reports from
the Danish Biological Station, XXII., 1914), in which he, on the
basis of various statements in the literature, gave a hypothetical
chart of the animal communities outside Denmark. In this
the North Sea is charted as chiefly a ‘“ Venws”-community, with
the “ Macoma baltica” community in some places along the
shore, and an area in the deeper parts of the North Sea, marked
with a 2, in which several indications in the literature suggested
the occurrence of a “ Brissopsis”-community. In the month
of July 1921 I was enabled to accompany the English Fishery
Research vessel ‘George Bligh’ on a cruise across the North Sea
from Lowestoft to Esbjerg, during which Dr. Petersen’s bottom
sampler, “ the Petersen Grab,” was used; by the courtesy of the
Director of Fishery Investigations, Dr. E. 8. Russell, I was given
permission to work out and publish the material procured.
Though the route followed did not touch the deeper parts of the
North Sea, as will be seen from the accompanying chart of the
stations (p. 30), and consequently cannot give information about
the existence or not of the “ Brissopsis” area mentioned above, still
the 20 stations worked during this cruise can be of service in
giving a preliminary orientation of the communities in the
Southern North Sea.
In the accompanying Valuation List the stations have been
arranged in communities. It will be seen that with some
hesitation I have referred the first two stations to the ‘‘ Venus ”-
community. These two stations contain, however, too few
bottom animals to settle the question. Among the characteristic
species of the “ Macoma baltica” community, which, with the
kind help of Captain Davis, I found in large quantities by digging
along the shore-line at ebb-tide in the harbour of Lowestoft, viz.,
Macoma baltica, Scrobicularia plana, Mya arenaria, and Cardiwm
edule, only one small Mya arenaria was found, at Station 2;
whereas a representative of the “ Epi-Fauna” of this community,
viz. Mytilus edulis, is present in numbers at this station. On the
other hand, a small Ophioglypha albida and two Natica aldert point
* Communicated by E. 8. Russexn, M.A., D.Se., F.Z.S.
LO I
8.0 G
Ue a3
|
G0 | @
10.0 | 1
WG | 1
OE eG
€.1 6
Hae OL
Pee. 03
‘VuaVy
JO Siete 81} 60 ST
wee wee wae 2.0 | OL
iO i ToalacOn 9
0 1 Ore peau
LO 1 it By |
oo ee = ee ee
TO 1 T AO VT
e i) |
Oe te Lee)
205 Ee 1k
(610) 9) es ate €
a0 jo | YG
THO G21 Sipe
TO) | ieee |
10.8 be ie OE
CO leGe SiG 69 | T
eee oe oS Sane
BO G1 8 8: SO | Ait
os Ico.0 ik
Ico.o | 1
Pa) |="
TRO yee Tec SL a ae
On I
gan) Z| s¢ se selina | 2,
Bee) 2S | Se eee S
3g | os 3 Byg| SS
Spe | val otk eM, am] @
“SINWOUTITA-WOIGUVOONTHOW
‘admis urojjoq ‘tw "bs 7.9 Yyrar ueyVy orem sepdurus Z
“1G
g
GA
Sey a
MWasoy DLMDUIQ09T
Se tas eee eepraep ey
punanUny wyaydg |
saaoad UyOURG ‘ds shyqyda xy
labuusp DOE |
“ pypnonu vppruoy |
: "ds svawanr }
“ds 20y70U wy
ao)
alan ncS BC)
ie oo
PD ea
‘6L ST LT‘9T ‘IT ‘OT “6 “8 “L ‘9 ‘GG ‘DP ‘ES
96
96 96 86 86 8€
epee ogee anal DAGALAZ 17797 ANT, fi
~ Deonipurtia puyoylip
9p] DEON | |
5 age SAE pe ““snsonxapy snun py
ns (ellshel elfesare “ant 66 ce
q@-ant
cae rio nae sosn " pqquh ppy1g.top }
SOnIBCC sijmpa SnyUyhTT
ocacon one DPUZUU DININAT |
surppnhunrs) a7..ngsy f
“ant (¢ 799227) HUE
“ant es
puLywb snua 4
sngnjqia xnuog |
“ant wndsoynis :
SEI
ce
DPVIUNAIGQNS VAJOD IAL
“Anl DiwwualD DATE |
ant’ o ss
~ myngng mane
we) Dorgouisitd us
“9979 Dugy |
“13 UL
YUSTOM
[e300
Yonoy
~ sarjeut ur yydeq
‘SONA
‘AINE TIS—TI9
darasn_
UOT BAS yore FW
CAOLSH MOT
| | fae) R =
Haws | 9 | S ay Dine ate 3 2 2 ep iad
2 £ 3 8 y [= ERs 3 3 3 ¥ SS ~ S
io i = eee a 5 See
| ot Ss * . “ w , n ;
cr = Car) M 9 e RN
we S = i) N = 2
° — | |) = PL es =I = a Ps
EG oe | iz a ¢g = =] inj
ct = “4 S H = b = a. Q SS
fa ps < .
RQ £ ) = 3 3 3 3 So ,
» = a we g 5
Be i oF cee g 2, =
. | By, ° ~ a 5 4 =
| NS RM > | Th n a M
oS 5 | + mo» 2 iS) S
iz 2 s = gb fo) ° = a5 S = - = S oO =] a .
= re = oa s © © 2 WoLLog {oO UTLOVAVAH
rie = D Oo = i= wa L
oe 5 nD | ue ise) ee n z
ree | 4 > | iS) & 2 . 4 Te |
55 | “ ©, | a s ss a = a 2 ea
= “ s — | S sy i=
S Mie ie ee | a = a
S S | 3 Be © © |
| | | Sao, lac
| ® 3 3 3 2° S) 32S |
d
S om
Sy Fs
| | Ss ESS | |
SS J
SCS. wel co a ae ie ae a IHOIAM TYLOY,
| i. sas He. ees ie | eH |) Gy 9% ‘ds wniaayy yy
pee ee Peg cof et ee eee gy Pg | gn | ao tt fucteettinetrtese ads maungngetog |
|
|
z anl snpsnyuiag snunbodngr §
Gin Se Sle a ES a aaa aa as GaP 2 aug Seay 3] RCL Seog signujsot snyauhysouazy |
I 2. eS eer Rot see | tre eee Jens eee ens ant snunpjaaisspa sagsh.op ;
a3 eer eis ‘antl sogn.undap snung10g |
““w“afiurds snoosougsv5 |
se | 06 BIIVUUN,) |
1.0 Om [eco se Le Ws] KO) I izes) Eb ey als Hy Ge ake ee i | oo eeprremTey) |
ee 0 ; srusofyyl vunryduy
F ! eects er £.0 | de leu. oo eyo. 2 ee ey ee ee 1.0 I Wagon ears saoboone snyprsnd snuphoouyor
aie ie (Wpaone ee ZOE ett eG ee ee ee ee ge ers He tay forseeteresens can fleds “
T I wee see on 2.0 WSL sae woe wee wae tee wae WS UL. see eae wae wee eee . Ung psod UUNIPADIOULY I HT
Simi
~ Tro |
S| |
O “wSrq) PLT | | a
aes eee eee | eee Jose. see eee OO B.0 G ee ove T I wee wee z I. tae . . ee vee eee COL? tee pee [eee ee ween ers ere CEE :
ee le ge mee Ot ne MR PR a es ow ee eee Sit :
. F.0 it I oe ae see tise | on see one dies Ser wee eee see eve ose aoe or Ser see bee aYsie tee oe [egeegmngani the opinpetaeeg3 ‘ds DIADUD) T |
oxoreca:
a
=)
ce
re
30 DR. H. BLEGVAD ON ANIMAL COMMUNITIES
to the “ Venus ”-community, if anything; so it is possible that
the hard sand and gravel bottom here in the strong current off
Lowestoft is a transition area between these two communities ;
at all events it is certainly a very poor area, exception made of
the Epi-fauna (to this also must be reckoned Stronglocentrotus
drobachiensis, the Hydroids, and Sabellaria alveolata, which last-
named species is not found in Danish waters inside the Skaw).
Stations 3-1] and 16-19 are typical ‘“‘ Venus ”-stations, with
the characteristic species Venus gallina, Tellina fabula, Mactra
Text-figure 1.
3
Chart showing Stations worked by the ‘ George Bligh,’ 6th-8th July, 1921.
Stations 1-2. Venus (?) Community.
aN 3-11, 16-19. Venws Community.
‘ 12-15. Echinocardium-filijormis Community.
Station 20. Abra Community.
subtruncata, Donax vittatus,and Hchinocardium cordatum. Among
the ‘‘ attendant species,” ¢.e. species frequently occurring in the
community in question, but which may also be found in others,
IN THE SOUTHERN NORTH SEA. 31
I may particularise Vatiea alderi, Ophelia limacina, Pectinaria
korent, and Hehinocyamus pusillus. In stations with no character-
istic animals, I have attached importance to these species, which
in conjunction with the depth, condition of bottom, and scarcity
of animals, all point to the ‘‘ Venws”-community. The occurrence
of Abra prismatica and Mactra elliptica in some stations is in-
dicative of the “ deep Venus” community, which has been found
in the deeper parts of the Danish ‘“ Venus”-community ; but as
the characteristic species Hchinocardium flavescens and Spatangus
purpureus, so typical of this community, are wanting, I can for
the present only designate these stations as ‘“‘ Venus ”-stations in
general.
Stations 12-15 are characterised by such animals as Amphiura
jiliformis, Turritella terebru, Auvinus flecwosus, and Hehinocardiwm
cordatum. Stations 12 and 13, however, are transitional from
the “ Venus ”-community, containing several of the characteristic
animals of this community but mostly as small specimens, and
only a few Amphiura. But as a whole these stations must be
reckoned as belonging to the “ Hchinocardiwm-filiformis” com-
munity. The bottom, consisting previously of hard sand or
eravel, is here softer, mixed with detritus and clay. ‘This
community is not represented in Dr. Petersen’s hypothetical
chart in Rept, X XIT. (on this point see remarks in Rept. XXIII.,
1915, p. 10, at the top) ; it was charted in Rept. XXI. as the
& Hehinocardiua- Pur ritella ’-community, which 1s widely distri-
buted in the Kattegat, Skagerrak, and Christiania Fjord. Without
doubt it will later become apparent that this community covers
big areas in the deeper parts of the North Sea.
The last station, No. 20, in the neighbourhood of Graa-deep
light-vessel, is a typical “Abra ”-station, with Abra alba, Nucula
nitida, and fragments of Hchinocardium cordatum. The existence
here of an “Abra”-community was already proved in 1912 by
some few samples taken by the ‘ Michael Sars’ in the neighbour-
hood of Graa-deep and Horns Reef.
We have thus met with four different communities :—a ‘“J/acoma
baltica”’-community along the shore in Lowestoft Harbour; a
“« Venus ”-community in the greater part of the investigated area
of the North Sea, with depths most frequently between 9 and
38 m. and a pure sand and gravel bottom; an Kchinocardiwm
jiliformis community in depths of 38-41 m. with a dark, soft
sandy bottom ; and finally, an Abra-community near the shores
of Jutland. Possibly this last-named community also will be
discovered near Lowestoft, as ina single sample taken with the
grab in Lowestoft Harbour, in a depth of 3 m.at ebb tide, I found
one adult Abra alba. Butit is beyond doubt that the greater part
of the bottom fauna of the North Sea below 38 m. will turn out
to belong to ‘“ Venws”-community, as indicated in Dr. Petersen’s
chart. ‘his is confirmed also by the few samples taken in former
years with the bottom sampler off Thyboron (Report XXI.,
Appendix pp. 63-66), and by the numerous samples which
J
.
32 ON ANIMAL COMMUNITIES IN THE SOUTHERN NORTH SEA.
according to the friendly communication of Dr. Russell, have
been taken with the bottom sampler this year on the Dogger Bank.
As to the number of animals per square mile, it will be seen from
the accompanying list (pp. 28-29) that, setting aside the Epi- Fauna,
we have to do with a comparatively poor fauna. For instance, the
“Venus”’-stations together give only 26°3 gr. per 2°6 sq. m. or
10-1 gr. per sq.m. But on the other hand, most of the animals
are useful as being excellent fish-food. Compared with the
‘ Venus”-community in the Kattegat (Rept. XXI., Appendix,
pp- 9-18), it will be seen that the apparent richness of the latter
is due mainly to big echinoderms such as Hehinocardium and
Ophiurids, or big lamellibranchs such as Cyprina or Aiacoma
calcarea, useless as fish-food.
On the other hand, we may find in the North Sea as well as in
the Kattegat small areas within the “ Venuws’”-community, in-
habited by large numbers of Mactra subtruncata (see, for instance,
St. 8 in Rept. XXI., Appendix, p. 65, with over 2 kg. per sq. m.,
also among the samples taken by the ‘George Bligh’ this year
on the Dogger Bank, several of which contain some hundreds of
specimens per 0°2 sq. m.). It would be of the greatest interest
to know something further about the distribution and abundance
of this most important food species, at different times of the year
and at different places in the North Sea. At the same time
continued investigation of the stomach-content of the food-fishes
is called for, following up the excellent work of Todd (Second
Report on the Food of Fishes, North Sea Investigation Committee,
Southern Area, Second Report, Part I., 1904-5). From his
paper it would appear that the food of the fish in the south-
western North Sea is composed mainly of “ Venuws’’-animals.
But, as mentioned above, we have in the deeper central parts of
the North Sea an extended area, which is still a terra incogita
and still awaits charting.
ny
ath vies
Waitt
Lee AY. 5
>, Z S, Il, SSVERRUEIES, Jel
DENDRONEPHTHYA CLAVATA, Kuklenthal.
PZ S, Il, SVERIRWEES, Pl, Ul,
DENDRONEPHTHYA GIGANTEA, Verrill.
Ri Zo So MI, SVR WMS, Il. IO!
DENDRONEPHTHYA ANNECTENS, sp. n.
EVOLUTION WITHIN THE GENUS DENDRONEPHTHYA. 33
4, Evolution within the Genus Dendronephthya (Spongodes)
(Aleyonaria), with descriptions of a number of species.
By W. Rae Suerrirrs, M.A., D.Sc., F.LS., Lecturer
in Zoology, University College, Southampton *.
[Received August 10, 1921: Read November 8, 1921.1]
(Plates I.-III.f ; Text-figures 1-30.)
Part [.—GeENERAL.
§ 1. All who have worked at Aleyonarians will agree as to the
difficulties presented by the genus Dendronephthya or Spongodes.
For here we have to deal with a multitude of species within
a relatively narrow range. Thus Kiikenthal, in his * Versuch
emer Revision der Alcyonarien: II. Die Familie der Neph-
thyiden, 2. Teil’ 1905, deals with no fewer than eighty-seven
species ; and Henderson, in the ‘ Aleyonarians of the Indian
Ocean,’ Part II. 1909, with another series of sixty different from
the former, And yet in both cases the species are described in
minute detail.
The continual experience in investigating a collection of repre-
sentatives of this genus is that, in spite of an initial determi-
nation to refrain from adding to the already large number of
described species, one is forced to do so. And there is no
denying that each of these new forms has a distinct individuality.
This experience inevitably raises a number of ztiological questions
which it may be useful to state, although they cannot be more
than partially answered. Some of them at least could be replied
to by an investigator in a good locality, having at his command
a large number of specimens of any given form.
§ 2. The outstanding phenomena which present themselves are,
apart from the multitudinous species, the following :—
A. That the specific distinctions are all of a relatively trivial
sort, such as mode of branching, grouping of polyps, length of
polyp stalk, strength of supporting bundle, presence or absence
of a definite ‘ crown,” the number and the arrangement of the
spicules in the anthocodial points, the distinction between the
cortex of the polyparium and that of the “sterile stalk,” and the
nature of the spiculation of the canal-walls.
B. That the species differ one from the other to a large extent
in the congeries or collocation of such characters as we have
mentioned ; that is to say, two species with similar anthocodial
armature may differ in the mode of branching, and vice versa.
C. That within the limits of a colony there is, in most cases,
* Communicated by Prof. J. ARTHUR THomson, M.A., LL.D.
+ For explanation of the Plates see p. 77.
Proc. Zoou. Soc,— 1922, No. ITI. 3
34 DR. W. RAH SHERRIFFS ON EVOLUTION
relatively little variability of architecture, though there are, of
course, quantitative differences in size of spicules, strength of
supporting bundle, and the like, which are merely growth
characters. I must emphasize the saving clause in: most cases,
in view of the occurrence of a number of very variable species,
such as D. gigantea, where there is great variability of antho-
codial armature in one and the same colony. In the second part
of this paper particular attention is paid to the interesting form
referred to; and it does not stand alone.
§ 3. The multiplicity of form detail within a narrow range
admits of various interpretations :—
I.(a). It may be urged that the observed differences are
individual, not specific, characters. If the observed differences
are purely individual, they may be either extrinsic modifications
or of the nature of intrinsic variations. And before considering
these alternatives separately, I may note the general answer
that in many cases a considerable number of specimens from the
same or different localities agree together. ‘Thus Kikenthal
speaks of twenty specimens of D. savignyi, twelve of D. gigantea,
ten of D. rubra, seven of D. pumilio; while I have myself got
twelve of D. ehrenbergi, ten of D. gigantea, ten of D. clavata, eight
of D. habereri, and the same number of D. hyalina.
Therefore the general suggestion that observed differences are
purely individual does not apply in every case. At the same
time, the unsatisfactoriness of basing a new species on a single
specimen when the specific characters are of such a tenuous
description is plain.
(6) It may be said that the observed’ differences are of the
nature of exogenous modifications correlated with environmental
peculiarities. Against a too ready acceptance of this easy
solution is the occurrence of the same form in widely separated
localities. For example, D. robusta is reported from Ternate,
Borneo, Zanzibar, and Madagascar; D. florida from Hong Kong,
the Philippines, and Port Jackson (Australia).
It may also be pointed out that certain factors in the
immediate environment in different parts of a large colony must
show considerable diversity, cf. shelter; and yet, as already
pointed out, there is rarely diversity in a colony except in colour
and the like.
(c) It may be, however, that the observed differences are
expressions of individual intrinsic variability, not correlated with
any peculiarities of environment. There is no way of testing this
interpretation until numerous similar specimens from the same
locality and of the same age and size are examined by some
investigator with abundance of living material at his disposal.
II. It may be that the multiplicity of distinct forms within a
narrow range is due to cross-fertilization between allied species.
a |
WITHIN THE GENUS DENDRONEPHTHYA. 3E
Tt is quite conceivable that species which have attained individu-
ality and fixity in the course of isolation and inbreeding may be
brought into contiguity by subsequent spreading or removal of
barriers. In such a ease, analogy points to the likelihood of
numerous new patterns arising by permutations and combinations
of the previously segregated species. This hypothesis is certainly
suggested by the fact that two 0 species may agree in anthocodial
armature and yet differ in mode of branching, or may agree in
having a foliaceous collar and yet differ in the nature of the
supporting bundle. In short, in the distinctions between species
there is a distinct suggestion of the shuffling of unit characters.
III. There seems to be a third possible interpretation—namely,
a mutation of species apart from any hybridizing influence. It
may be that certain widespread and strongly established species
such as D. gigantea, D. chrenbergi, and D. brevirama have been
the stocks from which mutations have been thrown off after the
fashion of Qnothera lamarckiana.
An indirect argument in favour of this interpretation may
perhaps be found in the possibility of discriminating similar
radiations of evolution within the three great groups : Glomerates,
Divaricates, and Umbellates. A move direct argument may be
found in the variability of certain species such as D. gigantea, to
which reference has already been made. As Kiukenthal remarks,
“ Aus diesen Beschreibungen geht hervor, dass D. gigantea eine
in ihrem Aufbau sehr variable Art ist.” (‘ Revision,’ p. 553.)
§ 4. Kukenthal has divided this difficult genus into the three
main groups: (1.) Glomeratie ; (II.) Divaricatee ; (III. ) Umbellatee,
giving precision to similar suggestions by previous workers such
as Holm.
I. The Glomerate are characterized by :—
(a) the comparatively slight branching of the polyparium ;
(6) the grouping of numerous bundles of polyps into roundish
bunches which make the surface of the polyparium
entirely irregular.
There is a marked definiteness about the Glomerate division
which suggests “naturalness,” and makes it easy to refer a
species to the group. Text-fig. 1 is a diagrammatic illustration
of what is meant by the Glomerate habit of growth.
If. The Divaricate are characterized by :—
(a) the profuse branching of the polyparium ;
(6) the length and slenderness of the twigs ;
(c) the divergent separateness of the polyp bundles ;
(d) the absence of anything that can be called een of the
Glomerate type or umbelg of the Umbellate type.
It should be noted that a Divaricate polyparium may have a
continuous contour like that of a well-pruned tree (see diagram).
2%
Text-figure |.
A typical Glomerate.
Text-figure 2.
A typical Divaricate.
EVOLUTION WILHIN THE GENUS DENDRONEPHTHYA., 37
IIT, The Umbellate are characterized by :—
(a) the umbel-like or sometimes corymb-like aggregates formed
by the terminal twigs, the heads of the umbels being
bundles of polyps;
(b) the disposition of all or most of the polyp heads on the
surface of the colony.
Text-figure 3.
A typical Umbellate.
Thus the Umbellate differ from the Glomeratze, and agree with
the Divaricate in showing much minor branchiug. They differ
from the Divaricatze in the presence of umbels or corymbs of
polyp bundles and in the entirely superficial arrangement of
the polyp heads. As this continuous superficial disposition
is obviously advantageous—giving all the polyps equal expo-
sure for nutritive and respiratory purposes—we may regard
the Umbellate as the latest expression of the evolution of the
Dendronephthya polypavium. Moreover, some Umbellatie pass
through a Divaricate stage.
Of minor importance is the question whether or not the umbels
combine into large secondary bunches rising like hillocks on the
surface of the polyparium. Nor can I attach more than slight
inportance to the shape of the polyparium whether flattened or
quite symmetrical, whether spherical or disc-like, and so on, for
these features probably depend to a large extent on local environ-
mental conditions.
It may be noted that a large and very handsome colony of
D. annectens was, as a whole, strikingly divaricate at first sight,
but a closer inspection soon showed that it was an Umbellate
having the terminal twigs massed into distinct umbels. (See
Plate ITT.)
38 DR. W. RAE SHERRIFFS ON EVOLUTION
$5. Before proceeding to consider the specific characters
selected for examination and criticism, it will be well to recognize
clearly the typical architecture of a Dendronephthya polyp as
Text-figure 4.
Spicules or
tentacle
Anthacodia/
Intermediate spreuses
spoicule pear) 3)
} Crown (©)
Accessory
spicule
-- ——- —- =
Supporting
b6uUNnd/e
ages (main)
spicules
Armature of lateral point of Polyp.
regards its external spiculation (text-fig. 4). When every por-
tion is represented, the ascending order above the polyp stalk
is as follows:
(1) The spicules of the supporting bundle (Stutzbiindel), several
of which are usually very large (often up to 3 or 4mm. in
length) and often projecting beyond the polyp for a
distance of 0°5-1-5mm. It is useful to distinguish in the
supporting bundle the main spicules and the accessory
spicules, the latter often mingling gradually with those of
the polyp stalk.
(2) The “crown,” consisting of a few rows, very commonly
three, of spicules arranged horizontally round the base of
the anthocodial part of the polyp.
(3) The anthocodial armature, in the strict sense, which
consists mainly of the spicules entering into the com-
position of each of the eight points. Between each pair
of points tiny spicules are frequently found. These I term
intermediates and discount their value because they are
so variable.
(4) Very minute spicules on the tentacles, the nature and
number of which seem to be quite unimportant.
WITHIN THE GENUS DENDRONEPHTHYA. a9
§ 6. If, as I think probable, the Glomerates, Divaricates, and
Umbellates represent three distinct stocks or lines of evolution,
the particular problem is to discover whether similar morpho-
logical radiations can be discerned in each of these main
divisions. A prior question is—What characters can be re-
garded as having real morphological importance ?
1. Negatively, it is quite plain that little emphasis can be laid
on the size of the colony, looseness or density of branching,
amount of sterile stalk or colour.
2. Positively, the features which seem most indicative of
relationship are in order of importance :—
(1) the architecture of the anthocodial armature—e. g., whether
crown and points are distinct, the number of spicules in
each point, their degree of projection and their differenti-
ation within the point ;
(2) the differentiation of the supporting bundle ;
(3) whether the polyps are marked by long stalks or are very
short-stalked ;
(4) characteristic features in the spiculation, such as the occur-
rence of particular forms in special regions of the colony
e.g., the “sterile stalk,” canal-walls.
§ 7. In studying the possible relationships of different species,
I make the following provisional assumptions :—
(a) That the presence of very numerous small spicules in the
anthocodial points is more primitive than a reduced number
of large spicules.
(6) That the absence of a definite “‘ crown ” is more primitive
than its differentiation.
(c) That a slightly differentiated supporting bundle with a
large number of spindles tending to form a sheath rather than
a buttress is a more primitive condition than the differentiation
of a bundle out of a few large spicules. And here it may be
noticed that in a few species—e. g., D. clavata, the supporting
bundle is so slight that the polyps may at first sight be mistaken
for those of a Hunephthya, where there is no supporting bundle
whatever.
(d) That the more primitive forms show :—
(1) less variety of spicule: i.e., are predominantly provided
with spindies, the characteristic spicule, and less variety
of spicule in the anthocodial point ;
(2) less localization of particular forms of spicule in parti-
cular areas. Thus there can be no doubt that the
presence of special interlocking spicules in the ‘sterile’
stalk” is a later differentiation ;
(3) the presence of few spicules in the canal-wall.
Thus D. piitéeri with a multitude of eanal-wall spicules is in other
respects highly specialized. Conversely, it is interesting that
40 DR. W. RAE SHERRIFFS ON EVOLUTION
D. clavata with no canal-wall spicules belongs to what I would
call the more primitive types. So careful a worker as Kikenthal
attaches great importance to the canal-wall system of spicules,
and it is obviously important to inquire what correlation there is
between the primitiveness of a species and the number and
nature of the canal-wall spicules. The unsatisfactory feature
here is that the examination of the canal-wall spiculation has not
been made uniformly by the various investigators. Minute
spicules which escape attention in
a fragment of canal - wall when Text-fig. 5.
boiled down may be found by the
other method of examining a piece
intact. But one is not uniformly Ih
successful with the sample examined.
§8. Bearing in mind the first of the
above assumptions, it is possible to ex-
pand it so as to reduce all the different 7 KK
Species to one or other of the follow- Laer gw,
ing six grades, ranging from the most aaa
primitive (1.) to the most specialized a
(VI.) :—
VI. Only one pair of spicules, or
even but a single spicule re-
placing all the others in each Semmes
point above the crown ; ‘Sas
VY. reduction of the point that one
pair (or, it may ae one of the 4
? ?
uppermost pair) predominates ees Y N
over the others, which are not aE
above three pairs in total
number above the crown ; UN
GN
IV. one pair or one spicule predo- on IN
minating over 3—4 others, no OT
crown ; a”
IIT. about 4-6 pairs, the uppermost
slightly specialized, no crown ; ‘
II. about 6-8 pairs, strong and aN
uniform, no crown ; ZX
° . ° WW t 4
‘I. about 8-12 pairs in each poin t 4S
numerous and small, rising \\
from indefiniteness to definite - we
ness, no crown.
—
==»)
Ui
§9. The suggestion has been made by critics of Dendroneph-
ihya species that the differences between the Glomerate, Divari-
cate, and Umbellate groups are not of taxonomic value, but
4
WITHIN THE GENUS DENDRONEPHTHYA, 4)
depend on conditions of growth. On this view, the occurrence
of a species with the same anthocodial armature in each of the
three groups would not mean that a similar specialization had
occurred three times on different lines of evolution; it would
mean that one and the same species occurred in three different
vegetative forms—Glomerate, Divaricate, and Umbellate, which
depended on growth conditions.
For various reasons this view is improbable:—(1) In many
cases the Glomerate, Divaricate, or Umbellate mode of growth
is clearly expressed in young forms 1-3 em. high. (2) When
exactly the same anthocodial armature occurs in two or all of
the three groups, the two or three species in question differ in
other details, such as the length of the polyp stalk, the nature of
the supporting bundle, and the spiculation. This does not look
like the occurrence of the same species in three vegetative guises.
There are Umbellate colonies, however, which seem to pass
through a Divaricate stage when they are young and small, but
these are in my experience fewer than forms which are Umbel-
late from near the start.
Text-figure 6.
Glomereate
UmbeHate —
Divaric32te
In support of the view that a colony usually shows very early
whether it is going to be Glomerate, Divaricate, or Umbellate,
I have given a series of diagrammatic representations of a
dozen small colonies drawn .to actual size. It goes without
saying that the definite architecture cannot be expressed until
there are quite a number of polyps in the colony. But the limit
is uncertain. The central figure of the group depicts the
primitive, quite undifferentiated colony from which scon the
distinctive form embodies itself. Above it to the left are
Glomerates, to the right Umbellates, while below is a row of
Divaricates.
10. Starting with the Glomerate, I suggest that the fol-
lowing should be regarded as illustrations of primitive types :—
1. In the savignyt group :—
D. argentea with irregular arrangement—most primitive of all,
49, DR. W. RAK SHERRIFFS ON EVOLUTION
with ventrally and laterally nothing more than minute cylindrical
corpuscles. D. fusca with indefinite double rows of about 6 pairs—
on a slightly higher level, but also showing primitiveness in
numerous quite small elements. D. savignyi with indefinite
steep double rows reduced to 6—somewhat higher, but also
showing, like the preceding, very numerous small cylindrical
bodies.
2. In the hemprichi group :—
D. clavata, where occur 10-12 pairs in each double row and a
very weak supporting bundle. It may be noted that in this
group we have a gradual reduction in the number of spicules
composing each point 5-7, 5-6, 4, 3, 2 and 1, reaching a climax
in forms like D. punicea, where are found a couple of slightly
hockey-club like spicules occupying the whole area which in
D. clavata, for instance, shows 10-12 pairs.
3. In the studeri group :—
Here we have, on the whole, a more differentiated group with
forms like D. mayi with 6-8 pairs in each point to be ranked
as relatively primitive compared with the highly-specialized
D, kollikert.
§ 11. So with the Divaricates, on my view the series begins
with forms like D. japonica with 8 pairs in each point, none
protruding—a very primitive arrangement—and ends with
D. suensoni, which has but one pair. As intermediate between
grade IT., which I define as having numerous pairs regularly
arranged in each point (grade I. possessing very numerous
irregular pairs), and grade III., with 5-6 equal pairs, we have
cases like D. armata, where the lateral points go up to 9 pairs.
There must also be noticed D. cerulea inclining to be primitive in
minuteness but not in number of spicules, while D. Alunzingert
may be considered as rising out of grade II. because one of the
uppermost pairs is very prominent.
It is interesting to see that in so many of Kiikenthal’s minor
groups his arrangement of the species ends in one pair, so that
the anthocodial armature, which I hold to be the evolutionary
key, is, as used by Kiikenthal, very probably just diagnostic.
Again, in the rigida group, D. microspiculata is undoubtedly
primitive with 6-8 pairs, though the uppermost again is slightly
larger and projecting.
§ 12. In the third great division—the Umbellates—grade I. is
represented by D. australis with its very numerous, small polyp
spicules, grade IT. perhaps by D. wmbellulifera with 6-8 pairs and
the practical absence of a supporting bundle.
In the dendrophyta group, D. villosa belongs to grade I. with
many spicules in each double row, while grade II. in the florida
group may be illustrated by D. brevirama (with 6-8) and D. florida
(with 5-7).
WITHIN THE GENUS DENDRONEPHTHYA. 43
D. pectinata must be regarded as the climax in the line of
evolution.
Again in the rubra group, grade I. is well represented by
D. repens, while in the spinulosa group, grade II. is probably
represented by D. spinulosa itself with 6-8 pairs very small and
not projecting, even better by this than by D. flabellifera where
we have 8-9, but in which I recognize what I regard as the
beginning of differentiation: viz., a specialization of the highest
pair.
In tabular form my conclusions may be represented as
follows :—
I. 10. JON
GRADE. GLOMERATES, DIVARICATES. UMBELLATES.
c ph Ge Az) he Ul Skee Cae | eid:
(| D. punicea (Stud.)*. ( -D. piitteri (Kiik.). ( D. lutea (Kik.).
| D. carnea (Wr. &Stud.). | D. swensoni (Holm). || D. longicaulis (Kiuk.).
VL4 D. deederleini (Ktik.). | D. orientalis (Hend.). } | D. simplex (sp. n.).
: < | D. flammea (sp. u.). |
| | | D. cervicornis(Wr. &Stud.). |
U L| D. laxa (Wr. & Stud.). Ne
v.{ ' D. eburnea (Kiik.). ; | D. coronata(Wr.&Stud.).
(| D. spinifera (Holm), ( D. mollis (Holm). ‘a D. disciformis (Kiuk.).
| | D. kéllikeri (Ixik.). | _ D. cirsium (Kiik.). | D. rubra (May).
IV J ) D. involuta (Kiik.). } D. pumilio (Studer).
a} | D. mirabilis (Hend.). D. nigrescens (Kiik.).
| | | D. collaris (Wr. & Stud.).
lL L |. D. sinensis (Piit.).
(| D. studeri (Ridley). ( j ~ DD. dendrophyta (Stud.).
lll D. gigantea (Verr.). \ D. hyalina (Kiik.).
j l / D. mexicana (Kiik.).
. ; D. florida (Esper).
( D. mayi (ik.). ( D. klunzingeri (Stud.). ( D. brevirama (Burch.).
D. hemprichi (Klunz.). | D. ehrenbergi (Kik.). | D. umbellulifera (Kiik.).
IL.< ~ D. japonica (Kuk.). <~ D.spinulosa (Gray).
| j | D. microspiculata (Put.). D. habereri (Kiik.).
Uy D. erinacea (Kuk.). D. annectens (sp. n.).
| D. argentea (Kik.). ( D. villosa (Kiik.).
1) D. clavata (Kiik.). —— / D. australis (Kiuk.).
(l _ D. repens (Kiik.).
* The species printed in italics are those worked out from the Dendronephthya collection
of the ‘Siboga ’ Expedition.
As a convenient means of expressing the composition of the
anthocodial point, I have drawn up an anthocodial formula on
the following lines:—The spicules of the point are denoted by
“P” if big and strong, and by ‘‘p” if small and weak; those
of the crown by “Cr”, and the supporting bundle by the letters
““S.B.” preceded by a qualifying adjective, such as weak, medium, ~
very strong, etc. In this way the anthocodial grade and formula
of D. annectens, for example, can be set forth briefly as :
IJ.=8 p+0Cr+very weak S.B.
44. DR. W. RAE SHERRIFFS ON EVOLUTION
Part I1.—DrscrIPTION OF SPECIES TAKEN BY THE
‘S1poGaA’ EXPEDITION.
TABLE OF SPECIES.
Genus DenpronepuTeya (Kik.).
| Number
| examined.
|
\
Locality.
I. GLOMERATA.
| Station 302.
Stations 165, 213, 240
261, 303. 5 |
Station not marked.
. DIVARICAT A.
| Stations 164, 258.
| Station 164.
‘ | Stations AmboinaReef,
310 B, 366. ;
5 BOR ETO:
| es 99, 282.
| Station 91,Macaras Reef.
per 5 VRe, 19R
% 315.
;
bs 125, East of
Polloe Weh. §
. UMBELLAT A.
5 3D), Cllawwennay (IRE) ga of Soe ono 10
ID, eae) (WEN) oon oss vss anavo- 10
Bs ID), joe) (SiC) soeces che sad sue 1
I
A. I). ehrenbergi (Kuk.):...........| 12
5. D. suensoni (Holm) ........ ...... 2
6. D. mollis (Holm).............00-.1 4
7. D. cervicornis (Wr. & Stud.)... 2
8. D. mirabilis (Hend.) ............| 2
@), JD), iilerravone, (js We) aco vennse soo 00: 2
10. D. klunzingeri (Stud.)............ 2
11. D. microspiculata (Piitter)......) 2
12. D. orientalis (Hend.) ............) 4,
Ill
13. D. collaris (Wr. & Stud.) ...... 2
14. D. longicaulis (Kik.) ..,......... 1
15. D. disciformis (Kwk.) ........... 6
UGS JD), Ineoeswer (UGH) Sone. Gencnde: 8
17. D. pumilic (Studer) ........ 4,
18. D. coronata (Wr. & Stud.)...... 3
19. D. brevirama (Burch.) 6
20. D. anmectens (sp: 1.) ... ....... il
PAL, 1B), smajolas (Gos Ws)) sonsst ceo ooses 1
22. D. hyalina (Kiuk.) 8
400) 925) oo eee a 93
Station 99.
310.
Stations 99, 164.
» 258, 282.
|, 99, 258, 315.
Station not marked.
Stations 164, 258.
Station not marked.
- not marked.
Stations 257, 318, 321.
J. GLOMERATAL.
‘1. Dendronephthya clavata (Kiikenthal).
Previously reported
from
| S.W. of Japan.
)
|
)
Japan, Hong Kong
Japan.
Red Sea.
Japan.
Japan.
Lifu, Funafuti, Kei.
Andamans.
Red Sea.
| Philippines, Hong Kong,
Amboina.
| Andamans.
Kei.
Japan.
China Sea.
Japan.
Japan.
Torres Straits.
Torres Straits, China Sea.
Pescadores.
(Plate I.)
Described by Kiikenthal, ‘Revision der Alcyonarien,’ pp. 549-6
(1905).
Draenosis :—Glomerate; with great bare gap between the
collar and the polyp-bearing area ; polyps in dense groups at the
ends of the terminal branches: polyp stalks very long ; support-
ing bundle weak; point spicules 7-8 pairs, short and uniform ;
no distinct crown; grade I.; tentacles with tiny, flat, Jagged
spicules; the foliaceous collar with large curved spindles ang
WITHIN THE GENUS DENDRONEPHTHYA, AD
small, quaint, flattened forms; colour: general cortex and sterile
stalk yellow, collar vivid red, anthocodial armature whitish,
twigs and polyp stalks with red spicules.
* ANTHOCODIAL GRADE AND ForMULA:—
L=aP+0Cr 4+ weak S.B.
Descriptive Noves :—
1. Colony as a whole. Ten specimens of a peculiar form, the
smallest 15 mm. in height, the largest 17 em. ‘They are remark-
able in showing, especially in the larger forms, a great bare gap
between the collar and the loosely glomerate polyp-bearing area.
In the largest specimen this interval extends for about 6 em.
Here and elsewhere it is extremely limp, and it looks as if the
polyp-bearing area had rested on the mud. In the upper part of
the colony there are numerous galls due to and containing small
Balanide, and the polyps are very muddy. It should be noted as
a feature that the upper part of the polyparium is distinctly top-
heavy; in the largest specimen it has a breadth of 10 em.,
whereas the stalk bearing it is only i em.across. The colony
broadens again at the very foliaceous collar and measures 6 em.
from side to side. Below that the sterile stalk, which is markedly
flattened, measures 3°5 cm. in breadth. There are numerous
stolons.
2. Branching. The three largest spe-
cimens show two lateral short branches esate Glee ae
arising immediately above the collar, fe
but not affecting the bareness of the gap pballag
alluded to. 6 ulin
3. Colouring. The twigs and polyp 7)
stalks bear red spicules; the anthocodial J
armature is whitish or transparent; the 4,
general cortex is yellowish; the collar is 4
‘of a very conspicuous red; the sterile Y
stalk has the same colour as the general
cortex.
4. Polyp stalks and their spicules.
The length of the polyp stalks is very
ereat—often reaching 3 mm. in the
collar and about 1:5 mm. in the upper
polyparium. The spicules are red and D. clavata Kiik.
very thorny.
5. Polyps. Found in dense groups at the tips of the end
branches.
6. Polyp spicules. The anthocodial architecture is primitive,
having 7 or 8 pairs of very uniform short rod-like spindles in
each point and no distinct crown. The supporting bundle is
* In the anthocodial formule the point spicules are denoted by “P” if big and
strong, and by “ p” if small and weak; the crown spicules by “Cr,” and the supporting
bundle by “S.B.” preceded by a qualifying adjective (ef. page 43).
46 DR. W. RAE SHERRIFFS ON EVOLUTION
very weakly developed and shows only occasionally a projecting
needle.
7. Other spicules. Those of the sterile stalk are almost en-
tirely small, star-shaped forms. In the foliaceous collar we find
(1) large, curved, almost smooth spindles up to 5 mm. in length;
(2) quaint, flattened forms, like biscuit-fingers in shape, not
exceeding 1 mm.; (3) the small red, rough spicules of the antho-
codia ; and (4) the tiny, flat, jagged ones of the tentacles.
SALIENT FEATURES :—The peculiar mode of growth (which seems
to be unique), the limpness of the whole colony, the length of the
polyp stalks.
I refer my specimens to D. clavata, although Kikenthal’s
description does not mention any growth peculiarity such as has
been noted here. The important points of agreement are the
following :—
(1) a glomerate mode of branching (Glomerate), with the polyp-
bearing twigs predominating over the stem and main
branches in building up the polyparium (hemprichi
group);
(2) a primitive anthocodial armature with 7-8 (D. clavata
10-12) pairs of very uniform, short, rod-like spindles in
each point; no distinct crown ;
(3) a very weakly developed supporting bundle which only
occasionally shows a projecting needle ;
(4) a long polyp stalk of from 1-2 mm, (about 1 mm. in
D. clavata) ;
(5) a well-developed foliaceous collar ;
(6) an almost exclusive occurrence of small, plump stars and
double stars in the cortex of the sterile stalk ;
(7) a practically complete absence of spicules from the canal-
walls. With agreements so numerous it seems entirely
justifiable to disregard the extraordinary growth pecu-
liarities.
Locauiry :—Station 302.
2. Dendronephthya gigantea (Verr.). (Plate IT.)
Described by Verrill, Bull. Mus. Comp. Zool. Cambridge, p. 40
(1864).
DiaGnosts :—Glomerate ; with numerous rounded, polyp-
bearing masses, the lowest branches slightly foliaceous; polyps
densely arranged in characteristic hemispherical masses ; polyp
stalks short, under 1 mm.; supporting bundle very strong ; point
spicules very variable, 1— 6 pairs; no crown; grade IV.; . spicules
of the upper cortex are large (4 mm.), bovered with blunt warts
and lie transversely, lower cortex and canal-walls have stouter
and shorter spindles, triradiates and multiradiates; colour : cortex
white, polyps deep orange.
WITHIN THE GENUS DENDRONEPHTHYA. 47
ANTHOCODIAL GRADE AND FoRMULA :—
III. = (1—6) P + 0 Cr + very strong 8.B.
Descriptive Noves :—
1. Colony as a whole. ‘Two handsome orange-coloured colonies,
the largest 28 cm. wide and 20 em. high, eight others smaller
in size.
2. Branching. Markedly glomerate. ‘The numerous rounded,
polyp-bearing, boss-like masses predominate over the stem and
main branches. Therefore the colonies must be included in
Kiikenthal’s hempricht group. The lowest branches of the colony
show slight foliation.
3. Colouring. Generally deep orange-coloured polyps and
white cortex. The colony from station 240, however, showed
quite a different colour-scheme, viz. bright red spindles in the
supporting bundle and anthocodiz, but the general cortex greyish
with opaque, white, almost porcellanous spindles.
4, Polyp stalks and their spicules. Short, under | mm.
5. Polyps. Densely arranged in very characteristic plump hem1i-
spherical masses of 7 or so.
6. Polyp spicules. The anthocodial architecture without enter-
ing into details, may be noted as having the 8 points often elon-
gated with the tips meeting above the polyp. Another feature
is the practical absence of any true crown. Kiikenthal notes in
his definition of D. gigantea that each of the 8 points has 5-6
pairs of spicules, the uppermost much larger than the others and
markedly projecting. It is easy enough to find on the specimens
anthocodize which exactly correspond with this description.
Text-fig. 8. 3 Text-fig. 9.
Y AAA
D. gigantea Verr. D. gigantea Very.
Yet on the same colony may be found points with 5, 4, 3, or
2 pairs, and finally even a single pair, abutting against a few
transitional small spindles leading on to the supporting bundle.
Great care was taken to avoid damaged or disturbed antho-
codiz, and the annexed diagrams show a series of frequently
recurring types of architecture.
A general feature may be noted that in the great majority of
eases the two uppermost predominate, and that when the number
48 DR. W. RAE SHERRIFFS ON EVOLUTION
is reduced to 2 pairs or to 1 pair, these occupy the same space as
3-6 rows. The larger numbers 5-6 were certainly less frequent
than the rest.
There is not in this case any confusion between what might be
counted to points and what to crown, for in no case did I see
what in other species is called a crown. It is also striking that
after an examination of scores of anthocodiz hardly any case of
intermediate spicules between the points was found.
The supporting bundle is enormously developed. Its largest
spicules may attain to a length of 6 mm. and project for 1-5 mm.
These large spindles are densely covered with minute, blunt,
thorns, except for 0°6 mm. at the projecting tip, which is smooth,
as is also the case in LD. giganteé. Very marked in the large
supporting-bundle spicules of the colony (Station 240) is the
disappearance of the red colour from the proximal third of most
of the largest. The same tendency is to be recognized in the
yellow colonies.
7. Other spicules. The spindles of the upper cortex are often
up to 4 mm. in length. They are covered with large, blunt,
truncate and sometimes compound warts, and they lie trans-
versely. Many show an abrupt narrowing at one end. In the
lower cortex and canal-walls there are numerous stouter and
shorter spindles exceeding 1 mm. in length and bearing crowded,
rough tubercles often compound. Large triradiates and multi-
radiates also occur, besides bracket-like and quite irregular forms
approaching the stellate type. A few curiously starfish-like forms
occur, and triradiates with one arm exuberantly branched. Many
much smaller counterparts of the large types occur. There are
also occasional long, narrow spindles.
Within the hemprichi group the specimens agree best with
D. gigantea, though this may not be obvious at first sight. Many
of the anthocodize show but one pair of spicules in each point,
which suggests approximation to D. carnea, D. dederleimi, and
D. punicea. From the first they are at once separated by the
massive spicules, which are visible from a distance and lie in very
regular transverse rows. This conflicts too seriously with the
original description by Wright and Studer (p. 196), where it is
said: “The stem and branches ..... are leathery and thickly
packed with fine spicules. The latter are not recognisable by the
unassisted eye and lie scattered confusedly in several layers.”
Both from D. dederleint and from D. punicea they are sufficiently
separated by the enormous strength of the supporting bundle.
Depvuctrion :—The specimens agree more closely with D. gigantea
than with any other Glomerate known to me, and not least in
showing quite an unusual degree of variability. Among the
- important features of agreement the following may be noted :—
(1) the very characteristic plump hemispherical masses closely
beset with polyps ;
(2) the supporting bundle of great strength and showing many
WITHIN THE GENUS DPNDRONEPHTHYA. AQ
spindles of over 4 mm. (the maximum size mentioned for
D. gigantea) and projecting for 1:5 mm.
(3) the large spindles of the supporting nual densely covered
with minute, blunt thorns, except for the last half milli-
metre at the projecting tip, which is smooth as in
D. gigantea ;
(4) the spindles of the upper cortex frequently reach 4 mm. in
length, are covered with large, blunt warts, and lie trans-
versely ;
(5) the spicules of the lower cortex and of the canal-walls;
(6) the eight anthocodial points are often elongated and ‘have
the tips meeting over the polyp;
(7) the lowest Michies of the colony show slight foliation ;
(8) the colour-scheme of deep orange polyps and white cortex
was found in some of the described species of D. gigantea.
LOCALITIES :—
Station 165 (3 specimens). Station 240 (3 specimens).
» 213 (1 specimen). », 261 (2 specimens).
., 03 (3 specimens),
3. Dendronephthya punicea (Studer).
Described by Studer, Ann. Mag. Nat. Hist. (6) vol. i. p.70 (1888).
Diaenosis :—Glomerate; bundles of polyps in rounded clusters ;
main stem and main branches little developed; polyps closely
crowded together, polyp stalks medium ; ; Supporting bundle strong ;
point spicules ive only, strong and converging ; crown of alot
three rows of spicules irr eoularly disposed ; grade VII.; spicules:
upper cortex has big, strong, thick spindles (2 mm.), lower cortex
smaller and more fborny forms, including roundish and stellate
types; colour: branches and all spicules scarlet; polyps pale
yellow.
ANTHOCODIAL GRADE AND ForMULA :—
VI. =2P+4 3Cr + strong8.B.
Descriptive Noves :-—
1. Colony as a whole. A handsome Glomerate colony, with the
bundles of polyps forming rounded clusters, often well separated
from one another.
2. Branching. ‘There is relatively little development of the
main stem and main branches as compared with the stalks of the
bundles of polyps composing the clusters. It is therefore one of
the hempricht group.
3. Colouring. Branches and all spicules scarlet while the
polyps themselves are pale yellow.
A. Polyp stalks and their spicules. The polyp stalks measure
almost 1 mm. in length.
5. Polyps closely crowded together.
6. Polyp spicules. The anthocodial armature has each point
Proc. Zoou. Soc.— 1922, No. LV. 4
50 DR. W. RAE SHERRIFFS ON EVOLUTION
consisting of two strong, very thorny, converging spicules bent at
thé base in hockey-club-like fashion, one slightly larger than the
other. Between adjacent points lie a
pair of much smaller, straight spindles Text-fig. 10.
longitudinally disposed, but in some
eases there seem to be two pairs (thus
connecting with D. dederlein). Below (7 AX)
the points there is a crown of horizontal
spindles in two or three rows, rather Zo
irregularly disposed.
The supporting bundle is strongly
developed, especially as regards the three
uppermost spindles, the lower median
one of which projects for almost D. punicea Stud.
1 mm.
Depuction :—Among the members of the hempricht group
there are several with only one pair of spicules in each of the
anthocodial points, and beside these this specimen must be
ranked.
It agrees very closely with D. punicea in the following
features :—
(1) each anthocodial point composed of two spicules, one
slightly larger than the other ;
(2) between adjacent points usually a single pair of small
intermediate spicules ; ;
(3) below the points the crown of horizontal spindles is present
with two or three rows ;
(4) the strong supporting bundle.
Locaurty :—Station not marked.
Il. DIVARICATA.
4. Dendronephthya ehrenbergi (Kiik.).
Described by Kiikenthal, ‘ Korallentiere des Rothen Meeres,’
p- 56 (1904).
Draenosts :—Divaricate; contours irregular; polyps in little
groups of 5-8; polyp stalks short; supporting bundle medium ;
point spicules 6 pairs; crown absent; grade ITI.; spicules: canal-
walls show few bent rough spindles, the stalk cortex has very
distinctive long curved red spindles and irregular clubs, discs, and
spheres ; colour: rich red in the coonenchyma, anthocodial spicules
grey-yellow, polyp spicules of the points white.
ANTHOCODIAL GRADE AND KorMULA :—
Il. =6p + 0Cr + medium 8.B.
Descriptive Norres :—
1. Colony as a whole. Ten specimens in all, including two
fine specimens agreeing closely with Kiikenthal’s definition of
this species, the largest 40 cm. in length.
WITHIN THE GENUS DENDRONEPHTHYA, Hil
2. Branching. Markedly divaricate but not markedly flat-
tened. The contour of the colony is irregular. These features
point to the swensoni section, and in that section the only species
to which the specimens could be referred is D. ehrenbergi. The
mode of branching is the same, and the resemblance of these
specimens to Kiikenthal’s coloured figures is striking except that
they have much less of a sterile stalk.
3. Colouring. Rich red in the ecenenchyma and with greyish-
yellow anthocodial spicules.
4. Polyp stalks short. Text-fig. 11.
5. Polyps arranged in little divaricate bat
groups of about 5-8.
6. Polyp spicules. There are in most ‘i
6 pairs of white spicules arranged en A y Xe gh
chevron in each of the anthocodial lines :
in Kukenthal’s forms there were 4—5.
There is no very distinct crown. The
supporting bundle consists of thick,
rough red spindles (up to 1:5 mm.),
projecting freely for about 0°5 mm.
7. Other spicules. The canal-walls
show a few bent, rough spindles. The
spicules of the stalk cortex are mostly
very distinctive, long (up to 3°5 mm.),
eurved, red spindles, also ovals and D. ehrenbergi Kik.
irregular clubs covered with massive
tubercles mainly with very characteristic rough blunt heads.
There are also very rough discs, approaching spheres, covered
with the same massive tubercles. <A few triradiate forms occur
in the lower cortex. Not very numerous, there are also minute,
colourless, irregular capstans.
Depuctrion :—The specimens agree with D. ehrenbergi in
having :—
(1) 6 pairs of spicules, in most cases, in the anthocodie ;
(2) supporting bundle of rough spindles slightly projecting ;
(3) polyps in little divaricate groups of about 5-8 ;
(4) spiculation of the canal walls ;
(5) mode of branching.
Locauitrss :—-Stations 164, 258.
5. Dendronephthya suensoni (Holm).
Described by Holm, Zool. Jahrb. viii. Syst. pp. 35-37 (1895).
Dracnosis :—Divaricate; outline irregular; not obviously
flattened ; polyps in little groups (4-10), distinctly scattered ;
polyp stalks medium ; supporting bundle medium; point spicules
one pair only, of which one member is a long projecting curved
spindle associated with a much smaller one at its base; crown of
some 6 rows of horizontally disposed spindles; grade VI.;
4*
52 DR. W. RAE SHERRIFFS ON EVOLUTION
spicules: canal-walls show numerous forms with greatly deve-
loped thorns ; colour: cortex and polyps have deep red spicules,
polyps grey-yellow.
ANTHOCODIAL GRADE AND ForMULA :—
VI.=1I+1P+4+4-6Cr + medium 8.B.
Descriptive Norgs :—
1. Colony as a whole. Two handsome colonies agreeing very
well with Holm’s figures.
2. Branching. Markedly divaricate, of irregular outline, not
obviously flattened, and should be referred to Kukenthal’s
suensont group of the Divaricate.
3. Colouring. Deep rose-red spicules Taste 1D,
on cortex and polyps; polyps themselves =
apart from the spicules ave yellowish
Bley
ore
4. Polyp stalks about 1:3 mm. long.
5. Polyps in little groups (4-10) )
distinctly scattered. ( ‘
6. Polyp spicules. Anthocodiz show a=
8 points, each consisting of a long, Go S
projecting curved spindle with a much =z NS
smaller one at its base, the two being ANS
obviously members of a pair. These
points rise from a crown of about 6 rows
of horizontally disposed curved spindles. D. suensoni Holm
The supporting bundle consists of only
three strong spindles up to 3 mm. long, one of which projects for
only a short distance.
7. Other spicules. The canal-wall contains numerous forms
with conspicuously developed thorns.
Locatity :—Station 164.
6. Dendronephthya mollis (Holm).
Described by Holm, Zool. Jahrb. vii. Syst. pp. 51-53 (1895).
Draenosis :—Divaricate; contour regular; not flattened ;
polyps in groups of about 10; polyp stalks medium ; suppor ting
bundle very strong; point spicules 4 pairs with uppermost pair
(or one of that pair only) strong and projecting; no crown;
gerade IV; spicules: stalk has rough spindles, triradiates and
quadriradiates ; colour: general surface white, anthocodial
armature and supporting bundles red-brown.
ANTHOCODIAL GRADE AND ForMULA :—
IV.=1-2P+3p+40Cr + very strong 8.B.
DescriprivE Nores :—
1. Colony as a whole. Four colonies of white general surface,
two of which are obviously young and are fragmentary.
2. Branching. WDivaricate, regular outline, not flattened.
WITHIN THE GENUS DENDRONEPHTHYA. ay]
Should be referred to Kiikenthal’s Divaricate and section
diwaricata, within which they approach D. mollis.
3. Colouring. General surface white, anthocodial armature
and supporting bundles reddish brown in the largest specimen.
In the two younger colonies the anthocodial and supporting-
bundle spicules are amber-yellow. The fourth colony is ivory-
white.
4. Polyp stalks about 1 mm. long. Text-fig. 13.
d. Polyps arranged in groups of about
ten in number.
6. Polyp spicules. One of the upper-
most pair of spicules in at least some of eo
the 8 points is strong, out of proportion iw, “ee
to the others and projects very markedly
beyond the polyp. The other spindles YN
of the points are small and converging, Y WN
and there may be 4 pairs of them. In
many cases the large projecting spindle Ss
has fallen off, and its attachment seems
to be very loose.
The supporting bundle is very strongly D. mollis Holm
developed and 38 or so of its component
spicules project beyond the polyp for over 1 mm. The spindles
are densely and regularly covered with very small blunt tubercles,
while here and there almost smooth forms occur.
7. Other spicules. Those of the short stalk include, besides
spindles much rougher than those of the anthocodie and more
curved, various triradiate and smaller irregular quadriradiate
forms.
Locatiry :—Amboina Reef.
The two young divaricate colonies with amber-yellow antho-
codial and supporting-bundle spicules have distinct flattening,
and show what we regard as essential characteristics of D. mollis,
viz :—
(1) a very marked projection of one of the uppermost spicules
of each point in the anthocodial armature ;
(2) the presence of 3 or 4 other pairs of smaller spicules
converging en chevron ;
(3) the practical absence of any crown ;
(4) the very strong supporting bundle with 3 specially strong
spicules, one of which projects beyond the others for
about one millimetre ;
(5) the dense regular covering of the anthocodial, supporting-
bundle, and ccnenchyma spicules with short blunt
tubercles ;
(6) the prevalence on the short stalk of much rougher spindles
(straight, curved, and much curved), with much longer and
more distant tubercles, and along with these numerous,
irregularly shaped, roughly tuberculate forms.
54 DR. W. RAE SHERRIFFS ON EVOLUTION
A few spindles occur with asymmetrical, bifid ends. The polyp
stalks are longer than in the other forms.
Locatiry :—Station 310 -B.
The fragmentary, ivory-white colony, without a stalk, divari-
cate in type, in its branches inclined to be flattened, closely
approaches D. mollis in showing :—
(1) a very marked projection of one of the uppermost spicules
of each point ;
(2) the others (usually 3 pairs in number) much smaller ;
(3) a very strong supporting bundle with 3 specially strong
spicules, one of which projeets for at least 1 mm. ;
(4) the dense regular covering of the spicules with short,
blunt tubercles.
Most of the spicules are curved spindles, many boomerang-like;
bifid and trifid forms occasionally occur; here and there one is
almost quite smooth as also occurs in D. mollis.
LocaLiry :—Station 366.
Depuction :—It appears to me that the most striking feature
of this species is the exaggeration of one of the terminal spicules
of certain anthocodial rows, but I emphasise the point that many
of the heads show not a trace of this.
7. Dendronephthya cervicornis (Wright & Studer).
Described by Wright & Studer, Rep. Sc. Res. ‘ Challenger,’
vol. xxxi. pp. 220-221 (1889).
Diaenosis :—Divaricate; rigid; outline irregular; distinctly
flattened; polyps loosely spaced ; polyp stalks long; supporting
bundle very strong; point spicules one pair only, one member of
which is stronger than the other and projects somewhat; crown
of about 5 rows closely packed together; grade VI.; spicules ;
stalk surface has numerous rough multiradiate interlocking
discs ; colour: bright orange as a whole.
ANTHOCODIAL GRADE AND FORMULA :—
VI. =2P + 4-5 Cr + strone 8.B.
Descriptive Notes :—
1. Colony as a whole. An approximately rigid, bright orange,
young colony.
2. Branching. Markedly divaricate; outline irregular and
flattened ; should therefore be referred to Ktikenthal’s cervicornis
group in which, however, the only species that it approaches
closely is D. cervicornis itself.
3. Colouring. Bright orange as a whole.
A. Polyp stalks long.
5. Polyps divergent and loosely spaced.
6. Polyp spicules. The composition of the anthocodial armature
WITHIN THE GENUS DENDRONEPHTIYA. 55)
shows 4-5 horizontal spindles closely packed in a crown and of a
pair of converging spindles in each of the 8 points, one member
of the pair often larger than the other
and projecting a little beyond the polyp. Text-fig. 14.
The supporting bundle is very strong,
and in it one spindle may attain a length
of 3 mm.
7. Other spicules. The surface of the
stalk is densely covered with roughly
tuberculate multiradiate discs, often
with a central knob, which fit into each
other.
Depuction :—This specimen does not
agree with previous descriptions in
colouring or in showing foliaceous basal
branches, yet I think it should be
ranked within the heading of D. cervi-
cornis. D. cervicornis Wr. & Stud.
Locatiries :—Stations 305, 310.
8. Dendronephthya mirabilis (Henderson).
Described by Henderson in ‘ Alcyonarians of the Indian
Ocean,’ Part Il. pp. 49-51 (1909).
Diagnosis :— Divaricate; loose ; contour irregular ; distinctly
flattened ; polyps in small bundles of about 6; polyp stalks
medium; supporting bundle medium; point spicules 5 pairs,
with the uppermost pair slightly projecting; crown absent; —
grade IV.; spicules: many of the stalk spicules characteristic
in appearance because of the length and strength of the pro-
tuberances ; colour: colony as a whole white, polyp stalks,
point spicules, and supporting bundle amber-yellow.
ANTHOCODIAL GRADE AND FORMULA :—
IV.= 5p + 0 Or + medium 8.B.
Descriprivi Norms :—
1. Colony as a whole. A white, divaricate colony,
2. Branching. Loose, divaricate, with irregular contour and
very distinct flattening. It should therefore be referred to
Kiikenthal’s group Divaricate, subgroup cervicornis. Yet it
agrees in several respects with Henderson’s D. mirabilis, which
he places in the rigzda group.
3. Colouring. Polyp stalks, spicules of the anthocodial arma-
ture and of the supporting bundle amber-yellow.
4. Polyp stalks not more than 1 mm. long.
5. Polyps occur in small bundles of about 6 in number.
6. Polyp spicules. The anthocodial armature consists of
8 double rows of about 5 pairs, converging steeply, and with the
uppermost pair slightly projecting. This does not agree well
56 DR. W. RAE SHERRIFFS ON EVOLUTION
with Henderson’s description and figure. The supporting bundle
is moderately developed and has about 3 projecting spindles,
one of them slightly longer than the others. The projecting
portion is about a third of a millimetre in length.
7. Other spicules. What is most cha-
racteristic of this specimen and Hender- Text-fig. 15.
son’s is the appearance of many of the
stalk spicules. They are spindles straight
and curved, spindles approaching clubs
and very rough at the broad end, irre-
gular bodies and many quadriradiates,
all marked by the length and strength
of their spines and protuberances, which
are sometimes compound and often tu-
berculate. Some of the curved spindles
have these prominent processes especially
strong about the middle, while others
are very markedly unilateral, with a few
tubercles on one side and a great array 1D), aprcidaipe lend).
of long, often divaricate processes on the
other. ‘These unilateral processes are strongest at one end of
the spicule and wane towards the other. Among the extra-
ordinary forms, which present an appearance unusual in the
genus, there are commonplace straight spindles with small regular
spines and tubercles.
LocALITIES :—Stations 99 and 282.
9. Dendronephthya flammea (sp. n.).
Diaenosis :—Divaricate; outline quite irregular; markedly
flattened ; lower branches somewhat foliaceous; polyps in small
branches of 3-6, but many isolated polyps occur; polyp stalks
very short ; supporting bundle strong; point spicules one strong
pair; crown of 4-5 rows of straight rough spindles; grade VI.;
spicules: sterile stalk has large rough yellowish spindles along
with small red interlocking forms; no spicules found in the
canal-walls; colour: brilliant red as a whole, spicules of th
branches and twigs deep red.
ANTHOCODIAL GRADE AND FoRMULA :-—
VI.= 2 P + 3-4 Cr + strong 8.B.
Descriptive NorEs :—
1. Colony as a whole. Two colonies, the larger measuring
17 em. in height, including the 3°5 em. sterile stalk, by 10 em.
in breadth.
2. Branching. Divaricate, quite irregular in contour, markedly
flattened. The lower branches are somewhat foliaceous, but do
not forma collar. It ought to be referred to the cervicornis group
of Kiikenthal’s Divaricatee and to the subdivision with short
polyp stalks ; but it does not agree with any of the three species
WITHIN THE GENUS DENDRONEPHTHYA. 57
there included, nor with any other Divaricatse approaching this
group. Hach of the half-dozen main branches ramifies irregularly
in a flabellate fashion, but there tends to be a predominance of
secondary branches to the one surface, though the surface is
richly beset with small, almost sessile bundles of polyps, which
also occur all over the cortex. These small bundles may, of
course, be young stages of twigs.
3. Colouring. Predominantly of a brilliant red. The cortex
is vividly coloured except at the beginning of the polyparium,
where it is almost white. On the branches and twigs all the
spicules are deep red.
4. Polyp stalks very short.
5. Polyps. A somewhat marked feature is the small number
(3-6) of polyps in each bundle and the frequency of isolated
single polyps.
6. Polyp spicules. The anthocodial Moxtofee lG.
armature consists of 8 strong points, Se
each composed of 2 rough hockey-club
spicules rising from a strongly-developed
crown of 4-5 practically horizontal rough
spindles. A deceptive appearance arises
im many cases where the non-retracted
state of the polyp brings the crown
spicules into a chevron arrangement in
a line with the upper pair of what I
take to be the point proper. The two
hockey clubs, which form a narrow
isosceles triangle, are seen in other cases
in the normal relation of points to
crown, and it may be noted that they
are distinctly larger than those beneath
them. Between every two points there
are usually two intermediates. On the whole in anthocodial
structure the arrangement is nearest to that of D. cirsiwm. ‘The
anthocodial spicules are almost white, but show a faint yellowish
tinge. They are characteristically rough, with relatively large,
very blunt tubercles, not densely crowded.
The supporting bundle is well developed and inclined to be of
the sheath type, showing 3 spindles much larger than the rest
that is to say, a median and two laterals abutted by numerous
smaller ones, which show a tendency to be in parallel lines sloping
towards the support. The median spindle usually projects for
about 0-5 mm., and the two laterals project sometimes a very
little. The supporting sheath spicules are spindles often curved
and densely covered with short, rounded tubercles.
7. Other spicules. On the sterile stalk are large, broad, rough, |
yellowish spindles connected by small interlocking irregular
reddish forms which produce an arenaceous appearance. ‘The
cortex here bears a variety of spicules, all thickly covered with
rather massive, blunt, rough tubercles, sometimes compound and
D, flammea, sp. n.
58 DR. W. RAE SHERRIFFS ON EVOLUTION
well suited for interlocking. They include the following forms :—
(a) long stout spindles, slightly curved ; (6) spindles expanding at
one end in a pseudo-club fashion ; (c) a few large triradiates ;
(d) numerous irregular triradiates ; (¢) smaller triradiates with a
rough boss rising at right angles to the rays; (f) long-armed
quadriradiates ; (g) knobbed spherical ‘“ astroscleres.”
The spicules of the cortex in general are spindles densely
covered with relatively larger, rough-tipped, rounded tubercles.
Especially on the shorter spindles do these tubercles stand out to
a notable height like mammille, projecting to about one-fifth of
the breadth of the spindle.
On the branches and twigs all the spicules are deep red, and
there are numerous large spindles which tend to project on the
surface in jagged fashion, being on the whole transversely
disposed.
No spicules were to be found in the canal-walls.
The following ineasurements were taken :-—
(a) Supporting-bundle spicules ..............:.... Up to 2 mm.
(6) General upper-cortex spicules ............... Up to 4 mm.
(c) Largest spindles of the cortex of the
Stétletshallke ak cawa ty cascereuenn. ace ean eee Up to 2 mm.
Locauiry :—Station 91. Macaras Reef in 27 fathoms.
10. Dendronephthya klunzingeri (Stud.).
Described by Studer in Ann. & Mag. Nat. Hist. (6) vol. i. p.72.
Dracnosis :—Divaricate ; contour discontinuous ; polyps 10-12
in each bundle, which resembles a close corymb; polyp stalks
very short; supporting bundle strong ; point spicules 8-10 close
together, none projecting; no crown; grade II.; spicules: in
the stem loosely disposed slender spindles minutely spinulose, in
the cortex of the sterile stalk spindles are more thorny and
include triradiates and quadriradiates; colour: white on the
whole, but with red-yellow twigs and polyp stalks.
ANTHOCODIAL GRADE AND FoRMULA :—
Il. = 8-10 p + 0 Cr + strong 8.B.
DeEscRIPTIVE NorEs :—
1. Colony as a whole. Markedly divaricate; contour dis-
continuous. A single, young colony.
2. Branching. Branches of varied lengths and growing in one
plane. The colony therefore ought to be referred to Kiikenthal’s
““B” section of the group Divaricate, and certainly in the
neighbourhood of D. klunzingeri.
2. Colouring. Predominantly reddish yellow in the twigs and
polyp stalks, elsewhere whitish.
4. Polyp stalks very short.
5. Polyps. Each bundle of polyps is rather like a close corymb
and contains 10-12 heads. The tentacles are soft.
WITHIN THE GENUS DENDRONEPHTHYA., 59
6. Polyp spicules. The anthocodial armature of 8-10 pairs of
white spicules, closely opposed en chevron with no more than a
hint of projection, is the salient feature
of the colony. Owing to their white Text-fig. 17.
colour these spicules stand out very
conspicuously. Their surface is covered dt...
irregularly with minute tubercles, which “yf NG
are much more crowded towards the EN
tips. 4, SS
The supporting bundle is strongly st NY
developed, its largest spicules projecting Y NY
NX
for 0°75 mm. BAZ
7. Other spicules. The stem has ——
leosely disposed spindles, many of them
having only minute and rather distant
spinules. These tend to be slender.
Those of the surface of the sterile stalk D. klunzingeri Stud.
show a stronger development of thorns.
There are, besides spindles, many irregular triradiates, quadri-
radiates, and brackets.
Kiikenthal lays stress on the great reduction of the stevile
stalk. This is far from the case in this specimen, where it
oceupies 1°5 cm. out of a total height of 4em. From what I
have seen in other cases, I do not think that the proportion of
stalk to polyparium is of much importance.
Kiikenthal also notes that the outermost polyps bore markedly
projecting spicules at the tips of the points. Of this feature this
specimen shows-no more than a trace; but on account of the
characteristic anthocodial armature, I cannot separate it from
D. klunzingeri.
Locautry :-—Station 164 EH.
11. Dendronephthya microspiculata (Pitter).
Described by Piitter, ‘ Aleyonaceen des Breslauer Museums °
in Zool. Jahrb. xii. Syst. pp. 459-460 (1900).
Diaenosis :—Divaricate ; firm; long stalked ; lowest branches
slightly foliaceous ; polyps in divergent groups of 4-10; polyp
stalks long ; supporting bundle strong; point spicules 6—8, small,
close-set, the uppermost pair predominating and slightly pro-
jecting ; no crown; grade II.; spicules: in the tentacles a
double row of transversely arranged narrow, jagged, reddish
spicules; in the cortex of the sterile stalk thick, bent, warty
spindles and quadrangular bodies; colour: stalk, supporting
bundle, polyp stalk, and tentacles a fine red; general cortex white
flushed with red ; anthocodial spicules white.
ANTHOCODIAL GRADE AND FORMULA :—
Il. =I1P + 5-7 p+ 0 Cr + strong 8.B.
DEscRIPTIVE NotTEs :—
1. Colony as a whole. A somewhat firm young colony with
60 DR. W. RAE SHERRIFFS ON EVOLUTION
a long, limp stalk; total length 5 em., of which 1:5 is poly-
parium.
2. Branching. The lowest branches show only hints of being
foliaceous.
3. Colour. A fine coral-red in stalk, Text-fig. 18.
supporting bundle, polyp stalk, and ten-
tacles; the general cortex of the poly-
parium is white flushed with red, and
the anthocodial spicules are white.
4. Polyp stalks long.
5. Polyps in divergent groups of 4-10.
6. Polyp spicules. The armature of
the anthocodiz consists of 8 points of
small spicules, very compactly disposed,
6-8 en chevron in each point; the upper-
most pair in each row predominate and
may project slightly.
The supporting bundle is strongly D. microspiculata Pitter
developed, and one strong central
spindle projects for 1 mm.; the spicules here are thickly but
finely spinose.
7. Other spicules. ‘The tentacles show a double dorsal row of
transversely disposed, narrow, jagged, reddish spicules ; the cortex
of the sterile stalk has thick spindles, usually bent, covered with
large warts, often compound. Along with these are smaller bent
spindles, often with large prongs on the convex side. There are
also numerous bright red irregularly quadrangular or knob-like
corpuscles, which may be vaguely called “stars.” These are
well figured by Pitter. All these stalk spicules with prongs are
suited for interlocking.
Perhaps the most characteristic features are (1) the numerous
compact chevron pairs of the anthocodial points, and (2) the
quadrangular “ stars.”
Locaurry :—Station 315.
12. Dendronephthya orientalis (Hend.).
Described by Henderson in ‘ Aleyonarians of the Indian
Ocean,’ Part II. pp. 30-32 (1909).
Draenosts :—Divaricate; no regular outline; somewhat flat-
tened ; polyps in loose clusters; polyp stalks long ; supporting
bundle strong ; point spicules 1 pair only, of which one member
is the larger and markedly projects; crown of 3 loose horizontal
rows; grade VI.; spicules: tentacles conspicuously armoured,
in the cortex very conspicuous strong spindles and numerous
small warty triradiates and multiradiates ; colour : general surface
white, anthocodial and supporting-bundle spicules pink.
ANTHOCODIAL GRADE AND ForMULA :—
VI. =1P + 3 Cr + strong S.B.
WITHIN THE GENUS DENDRONEPHTHYA. 61
DeEscRIPTIVE NotTEs :-—
1. Oolony as a whole. Divaricate ; without regular outline ;
somewhat flattened ; with loose clusters of polyps ; agrees generally
with Henderson’s description. 4 colonies in all.
2. Branching. Short.
8 Colour. General surface white, with the spicules of the
anthocodie and supporting bundles coral-pink.
4. Polyp stalks. Long.
5. Polyps in loose clusters of 6 or so. Text-fig. 19.
6. Polyp spicules. The anthocodial
8 points consist of two curved spindles,
one always larger than the other and
projecting markedly beyond the polyp.
The crown shows 3 rather loose hori-
zontal rows, while below these and pro-
jecting outwards there are 8 stronger
spindles approaching in size those of the
points.
The supporting bundle is strongly
developed, and consists of one very pre-
dominant spindle based in numerous
small ones.
7. Other spicules. On the aboral sur-
face of the tentacles the armature is
very conspicuous. Very characteristic D. orientalis Hend.
in the cortex among the strong spindles
are numerous small, irregular, warty spicules, often triradiate or
with irregular rays in different directions. Among them are
minute warty capstans like those figured by Henderson for the
canal-walls.
Locatrry :—Station 125.
Another specimen with the same anthocodial armature, notably
two unequal curved spindles rising in each point from a crown
of 2-3 longitudinal rows, has a regular outline and only a slight
hint of flattening. The spicules are coarsely tuberculate spindles
and numerous irregular warty discs.
The spicules of the anthocodial and supporting bundles are
amber-red and those of the general surface are golden yellow.
Locatity :—Station. East of Polloe Weh.
Ill. UMBELLATA.
13. Dendronephthya collaris (Wr. & Stud.).
Described by Wright & Studer in Rep. Se. Res. ‘ Challenger,’
vol. xxxi. pp. 208-210 (1889).
Draanosis.—Umbellate ; contour irregular ; basal branches
foliaceous and form a collar; polyps arranged in small roundish
62 DR. W. RAE SHERRIFFS ON EVOLUTION
masses: polyp stalks long; supporting bundle very strong ; point
spicules 5-6 with the uppermost pair stronger and projecting ;
no crown; grade LV.; spicules: very abundant spindles on the
polyp stalks, of the polyps not on the collar sigmoid spindles, and
of the collar irregular interlocking quadrangular “ crosses”;
colour: polyps of the foliaeeous collar bright red, rest of the
polyps white.
ANTHOCODIAL GRADE AND ForMULA :—
IV. = 5-6 p + 0 Cr + very strong §.B.
Descriptive Norns :—
1. Colony as a whole. Two colonies of which the larger
measures, as regards the polyparium, 9°5 cm. high, 12 em. broad,
and 5em. in thickness.
2. Branching. Umbellate, referable to Kiikenthal’s section
Umbellate, subsection with the umbels forming large hemi-
spherical masses, and to the group collaris, where the contour is
broken by the clefts between the masses. Quite divergent,
however, in having only a short stalk, but this is probably only
a growth condition related to the nature of the substratum. At
the top of the short stalk the basal branches are foliaceous and
form a collar.
3. Colouring. Polyps of the foliaceous collar are bright red ;
those of the rest of the colony white (not purple as in the
‘Challenger,’ specimen), but some of them show a tinge of red.
4, Polyp stalks are long. "
5. Polyps arranged in small circular
masses.
6. Polyp spicules. The anthocodial
armature shows 8 double rows of 5-6
converging pairs of spindles, and one at_
the top of each row may be much
stronger than its neighbour and project
beyond the polyp. These 8 projecting
spines catch the eye at once.
The supporting bundle is very strong,
with one spindle projecting far beyond
the polyp.
7. Other spicules. The predominant
spicules of the polyps not on the collar _—D. coliaris Wr. & Stud.
are spinulate slightly sigmoid spindles,
the “‘f’s” of the ‘Challenger’ description. The abundant spindles
on the polyp stalks are almost en chevron. The characteristic
spicules of the collar region are irregularly quadrangular bodies
with very rough protuberances which secure interlocking. They
correspond to what are called “crosses” by Wright and Studer.
There are also spindles enlarged at one end into very rough
clubs, curved spindles with very strong protuberances on the
convex side, and irregular bodies like teeth with many roots.
WIULHIN THE GENUS DENDRONEPHTHYA. 63
Locauity :—Station 99.
A second colony, the larger and flattened from side to side,
may also be referred to D. collaris. Here the polyparium is very
markedly transverse oval. There is no hint of any projection
from the anthocodial points (which consists of 5-6 pairs en
chevron), and the lower cortex shows, among other forms,
numerous flattened, spindle-shaped, triradiate, boomerang-like,
and irregular spicules, which have not been noticed in the other
colonies studied. Very marked also are the length and strength
of the prongs arising from some of the curved spindles, pseudo
clubs, and irregular multiradiate bodies.
Depuction :—If one disregards the detail of growth on which
Kikenthal partly bases the group collaris, namely that the
hemispherical bunches are quite superficial, one should place
D. collaris close beside D. habereri, which has more, though
similar, spicules in its anthocodial points and a different kind of
spiculation on the cortex of the sterile stalk.
14. Dendronephthya longicaulis (Kiik.).
Described by Kikenthal in ‘ Versuch einer Revision der
Alcyonarien,’ li. pp. 633-634 (1905).
Dracnosis :— Umbellate; very long stalked; somewhat flattened
polyparium ; basal branches foliaceous and forma collar ; polyps
in small, loose umbellate groups; polyp stalks usually short, but
may reach 2 mm. long; supporting bundle weak; point spicules
one pair only, one of which is stronger; crown of 4-6; grade V1.;
spicules: cortex of the branches covered with stout spindles
bluntly tuberculate, cortex of the stalk with many stellate forms ;
colour: general cortex has the surface pink-white, poly p-stalks
red-brown, anthocodial spicules almost translucent.
ANTHOCODIAL GRADE AND KorRMULA :—
VI.=141P + 4-6 Cr + weak S.B.
DescriptivE Notes :—
1. Colony as @ whole. A single, peculiar specimen unlike any
other in the collection; marked by a long flexible stalk (10 cm.
out of a total of 13); of uniform diameter (1°3cm.); and bearing
a rather loose umbellate polyparium, somewhat flattened.
2. Branching. The basal branches of the polyparium are
foliaceous and form a collar turned downwards. There are but
few branches extending transversely, so that the top of the
polyparium appears rather truncate.
3. Colouring. General surface of the cortex is pinkish white,
the polyp stalks reddish brown, while the anthocodial spicules
are almost translucent.
4. Polyp stalks are on an average about 1 mm. long, but a
length of 2 mm. may be attained. They appear to be bare on
the ventral side.
5, Polyps. The bundles of polyps are disposed in small loose
64 DR. W. RAE SHERRIFFS ON EVOLUTION
umbellate groups. The upper part of the stalk, however, as
Kiikenthal describes, shows an incrustation of minute polyps.
6. Polyp spicules. The anthocodial
armature conforms with Kikenthal’s Text-fig. 21.
figure, but I should describe it some- a.
what differently, for it seems to me to Bee.
have in each true point only one pair, a Ss
one member of which is distinctly larger
than the other, all the other (4—6) spi-
cules being referable to the crown. BES.
Kiikenthal describes two closely opposed as
spicules forming the upper part of the = We
point, below 1-2 pairs of horizontals and Y \
below that again 3-4 converging pairs. =
On my view, however, each point con- _—
sists of a pair of spicules,
The supporting bundle is very short, D. longicaulis Kiik.
scarcely projecting.
7. Other spicules. The cortex of the branches is closely covered
with strong spindles of typical shape and blunt tubercles—
a feature in which this specimen does not conform to Kiikenthal’s
description of D. longicaulis, where “the branch-cortex contains
separate longitudinally disposed very narrow spindles with sharp
thorns up to 2 mm. in length.”
On the other hand, the cortex of the stalk shows a close array
of stars as Kiikenthal describes, and with the same average
diameter of 0°-1 mm. Among the stars are a few broad, short,
densely tuberculate. spindles.
Locauiry :—Station 310.
15. Dendronephthya disciformis (Kiik.).
Described by Kiikenthal in ‘ Versuch einer Revision der
Alcyonarien,’ ii. pp. 636-638 (1905).
Diagnosis :—Umbellate; compact; disc-like ; outline uniform ;
polyps in bundles of 6-10; polyp stalks medium; supporting
bundle very strong; point spicules about 5 pairs, with the upper-
most larger spicule predominating and projecting ; no crown;
grade IV.; spicules: in the upper cortex long, curved spindles
(2mm.), in the lower cortex stout spindles, irregular stellate
bodies for interlocking, and large triradiates and quadriradiates ;
colour: generally pink and orange, supporting bundles red.
ANTHOCODIAL GRADE AND ForRMULA:—
IV.=14+4P+40 Cr + very strong 8.B.
DrscriIPrivE Nores :—
1. Colony as a whole. Six specimens. Hach a compact colony
of agenerally uniform outline, with its umbels arranged in larger
hemispherical masses which are superficially in contact, forming
WITHIN THE GENUS DENDRONEPHTHYA. 65
a uniform outline. Below the surface are large spaces between
the main branches, and some of these are tenanted by Decapod
Crustaceans about 1:5 em. in length. The colony is slightly
dise-shaped, and has a very short, broad sterile stalk.
2. Branching. From the uniform outline and the arrange-
ment of the umbels in large hemispherical masses, it should
evidently be referred to Kiikenthal’s section Umbellate, first
division, group discifornus.
3. Colouring. Generally of a pink and orange colour; the
supporting bundle red.
4, Polyp stalks not over 1 mm. long. Text-fig. 22.
5. Polyps in small bundles of 6-10.
6. Polyp spicules. In the anthocodial
armature there are 4-5 pairs of con-
verging spindles en chevron in each of
the 8 points, and above these a much
larger converging spindle rising almost
0-5 mm. beyond the polyp. In rare
cases two spindles share in the pro-
jection, which is very characteristic.
Below the points there is a vague crown,
and there are 2-4 small intermediates
between the points. All these spicules
are whitish. pn
In the supporting bundle about three ;
very strong dark red densely spinose D. disciformis Kiik.
curved spindles project beyond the an-
thocodiz, one being stronger than the others, and these are
based in smaller but still strong spindles. The largest spindle
measures 4 mm. in length, and shows a distinctly smooth tip
(cf. D. gigantea).
7. Other spicules. The upper cortex is densely covered with
long curved spindles, mostly in longitudinal arrangement, many
of them 2 mm. in length. Some are red, some yellowish, while
others are colourless. Only some of them show the distant, fine
spines that Kiikenthal describes; most are covered somewhat
densely.
The lower cortex shows (a) stout spindles with long conical
tubercles ; (b) very numerous small irregular stars with longish
prongs, which may again be divided and are suited for inter-
locking; (c) large triradiates and quadriradiates covered with
strong tubercles, sometimes massive and compound. A typical
triradiate is about 1 mm. in maximum length, which is much
larger than Kiikenthal notes. But, of course, they occur in
many sizes.
SALIENT FEATURES.—We should say that this species is best
marked by (1) the strongly projecting 8 spindles at the tops of
the points, (2) the strong red supporting bundle with its longest
spindle smooth at the tip, and (3) the compact disc-like mode of
growth in the full-grown colonies. It should also be noted that
Proc. Zoon. Soc.—1922, No. V. 5
66 DR. W. RAE SHERRIFFS ON EVOLU'TION
there is considerable variety in the anthocodial points, especially
with regard to the relation between the two terminal or the
single terminal and the two or three rows immediately below.
Locauiries :—Stations 99, 164.
16. Dendronephthya habereri (Kiik.).
Described by Kiikenthal in ‘Versuch einer Revision der
Alecyonarien,’ ii. pp. 638-641 (1905).
Draagnosis :—Umbellate ; compact; stiff; almost uniform con-
tour; polyparizm very brittle; lowest branches foliaceous ;
polyps in small groups; polyp stalks very short; supporting
bundle strong ; point spicules over 7 pairs, short, not projecting ;
no crown; grade II.; spicules: lower cortex has very thick,
rough spindles, triradiates, quadriradiates, and irregular bodies,
all with compound tubercles suited for interlocking; colour:
polyps yellowish, anthocedial armature opal-white, supporting
bundle and upper cortex spicules pale pink, others whitish and
colourless.
ANTHOCODIAL GRADE AND FORMULA :—
Il.=7+ p+ 0Cr + strong 8.B.
Descriptive Nores :-—
1. Colony as a whole. Hight colonies; umbellate; of very
compact build, stiff; with a grouping of the umbels in hemi-
spherical masses which, however, are so close together that an
almost uniform contour results. The polyparium is 45cm. in
height, 7 em. in width, and 4:5 cm. in thickness, in one instance,
and is very brittle.
2. Branching. Nearer inspection shows that the colony is
composed of about a dozen hemispherical masses rising from a
short, broad, flattened stalk, with few short, thick branches, the
lowest of which are foliaceous. It should therefore be referred
to Kiikenthal’s group A, division 2, and to the disciformis group,
where the polyparium is flattened and has an oval shape with the
largest axis horizontal (“‘queroval”). Within this group the
specimens agree with D. habereri. _ Although considerable por-
tions of the short, broad stem and branches are bare, these are
enly seen on close examination, for the umbels, which all come to
the surface, are practically continuous. The sterile stalk has its
long axis in line with that of the polyparium.
3. Colouring. The polyps are yellowish, the anthocodial arma-
ture opal-white, the spicules of the supporting bundle and upper
cortex are pale pink; the others are whitish and colourless.
4, Polyp stalks very short.
5. Polyps yellowish, in small groups.
6. Polyp spicules. The anthocodial armature consists of at
least 7 pairs of converging short spindles in each point. These
point spicules, which may not project, are about 0°2 mm in length,
WILHIN THE GENUS DENDRONEPHTHYA. 67
and are densely covered with minute blunt cones. There is no
distinct evidence of a crown.
The supporting bundle is strongly de- Text-fig. 25.
veloped, and includes numerous curved
spindles of various sizes up to 3-5 mm. Pe
in length. 3 or 4 spindles, often about ae di
3 mm. long, make the apex of a bundle,
and one is stronger than the others AN
and projects beyond the polyp for over A); WS
05mm. The other spindles of the sup- Zr.
porting bundle are very much smaller. Y
All show a very dense covering of high,
blunt cones.
7. Other spicules. The lower cortex is
densely packed with very thick, rough
spindles, triradiates, quadriradiates, irre-
gular bodies, and small, almost globular
forms, with one particularly strong boss D. haberers Weak,
and numerous jagged points. The com-
pound tubercles of all these spicules are well suited for inter-
locking. Some of the spindles are irregularly expanded at one
end, and might be called club-like in a loose usage of that term.
Locaurry :—Station 258.
One specimen shows an undamaged sterile stalk, which has its
long diameter in the same plane as the slightly flattened poly-
parium. The stalk is bright crimson, and shows very markedly
a multitude of irregular, globular, slightly stellate small spicules
which might perhaps be called “‘ astroscleres.”
Some of the anthocodial points show 8-10 rather irregularly
converging pairs of rough spindles. There is no definite crown.
The degree to which the supporting bundle projects varies con-
siderably in different parts of the colony.
LocaLiry :—Station 282.
Another colony, marked by a very short sterile stalk, a trans-
verse oval mode of growth and little indication of foliate basal
branches, shows an interesting colour variation which catches the
eye at once. The 8 anthocodial points consist of 7 or more
converging pairs of short spindles, but the upper half of each
point has bright red spicules, while the lower half and the inter-
mediates are white. In most of the polyps this contrast is very
pronounced, but in some cases the red spicules predominate, anc
in other cases there is a slight mixture of the two colours.
Locauiry :—Station not marked.
5
68 DR. W. RAE SHERRIFEFS ON EVOLUTION
17. Dendronephthya pumilio (Studer).
Described by Studer in Ann. Mag. Nat. Hist. (6) vol. 1. pp. 70—
71 (1888).
Dracnosis:—Umbellate; branches in rounded bunches closely
packed; outline uniform; general shape flat to globose; polyps
on the terminal twigs; polyp stalks long; supporting bundle
very strong; point spicules about 5 pairs, the uppermost
elongated ; no crown; grade IV.; spicules: lower cortex has
thick, bent, thorny spindles but no distinct stars; colour:
generally yellowish white, polyps dark red, the tentacles densely
crowded with minute rod-like reddish spicules.
ANTHOCODIAL GRADE AND FoRMULA :—
IV.=1+4p4 0 Cr +4 very strong 8.B.
Descriptive Noves :-—
1. Colony as a whole. Four interesting specimens at different
stages of growth, the largest having the following dimensions :—
Total height 16 cm., height of sterile stalk 6-5 cm., breadth of
polyparium 9°5em. The colony is very markedly umbellate in
its largest forms, but less so in the younger forms, which tend
to be much flattened.
2. Branching. ‘The branches form rounded bunches, closely
packed together and forming in the larger colonies a markedly
uniform contour. The general shape varies from flat to globose,
and I do not think that any importance can be attached to
this.
3. Colouring. Yellow white, polyps ieee ar 6),
dark red. See capa
4, Polyp stalks long. re,
5. Polyps arranged on the terminal oa
twigs. a a
6. Polyp spicules. The anthocodial
armature is marked by the elongation of
one or both of the terminal spicules of 1
each point, the others, to the number
of 2 or 3 pairs, being markedly sub- GN
ordinate. This ismy grade 1V. There
is no crown. NE
The tentacles are long and heavily
armoured, bearing dorsally a crowd,
sometimes bilinear, of minute oval or D. pumilio Stud.
vod-like reddish spicules.
The supporting bundle is very strong, the largest spindle often
projecting for 1mm, ‘The spicules are finely and thickly spinose,
and often show smooth points. In many cases one polyp and its
supporting bundle may be seen to predominate markedly over
the others in the group.
7. Other spicules. ‘The spicules of the lower cortex are mostly
thick spindles, usually bent, and densely covered with strong,
WITHIN THE GENUS DENDRONEPHTHYA, 69
blunt thorns, often compound. There are also numerous irregular
bodies, but no distinct stars were seen.
Depuction :—I agree with Kiikenthal in regarding D. pumilio
as near to J. spinosa if not identical with it.
Localities :—Stations 99, 258, 315.
18. Dendronephthya coronata (Wr. & Stud.).
Described by Wright & Studer in Rep. Sc.- Res. ‘ Challenger,’
vol. xxxi. pp. 212-213 (1889).
Draenosis :—Umbellate; polyps not grouped in small hemi-
spherical heads; outline uniform; polyps im small loose bundles
of 3-5; polyp stalks medium; supporting bundle very strong ;
point spicules 3-4, the upper pair markedly projecting ; crown of
three rows of roughly horizontal spindles; grade V.; spicules:
cortex of the stem shows stout, thick spindles strongly tuberculate,
also triradiates, quadiradiates, and irregular forms; colour :
generally whitish, polyp spicules clear amber-yellow, supporting
bundles strong purplish red.
ANTHOCODIAL GRADE AND FoRMULA :—
V.=3-4p+3 Cr + very strong 8.B.
DeEscrIPTIvE NorTes :—
1. Colony as « whole. Three colonies with polyps in distinct
umbels, but not grouped in small hemispherical heads and forming
as a whole a fairly uniform outline, agree in general with
D. coronata, especially as revised by Kiikenthal. They belong to
his florida group of Umbellate, being slightly flattened in a lon-
gitudinal oval and showing a convex upper surface to the umbels.
2. Branching. The branches of the stem lead to the formation
of large hemispherical masses of polyps.
3. Colouring. The general colour of Text-fig. 25.
the colonies is whitish, tending at the :
base to become red. One of the speci-
mens showed a slightly different colour-
scheme, having whitish instead of amber
polyp spicules, but the same tendency to
purplish red.
4. Polyp stalks never attain a length
of 2 mm.
5. Polyps in small, loose bundles of
3-5.
6. Polyp spicules are clear amber-
yellow. The anthocodial armature shows
3-4 pairs of curved, thorny spindles _D. coronata Wr. & Stud.
en chevron, the upper pair markedly pro-
jecting over the base of the tentacles, below these is a crown of
several rows of roughly horizontal spindles, and below these
again are larger purplish spicules forming par5 of the supporting
bundle.
70 DR. W. RAE SHERRIFFS ON EVOLUTION
This supporting bundle is very strong, and one spindle over
3mm. in length projects for a distance of a little over 1mm.
beyond the polyp head; it is flanked basally on each side by
one or more almost equally strong, and lateral to these are smaller
spicules. All the spicules of the supporting bundle are strong
purplish red.
One specimen showed frequently the occurrence of one project-
ing spicule at the tip of the points, or of two spicules markedly
unequal. A search revealed, however, especially in the collar
region, the typically equal pair characteristic of the species.
7. Other spicules. The cortex of the stem shows stout, thick
spindles with strong tubercles, many markedly compound, trun-
cated, and along with these there are triradiates and quadri-
radiates besides smaller exceedingly irregular warty bodies, often
like the broken heads of clubs.
Locauiry :—Station not marked.
19. Dendronephthya brevirama (Burcharat).
Described by Burchardt in ‘Aleyonaceen von Thursday-Island
und von Amboina’ in Semon, Forschungsreisen, V.5; pp. 438-459
(1898).
Draenosis:—Umbellate; umbels not combining in hemi-
spherical masses; contour almost continuous; flattened in one
plane; surface of the individual umbels varies from convex to
coneave ; lowest branches slightly foliaceous and form an inter-
rupted collar; polyps in small bundles; polyp stalks short;
supporting bundle strong; point spicules 6-8: converging pairs of
small spicules, none projecting; no crown; grade II.; spicules:
cortex of branches shows long bright red spindles spinulose, cortex
of stalk has coarsely tuberculate broad spindles, triradiates, and
irregular discs: colour: general surface yellow-white, polyps
yellow, branches with strong purplish spiculation.
ANTHOCODIAL GRADE AND ForRMULA :-—
Il. = 6-8 p + 0Cr + strong 8.B.
DESCRIPTIVE NOTES :—
1. Colony as a whole. Six colonies, of which the largest measured
19cm. in height by 18cm. in width. The umbels do not combine
in hemispherical masses; the contour of the polyparium is almost
continuous; there is a considerable flattening in one plane; the
surface of the individual umbels varies from convex to econcave—
a point to which I cannot attach importance. The specimen
should be referred to Kiikenthal’s florida group and therein to
within the species D. brevirama.
2. Branching. The lowest branches are slightly foliaceous, and
form an interrupted collar.
3. Colouring. The general surface is yellow-white, the polyps
yellow, the branches showing strong purplish spiculation. -
“I
=
WITHIN THE GENUS DENDRONEPHTHYA.
4. Polyp stalks ave short, a little less than 1 mm.
5. Polyps in small bundles.
6. Polyp spicules. The anthocodial Text-fig. 26.
armature of 8 points shows 6-8 con- Ps
verging pairs of small spicules, none oUt\y ss
projecting, and no very definite crown, Y \ .
The supporting bundle has one strong 4
spindle up to 3 mm. in length, pre-
dominating and projecting for about
0°75 mm.
7. Other spicules. The cortex of the
branches shows numerous long, straight, Ar
or curved bright red spindles, covered WN
with short, rather delicate, sharply
conical spinules, particularly crowded at
the ends. The cortex of the stalk region
shows (a) numerous coarsely tubercu- LD breesranin Bact
late broad spindles, straight or curved ;
(6) asymmetrical bent spindles, with the tubercles stronger on
‘the convex side; (¢) spindles with one end broadened out ;
d) irregular tuberculate discs, some almost spherical, others
almost stellate; (e) regular and irregular triradiates (some very
massive).
The canal-walls in the stalk region show numerous strong
triradiates and curiously irregular quadriradiates, all roughly
tuberculated, while besides these are minute irregularly radiate
forms inclining to be smooth.
Locatrry :—Station not marked.
A much smaller younger colony, of which the polyparium
measures 4 em. by 4°6 cm., is worthy of notice because of certain
slight divergences. The umbellate character of the twigs is
much less pronounced, the whole branching is looser, the upper-
most pair of spindles in each point sometimes projects to a very
slight extent. These do’not, however, differ from the others in
the way characteristic of D. coronata, where the projection
beyond the polyp is, of course, very marked. The specimen
suggests that young stages of members of the Umbellate may
present a somewhat Divaricate appearance.
On the other hand, the collection includes another colony, of
which the polyparium measures 4:7 em. by 4:7 em. in maximum
height and breadth, which is interesting in showing the very
opposite vegetative characters; for it is markedly Umbellate
in the branching of the twigs and extremely dense in the general
branching, so presenting a uniform compact surface. Yet the
spiculation and the anthocodial armature are identical. In the
72 DR. W. RAE SHERRIFFS ON EVOLUTION
recesses of the polyparium there lay a large Ophiuroid, with its
arms extending in different directions and raising puzzling ques-
tions as to entrance and exit; for the whole surface of the
polyparium is like a thick-set hedge without any gaps.
Locatiry :—Station 164.
Still another colony, at first sight divergent, agrees so
thoroughly in anthocodial architecture and spiculation that
separation appears to me impossible. The somewhat looser
group of umbels show a marked tendency to grouping in hemi-
spherical bunches, especially on one side of the colony; the
contour is somewhat broken up; the lowest branches are slightly
foliaceous; the sterile stalk has been broken off, leaving only a
short stump; the general colouring is yellow-white, with white
spindles in the polyps and faint rose tips to many of the sup-
porting bundles. Yet an examination of the anthocodie shows
the characteristic composition of the points, about 7 pairs of
converging spicules, of which the uppermost pair may be slightly
larger but do not project. Again, the lower cortex contains
massive spindles with high compound warts, a few massive
triradiates, many rough spindles curved at one end like a simple
bugle, many pseudo-clubs, and very numerous small knobbed and
bossed forms like astrospheres. In the canal-wall there are among
rough types a number of characteristic, almost smooth, delicate,
flat triradiates, quadriradiates, and approximate stars.
Depucrion: — We have here another illustration of the
inadvisability of attaching much importance to the vegetative
characters.
In anthocodial architecture this species comes near D. florida,
but the shortness of the polyp stalks is too emphatic to be
neglected.
Locatrry :—Station 258.
20. Dendronephthya annectens (sp. n.). (Plate III.)
Dracnosis :— Umbellate; much branched, but few large
branches; distinctly flattened; contour not uniform; basal
branches foliaceous; polyps in small bundles of 7-12; polyp
stalks very short; supporting bundle very weak; point spicules
about 8 pairs, equal in size; crown absent; grade I1.; spicules:
short sterile stalk has irregularly branched, almost stellate,
bodies; spindles of the branches often very large (6mm.);
colourless spindles in the general cortex and stars at the base;
colour: exposed areas on the branches white; twigs have large
orange and crimson spindles, supporting bundles either orange or
crimson, polyp spicules white.
ANTHOCODIAL GRADE AND KoRMULA :—
Il. =8 p+ 0 Cr + very weak 8.B.
WITHIN THE GENUS DENDRONEPHTHYA. 73
Descriptive Nores :—
1. Colony as a whole. Very handsome umbellate colony with
profuse root-work; hardly visible sterile stalk; much branched
and, in its general aspect, distinctly flattened polyparium. It is
32 cm. in maximum height and 40 cm. in maximum breadth,
with an approximate thickness of. 6em. The contour of the
whole colony is markedly interrupted.
2. Branching. There is a general division into tiree strong
branches each of which tends to lose the flattened aspect,
broadening out into an exuberant corymbose head. The polyp-
bearing twigs are distinct umbels. There are no definite large
hemispherical bunches. Large areas on the branches are bare
and exposed. The basal branches are foliaceous, but do not form
a complete circle. The main branches give off numerous second-
ary and tertiary branches, which eventually break up in fairly
regular dichotomy into minor corymbs which bring the crowded
polyps more or less up to one level.
3. Colouring. The exposed areas on the branches are covered
with irregularly scattered spindles, visible to the naked eye and
giving the surface a glistening white appearance. On the twigs
are large orange and crimson spindles, standing out conspicu-
ously against the general white of the cortex. There seems to be
no regularity in the distribution of the two colours, except that
any one group of twigs is either orange or crimson. The
supporting bundles are also either orange or crimson, but the
polyp spicules are white.
Text-fig. 27. Text-fig. 28.
LZ //
D. annectens, sp. n. D. annectens, sp. n., ensheathing type of
supporting bundle: seen from below.
4. Polyp stalks are very short.
5. Polyps. The polyps themselves occur in smail bundles of
7-12, and the stalk of each bundle is long.
6. Polyp spicules. The anthocodial armature consists of about
8 pairs of spicules en chevron on each of the eight points, all
practically of the same size, about 0-13mm. Below the point
74 DR. W. RAE SHERRIFFS ON EVOLUTION
there is no true crown, but there are small white spindles on the
soft polyp stalk, lying irregularly but on the whole transversely
and quite distinct from the coloured spindles of the supporting
sheath which lie longitudinally. It is plain that the uppermost
rows of these transverse spindles might furnish the raw materials
of a crown.
The supporting bundle is of the ensheathing type and is often
very inconspicuous. The typical form is a curved triangle
around the polyp stalk, composed altogether of about a dozen
spicules, and those composing the tip are not conspicuously longer
than the rest. It is what might be called a slightly differentiated
supporting sheath rather than a supporting bundle, and its
comparatively small spindles must not be mixed up with the
large supports of the common stalk of the common bundle. The
supporting bundle only occasionally projects a little beyond
the polyp. its component spicules do not attain a length of
more than 0°75 mm,
7. Other spicules. Very noteworthy is the spiculation of the
short, sterile stalk, where for a very limited area the spindles are
replaced by irregularly branched, almost stellate, bodies, which
interlock and give the surface an arenaceous appearance. Many
of the branchlet spindles are striking in their size, reaching a
length of 6mm. They are densely covered with truncated
columnar tubercles which form regular curved transverse rows,
recalling the septa of an elongated /ungia coral. The truncated
tops are very rough and sometimes compound.
The colourless spindles of the general cortex are partly like
small editions of those just mentioned, and partly narrower forms
with distant and relatively few tubercles. They range from
1-2-5>mm. The small “stellate” forms at the base are very
irregular in shape. Many are almost globular; many are minute
quadriradiates; many show one predominant boss among the
radiating knobs ; while many again correspond exactly with those
figured by Kiikenthal for D. japonica (fig. Lb) *.
Very few canal-wall spicules could be found. Those that
occurred very sparingly were narrow rodlets with a few distinct
tubercles.
Depuction :— Although this splendid specimen has no uniform
contour it presents many resemblances to D. brevirama (Burch.),
such as :
(1) the anthocodial armature of 6-8 spicules in each point, the
uppermost not projecting ;
(2) the short polyp stalk ;
3) the numerous 2 mm. spindles of the general eortex ;
(4) the crowded stellate bodies of the lower cortex ;
(5) in the polyp-bearing twigs a corymbose - dichotomous
branching, almost identical with that of specimens which
agree in detail with D. brevirama.
* Kiikenthal, “Versuch einer Revision der Alcynarien.” Zool. Jahrbch. Jena.
System. vol. xxi. p. 576.
WITHIN THE GENUS DENDRONEPHTHYA. 75
_ Yet this specimen differs deeply from D. brevirama in the
following respects :
(a) it has a different type of supporting bundle, namely a
sheath, instead of showing one markedly predominating
needle ;
(6) it has practically no spicules in the canal-walls ;
(c) it has not more than ‘a hint of triradiates.
It therefore seems best to establish a new species, closely like
D. brevirama (an Umbellate) in some respects and very like
D, japonica (a Divaricate) in others.
Locaurry :—Station not marked.
21. Dendronephthya simplex (sp. n.).
Draenosis :—Umbellate; compact; slightly flattened umbels
not combined into large hemispherical masses; lower branches
foliaceous ; contour uniform ; polyps compact; polyp stalks short ;
supporting bundle strong; point spicules, 2, long converging
only; crown of some 3 rows of curved spindles; grade VI.;
spicules: cortex has stout tuberculate spindles but no stars;
colour: generally drab-yellow.
ANTHOCODIAL GRADE AND KoRMULA :—
VI. = 1 or2 P+ 3 Cr + strong S.B.
Descriptive NOTES :—-
1. Colony asa whole. The entire colony is 9 em. high, of which
2'5 em. go to the sterile stalk. The breadth of the polyparium
is6mm. The polyp-bearing twigs form small umbels, but these
do not combine in larger hemispherical masses. The specimen
should therefore be referred to [Ciken-
thal’s second group of the Umbellatz exif. 29:
and to the florida group, though the S
shape of the polyparium is rather cir-
cular than long oval. There is a uniform,
very slightly interrupted contour to the
whole colony, which has a somewhat
flattened circular shape. The general
appearance is compact. Practically the
whole surface is covered by the polyps
on their umbels, and no polyps occur
except on the surface.
2. Branching. Some of the lower
branches are foliaceous.
3. Colouring. On the whole the
colour is drab-yellow. D. simplex, sp. 0.
4. Polyp stalks. Short.
5. Polyps in compact bundles of 10 or so, dense corymbs.
6. Polyp spicules. The anthocodial armature consists of 8
points of 2 long converging spindles each, and it may be said at
76 DR. W. RAE SHERRIFFS OF EVOLUTION
once that it is almost precisely that of D. piitteri (Divaricate)
and not far from D. lutea (Umbellate, but with interrupted
contour). The length of the point spicules is about 0°75 mm.,
but one is often slightly smaller than the other. Between the
points there lies a pair of much smaller intermediates. At the
base of the points there is a crown of about 3 rows of curved
spindles, but some of these are often displaced so as to appear
like the lower point spicules in D. lutea, in regard to which
Kikenthal notes that an approach to a horizontal position
sometimes occurs. This dislocation of what is really a horizontal
band suggests the need of examining numerous polyps.
There is a strongly developed supporting bundle, the largest
spicule of which, about 2 mm. in length, projects beyond the
polyp for about 0°75 mm. It is abutted by two strong spindles
on each side and by a number of smaller ones transitional to the
polyp. The supporting-bundle spicules are slightly curved
spindles, densely covered with relatively minute blunt and narrow
cones. The spindles of the 8 points are similar, but show the
characteristic hockey-club-like bend at the base.
7. Other spicules. All over the cortex there is a felting of
more irregular and stouter spindles, often “‘f” shaped, with
larger and often compound blunt tubercles. No stellate forms
occur, but very numerous, quite irregular types are found which
might be derived from spindles and club-like ends or from
boomerangs. Their compound warts are often very strong and
stronger on one side.
Locaity :—Station not marked.
22. Dendronephthya hyalina (Kiik.).
Described by Kiikenthal in ‘Versuch einer Revision der
Aleyonarien,’ ii. pp. 688-689 (1905).
D1acnosis:— Umbellate; flaccid; hyaline; umbels not in
large hemispherical groups; outline not uniform; distinct foli-
aceous collar; polyps loosely aggregated ; polyp stalks medium;
supporting bundle strong; point spicules 4—5 pairs, without pro-
jection of the top pair; crown absent; grade III.; spicules: sterile
stalk has numerous stellate and multiradiate forms; colour:
cortex very hyaline, with loosely disposed white spicules.
ANTHOCODIAL GRADE AND ForMULA :—
III. = 4-5 p+ 0 Cr 4 strong 8.B.
Descriptive Noves :—
1. Colony as a whole. Eight young colonies, the largest of
which measures 6°5 cm. high. They are very flaccid in texture
and hyaline in appearance. ‘The youngest forms show little hint
of umbellate branching, but this is well marked in the biggest.
As the umbels do not compose large hemispherical groups, the
WITHIN THE GENUS DENDRONEPHTHYA. ve
specimens should be referred to section B of Kiikenthal’s Umbel-
late, and the presence of unequally long branches resulting in no
very uniform outline points to the spinulosa group.
2. Branching. There is a ‘distinct
foliaceous collar. Text-fig. 30.
3. Colouring. Cortex very hyaline
spicules.
4, Polyp stalks. Slightly over 1 mm.
long.
d. Polyps. Few are in good con- \S
dition, owing to the colonies having ZX
been much compressed in most case
6. Polyp spicules. The anthocodial ———
points consist of 4-5 pairs en chevron,
with almost no projection of the upper-
most pair and almost no crown. D. hyalina Kik.
developed, and the strongest spicule may project for half a
millimetre.
7. Other spicules. The sterile stalk shows numerous stellate
and multiradiate spicules.
and bearing loosely disposed white
small groups of 3 or 4.
The supporting bundle is fairly well
Locaniries :—Stations 257, 318,321. Saleyer, 10-25 fms.
EXPLANATION OF THE PLATES.
Prare I. Dendronephthya clavata Kiikenthal.
Prate II. Dendronephthya gigantea Verrill.
PuateE III. Dendronephthya annectens, sp. n.
Obes Meth
MeMiy As Wao
. cilia
edrr lated
ore . a ta
ee
i dopeonayg me
aus vier a
2) Ra, en ih a
awh
i M
aes ee aur
Peed! bik ie oye
a
ru es Diy ty bned An ey
hs ee Ath
hee aa
+
Nae a) ahs EBs \ se eis
1922. Balfour-Browne. PI. [.
ll I2
Life history of the
WATER BEETLE (PELOBIUS TARDUS) HERBST.
VITTY & SEABORNE, LTD.,
LONDON.
P.Z.S. 1922. Balfour-Browne. PI. Il.
| milinelic.
Life history of the
WATER BEETLE (PELOBIUS TARDUS) HERBST.
VITTY & SEABORNE, LTD.,
LONDON,
‘ d r
a Se em
P.Z.S. 1922. Balfour-Browne. PI. Ill.
Life history of the
WATER BEETLE (PELOBIUS TARDUS) HERBST
VITTY & SEABORNE, LTD.,
LONDON,
THE LIFE-HISTORY OF THE WATER-BEETLE. 74)
The Life-History of the Water-Beetle Pelobius tardus
Herbst. By Frank Batrour-Browns, M.A. (Oxon. et
Cantab.), F.R.S.H., F.Z.8., F.E.S., Lecturer in Zoology
(Entomology) in the University of Cambridge.
Or
Received January 10, 1922: Read March 7, 1922.]
(Plates I.-ITT.)
Sunumary.
The paper has been paragraphed under the following
headings :—
1. The family Pelobiide. p. 79.
2. Britannic distribution of P. tardus. p. 79.
3. The Imago.
(a) Habitat and Habits. p. 82.
(6) Longevity in artificial environment. p. 83.
(c) Stridulation. p. 83.
(d) Sexual differences. p. 84.
4, The Life-history.
(a) Oviposition. p. 84.
(6) Incubation. p. 85.
(c) Vital staining of Embryo. p. 85.
(d) The Larva. p. 87.
(e) Food of the Larva. p. 89.
(f) Stomodeum of the Larva. p. 90.
(g) Habits of the full-grown Larva. p. 92.
. The Life-cycle. p. 95.
On
(1) The Family Pelobiide.
The family Pelobiidz contains the single genus Pelobiuws and
only four species, one of which is European, one Chinese, and
two Australian, The family holds a somewhat isolated position,
being related to the Carabide and showing similarities of struc-
ture with Dytiscide and other aquatic beetles of the series
Adephaga, these being due, according to Sharp (10)*, not to
close relationship but to adaptation to life in the water.
(2) Britannic Distribution.
The single Huropean species is not uncommon in Britain, but
belongs to the Southern or “ English” type, as defined by
Watson (16), being chiefly found in the Southern and Eastern
counties of England. It has been recorded from one Scottish,
* The numbers refer to the bibliography at the end of the paper.
80 MR. F. BALFOUR-BROWNE ON THE LIFE-HISTORY
three Irish, and thirty, out of the seventy, English county and
vice-county divisions.
The only Scottish record, for Midlothian, occurs in Duncan’s
Catalogue (3), where the species is described as “ very scarce,
Pond at Coates” on the authority of Sir P. Walker. ‘The same
record is repeated in the ‘ Entomologia Edinensis’ (17), but neither
Andrew Murray (9) nor Sharp (11) refers to it.
For the North of England there are two isolated records for the
species, one for Northumberland South and the other for Cum-
berland. The former is given by Stephens (15) on the authority
of G. Wailes: ‘‘ Once near Newcastle by Mr. Hewitson.” Bold’s
comment that this is ‘‘ probably erroneous as it has never since
been met with” (4) is quite valueless. The Cumberland record is
bare, being a note by James McDougall (8), who mentions having
taken the species at Wigton, and although I have made enquiries
T can find out nothing further about it.
In the case of many species, records for localities far beyond
the normal range are frequently regarded as due to errors in
identification, but Pelobius is not a beetle likely to be confounded
with any other, so that, unless a mistake has been made in
labelling, records for this species may be regarded as correct,
and are probably to be accounted for on the assumption that
individuals have, from time to time, wandered northwards. In
such a manner might the species extend its range, but in this
case the attempts to extend northward seem to have failed up to
the present.
The extreme northern range limit seems to be Yorks S8.E.,
where the species is ‘‘ fairly common in the stagnant pools near
Withernsea,” according to H. W. Baker of Hull(4). The records
for Yorks S.W., Lanes 8., and Derby may or may not be for
single specimens in each case. The Yorks $.W. record appears
in the Victoria County History (2) as “ Doncaster (J. Wilcock).”
The Lanes. record is an old one given by W. EH. Sharp (12) as
“ Rufford (Gregson),” though, for some unexplained reason, this
record is omitted in Sharp’s ievisseltists of 1906 (18) and 1908 (14).
The fact that Jahn (6) mentions “ two specimens in a pool near
Cheadle ” seems to suggest that here again we are within the
normal range of the species; and although records even south of
this are scarcee—in many cases only one in a county or vice-
county,—there is little doubt but that the species is fairly
common in the eastern and southern parts of the country. The
accompanying ‘“typo-map” will give a general idea of the
distribution of the species, and the letters representing the
county and vice-county names will be sufficiently intelligible, as
they are more or less in correct geographical position.
Pelobius is scarcely to be described as established in Ireland,
since, in the case of both Cork Mid. (7) and Clare (7), only one
specimen was found, and possibly the Wicklow record, ‘near
Lara,” in the Haliday MS. (5) also refers to a single specimen. I
have worked many of the Irish counties fairly thoroughly, and
OF THE WATER-BEETLE PELOBIUS TARDUS. 81
ok !
TYPOMAP SHOWING BRITANNIC DISTRIBUTION OF PELOBIUS TARDUS Herbst.
SI
or oe
NS CA
HB Ss
RW RE EL BF AN
WI El PN AS
AM PM FF KI
DN SG PC KF
CT RE LE ED HD
AY LA PE BW NN
WD ED LD AN WT KB DF SK RX SN
FE TY AR DO CU WL NY DM
WM SL LE MO Me ML MY EY
EM RO CV LH SE MVS
WONG LE WH WE ° GR DB FT CH DY NM LS
SG KC KD DU MN MG SP ST LR CB WN EN
CL NT Qc OW WI CD RA HF WO WW NO HU WSES
NK LK ST KK WX PB CM BR GE OX BX BD HT NE
SK MC EC WA GM MMGWNW BK MX SE
wc NS SW NH SR WK EK
EG. SD IW
SC WC
. ; Seaha F :
Underlined letters indicate County and vice-County from which Pelobius has been recorded.
Bi.=Berks. CB.=Cambridge. CL.=Clare. CU.=Cumberland. DT.=Dorset. DY.=Derby.
ED. (Scot.)=Edinburgh. EK.=EHast Kent. EN.=East Norfolk. ES.=BHast Suffoll.
EX.=East Sussex. GM.=Glamorgan. HF .= Hereford. IW.=1. of Wight.
LN.=Lincoln, North. LR.=Leicester + Rutland. MC.=Mid Cork. MX.=Middlesex.
NE.=North Essex. NH.=North Hants. NS. (Eng.)=North Somerset. SE.=South Essex,
SH.=South Hants. SL. (Eng.)=South Lanes. SN.=South Northumberland. S.=Surrey.
ST. (Eng.)=Stafford. SY.=South-Hast Yorks. ‘WI. (Ive.)= Wicklow. WK.=West Reo
WN.=West Norfolk. WO.=Worcester. WW.=Warwick. WY.=South-West Yorks.
Proc. Zoou. Soc.—1922, No. VI. 6
82 MR. F. BALFOUR-BROWNE ON THE LIFE-HISTORY
although in some of these, e.g. Wexford, there were many ideal
ponds, and although I was specially looking for this species, I
never found a specimen. I therefore regard the Irish individuals
as having strayed from the English breeding centres, the species
so far having failed to establish itself west of the Irish Sea.
(8) The Imago. (a) Habitat and Habits.
Pelobius is a pond species, never, in my experience, occurring
in drains, lakes, or running water, but it seems to be very par-
ticular as to the type of pond it frequents. It is apparently only
found in ponds where the bottom is covered with a fine ‘ ooze,”
so that gravel- or marl-pits are ideal habitats. In some cases the
water of such ponds is clear and bright, while in others the pond
is a favourite haunt of cattle during hot weather, the water then
becoming thick and turbid and containing a high percentage of
ammonia. The cattle pond is quite as thickly populated as the
clean pond, so that it is the “ooze” which seems to be the im-
portant characteristic of the habitat.
In this ‘‘ ooze” the beetle frequently buries itself, so that only
the apex of the body, often with a protruding air-bubble, is
visible, and, during the summer, it may remain thus buried for
even more than thirty minutes at a time, coming up rapidly to
the surface to renew its air-supply, which takes from five to ten
seconds, and then once more disappearing in the mud.
By keeping the beetles in tumblers of water half filled with
pond mud, I have found that at irregular intervals the air-bubble
protruding from the apex of the body is released, and another
then appears in its place and gradually increases in size. At first
I was satisfied to explain this on the supposition that these were
bubbles of exhausted air, but I can find no statement as to the
mechanism of respiration in the Water-Beetle, or as to how the
sub-elytral air is utilized; and I am not now inclined to speculate
on the subject beyond saying that possibly the released bubbles
ave merely due to the expansion of the air in the sub-elytral air-
space, due either to the body temperature of the insect or to the
temperature or pressure of the mud.
The food of the imago consists of insect larva and worms, and
while I was working at the life-history I kept my specimens in
tumblers, feeding them mainly upon chopped-up earthworm, which
they took readily. Agrionid dragonfly nymphs, Chironomus
larve, Sialis larvee, and various other forms, were readily
devoured, but more active types, such as Chloeon, could usually
keep out of the way.
The mouth-parts are typically mandibulate, and in nature the
insect hunts for its food, swimming over the surface of the ground
and poking into hollows and under stones. It seems to have a
good sense of smell, since, in swimming over the surface of the
mud, it will suddenly check and dive downwards; and, in such
circumstances, if the beetle is dug up, it usuaily has some food
OF THE WATER-BEETLE PELOBIUS TARDUS. 83
between its jaws. Also, in my tumblers, the beetles immediately
became very active when a piece of worm was quietly dropped in,
even when they could not see it, and they would at once begin to
hunt round until they found it.
3 (b) Longevity of imago in artificial environment.
In some of my tumblers I kept pairs, in. others single indi-
viduals, each tumbler being supplied with a smali piece of
water-weed, usually ZVodea, and also with a few pebbles to give
foothold. The water was changed daily or on alternate days,
according to circumstances, and I had no difficulty in keeping
the beetles alive for months and even for more than a year in
this way. Deaths that occurred were almost always among the
pairs, and the male succumbed more frequently than the female.
Presumably the cause of death in these cases was overcrowding,
six or seven ounces of water, even when changed daily, not being
sufticient for two individuals.
[ had previously experienced this difficulty in keeping several
individuals of a species in a confined space. For instance,
Hydrobius fuscipes L., a smaller beetle than Pelobius, lived much.
longer when one was kept in each tumbler than when two or
three were sokept. Of twelve individuals of such a small species
as Bidessus ninutissimus Germ. only six survived after a month,
two after three months, and one shortly after that, the last
survivor living for many months. At the same time six indi-
viduals of this same species, each in a separate tumbler, survived
for periods varying from one to three years, the water in these
tumblers never being changed, only a little bemg added from
time to time to make up loss due to evaporation.
T have mentioned that, in the case of Pelobiws, when one pair
was confined in a tumbler the male usually died first, which
suggests that the male is not so hardy as thefemale. Experiments
showed that the female lives longer than the male. As I was
unable to find the details of about a dozen experiments carried
out in 1913—before the work described in this paper was com-
menced—I set aside four tumblers in May 1920, each containing
much decaying leaf material rich in Chironomus larvee and various
other forms of life. In each tumbler I placed one Pelobiws, using
for the experiment two males and two females. A small piece
of Elodea was placed in each tumbler to keep the water fresh, and
each tumbler was covered with a piece of glass to reduce evapo-
ration. Nothing was done to these tumblers until May 1921,
by which time, out of the six ounces of water originally placed
in each of them, only about three remained. The two females
were, alive but both the males were dead.
3 (c) Stridulation.
Pelobius used to be sold in St. Martin’s Lane, London, under
the name of the ‘“Squeak beetle,” owing to its being able to
§*
84 MR. F, BALFOUR-BROWNE ON THE LIFE-HISTORY
make a loud, strident noise, which it does by rubbing the apex of
the abdomen. against the underside of the apices of Sue elytra, in
which position is a file-like structure (vide Pl. FIT. fig. 6). Both
sexes are alike inthis. The beetle ‘‘ squeaks ” when it is alarmed
or annoyed; for instance, when it is caught in the water-net or
held between finger and thumb. In my tumblers, if one
individual tried to seize a piece of worm upon which another
was feeding, the latter “squeaked,” the squeak in this case
presumably being equivalent to the growl of a dog with a bone.
3 (d) Sexual differences.
There is very little difference in outward appearance between
the sexes, the larger tarsal pads of the male being about the only
one. The female is slightly larger than the male on the average
and perhaps slightly broader in ‘proportion to length.
(4) Life-history.
The beetle is to be found in its habitat from March until
October. In the latter month it descends into the mud in the
bottom of the pond to hibernate, apparently remaining there
without moving, its legs and antenne folded in against the body,
until the following spring. Early in March, or possibly some-
times late in February, it reappears and towards the end of
March oviposition commences.
4A (a) Oviposition.
In the labovatory my earliest date for the appearance of eggs
is March 23rd, but under natural conditions eggs are very scarce
until about the second week of April, when they become common.
Whereas Dytiseid eggs are buried in the tissues of plants, the
females possessing piercing ovipositors, Pelobius eggs are laid
upon the surface of sub-aquatic vegetation. In my tumblers and
tubs the eggs were laid in rows, end to end, sometimes as many
as eight to ten in a row, though odd eggs were also to be found.
When about to oviposit the female lies along the plant stem,
clasping it with her legs so that the apex of her abdomen touches
the stem. The egg, mon is blunt oval in form, 15 mm. long
by -87 mm. broad and equally curved at both ends is very clear
at the moment of extrusion and enclosed in a thin gelatinous
envelope which at once adheres to the plant. This envelope
quickly absorbs water and swells up into a thick ‘protective
covering.
Having laid an ege the beetle moves slowly forward, waving
her antenne in the water and usually touching the stem with
her palpi. She moves just far enough to permit the next egg to
emerge so that it will touch the end of the previous egg, ane in
this way she works along the stem, producing a string of eggs.
OF THE WATER-BEETLE PELOBIUS TARDUS. 85
I thought at first that the beetle judged her distance by means
of the palpi feeling their way, but, as I have mentioned, these
organs do not invariably keep contact with the stem, while the
gonapophyses do, so that. it appears as if these latter decide how
far forward the beetle is to move after depositing an egg.
I have not been able to determine the total number cf eggs
laid by a female, the greatest number of which I have a record
being twenty ; neither do I know whether an individual lays all
her eggs during one short period or whether she lays a number
of batches.
The period of oviposition for the species is a long one, lasting
from March until July. I have not actually found eggs later
than June 15th, but I have found very young larve in August
under rather interesting circumstances.
On August 5th, 1915, in a disused chalk pit at Beckham in
East Norfolk, where there were several shallow clear-water
ponds, I found various stages of the beetle. There were old
males and females, newly-emerged soft males and females, full-
grown larve and a few larvee not more than ten days or a
fortnight old. As incubation in the warmest period of the year
occupies only nine days, the eggs from which these larvee hatched
must have been laid in July.
4 (b) Incubation.
The earliest laid eggs took twenty-five days to hatch, but the
incubation period gradually became shorter until, in June, nine
or ten days became normal. Eggs laid early in April and
placed in an incubator at summer temperature hatched in nine
or ten days, so that temperature is evidently the controlling
factor in the length of the incubation period. J shall return to
this point later on.
So far as 1 can find, there is no special hatching apparatus
such as thatin the larva of Dytiscus (1), the shell ripping from end
to end along an irregular line and apparently always along the
ventral side of the embryo, however it may happen to be lying
in the egg. A small pair of spines are certainly visible on the
head between the eyes, near the position of the hatching spines
of Dytiscus lapponicus, but they are not functional, as is evident
from the manner in which the shell rips, and I imagine that, if
anything weakens the latter along the breaking line, it must be
the tarsal claws.
4 (c) Vital Staining of the Embryo.
In order to observe the developing embryo I removed many of
the eggs from the water-plants upon which they had been laid
and kept them upon cotton-wool saturated with water. I have
used this method with the eggs of several other water insects—
e.g., Agvionid dragonflies and others, Ranatra linearis, and
86 MR. F. BALFOUR-BROWNE ON THE LIFE-HISTORY
various water-beetles both Hydradephaga and Hydrophilide ; and
as a rule they do very well, but occasionally an egg shows signs
of being unhealthy, that is, the surface becomes the home of
sitter hens of minute roneran and Alege. Such an egg if left to
itself invariably dies, but I discovered a way of treating it which
usually saved it, and this was by saturating the cotton- eae with
a solution of methylene blue. Any moderate strength seemed to
be suitable, and the methylene blue killed off the Protozoa or Alge
without injuring the egg. But the effect of the methylene blue
was not confined to destroying the Protozoa and Algz on the
ege-shell. It did not in any way injuriously affect the developing
embryo, but certain parts of the living tissues took up the stain
and stood out bright blue. The staining varied with different
individuals, with the same individual at different times and
possibly also with different strengths of methylene blue, but in all
eases the embryonic appendages of the first abdominal segment,
which are well developed in this species just as they ave in
Dytiscus lapponicus, took up the stain (v. Pl. I. figs. 9,10, & 11).
Otherwise the staining was very irregular. In the case of one
larva just ready to hatch, the dorsal pharyngeal muscles and the -
malpighian tubules were stained, and on the left side of each
thoracic segment and of the first two abdominal segments a short
line, curved in the thoracic, straight in the abdominal segments,
was also stained. At one end this line came to the surface of the
embryo and suggested a spiracle, although there is no pro-thoracic
spiracle in the larva and the spots were above the real meso- and
meta-thoracic spiracles which were also stained, Incidentally it
may be mentioned here that none of the spiracles are functional
in the larva. In the abdomen of this specimen parts of the
dorsal longitudinal muscle system under the terga were stained,
but not evenly, certain strands having taken the stain, others
not having taken it. In the alimentary canal was some greenish
fluid which I took to be methylene blue changed by the action of
the secretions.
I give this example merely to show the strange effect of
methylene blue, and without making any attempt to explain it,
T assume that the parts which stain are in some way active at, the
time they stain. For instance, possibly the one-sided staining
of the dorsai longitudinal muscle system in the specimen referred
to may have indicated that those muscle-strands were doing work
while the corresponding strands upon the other side were doing
none. If this suggestion is sound it mee perhaps be possible,
by watching any embryo develop in an egg kept in methylene
blue, to recognize various parts functioning a different times.
The staining of the first abdominal appen \dages, which disappear
after hatching, is also of some interest if this suggestion is correct.
These appendages first appear when the embryo is about three
days old and after the three pairs of thoracic appendages, but
they are more distinct than any of the following abdominal
appendages which rapidly disappear, and they persist, as more
OF THE WATER-BEETLE PELOBIUS TARDUS, 87
or less globular structures with a deep cup-like pit in them, right
up to the time of the escape of the larva from the egg, and can be
seen, especially after the staining referred to, after the larva has
hatched, disappearing as it expands.
I had somewhat similar experiences as to the staining powers
of methylene blue in the case of the embryos of Dytiscus lapponicus,
and I used to run a kind of hospital of methylene-blue tumblers
for the eggs. But in the case of that species [ found another use
for methylene blue. When rearing the larve 1 found that I
could not keep them alive in tumblers, one in each, unless the
water was so charged with the stain that it looked like ink, in
which case the larvee grew up quite successfully. This solution
was changed every two or three days, but in no case was I suc-
cessful with the larvee in tumblers of plain water, even when I
changed the water daily. Possibly the explanation in this case
is that the methylene blue oxidized the fecal products and
remnants of food of the larve and made them innocuous.
4(d) The Larva.
The larva, on hatching, is colourless and possesses a long
medio-dorsal spine ou the last abdominal segment and two
lateral cerei. Within the egg these structures are doubled
beneath the embryo, the last abdominal segments occupying the
end of the egg. On escaping from the egg, the larva at once
straightens out, and these appendages are at first about equal to
the body-length, though before the end of the first stage the body
has increased in length while the appendages remain the same.
Within an hour the chitin hardens and the pigment has
appeared and darkened. .The main colour of the larva is
brownish green, but there is a pattern of pale yellow, and this
varies in different individuals, some of which might be described
as almost black with a few light markings, sale others are pale
yellowish with a few dark markings. The colouring apparently
does not vary much during the life of the individual—that is,
one which is dark in the first stage will be dark in its final stage.
An infinite number of diagrams could be given to show the
range from dark to light, but I have selected five specimens, two
at the end of their first stage (Nos. 1 and 2) and three in their
third stage (Nos. 3, 4, and 5) to show the range (vide Pl. I.
figs. 1 to 5).
A study of these diagrams will make it clear that, although
variations occur, the disappearance of the dark pigment in the
lighter individuals is not a phenomenon which may happen at
any spot on the surface of a segment, but occurs in more or less
definite areas; and this is seen even better by comparing a
number of individuals of about the same degree of darkness or
lightness when a rather surprising uniformity of pattern is
recognizable.
Where i is, I think, some very interesting work to be done upon
88 MR. Fe BALFOUR-BROWNE ON THE LIFE-HISTORY
the variations in colour-pattern within a species*. I have only
paid slight attention to the subject, and only in the case of water-
beetles. In many of the Hydradephaga one finds a dark ground-
colour marked with yellow lines or spots, and there is usually
considerable variation to be found within a species, some indi-
viduals showing more yellow and others being almost without
it. Such species as dgabus abbreviatus ¥., A. didymus Ol., and
Platambus maculatus L. are most interesting studies, while in
such a genus as Veronectes four out of the five Britannic species
give excellent material for research.
To return to Pelobius. In the newly-hatched larva the head is
more or less triangular in shape and the prothorax is but little
narrower than the head, the body thereafter tapering away to
the Jast abdominal segment, which is only about one-fifth the
width of the head,
The larva is heavier than water and breathes subaquatically
by means of a series of filamentous gills attached at the sides of
the sterna of the thoracic and first three abdominal segments.
At the inside of the base of each of the front legs is a tuft of
three gill-filaments. Four rather larger filaments lie inside the
bases of the middle legs, and the two posterior legs have each a
tuft of three gills situated as in the case of the anterior legs.
The abdominal gills consist each of a pair of filaments arising
from a common base. and one of these is attached on each side to
the posterior edge of the sternite, those of the first abdominal
segment lying posterior to the pair of embryonic appendages
already referred to.
The gills are tracheate, with a single unbranched tube running
the length of each filament.
The legs are long and delicate. The coxa is large, the
trochanter consists of two small segments, and the femur, tibia,
and tarsus are each of one segment, the tarsus bearing a pair of
claws almost as long as the segment. The tarsus in each case,
but more especially of the second and third pairs of legs, is
strongly feathered, and therefore forms an efficient swimming
organ; but it has also another function. Gilled insect nymphs
and larve, resting in stagnant water, usually adopt some means
of cireulating the water about the respiratory apparatus.
Agrionid dragonfly nymphs will frequently lash the abdomen
from side to side; the nymphs of the Ephemerid, Chlocon, flap
the lamellate gills ranged along the sides of the abdomen, and the
larvee of eertain Perlids “dance” up and down, raising and
lowering the body between the legs. The larvee of Pelobius use
their feathered legs as fans, the middle leg on one side and the
anterior and posterior legs on the other being raised off the
* Vide Brown, Annetta F., “ Evolution of Colour-pattern in Lithocolletis.”
Journ. Acad. Nat. Sci. Philadelphia, xvi. (1914) pp. 105-165 ; and Palmer, Miriam 4.,
“Some Notes on Heredity in the Coccinellid genus Adalia Mulsant.” Ann. Ent.
Soc. America, iv. (1911) pp. 283-808.
OF THE WATER-BEETLE PELOBIUS TARDUS. 89
support and rapidly vibrated so that a current of water passes
under the body and bathes the gills.
The first stage of larval life occupies from six to eleven days,
seven or eight days being apparently the usual period, and the
larva then moults.
There is little difference in general appearance between first
and second stage larvee except that the head is more rounded
and less triangular in shape, and the three “ tail” processes are
somewhat shorter in proportion to the body-length, but some
change has taken place in the number of gill-filaments. Those
of the prothorax remain as in the first stage, but there are now
six filaments round the base of each of the middle legs and five
round the base of each of the hind legs. The only change in the
abdominal gills is the appearance of an additional pair of very
small ones on the first segment, one gill towards either side of
the sternite.
This stage lasts from nine to thirteen days, when the larva
once more moults, reaching its third and final stage.
In the third stage again the body-length increases more than
that of the “ tail’’ processes, which are now only about one-third
the body-length, and the gills are also more complex in that each
filament shows several transverse constrictions, as 1f an attempt
had been made to produce segmented filaments such as are seen
in the larva of Sialis. As to numbers of filaments, no change
takes place on the pro- or meso-thoracic segments, but an
additional pair appears on the metathorax, one usually very
‘short gill appearing anterior to and outside each. of the posterior
legs. The abdominal gills do not again increase in number, but
remain as in the second stage. A diagram showing the arrange-
ment of the gills will be found on PI. IIT. fig. 4.
In all stages of the larva nine pairs of spiracles are to be
found, two thoracic and seven abdominal, but, as has been
mentioned previously, these all remain closed and functionless
throughout the larval period.
As a rule it was quite easy to determine when a larva had
moulted, because the cast skin would be visible in the tumbler or,
‘if the moult had just taken place, the larva would be white,
though the pigment appears very quickly after the ecdysis; but
individual larve possessed idiosyncrasies, some always retiring
into the mud just before moulting and not appearing again until
their colours had been restored. JI have records of quite a
number of larvee, amongst my earlier batches, which apparently
erew up without moulting at all, and it was not until I discovered
skins amongst the mud at the bottom of the tumblers that I
realized what had happened.
4 (e) The Food of the Larva.
I had considerable difficulty at first in keeping the larve alive.
I adopted the same method as that used in the case of the
90 MR. F. BALFOUR-BROWNE ON THE LIFE-HISTORY
Agrionid dragonfly, Hydrobius fuscipes and Dytiscus, keeping
records of individuals, each in a separate tumbler, but I could
find no food which they would eat. The newly-hatched Pelobius
larva is only about 2 zm. long in body-length, so that minute
food was obviously necessary, but Paramecium, upon which I
fed the newly-hateched dragonfly nymphs, had no attraction for
them, and minute alge, Cladocera, and Copepoda were equally
useless. I noticed, when I placed a larva in a tumbler with a
layer of mud at the bottom, that this mud was continually
examined and even burrowed into, which suggested that the
necessary food was some mud-inhabiting species. As, however,
these larvee also died, I made a journey to one of the ponds where
the beetle occurred and brought home a supply of mud from
there, and, on using this in the tumblers, it was at once obvious
that the food-problem was solved. This mud was swarming with
Tubifex, and it was most interesting to watch the larva hunting
for and capturing its prey. It would move slowly over the
surface, touching it with its antenne and palpi, and, on dis-
covering a Zubifex burrow, the larva would either at once leap
into it, sometimes disappearing except for the apices of its
“tails,” or it would sit and watch, like a cat at a mouse-hole, its
action presumably depending upon the position of the worm in
the burrow. Sometimes the snatch at the prey would fail and
the larva would move elsewhere, but with these first-stage larvee
it was difficult to decide whether the pounce had been successful,
as it was rare for. the larva to appear from the burrow with the
worm in its mouth. With the larger larve one could see the
struggle, the worm trying to withdraw to the bottom of the burrow,
and the larva, as it were, sitting back upon its haunches and
pulling with all its might to get the worm out. In the case of
these larger larvee, however, the feeding was sometimes done in
the burrow.
The feeding process is rather peculiar; the larva sits motionless
with most of the worm projecting from its mouth and wriggling
furiously, the larva to all appearance doing nothing; suddenly,
however, the latter makes a gulp, and a little of the projecting
part of the worm disappears into the mouth. After a short
interval another gulp takes place, and thus by a series of gulps
the worm disappears down the “throat.” The larva apparently
always seizes the worm by the end, never by the middle.
So far as I could find, Zubifex is the only food of the larval
Pelobius, as, although I tried various other materials and could
get the larve to eat Chironomus larve, I never succeeded in
rearing one on any diet other than the little red worm.
4 (f) The Stomodeum of the Larva.
The guping down of the worm struck me as rather extra-
ordinary. There was no chewing or biting, and yet an examination
of the contents of the alimentary canal always showed chewed-up
OF THE WATER-BEETLE PELOBIUS TARDUS, 91
feod material. An examination of the stomodeum, however,
explained the mystery.
The mouth-parts consist of a pair of sharply-pointed mandibles
which overlap when closed, the right overlying the left. The
maxille are palp-like in appearance, the maxilla itself forming
the basal part of the palp, and, when at rest, the whole structure
is contracted by being telescoped within the base (v. Pl. IL.
figs. 1 to 4). The labium has an apical piece, which, from its
appearance, | will call the “scoop ” (s), and which, when at rest,
is contracted within the basal piece which bears the two short
labial palpi (v, Pl. II. figs. 1 to 4).
When the floor of the mouth is examined (v. Pl. II. fig. 1),
it will be seen that at the base of the extensible part of the labium
is a small, projecting piece in the median line. This piece, which
I will call the “ flap” (/), rises up in the “scoop,” and when the
mouth is closed this flap stands up, and the lobes of the epi-
pharynx—this structure being split like a hair-lip—descend
from above and thus block the entrance. The anterior part
of the floor of the mouth is covered with minute projections
arranged more or less in rows radiating from a deep groove,
while ‘the anterior part of the scoop is similarly studded. In the
posterior region of the floor of the mouth is a dark brown
chitinous plate (hyp) with a raised median piece posterior to it,
and J regard these structures as representing the hypopharynx.
On either side of the chitinous plate there projects inwards a
membranous lobe (mal) which is extremely difficult to make
out; L examined a large number of specimens before I could
satisfy myself that it really existed. Possibly these lobes
represent the maxillule.
Immediately behind the hypopharynx the mouth contracts into
the gullet, which is very short and opens out at once into the
cesophagus (v. Pl. I. fig.6). This again is very short, and there
is no pro-ventriculus clearly marked off at its posterior end, but
this region is armed with eight dark-coloured spines (v. Pl. I.
figs. 5 & 6), which project forward in the walls and posteriorly
are enlarged. The inner faces of these spines are armed with
strong, backwardly turned teeth which are specially numerous on
the enlarged bases, and evidently form the grinding-mill by means
of which the food is broken up.
With the exception of a pair on each side whose bases are
fused together, each spine is independent of the others. The
median ventral one is the longest, and extends forwards for about
two-thirds the length of the cesophagus, none of the other spines
extending more than half the length of the ceesophagus.
To the esophageal walls, between the pro-ventricular spines,
are attached muscles which run to the head capsule, so that the
cesophagus is capable of considerable expansion and contraction.
In this case, therefore, almost the whole stomodeum acts as a
sucking-pump, there being no pharynx marked off for this purpose,
as is usually the case.
5/3} MR. F. BALFOUR-BROWNE ON THE LIFE-HISTORY
The mechanism of all this complex apparatus seems to be as
follows :—The Pelobius larva pounces upon the Tubifex and, by
expanding the csophagus, sucks the end of the worm into the
stomodeum, and the mandibles then close so that the prey is held
either between the two, or more usually, between the upper and
the epipharynx. I think, therefore, that it usually passes into
the mouth between the two lobes of the epipharynx (v. Pl. I.
figs. 11 & 12).
At the moment of seizing upon the worm the maxille and
labium are shot forward and the prey is received into the “scoop, ’
and these parts then retract, the flap in the “scoop” pressing
the worm upwards into the epipharyngeal slit, and the closing
of the jaws bending the body of the worm upwards so that it
is completely jammed and cannot move backwards or forwards
(v. Pl. I. fig. 12). At the first suck, the end of the worm pre-
sumably reaches well down into the pro-ventricular mill, where
it is well bruised, and, when this portion has been sufficiently
crushed, the hold upon the worm is momentarily relaxed by the
opening of the jaws and the shooting forward of the maxille and
labium ; the sucking action is repeated, and once again the mouth-
parts jam the body until another section has been crushed by
the pro-ventricular mill.
4 (g) Habits of the full-grown Larva.
The larva is full-fed about twenty-two days after the second
moult, by which time it measures about 15 mm. in length, the
median “ tail” adding another 5 mm., and it is now ready to
leave the water. In my tumblers it was always evident when
this stage was reached, because the larva would swim round and
round at the surface and endeavour to climb up the glass.
As I wished to watch the work of the larva in forming its
pupal cell in the earth, I tried, with this species, the same
method as I had used successfully with Dytiscus lapponizus—that
is, | placed the larve in an earth-bottomed vivarium with a glass
side, made an artificial burrow in the earth against the glass, and
carefully directed the steps of the larva to the entrance of the
burrow, closing the mouth of the latter when the larva had
entered. With this species, however, the method did not work
so successfully, as the iarva, having reached the end of the
artificial burrow, usually continued it on its own account and as
a rule in a direction away from the glass. Digging up such larvee
and starting them again seemed to discourage them, and they
were apt to sulk and make no cell at all.
I therefore devised a special glass cell consisting of two Jantern-
slide cover-glasses. Between these two, at adjacent corners, two
small pieces of cork, half an inch thick, were placed so that, along
one side, these two plates could not come into contact. Along the
opposite side the two plates were brought into contact and held
OF THE WATER-BEETLE PELOBIUS TARDUS. 93
together by string or narrow adhesive tape. The cell was formed
between these two glasses, a strip of cotton-wool filling up the
open ends between the two plates and a similar strip being pushed
down into the narrow angle where the two plates came together,
and this cell was then filled up with sand (v, Pl. III. fig. 9).
By keeping the cotton-wool moist by occasional wetting, the sand
was also kept moist and in a suitable condition for the require-
ments of the larva or pupa of Pelobius.
An artificial burrow was then made perpendicularly downwards
in the sand with a pencil, which, of course, after penetrating the
sand for about 2 inches, came into contact on either side with the
glass plate ; and into this burrow a larva was pushed uncere-
moniously, the entrance to the burrow being then closed by
covering the surface of the sand with a narrow strip of glass.
The larva was thus in such a position that whichever way it
worked it was compelled to form its cell in contact with
the glass.
These glass cells were kept in the dark so that the larva should
not be worried by the light, but I found that when once the larva
had accepted the situation, the light did not affect it unless it was
very intense.
Tn starting its burrow, the larva digs itself in head first, and
having reached the depth at which it is going to make its cell,
a depth which varies from 2 inches to at least 5 under different
soil conditions, it moves its head from side to side and, with its
jaws, chews up the pellets of earth, causing the grains to pack
more closely, and in this way it forms a space around itself.
Practically the whole space of the cell is produced by this re-
arrangement of the minute particles of soil, though the entrance
from the tunnel becomes closed during the operation.
At first the larva, in a straight tunnel, is rather confined in its
movements, but, as soon as space permits, it changes its position
and sits up, alternately chewing up the pellets of soil and
pressing them into the walls, using its head asa 1am. The three
“tail ? processes, which one would expect to be very much in the
way in the confined space, become very useful. They are bent
over the back, like a Squirrel’s tail, and wedged against the wall
of the cell, and on them the body turns about freely in all
directions {v. Pl. HI. figs. 1 & 2).
The work of constructing the cell usually took from twelve to
fifteen hours, though sometimes two or three days seemed to be
necessary; but in these cases the larva seemed to work inter-
mittently and without enthusiasm, whereas in most cases great
energy was shown and no rest taken until the cell was completed.
After the completion of the cell the larva rests for from six to
eleven days (v. Pl. Ill. fig. 3) before it changes into a white
pupa of the usual Coleopterous type. The pupal period lasts”
about sixteen days, and the imago, white at first, takes twelve
hours or longer to attain its colours, and remains in the cellfor a
week or more before breaking out.
94 MR. F. BALFOUR-BROWNE ON THE LIFE-HISTORY
(5) The Life-Cycle.
Thus from the hatching of the egg to the appearance of the
imago occupies from nine to about fifteen weeks. On this basis,
the earliest imagines should appear about the middle of June,
and this raises the question whether eggs laid in July and
August may not be the commencement of a second annual
generation. From an examination of the ovaries of a few
beetles in August which emerged in the previous June, I am
inclined to think that there is normally only one generation a
year. Perhaps also the longevity of the beetle, at any rate of
the female, which will live for three years and perhaps longer,
suggests a one-year cycle.
On the other hand, the possibility of a few precocious indivi-
duals breeding soon after emergence must not be lost sight of
(and also the possibility of old females laying a second batch of
eges must be kept in mind), though it does not follow that the
resulting larvee would complete their metamorphosis.
I make this suggestion that a second generation may be
commenced, even if not completed, because of a number of
experiments I made some years ago with a few species of
Hydrophilid beetles. By keeping these in tumblers in an
incubator at a high summer temperature, 19° to 21°C. (66° to
70° F.), I found that it was possible to get egg-cocoons from
some of them (e. g¢., Hydrobius fuscipes L. and Philhydrus mari-
timus Thoms.) in November, after only a month of treatment—
ege-cocoons which, under normal conditions, would not have
been produced until the following April or May. A hot June
and July might therefore cause the ovaries of Pelobius to mature
rapidly, and this would account for the young larve which I
found in August at Beckham.
The life-history of Pelobius is interesting because of the
eill-bearing larva, which does not require to come to the surface
to renew its air-supply. In this respect it differs entirely from
the larve of the Dytiscids and Hydrophilids, and resembles those
of Gyrinids and Haliplids, but like, apparently, all other aquatic
beetles, the pupa requires free air, and therefore, like the majority
of them, the full-grown larva leaves the water and burrows into
the soil, where it forms its pupal cell.
The change on the part of the larva from an aquatic to a
terrestrial habit does not involve a change in the respiratory
apparatus, although I had expected one or more pairs of spiracles
to open during the last larval stage, as happens in the last stage:
of the nymph of the dragonfly. But, whereas the dragonfly
nymph comes from the water into the air with the object of
drying itself, the Pelobiws larva leaves the water and quickly
burrows into the earth, and therefore, presumably, is under
sufficiently damp conditions to allow the gills to function until
the larval skin is thrown off.
OF THE WATER-BEETLE PELOBIUS TARDUS, 95
This work on the life-history of Pelobius was begun in 1915 as
the commencement of a study of subaquatie beetle larve, my
intention then being to work out the life-histories of a Gyrinid
and an Haliplid. The work on Pelobius was almost completed
by the autumn of that year, when military duties took me away
from Cambridge.
Since my return to entomological work, time has not permitted
me to continue on this line of research, and I have therefore
merely completed a number of observations on Pelobius and
written up the life-history of the beetle.
Bibliography.
. Batrour-Browne, F.—‘*‘ The Life-history of a Water-beetle.”
Journal Royal Institution, 1913.
Bayrorp, E. G., and Lawson, THompson M.-—Yorkshire:
Victoria County History. Coleoptera. 1907.
. Duncan, J.—‘* Cat. of the Coieopterous Insects found in the
neighbourhood of Kdinburgh.” Mem. Wernerian Soe. vi.
pp. 443-538. 1826-31.
. Fowxer, W. W.—‘ Pelobins tardus Hbst.in Yorks.” Natura-
list, xvil. p. 234. 1892.
. Hauipay, A. H.—‘‘ MS. Notes on Irish Coleoptera.’’ Publ.
in Proc. Belfast Nat. Field Club, Appendix viii, 1885.
JAHN, L. H.—‘‘ Coleoptera occurring in North Staffordshire.
Prelim. List.” North Staffs. Field Club Ann. Rep. and
Trans, 1904-5, xxxix. p. 73 et seq.
Jounson, F. W., and Hatsert, J. N.—‘A List of the Beetles
of Ireland.” Proc. Roy. Irish Acad. ser. 3, vi. 1901.
. McDoveatt.—Nat. Hist. Journal. York and School Register,
vi. 1880-82.
. Murray, A.—Coleoptera of Scotland. 1853.
. SHarp, D.—‘‘ Dytiscide.” Sci. Trans. Roy. Dublin Soe.
ser. 2,11. 1880-82.
‘“‘ The Coleoptera of Scotland.” Scottish Natura-
list, 1. to iv. 1871-78.
29
. Saarp, W. E.—“ The Hydradephaga of Lancashire and
Cheshire” (Ent. Soc. of Lancs. and
Cheshire), 1892.
Victoria County History: Lancashire.
1906.
“ Coleoptera of Lancashire and Cheshire ”
(Lanes. and Cheshire Ent. Soc. 31st
Ann. Rep. and Proce. for 1907). 1908.
. SrepHens, J. F.—IIl. Brit. Ent., Mand. ii. App. 1829.
. Watson, H. C.—‘‘ Cybele Britannica.” 1847-59. 4 vols.
and Supp. 1. 1860.
. Witson,J,,and Duncan, J.—‘‘ Entomologia Edinensis.” 1844.
96
MR. F. BALFOUR-BROWNE ON THE LIFE-HISTORY
EXPLANATION OF THE PLATES.
Puate I.
Figs. 1-5. Colour-patterns of Pelobius larve. Figs. 1 and 2 are taken from larvae
Fig.
Pig.
Wig.
near the end of their first stage, and Fe. 3, 4, and 5 from larvee in their
third stage. The specimens were merely selected tu show the range of
pattern, there being no noticeable difference in different stages of the
same larva.
Diagram of alimentary canal of a Pelobius larva to show proportionate
lengths of Stomodzeum (8), Proctodeum (P), and mid-gut (MG).
The egg of Pelobius 24 to 36 hours old. The embryo has begun to appear.
Note the large head-lobes, the first signs of the neural canal and the
post. end submer ged in the ‘yolk, which, : at this stage, is not segmented.
. The embryo about 3 days old (in June when incubation lasts only 9 or 10
days), showing the appendages appearing, the antennz between the head-
lobes and the first abdominal appendages, the last of the series at this
particular stage. The yolk is now segmented and the amnion is
completely formed.
. The embryo 4 to 5 days old (in June); the amnion still intact. The first
abdominal appendages very large and distinct. The depressions at the
sides of the thorax and first abdominal segments are, presumably, the
commencements of tracheal-tube formatiun, though the fact that there
is no prothoracic spiracle perhaps makes this doubtful.
The embryo 7 or 8 days old (in June). The eyes have just developed, the
membranes have ruptured, rolled back, and disappeared within the dorsal
region of the embryo. ‘he first abdominal appendages are now smaller. .
Front view of head of a third-stage larva to show the relative positions of
parts. CL=Clypeus. LBR=the labral ridge, beneath which are the
two epipharyngeal lobes, EP. F is the “flap” standing up in the
“scoop” of the labium, LAB. LAB.P=labial palp. Mx=mazxilla,
sunk in its base. Mx.P=Maxillary palp. M=Mentum. GU=Gular
sclerite of head capsule. ANT=Antenna. O=Ocelli. MDB=Mandibles
which are in a position for seizing the prey.
Front view of head as in fig. 11, except that a worm is in the mouth
and the mandibles have closed upon it. This diagram shows how the
worm is bent upwards by the mandibles and thus jammed so that it
cannot move backwards or forwards.
Prate II.
Mouth-parts of a third-stage larva with labrum, epipharynx, and “roof”
of mouth removed to show the relative positions of labium (LAB),
maxilla (Mx), fully extended, and mandibles (Mdb). S=the “scoop” fully
extended and F, the “flap” in the “scoop.” Mxl possibly represents the
maxillule. Hyp, a dark chitinous plate in the floor of the mouth just
where 1t narrows to the “ gullet,” presumably represents the hypopharynx.
Note the four holes in this plate which seem to be always present.
Labium and maxille as in fig. 1, but with the “scoup” fully retracted and
the maxillee almost fully retracted within their bases.
Ventral view of labium and maxilla, showing the “scoop ” fully extended
and one maxilla extended and the other fully retracted within its base.
M=mentum, not clearly marked off from the bases of the maxillz, but
recognizable. G=the gular sclerite.
Ventral view of labium as before, but with the scoop fully retracted. It
will be seen therefore that the scoop telescopes into the rest of the labium
just as the maxille do into their bases. 4
Vigs. 5 & 6. Views of the pro-ventricular spines of a third-stage larva, in situ,
showing the backwardly directed ‘“‘ teeth” and hairs on the inner faces.
Fig. 5 is a view from above and fig. 6 is a lateral view. These two
figures are from: camera lucida drawings, and the millimetie scale
indicates their magnification.
OF THE WATER-BEETLE PELOBIUS TARDUS. 97
Prats III,
Figs. 1 & 2 are from sketches of the positions adopted by the larva at work when
constructing its cell.
‘ig. 3 shows the position of the larva resting in the completed cell, waiting to
change into a pupa,
i?)
19
Fic. 4. Ventral view of a third-stage larva, showing the gills on the left side of the
thoracic and right side of the abdominal segments, the others having
beer omitted for the sake of clearness. This diagram is based upon a
camera lucida drawing, Cc=Coxal cavity. Sp=the closed meso- and
meta-thoracic spiracles.
Fig. 5. A diagram of one of the glass cells used to enable the work of cell con-
struction by the larva to be watched.
Fig. 6. Under side of the right elytron of a male specimen of the beetle to show the
file, upon which the apex of the abdomen rubs to produce the “ squeak.’’
There is a file on each elytron, and the files are alike in the two sexes.
Proc. Zoou. Soc.—1922, No VII i
. Brie as
i on a er « ’ i sae WEL
oS isc Fo cenig ia
& Efe hi
iba
ON THE VAGUS NERVES OF THE EDENTATA G9
6. On the Vagus and Sympathetic Nerves of the Hdentata.
By Caarues F. Sonntag, M.D., F.Z.S., Anatomist to
the Society.
[Received December 28, 1921: Read February 21, 1922. |
(Text-figures 1-5.)
In a former paper (2) I showed that the cervical parts of the
vagus and sympathetic nerves are fused to form vago-sympathetic
cords (type 1), or are simply linked together by communicating
branches (type 2). And these forms are distributed in the
Edentata as follows :—
Type 1 :—Bradypus tridactylus.
Type 2 :—Huphractus villosus and Tatusia novemeincta.
In Tanandua tetradactyla both forms are present in the neck
of the same animal, the first occurring on the right side and the
second on the left.
In all species the course of the vagi from the foramen lacerum
posticum to the point where they reach the sides of the cesophagus
in the posterior part of the thorax is similar to that in the
Marsupialia (1,2). The relations differ, however, for the branches
of the aortic arch are more frequently similar to those in Homo.
Consequently the relations resemble those described in Anthro-
potomy. The branches also differ in many respects from those
in the Marsupialia.
Tamandua tetradactyla (text-figs. 1 and 2).
The vagus nerves communicate with the glosso-pharyngeal,
spinal accessory and hypoglossal nerves by very fine filaments, but
no branches run to the sympathetic in the anterior part of the
neck. And neither vagus has any trace of the ganglion nodosum
in the neck, but they are flatter and wider in the foramen lacerum
posticum. The usual branches of distribution are given off, but
these have a longer course, and are more easily studied than in
most Mammals. They form thick bands, with the spinal accessory
_ nerves in the lacerate foramina.
The pharyngeal nerve (a) is the largest cervical branch. It
runs along the side of the pharynx and wsophagus and supplies
both. It forms a plexus with branches of the glosso-pharyngeal
nerve (ix), but no branches of the sympathetic are seen running
into it. On the right side a branch (/.6.v) of the hypoglossal
nerve (xii) communicates with the pharyngeal branch ; and this
nerve may correspond to what is termed the “ lingual branch of
the vagus” in human anatomy (8).
The superior laryngeal nerve (6) divides into three branches.
One gains the interior of the larynx through the thyro-hyoid
100 DR. C. F. SONNTAG ON THE VAGUS AND
interval, and the others end in the extrinsic laryngeal muscles.
It does not communicate with the sympathetic.
Text-figure 1.
The cervical and anterior thoracic parts of the vagus and sympathetic nerves in
Tamandua tetradactyla. Sg.b.: branch of hypoglossal nerve to sterno-
glossus; V-S: vago-sympathetic cord. Other letters in text. The
nerves have been separated for demonstration purposes so that the super-
ficial and deep cardiac plexuses appear in line.
The right recurrent nerve (d) has the usual origin, course, and
relations. It communicates freely with the sympathetic ( S) and
the superficial cardiac plexus, and is brought into communication
SYMPATHELIC NERVES OF HH BDENTATA. 101
with the le/t recurrent nerve (e) thereby. The latter also com-
municates wth the lower cardiac branch of the left vagus. But
no direct branch runs between the two recurrent nerves.
The Cardiac Nerves (jf) :—The left vagus gives two fine nerves
to the superficial cardiac plexus (S.C.P), wherein they communi-
cate with the cardiac branch from the inferior cervical ganglion
of the left sympathetic. But no ganglia are found in connection
with them. The right vagus gives two large and two fine filaments
to the deep cardiac plexus (D.C.P), and fine filaments to the
superficial one. Reddish-brown ganglia are connected to the
nerves. ‘There are no upper or middle cervical cardiac nerves.
The pulmonary nerves (g) come from the vagus ou the left
side, and from the vagus and deep cardiac plexus on the right.
In the posterior part of the thorax branches are given off to the
esophagus and aorta, and the esophageal plexus (O.P) is a link
between the vagi, for there are no direct branches connecting
them. And the main cords run through the diaphragm on the
sides of the cesophagus.
Abdominal Parts of the Vagi:—The left vagus runs along the
ventral surface of the stomach close to the lesser curvature.
The right one crosses behind the cesophagus and breaks up into
a number of filaments. These reunite and the vagus ends in the
right part of the cceliac plexus. Several gastric nerves (j) run
to the dorsal aspeet of the stomach, and communicating twigs
run to the hepatic (7) and splenic (£) plexuses.
f did not observe any communications between the vagus and
phrenic nerves; but fibres may get from the vagus into the
phrenic sympathetic plexus and be carried up to the diaphragm
to anastomose with the phrenic nerve.
The Cervical Sympathetic :—The superior and middle cervical
ganglia are absent, but the inferior cervical ganglia (1.C.G) are
large and round. They give off the cardiac nerves (c.b.s),
thymic branches (¢.0), subclavian (S.V.B) and phrenic commu-
nicating (¢.p.n) nerves. The left cardiac nerve ends in the
superficial cardiac plexus. The right cardiac branch ends in the
deep plexus. The right sympathetic communicates with the
superficial cardiac plexus and left recurrent nerve.
The Cardiac Plexus surrounds the aorta and its branches,
supplies them and the heart, and gives twigs to the right
pulmonary plexuses. It consists of superficial (S.C.P) and deep
(D.C.P) parts. The former lies on the ventral surface of the
aortic arch. It receives branches from both inferior cervical
ganglia, two branches from the left vagus, and twigs from both
recurrent nerves. The latter lies between the aortic arch and
innominate artery ventrally, and the trachea dorsally, and it
contains three reddish-brown ganglia (C.P.G). It receives
fibres from the right inferior cervical ganglion, and four branches
from the right vagus. Branches are given off to the trachea (¢.p),
heart, and right lung. Both plexuses communicate by fine
nerves.
102 DR. C. F. SONNTAG ON THE VAGUS AND
The thoracic cords have few ganglia. The first left ganglion
(G.S) is long, and connected by the Annulus of Vieussens (A.V)
to the inferior cervical ganglion. When ganglia are present
they lie on the necks of the ribs. The left cord is tortuous
and diminishes in thickness from before backwards; it united
Text-figure 2.
The posterior thoracic and abdominal parts of the vagus and sympathetic nerves
in Tamandua tetradactyla (A), and the fused vagus and spinal accessory
nerves in the lacerate foramina (B). The vagi have been drawn to the
right so as to extend the cceliac plexus. Letters in text.
posteriorly by two filaments with a branch running up to the
aorta at the upper end of the descending part. The right
sympathetic is larger than the left one, and is not connected to
SYMPATHETIC NERVES OF THE EDENTATA. 103
any ascending aortic branch. In addition to the communicating
nerves to the intercostals, and filaments to the esophagus, sorta
and mediastinum, splanchnic nerves are given off. .
The Solar or Celiac Plexus (text- tig. 2) consists of several
ganglia and communicating bundles of fibres, and is divisible
into upper and lower parts. It is connected to the sympathetic
cords by splanchnic nerves (4); these are three in number, but
the lowest is really a bundle of fibres. It lies between the layers
of the mesentery, along the course of the cceliac axis, and is not
set across the vertebral “column, The following are the branches
of distribution :—
A. From the upper part of the plexus.
1. Hepatic plexus (i) which communicates with the gastric
branches of the right vagus (7) by nerves meeting in a small
ganglion.
2. Splenic plexus (k) communicating with the right vagus 2a
supplying spleen and pancreas.
3. Phrenic plexus (/) communicating with the phrenic nerve in
the diaphragm.
B. From the lower part of the plexus.
1. Right renal plexus (mn).
2. Communicating branches to the splanchnic nerves and aortic
plexus (1).
The Aortic Plexus (p) accompanies the aorta and its branches.
It communicates with the splanchnic nerves, sympathetic cords
(RS and LS), and their common ganglion. And it gives off the
left renal plexus (0).
The abdominal parts of the sympathetic have few ganglia. They
gradually diminish in size and meet in a common ganglion in
front of the sacrum. From the latter nerve plexuses run to the
pelvic viscera.
Huphractus villosus (text-figs. 3 and 4).
The vagus and sympathetic nerves differ in many ways from
those in Tamandua.
The Vagus Nerves:—The ganglion nodosum (G.N) is present
on both sides. And the branches of communication to the other
cranial nerves and the superior cervical ganglion of the sympa-
thetic are well marked. In the middle of the neck a thick
communicating branch connects the right vagus and sympathetic.
The pharyngeal branch (a) arises above he ganglion nodosum,
and no nerve connects it to the hypoglossal nerve.
The superior laryngeal nerve (6) is a thick branch arising orn
the centre of the ganglion nodosum. It divides into a thick
internal laryngeal nerve (i./.2) and a thin external laryngeal
nerve (e./.2) which receives a loop from the superior cervical
104 DR. C. F. SONNTAG ON THE VAGUS AND
ganglion of the sympathetic. Two small ascending branches pass
to ganglia whence fibres run up to, and along the course of the
arteries. At the level of these ganglia the common carotid artery
divides into thyroid, ectocarotid, entocarotid,and occipital arteries,
and fibres from the ganglia accompany them. The ganglia also
communicate with the superior cervical ganglion of the sympa-
thetic.
Text-figure 3.
DY ocr
The cervical aud anterior thoracic parts of the vagus and sympathetic nerves in
Euphractus villosus. The superior cervical ganglion (S.C.G) has been
cut down as it is very large and round and masks other structures.
Letters in text.
The Cardiac Nerves (f):—The two cardiac branches of the left
vagus end in a well-marked ganglion lying in front of the lower
SYMPATHETIC NERVES OF THE EDENTATA. 105
border of the aortic arch. Fibres radiate from it over the arch
and its branches, and constitute the superficial cardiac plexus. A
branch of the left recurrent nerve (¢) ends in the plexus, but the
recurrent and lower left cardiac nerves do not communicate as in
Tamandua. The right vagus gives two branches to the deep
cardiac plexus, but none to the superficial one.
The right recurrent nerve (d) has the usual origin, course, and
relations. It communicates with the sympathetic, and a fine
branch crossing the front of the trachea, connects it to the left
nerve.
The pulmonary nerves (g) are given off as the vagi reach and
pass behind the roots of the lungs. And they are connected by
fine nerves to the deep cardiac plexus.
The posterior thoracic parts of the vagi differ from those in
Tamandua. The vight vagus runs along the ventral aspect in
the csophagus. The left nerve runs along the dorsal surface of
the esophagus and communicates with the right one. It gives
ascending, transverse, and descending branches to the ceesophagus
and aortic plexus. Jn the abdomen the right vagus breaks up
into a number of branches which run along the ventral surface
of the stomach close to the lesser curvature. The left vagus
gives several twigs to the celiac plexus and its offshoots, and
ends in the dorsal wall of the stomach.
The Cervical Sympathetic:—The superior cervical ganglion
(S.C.G) is large and round. It communicates with the cranial
nerves in its vicinity, and sends a loop to the external laryngeal
nerve (el.n). Twigs of communication run to the ganglia
connected to the laryngeal nerve. The internal carotid nerve
(i.c.n) is given off as usual to accompany the internal carotid
artery into the skull. The mddle cervical ganglia are absent,
and the inferior cervical ganglia (1.C.G) are smaller and narrower
than those in Zamandua. And the cardiac nerves are similar to
those in the latter.
The Cardiac Plewuses (S.C.P and D.C.P) differ from those in
Tamandua. The superficial one is formed entirely by the left
vagus and sympathetic, and has a well-marked ganglion of
Wrisberg (G.W). The deep plexus is’ composed of branches of
the right vagus and sympathetic, and its solitary ganglion is
applied to the wall of the right auricular appendix. The two
plexuses are united by fine communicating branches.
The thoracic cords contain few ganglia as in Vamandua.
They increase in thickness from before backwards, and the
splanchnic nerves (/) are given off from an enlargement. There
is no branch comparable to the ascending aortic branch in
Tamandua. The abdominal part has few ganglia, and its rami
communicantes to the lumbar nerves (¢./.p) are well marked.
The Celiac Plewus (text-fig. 4) consists of right and left halves
connected by bundles of fibres, and united to the sympathetic
cords by the splanchnic nerves (A). It lies between the layers
of the common mesentery, along the course of the cceliac axis.
106 DR. C. F. SONNTAG ON THE VAGUS AND
It differs from that in Zamandua in that it receives branches of
the left vagus, whereas that of the latter receives the right vagus.
Text-figure 4.
f IV, L.Ve a./).
0.7).
boo
a,
XQ
LX |
MSO
a
UW
Fie S./77.p. cp.
cat : : : : , ;
I'he posterior thoracic and abdominal parts of the vagus and left sympathetic
nerves in Huphractus villosus. «a.n.: aortic nerves; 0.2.: cesophageal
nerves. Other letters in text.
There is no division into upper and lower parts. The following
are the branches connected to it :—
A. To the right half of the plexus :—
. Communicating to left vagus (c./.v).
. Coronary plexus (q).
Hepatic plexus (7).
Splenic plexus (4).
Right renal plexus (m).
Ort Co be
SYMPATHETIC NERVES OF 'THE EDENTATA. 107
B. To the left half of the plexus :—
1. Superior mesenteric plexus (s.m.p).
2. Aortic plexus (p).
3. Left renal plexus (0).
4, Phrenic plexus (/).
5. Communicating to left vagus (c.1.v).
The arrangement differs, therefore, from that in Tamandua.
Tatusia novemeincta (text-fig. 5).
The Vagus Nerves:—Both ganglia nodosa are present, and
communicating branches run to the superior cervical ganglion of
the sympathetic and other cranial nerves in its vicinity ; and the
branch to the spinal accessory is very prominent (¢.s.a). The
pharyngeal branch (a) resembles that in Huphractus. No
branches run to the sympathetic trunk.
Text-figure 5.
A.
The vagus and sympathetic in Tatusia novemcineta. A. Upper cervical part;
B: posterior thoracic and abdominal parts of the vagi. Description in
text.
The superior laryngeal nerve (b) forms a loop which commu-
nicates with the sympathetic cord distal to the superior cervical
ganglion; but no vascular nerves run upwards from it, as in
Euphractus. No lingual branch of the vagus is present as in
Tamandua.
The Cardiac Plewuses have the usual mode of formation, but
the superficial one has no Ganglion of Wrisberg.
In the posterior part of the thorax the left vagus divides into
two. One branch runs along the dorsal aspect of the cesophagus,
communicates with the celiac plexus and ends on the dorsal
gastric wall. The other branch runs along the ventral aspect of
the esophagus, communicates with the right vagus and ends on
108 ON THE VAGUS NERVES OF THE EDENTATA.
the ventral wall of the stomach. The right vagus runs along the
lesser curvature of the stomach to the pyloric region. Both vagi
send branches to the cesophagus.
The Caliae Plewus resembles that in Huphractus in most
points.
Summary and Conclusions.
1. The course of the vagus nerves resembles that in the
Marsupialia, but their thoracic relations are similar to those in
Man.
2. The superior cervical sympathetic ganglia are present in the
neck in Muphractus and Tatusia, but are represented by wide
expansions within the foramen lacerum posticum in Zumandua.
3. In Tamandua the pharyngeal plexus is more complicated
than that in Huphractus or Tatusia, and it is the only species in
which the lingual branch of the vagus is present.
4. In Huphractus the nerves accompanying the branches of the
common carotid artery are larger and more numerous than in the
other genera.
5. No Kdentate examined by me has a middle cervical sympa-
thetic ganglion, and the thoracic ganglia are few in number.
6. The cardiac plexuses are built on the same plan, but there
are variations in detail in their communications and ganglia in
the various species.
7. Hach species exhibits a different arrangement of the vagi in
the posterior part of the thorax, and their intercon:munications
are variable.
8. The solar plexus in Zamandua differs in its arrangements
and distribution from that in Huphractus and Tatusia. It does
not lie across the vertebral column, but is situated within the
mesentery along the course of the celiac axis.
9. The splanchnic nerves are well-developed in all species, but
do not all arise from ganglia.
10. There is no separate depressor nerve, and no external
communication between the superior and recurrent laryngeal
nerves as in Hyrax capensis and Man.
Bibliography.
1. Sonnac, C. F.—Proc. Zool. Soc. London, 1921, pp. 572-575.
2. Sonnrac, C. F.—Proc. Zool. Soc. London, 1921, pp. 873-876.
3. CunnincHam, D. J.—Text-book of Anatomy, p. 655.
ts
~)
— =
ae
Stephenson.
1922.
P.ZS.
heeta.
igoc
Morphology of Indian Ol
VITTY & SEABORNE, LTD.,
LONDON.
ON INDIAN OLIGOCH ATA. 109
7. Contributions to the Morphology, Classification, and
Zoogeography of Indian Oligocheta. By J. StepHEN-
peagrapnay $ )
son, M.B., D.Se., F.Z.S., Lecturer in Zoology in the
University of Edinburgh.
| Received October 29, 1921: Read February 7, 1922.]
(Plate I.* Text-figures 1-19.)
ContTENTs. Page
IV. On the diffuse production of Sexual Cells in a species of
Chetogaster (fam. Naididz)..............0..0... . 109
V. On Drawida japonica (Mchlsn.), 2 Contribution to the
Anatomy of the Moniligastride ....0....0.....ccccccceeeeeeeeee 119
VI. On the Relationships of the Genera of Moniligastride ;
with some Considerations on the Origin of Terrestrial
Oliwoahastape cents. ticscce es cc ke dentens saaonas meee eae ee eS
1V.—ON THE DIFFUSE PRODUCTION OF SEXUAL CELLS IN A SPECIES
OF CH #£TOGASTER (FAM. NAIDID#).
The Naidide are a family of fresh-water Oligocheta, of small
size, in which the usual mode of reproduction is asexual, by
fission. Sexual reproduction probably occurs, at times, in all;
but in many, perhaps the majority, of species, and in some whole
genera, it has not as yet been observed. Where it has been
described (except in the genus Pristina, where the organs are two
segments further back) the testes and male funnels are in segment
v, the atria and male apertures, and also the ovaries, in segment
vi. Except in Chetogaster, a single sperm-sac is constituted by
a posterior bulging of septum 5/6, and an ovisac, which envelopes
the sperm-sac, by a similar protrusion of 6/7; the sexual cells
given off from the gonads enter the sacs, where they ripen, and
ultimately again leave the sacs to be discharged.
Chetogaster is a somewhat aberrant genus of the Naidide,
which was separated as a distinct family (Chetogastride) by
Vejdovsky. It agrees, however, with the other genera of the
Naididz in the prevalence of asexual multiplication, and (so far
as hitherto known) with the majority (all except Pristina) in the
position of the gonads in segments v and vi. Thus Vejdovsky
(6) figures and describes the testes and ovaries of C. diaphanus
in these segments; while Beddard (1) says that “the sperma-
thece, testes, &c., are in the same segments in both the‘ families’
Naidomorpha and Chetogastride.” There appear to be no
detailed descriptions of the sexual organs in any species of later
date than these monographs, except my own of C. orientalis (4).
The species of Chetogaster which is commonest in Lahore in
N. India is that just mentioned, which I described first (3) as
C. pellucidus, but later (the specific name being preoccupied)
called orientalis (4). In this second paper I gave a description
* For explanation of Plate I., see page 132,
110 DR, J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,
of the genital organs, but although I had a number of specimens,
including one in a very early stage, I did not discover the testes
(‘the testes seem to disappear early, and I could not distinguish
them in a specimen which showed sperm-morule in all stages of
development, but no other genital organs, male or female, except
the ovaries in an early stage”).
Since that date I have again met with numbers of sexual
specimens of this species, and have always looked for the testes,
but have never found them, even in the earliest stages of
sexuality. On the contrary, careful examination of the living
animal (which is very transparent), of stained specimens, and of
sections, have united to convince me that the male cells are
produced in various parts of the body; and not only so—their
production is not limited to the parent animal of the chain, but
is spread over the whole series of individuals, and occurs in the
youngest members of the chain, which are scarcely recognizable
as separate entities, as well as in the most advanced.
The situations where the male cells arise are mainly as
follows :—
(1) On strands passing between the alimentary canal and
parietes. The genus as a whole, and certainly this species, is
remarkable for the incompleteness of the septa; in this species
there is nowhere a definite diaphragm across the coelomic space ;
and many of the strands, in various parts of the animal, from
which some of the sexual cells arise, should no doubt be regarded
as representing septa.
(2) On the inner surface of the body-wall, often in the angle
where one of these strands joins the parietes.
(3) In the new cellular-tissue which is formed in abundance
as a ring round the body at the site of the future divisions of
the animal. This cellular tissue gives rise to a number of new
segments; the region is hence called the budding zone, and the
production of this embryonic tissue is the first sign of subsequent
fission.
I propose to select as an example for description one out of
numerous observations, and then to add some account of the
appearances in sections, concluding with a few general remarks.
I may say that I have been particularly alive to one (I think
the only) source of possible error—namely, that the testes might
be quite inconspicuous masses in the normal position, that the
sexual cells might be detached very early in their history, and
the developing morule, wandering widely through the body,
become attached to the body-wall or to strands in various
regions, so simulating the production of morule in these
situations. This would, @ priori, not be impossible, for tbe
following reason :—There are, in this species, no sperm-sacs or
ovisacs, and this seems to depend on the incompleteness of the
septa, of certain of which the sacs, in other genera, are backward
protrusions; the sexual products, mstead of being received into
AND ZOOGEOGRAPHY OF INDIAN OLIGOCH ATA. 111
sacs, must therefore ripen in the general body-cavity ; and the
incompleteness of the septa allows them to become widely
distributed, both through the parent animal and the posterior
components of the chain. I believe, however, that the obserya-
tions I have made, only a part of which I record below, show
that this is not the case, and that the cells are really produced in
all parts of the chain.
Whether the ovaries also are similarly non-localized I cannot
say. I have described the ovaries as definite organs—as cellular
aggregates or as hyaline masses of nucleated protoplasm in
which cell outlines were not visible, suspended in the ventral
part of the body-cavity on fine strands in front of the level of
the sete of segment vi (7. e., about the position where septum 5/6
would normally be). The ovaries of C. diaphanus are several
times figured by Vejdovsky as definite spherical or pear-shaped
masses: they had previously been described by Lankester, and
were seen by Vejdovsky also, in C. limnei. Ova are, however,
sometimes met with in the hinder animals of a chain of C. orien-
talis; in their early stages, moreover, male and female cells are
indistinguishable; and it is possible that in some of the examples
which I assume below to be the early stages of the male
products, the cells may be young ova.
Text-figure 1.
To illustrate sites of production of genital cells in a specimen of
Chetogaster orientalis.
The above figure represents in outline a specimen of Chetogaster
orientalis, in which the process of fission has demarcated eight
future individuals, including the anterior one which we may
regard as the parent. The first constriction probably appeared
between the four anterior and the four posterior components of
the chain, and the first break in the chain would subsequently
take place here; the fifth individual was at the time of observa-
tion the most completely differentiated (after the most anterior
or parent). The figures and their reference lines indicate the
places where the sexual cells were visible.
None were seen at any stage of development in the living
animal in the front part of the parent, nor were gonads distin-
euishable; subsequently, after staining, examination in cedar oil
showed a small morula, of about four cells, ventrally by the side
112 DR. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,
of the nerve cord, almost at the level of the anterior border of
the crop*. At 1, two small morula were visible, attached to
strands in the dorsal region of the body-cavity, and moving with
the strands. At 2, ventrally situated at the level of the hinder
part of the cromaehy were a number of morule of large size, of
which one or more were quite free. At 3 were a considerable
number, apparently fixed to strands. At 4, in the next indi-
vidual, were a large number of morule behind the developing
pharynx and oesophagus, all seeming fixed to numerous strands
in this region; this attachment seemed also gene beyond doubt
on subsequent examination in cedar oil. At 5 were seen three
morule on a level with the anterior part of the crop, on strands,
to which they remained attached even on moderately violent
movements of the animal under the coverslip; one or two others
in this component of the chain were quite free. At 6 were two
morules, also apparently attached to strands; and this was con-
firmed by subsequent examination in cedar oil under the immer-
sion lens. At 7 were a number, one showing commencing
differentiation into central (granular) and peripheral (cellular)
parts ; two at least of these, well seen, were obviously, one would
say, fixed, and not merely entangled in the strands.
In the above example it happens that all the morulz in course
of development were situated on strands in the body-cavity; and
such strands appear to be the commonest site of their pro-
duction. JI have repeatedly seen the morulz on the strands keep
their attachment in spite of the animal’s movements and conse-
quent swaying of the strands to and fro—indeed, I have no note
of ever seeing them shaken off. These strands are frequently
situated in the anterior region of young individuals, towards the
front end or about the middle of what will be the crop, sometimes
behind its middle, or behind the stomach; as exemplified above,
they may be either dorsal or ventral to the alimentary tube; not
infrequently they are in the angle between the strand and the
body- wall, and might be described with equal correctness as
arising from the parietes.
The greatest production of morule may not be in the parent
animal—perhaps seldom is so. I have, for example, noted the
production as being most active in the individual behind the
primary fission zone, and in the one succeeding this, and as being
small in the anterior half of the chain. It Will be Hondereeaod!
that besides the fixed and not fully-developed morule, there are
other larger ones which float freely in the body-cavity.
A curious and perhaps pathological condition was met with in
some specimens which had been in a tube in the laboratory for a
* The alimentary canal in Chetogaster shows a considerable degree of differentia-
tion ; the pharyux may be said, in this species, to occupy the first three segments,
the esophagus the fourth, the crop the fifth to part of the seyenth, the stomach
most of the seventh and the eighth, and the intestine (very short compared with the
part of the tube so called in other Naididz) the few remaining segments.
AND ZOOGEOGRAPHY OF INDIAN OLIGOCHAETA. US
long time, and many of which must have died. Very numerous
small morule were budding from the body-wall, to which they
were all, at the stage examined, adherent. Some were also
budding from the surface of the alimentary canal; and here and
there some were seen in the angle between strands and parietes.
There was no special production in the proper genital segments
of the parent animal.
Sections 7 thick, stained in Delafield’s hematoxylin and
differentiated by a short immersion in iron-alum, dispel any
doubts as to the actual formation of morule on the strands; they
show that sexual cells may also be produced from the lining of
the body-wall, as well as—which is interesting—from the newly-
forming ring of cellular proliferation around the site of each
future fission. They also allow the history of the developing
morule to be traced.
Text-figs. 2-6, all taken from longitudinal sections, will illus-
trate the various situations of origin, and will supply the place
of further description.
Text-figure 2.
Proliferation of sexual cells from body-wall, near a zone of budding in a
posterior individual of a chain. X 1000.
The early stages in the production of morule may be shortly
summarized as follows :—
(1) The sexual cells when first recognizable as such (text-fig. 2
possess large, round nuclei, close together and with a diameter of
7m or vather less; a protoplasmic covering is scarcely dis-
tinguishable; in the periphery of the cell the chromatin is
scattered, but (as seen in sections) it forms a ring near the
middle, the centre of the ring being clear (text-fig. 2), or more
Proc. Zoou. Soc.— 1922, No. VIII. , 8
114
DR, J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,
Text-figure 3. Text-figure 4.
e.
A group of sexual cells forming on Formation of sexual cells from the cell-
a strand just infront of a zone ‘proliferation of a budding zone.
of budding in the hinder part The subsequent separation of the
ofachain. X 900. individuals would have taken place
between the two groups of pro-
liferating cells. 700.
Text-figure 5. Text-figure 6.
+ SOL.
Young morula on a strand in the Older morula, still attached to a strand,
body-cavity. > 850. traversing the ccelom, in neighbourhood
of a zone of budding. X 4380.
Reference letters for all the above text-figs (2-6) :—
e. surface epithelium: g. sexual cells; m. muscle strands of body-wall;
mor., morula; p. peritoneal cells; s. strands traversing ccelom ; 2. cell-
proliferation of budding zone.
AND ZOOGEOGRAPHY OF INDIAN OLIGOCH AIA. 115
usually (perhaps at a later stage) shows a smaller ring-like aggre-
gate in the centre (text-figs. 3 & 4). ‘hese characters s distinguish
the young sexual cells from the ordinary peritoneal cells. and
from the cells of the proliferations in the budding zones; ‘in
these the nuclei are on an average-4°3 to 5m in diameter, anid
the chromatin has no character “istic arrangement.
(2) The next stage, or young monte: ‘consists of an ovoid
ageregate of cells, en appear to be Tittle else than naked
nuclei (text-fig. 5); these are still of about the same” size, with
the clear ore and central aggreeate of chromatin.
(3) The cells become peripheral, leaving a granular mass in the
middle of the morula; cell boundaries are not distinguishable,
but the peripheral vegion of nuclei is marked off from the central
eranular mass (text- fis. 6). The characteristic enone or
the chromatin disappear S.
(4) The morulw, attached up. to this stage, may now become
free.
General hemarks.
Chetogaster orientalis appears to be the only Oligochete in
which a diffuse production of sexual. cells, similar-to that of the
Polycheeta, has hitherto been deseribed. It is possible, however,
that the phenomenon may occur in other species of the genus,
though the statements of previous authors (in particular Vej-
dovsky, cf. ant.) seem at first sight to negative this.
The testes ef C. diaphanus are figured by Vejdovsky (pl. v.
fig. 4) as rather loose masses of cells some little distanee behind
the septum, and are said in the text to arise on the first pair of
nerves given off from the ventral nerve cord. ‘The cells ave
loosely aggregated, and not bounded by a peritoneal membrane ;
they grow rapidly to four times their original size, become free
in the body-cavity, and then soon aitach themselves to the var 10us
organs and go through their further development—for example,
on the ventral nerve cord, or on the alimentary canal. The
later stages again become free. The original testes quickly
disappear.
Tt will be noted that.the mass of cells which Vejdovsky calls
the testes are not in the normal position of these organs; and it
seems possible that: they are only two among the numerous
groups of sexual cells which are, according to my observations on
CO. orientalis, scattered about the body. Probably also, in view
of the results of my sections of C. orientalis, the deyeloping g male.
cells which Vejdovsky describes as having become attached to
various organs really originated from their peritoneal covering,
in the same way that they originate, according to the above
descriptions, from strands, from the body- -wall, and from the
proliferating tissue of the budding. ZONES.
In the Polycheta the genital organs are as a rule not: strictly
localized ; the gonads are proliferations of peritoneal cells which
originate on the parietes, on the blood-vessels, or on the alimentary
8 **
116 DR. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,
canal throughout a considerable region of the body. In the
Oligocheta, on the contrary, this proliferation is restricted,
and the production of sexual cells is confined, in general, to one
or two testicular segments and a single ovarian segment in the
anterior part of the body; the gonads, moreover, arise from a
narrowly circumscribed area on the posterior face of the septa.
Tt is probable that the Oligochzta have originated from some
group of Polycheta; the evolution of the gonads has therefore
been from a non - localized to a more definitely localized
condition.
in Chetogaster orientalis the gonads (the male, at least) have
again lost their localized character; they are no longer definitely
circumscribed proliferations of peritoneal cells in a constant
position. That this condition, which is not unlike that of the
Polycheta, represents a regression, and not the persistence of a
primitive state, may be taken as certain ; Chetogaster is the
most modified genus of the Naididee—of the numerous characters
in which it differs from the rest of the family, not one can be
thought of as primitive; and I believe the Naidide to be
themselves far from primitive among the Oligocheta.
We may for the sake of clearness express the facts in a
somewhat different way. In certain animals the first visible
differentiation in development is that of the germ-cells; in
Ascaris, for example, as is well known, the peculiarities of their
chromatin constitution enable them to be distinguished from the
tirst. And it is possible that a similar ‘‘germ-track” exists in
all animals, but that we are unable, with our present means of
investigation, to distinguish it in all—at least in its earlier
stages. It is possible, in other words, that in all animals the
germinal material (germ-plasm) is set apart from the first in
certain cells and their descendants, and that in the shaping of
the organism these find their way, sometimes by fairly extensive
wanderings, to their station in the gonads, or rather to their
particular station in the ccelomic lining membrane (in the
Triploblastica) where, by their proliferation, they give rise to
the gonads. In the Polycheeta their distribution is wide; in
the Oligocheta extremely circumscribed ; in Chetogaster orien
talis it has again become wide.
With what pec elay of structure or habit, or with what
feature of the past history of Cheetogaster can this change be
correlated ? I can think of only one that is at all likely, and
even here I do not think it is easy to suggest how the change
has been brought about.
I have pr eviously suggested (5) that Chetogaster is descended
from parasitic ancestors. I do not wish to repeat the argument
at length; I will only say that Chetogaster, alone of the Naidide,
never shows the phenomenon of an ascending current of water
beginning at the anus and impelled up the. intestine by anti-
per Tistalsis and ascending ciliary action ; and that the loss of cilia
and of the ciliary function of the intestine is presumably due
AND ZOOGEOGRAPHY OF INDIAN OLIGOCH ETA. ARG
(using the principle which has in recent years been illustrated
by Dollo and Abel) to a different habit of life, which has left its
mark on the race; that this different habit must have been a
life in some other medium than that of water—i.e., it must have
been either terrestrial ov parasitic; that the first of these is
entirely inadmissible, and we are thus thrown back on the
second. Also that a number of the other peculiar features of
the genus, of which there are many, are explicable in a similar
way; and finally that Chetogaster at the present day is still in
soine cases an ecto- or even an endoparasite (in the pulmonary
chamber and liver of fresh-water Gastropods), while in others it
is a commensal; and in still others, though free-living, it (alone
among the Naidide, so far as I know) feeds predominantly on
animal food *.
Exactly how or why parasitism should have produced this
change in the distribution of the germ-cells, it is not easy to say.
Parasitism, of course, places a larger supply of material at the-
disposal of the reproductive activities; but this would not
necessarily lead to a different distribution—to a widening or
scattering of the germ-track. J can only suggest that there is
possibly some connection between past and present habits of
nutrition of Chetogaster orientalis and a change—a scattering—
of the germ-track.
The affinities of the Hirudinea with the Oligocheta have been
recognized for some time (compare, for example, the opening
paragraph of the chapter on Leeches, by Beddard, in the
‘Cambridge Natural History,’ of date 1896), though text-books
still often separate the two groups widely. Thus in the
arrangement adopted by Parker and Haswell the Gephyrea and
Archiannelida intervene between the Chetopoda and Hirudinea.
Michaelsen (2) has recently set forth the evidences for a close
relationship between the Oligocheta and the Leeches in detail.
The argument rests on the interpretation of certain structures, in
Sudanese and Sumatran species of leeches, as spermatheca which
communicate with the alimentary canal,—comparable there-
fore with those of many Enchytreids; on a consideration of the
Branchiobdellide and of Acanthobdella as intermediate forms
(though not necessarily in the direct line of descent of the
Leeches); and on the demonstration that there are no pecu-
liarities of the organization of Leeches that are not found, or at
least foreshadowed, in certain Oligocheta. Detaching the
Oligochta from their association with the Polycheta in the
* T may mention an additional observation, made since the date of my former
paper. I met with a specimen of Chetogaster orientalis endeavouring to consume
a whole Nais—a worm about as big as itself ; the end of the Nais had: entered the
pharynx of the Chetogaster, and I saw it drawn on into the region of the crop; the |
part of the Nais outside the Chetogaster was wriggling. On transferring the two
animals to a slide they unfortunately became separated—the Nais without its head.
The Chetogaster would not take in the Nais again, though I put them in contact
several times. }
Rotifers also are a very common food of Chetogaster orientalis.
118 =r. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,
Class Ghetopeds, Michaelsen unites them with the Hirudinea in
a new class, Cliteilata. ‘I consider the Hirudinea,” says he,
“trankly as Lumbriculide which have undergone special modifi-.
cations in adaptatton to a carnivorous habit of life.” It might
be justifiable, He thinks, to include the Leeches as a family, of
Oligocheta, though as we have seen he does not actually do this.
The charatter of the Leeches which is of interest from the:
point of view of the present paper is the large number of testes.
Michaelsen has no difficulty in showing that, as in the case of
other peculiarities, this too is foreshadowed in the Oligocheeta ;
for example, Lamprodrilus satyriscus, a Lumbriculid, has either
three or four pairs of testes. The position of the testes behind
the ovaries, however, offers greater difficulties; Michaelsen does’
not wish to assume a sudden dislocation backwards of the testes,
in such a manter that they “hop over” the ovaries. A trans-
lation of the.testes backwards can, he thinks, only take place
through the intermediation of a stage in which the series of
gonads are hermaphrodite. Hermaphrodite gonads have often
been observed; for example, in specimens which show an abnor-
mally large number of genital glands (“supernumerary gonads ”
—such series may extend as far back as segm. xviii), the anterior
are often testes;the middle ones: hermaphrodite, and the posterior
Ovaries; though Beddard in a Uvocheta (= Pontoscolea corethrurus)
found the hindmost gonads to be testes.
It seenis to me, honmere, that such a condition as that described
above in Cheetogaster orientalis furnishes, perhaps, a more likely
origin for the multiple testes of the Hirudinea. On such an_
assumption there would be no necessity to invoke a series of
hermaphrodite. supernumerary gonads—a double abnormality
where it occurs to-day—as a transitional stage. In Cheetogaster
orientalis male cells are produced abundantly, in every cau
individual, in the segments behind the ovaries; a stricter locali-_
ation aranelie result in segmentally arranged Hegban. And the,
ae cause Which we can surmise to have been active in the case.
of Chetogaster, orientalis—the adoption of carnivorous or para-'
sitic habits—must also have existed in the ancestors of the’
Leeches.
References to Literature.
1. Bepparp, F. E.—A Monograph of the Order of Oligochxta. Oxford, 1895.
2. Micwartsen, W.—Uber die Beziehungen der Hirudineen zu den Oligochiten.-
Mitth. Zool. Mus. Hamburg, vol. xxxvi. 1919, p. 131.
3. STEPHENSON, J.—Descriptions of two fresh-water Oligochete worms from the
Punjab.’ Rec. Ind. Mus.,'vol. i. 1907.
4. , ‘Studies on the aquatic Onvecneel of the eons Rec. Ind. °
i Mus., vol. v. 1910.
5. % On Intestinal Respiration in Annelids : with Considerations
on the Origin and Evolution of the Vascular System in
that Group. Trans. Roy. Soc. Edin., vol. xlix. pt. iii.
1913.
6. Vrspovsxy, ¥.—System und Morphologie der Oligochiaten. Prag, 1884.
AND ZOOGEOGRAPHY OF INDIAN OLIGOCHATA. 119
V.—On DrawipdA s4Ponrca (Mchlsn.); A ContRIBUTION TO THE
ANATOMY OF THE MoniLicasTRip&.
The Moniligastride are a family of Oligocheta with a rather
restricted range. Hxcept for the recently discovered Syngeno-
drilus, which differs very considerably from all other members of
the family, they are endemic only in India’(mainly the South),
Bur ma, and ee islands of the Malay Archipelago; a few
species of the largest and best known genus, Drawida, however,
have been able to spread more widely, : and have been found in
Japan, China, and certain islands cf the Pacific. The writings
of Beddard, Benham, Bourne, Rosa, and Michaelsen have sitet
dated the anatomy of the several genera, and in a number of
cases, by means of sections, the finer structure also. Since,
however, the worms occur for the most part in localities remote
from centres of population, writers on the group have as a rule
been dependent for their material on collections made by others,
without, it is to be supposed, any special methods of fixation.
While such material is adequate for systematic work (for which
it has generally been used), it is often quite hopeless as a basis
for more minute investigations. The only students of the group
who have had the opportunity of fixing living material for
themselves appear to be Beddard (7), who received specimens of
Drawida bahamensis alive from Kew Gardens, and Bourne (11),
who colleeted his own material on the Nilgiris in Southern India.
These authors do not, however, indicate the methods they employed
for fixation.
A few years ago one of my pupils, Mr. G. 8. Thapar, M.8c.,
now Professor at the Canning College,. Lucknow, brought back
with him to Lahore from Murree in the W. Himalayas a
considerable number of living specimens of Drawida japonicus,
one of the per egrine species of the genus. These were kept for
some days 1n the laboratory in scraps of moist blotting paper,
and the alimentary canal having been freed in this way from
earth, they were, after Harcotization: fixed in Zenker’s solution.
Some specimens were dissected, . and others sectioned, both
transversely and longitudinally, and the sections stained with
Delafield’s hematoxylin followed by eosin. The chief attention
has been devoted to the nephridia and the genital organs. —
Of the nephridia‘in the genus Drawida, or in the family, we
possess nothing that can be called.a detailed deser iption.
The general : anatomy of the reproductive organs in the genus
is, however, well known, and may be summarized as follows : —
The testes are one pair, enclosed each with its funnel in paired
testis sacs, which have a peculiar position ; they are suspended
by septum 9/10, fairly high up in the body- cavity, projecting
sometimes more on the poster ior, sometimes more on the anterior,
side, sometimes equally on both sides of the septum. ‘The vas
deferens leaves the sac, and descends, with many twists and coils,
120 DR. J. StEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,
on one or other or both sides of the septum; on reaching the
ventral body-wall it runs backwards for a short distance and
joins the prostate, a glandular mass in segm. x of cylindrical
or rounded shape, with a central lumen, which discharges to the
exterior in the groove between segms. x and Xi.
The ovaries are on the posterior face oe septum 10/ ‘11, the
funnels on the anterior face of septum 11/12; the short oviducts
reach the surface in groove 11/12. A large pair of ovisacs—
backward pouchings of septum 11/12—extend backwards through,
as a rule, several segments. The spermathece are ovoid sacs
attached to the hinder face of septum 7/8; a winding duct leads
down the septum to the body-wall, and ends in the corresponding
groove; just before it enters the body-wail there is often a
muscular sac, the spermathecal atrium, of moderate size, or small,
or it may be even minute, connected with it.
In the account which follows of the minuter structure of the
various organs, references will be made where necessary to the
previous work of Beddard (2, 3, 6, 7), Bourne (11), and Benham
(10), who used serial sections in their investigations, and to that
of Michaelsen, whose systematic papers (13, as, 16) contain the
record of important anatomical details.
(1) Zhe Nephridia.
The nephridia (Pl. I. fig. 1) are attached to the posterior faces
of the septa; in the middle of the body each has a total length
of about 1 mm.
Each is divisible on inspection into four portions :—(i.) The
anteseptal portion with the nephrostome; (i.) the twisted
nephridial tube; (iii.) larger than the last portion and more
dorsally situated, an oval and vertically elongated sac, occupying
the mid-lateral region as it lies on the body-wall in the dissection ;
(iv.) projecting from the lower end of the sac is a stiff, straight
terminal duct.
The nephrostome (7) is 45—50 pw in Hinmiciee and has a marginal
rosette of about 20 ciliated cells.
From the nephrostome leads the nephrostomial canal (7.c.),
8-10 » in diameter, which pierces the septum, and after a straight
course of about 170 « becomes the spiral coil.
The portion of the organ which was roughly distinguished
above as the twisted nephridial tube can be divided on closer
inspection into three parts, which may be named the spiral coil,
the twisted loop, and the spindle-shaped portion. The diameter
of the tube throughout this part of its course is 40-45 p ; it is of
a slightly brownish colour. The lumen is wide, but intracellular
—bored through the middle of a single series of cells (drain- pipe
arrangement). ‘The appearance of both longitudinal, transverse,
and optical sections is at first sight misleading; the drain-pipe
series of cells appears to be free within the lumen of a larger
AND ZOOGEOGRAPHY OF INDIAN OLIGOCH ETA‘ 121
tube. This larger tube, however, is easily recognized as the peri-
toneal covering ; it consists of separate cells, shortly columnar in
shape, with the nuclei peripherally situated.
The spiral coil (coil) lies more ventrally than the rest of the
organ ; it is altogether ventral to the lower end of the oval sac.
'The spiral consists of about two turns; its apex is below, so that
the nephridial tube followed from the nephrostome first descends
to the apex, and then ascends as the returning limb of the spiral.
On quitting the spiral coil it becomes the twisted loop, a dorsal
continuation of the coil, which lies by the side of the lower por-
tion of the oval sac. Thus roughly half the vertical extent of the
twisted nephridial tube consists of the spiral coil which is below
the level of the oval sac, and half of the twisted loop, situated
alongside the lower part of the sac; the total extent of the twisted
nephridial tube is.about ‘55 mm. The twisted loop is retained
by a mesentery which joins that of the oval sac.
The twisted loop (Joop) contracts to 12 » in diameter at its end,
and joins the oval sac by the intervention of a spindle-shaped
portion (spin.). This part of the organ is 80-100 » in length and
40 w in greatest thickness; it is perfectly circular in transverse
section, has thick (7) walls, and a lumen which may be widely
patent, or may be almost blocked by faintly pink-staining
nucleated cells.
The oval sac (sac) is much elongated dorso-ventrally, gently
curved round the alimentary canal, and somewhat flattened ; it is
the largest and most conspicuous part of the whole apparatus,
being °65-"73 mm. in length and -17 mm. or more in width (in-
cluding its peritoneal covering). It has a very bulky coat of
high, clear peritoneal cells (p.), 40 ~, 60 » or more in height; in
one case the peritoneal coat formed a layer of cells thicker
(-18 mm.) than the whole of the included sac. ‘The nuclei of the
peritoneal cells are at various levels.
The muscular coat (m.) of the sac is remarkably thick—12 p~—
and consists principally of circular fibres. Inside the muscular
coat is a thin but very obvious connective-tissue coat, staining an
intense blue with hematoxylin, about 1» thick. The epithelial
lining (ep.) consists of lightly staining low columnar or cubical
cells; its height is about 12 ~—-about as thick as the muscular
coat.
A mesentery, about °32 mm. in width, passes from one border
of the sac to the septum; it consists of a layer of blue-staining
connective tissue, continuous with that of the wall of the sac and
with that of the septum, covered on both faces by peritoneal
cells.
The spindle-shaped portion is similar in its structure and its
mesenteric relations to the sae.
The terminal duct (d.) has a length of *3-—33 mm. in its course
from the ventral end of the oval sac to the body-wall, and its
diameter is 40 u. It comes off just to one side of the ventral end
222 DR J, STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,
of the oval sac, is straight and stiff, and- makes a right angle or
rather less with the general direction of the sac (except where
the duct has the lower opening, v. if.).
In whole preparations of the nephridium the ectal end, where
it is broken off from the body-wall, always appears trumpet-
shaped, the end of the trumpet being oblique and the dorsal lip
the more prominent (é., fig. 1). The duct is covered by a layer
of cubical peritoneal cells; the remaining layers are the same as
those of the oval sac, but all are thinner; the lining epithelium
consists of approximately cubical slightly staining cells.
The duct is attached by a narrow mesentery to the posterior
face of the septum. Reaching the angle between the septum
and the parietes 16 cilates somewhat, the dilatation corresponding
to the trumpet-shaped ending of the duct as seen in the isolated
nephridium (v. sup.).
The duct may reach the hady- wall in one: e two situations—
either at the level of the lateral or of the ventral sete; in the
first case it makes an angle rather less than a right angle with
the axis of the oval sae, Sanita in the second it appears as a direct
downward continuation of the sac. At both these levels the
longitudinal muscular coat shows an interruption in transverse
sections ;- the . trumpet- shaped dilatation rests ther efore on the
circular muséular coat.
-If the duct has the lower position, it penetrates the parietes
forthwith; the blue-staining layer becomes continuous with the
similar layer of the body-wall, and the epithelium of the duct
with the superficial epithelium of the body. The same may
happen if the duct reaches the body-wall at the level of the
lateral setee—the duct may penetrate the parietes immediately,
in a slightly upward direction. Often, however, the duct
continues in the body-wall in an upward direction, and reaches a
level not very far from the mid-dorsal line—a level corresponding
approximately on the dorsal to that of the ventral sete on the
ventral side—before it comes to the surface.
In the latter case the muscular layer of the duct ceases to be
distinguishable; the blue-staining layer becomes continuous with
a very marked similarly staining layer between the circular and
iovnetntanoel muscular coats of the body-wall; and the epithelial
layer of the duct is only distinct at the beginning and end of its
course within the wall. For the- greater. part of this course the
duct appears rather like a split in the_body-wall immediately
internal to the circular. muscular layer; the epithelial layer is
difficult to make out (except at the tyo ends), and the lumen of
the duct appears in most parts to be ‘bounded by a very definite
wall of the blue-staining connective tissue.
1 give below the positions of the endings of Ab aerial
ducts in a consecutive series of eleven segments, both on the
right and left sides. . It will be noted that there are no apertures
in the lowest position (on the level of the ventral sete) on the
left side and none in the dorsal position on the right side.
AND ZOOGEOGRAPHY OF INDIAN OLIGOCH ATA. 123
(Upp.=uppermost position, not far from mid-dorsal line; mid.=
middle position, at level of lateral setee; low.=lowest position,
at level of ventral sete.)
Right. | Left.
maids, | | upp.
low.7 mid,
mid, upp.
low. upp.
mid, upp.
mid. mid.
, low. upp.
mid. mid.
mid. ‘mid.
mid. upp.
mid, upp.
(2) The Testis Sacs (Sperm-sacs).
Beddard describes the sperm-sacs of D. barwelli (6). He.
makes the interesting suggestion contained in the following
extract (postscript to 6):—‘‘ The remarkable partial obliteration
of a segment (the xilith) which Michaelsen has recently described
in Nemertodrilus griseus, suggests that something of the same
kind may have occurred in the Moniligastride, the supposed.
sperm-sacs may be all that 1s left of the ccelom belonging to the
segment which contains the testes. This is, of course, no more:
than a suggestion; but the varying position of the essential
organs in the Oligocheta requires, as I point out in a forthcoming
number of the ‘ Quarterly Journal of Microscopical Science,’ some
possibility of the intercalation or excalation of segments at the
head end.” The sacs are also described by Benham in DP. indi-
cus (10), where the eavity is traversed ky muscular fibres passing
in different directions ; and by Bourne in D. grandis (11),
As in all species of the genus Drawida, the testis sacs,
containing the testes, male funnels, and ripening spermatozoa,
are in the present species a single pair, suspended by septum 910.
In some species of the genus they, project about equally’ into
segms. ix and x, in some they project forwards into ix more than
backwards into'x; often there are differences on the two sides—
individual differences, not specific. In the present species the
sacs project backwards more than forwards. The appearance is
somewhat as if the septum had split, the sacs being cavities
within it (fig. 2). | aaa
‘Both anterior and posterior walis of the sac have the usual
peritoneal (p.), connective-tissue, and muscular (m.) components.
The inner lining of the sac is in places a cubical or flattened
epithelium (ep.) ; over the greater part of the wail, however, the
lining is not distinguishable from the mass of cells which fills the
cavity. This mass stains of a pinkish colour with eosin (g.), cell
124 DR. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,
outlines are not visible in it, and there are numerous lightly
staining nuclei (n.); there are also many yellow granules
resembling chloragogen, often aggregated into considerable
masses (a’.). Blood-vessels (0.v.) traverse the cavity; in some
eases the chloragogen-like granules are in—and indeed block—
the blood-vessels.
The male funnel is a region of high columnar epithelium (/-)
surrounding the exit of the vas deferens from the sac; the cells
are exceptionally high near the opening of the vas, and become
lower by degrees at the periphery of the patch; their nuclei are
much elongated, and are placed at or above the middle of the
height of the cells. The vas deferens (v. def.) after emerging is
attached in numerous closely-set short loops on the posterior side
of the septum.
The testis (¢.) is a cellular proliferation just anterior to the
opening of the vas deferens; it is thus implanted actually on the
funnel. Its attachment is narrow, but the cell-mass spreads out
considerably inside the sac. Near its attachment the nuclei are
small, and stain deeply ; in the middle portion of the organ the
nuclei are closely packed, with the chromatin in discrete rounded
particles ; more peripherally, cell outlines, which are wanting in
the more central (or proximal) parts of the testis, appear ; and
lastly, discrete cells are detached from the surface of the organ.
There were no ripe, or even ripening, spermatozoa in the sacs (the
Moniligastride are apparently characterized by a seasonal sexual
maturity, like the Limicole, and unlike most earthworms; the
clitellum, for example, is often absent or very poorly marked) ;
moreover, it is pretty evident that the pinkly staining mass, with
small lightly staining nuclei, which fills out the greater part of
the sac, is not directly concerned in the formation of spermatozoa.
(3) The Prostates.
There already exist a number of observations on the structure
of the prostates in the genus Drawida, but these are not concor-
dant throughout. The lining epithelium, the layer of muscular -
fibres, and the “ gland cells” are distinguished in all descriptions;
but authors are not agreed as to whether the gland cells are to
be referred to the peritoneal or to the epithelial layer.
Beddard, in a first communication on D. barwelli (2), states
that outside the layer of muscular fibres there is a layer of pecu-
liarly modified peritoneal cells. In a later account (3) this
glandular tissue is described as consisting of large granular cells
separated into groups by partitions; each cell is prolonged into a
fine process, which extends at least as far as the muscular wall;
indeed, says the author, it is difficult to believe that the cells do
not in some way or other reach the lumen of the atrium and
there discharge their secretion. Nevertheless, this layer of gland
cells is the modified peritoneal layer, and there is no peritoneal
coat outside it.
AND ZOOGEOGRAPAY OF INDIAN OLIGOCH ILA. 125
In a third paper (6) Beddard refers toa recent paper of Rosa’s
on Desmogaster doric, ~ member of another genus of Monili-
gastride (19); there is here the same annular layer of muscular
fibres outside the lining epithelium, and in addition muscular
fibres are interspersed between the bundles of glandular cells ;
a delicate peritoneal investment surrounds the whole. Rosa,
however, states that the gland cells empty into the lumen of the
prostate by long ducts, and the figure also indicates that these
ducts—apparently prolongations of the cells—pierce the layer of
circular muscular fibres ; nevertheless, Beddard thinks it 1s still
possible to refer all that lies outside of the lining epithelium to
the peritoneum (using “ peritoneum” in a wide sense, Beddard
speaks of the muscular layer also as having been developed from
the peritoneum).
Beddard also gives an account of the prostate of D. bahamensis
(7). The gland-cells le in groups in the interstices of a some-
what loosely arranged muscular mass; the ducts of the cells—
simply filiform prolongations of their substance—pierce the
muscular layer in bundles, and evidently pour their secretion
into the Jumen. ‘There is no peritoneal layer outside.
Benham examined MJoniligaster indicus (=Drawida robusta)
(10); the specimens were badly preserved. The necks of the
glandular cells pass through the muscular coat nearly up to the
epithelium, but not quite. The epithelium is traversed by
narrow, clear tubes resembling ducts, which differ from the
epithelial cells in having no inner boundary. There is a ccelomic
epithelium outside; hence the gland cells are not peritoneal, and
must belong to the layer.of epithelial cells. Wise
Bourne, in D. grandis (11), found the gland cells arranged in
groups, each cell, nowever, sending its duct to take its place
among the columnar cells of the epithelial lining. There are two
layers of muscle—one overlying the epithelial layer and within
the glandular layer, and another outside the glandular layer; a
layer. of coelomic epithelium invests the whole.
There is thus an extraordinary variety in the accounts of this
organ. The layer of gland cells is said to be separate from the
epithelial layer, and to send prolongations as ducts through the
epithelium to the lumen; to have outside it a peritoneal invest-
ment, and to have no such layer; and to belong to the epithelial
layer, or to the peritoneal layer. In the present species there
seems to be no doubt that there is a peritoneal layer outside the
layer of glandular cells; but equally definitely these latter seem
to be altogether cut off from the epithelial layer and the lumen
of the organ. JI am disposed to believe that the cells belong to
the peritoneum rather than to the lining epithelium; and to
agree with Beddard, who would hold that the presence of a
flattened peritoneal layer outside the glandular cells does not
preclude the peritoneal origin of these latter.
In the present species the prostates (fig. 3) are situated in
126 DR. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,
seem. x. Hach is a short, thick cylinder, with a soft, white
lobulated surface and a central canal opening on the surface in
groove 10/11.
The outermost coat is’ by far the most bulky; it is composed
of large cells (prost.) with distinct outlines, the cytoplasm of
which stains moderately with hematoxylin. The cell substance
isin general granular, the nuclei (mwe.) are small, homogeneous,
of various shapes, often peripherally situated, and do not stain
much more deeply than the cytoplasm. In one specimen there
are numerous vacuoles (#.) in the cells, each with a small, round,
homogeneous, lightly staining mass in the centre.
In thickness this ‘layer is 120 » or more. The cells do not
ierce the muscular and connective-tissue layers to join the
epithelial lining of the prostatic lumen; some cells are sessile on
the muscular layer, others have their rounded inner ends slightly
separated from it (ef. fig. 3). There can, I think, be no real
doubt that the layer of large cells is not a bursting of the lining
epithelium through the muscular coat.
Outside this layer there is often to be seen a peritoneal coat
as a very thin blue-staining membrane—a mere line—with here
and there more or less flattened nuclei, often ‘apparently
degenerate. ‘In some cases the membrane spans the angle
between the outer ends of neighbouring cells. Sometimes a few
muscular fibres may be seen outside the layer of large cells.
The peritoneal layer may also carry blood-vessels of considerable
size, some branches of which penetrate the layer of large cells.
From what are the cells of this layer derived? The presence
of a peritoneal layer surrounding them would seem to preclude
the possibility of their being themselves peritoneal in origin, as I
at first supposed; while the fact that they are totally cut off
from the lining epithelium—separated from it by the museular
layer—seems to shut out the possibility of referring them to the
epithelial layer.
I believe that they are peritoneal in origin, however; the fact
that they are covered in places by a flattened peritoneal epi-
thelium does not necessarily negative this view. A peritoneal
membrane persists over the layer of large cells in the same way
that such a membrane persists over the lobes of the testes in
species of Lumbricillus: the testes are proliferations of the
celomic epithelium, which in this genus (as in Ocnerodrilus) ave
covered in at first by a delicate peritoneal membrane.
(4) The Vas Deferens.
The vas deferens descends from the testis sac to the ventral
body-wall, and in doing so forms a number of coils on both sides
of septum 9/10. In this part of its course its diameter is about
40»; its lining epithelium consists of cells staining pink with
eosin, which almost occlude the lumen and whose nuclei are
situated below the middle of their height; it possesses also a
AND ZOOGEOGRAPHY OF INDIAN OLIGOCH ETA. 127
well-marked connective-tissue (blue-staining) coat, 2°5 yu thick
and rather loose in texture.
Having reached the parietes it runs backwards for a short
distance, and then, sinking into the body-wall, lies in the blue-
staining connective tissue between the two muscular layers ;
here its diameter is 30 ,, and its connective-tissue coat is still
moderately thick. Arrived at the base of the prostate, it enters
this organ at its lower end, and runs upward outside the
muscular coat of the prostate and within the layer of large cells,
in a direction parallel to the prostatic lumen; its diameter is
now 20, and it still possesses a deeply staining connective-
tissue layer outside its. epithelium. Sinking “through the
muscular layer ot the prostate, the rest of its course lies. within
that coat, in the lining epithelium (fig. 3, v. def.); it finally opens
into the. prostatic lumen at the ental extremity of the latter.
In this‘last part of its course it loses its blue- -staining layer and
becomes a small, round tube of cubical cells, 18 u in diameter.
The actual relations ‘of vas and prostate, are thus disguised,
inasmuch as the vas, entering the prostate at the base ‘of the
latter, and ascending ath its wall to the apex, is hidden from
view during this part of its course. In a species of Drawida
(D. raui), eich! I have recently described (26), and which is in
respect of its male apparatus the most primitive of the genus (it
possesses two pairs of prostates, an ancestral character), “ the vas
deferens, joining the prostate below, can be seen running up its
surface towards the free upper end. Dann Moniligaster deshian yest
the prostates are very large and sausage-shaped, and extend back
through several segments ;° the vas deferens passes back along it
to fuse with ‘it some little distance from its ental end; and
presumably sections would show that here, too, the mae is a
separate tube as far as the ental end of the prostate, where the
one presumably passes into the other; in two varieties of this
species the vas can be seen in an ordinary dissection to enter the
ental end of the prostate.
In certain species of the genus Drawida, therefore, and more
obviously in the genus Monzlig iguster, the prostate i is to be regarded
as the thickenéd « continuation of the vas deferens surroundéd: by
a. bulky mass of peritoneal cells. These cells give the organ thé
soft and papillose surface which frequently characterizes Goseoln
other species of Drawida, however, the surface of the prostates
is firm and shining—an appearance which is associated with a
firm muscular investment; cither there is here no soft covering
of swollen peritoneal cells, or the layer is so intermixed with and
covered by muscular fibres as to give a firm and resistant
surface.
(5) The Ovarian Chamber and its Contents.
The first indication that there is anything peculiar in the
septa which come into relation with the female organs in the
128 DR. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,
genus Drawida occurs, perhaps, in Benham’s description of
Moniligaster indicus (= Drawida robusta Bourne), where it is
said (10) that septum 10/11 is thrown back a whole somite, and
is confluent with 11/12 along a certain part of its course laterally,
but becomes separate towards the middle of the body (7. e., pro-
bably, towards the mid-dorsal and mid-ventral lines). This is,
however, not the condition of the ‘ovarian chamber” which I
wish to describe.
Bourne (11) describes an arrangement of the septa which
results in the cavity of segment x1 being reduced to two sacs ;
one of these contains the ovaries and nephridia of the segment,
and into it the oviducts open internally ; the other sac contains
the portions of the dorsal vessel, alimentary canal, and ventral
vessel which belong properly to the segment. ‘Ihe description is
not clear, nor does the figure solve the difficulties.
Michaelsen (18), describing D. willsi and D, nepalensis, speaks
(in the latter case with doubt) of an ovarian chamber formed by
septa 10/11 and 11/12, and a fine connecting membrane; the
chamber is thus a separated-off portion of segment xi. In a
subsequent paper (14) he seems inclined to recognize the ovarian
chamber as itself the xith segment in D. travancorensis, mattha,
and ghatensis. Cognetti states (11 a) that D. fakir has the
ovaries enclosed in a thin-walled peri-cesophageal capsule.
I have myself recorded the relations of the female organs in a
number of species (21, 22, 23, 24, 25), and have found that the
condition is not the same in all. Some species have nothing note-
worthy in the arrangement of the septa in this region (D. pellu-
cidus var. stewarti, D. kanarensis). The complete ovarian
chamber, found in D. rotungana, kempi, robusta, rangamatiana,
nepalensis, ghatensis, decourcyt, barwellt var. impertusus, and in
Moniligaster deshayesi, is produced by the two septa which bound
the somite containing the ovaries and funnels coming together
and fusing, except over a horseshoe-shaped space which spans the
alimentary canal above ; the canal passes through an opening in
a membrane which represents the fused septa 10/11 and 11/12,
and a needle can be passed above it and below the ovarian cham-
ber without doing any injury to the chamber or any organ. In
a few species (D. hodgarti, afinis, papillifer, chalakudiana) the
septa concerned in the formation of the chamber are not fused in
the dorsal part of their extent; the consequence is that while the
chamber is formed below in the manner just explained, and spans
the alimentary canal as’ described, it reaches and is bounded by
the dorsal parietes above. In two species (D. brunnea and
parambikulamana) the septa meet above exactly at the dorsal
parietes, so that the chamber is just and only just excluded from
the dorsal body-wall.
As shown by sections, the condition (in this worm at least) is
somewhat more complicated than was imagined, on the basis of
dissections, by either Michaelsen or myself.
In the present species (fig..4) there is dorsally in place of
AND ZOOGEOGRAPHY OF INDIAN OLIGOCHAETA, . 129
septa 10/11 and 11/12 only a single septum, which is to be
regarded as a product of the fusion of the two. More ventrally
these separate, so that in a longitudinal section taken to one side
of the alimentary canal, segm. xi appears as a wedge-like space
with its apex upwards and base at the pavietes. This wedge-
like space is continuous above the alimentary canal with the
similar space on the other side.
There are, however, other complications in the form of this
ehamber which are difficult to describe in a few words. If the
above were a complete description, it is obvious that segm. xi
would be an ordinary segment, except that its bounding septa
would be fused dorsally; and it would contain a section of the
alimentary tube, dorsal and ventral vessels, and ventral nerve
cord. What actually happens may be described by saying that
(a) throughout the whole of the gizzard and reproductive regions
of the worm the septa are incomplete in their mid-ventral por-
tions—around the ventral vessel and ventral nerve cord—as if a
series of arches were cut out of them; (6) that all round the
alimentary canal, between septa 10/11 and 11/12, a ring-shaped
partition isolates an annular ccelomic space (figs. 4 & 6, ann.)
round the tube; this annular cavity contains also the dorsal
vessel (d.v.), but the ventral vessel is below it; (c) longitudinal
and almost vertical partitions are also established, extending in
the ventro-lateral region between septa 10/11 and 11/12, and thus
isolating the mid-ventral portion of the coelomic cavity here
(fig. 6, v.ch.).
The cavity of segm. xi is thus altogether absent dorsally ; and
what exists of 1t is divided up as follows:—(i.) An annular space
immediately surrounding the alimentary canal and containing
also the dorsal vessel ; (i1.) a larger space, the ovarian chamber
(figs. 4 & 6, o.c.), surrounding this in turn, of which (111.) the
mid-ventral portion is isolated by longitudinal partitions on
each side of the ventral vessel and ventral nerve cord, but is
continuous in front and behind with the ccelomic cavities of
neighbouring segments *.
The ovarian chamber contains the ovaries with their funnels,
and the nephridia. The ovisae (fig. 4, sac) is an evagination of
its posterior wall. The ovaries are proliferations of the epithelium
covering the anterior wall of the chamber. The funnels (/) are
constituted by modifications of the epithelium of the posterior
wall.
The walls of the chamber have the usual constitution of
septa—peritoneum on the outside, muscular and connective
tissue composing their substance. The inside of the chamber is
* There are other peculiarities of the septa in the neighbouring regions which are
difficult to make out. Thus the floor of the annular chamber round the gut in
segm. xi is continued forward and backward as a sheet of tissue which extends from
behind the gizzard region to the spermatiecal segment. This sheet may have free
lateral margins, or the lateral margins may be bent upwards to join the ventro-lateral
wail of the cesophagus.
Proc. Zoou. Soc.—1922, No. 1X. 9
130 DR. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,
lined by a layer of cubical epithelium, modified as follows in
certain regions :
(1) The funnel—a long, rather curved groove on the posterior
wall, extending from the roof of the chamber downwards to the
beginning of the oviduct. The cells composing it are high and
narrow, with much elongated nuclei situated above the middle of
the height of the cells; the cytoplasm is granular; the free
surface is bordered by a layer of rodlets (‘ Stiibchen ”’).
(2) The ovary, on the anterior wall, consisting of a number of
finger-like lobes attached along an oblique area on each side of
the alimentary canal; the upper portion of this area is quite
narrow, but it evadually widens below.
(3) Various “proliferations of small cells, with deeply staining
nuclei; these constitute somewhat indefinite heaps which do not
seem to be on the way to develop into ova, since the characters
of the cells are the same at the periphery of the mass as deeply
in its substance. These masses occur on both anterior and pos-
terior walls (one is cut dorsal to the gut on the posterior wall of
the chamber in the figure), and also at various places within the
ovisacs.
In another specimen the ovisac was ial filled by pinkish-
yellow (yolk) granules, amongst which nuclei were occasionally
seen, and a few of the small cells mentioned above. A thick
layer of the grains replaces the coat of proliferating cells on the
inner Sumtaces of the wall of the ovisac. In the chamber and
ovisacs there may be numerous small yellow masses like those in
the testis sacs ; these frequently have a number of dark granules
within them.
(6) The Spermatiece.
The spermathecal apparatus in the genus consists of a spherical
or ovoid ampulla, attached to the hinder surface of septum 7/8,
and communicating with the exterior by a long, coiling duct; the
duct leads downwards on the septum, and ends in an atrial
chamber which debouches to the exterior in the intersegmental
fissure. The atrium is the variable part of the apparatus; it may
be of considerable size, and either simple in form, projecting to
one side or other of the septum where this joins the ventral
parietes; or bifid, one horn projecting on one side and one on the
other side of the septum; or it may be small—even invisible in
dissection, owing to its being embedded in the body-wall; or
possibly, in some cases, 1t may be altogether absent.
In the present species the ampulla consists of the usual coats.
The lining epithelium is shortly columnar over the vault of the
sac and away from the origin of the duct, and the nuclei are
round; it is high, and may be very high, on the lower part of the
sac near the origin of the duct, and on the anterior wall, and the
nuclei are correspondingly elongated. All the cells are provided
with rather short cilia. Embedded in the much elongated
epithelium at one part of the wall was a quantity of bright yellow
AND ZOOGEOGRAPHY OF INDIAN OLIGOCHATA, W331
matter, lighter in tint than that in the testis sac, in granules,
and also in larger spherical aggregations up to 20 4 in diameter.
The ampulla is attached by a connective-tissue mesentery to the
septum.
The first part of the duct is wide—125 ~»—and has a consider-
able lumen ; it is lined by moderately high columnar epithelium,
with cilia which spring in bunches from each cell, ‘The neck by
which it is joined to the ampulla is narrower, 60 « in diameter
and similarly ciliated. The duct is also suspended by a mesentery.
The coiled part of the duct is narrower, with a diameter of
D0; 161s remarkable for the great thickness of its muscular
coat—15 »—much more marked than in the previous portions of
the apparatus ; the fibres are mostly circular, The epithelium is
cubical and non-ciliated ; a lumen is not visible. The mesentery,
with marked muscular fibrils, branches to its attachment to the
various coils.
The atrium in this species is small, and is contained within the
lower part of the septum; it narrows downwards to pierce the
body-wall. ‘The duct joins it near the apex; but it is included
for some distance within a common connective and muscular coat
before the lumina coaiesce. The epithelium of the atrium is high
and irregular, and hence the lumen is small, and irregular also;
the nuclei are situated towards the base of the cells; there are no
cilia. The muscular coat is independent of the musculature of
tlie septum. In the body-wall the epithelium is lower, columnar,
non-ciliated ; the muscular coat is here apparently derived from
the circular fibres of the body-wall; the diameter of the tube,
including its muscular coat, is 85 pw.
(7) Accessory (Copulatory) Glands.
The sexual animal is characterized by a number of markings,
each a transversely oval patch, slightly raised above the general
surface, with a circular groove in its centre; these are two. three
or four in number, may ‘be paired or unpaired, and may occur on
the ventral surface of segms. Vil, vill, 1x, and xii, or any of these.
Corresponding to each of these markings there j is found inter-
nally a small, almost spherical mass, attached to the body-wall by
a short stalk-like portion. On investigation by sections they pre-
sent the following characters (fig. 5):—
Each is about -42 mm. in diameter, and is solid, its centre con-
sisting of a mass of cells. The outer thin coat of peritoneum is
in places perceptible only with difficulty; it appears as an ex-
tremely fine membrane with slight thickenings here and there
which represent much flattened nuclei; in places it carries blood-
vessels. The muscular coat (m.) is very well marked—9 p thick ;
its fibres are derived from both the muscular layers of the body-
wall. There is no special (blue-staining) connective-tissue layer,
though there are traces of this tissue in between the muscular
fibres, as in the body-wall.
The cells which All up the interior of the organ (c.) are large
i ho} if fS) 5)
132 DR. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,
and stain a light pink; the nuclei, 3-4 u in diameter, do not stain
deeply; the cell contents are uniformly granular. Cell outlines
are present; the cells in the depth of the. organ appear polyg gonal
in section, while those nearer the mouth are elongated in the
direction of the neck, and at the mouth these are continuous
with the surface epithelium. It may be that all the cells of the
interior reach the surface.
The figure of one of these organs shows a remarkable similarity
toa simple type of spermatheca ; and it does not seem impossible
that spermathece may have arisen from such structures.
(8) Rudimentary (1.e., Incipient) Gizzards.
Invisible in dissection, but revealed in the microscopic examina-
tion of sections, is an incipient gizzard belonging to segm. X1
(fig. 4, rad.), and, more rudimentary still, one in x. Notable is
the thickening of the connective-tissue layer in the incipient
gizzards, outside the layer of the strengthened circular muscular
fibres.
(For references to literature see end of next article.)
Heplanation of Plate J.
.1. Nephridium of Drawida japonica, X70. coil, spiral coil; d., duct;
ep., epithelial lining of sac; Joop, twisted loop; m., muscular coat of sac ;
n., nephrostome; 2. c., nephrostomial canal; p., peritoneal coat of sac;
sac, oval sac; spin., spindle-shaped portion ; ¢7., trumpet-shaped expan-
sion of duct at junction with body-wall.
. 2. Section through portion of testis sac, X100. 6.v., blood-vessels ; e., cells in
sac; ep., epithelial lining of sac; f£, elongated epithelium of funnel ;
g., pink-staining ground-substance of sac contents; m., muscular fibres ;
m., nuclei in ground-substance ; sept., septum 9/10; p., peritoneal coat of
sac; ¢., testis; v. def., vas deferens; 2., yellow or brown chloragogen-like
erains and masses.
r, 8. Section of prostate, somewhat oblique, X50. The covering layer of flattened
peritoneal cells, if and where present, is not to be seen at this magnifica-
tion. ats., atrium in centre of prostatic cells, with epithelial and
muscular coats; wuc., nuclei of prostatic cells; prost., prostatic cells;
wv. def., vas deferens in wall of atrium ; w., vacuoles with central particle.
Fic. 4. Longitudinal section, rather oblique, through ovarian chamber and neigh-
bouring segments (semi-diagrammatic), 20. ann., annular chamber
round gut; d.v., dorsal vessel; ep., epithelium of cwsophagus; ~, female
funnel ; giz., muscular thickening of cesophageal wall, constituting the
gizzard of segm. xii; par., parietes, with epithelial, circular, and longi-
tudinal muscular layers; o.¢., ovarian chamber; rzd., rudimentary
vizzard of segm. xi; sac, ovisacs; 10/11, 11/12, 12/13, the "corresponding
septa.
Fig. 5. Section of accessory (copulatory) gland, X50. ¢., cells filling up interior
of gland; e.m., circular muscular Rabe ot body- ‘vals ce.m.’, two strands
of circular muscle traver sing plug of cells at mouth of gland ; ep., surface
epithelium ; m., muscular coat of gland (the peritoneal coat not visible at
» this magnification); sept., septum 9/10; ¢., trabecular (blue-staining)
connective tissue, the longitudinal muscular layer of the body-wall being
absent here.
Fig. 6. Transverse section through ovarian chamber (semi-diagrammatic), x12.
alim., alimentary canal; ann., annular chamner round gut; d.v., dorsal
vessel; o.¢., ovarian chamber; ova, masses of ova, cut tangentially to
attachment of ovary to anterior wall of chamber; paz., parietes; x., upper
limit of ovarian chamber; v.ch., ventral chamber; v.7.¢c., ventral nerve
cord; v.v., ventral yessel; y., yolk masses.
es
gg
Fi
gg
a
go
AND ZOOGEOGRAPHY OF INDIAN OLIGOCH ATA. WAS
VI.—On rue RELATIONSHIPS OF THE GENERA OF MONILIGASTRID#;
WITH SOME CONSIDERATIONS ON ‘tHE ORIGIN oF TERRESTRIAL
OLIGOCH ETA. —
(1) The Testis Sacs of the Oligocheta. .
The primitive condition of the male organs of the Oligocheta
is that seen in most of the lower families as well as in many
Megascolecide and Lumbricide. The testes are attached to the
anterior wall of their segment, the funnels to the posterior; the
sperm cells are shed into the ccelomie cavity at an early stage of
their development, and ripen in the general cavity of the
segment, or more usually in cxeal sacs, the seminal vesicles,
formed by bulgings of the septa (text-fig. 7). In this case there
are no testis sacs.
Text-figure 7.
as aes ;
SV t f ud.
Primitive condition; no testis sacs. f, male funnel; s.v., seminal vesicle
(shown in-seems. ix and xii); ¢., testis; v.d., vas deferens.
Not infrequently, however, it happens that the lower part of
the segment, where the testes and funnels lie, is cut off by a
partition from the rest of the cavity of the segment. ‘Ihe
Vext-figure 8.
The ordinary form of testis sacs, e.g. as in Lwmbricus. References as before ;
in addition, ¢.sac., testis sac (median seminal vesicle).
organs are then contained in special com partments of the ccelom,
the testis sacs—either a single one in each testicular segment,
134 DR. J, STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,
containing the organs of both sides, or a pair in each segment,
one for the or ans vot the right and one for those of the left side.
Thus in the common European genus Lumbsicus there is a single
testis sac (often called the median seminal vesicle) in each of the
testis segments ; while in many species of the common Indian
genera, Pheretima and Hutypheus, the sacs are paired (text-
fig. 8).
In the genus Ocnerodrilus, and in a few species of Enchytreus,
testis sacs of a different type are found. The sacs enclose the
testis, and are attached like the latter to the anterior septum of
the segment, projecting freely into its cavity; the funnels,
however, are outside the sac, and, as usual, on the posterior
septum of the segment; it is evident that the sac must rupture
before the spermatozoa can enter the funnel. In this case the
sac can hardly be anything else than the superficial layer of the
proliferating peritoneal cells which give rise to the gonad; this
layer separates off as a connected membrane, beneath which the
genital cells are shed and ripen (text-fig. 9)*. The lobes of the
Text-figure 9.
Testis sacs of Ocnerodrilus occidentalis. References as before.
testes of some species of Luwmbricillus are also (at any rate in
their young state) surrounded by a thin membrane, within
which the genital cells are cast off and undergo some of their
development.
Lastly, in all the Moniligastride the testis sacs are paired
vesicles, suspended by the septa in such a way that they project
into the segment in front or into the segment behind, or (more
usually) into both; the degree of projection forwards or
backwards varies in different individuals of the same species,
and sometimes on the two sides of the same individual. Within
each sac—usually perhaps on its floor towards the anterior wall—
is the testis, a proliferation of the epithelial lining of the sac;
the funnel is also included in the sac, and is situated on the
* T donot know of any observations on migration of genital cells in the Oligo-
cheeta. If such took place, it might be possible to interpret the sac wall in these,
cases as the peritoneum, raised up and distended by the ripening germ cells which
had taken up their position beneath it.
AND ZOOGEOGRAPHY OF INDIAN OLIGOCH ATA. 135
floor near the !testis, sometimes actually surrounding the
attachment of the testis as in Drawida japonica. The vas
deferens leads off from the funnel, and descends on the septum
to the ventral body-wall (text-fig. 10).
Text-figure 10.
Testis sac of Drawida. References as before; in addition, p7’., prostate.+ |
(2) The Testis Sacs of the Moniligastride considered as
Segments.
Leaving aside for the moment the Moniligastride, it will be
seen from the above that throughout the Oligocheta the testis is
morphologically on the posterior face of the anterior septum of
its segment, of the peritoneal covering of which it is a proli-
feration; and the funnel faces it on the anterior face of the
posterior septum, of the peritoneal covering of which it is a
modification. On now comparing the figure which represents the
condition in the Moniligastride (text-fig. 10) it is evident that
the most feasible explanation of the nature of the testis sacs of
this family is that which would regard them as the equivalent of
segments, of which the anterior and posterior walls have fused
together for the greater part of their extent.
The cavity of the sacs cannot be a mere split in the septum,
since it is lined throughout with epithelium. Following the
usual rule, the cavity must be ccelomic, and the epithelium
peritoneal epithelium.
In Syngenodrilus, to which reference will again be made in the
sequel, there is given off from the posterior testis sac a seminal
vesicle (text-fig. 16) in precisely the same way as the seminal
vesicle is given off from the testis segment in the more primitive
forms (cf. text-fig. 7).
Again, we have seen in the foregoing article how, in the
genera Moniligaster and Drawida, there is a tendency to shut off
the ovaries and their funnels in special chambers. These
chambers represent segment xi, and various stages in the gradual
limitation of the extent of this segment can be traced in actually
existing forms. The ovarian chamber contains the ovaries and
126 DR. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,
their funnels. and the alimentary canal is excluded. The testis
sacs have similar relations; and if the ovarian chamber were to
lose its nephridia, and to become divided into two, one for the
organs of each side, we should have an identical condition
reproduced. It is difficult to resist the conclusion that the testis
saes owe their origin to the same process which we can see and
follow in the case of the ovarian segment.
Beddard, so long ago as 1891 (6), suggested this origin for the
testis sacs ; he thinks they ‘“ may be all that is left of the ccelom
belonging to the segment which contains the testes.” He adds:
‘“« This is of course no more than a suggestion”; and, so far as I
know, the suggestion has never since been considered. But
I believe that the above considerations show with some degree of
probability that this is really the true nature of the saes.
(3) The Relationships of the Genera of Moniligastride.
In the present section I omit the consideration of the recently
discovered genus Syngenodrilus, to which reference will be made
later, and confine myself to the previously known genera of
the family.
The most primitive of the four genera is Desmogaster, in which
the sexual organs are arranged as shown in the accompanying
figure (text-fig.11). I use letters to indicate the several segments,
since It is easier thus to make comparisons between this and
other genera.
Text-figure 11,
7 CHR.) Lee eS on Si ovls
Sptn. L OSA MILE VEO. Of.
Desmogaster. 1.h., last heart; m.f.. male funnel; o., ovary; o.f., ovarian
funnel ; ovis., ovisac; spth., spermatheca; ¢., testis; ¢.sac., testis sac;
w.d., vas deferens.
There may be either one or two pairs of spermathece ; two
pairs are shown in the figure, but if only one pair is present it
may be either the anterior or the posterior.
Hupolygaster is derived from Desmogaster by the disappearance
of the posterior set of male organs and the anterior sperma-
thece (the anterior spermathecee are the complement of the
posterior male organs; these face each other when the worms
AND ZOOGEOGRAPHY OF INDIAN OLIGOCHAETA. 137
are apposed in copulation); otherwise the figures are identical
(text-fig. 12) :—
Text-figure 12.
Eupolygaster.
In Drawida and Moniligaster the figure is as below (text-
fig. 13).
Text-figure 13.
Drawida and Moniligaster.
The ovarian chamber corresponds to the ovarian segment in
the other genera; as in Hupolygaster, there is a single pair
of spermathece and a single set of male organs. But there is
a further reduction still—another segment has disappeared.
Assuming for the moment that the similarly lettered segments
in text-figs. 11 and 13 correspond, ¢ with cand g with g, it will
be seen that the single testis sac of Drawida is left to correspond
with the two testis sacs and the intervening segment e of Desmo-
gaster ; in other words, there has been a squeezing together of
the segments, and the two testis sacs have fused. This fusion
probably gave at first a single sac with a vertical diaphragm ;
later the diaphragm disappeared (cf. text-fig. 14, c). In thus
fusing, the testis sacs have annihilated the space e—in other
words, the cavity of the segment between them; the most
posterior heart, which was in this space, has also disappeared,
and hence in these two genera, Drawida and Moniligaster, the
last heart is in the segment in front of the testis sac. The
actually existing testis, funnel and duct of these genera may not
improbably be the testis of the original anterior sac and the
funnel and duct of the posterior.
Michaelsen (12) imagines the genus Drawida to have arisen in
a rather different way—by the disappearance of the anterior set
138 DR, J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,
of male organs of Desmogaster and the dislocation forwards of all
the other sexual organs. Though the difference between the two
views is not great, | prefer the above wording, for the following
reasons :-——
First, the approximation of the septa in the region of the
testes is observed in other genera ; it has been noted by Michael-
sen in some species of Acanthodrilus (15); it occurs in Hutypheus,
where septa 8/9-10/11 are usually very close together (testes and
funnels have, however, in this genus disappear ed from segment x);
it 1s seen also in Hoplocheetela' (Stephenson, 24), where septa 9/10,
10/11, and 11/12 may be so closely approximated as to appear at
first like one hugely thickened septum.
Next (and this ‘I am inclined to regard as important), the
process that I have assumed for Drawida is apparently seen in
progress in Syngenodrilus (cf. text-fig. 16), where the two testis
sacs are contiguous, indeed fused, and the intervening segment
has been squeezed out of existence ventrally, though a portion
persists dorsally.
Lastly, the trabeculee which Benham describes as crossing the
testis sac in Moniligaster indicus (= Drawida robusta) (10) may
possibly represent the remains of the septum which, on my view,
at first separated the cavities of the two sacs after they had come
together. It may be added that the existence of a rudimentary
second prostate, in front of that belonging to the normal male
apparatus, in Drawida willst (Michaelsen, 13), and of a similarly
situated well-developed second prostate in D). raui (Stephenson,
26), shows only that the actual funnel and vas deferens of
Drawida are the equivalent of the posterior pair of Desmogaster ;
the testis may quite well be the anterior testis, as I have supposed,
and the testis sac the product of fusion of the original two.*
(4) The Numbering of the Segments in Desmogaster.
The segments 7, g, h, f, e (text-figs. 11 & 12) correspond in Des-
mogaster and Hupolygaster; and on the view here advocated, 2
and f# correspond in Desmogaster and Drawida (text-figs. 11 & 13).
Now 7i-e are segments vili-xi, with the anterior testis sac, In
Desmogaster, and vii-x, with the testis sac, in Hupolygaster ;
i and f# are vill and ix in Desmogaster, vii and vili in Hupoly-
gaster and Drawida. That is to say, the whole of the sexual
organs are one segment further forward than they ought to be in
Hupolygaster and Drawida.
It would be easy arbitrarily to suppose that a segment in the
anterior part of the body had somehow been lost in “the ancestors
of these two genera, or, with Michaelsen, to assume a “ dislocation
forwards” of the sexual organs (which comes to much the same
* It may be said that if it is the posterior vas deferens which persists in Drawida,
the persistent: spermatheca should be the anterior and not the posterior. This is not
necessary, however; as already stated, there are species of Desmogaster with two
pairs or with only one pair of spermathece ; in the latter case, the single pair of
spermathecee may be the anterior or the posterior. In D. schildi it is the posterior ;
if such a form were to undergo the assumed squeezing together and fusion of the
testis sacs, the exact condition of Drawida and Moniligaster would be arrived at.
AND ZOOGHOGRAPHY OF INDIAN OLIGOCH ATA. 139
thing). But there is no reason to suppose that Hupolygaster and
Drawida had a common origin—the reverse in fact; and if they
originated separately, this curious dislocation must have occurred
twice independently.
I believe the explanation to be that the accepted numbering of
the segments in Desmogaster is incorrect, and that the numbers
given to all the segments should be diminished by one. This
view requires some justification.
There are three means of establishing the enumeration of the
segments at the anterior end of the body—the sete, which
commonly begin in segment 11; the septa, which are, however,
invariably wanting in the anterior segments ; and the annulation
of the body. Of the four species of Desmogaster, the sete are
invisible in the anterior segments in 2). dorie ; they do not begin
till segment vili (laterally not till ix) in D. schildi; they do not
begin till segment 1x in J). giardi ; and we have no information
concerning them in D. horsti, except that they are minute. The
sete are thus useless for the determination of segments.
As to the septa, in all species the extreme displacement of
the septa in the anterior part of the body is remarked by the
authors. In D. horsti the tenth septum (presumably the meaning
is septum 10/11) is either absent, or displaced backwards to the
level of 11/12; if we suppose it to be absent, the numbering of
the segments is brought up to that which is generally accepted
for the genus, while if we make no such supposition, the numbering
of the segments is apparently that for which I am contending.
Concerning the annulation, we only know that in D. horsti
“the two anterior segments have their longitudinal diameter
smaller than that of the succeeding ones”—which appears to
admit of the supposition that they really constitute a single
segment; and that in D. schildi segments i-iii are simple while
iv-xll are biannular. But since there are no sete near the
anterior end of this worm, the distinction of primary end secon-
dary grooves is conceivably somewhat arbitrary, and a slight
difference in estimating the value of the grooves would reduce
the number of segments by one. It is worth noticing that in a
species of Hupolygaster (HL. browni, 13) Michaelsen is in doubt
regarding the numbering of the segments because of the secondary
annulation, sete being absent here also from the anterior
- segments.
Naturally, when the numbering of the segments had once been
determined in the first species of Desmoyaster (rightly or wrongly),
subsequent writers describing new species would, if in any doubt,
bring their numbering to correspond, as Michaelsen does in the
example of the Hupolygaster just quoted *.
* The first species of Desmogaster to be described was D. dorie (Rosa, 19).
There seem to be inconsistencies in the data there given. ‘he ovaries are said to be
im segm. xiil; but the funnels to be on anterior face of septum 14/15—7.e., in sezment
xiv. The spermatheceze (there is only a single pair in this species) are first stated to
be in viii, but further on to be on the posterior face of septum 6/7—1.e., in vil. This
would seem to indicate that the estimation of the segments was not easy, and that
the author himself came to different results at different times.
140 DR. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,
Since, therefore, the criteria for estimating the numbering of
the segments in the anterior part of the body of Desmogaster
allow of results of only doubtful value, I believe we are justified
in taking a numbering which corresponds—allowing for the
evolution described above—with that of the other genera of the
family. In any case, if subsequent investigations, directed
specially towards this point, should show that the accepted
numbering is correct, I should prefer to account for it by
assuming an increase in the number of segments in the anterior
part of Desmogaster after the stocks of Hupolygaster and Drawida
had been given off, rather than by an independently produced
elimination of a segment, or a dislocation forwards of all the
organs, in these two genera.
The following figures show the relations of the three genera ;
Text-figure 14.
A. Desmogaster; B. Eupolygaster ; C. Drawida and Moniligaster.
The actual segments ave numbered, and the relations indicated,
AND ZOOGEOGRAPHY OF INDIAN OLIGOCH ATA. 141
the letters show the morphological correspondences, and the
roman numerals show the actual segments (revised as above im
the case of Desmogaster).
(5) The Relationship of Moniligaster to Drawida.
The genera Drawida and Moniligaster differ only in the fact
that Moniligaster has, and Drawida has not, connected with the
ectal end of the spermathecal duct, a bilobed glandular appendage,
of which each lobe consists of a number of closely-packed branches.
In Drawida there may be a swelling on the end of the duct, or at
most a relatively small sac-like atrial diverticulum.
Michaelsen (18) takes Drawida for the ancestor, Moniligaster
for the descendant; he traces the history of the at first simple
duct in Drawida. In some species it “widens and is transformed
into a muscular coat. In other species of this genus this widened
muscular distal (2. ¢., ectal) end grows into a real muscular atrial
chamber, which further on enlarges at one side to form a
separate blind sae, depending into the second segment, or, in the
species D. robusta and its subspecies, at two sides, forming two
outgrowths, one depending into the seventh segment, the other
into the eighth, the two outgrowths being separated by the
septum 7/8,-and the duct of the pouch entering the atrial
chamber at the angle between these two outgrowths. The
structure found in the species of Moniligaster may be compared
with this structure in D. robusta. The two outgrowths of the
atrial cavity in the latter species are continued in Moniligaster
each into a large gland, consisting of a large, much-branched
tube, the branches and twigs of which are packed together and
enveloped in a peritoneal membrane.”
This derivation does not seem easy tome; I should prefer to
read the series (if indeed it is a series) in the reverse direction.
T cannot see in Moniligaster any atrial chamber at all—only two
glands, with ducts which unite, like the limbs of a Y, to form a
single canal. This is well shown in Perrier’s original plate (18) ;
my own description of the condition in Perrier’s species 1s
similar (22):—‘ The stalk (atrium) is relatively narrow, and
pifurcates, one branch going to each half of the mass. Neither
the stalk nor its two branches are in any sense sacs; the
appearance is that of two ducts proceeding one from each half of
the mamillated glandular mass, which unite to form a common
duct; this common duct is less than half as long as the glandular
part and nowhere dilated.” Itseems difficult to imagine a
slight swelling of the end of the duct dilating, becoming a sac,
bifureating, and then sprouting glands from each horn, itself
concomitantly narrowing to a Y-shaped duct. If the series
really is a natural one, it would be easier to read it the other way —
as a gradual reduction, of the glands toa non-functional vestige
and their final disappearance. It is possible that the glands ave
entirely independent of the spermathece in origin; compare
142 Dh. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,
the accessory spermathecal glands in Drawida schunkarai
(Michaelsen, 16); similarly there are associated yet independent
glands in the genera Hoplochetella and Erythreodrilus, belonging
to another family, the Megascolecide.
(6) Lhe Origin of the Moniligastride.
If the testis sacs represent segments, they must have existed
as such in the ancestors of the present genera. Let us take
Desmogaster, as the most archaic of existing genera of Monili-
gastride, and expand the testis sacs into segments; we get a
form which may be represented as in the accompanying figure
(text-fig. 15).
Text-figure 15.
Desmogaster, with the testis sacs expanded to appear as segments.
Now let us take the remarkable genus Syngenodrilus, described
by Smith and Green (20), and separated by them as a distinct
subfamily. The actual arrangement of the sacs and septa is
approximately represented in the subjoined figure (text-fig. 16),
which is a diagrammatic copy of the section illustrated in the
original paper.
Text-figure 16.
Vil VIM
Syngenodrilus. The external segmentation is indicated. Pv., prostate.
Note the three pairs of prostate glands, which, however, are none
of them in close association with the male ducts or pores; it is
quite possible, therefore, that these prostates of Syngenodrilus
have nothing to do with the structures that go under the same
name in the other genera of the family. If we expand this
figure, so as to separate the testis sacs (which are close up against
AND ZOOGEOGRAPHY OF INDIAN OLIGOCH2TA. 143
one another) and correct the extreme displacements of the septa,
we get the following (the seminal vesicle may be neglected, as being
only a diverticulum of the posterior testis sac) (text-fig, 17).
Text-figure 17.
Syngenodrilus, with the testis sacs separated from one another.
The condition is similar to that in Desmogaster, except that the
spermathece are a segment further back, and that there are three
pairs of prostates. If now we expand the testis sacs into segments,
we get a form with the following characters (text-fig. 18) :—
Text-figure 18.
Syngenodrilus, with the testis sacs expanded so as to appear as segments.
The figure is identical, except for the characters already mentioned,
with that of the ‘“‘ expanded ” Desmogaster.
Probably neither text-fig. 15 nor 18 represents a form that
actually existed. In the first place, there is a gap of a segment
between the two pairs of testes; and in the second place, the
apertures of the spermathecze are separated by only one segment,
while those of the male apparatus are separated by two segments,
and hence the spermathecal apertures would not coincide with
the male pores of another worm in copulation.
It has been pointed out by Michaelsen (12) that in all the lower
families of Oligocheta the gonads follow each other in successive
segments; the ovarian segment immediately follows the testicular
segment or segments; and the normal arrangement in the higher
families (testes in segments x and xi, ovaries in xiii) is to be
explained by the dropping- -out of a pair of ovaries in xii. The
original condition—two pairs of testes followed by two pairs of .
ovaries—still exists in the genus Haplotuxis; and hence Michael-
sen (12, 17) and Beddard (9) look on this genus as the ancestor
of the higher Oligocheta.
144 DR. J, STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,
To get a complete series of gonads in the ancestors of the
Moniligastride, however, we must supply a third pair of testes
in the vacant segment between the two pairs already present, as
well as a pair of ovaries between the last testicular segment and
the actual ovarian segment, ‘That the insertion of the third pair
of testes is Justified is seen from the presence of three pairs of
prostates in Syngenodrilus. One further addition is to be made—
three pairs of male pores imply three pairs of spermathecal pores
apposed tothem in copulation. The diagram of the ancestor thus
becomes that shown in text-fig. 19.
Text-figure 19.
The ancestral terrestrial Oligochzte, with complete series of gonads and
spermathece.
The above diagram probably represents an animal in which as
yet the moniliform series of gizzards characteristic of the family
had not been evolved. The gizzards are in front of the testis
segments in Syngenodrilus, behind them in the other genera:
that is, the muscular wall of the esophagus has been strengthened
in one place in one subfamily, in another place in the other.
Syngenodrilus (text-fig. 16) is derived from the above form by
a contraction, or huddiing together, of the testis segments in
such a way that the middle one is almost squeezed out of
existence and the other two become testis sacs, by a corresponding
reduction of the number of spermathece to two pairs opening in
7/8 and 8/9, and by the disappearance of the anterior pair of
ovaries with their funnels and ducts.
Desmogaster (text-fig. 11) is derived from the common ancestor
by the contraction of the segments of the first and third pairs
of testes to form testis sacs, and the disappearance of the middle
pairs of male organs; by a corresponding diminution in the
number of spermathece, the remaining pairs cpening in 6/7 and
7/8; and by the disappearance, as in Syngenodrilus, of the
anterior ovary with its funnel and duct.
Eupolygaster (text-fig. 12) came into existence by the further
disappearance of the posterior pair of male organs and the anterior
pair of spermathece of Desmogaster.
Drawida and Moniligaster (text-fig. 13) originated from a form
similar to Desmogaster by a contraction or huddling together of
the testis segments like that in Syngenodrilus, but carried further,
AND ZOOGEOGRAPHY OF INDIAN OLIGOCH ATA. 145
so that the testis sacs fuse, extinguishing the intervening
segment; the spermathece are again reduced to one pair.
(7) Lhe Origin of Terrestrial Oligocheta.
From an ancestor having the characters illustrated above
(text-fig. 19), then, we can derive the various genera of Monili-
gastride without, as it seems to me, forcing the facts in any way,
and relying only on processes which can be illustrated from
actual examples of the genera themselves or of other families of
Oligocheeta.
I was, however, surprised, and I must confess to feeling
pleased, to find that the diagram arrived at as above gives in a
very natural way a starting-point for the other families of
terrestrial Oligocheta as well. These all agree in having,
primarily, two pairs of testes in segments x and xi, and one pair
of ovaries in xiii; the primary number of spermathece in the
Megascolecide is two pairs, opening in grooves 7/8 and 8/9. For
the Glossoscolecide and Lumbricide it is not so easy to give the
primitive position of the spermathece ; perhaps all that can be
said is that the Glossoscolecine, probably the ancestral group of
these two families, typically have spermathece which open in
front of the testis segments (Michaelsen, 17).
The arrangements of the gonads and their funnels im the
higher families is thus obtained simply by the disappearance of
the posterior pairs of testes and ovaries respectively ; and the
arrangement of the spermathecee—in the Megascolecide at any
rate—is arrived at by the disappearance of the anterior of the
three pairs of the ancestral form (the anterior pair of sperma-
thece would correspond, as already explained, to the posterior
pair of male openings). The primary position of the spermathece
in the Glossoscolecides and Lumbricide is, as has just been
indicated, scarcely ascertained with precision.
The occurrenee, in existing earthworms, of gonads or of traces
of gonads, in the segments in which, on the present theory, they
are supposed to have existed regularly in the ancestor, confirms
the view here taken. Woodward (28) has shown that in the
common English species the presence of additional pairs of
ovaries in segment xiv is muca commoner than has been
supposed, and that additional testes are not rare. There are
also cases where the supernumerary ovaries are not limited to
one pair; Woodward (27) examined a specimen with seven
pairs.
Confining ourselves, however, to cases where the condition is
definitely that of the presumed ancestor—one extra pair of
ovaries in segment xlv—we have the following records :—Bergh
(quoted by Woodward) described three cases in species of Laum-
bricus. Woodward's own investigation shows nine cases of this
condition out of fifty worms examined. Beddard, in an
examination of a number of specimens of Perronya excavatus GUE
Proc. Zoou. Soc.—1922, No. X. 10
146 DR. J.STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,
found a number of cases where there were additional female
pores one segment behind the normal, and where therefore there
were probably additional ovaries and oviducts internally to
correspond.
The testes which we suppose to have disappeared from
segment xii also sometimes crop up again. Thus Woodward (28)
found testes here in three cases out of fifty examined. Beddard
(4) found a pair of testes in segment xii in Pontoscolex corethrurus
(in this case the gonads of segment xiii also appeared to be
testes).
On the other hand, the gonads of segment xii, which ought,
when they reappear (if the common ancestor of earthworms was
constituted as above supposed), to be testes, are when present
sometimes ovaries. Beddard (/.c.) found this to be so in some
examples of Pontoscolex; it is so (if we may draw conclusions
from the position of the female pores) in some specimens of
Perionyx excavatus (Beddard, J. c.); and Woodward had six cases
of this condition out of the fifty specimens of British earthworms
which he examined.
In a number of cases the nature of the gonad in segment xii
is indeterminable, or the evidence is inconclusive. Beddard
(l.c.) believes that the same gland in Pontoscolex may produce
either ova or spermatozoa. In one specimen examined micro-
scopically by Woodward the gland was actually hermaphrodite.
In three very young specimens (? of Lumbricus rubellus) a gonad
was present on each side in segment xii, but at that stage could
not be identified as definitely testis or ovary (Woodward suggests
that this pair is always developed in the embryo). Beddard
(5, 8) found this pair of glands present in embryos of Octochetus
multiporus, but of course could not identify it, at that stage, as
either testis or ovary. Beddard also notes (9) that two pairs of
ovisacs have been found both by Horst and himself in certain.
species of Perichetide, in segments xiii and xiv; these would
correspond to ovaries in xii and xi; Beddard remarks on
certain resemblances of these organs to sperm-sacs, and, since
neither gonads were found in segment xli nor ova in the corre-
sponding sacs, it seems not improbable that the organs in
segment xiii were vestigial sperm-sacs, and the gonads which
formerly existed in segment xii were testes.
There is thus no doubt that ovaries occur not infrequently at
the present day in segment xiv, and that segment xii also
frequently contains a pair of gonads, which in some cases are
ovaries, in some cases testes, and in some cases their nature is
equivocal. According to our view they were, in the ancestral
earthworms, regularly testes.
The more usual view of the origin of the terrestrial Oligocheta
(Beddard, 9; Michaelsen, 12, 17) is rather different. The
Phreoryctide, with two pairs of testes and two pairs of ovaries, in
AND ZOOGEOGRAPHY OF INDIAN OLIGOCHATA. 147
segments x—xi and xii-xiii respectively. and with an oesophageal
gizzard, are held to be the present-day representatives of the
common ancestor. The Phreoryetide are probably in turn to be
derived from the Lumbriculidee, where the number of gonads is
variable, and also their position; Lamprodrilus satyriscus has
three or four pairs of testes, and other forms—e. g., Lumbriculus
variegatus—have more than one pair of ovaries.
But on the above hypothesis of the origin of the terrestrial
families of Oligocheta we require as our starting-point a form
with a series of five pairs of gonads—three pairs of testes and two
pairs of ovaries; while the discovery of Syngenodrilus makes it
probable that the ancestor of the Moniligastride had no special
thickening of either cesophagus or anterior intestine—the gizzards
developed independently in the two subfamilies, in the Syn-
genodriline in front of and in the Moniligastride behind
the testicular segments. It is obvious, therefore, that the
Phreoryctids are already too specialized for our purpose; the
ancestral form as here conceived has a larger number of gonads,
and is without an oesophageal gizzard. The ancestral group more
nearly resembled the Lumbriculide than the Phreoryctide; the
latter are not in the direct line of descent of the earthworms.
The Moniligastride, which, as Beddard has repeatedly insisted,
vetain many of the characters of the Limicoline section (clitellum
one cell only in thickness, non-union of the vasa deferentia of the
same side, vasa deferentia passing over only one segment), arose
independently of the other earthworms by the disappearance of
testes from segment xi and of ovaries from xili, and the con-
traction of the original segments x and xii to form testis sacs;
while the common stem of the Megascolecide, Glossoscolecide,
and Lumbricide originated by the disappearance of testes from
xii and ovaries from xiv, the lengthening of the vasa deferentia,
aud the union of those of the same side.
References to Literature.
1. Bepparp, F. E.—Descriptions cf some new or little-known Farthworms,
together with an account of the Variations in Structure
exhibited by Perionyx excavatus K.P. Proc. Zool. Soc.,
1886, p. 298.
2. a Note on the Reproductive Organs of Moniligaster. Zool.
Anz., ix. 1887, p. 679.
3. 44 On the Structure of three new species of Harthworms... .
Quart. Journ. Mic. Sci., xxix. 1888, p. 119.
4. bs On certain points in the Structure of Urocheta .P., and
Dichogaster, nov. gen., with further remarks on the
Nephridia of Earthworms. Quart. Journ. Mic. Sci.,
xxix. 1888, p. 235. ;
5, as On the Homology between Genital Ducts and Nephridia in
the Oligocheta. Proc. Roy. Soc., xlviii. 1890, p. 452.
6. ' Observations upon the Structure of a Genus of Oligocheta
belonging to the Limicoline Section. Trans. Roy. Soc.
Wdin., xxxvi. 1891, p. 7.
10#
148 ON INDIAN OLIGOCH@TA,
7. Bepparp, Ff. H.—On some new species of HKarthworms from various parts of
the World. Proc. Zool. Soc., 1892, p. 693.
8. 5 Researches into the Embryology of the Oligocheta. I. On
certain points in the Development of Acanthodrilus
multiporus. Quart. Journ. Mic. Sci., xxxiii. 1892, p. 497.
9. A Monograph of the Order of Oligocheta. Oxford, 1895.
10. Bile W. B.—Description of a new species of Moniligaster from India.
Quart. Journ. Mie. Sci., 1893, p. 363.
11. Bourne, A. G.—On Moniligaster grandis A. G. B., from the Nilgiris, S.
India.... Quart. Journ. Mic. Sci., xxxvi. 1894, p. 316.
lla. Cognerrt pE Marrttts, L.—A Contribution to our knowledge of the Oligo-
cheta of Travancore. Ann. Mag. Nat. Hist., ser. 8, vii.
1911, p. 494.
12. Micwartsen, W.—Die Geographische Verbreitung der Oligochaten. Berlin,
1908.
13. k The Oligochseta of India, Nepal, Ceylon, Burma, and the
Andaman Islands. Mem. Ind. Mus., i. 1909, p. 101.
14. nd Die Oligochitenfauna der vorderindisch-ceylonischen
Region. Abh. Naturw. Verein Hamburg, xix. 1910, p. 1.
15. Die Oligochaten von Neu Caledonien und den benachbarten
Inselgruppen. In: F. Sarasin und ‘I’. Roux, Nova Cale-
donia, A, Zoologie, Bd. 1. Wiesbaden, 1913, p. 171.
16. sy Oligochaten von Travancore und Borneo. Mt. Mus. Ham-
burg, xxx. 1913.
17. 6 Die Lumbriciden, mit besonderer Berticksichtigung der
bisher als Familie Glossoscolecide zusammengefassten
Unterfamilie. Zool. Jahrb., Abt. f. Syst., xli. 1917, p. 1.
18. Perrier, E.—Recherches pour servir 2 |’ Histoire des Lombriciens terrestres.
Nouv. Arch. Mus. Paris, viii. 1872.
19. Rosa, D.—Moniligastridi, Geoscolecidi ed Hudrilidi (Viaggio di Leonardo Fea
in Birmania e Regioni vicine). Ann. Mus. Genova,
ser. 2 a, ix. (xxix.), 1890, p. 368.
20. Suita, F., and Brsstz R. Green.—Descriptions of New African Earthworms,
including a new genus of Moniligastride. Proce. U.S.
Nat. Mus., lv. 1919, p. 145.
21. SrrpHEnson, J.—Oligocheta, in: Zoological Results of the Abor Expedition,
1911-12. Ree. Ind. Mus., viii. 1914, p. 365.
22. fs On some Indian Oligocheta, mainly from Southern India
and Ceylon. Mem. Ind. Mus., vi. 1915, p. 35.
23. er On a Collection of Oligocheta belonging to the Indian
Museum. Ree. Ind. Mus., xii. 1916, p. 299.
24. ra On a Collection of Oligochzeta from various parts of India
and Further India. Ree. ind. Mus., xiii. 1917, p. 353.
25. 53 On a Collection of Oligocheta from the lesser-known parts
of India and from Eastern Persia. Mem. Ind. Mus,,
vil. 1920.
26. 7p Oligochzta from Manipur, the Laccadive Islands, Mysore,
and other parts of India. Rec. Ind. Mus., xvii. 1921.
27. Woopwarp, M. F.—Description of an abnormal Earthworm possessing Seven
Pairs of Ovaries. Proc. Zool. Soc., 1892, p. 184.
28. 3 Further Observations on Variations in the Genitalia of
British Karthworms. Proc. Zool. Soc., 1913, p. 319.
ON THE VAGUS NERVES OF HYRAX CAPENSIS. 149
8. On the Vagus and Sympathetic Nerves of Hyrawx capensis.
By Caaries F. Sonnvrac, M.D., F.Z.8., Anatomist to
the Society.
[Received January 23, 1921; Read February 21, 1922.]
(Text-figures 6-8.)
The present paper is based on the examination of several speci-
mens of Hyrax capensis, both immature and fully adult, which
died in the Society’s Gardens. Variations were observed in the
branches of their vagus and sympathetic nerves; and the con-
ditions present in some of the animals are more complex than
those already described in my papers on the Marsupialia (8, 4)
and Kdentata (5). ‘They are also more complex than those in
many animals belonging to other orders.
The anterior cervical parts of the nerves are placed deeply, and
it is necessary to remove the wide ascending rami of the mandible
to gain access to them and their branches.
In all examples the cervical parts of the vagus and sympathetic
nerves are fused on both sides to form vago-sympathetic cords,
the union taking place at the level of the middle of the thyroid
cartilage. The cords are resolved again into their component
elements at the root of the neck, but the separation is usually
higher on the left side. After they have parted, however,
branches of communication may run between them.
Tar Vagus NERVES.
Course:—As it emerges from the foramen lacerum posticum
each vagus has the usual relations to the glosso-pharyngeal, spinal -
accessory and hypoglossal nerves, and it communicates with them
and the superior cervical ganglion by well-marked branches
(text-fig. 6. C1x, Cx1, Cxir and S.0.G). No ganglion nodosum is
present in the neck, as in Zamandua tetradactyla (4), or within
the foramen lacerum posticum. I+ then courses postero-mesially,
gives off communicating branches to the cervical plexus (C.C.P)
and unites with the cervical sympathetic to form the vago-
sympathetic cord (V—S) at the level of the middle of the thyroid
cartilage. And the nerve up to this point is internal to the
mandible. The vago-sympathetic cords break up again in the
posterior third of the neck; the vagus half runs still postero-
mesially and enters the thorax. Owing to the high level reached
by the heart and aorta the first part of the intra-thoracic course
is short, but the calibre of the left nerve remains thicker than
that of the right one.
Behind the roots of the lungs the vagi are united by a sinuous
cord which is as thick as the left vagus (text-fig. 7, br), and the
nerves emerge again from its extremities, so the arrangement is
H-shaped. The left vagus is replaced by vight and left branches.
150 DR. C. F. SONNTAG ON THE VAGUS AND
The former runs along the ventral border of the cesophagus,
passes through the cesophageal opening in the diaphragm and
ramifies over the anterior and ventral aspects of the body and
fundus of the stomach. The latter runs along the dorsal border
of the cesophagus and unites with the right vagus which deseribes
a wide curve to meet it; and the combined trunk ends on the
dorsal aspect of the stomach. Numerous communicating branches
link the vagi to one another and to the cceliac plexus, and some
go to the duodenum.
Text-figure 6.
The Cervical and Upper Thoracic Parts of the Vagus and Sympathetic Nerves
of Hyrax capensis. V: vagus; S: sympathetic; Mn: line of mandible.
Other letters in text.
Branches :—Numerous branches of communication and distri-
bution are again given off, many of which are very complex,
~ The pharyngeal nerves (text-fig. 6,a) are given off from the
trunk of the vagus and the loop (C) connecting its superior and
recurrent laryngeal branches. They form a plexus with branches
of the glosso-pharyngeal nerve and sympathetic. In some speci-
mens the laryngeal nerves do not form a loop which gives off
pharyngeal branches.
SYMPATHEDIC NERVES OF HYRAX CAPENSIS. 151
The superior laryngeal nerve (6) is replaced by a strong cord
which unites with one of the divisions of the recurrent nerve.
This cord gives off internal laryngeal (7./.n), two external laryn-
geal (eli), two thyroid (¢.y.n), and three external pharyngeal
(a) nerves. Small branches (v.n) accompany the common carotid
artery. The internal laryngeal branch enters the larynx through
the thyro-hyoid interval.
The left recurrent nerve (e) is given off in the common position.
It encircles the aortic arch and runs antero-laterally through the
neck. In the middle of the neck it shows a fusiform expansion
(1) and divides into a mesial nerve (R.L.N), which is the true
inferior laryngeal nerve, and a lateral cord (C) which unites with
the superior laryngeal nerve to form a loop, whose upper part
has been described above. Numerous tracheal (TN) and cesopha-
geal nerves (ON) are given off. They form plexuses of which the
tracheal one is the more superficial. And the esophageal plexus
anastomoses with the cesophageal branches of the right recurrent
nerve (d). The latter also forms a loop with the right superior
laryngeal nerve, but there is no fusiform expansion. And both
recurrent nerves give off cardiac branches (C.N) which are thicker
and more numerous on the left side.
In some specimens these loops are absent, and there is no
external communication between the superior and recurrent
laryngeal nerves.
The existence of anastomoses between the laryngeal nerves 1
Man has long been known. And Landois and Stirling (2) sum-
marised our knowledge as follows :—‘“ A connecting branch runs
from the superior laryngeal to the inferior (the anastomosis of
Galen), which occasionally gives off sensory branches to the upper
half of the trachea (sometimes to the iarynx?%); perhaps also to
the cesophagus (ZLonget), and sensory fibres (?) for the muscles of
the larynx supplied by the recurrent laryngeal. According to
Francois Frank, sensory fibres pass by this anastomosis from the
recurrent into the superior laryngeal. According to Waller and
Burekhard, the motor fibres of both laryngeal nerves are all de-
rived from the accessorius, while Chauveau maintains that the
crico-thyroid is an exception.” In a recent paper Dilworth (1)
described and figured a well-marked branch running between the
recurrent and internal laryngeal nerves, and says: “*.... the
laryngeal nerves are really a plexus of nerves. Just as the vagus
breaks up into its various plexuses in the body, it does the same
in the larynx. It isa highly modified plexus. I would further
suggest that it arose by the larynx separating a strand of fibres
from the vagus—that this strand is represented by the continuous
nerve joining the internal and recurrent laryngeal, and that the
separation from this strand of further fibres forms the various
nerves of the larnyx.” The conditions shown in text-fig. 6 sup-
ort Dilworth’s views, and I would extend his views of the origin
ot the laryngeal nerves from the continuous strand to account for
the origin of the tracheal, cesophageal, and thyroid nerves.
152 DR. C. F. SONNTAG ON THE VAGUS AND
The Cardiac Nerves (text-fig. 6,7) arise on the left side from the
vagus distal to the origin of the left recurrent nerve, but the
right ones arise from the vagus and.its recurrent branch. They
run to the cardiac plexus wherein they become associated with
the sympathetic (c.B.s). .
The wsophageal nerves (text-figs. 6, on, 7,0N, and 8, vB) arise in
the neck from the cords uniting the laryngeal nerves. In the
upper part of the thorax they are branches of the right vagus,
but they come from the two divisions of the left vagus distal to
the roots of the lungs.
The tracheal nerves (text-figs. 6,1.N and 7,1.N) arise from the
laryngeal cords in the neck, but its terminal part and the main
bronchi receive their nerve-supply from the thick bridge between
the vagi (67).
Text-figure 7.
The Lower Thoracic and Abdominal Parts of the Vagus Nerves of Hyraa
capensis. L.V: parts of left vagus; R.V: right vagus. Other letters
in text.
The pulmonary plecuses (text-fig. ight one is
formed by two branches from the ee vagus and communi-
cating twigs from the deep cardiac plexus. But the left one
consists of many branches from the bridge between the two vagi.
The latter sends a branch to the aortic plewus (A.P), which receives
many twigs from the left vagus. And no ganglia were found in
the pulmonary and aortic plexuses.
The Gastric Nerves (text-fig. 7):—The left vagus reaches the
stomach along the ventral border of the cesophagus and breaks
up into two branches. The first gives off twigs which run along
SYMPATHETIC NERVES OF HYRAX CAPENSIS. 153
the lesser curvature, and the latter supplies the fundus. Both
groups anastomose with branches from the right vagus, and with
the celiac plexus. The cord formed by the right vagus and part
of the left one supplies the dorsal aspect of the stomach almost to
the pylorus. Its branches anastomose with those of the lett
vagus, and twigs from the celiac plexus. A well-marked branch
(splanchnic?) runs to the left sympathetic cord (A). Many of
the cesophageal nerves run posteriorly and anastomose with both
vagi, so there is a close network round the lower end of the
cesophagus and over the left part of the stomach.
Tur SyMPATHETIC NERVES.
The Superior Cervical Ganglion (text-fig. 6,8.c.¢) is round or
oval, and flat. It gives off an internal carotid nerve of con-
siderable length (1.c.N) which accompanies the internal carotid
artery into the skull. The nerve communicates with the glosso-
pharyngeal and hypoglossal nerves, and the ganglion is connected
to the vagus, cervical plexus, and loop between the superior and
recurrent laryngeal nerves. But no connection exists between
either and the spinal accessory nerve. From the posterior pole
of the ganglion the sympathetic cord runs laterally and joins the
vagus at the level of the middle of the thyroid cartilage to form
the vago-sympathetic cord.
In Ayraaz, as in all animals possessing the vago-sympathetic
cord, no middle cervical ganglion is present, and no direct rami
communicantes run to the middle cervical nerves.
The sympathetic separates again in the posterior part of the
neck, but communicates with the vagus after the partition. It
exhibits a well-marked inferior cervical ganglion (I.C.G) on the
left side, but none on the right. From the ganglion branches of
communication run to the brachial plexus, but I was unable to
detect any communication between the right sympathetic and
brachial nerves, or between either cord and phrenic nerves. The
right sympathetic gives off a branch which accompanies the
vertebral artery (V.A.N), and a loop connects it to the right
recurrent nerve. And as no branch runs directly to the cardiac
plexus, the right recurrent and right vagus nerves conduct all
the right sympathetic filaments to the heart.
The left inferior cervical ganglion gives off a medium-sized
cardiac branch (c.B.8) which runs almost parallel to the left
recurrent nerve, passes to the dorsal aspect of the aortic arch and
ends in the deep cardiac plexus.
The Annulus of Viewssens (A.V) is present only on the left
side, and the right sympathetic cord passes in front of the right
subclavian artery.
The left sympathetic thoracic cord (text-fig. 8, u.s) has few
ganglia in its anterior part, and it forms loops. At the level of
the middle of the root of the left lung it divides into lateral and
mesial divisions. The former possesses a loop anda small ganglion,
154 DR. C. F. SONNTAG ON THE VAGUS AND
and is continued as a thick nerve which ends in the celiac gang-
lion (text-fig. 8, c.¢). The latter ends in a large ganglion whence
two nerves emerge. One joins the lateral division and the other
becomes the gangliated cord of the sympathetic. In the thorax
it gives branches to the aortic plexus(A.P) and a long branch runs
to the ceeliac ganglion. The lateral division of the cord, and the
eceliac branch of its mesial half constitute splanchnic nerves
(Sp.N).
Text-figure 8.
The Thoracic and Abdominal Sympathetic of Hyrax capensis. V.B: aortic
and cesophageal branches of the vagi; L.D: line of diaphragm. Other
letters in text.
The abdominal part of the left sympathetic (text-figs. 7 and 8)
possesses four ganglia. It gives off two groups of lateral branches
to the semilunar ganglion, rami communicantes to the lumbar
nerves, and branches to the aortic plexus. It has a great ten-
dency to subdivision.
The thoracic part of the right sympathetic (text-fig. 8, R.s) has
few ganglia in the anterior part, and its posterior part is very
complicated, but not so much as the left cord. It divides into
mesial and lateral parts and these are united at intervals by
common ganglia or communicating branches. The lateral
division ultimately continues the cord back to the sacrum. The
mesial division is thick and strong. Branches of the cord run to
SYMPATHETIC NERVES OF HYRAX CAPENSIS. 155
the aortic plexus (A.P), vena azygos major (Az.N), thoracic nerves,
and left sympathetic.
Jn the abdomen the mesial division divides into two, and the
following are the fibres of distribution :—
. To the eceliac ganglion (A).
. Hepatic plexus (contained in A).
. Phrenic plexus (contained in A).
. Fibres to the right renal plexus (R.R.P),.
. Fibres to Meckel’s Tract and Duodenum (M.B).
Fibres to the colon (C.B).
The abdominal part lies close to the left sympathetic and fibres
connect them. It gives off the right renal plexus (R.R.P), right
spermatic plexus (R.S.P), and filaments to the aortic plexus.
The Cardiac Plexus consists almost entirely of the deep part,
the superficial plexus consisting only of a few filaments derived
from it. ‘The nerves entering into it are :—
1. Three branches of the left vagus.
2. A twig from the left recurrent nerve.
3. A branch from the inferior cervical ganglion of the left
sympathetic.
4, Two branches from the right reeurrent nerve.
5. Two branches from the right vagus.
No ganglia are present, nor are there separate depressor nerves ;
but these may be contained in the loops uniting the superior and
recurrent laryngeal nerves. Oftshoots of the plexus can be traced
into the pulmonary and tr: acheal plexuses.
The Aortic Plexws (A.P) derives its fibres from the two sympa-
thetic cords, the bridge between the vagi in the thorax, and the
solar plexus.
The Solar Plexus (text-fig.7):—In the upper part of the
abdomen dorsad of the stomach there is a broad band of fibres
with a large reddish-brown semilunar ganglion at its left
extremity. The fibres are closely packed and have wide con-
nections as foilows :—
1. The splanchnic nerves from the left sympathetic cord
entering the anterior pole of the ganglion (A).
2. Two bundles of fibres from the left sympathetic cord
entering the mesial border of the ganglion (B).
3. Fibres from the right vagus to the large left splanchnic
nerve (C).
4. Fibres from the right vagus to the semilunar ganglion (D).
5. Fibres from the left vagus to the semilunar ganglion (E).
6. Fibres from the right vagus running into the plexus and
turning down to the splenic plexus (F).
7. Fibres from the semilunar ganglion to the splenic plexus (G).
8. Branches from the solar to the splenic plexus (1H).
9. Branches from the right sympathetic to the semilunar
ganglion (1).
10. Hepatic plexus (J) giving off pyloro-duodenal nerves (K).
11. Diaphragmatic plexus (L).
HOU Co bo
156 ON 'THE VAGUS NERVES OF HYRAX CAPENSIS,
SuMMARY AND CONCLUSIONS.
1. In all examples of Hyrax capensis the cervical parts of the
vagus and sympathetic nerves are fused. And the ganglion
nodosum is frequently absent in the neck.
2. The recurrent and superior laryngeal nerves are frequently,
but not always, connected by a loop.
3. There is no separate depressor nerve.
4, The internal carotid nerve has a very long cervical course.
5. The right recurrent nerve has well-marked cardiac branches.
6. The left sympathetic alone has an inferior cervical ganglion
and an Annulus of Vieussens.
7. The posterior thoracic parts of the vagi have a complicated
arrangement.
8. The thoracic sympathetic cords have few ganglia, and there
is only one semilunar ganglion.
9, The right sympathetic is distributed to the colon, and the
left one and cceliac plexus supply the small intestines.
BIBLIOGRAPHY.
1. Ditwortu, T. F. M.—Journal of Anatomy, 1921, pp. 48-52.
2. Lanpots and Srrrtinc.—A Text-book of Human Physio-
logy, vol. ii.
3. Sonntac, C. F.— Proc. Zool. Soc. London, 1921, pp. 572-575.
4. 3 Proc. Zool. Soc. London, 1921, pp. 873-876,
5. - Proc. Zool. Soc. London, 1922, pp. 99-180.
#
‘SINZOILIV SINOUHODOIdVH
‘dur yany ‘UTL F8 Tap weet 'P
AL Wel IN oka Gia Gillis) “arta
Nad
ae
m4
SS Gh GUN etal se a IN Ose Gta ve
‘dit yyny ‘WMT ZO Tap users Pp
NS
\
\
SUI UciON Vane ttie:G: Se SE)
at
Chatien
Srone:
Seer
ES es
ie
eo
ae
vate
e
ie)
- ‘OQNHIVNDSOWOVN HS Xd LNaAd SINOYUHOOTdVH T
-durr w4ny :
‘USTT 49 TSP User 'C
Title NV Daa oGcels 42d
ae
ST LTeOeroONOT Hw VOUlvol love SstNOadn oO rary hee :
‘dart HFT FO TSP Wear ©
Atic NVOae 6cetS 4c
ON THE CICHLID FISHES OF LAKE VICTORIA, Lae
9, The Cichlid Fishes of Lake Victoria. By C. Tarn Rraan,
M.A., F.RS., F.Z.8., Keeper of Zoology in the British
Museum (Natural History).
[Received January 6, 1922: Read February 21, 1922. |
(Plates I-IV.*; Text-figures 1-14.)
A revision of the Cichlide of Lake Victoria is a much more
difficult task than that of revising the Cichlide of Nyassa,
inasmuch as the types of all the described species from Nyassa
are in the collection of the British Museum (Natural History),
whereas a number of the types from Lake Victoria are in Paris,
Berlin, and Genoa.
Dr. Pellegrin has kindly sent me notes on two or three species
in the collection of the Paris Museum about which I had some
doubts, and I am indebted to Mr. Boulenger for photographs
of the types of Haplochromis nuchisquamulatus, H. obliquidens,
H, longirostris, and H. sauwvagei, which were sent to him by the
late Professor Hilgendorf in 1898, and to Dr. Pappenheim for
information about the first two of these species.
In Boulenger’s ‘Catalogue of African Freshwater Fishes’ 47
species of Cichlids are described from Lake Victoria (including
the Victoria Nile, Lakes Salisbury, Kioga, etc.). The number
in the present revision is practically the same (50), but, as no less
than 18 species are described below as new and some others
formerly regarded as synonyms have been re-established, it will
be evident that a considerable number of species recognized by
Boulenger have been eliminated.
A summary of what is known of the Cichlide of the great
African lakes may now be given ‘.
From Lake Albert only three species have been recorded
(Tilapia wilotica, Haplochromis wingatu, and H, multicolor), all
of which occur also in the Bahr-el-Gebel.
Tilapia nilotica is found in Lakes Edward and Kivu; in addition,
Lake Edward has a species of Tilapia (1'. eduardiana) which is
closely related to 7. variabilis of Lake Victoria, 5 endemic species
of Huplochromis, 4 apparently related to H. cinerews and the fifth
to H. spekii, of Lake Victoria, and an endemic monotypic genus,
Schubotzia, distinguished from Haplochromis by the dentition.
Kivu has five species of Haplochromis peculiar to the lake, of
which two seem to be related to H. cinereus.
From Tanganyika 89 species belonging to 37 genera are
known. Except 7%lapia nilotica, all these are endemic and the
majority of them belong to endemic genera. Nearly all the
Tanganyika Cichlid belong to genera which may have evolved —
* For explanation of the Plates see p. 191.
+ For Cichlide of Lakes Edward and Kivu, see Ann. & Mag. Nat. Hist. (9) viii.
1921, p. 632; of Tanganyika, Ann. & Mag. Nat. Hist. (9) v. 1920, p.30; of Nyassa,
P. Z.S. 1921, p. 678.
158 MR. C. TATE REGAN ON THE
in the lake from two ancestral types, Limnotilapia and Haplo-
chromis; but, although there are a number of genera which
appear to be derived from Haplochromis, that genus itself is
represented by two species only.
From Nyassa 84 Cichlide belonging to 14 genera* have been
described, nearly all endemic. The most striking feature is the
presence of 53 endemic species of Haplochromis, which appear to
form a natural group and have evidently evolved in the lake
from one or a few ancestral forms. Of the endemic genera four
(with 12 species) are evidently derived from forms closely related
to species of Haplochromis now living in the lake, and two more
(Cynotilapia, Pseudotropheus) are related to Haplochromis. Ot
the others Hemitilapia and Otopharynx are near Tilapia, Chilo-
tilapia is Otopharynx specialized, and Corematodus is mainly
distinguished from the endemic Tilapia sqwamipinnis by its
mouth and dentition. The indications are that the endemic
Nyassa Cichlids have originated in the lake from about haltf-a-
dozen ancestral forms.
In Lake Victoria there are 50 species of Cichlid, all peculiar
to the lake except Tilapia zillii, a Nilotic species. The endemic
species area Tilapia, 44 Haplochromis, and 4 monotypic genera
distinguished from Haplochromis by peculiarities of the dentition,
or in one by an increased number of anal spines.
The species of Haplochromis exhibit almost as great a diversity
as in Nyassa, yet there are certain features which enabie one to
say almost at a glance to which lake a species belongs. In most
of the Nyassa Haplochromis the caudal fin is more or less
distinctly emarginate, and is covered with small scales in the
adult fish, whereas in most of the Victoria species it 1s rounded
or truncate and is scaly only in the basal half. In the Nyassa
Haplochromis a few distinctive types of coloration are prevalent—
for example, 15 species have a dark band on each side from nape
to caudal fin, and several others have four dark spots on each
side; in most of the Victoria Haplochromis, when markings are
present, they take the form of regular dark cross-bars with a
dark band from head to caudal fin and another above the lateral
line.
There can be little doubt that Haplochromis cinereus, a small
species, with rather short decurved snout and moderately small
and slightly obliqne mouth, is a generalized type; species with
these characters are widely distributed in African rivers. A
number of species in Lake Victoria are closely related to
H. cinereus, but differ in the dentition of the jaws or of the
pharyngeals, in haying thicker lips, more gill-rakers, a larger
eye, etc.
Haplochromis serranus and its allies are somewhat larger
* In my paper I recognized 15, but I now find that Astatotilapia cannot be
maintained as distinct from Haplochromis, several of the Lake Victoria species
having enlarged teeth at the ends of the premaxillaries more or Jess developed and
not at all constant.
CICHLID FISHES OF LAKE VICTORIA. 159
species, with a larger mouth and more prominent lower jaw; in
some species of this group the mouth is oblique, and in some the
outer teeth are strong and spaced. Finally, there are species
with a strongly projecting lower jaw, in some of which the cleft
of the mouth is nearly vertical.
One species, H. ishmaeli, is of peculiar interest; it is so like
H. cinereus that the types (12 specimens) include 6 examples of
the latter, but it differs in having much larger and more massive
pharyngeals, with the teeth large armel blunt, ihn eas in H, cinereus
they are small and slender. ie have described a similar pair of
species (/7. tetrastigma and H. placodon) from Nyassa. A remark-
able group of three species includes Haplochromis sauvagei and
the monotypic genera Macropleurodus and Hoplotilapia, which
scarcely differ from each other except for the considerable differ-
ences in dentition, and seem to illustrate Cope’s paradoxical
view that a species may persist through several genera. It is
especially interesting to note that the normal markings are those
described above as characteristic of many Victoria species, but
that in addition all three are known to possess a ‘ bicolor” form,
a sort of piebald, in which the dark pigment is concentrated into
a number of blotches and irregular cross-bars which extend on to
the vertical fins.
Another interesting example of ahs same species in different
genera is provided by Haplochromis annectens and Platyteniodus
degent ; ; here the former shows a slght departure from the
normal Haplochromis dentition towards the Platyteniodus type.
The conclusion is that in Victoria most of the endemic
Cichlide have evolved in the lake from a type very similar to
Haplochromis cinereus, and that the primary evolution has been
in the size and shape of the mouth, and the number, structure,
and arrangement of the teeth, doubtless in adaptation to different
kinds of food and different methods of feeding. The facts seem
more in harmony with Gulick’s theory of habitudinal segregation
than with the modern idea of evolution by accidental mutations.
The Cichlide of Tanganyika and Nyassa lead to the same
conclusion.
In my paper on the Nyassa Cichlide I called attention to two
striking examples of convergent evolution, Pseudotropheus and
Aulonocara of Nyassa, respectively showing a great resemblance
to Tropheus and Trematocara of Tanganyika. Lake Victoria
provides some more examples : Haplochromés chilotes has the lips
thick and produced into lobes as in Lobochilotes of Tanganyika ;
ff, ishmaeli has the pharyngeals massive and provided with large
blunt teeth as in H. placodon of Nyassa. Further, lomaginiies
obliquidens and Macropleurodus bear a resemblance to Nyassa
genera in their dentition, the former to Hemitilapia, the latter
£0 Chilotilapia.
If the degree of differentiation be taken as a guide, one may
form the conclusion that the Cichlid have inhabited Tanganyika,
longer than Nyassa, and Nyassa longer than Victoria.
160 MR. C. TATE REGAN ON THE
From what has been said above as to the evolution and
relationships of the Cichlidz of Victoria, it will be evident that
I do not regard the classification here proposed as entirely
satisfactory. A number of divergent species are placed in
Haplochromis and a few extreme types are regarded as generically
distinct, although the close relationship of each to a species of
Haplochronis is obvious. At present I am not in a position to
improve this arrangement.
Synopsis of the Genera.
ES SEMICE COOOL |: cooooobopscdbnonooude-iancdoto seqounssceadaueaan | Ibon tLe INA.
Il. Scales ctenoid.
A. Teeth in 2 or more series anteriorly ; a single series on each side of upper
jaw.
SHAVUOT sassso svavcacongoanneued esseggooncdenapsacauaasasonocana sc, Bn JelayNIOGorOaS:
4to 6 anal spines ........ Levesseeseee. 8. Astatoreochromis.
B. Several series of teeth on each side of upper jaw.
1. Upper jaw with an outer series of enlarged teeth and several inner series
of small teeth anteriorly, and 3 or 4 series of enlarged teeth laterally.
4. Macropleurodus.
2. Teeth small, conical, in bands.
Bands rather broad, narrower at the sides than in front ...... 5. Hoplotilapia.
Bands very broad, upper broader at the sides than in front... 6. Platyteniodus.
1, Trnapra A. Smith, 1840.
Pharyngeal apophysis formed by parasphenoid only. Scales
eycloid. An outer series of bicuspid teeth and several inner
series of tricuspid teeth.
Africa and Syria.
1. TILAPIA VARIABILIS Bouleng., 1906.
Tilapia variabilis Bouleng. Cat. Afr. Fish. 11. p. 167, fig. 108.
Teeth in 4 to 8 series, 50 to 120 in outer series of upper jaw.
17 to 19 gill-rakers on lower part of anterior arch. Dorsal
XVI-XVIIT 10-13. . Anal III (IV) 10-11. Pectoral longer
than head.
Total length 320 mm.
L. Victoria and Victoria Nile.
2. TILAPIA ZILLIT Gerv., 1848.
Tilapia zillii Bouleng. Cat. Afr. Fish. iu. p. 197, fig. 126.
Teeth in 3 to 6 series, 20 to 60 in outer series of upper jaw.
8 to 11 gill-rakers on lower part of anterior arch. Dorsal XIV—
XVI 10-13. Anal IIT 7-10. Pectoral usually not longer than
head.
Total length 290 mm.
Syria to the Niger and the Victoria Nile.
2. Haprocuromis Hilgendorf, 1888.
Pharyngeal apophysis formed by parasphenoid in middle and
basioccipital at sides. Scales ctenoid. Teeth in 2 or more series
CICHLID FISHES OF LAKE VICTORIA. 161
anteriorly, becoming a single series laterally, conical or com-
pressed, unicuspid, or outer mostly bicuspid and inner tricuspid.
Africa and Syria.
All the species described below are peculiar to L. Victoria and
the Victoria Nile. Haplochromis obliquidens, the type species,
has a dentition unlike that of most of the species which have
been placed in this genus, but the discovery of a species in
L, Kivu (2. astatodon) some individuals of which have a typical
Ctenochromis dentition, whilst others approximate to H. obli-
quidens, makes 1t possible to regard this difference as only sub-
generic and to still include species with conical or cuspidate
teeth in Haplochromis.
Synopsis of the L. Victoria Species.
1. Teeth slender, cuspidate, in 5 to 8 series, the inner well developed and not
separated by a distinct interspace from the outermost. (Weochromis Regan.)
Snout usually a little projecting beyond lower jaw, which is
short and broad, about + lengthof head ©....... . L. nigricans.
Jaws equal or lower slightly projecting, relatively slender,
avout --slenathwommead. oa. chsscreck ec section cose stone ecnceeer 2. nuchisquamulatus.
If. Outermost series of teeth conical or bicuspid, enlarged, separated by an inter-
space from the smaller inner teeth. (Ctenochromis Ptefter.)
A. Jaws equal anteriorly, or lower shorter than upper, or rarely slightly pro-
jecting.
1. Lips not produced into lobes.
a. Interorbital width not more than 4 length of head.
a. Pharyngeal teeth slender.
%* Lower jaw not shorter than upper; lips normal.
Caudal rounded; caudal peduncle as long as deep; maxillary
extending to below anterior part of eye ............0....00.
Caudal subtruncate; caudal peduncle longer than deep ;
maxillary extending to below anterior edge of eye ......
Caudal truncate; 11 or 12 gill-rakers on lower part of
ANGELIOTRELG lyases gen ieee va Nae SAUL at craet 0 atuaveincye ane aes
Caudal truncate ; 9 or 10 gill-rakers; eye 3 in head, twice
preorbital depth
Caudal tr teats 7 to 10 gill- rakers ; g ey e - more e than 311 in 1 head
* Lower jaw not shorter than upper; lips thick.
Co
. gestrt.
. nubilus.
. melanopus.
. Macrops.
. cinereus.
NOD OO
Teeth in 4 to 8 on 30 to 46 in outer series of upper jaw... 8. sawvagei.
Teeth in 3 to 5 series, 20 to 24 in outer series of upper jaw... 9. erassilabris.
*k* Lower jaw shorter than upper............... 10. annectens.
8. Middle pharyngeal teeth stout, subconical... 11. humilior.
y. Pharyngeal teeth large, obtuse .........:........ 12. ishmaeli.
b. Interorbital width more than + length of head. 13. obesus.
2. Lips thick, each produced into a lobe anteriorly .. 14. chilotes.
B. Lower jaw distinctly, but usually not strongly, ree
A. Outer teeth numerous, close together.
1. Maxillary not extending to below eye ............ 15. teniatus.
2. Maxillary extending to below eye, or nearly.
a. Hye + length of head or more (in ee of
100 to 110 mm.).. .. 16. martini.
b. Hye less than 4 leng th of head, ‘except i in the very young.
Gan Pr; emaxillary pedicels extending to be-
tween anterior edges of orbits ............... 17. nigrescens.
. Premaxillary pedicels not reaching orbits.
* Caudal peduncle 15 to 2 as long as deep.
D. XIV-XV 9-10. 8 or 9 gill-rakers on lower part of
anterior arch ......... mabhedeasaenunecane! Ish (MeCN anmpas.
Proc. Zoou. Soc. 1922, D No. xc il
162 MR. C. TATE REGAN ON THE
D. XIV-XV 8-10. 10 to 12 gill-rakers on lower part of
AMPETION ATC cele. sau ete each ete EE EERE ee ee LOM anieno don:
D. XV-XVII 9-10. 9 to 11 gill-rakers on lower part of
BIMAEVKOT CANO 5.5 snsnn500 90° ; Nes 20. guiarti.
** Caudal padaneiens once Sis re as done as deep; 5 series of scales on
cheek.
Depth of prorbital not greater than diameter of eye (in a
specimen of 200 mm. ie .. 21. serranus.
Depth of preorbital considerably. ereater than diameter of
eye (in a specimen of 220 mm. aie ane Aed Suanbonadesouacans ie CNC HIS
*** Caudal peduncle ig as =p loins as deep; 6 or 7 series of scales on
(B)INGIS) Fearn npalsoacoumdand bn ab Bando Boe cacaotaatd esate eMC K DONATI
B. Outer teeth rather strong, set well apart.
Pectoral not reaching anal; caudal subtruncate; mouth
little oblique ...... . 24, bayoni.
weet reaching anal ; “caudal tr ‘ancate ; F “mouth ‘oblique .. 25. macrodon.
. Lower jaw strongly projecting.
j gs 2} $
i Pectoral reaching anal; caudal truncate; mouth moderately oblique;
maxillary not extending to below eye.
Eye 5 in head (in a specimen of 170 mm.) ..................... 26. prognathus.
Hye 4 in head (in a specimen of 180 mm.) ..................... 27. maculipinna.
B, Pectoral reaching anal, or nearly; caudal rounded or subtruncate ; maxillary
reaching vertical-from anterior edge of eye.
1. Mouth ‘moderately oblique ; maxillary extending to below eye.
a. Premaxillary pedicels ending above nostrils ... 28. dichrourus.
6. Premaxillary pedicels ending well behind nostrils.
Maxillary extending to below anterior Re of eye; caudal
peduncle longer than deep .......... 29. spekit.
Maxillary barely | reaching vertical from anterior ‘edge of
eye; caudal peduncle : asilonetasideep) weresaeee cere eens 30. serranoides.
. Mouth very oblique.
Maxillax y reaching vertical from anterior edge of eye.......... 31. acutirostris.
Maxillary not reaching vertical from anterior edge of eye . 382. plagiostoma.
C. Pectoral not reaching anal; mouth not very ppiGnee lower jaw very
prominent, with the anterior teeth exposed.
Outer teeth forming a close-set series; maxillary reaching
vertical from anterior edge of eye ............... ec ee cess eee 33. macrognathus.
Outer teeth strong, spaced; maxillary not quite reaching
to below eye ...... 34. dentex.
Outer teeth strong, spaced ; “maxillary not nearly reaching
tovbelogrueyien 2) phe ee Aiea ed elo Muar Ulan, Une We ee 35. mento.
D. Pectoral not reaching anal; mouth very oblique.
1. 8-10 scales from origin of dorsal to lateral line... 36. cavifrons.
2. 5 to 7 scales from origin of dorsal to lateral line. :
a. Depth of body 22 in the length ..................... 87. orthostoma.
6. Depth of body 3 to 44 in length,
a, Caudal pedunele 14 to 13 as long as deep.
Lower jaw projecting upwards above end of snout ............ 38. xenostoma.
Lower jaw not projecting above end of snout; maxillary not
quite reaching vertical from anterior edge of eye ...... 39. pellegrini.
Lower jaw not projecting above end of snout; seek
ending midway between nostril and eye ned ane .. 40. argenteus.
(3. Caudal peduncle 13 to 21 as lea as JED.
Snout 13 to twice diameter of eye (in specimens of 100 to
USO son Tae)) Were saben acon haat aanre nates accoce aba melnchucscsatemdere 2 llat ory. URa arn
Snout 1; to 13 diameter of eye a ieee of 115 to
160k) AB igdishSihik cohrseiia nds HABE REL REALE LEE. ILO CLCULECA LOC
ILI. Outer teeth few and ee with ce anterior cusp and mdistinct posterior
cusp (Bayonia Bouleng.) Je Sek ee Mase ee BA’ aren OwOr.
IV. Teeth slender, distally expanded and compressed, outer obliquely truncated
(Haplochromis ELil orercl) is VEE aga meme natn OL obliquidens.
CICHLID FISHES OF LAKE VICTORIA. 163
1. HAPLocHROMIS NigRIcANS Bouleng., 1906.
Tilapia wigricans (part.) Bouleng. Cat. Afr. Fish. iii. p. 240,
fig. 160.
Tilapia simotes Bouleng. t.c. p. 242, fig. 161.
9
Depth of body 23 to 3 in length, length of head 3 to 33.
Snout decurved, about as long as diameter of eye, which is 31 to
33 in length of head, greater than preorbital depth, equal
to interorbital width. Jaws equal or lower jaw a little the
shorter, short and broad, about 4 length of head; maxillary
reaching vertical from anterior edge of eye or a little beyond ;
teeth slender, cuspidate, in 6 to 8 series, outer but little enlarged
and not separated by a distinct interspace from inner; 40 to 70
in outer series of upper jaw. 3 or 4 series of scales on cheek.
7 to 10 gill-rakers on lower part of anterior arch. Pharyngeal
teeth small. 31 to 33 scales in a longitudinal series, 5 or 7 from
origin of dorsal to lateral line. Dorsal XV-XVI 9-10; last
spine 3 length of head. Anal IIIT 8-9; third spine nearly as
long as last dorsal. Pectoral a little shorter than head, not
reaching anal. Caudal truncate or subtruncate. Caudal peduncle
as long as or a little longer than deep. Brownish or greyish,
with traces of blackish cross-bars; males with 2 to 4 ocelli on
anal.
Nine specimens, including types of 7. nigricans and 7’. simotes,
70 to 145 mm. in total length, and a skeleton.
Boulenger’s figures show 7. wigricans as having quite a
different physiognomy from 7’. simotes, but this is because the
head is directed upwards in the former.
2. HAPLOCHROMIS NUCHISQUAMULATUS Hilgendorf, 1888.
Chromis nuchisquamulatus Hilgend. Sitzb. Ges. naturf. Fr.
Berlin, 1888, p. 76. 9
Ctenochromis nuchisquamulatus Pfeffer, Thierw. O.-Afr. Fische,
p. 14 (1896).
Tilapia ngricans (part.) Bonleng. Cat. Afr. Fish. iii. p. 240.
Haplochromis nuchisquamulatus (part.) Bouleng. t.c. p. 290.
Depth of body 23 in length, length of head 3 to 31. Snout
with straight profile, about as long as diameter of eye, which is
33 to 33 in length of head, greater than preorbital depth, equal
to or greater than depth of cheek; interorbital width 31 to 34 in
length of head. Jaws equal or lower slightly projecting ; rami
of lower jaw longer and more slender than in H. nigricans,
about 2 length of head. Teeth as in the preceding species, in 5 to
8 series, 50 to 80 in outer series of upper jaw. 2 or 3 series of
seales on cheek. 9 to 11 gill-rakers on lower part of anterior
arch.. Pharyngeal teeth small. 31 to 33 scales in a longitudinal
series, 5 or 6 from origin of dorsal to lateral line. Dorsal XV—
XVI 9-10; last spine 4 length of head. Anal III 9-10; third
spine 2 to 4 head. Pectoral as long as or a little shorter than
Tales
164 MR. C. TATE REGAN ON THE
head, nearly or quite reaching anal. Caudal truncate. Caudal
peduncle 14 to 14 as long as “deep. Silvery or greyish, with or
without 8 dark cross- bars an opercular spot; sometimes a bar
below eye.
Lake Victoria.
Two specimens, 100 and 145 mm. jong, from Kakindu (Bayon)
and Buddu Coast (Stmon), and two Sivelecone.
Very near 1. nigricans ; the difference in the structure of the
lower jaw is well seen on comparing the skeletons.
3. Hapiocnromis Gestri Bouleng., 1911.
Haplochromis desjontainesti (part.) Bouleng. Cat. Afr, Fish. 11.
p. 302.
: Paratilapia ae (part.) Bouleng. t. c. p. 318, Hea Abid
Depth of body 22 to 3 in length, length of head 22 to 3, Snout
with straight or convex profile, ‘from as ; long as to i diameter of
eye, which is 34 to 43 in length of head, greater than preorbital
depth, in adults less then depth of cheek ; interorbital with 33 to
44 in length of head. Jaws equal anteriorly or lower slightly
projecting ; premaxillary pedicels not quite reaching aD between
orbits; maxillary extending to below anterior 4 or 3 of eye;
teeth in 2 to 4 series, conical in adult, 34 to 60 in gue series of
upper jaw.. 3 to 5 series of scales on cheek. 8 gill-rakers on lower
part of anterior arch. Pharyngeal teeth caval 30 to 33 scales:
in a longitudinal series, 5 or 6 from origin or dorsal to lateral
line. Dorsal XV-X VI 8-10; last spine 2 to 4 oe of head.
Anal TIT 8-10; third spime 2 ‘or 2 head. Pectoral § to 4 head,
reaching vent. Caudal rounded. Caudal medtomalle as ions as
deep. Often a dark band from opercular spot to base of coal.
sometimes a second above lateral line; a bar below eye usually
present ; soft dorsal and caudal with or without spots; males
with blackish pelvic fins and oeelli on the anal.
Several specimens, 70 to 160 min. long.
Orit
4, HapLOCcHROMIS NUBILUS Bouleng., 1906.
2 Paratilapia victoriana Pellegrin, Bull. Soc. Zool. France, 1903,
p. 185, and Mem. xvii. 1905, p. 182, pl. xvii. fig. 3.
Pilapia nubila Bouleng. Cie Air. Wish. ps 2307 nes l55.
Haplochromis nuchisquamulatus (part.) Bouleng. t. c. p. 290,
mes IEE
Haplochromis desfontaineswt (part.) Bouleng. t. ¢. i 302.
Depth of body 24 to 3 in length, length of head 25 to 3. Snout
as long as or a Tittle longer than diameter of eye, which is 3 to 4
in length of head, greater than preorbital depth, in adult equal
to depth of chesle® : ee Suonlbitall width 354 to 4 in seme i of head.
Jaws equal anteriorly, or lower feebly projecting; maxillary
extending to below anterior edge of eye; teeth es or some
conical, in 2 to 5 series, 40 to 603 in cuter series of upper jaw. 3 or
4 series of scales on ‘cheek. 8 to 10 gill-rakers on lower part
CICHLID FISHES OF LAKE VICTORIA. 165
of anterior arch. Pharyngeal teeth small. 30 to 32 scales in a
longitudinal series, 4 to 6 from origin of dorsal to. lateral line.
Dorsal XV—X VI 8-10; last spine } to 4 length of head. Anal
IIL 8-10; third spine as long as or a little shorter than last
dorsal. Pectoral ? to as long as head, reaching vent or anal fin.
Caudal subtruncate. Caudal peduncle longer than deep. Oliva-
ceous or greyish to blackish, uniform or with dark cross-bars ;
often an opercular spot and a dark bar below eye; anai fin with
ocelli in males.
Numerous examples up to 125 mm. in total length.
Paratilapia victoriana Pellegrin appears to be very closely
related to, if not identical with, 7. nabilus, but if the figure
be accurate the pectoral fin is longer than in any examples of
H. nubilus that I have examined, reaching the middle of the
anal.
5. HAPLOCHROMIS MELANOPUS, Sp. n. (Text-fig. 1.)
Tilapia lacrimosa (part.) Boulenz. Cat. Afr. Fish. iii. p. 234.
Haplochromis stanleyt (part.) Bouleng. t. ec. p. 295.
Depth of body 2% to 3 in length, length of head 3 to 34. Snout
decurved, as longas or a little shorter than diameter of eye, which
Text-figure 1.
Sf SEK
Haplochromis melanopus, sp. 1.
is 3 to 32 in length of head, greater than depth of cheek, not
twice preorbital depth; interorbital width 33 to 37 in length of
head. Jaws equal anteriorly; maxillary extending to below
anterior edge of eye; teeth cuspidate or some conical, in 2 to 4
series, 50 to 65 in outer series of upper jaw. 3 to 4 series of scales
on cheek. 11 or 12 gill-rakers on lower part of anterior arch.
Pharyngeal teeth small. 33 scales in a jongitudinal series, 6 or 7
from origin of dorsal to lateral line. Dorsal XVI 8-9; last spine
2 to L length of head. Anal III 8-9; third spine } to nearly 4
length of head. Pectoral as long as or a little shorter than head,
166 MR. C. TATE REGAN ON THE
reaching vent or origin of anal. Caudal peduncle 1+ to 13 as
long as ‘deep. antl 6 more or less distinet dark cross- bars ;
sometinies an interrupted lateral band ; an opercular spot ; a dark
bar. below eye; soft dorsal and cane spotted ; pelvics blackish ;
1 to 3 ocelli on anal fin.
Seven specimens, 80 to 100 mm. long, from Entebbe and
Bunjako.
6. HApLocnromis MAcRors Bouleng., 1911.
? Astatotilapia jeannelli Pellegrin, Bull. Soc. Zool. France,
xxxvil. 1913, p. 313.
Tilapia macrops Bouleng. Cat. Afr. Fish. 141. p. 258, fig. 157.
Haplochromis stanleyi (part.) Bouleng. t. c. p. 295.
Depth of body 22 to 3 in length, length of head 3. Snout
shorter than diameter of eye, which is 3 in Tength of head, slightly
greater than iterorbital width, twice preorbital depth. Jaws
equal anteriorly; maxillary extending to vertical from anterior
edge of eye ; teeth in 4 or 5 series in upper jaw, 3 or 4 in lower,
outer bicuspid or some conical, 50 to 60 in outer series of upper
jaw. 9 series of scales on cheek. 9 or 10 gill-rakers on lower
part of anterior arch. Pharyngeal teeth small. 32 scales ina
longitudinal series, 5 or 6 from origin of dorsal to lateral line.
Dorsal XV 9; lasb spine 2 length of head. Anal III 10; third
spine as long as last dorsal. aanor al shorter than head, reaching
vent or origin of anal. Caudal truncate. Caudal peduncle longer
than deep. Silvery, with or without cross-bars and two dark
lateral bands, or uniformly blackish.
Three specimens, 85 to 105 mm. long.
7. HAPLOCHROMIS CINEREUS Bouleng., 1906.
Tilapia pallida (part.) Bouleng. Cat. Afr. Fish. 11. p. 231,
fig. 152.
Tilapia lacrimasa (part.) Bouleng. t. ¢. p. 234, fig. 154.
Haplochromis ishmaeli (part.} Bouleng. is (G5 Je ASS,
Paratiiapia victoriana (pent Boulene. t. ec. p. 341.
Paratilapia cinerea Bouleng. t. c. p. 344, fig. 232.
Pelmatochromis riponianus Bouleng. t. c. p. 411, fig. 281.
Pelmatochromis obesus (part.) Bouleng. t. ¢. p. 414.
Depth of body 2} to 34 in length, length of head 25 to 3}
Snout ou ved, nearly as long as or lon ger than diameter of eye,
which is 33 to Aa in length of head and greater than depth of
preeor mel aiterorbie a width 34 to 4 in length of head. Mouth
shehtly oblique ; jaws equal anteriorly ; maxillary extending to
below anterior margin of eye, or nearly; teeth in 3 to 6 series in
upper jaw, in 2 to 5 in lower, 36 to 70 in outer series of upper
jaw. Pharyngeal teeth small. 7 to 10 gill-rakers on lower part
of anterior arch. 30 to 34 scales in a longitudinal series, 6 or 7
from ogue of dorsal to lateral line. Dorsal XIV-XVI 8-10;
last spine 3 to 3 length of head. Anal ITI 8-10; third spine as
CICHLID FISHES OF LAKE VICTORIA. 167
long as or shorter than last dorsal. Pectoral as long as head, or
a little shorter, extending to origin or anterior part of anal.
Caudal truncate. Caudal peduncle 1} to 1? as long as deep.
Silvery or greyish, with or without dark cross-bars and a con-
tinuous or interrupted lateral hand; often bars across the snout
and a vertical one below the eye; males with pelvic fins dusky
and ocellar spots on anal fin.
Lake Victoria.
Numerous examples up to 140 mm. in total length.
8. HAPLOCHROMIS SAUVAGEL Pfeffer, 1896.
Ctenochromis sauvaget Pfeffer, Thierw. O.-Afr. Fische, p. 14.
Paratilapia granti (Bouleng., 1906) Bouleng. Cat. Afr. Fish.
i. p. 342, fig. 231.
Paratilapia crassilabris (part.) Bouleng. t. c. p. 345.
Paratilapia bicolor (part.) Bouleng. t. c. p. 346.
Paratilapia retrodens (part.) Bouleng. t. ¢. p. 347, fig. 235.
Depth of body 2% to 34 in length, length of head 22 to 33.
Snout as long as or longer than diameter of eye, which is 33 to
Ai in length of head, greater than przorbital depth, from a little
greater to a little less than depth of cheek; interorbital width 33
to 4 in length of head. Mouth as broadas long; lips thick; Jaws
equal anteriorly ; maxillary extending to below anterior edge of
eye, or not quite so far; teeth in 4 to 8 series, cuspidate or
conical, 30 to 46 in outer series of upper jaw. 3 to 5 series of
scales on cheek. 7 to 9 gill-rakers on lower part of anterior arch.
Pharyngeal teeth small. 32 to 34 scales in a longitudinal series,
6 or 7 from origin of dorsal to lateral line. Dorsal XTV—XVI 9—
10; last spine 4 to 4 length of head. Anal [iL 8-9; third spine
as long as or a little shorter than last dorsal. Pectoral as long as
ov a little shorter than head, reaching vent or origin of anal.
Caudal truncate. Caudal peduncle 17 to 13 as long as deep.
Body without markings or with regular dark cross-bars, or with
irregular blackish blotches and cross-bars extending on to vertical
fins; usually a dark lateral band and sometimes another above
lateral line; an opercular spot; often 2 bars across snout, another
between posterior margins of eyes, and another from eye to end of
maxillary; soft dorsal and caudal sometimes spotted ; males with
1 to 3 ocelli on anal fin.
Numerous specimens 80 to 150 mm. in total length.
9. HAPLOCHROMIS CRASSILABRIS Bouleng., 1906.
Paratilapia crassilabris (part.) Bouleng. Cat. Afr. Fish. 11.
p. 345, fig. 233.
Paratilapia retrodens (part.) Bouleng. t. c. p. 347.
Lepth of body 22 to 3 in length, length of head 3. Snout
from as long as to 14 diameter of eye, which is 32 to 44 in length
of head, greater than preorbital depth, about eyual to depth of
cheek; interorbital width 3) in length of head. Mouth broader
168 MR. C. TATE REGAN ON THE
than long; lips thick; jaws equal anteriorly; maxillary not ex-
tending to below eye; teeth in 3 to 5 series, outer strong, conical
in adult, 20 to 24 in outer series of upper jaw. 3 series of scales
on cheek. 7 to 9 gill-rakers on lower part of anterior arch.
Pharyngeal teeth small. 381 to 33 scales in a longitudinal series ;
6 or 7 from origin of dorsal to lateral line. Dorsal XV—XVI 8—
10; last spine from a little less to a little more than # length of
head. Anal III 8-9; third spine as long as or a little shorter
than last dorsal. Pectoral a little shorter than head, reaching
origin of anal. Caudal truncate or subtruncate. Caudal peduncle
11 to 12 as long as deep. An opercular spot; traces of dark
cross-bars and a dark lateral band.
Four specimens, 100 to 150 mm. long, from Entebbe; several
smaller examples are not included in the description.
10. HaPLocHROMIS ANNECTENS, sp.n. (Text-fig. 2.)
Haplochromis ishmaeli (paxt.) Bouleng. Cat. Afr. Fish, 111. p. 293.
Depth of body 2? in length, length of head 3. Snout decurved,
a little longer than diameter of eye, which is 4 in length of head,
Text-figure 2.
Haplochromis annectens, sp.n.
nearly twice preorbital depth, equal to depth of cheek; inter-
orbital width 33 in length of head. Lips thick; lower jaw shorter
than upper ; maxillary extending to below anterior edge of eye.
Teeth conical, in 4 or 5 series; outermost series enlarged, 36 in
upper jaw; inner series of upper jaw forming a band which
narrows slightly at the sides and then slightly increases in width at
each end. 4 series of scales on cheek. & gill-rakers on lower part
of anterior arch. Pharyngeal teeth small. 32 scales in a longitu-
dinal series, 7 from origin of dorsal to lateral line. Dorsal XVI9;
last spine 2 length of head. Anal III 9; third spine 2 head.
Pectoral as long as head, reaching anal. Caudal truncate.
CICHLID FISHES OF LAKE VICTORIA. 169
Caudal peduncle longer than deep. A: dark lateral band from
opercular spot to base of caudal; a dark bar below anterior part
of eye; pelvics blackish ; 3 ocelli on anal.
A single specimen, 180 mm. in total length, from Buddu Coast.
11. Hapiocuromis HumiLIoR Bouleng., 1909.
Tilapia humilior Bouleng. Cat. Afr. Fish. iii. p. 280, fig. 151.
Tilapia bayoni Bouleng. t. c. p. 240, fig. 159.
Depth of body 3 to 34 in length, length of head 3. Snout
decurved, as long as or a little shorter than diameter of eye,
which is 34 in length of head, nearly twice depth of preorbital,
greater than depth of cheek ; interorbital width 45 in length of
head. Jaws equal anteriorly; maxillary extending to below
anterior margin of eye or a little beyond; teeth cuspidate, in 4
to 6 series, 54 to 70 in outer series of upper jaw. 3 to 4 series
of scales on cheek. 7 or 8 gill-rakers on lower part of anterior
arch. Middle pharyngeal teeth rather stout, subconical. 33 or
34 scales in a longitudinal series, 7 from origin of dorsal to
lateral line; pectoral scales very small. Dorsal XV—XVI 9-10,
last spine 4 or nearly 3 length of head. Anal IIT 8-10; third
spine + to a little more than 2 length of head. Pectoral shorter
than head, not reaching anal. Caudal subtruncate. Caudal
peduncle 14 to 14 as long as deep. Silvery, brassy, or coppery ;
back darker; a dark bar below eye; males with 2 or 3 ocelli
on anal,
Three specimens, 100 to 130 mm. long, one of the types of the
species and two types of 7’. bayoni. The second specimen of
1’. humilior isin very poor condition and is not included in the
description.
12. HapnLocHroMis IsHMAELI Bouleng., 1906.
Tilapia pallida (part.) Bouleng. Cat. Afr. Fish. ii. p. 231.
Haplochromis ishmaeli (part.) Bouleng. t. c. p. 293, fig. 199.
Paratilapia victoriana (part.) Bouleng. t. c. p. 341, fig. 230.
Depth of body 23 to 3 in length, length of head 23 to 3.
Snout decurved, about as long as diameter of eye, which is 37 to
33 in length of head, considerably greater than preorbital depth,
equal to or greater than depth of cheek; interorbital width 33 to
4in head. Jaws equal anteriorly or lower very slightly pro-
jecting; maxillary extending to below anterior edge or anterior
2 of eye; teeth cuspidate or conical, in 3 or 4 series, 34 to 60 in
outer series of upper jaw. 3 or 4 series of scales on cheek. 8 or
9 gill-rakers on lower part of anterior arch. Lower pharyngeal
broad and massive, with strong blunt teeth. 31 to 33 scales in
a longitudinal series, 6 to 8 from origin of dorsal to lateral line.
Dorsal XV-XVI 9-10; last spine 4 to 2 length of head. Anal
ILL 8-9; third spine as long as or a little shorter than last
dorsal. Pectoral as long as head, extending to above anterior
part of anal. Caudal truncate. Caudal peduncle 13 to 13 as long
170 MR. C. TATE REGAN ON THE
as deep. Silvery, with or without dark cross-bars; sometimes an
interrupted lateral band ; an opercular spot ; usually a bar below
eye; soft dorsal and caudal sometimes spotted ; males with ocelli
on anal fin.
Several specimens, 90 to 135 mm. in total length.
The remarkable pharyngeal dentition might well be held to
justify the genus Labrochromis (Regan, 1920), were it not that in
all other characters the species is nearly identical with H. cinereus.
13. HarLocyromis opEsus Bouleng., 1906.
Pelmatochromis obesus (part.) Bouleng. Cat. Afr. Fish. ii.
p. 414, fig. 283.
Depth of body 2 in length, length of head 3. Snout longer
than diameter of eye, which is 4 in length of head, greater than
depth of preeorbital, less than depth of cheek ; interorbital width
25 in length of head. Mouth wide, oblique; lower jaw not pro-
jecting ; maxillary exposed, extending to below eye; teeth rather
stout, conical, biserial, about 50 in outer series of upper jaw.
4 series of scales on cheek. 10 gill-rakers on lower part of
anterior arch. Pharyngeal teeth small. 32 seales in a longitu-
dinal series, 7 from origin of dorsal to lateral line. Dorsal XV
9; last spine 2 length of head. Anal III 8; third spine 3 head.
Pectoral as long as head, reaching anai. Caudal truncate.
Caudal penduncle as long as deep. Traces of dark cross-bars ;
an opercular spot and a bar beiow eye; soft dorsal spotted; anal
fin with 3 ocelli (3).
One of the types, 150 mm. long.
An isolated species, for which I proposed the generic name
Lipochromis (‘ Annals,’ 1920).
14. HapLocHromis cutnores Bouleng., 1911.
Paratilapia chilotes Bouleng. Cat. Afr. Fish, ii. p. 338, fig. 228.
Depth of body 3 in length, length of head 22. Diameter of
eye 4 in Jength of head. Jaws equal anteriorly; lips very thick,
each produced anteriorly into a lobe; maxillary not extending to
below eye ; teeth small, conical. 3 series of scaleson cheek. 8 or
9 gill-rakers on lower part of anterior arch. 31 or 32 scales
in a longitudinal series. Dorsal XVI-XVIT 9. Anal III 8-9.
Pectoral 2 head. Caudal truncate. Dark cross-bars and a
lateral band.
Total length 98 mm.
15, HAPLocHROMIS THNIATUS, sp. n. (Text-fig. 3.)
Paratilapia prognatha (part.) Bouleng. Cat. Afr. Fish. i.
Depth of body 32 in length, length of head 23. Head 22
to 22 as long as broad; upper profile slightly concave. Snout 13
to 1? diameter of eye, which is 4 to 44 in length of head, greater
CICHLID FISHES OF LAKE VICTORIA. 7p
than preorbital depth, equal to or greater than depth of cheek ;
interorbital width 5 in length of head. Maxillary not extending
to below eye; lower jaw moderately projecting ; chin obtuse ;
teeth in 3 series in upper jaw, 2 or 3 in lower, some of the inner
tricuspid, outer series conical, 40 in upper jaw, anterior mode-
rately strong. 3 series of scales on cheek. 8 or 9 gill-rakers on
lower part of anterior arch. Dorsal XV-XVI 9; last spine
Text-figure 3.
Haplochromis teniatus, sp. n.
longest, 4 length of head; longest soft rays less than half length
of head. Anal ITI 9-10; third spine stronger than and as long
as last dorsal. Pectoral 2 length of head. Caudal truneate.
Caudal peduncle 12 as long as deep. 31 or 32 scales in a longi-
tudinal series, 6 from origin of dorsal to lateral line. Silvery,
back darker; an opercular spot; a dark band along middle of
side, another above lateral line; dorsal and caudal with small
spots.
Lake Victoria.
Two specimens, 95 and 115 mm. long, from Entebbe (Degen)
and Kavirondo Bay (Alluaud).
|
16. Haritocuromis MARTINI Bouleng., 1906.
Tilapia martini (part.) Bouleng. Cat. Afr. Fish. iii. p. 239,
‘fig. 158.
Depth of body 22 to 3 in length, length of head 22 to 3.
Upper profile of head convex; snout shorter than diameter of
eye, which is 22 to 3 in length of head, greater than interorbital
width, twice depth of preorbital. Lower jaw a little projecting;
maxillary extending to below anterior 4 of eye; teeth in 3 or 4
series, cuspidate, 50 to 70 in outer series of upper jaw. 4A or 5
series of scales on cheek. 8 or 9 gill-rakers on lower part of
anterior arch. Pharyngeal teeth small. 33 seales in a longitu-
dinal series, 6 from origin of dorsal to lateral line. Dorsal XV—
XVI 8-9; last spine } length of head. Anal Hi 8-9; third
spine as long as last dorsal. Pectoral as long as head, extending
1 We MR. C. TATE REGAN ON THE
to above anterior part of anal. Caudal truncate. Caudal
peduncle 13 to 13 as long as deep. A blackish opercular spot ;
a blackish stripe from head to caudal fin, a second above lateral
line.
Three of the types, 100-110 mm. long, from Bunjako.
17. HAPLOCHROMIS NIGRESCENS Pellegrin, 1909.
Astatotilapia nigrescens Pellegrin, Bull. Soc. Zool. France,
XxXxiv. p. 157, and Meém. xxii. 1910, p. 292, pl. xiv. fig. 3
? Astatotilapia roberti Pellegrin, Bull. Soc. Zool. France,
xxxvul. 1913, p. 312.
Haplochromis percoides (part.) Bouleng. Cat. Afr. Fish. i.
p. 296,
Paratilapia parvidens Bouleng. t. c. p. 323, fig. 215.
Paratilapia serranus ea ) Bouleng. (ty (6) 105 33 4,
Depth of body 22% to 3 in length, length of head 25 to 3.
Snout 13 to 13 diameter of eye, which is 4 to 42 in length of
head, greater than depth of preorbital, equal to or ereater than
depth ‘of cheek ; interorbital width 4 in length of head. Upper
profile of head straight or slightly concave; mouth moderately
oblique; premaxillary pedicels extending to between anterior
edges of orbits; maxillary extending to below anterior edge or
anterior + of eye; lower jaw usually a little projecting. Teeth
conical, or outer bicuspid and inner tricuspid, in 3 to 5 series in
upper jaw and 3 or 4 in lower, 40 to 55 in outer series of upper
jaw. Cheek with 4 series of scales. 8 or 9 gill-rakers on lower
part of anterior arch, seer aymgiee teeth slender. 52 or 33 scales
in a longitudinal series, 5 or 6 from first dorsal spine to lateral
line. Dorsal XV-XVI 9-10; last spine from 3 to nearly §
length of head. Anal III 8-9; third spime stronger than and
nearly as long as last dorsal. Pectoral 2 ? length of he: ad, reaching
vent or origin of anal. Caudal ee or subtr uncate. Caudal
peduncle 1 to 13 as long as deep. Coloration uniform, or a dark
lateral hand, or traces of 8 to 10 dark cross-bars; soft dorsal and
caudal sometimes with series of spots.
Five specimens, 95 to 140 mm. in total length, from the
Victoria Nile and Jinja (Bayon).
18. HapLocHROMIS FLAVIPINNIS Bouleng., 1906.
Haplochromis percoides (part.) Bouleng. Cat. Afr. Fish, 11
p. 296, fig. 201.
Pelmatochromis flavipinnis Bouleng. t. c. p. 418, fig. 286.
Depth of body 3 to 31 in length, length of head 27 GON oe
Snout 14 to 14 diameter of eye, which is 4 to 5 in fone) of head,
equal to or greater than przorbital depth, from #% to a little
greater than depth of cheek ; interorbital width 5 ia length of
head. Head twice as long as broad; upper profile concave y above
the eyes; mouth oblique; maxillary extending to below anterior
edge of eye, or nearly; lower jaw projecting; teeth in 3 or 4
CICHLID FISHES OF LAKE VICTORIA. Uys
series, outer conical or bicuspid, inner conical or tricuspid ; 45 to
60 in outer series of upper jaw. Cheek with 4 to 6 series of
scales. 8 or 9 gill-rakers on lower part of anterior arch. Pharyn-
geal teeth slender. 31 or 32 scales in a longitudinal series, 6 to
8 from first dorsal spine to lateral line. Dorsal XIV-XV 9-10;
last spine 4 length of head. Anal III 8-9; third spine as long
as or shorter than last dorsal. Pectoral 3 to # length of head,
not extending to above anal. Caudal subtruncate. Caudal
peduncle 14 as long as deep. Four broad dark eross- bars on body
and a dark spot at base of caudal fin ; sometimes a longitudinal
band connecting first two bars below lateral line and another
running forward from caudal spot; a bar between anterior edges
of eyes and sometimes another in front of it across snout; a
vertical bar below eye and another running upwards and back-
wards from posterior edge of eye; an opercular spot; fins yellow,
or dorsal and caudal dusky; one or two orange ocelli on anal fin
in males.
Three specimens, types of the species and of H. percoides, 85 to
145 mm. long.
19. HapLocHROMIS MICRODON Bouleng., 1906.
Tilapia lacrimosa (part.) Bouleng. Cat. Afr. Fish. 11. p. 234.
Haplochromis stanley: (part.) Bouleng. t. ec. p. 296, fig. 200.
Paratilapia serranus (part.) Bouleng. t. c. p. 334.
Pelmatochromis microdon Bouleng. t. ¢. p. 412, fig. 282.
Depth of body 3 to 3} in length, length of head about 3.
Upper profile of head straight or slightly concave; snout from a
little shorter than to !4 as long as diameter of eye, which is 3 to 4
in length of head, greater than depth of preorbital, equal to or
greater than depth of cheek ; interorbital width 32 to 4 in length
of head. Mouth oblique; lower jaw projecting; maxillary
reaching vertical from anterior edge of eye; teeth in 3 or 4
series in upper jaw, 2 to 4 in lower, 40 to 70 in outer series of
upper jaw. 3 or 4 series of scales on cheek. 11 or 12 gill-rakers
on lower part of anterior arch, the posterior much expanded.
30 to 33 scales in a longitudinal series, 44 to 6 between first
dorsal spine and lateral line. Dorsal XTV—XV 8-10; Jast spine
1 to 2 length of head; longest soft rays § to 2? length of head.
Anal 111 8-9; last spine as long as or a httle shorter and
stronger than last of dorsal. Pectoral as long as or a little
shorter than head, reaching anal. Caudal truncate or slightly
emareinate. Caudal peduncle 14 to 13 as long as deep. Silvery
or greyish, back olivaceous or brownish ; sides with or without
dark cross-bars and a dark lateral band; males with a dark bar
below eye, blackish pelvic fins and ocellar spots on anal.
Lake Victoria.
The specific name refers to the fact that the type, a specimen
of 175 mm., has very small teeth ; this I believe to be due to mal-
formation of the lower jaw, which does not bite against the
174 MR. C. TATE REGAN ON THE
upper ; in all other characters it agrees with seven specimens of
80 to 140 mm., including the type of H. stanleyi figured by
Boulenger.
20, Harnocuromis GuiArti Pellegrin, 1905.
Tilapia guiarti Pellegr. Mém. Soc. Zool. France, xvii. p. 184,
jOilaea iil waifee le
Tilapia perriert Pellegr. ib. xxii. 1910, p. 295, pl. xiv. fig. 4.
Tilapia pallida (part.) Bouleng. Cat. Afr. Fish. 111. p. 232.
Paratilapia longirostris (part.) Bouleng. t. c. p. 332.
Paratilapia serranus (part.) Bouleng. t.¢. p. 334, fig. 225
Par atilapia guiarti Bouleng. t. ¢. p. 236, fig. 226.
Depth of body 3 to 4 in length, length of head 3 to 33. Snout
from a little shorter than to twice diameter of eye, soit is 3
(young) to 54 in length of head, in adult equal to or less than
depth of cheek and equal to or nob much greater than depth of
preorbital ; interorbital width 3 3 to 41 in Jength of head.
Lower jaw more or less distinctly projecting ; sane pail ap nearly
or quite reaching vertical from anterior edge of eye, sometimes a
little beyond ; teeth cuspidate in young, conical in adult, in 3 to
5 series, 36 to 70 in outer series of upper jaw. 39 to 5 series of
seales on cheek. 9 to 11 gill-rakers on lower part of anterior
arch. Pharyngeal teeth slender. 32 to 34 scales in a longitu-
dinal series, 6 or 7 from origin of dorsal to lateral line. Dorsal
XV-XVII 8-10; last spine from + to more thau 2 length of
head. Anal III aL 10; third spine stronger and as long as or a
little shorter than last dorsal. Pectoral fr om # to nearly as long
as head, reaching vent or origin of anal. Caudal truncate or very
slightly emar einate, sometimes rounded below. Candal peduncle
12 to 2 as long as deep. Silvery or golden on sides, back darker ;
faint dark erogs-bars sometimes present ; often a blackish band
from opercular spot to base of caudal and a second above lateral
line; soft dorsal and caudal usually spotted; anal with ocelli in
males.
Numerous examples up to 225 mm. in total length.
21. HAPLOCHROMIS SERRANUS Pfeffer, 1896.
Hemichromis serranus Pfeffer, Thierw. O.-Afr. Fische, p. 23.
Pelmatochromis spehit (part.) Bouleng. Cat. Afr. Fish. 111. p. 416.
Depth of body 22 to 31 in length, length of head 22 to 23.
Head 2 to 24 as long as broad ; upper profile straight. Snout
13 to nearly twice diameter of eye, which is 4 to 5 in length of
head! equal to or greater than preorbital depth, equal to or less
than ‘depth of cheek; interorbital width 4 to 42 in length of head.
Mouth moderately oblique; maxillary reaching vertical from
anterior margin of ey ; lower jaw projecting ; teeth conical, or
inner ti icuspid, in 3 to 5 series in upper jaw and 2 to 4 in eee
40 to 80 in outer sevies of upper jaw. 5 series of scales on cheek.
8 or 9 gill-rakers on lower part of anterior arch. Pharyngeal
CICHLID FISHES OF LAKE VICTORIA, 175
teeth slender. 31 to 33 scales in a longitudinal series, 7 or 8
from origin of dorsal to lateral line; pectoral scales very small.
Dorsal XV—XVI 9-10; last spine longest, less (adult) or more
(young) than 4 length of head ; longest soft rays 3 to 2 length of
head. Anal IIT 8-9; third spine stronger and a little shorter
than last dorsal. Pectoral 3 to # length of head, nearly or
quite reaching anal; pelvics reaching vent or origin of anal.
Caudal truncate or subtruncate. Caudal peduncle a little longer
than deep. A dark opercular spot, usually a more or less distinct
dark band from eye to caudal fin, another above lateral line, and
a dark stripe at base of dorsal; traces of dark cross-bars, some-
times a dark bar below eye; dorsal and caudal sometimes with
series of small dark spots. Pelvies and anal pale or dusky, some-
times with ocelli on posterior part of anal.
Four specimens, 125 to 200 mm. in total length.
22. HAPLOCHROMIS ALTIGENIS, sp. n. (PI. I.)
Paratilapia longirostris (part.) Bouleng. Cat. Afr. Fish. iii.
p- 332.
Pelmatochromis spekii (part.) Bouleng. t. ¢. p. 417.
Depth of body 3 in length, length of head 22 to 23. Head
2 to 27 as long as broad ; upper profileconvex. Snout morethan
twice as long as diameter of eye, which is 52 to 6 in length of
head, less than preeorbital depth and little more than 3 depth of
cheek ; interorbital width 43 to 43 in length of head. Maxillary
extending to below anterior margin of eye; lower jaw projecting ;
teeth conical, in 4 or 5 series in upper jaw and 3 or 4 in lower,
60 to 70 in outer series of upper jaw. 5 series of scales on cheek.
9 or 10 gill-rakers on lower part of anterior arch. Pharyngeal
teeth all slender. 32 or 33 scales in a longitudinal series, 7 or 8
from origin of dorsal to lateral line. Pectoral scales very small.
Dorsal XV 9-10; last spine longest, + to 2 length of head;
longest soft rays less than 3 length of head. Anal III 8-10;
third spine stronger than last dorsal, 2 to 7 length of head.
Pectoral length of head or less, reaching origin of anal or not.
Caudal subtruncate. Caudal peduncle longer than deep. Silvery;
back darker; a bar below eye; an opercular spot and a lateral
band more or less distinct. Vertical fins dusky, soft dorsal and
caudal with or without series of spots; pelvics blackish; 4 or 5
ocelli on posterior part of anal.
Two specimens, 220 and 235 mm. in total length, from Bunjako.
A specimen of 100 mm. from Bulolo, L. Kioga, is not included in
the description.
23. HAPLOCHROMIS SQUAMULATUS, nom. n.
Paratilapia pectoralis (not Ctenochromis pectoralis Pfeff.).
Bouleng. Ann. Mus. Genov. 1911, p. 66, pl. i. fig, 2, and Cat. Afr.
Fish. ui. p. 339, fig. 229.
Closely related to H. serranus and H. altigenis, with the snout
176 MR. C. TATE REGAN ON THE
more convex than the former and the preorbital narrower than
the latter. 6 or 7 series of scales on cheek.
Total length 185 mm. Ripon Falls.
HAPLocHROMIS BAYONI Bouleng., 1909.
Paratilapia bayoni Bouleng. Cat. Afr, Fish. 111. p. 337, fig. 2
Depth of body 34 in length, length of head 24, Snout as
curved, a little more than kos diameter of eye, which i is 53 in
length of head, equal to depth of preorbital, less than depth of
aihsel 3 interor one al width 44 in length of head. Mouth mode-
rately oblique ; lower jaw pr ojecting ; maxillary not extending to
below eye; teeth conical, in 4 series in ae jaw, 3 in lower:
outer teeth rather strong, set well apart. 4 series of scales on
cheek. 9 gill-rakers on lower part of anterior arch. Pharyngeal
teeth slender. 32 scales in a longitudinal series, 6 from oatsie of
dorsal to lateral line. Dorsal XVI 10; last spine 2 length
of head. Anal [11 8; third spine + 1head. Pectoral 2 length of
head, not reaching anal; pelvics reaching origin of anal. Caudal
subtruncate. Caudal peduncle longer than deep. An opercular
spot.
Here described from one of the types, probably a female, 160
mm. long. ‘The figured specimen, a male of 180 mm., differs in
having the first pelvic ray produced into a long flament and
in the. presence of two ocelli on the anal fin.
25. HAPLOCHROMIS MACRODON, sp. n. (Text-fig. 4.)
Pelmatochromis spekit (part.) Bouleng. Cat. Afr. Fish. iii. p. 417.
Depth of body 3 in length, length of head 25 to 23. Upper
profile of head somewhat concave. Snout 14 io 12 Ceamene: of
Text-figure 4,
Haplochromis macrodon, sp. 0.
eye, which is 43 to 43 in length of head, a little greater than
preorbital depth, equal to or a little less than depth of cheek ;
CICHLID FISHES OF LAKE VICTORIA. LG,
interorbital width 4 to 43 in length of head. Mouth oblique;
maxillary not extending to below eye; lower jaw projecting ;
teeth in 3 or 4 series in upper jaw, 2 or 3 in lower, outer conical,
strong, and set well apart anteriorly. Cheek with 3 or 4 series of
scales. 10 gill-rakers on lower part of anteriorarch. Pharyngeal
teeth slender. 31 to 33 scales in a longitudinal series, 5 or 6 from
origin of dorsal to lateral line. Dorsal XV 9-10; last spine
3 length of head or a little more. Anal ITI 8-10; third spine as
long as or a httle shorter than last dorsal. Pectoral ? head,
reaching origin of anal. Caudal truncate. Caudal peduncle 14
to 1§ as long as deep. Silvery; back darker; male with blackish
pelvic fins and 3 ocelli on anal.
Three specimens, 135 to 155 mm. jong, from Entebbe and
Munyonya.
26. HAPLOCHROMIS PROGNATHUS Pellegrin, 1905.
Paratilapia prognatha (part.) Bouleng. Cat. Afr. Fish. iii. p. 333,
fig. 224.
Depth of body 24 in length, length of head 22. Head 22 as
long as broad; upper profile slightly concave. Snout twice
diameter of eye, which is 5 in length of head, equal to preorbital
depth or interorbital width, ? depth of cheek. Mouth moderately
oblique; maxillary not extending to below eye; lower jaw
strongly projecting ; chin acute; teeth in 4 series in upper jaw,
3 in lower, some of the inner tricuspid, outer series conical; 60 in
upper jaw, anterior moderately strong. 3 series of scales on
cheek. 10 gill-rakers on lower part of anterior arch. 32 scales
in a longitudinal series, 5 from origin of dorsal to lateral line.
Dorsal XIV 10; last spine longest, 2 length of head; longest
soft rays 3 length of head. Anal III 9; third spine stronger
than and as long as last dorsal. Pectoral ? length of head,
extending toanal spines; pelvicsreaching anal. Caudal truncate.
Caudal peduncle 1} as long as deep. Silvery; back darker; an
opercular spot ; a dark bar below anterior part of eye.
One of the types, 170 mm. long, from Kavirondo Bay. A
specimen of 80 mm. from Hntebbe (Degen) seems to belong to
this species.
27. HAPLOCHROMIS MACULIPINNA Pellegr., 1913. (Text-fig. 5.)
Paratilapia maculipinna Pellegr. Bull. Soc. Zool: France, xxxvii.
p. dll.
Paratilapia prognatha (part.) Bouleng. Cat. Afr, Fish. iti. p. 333.
Depth of body equal to length of head, nearly 3 in length of
fish. Head 2 as long as broad ; profile slightly concave. Snout
1} diameter of eye, which is 4 in length of head, greater than
preorbital depth, equal to depth of cheek ; interorbital width 42
in length of head. Mouth obhique; lower jaw strongly projecting;
maxillary not quite extending to below eye: teeth triserial, some
Proc. Zoou. Soc.—1922, No. XII. 12
178 MR. C. TATE REGAN ON THE
of the inner tricuspid, outer series conical, 56 in upper jaw. 3 or
4 series of scales on cheek. 11 gill-rakers on lower part of anterior
arch. 33 scales in a longitudinal series, 7 from origin of dorsal
to lateral line. Dorsal XV 9; last spine longest, 3 length of
head; longest soft rays 3 length of head. Anal IIT 8; third
spine stronger than last dorsal, 4 length of head. Pectoral 2
length of head, extending to origin of anal; pelvies reaching
Text-figure 5.
Haplochromis maculipinna.
anal. Caudal truncate. Caudal peduncle 13 as long as deep.
Silvery ; back darker; an opercular spot anda lateral bana ; dark
spots on dorsal and caudal.
A specimen of 180 mm. from Bunjako.
The type, 156 mm. long, is described as having the eye 3+ in
length of head. :
28. HAPLOCHROMIS DICHROURUS, sp. n. (Text-fig. 6.)
Paratilapia serranus (part.) Bouleng. Cat. Afr. Fish. ii. p. 334.
Depth of body 33 in length, length of head 2%. Snout 12
diameter of eye, which is 5 inlength of head, equal to Saiecgrpiicl
width, greater than depth of przeorbital, less than depth of cheek.
Mouth moderately oblique ; premaxillary pedicels ending above
nostril ; maxillary extending to below anterior edge of eye;
lower jaw strongly projecting ; teeth in 3 series, outer Conical,
some inner tricuspid, 50 in outer series of upper jaw. 4 or 5
series of scales on cheek. 9 gill-rakers on lower part of anterior
arch. Pharyngeal teeth slender. Dorsal XVI 9; last spine
longest, less than 4 length of head; longest soft rays less than 3
length of head. Anal II] 8; third spine ? length of head.
Pectoral a little less than ? length of head, not quite reaching
anal. Caudal subtruncate. Caudal peduncle a little longer than
deep. 33 scales in a longitudinal series, 6 from first dorsal spine
to lateral line, 5 or 6 between pectoral and pelvic fins. Silvery ;
CICHLID FISHES OF LAKE VICTORIA. 179
back darker ; dark spots on snout and above and below eye; an
opercular spot; belly dusky; dorsal and upper half of caudal
Text-figure 6.
Haplochromis dichrourus, sp. n.
brown, anal and lower half of caudal bright red; an ocellus on
posterior part of anal; pelvics blackish.
Buganga, lL. Victoria (Degen).
A single specimen, 135 mm. in length.
29. HapLocHromis speKkit Bouleng., 1906.
Pelmatochromis svekii (part.) Bouleng. Cat. Afr. Fish. iii. p. 417,
fig. 285.
Depth of body 23 to 3 in length, length of head 22 to 23.
Head 27 to 22 as long as broad; upper profile straight or slightly
concave. Snout 14 to twice diameter of eye, which is 4 to 54 in
length of head, greater or less than preorbital depth, 1 to 12 in
depth of cheek ; interorbital width 4 (adult) to 5 (youn g)in length
of head. Maxillary extending to below anterior q of eye ; lower
jaw strongly projecting; teeth conical (adult) or outer bicuspid
and inner tricuspid (young), in 3 to 5 series in upper jaw, 2 to 4
in lower; 40 to 60 in outer series of upper jaw. Cheek with
4 or 5 series of scales. 9 gill-rakers on lower part of anterior arch.
Pharyngeal teeth slender. 31 or 32 scales in a longitudinal
series, 5 or 6 from origin of dorsal to lateral line. Dorsal XV
9-10; last spine longest, 3 (adult) to 2 (young) length of head;
longest soft rays a little more than 3 (@) or 2 (d) length of head.
Anal IIT 8-10; third spine 2 to 4 length of head, stronger than
7
7
last dorsal. Pectoral ? to 3 length of head, reaching origin of
anal or beyond ; pelvies reaching origin or anterior part of anal.
Caudal rounded or subtruncate. Caudal peduncle 11 to 13 aslong
as deep. Silvery; back darker; a dark opercular spot; usually
a dark lateral band from eye backwards, extending on caudal fin ;
a dark bar below anterior part of eye (¢); soft dorsal and caudal
12*
180 MR. C. TATE REGAN ON THE
sometimes spotted; pelvics and anal yellow (Q) or pelvics
blackish and anal greyish, with several ocelli posteriorly ( ¢).
A male of 235 mm. from Bunjako (specimen figured) ; a female
of 170 mm. from Entebbe, and four young (100-120 mm.) from
Entebbe and Jinja.
30. HAPLOCHROMIS SERRANOIDES, sp.n. (PI. IT.)
Paratilapia serranus (part.) Bouleng. Cat. Afr, Fish. iil. p. 354.
Pelmatochromis spekii (part.) Bouleng. t. c. 2 AIT.
Depth of body equal to length of Menlo ? in length of fish.
Snout with straight upper profile, 13 to twice diameter of eye,
which is 32 to Bd in leneth of head, cn adult less than depth of
prsorbital; interorbital “width 4 to 4% inlength of head. Mouth
moderately oblique; maxillary banal reaching vertical from
anterior edge of eye ; lower jaw projecting, moderately i in young
strongly 1 im ‘adult : teeth conical,in 3 or 4 series in upper jaw aa
2 or 3 in Jower, 40 to 60 in puter series of upper jaw. Cheek with
3 or 4 series of scales, once to 13 diameter of eye. 8 gill-rakers
and 2 rudiments on lower part of anterior arch. Pharyngeal
teeth slender. 382 scales in a longitudinal series, 5 or 6 from
origin of dorsal to lateral line. Dorsal XVI 9-10; last spine 4
to 2 3 length of head. Anal T{I 10-11; third spine 7 to more than
= head. Pectoral 2 to 2 = length of inewdl, reaching mene or origin
of anal. Caudal subtruncate. Caudal peduncle as long as deep.
Silvery or greyish; a dark opercular spot; spinous dorsal dusky ;
soft dorsal and anal dusky at the base, pale distally, the dark
colour with a well-defined undulating margin; caudal dusky at
base. Adult male with a blackish bar below the eye, blackish
pelvic fins, and ocelli on the anal fin.
Three specimens, 95 to 220 mm. in total length, from
Lake Victoria (Delmé Radcliffe) and between L. Kioja and
Murchison Falls (J/elland).
31. HAPLOCHROMIS ACUTIROSTRIS, sp. n. (Text-fig. 7.)
Paratilapia prognatha (part.) Bouleng. Cat. Afr. Fish. 111. p. 333.
Depth of body 3 to 3} in length, length iu head 22 to 24.
Snout 14 to 2 diameter of eye, which is 4 to 5Lin length of head,
in adult less than depth of preorbital or cheek; nterorbital
width 44 to 5in length of head. Mouth oblique, anteriorly above
level of eye; lower jaw strongly projecting; maxillary extending
to vertical from anterior edge of eye; teeth conical in adult,
some cuspidate in young, in 3 or 4 series in upper jaw and 2 or
3 in lower, 40 to 50 in outer series of upper jaw. Cheek with 4
to 6 series of scales. 8 to 10 gill-rakers on lower part of anterior
arch. Pharyngeal teeth slender. 81 to 33 scales in a longitu-
dinal series, 5 or 6 from origin of dorsal to lateral line. Dorsal
XV-XVI 8-10; last spine longest, 2 or a little less than ? length
of head, Anal IIT 8-10; third spinestronger and as long as and
CICHLID FISHES OF LAKE VICTORIA. 181
a little shorter than last dorsal. Pectoral # length of head,
nearly or quite reaching anal. Caudal rounded or subtruncate.
Caudal peduncle longer than deep. An opercular spot and a
Text-figure 7.
Haplochromis acutirostris, sp.u.
lateral band; vertical fing dusky, the soft dorsal and caudal
sometimes with clear spots; adult male with blackish bar below
eye, blackish pelvic fins and ocelli on anal fin.
Four specimens, 90 to 180 mm. in total length, from Bunjako
(Degen) and L. Salisbury (Jackson).
32. HaAPLOCHROMIS PLAGIOSTOMA, sp. n. (Text-fig. 8.)
Paratilapia longirostris (part.) Bouleng. Cat. Afr, Fish. 111. p. 332.
Depth of body 3 in length, length of head 27. Head a little
more than twice as long as broad; upper profile straight. Snout
Text-figure 8.
Haplochromis plagiostoma, sp. n.
14 diameter of eye, which is 43 in length of head, slightly greater
than preorbital depth, # depth of cheek; interorbital width 43
182 MR, C. TATE REGAN ON THE
in length of head. Mouth very oblique; lower jaw strongly
projecting ; maxillary not extending to below eye ; teeth conical,
4 series in upper jaw, 3 in lower; outer series regular, about 50
in upper jaw. 4 series of scales on cheek. 8 gill-rakers on lower
part of anterior arch. Pharyngeal teeth slender. Dorsal XV 9;
last spine longest, } length of head; longest soft rays 3 length of
head. Anal IIT 9; third spine stronger and a little shorter than
last dorsal. Pectoral ? length of head, reaching origin of anal ;
pelvies reaching vent. Caudal subtruncate, Caudal peduncle a
little longer than deep. 390 scales in a longitudinal series, 5 from
origin of dorsal to lateral line, 7 or 8 from base of pectoral to
middle of chest. Silvery; back darker, an opercular spot and a
dark lateral band.
A single specimen, 140 mm. long, from Bunjako.
33. HAPLOCHROMIS MACROGNATHUS, sp. n. (PI. ITI. fig. 2.)
Paratilapia longirostris (part.) Bouleng. Cat. Afr. Fish. ii. p.332.
Depth of body 32 in length, length of head 23. Head 3 times
as long as broad; upper profile nearly straight. Snout as long
as postorbital part of head, 24 diameter of eye, which is 6 in
length of head, a little less than preorbital depth, ? depth of
cheek; interorbital width 53} in Jength of head. Maxillary
extending to vertical from anterior margin of eye ; lower jaw very
strongly projecting, the anterior teeth exposed to the innermost
series; teeth conical, in 5 series in upper jaw and 4 in lower,
about 80 in cuter series of upper jaw. 5 series of scales on cheek.
9 gill-rakers on lower part of anterior arch. Pharyngeal teeth
slender. 32 scales in a longitudinal series, 6 from origin of
dorsal to lateral line. Dorsal XV 10; lastspine longest, # length
of head; longest soft rays 2 length of head. Anal III 9; third
spine stronger than last dorsal, } length of head. Pectoral 3
length of head, not reaching anal; peivies reaching vent. Caudal
subtruncate. Caudal peduncle 14 as long as deep. A dark
band connecting opercular spot with a spot on basal part of
caudal, another above lateral line and a dark stripe at base of
dorsal; dorsal and caudal with series of dark spots; pelvics
blackish ; two ocelli on posterior part of anal.
A single specimen, 195 mm. in total length, from Bunjako.
This species resembles H. mento in the strongly projecting
lower jaw, but differs in the much larger mouth, longer head,
shorter caudal peduncle, ete.
34, HapPLocHRoMIS DENTEX, sp.n. (PI. III. fig. 1.)
Paratilapia longirostris (part.) Bouleng. Cat. Afr. Fish. 11. p.332.
Depth of body 32 in length, length of head 23. Head 23
as long as broad; upper profile slightly convex. Snout nearly
twice diameter of eye, which is 5 in length of head, equal to
preorbital depth, less than depth of cheek; interorbital width 4
CICHLID FISHES OF LAKE VICTORIA. 183
in length of head. Snout decurved; mouth little oblique;
maxillary not far short of vertical from anterior margin of eye;
lower jaw strongly projecting, with the anterior teeth exposed.
Teeth conical, triserial, outer strong and set well apart anteriorly.
4 or 5 series of scales on cheek. 10 gill-rakers on lower part of
anterior arch. Pharyngeal-teeth slender. Dorsal XV 10; last
spine longest, 4 length of head; longest soft rays 3 length of head.
Anal IIT 9; third spine stronger than and as long as last dorsal.
Pectoral more than % length of head, ending above vent; pelvics
reaching origin of anal. Caudal slightly emarginate. Caudal
peduncle 12 as long as deep. 34 scales in a longitudinal series,
5 from origin of dorsal to lateral line, 10 from base of pectoral to
middle of chest. Silvery; back darker; an opercular spot and an
interrupted lateral band; dorsal and caudal greyish; pelvics and
anal yellow.
A specimen of 155 mm. from Sesse Isds. (Bayon). Two young
(60 and 80 mm.) from Entebbe may belong to this species.
30). HAPLOCHROMIS MENTO, Sp. n.
Paratilapia longirostris (part.) Bouleng. Cat. Afr. Fish. i.
p- 332, fig. 223.
Depth of body 34 in length, length of head 232. Head 2?
as long as broad ; upper profile slightly convex. Snout 23 diameter
of eye, which is 6 in length of head, equal to preorbital depth,
less than depth of cheek; interorbital width 44 in length of head.
Maxillary not nearly reaching vertical from anterior margin of
eye; lower jaw strongly projecting, with the anterior teeth
exposed; teeth conical, 5 series in upper jaw, 3 in lower,
outer strong and set well apart anteriorly. 4 series of scales
on cheek. 10 or 11 gill-rakers on lower part of anterior arch.
Pharyngeal teeth slender. 34 scales in a longitudinal series, 6 or
7 from origin of dorsal to lateral line. Dorsal XVI 10; last
spine longest, nearly 4 length of head ; longest soft rays 2 length
of head. Anal III 10; third spine stronger than last dorsal, 7
length of head. Pectoral 2 length of head, ending above vent ;
pelvics reaching origin of anal. Caudal rounded (?). Caudal
peduncle 1? as long as deep. Silvery ; back darker; an opercular
spot; pelvics dusky; 3 ocelli on posterior part of anal.
A specimen of 210 mm., from Bunjako.
36. HapLocHRoMIs CAvIFRONS Hilgendorf, 1888.
“Pelmatochromis cavifrons Bouleng. Cat. Afr. Fish. 1. p. 419,
fig. 287.
Depth of body about 3 in length, length of head 23 to 23.
Snout twice diameter cf eye, which is 5 to 53 in length of head,
nearly equal to preorbital depth, | depth of cheek ; interorbital
width 4 in length of head. Mouth very oblique; lower jaw
strongly projecting ; maxillary not quite reaching to below eye;
184 MR. CG. TATE REGAN ON THE
teeth mostly conical, in 4 or 5 series in upper jaw, 3 or 4 in lower,
60 to 70 in outer series of upper jaw. 5 or 6 series of scales on
cheek. 7 to 9 gill-rakers on lower part of anterior arch. 33 to
36 scales in a longitudinal series, 8 to 10 from origin of dorsal to
lateral line. Dorsal XV-XVI 8-10; last spine 3 head. Anal
ITI 8-9; third spine ? head. Pectoral 2 head, not reaching anal.
Caudal rounded or subtruncate. Caudal peduncle 1; to 14 as
long as deep. Body with numerous small irregular dark spots.
Seven specimens, up to 200 mm. long.
37. HAPLOCHROMIS ORTHOSTOMA, sp. n. (Text-fig. 9.)
Pelmatochronis spekii (part.) Bouleng. Cat. Afr. Fish. 111.
p. 417 (1915).
Depth of body nearly equal to length of head, 22 in length of
fish. Head 25 as long as broad; upper profile convex to above
anterior part of eye, thence straight. Snout longer than diameter
of eye, which is 5 in length of head, nearly equal to interorbital
width or preorbital depth. Mouth very oblique; maxillary not
quite reaching vertical from anterior edge of eye; lower jaw
strongly projecting, more than 4 length of head; teeth conical,
triserial, about 60 in outer series of upper Jaw. Cheek as deep
Text-figure 9.
Haplochromis orthostoma, sp. 1.
as long, with 4 or 5 series of scales. 9 gill-rakers on lower part of
anterior arch. Pharyngeal teeth slender. 33 scales in a longi-
tudinal series, 6 or 7 from origin of dorsal to lateral line. Dorsal
XV 9; last spine longest, 2 length of head. Anal IIE 8; third
spine stronger than and nearly as long as last dorsal. Pectoral
shorter than head, not reaching anal. Caudal rounded. Caudal
peduncle 13 as long as deep. Greyish; a dark bar below anterior
part of eye; pelvic fins blackish ; an ocellus on posterior part of
anal.
A single specimen, 115 mm. in total length, from Lake
Salisbury.
a
CICHLID FISHES OF LAKE VICTORIA. 185
38. HAPLOCHROMIS XENOSTOMA, sp.n. (‘Text-fig. 10.)
Paratilapia prognatha (part.) Bouleng. Cat. Afr. Fish. 111. p. 333.
Depth of body 34 to 32 in length, length of head 2 to 3.
Head 2% as long as broad; upper profile slightly concave. Snout
14 to 13 diameter of eye, which is 43 to 4% in length of head.
greater than prworbital depth, nearly equal to depth of cheek ;
interorbital width 42 in length of head. Mouth oblique; maxil-
lary not extending to below eye; lower jaw very prominent,
projecting upwards above end of snout; teeth in 3 series, inner
tricuspid, outer conical or some bicuspid, 50 to 56 in upper jaw,
Text-figure 10.
Haplochromis xenostoma, sp. i.
4 series of scales on cheek. 9 gill-rakers on lower part of anterior
arch. 33 scales in a longitudinal series, 6 from origin of dorsal
to lateral line. Dorsal XVI 8-9; last spine longest, ? length of
head; longest soft rays 2 length of head. Anal III 8; third
spine stronger than and as long as last dorsal. Pectoral 3 to 7
length of head, not reaching anal; pelvics nearly reaching vent.
Caudal subtruncate. Caudal peduncle 14 as long as deep.
An opercular spot and a lateral band ; small dark spots on soft
dorsal and caudal.
Two specimens, 105 and 125 mm. long, one collected by Sir
H. H. Johnston, the other from Entebbe (Degen).
39, HAPLOCHROMIS PELLEGRINI, sp. n. (Text-fig. 11.)
Paratilapia prognatha (part.) Bouleng. Cat. Afr. Fish. in.
p. 333,
Depth of body 32 in length, length of head 2? to 2%. Head
25 to 23 as long as broad ; upper profile slightly concave. Snout
12 diameter of eye, which is 43 in length of head, greater than
preorbital depth, equal to or a little less than depth of cheek,
equal to interorbital width. Mouth oblique; maxillary not far
short of vertical from anterior edge of eye; lower jaw strongly
projecting, but not above end of snout ; teeth conical, 4 series in
186 MR. C. TATE REGAN ON THE
upper jaw, 3 in lower, 60 in outer series of upper jaw. 3 or 4
series of scales on cheek. 8 or 9 gill-rakers on lower part of
anterior arch. 32 scales in a longitudinal series, 6 or 7 from
origin of dorsal to lateral line. Dorsal XIV 10; last spine
longest, 3 or a little more than 3 length of head: longest soft
rays 4 length of head. Anal III 9; third spine 2 or a little more
than 2 length of head. Pectoral 2 length of head, extending to
i
Text-figure 11.
Haplochromis pellegrini, sp. nu.
above vent; pelvics reaching origin of anal. Caudal truncate.
Caudal peduncle 14 as long as deep. An opercular spot; small
dark spots on dorsal and caudal; one specimen with 3 ocelli on
posterior part of anal.
Two specimens, 125 and 130 mm. long, from Entebbe (Degen).
Two others (75 mm.), from Entebbe, seem to belong to this
species ; they have XV 9 dorsal rays.
40. HAPLOCHROMIS ARGENTEUS, sp. n. (Text-fig. 12.)
Paratilapia longirostris (part.) Bouleng. Cat. Afr. Fish. iii.
p. 332.
Depth of body 3 to 33 in length, length of head 24 to 23.
Head 3 times as long as broad; upper profile slightly concave.
Snout 12 to 1? diameter of eye, which is about 44 in length of
head, slightly greater than preorbital depth, equal to depth
of cheek ; interorbital width about 5 in length of head. Mouth
oblique; lower jaw strongly projecting; maxillary not nearly
reaching vertical from anterior margin of eye ; 3 series of teeth in
upper jaw, 2 in lower, anterior inner teeth tricuspid, outer mostly
conical, some lateral ones bicuspid ; 46 to 60 in outer series of
upper jaw, the anterior rather strong. 3 or 4 series of scales on
cheek. 8 or 9 gill-rakers on lower part of anterior arch. Pharyn-
geal teeth slender. 32 or 33 scales in a longitudinal series, 5 or 6
from origin of dorsal to lateral line. Dorsal XV 9-10; last
CICHLID FISHES OF LAKE VICTORIA. 187
spine longest, a little less than + length of head. Anal III 9;
third spine stronger than and about as long as last dorsal.
Q » . =
Pectoral 2 to $ length of head, reaching vent. Caudal truncate.
Text-figure 12.
Haplochromis argenteus, sp. 1.
Caudal peduncle 14 as long as deep. Silvery; back darker ;
soft dorsal and caudal with or without series of small dark spots.
Two specimens, 115 and 140 mm. in total length, from
Bunjako.
4
fie.
to)
1. Hartocuromis Lonerrostris Hilgend., 1888. (Pl. IV.
i)
7)
Paratilapia longirostris (part.) Bouleng. Cat. Afr. Fish. 11.
p. 332.
Depth of body 34 to 33 in length, length of head 25 to 3.
Head 23 to 24 as long as broad; upper profile straight or shghtly
concave. Snout 12 to twice diameter of eye, which is 43 to 54
in length of head, equal to or a little more than preorbital depth,
equal to or a little less than depth of cheek ; interorbital width
4! to42 in length of head. Mouth oblique; lower jaw projecting ;
maxillary not extending to below eye; teeth conical (inner
tricuspid in young), 3 or 4 series in upper jaw, 2 or 3 in lower,
50 to 60 in outer series of upper jaw. 3 or 4 series of scales on
cheek. 10 or 11 gill-rakers on lower part of anterior arch.
Pharyngeal teeth slender. 33 scales in a longitudinal series, 6 or
7 from origin of dorsal to lateral line. Dorsal XV-XVI 9-10;
last spine longest, + to 3 length of head. Anal LIT 9; third spine
stronger than and as long as last dorsal. Pectoral 2to 7 length
of head, not reaching anal. Caudal truncate. Caudal peduncle
nearly twice as long as deep. Silvery; back darker; an oper-
cular spot ; soft dorsal and caudal sometimes spotted.
Three specimens, 100 to 160 mm. long.
188 MR. C. TATE REGAN ON THE
42, HAPLOCHROMIS GRACILICAUDA, sp. n. (PI. IV. fig. 1.)
Paratiiapia longirostris (part.) Bouleng. Cat. Afr, Fish. in.
p. 332.
Depth of body 4 to 4+ in length, length of head 3 to 33.
Snout 1! to 12 diameter of eye, which is 4 to 4% in length of head,
greater than preorbital depth ; interorbital width 47 in length of
head. Mouth oblique; lower jaw projecting; maxillary not
extending to below eye; teeth conical (cuspidate in young), tri-
serial, 50 to 60 in outer series of upper jaw. 3 or 4 series of
scales on cheek. 10 or 11 gill-rakers on lower part of anterior
arch, Pharyngeal teeth slender. 33 scales in a longitudinal
series, 6 from origin of dorsal to lateral line. Dorsal XVI 9-10;
last spine 2 to + length of head. Anal III 8-9; third spine as
long as or a little longer than last dorsal. Pectoral ? to + head,
not reaching anal. Caudal slightly emarginate. Caudal peduncle
2 to 23 as long as deep. Silvery; back darker; an opercular
spot; dorsal and caudal spotted.
Two specimens, 105 and 150 mm. long, from Bunjako and
Entebbe.
43. HAPLOCHROMIS XENODON Bouleng., 1911.
Bayonia xenodonta Bouleng. Cat. Afr. Fish. iii. p. 488, fig. 338.
Very near A. cinereus, distinguished principally by the
dentition. Outer teeth few and large, compressed, with long
anterior cusp directed inwards and posterior cusp very short or
indistinct.
Total length 90 mm.
44, HAPLOCHROMIS OBLIQUIDENS Hilgend., 1888.
Hemitilapia bayoni Bouleng. Cat, Afr. Fish. i. p. 491, fig, 840.
? Hemitilapia materfamilias (Pellegrin, 1913), Bouleng. t. c.
p. 341.
Very near H. nuchisquamulatus. Teeth in 4 to 6 series,
slender, distally expanded and compressed ; teeth of outer series
enlarged, obliquely truncated.
Total length 138 mm.
3. AsTATOREOCHROMIS Pellegrin, 1904.
As Haplochronvis, but with 4 to 6 anal spines.
L. Victoria.
ASTATOREOCHROMIS ALLUAUDI Pellegrin, 1904.
Haplochromis allwaudi Bouleng. Cat. Afr. Fish. ii. p. 305,
fig. 206.
Near H, gestri, especially distinguished by the increased number
CICHLID FISHES OF LAKE VICTORIA. 189
of dorsal and anal spines and the large blunt pharyngeal teeth.
Dorsal XVII-XIX 6-9. Anal IV—VI 6-9.
Total length 155 mm.
4, MACROPLEURODUS, gen. n.
Differs from Haplochromis in the dentition of the upper jaw,
which has an outer series of enlarged teeth and several inner
series of small teeth anteriorly and 3 or 4 series of enlarged teeth
laterally, which are exposed when the mouth is shut.
L. Victoria.
Text-figure 13.
Dentition of 1. Haplochromis sawvagii; 2. Macropleurodus bicolor ;
3. Hoplotilapia retrodens.
MAcropLEevRoDUS BIcoLoR Bouleng., 1906.
Paratilapia bicolor (part.) Bouleng. Cat. Afr. Fish. 11. p. 346,
fig. 234.
Paratilapia retrodens (part.) Bouleng. t. ¢. p. 347.
Depth of body 22 to 32 in length, length of head 3 to 33.
Snout decurved, as long as or a little longer than diameter of eye,
which is greater than preorbital depth and 4 in length of head ;
interorbital width 3 in head. Mouth wide; lower jaw a little
shorter than upper; maxillary extending to below anterior edge
of eye; 5 or 6 series of teeth in upper jaw, 4 or 5in lower. 3 or
4 series of scales on cheek. Gill-rakers short and stout, 7 or 8
on lower part ofanterior arch. Pharyngeal teeth small. 32 scales
in a longitudinal series, 6 or 7 from origin of dorsal to lateral
line. Dorsal XV 8-9; last spine ? length of head. Anal III 8-9.
Pectoral as long as head, reaching anal. Caudal truncate. Caudal
peduncle longer than deep. Olivaceous, with a faint dark lateral
band or with irregular dark cross-bars extending on to vertical
fins.
Lake Victoria.
Two specimens, 135 and 150 mm. long, from Bunjako.
I take the figured specimen of P. bicolor as the type and so
restrict the name to the species described above.
190 MR. C. TATE REGAN ON THE
5. Hoprormaria Hilgend., 1888.
Cnestrostoma Regan, 1920.
Differs from Haplochromis in the dentition. ‘Teeth small,
conical, in rather broad bands, which are well-developed on the
sides of the jaws, being formed of 3 or 4 series of teeth posteriorly.
L. Victoria.
HLoPLOrILAPIA RETRODENS Hilgend., 1888.
Paratilapia retrodens Hilgend. Sitzb. Ges. Nat. Fr. Berlin, 1888,
p. 76.
Hemichromis retrodens Pfeff. Nhierw. O.-Afr. Fische, p. 19 (1896).
Paratilapia polyodon Bouleng. Ann. Mus. Genova, (3) iv. 1909,
p. 306, fig., and Cat. Afr. Fish. i11. p. 849, fig. 236.
Paratilapia bicolor (part.) Bouleng. Cat. Afr. Fish. i. p. 346.
Lisp of body 24 to 3 in length, length of head 3. Snout once
to 14 diameter of eye, which is 34 to 4 in length of head, greater
sheen preorbital depth, about equal to depth of ‘cheek: inter orbital
width 8 to 34 in length of head. Jaws equal anteriorly ;
maxillary extending to below anterior edge of eye; teeth in 5 to
8 series in upper jaw, 6 to 10 in lower. 4 or 5 series of scales on
cheek. 8 gill-rakers on lower part of anterior arch. Pharyngeal
teeth small. 33 scales in a longitudinal series, 7 or 8 from orig
of dorsal to lateral line. Dorsal XV—X VI 9-11; last spine 4 to
2 length of head. Anal IIT 8-9. Pectoral as Jong as head,
reaching anal. Caudal truncate. Caudal peduncle longer than
deep. Traces of regular dark cross-bars and of a band along
middle of side and another above lateral line, or irregular blackish
cross-bars extending on to vertical fins.
Four specimens, 110 to 170 mm. in total length.
6. PLatyraniopus Bouleng., 1906.
Near Haplochromis, but jaws with very broad bands of small
Text-figure 14.
i 2
Dentition of 1. Haplochromis annectens ; 2. Platyteniodus degeni.
CICHLID FISHES OF LAKE VICTORIA. i91
conical teeth, that of the upper jaw broader at the sides than in
front.
L. Victoria.
PLATYTENIODUS DEGENI Bouleng., 1906.
Platyteniodus degeni Bouleng. Cat. Afr, Fish. iii. p. 426,
fig. 292.
Very near /aplochromis annectens, differing especially in the
dentition.
Total length 140 mm,
EXPLANATION OF THE PLATES.
Prater |. Haplochromis altigenis.
is Jif 3 serranoides.
5” WOT, ines, al, 53 dentex.
fig. 2. 3 macrognathus.
5 We alee, » gracilicauda.
fig. 2.
_ longirostris.
, ‘a ett Bali why
a hich, el ee, Reshegs sit
Hi Preaek: Tratgn sa grchcpa
- mai Pe Peas: EN a | :
i ete
iyi t
¥ ie :
Adis has edits Wie Sas ee :
ie 1 Sa aasateee Basia. ge Mes:
‘n: Raee me ae i tla ue
bs : PF i ie Ha suai
ON DIRECT DEVELOPMENT IN A DROMIID CRAB. 193
10. Direct Development in a Dromiid Crab.
By Srepaen K. Monteommry, B.A., B.Sc.*
[Received February 10, 1922; Read March 7, 1922.]
(Text-figures 1-3.)
The specimen here described is in the collection of the British
Museum (Natural History), and was submitted to me for
examination by the the kindness of Dr. W. 'T. Calman, to whom
thanks are also due for much assistance in studying it. The
specimen was presented to the Museum many years ago by the
late Dr. Henry Woodward. It is dried, and the only information
regarding its origin is the locality, “‘ Bass’s Strait,” given on the
label.
The specimen is an adult female Petalomera lateralis (Gray),
a species referred to by Haswell as “ very common ” in Australian
seas.
Under the abdomen are carried about 20 young in a post-larval
stage, which, although possessing the same general form as the
adult, differs from it in many details.
A. similar occurrence is recorded by Miss Rathbun § in a paper
to this Society in the case of the Oxyrhynch Vawioides serpulifera
(Guérin); this is the only other available record of such a case
among marine Brachyura.
The adult agrees entirely with the descriptions, and the figure
of Stimpson ||. It is figured here for comparison with the young.
Only one stage of development has been observed in this ease,
not two as in the case of WV. serpulifera.
The carapace of the young crab is longer than broad, the
measurements being about 175mm. long and 1:2 mm. broad, is
flatly convex, much flatter than in the adult, and in the centre
shows, under a high magnification, a minute reticulation. The
regions are well marked, again in contrast to the adult, and there
are numerous very fine hair-like spines on the upper surface.
The front consists of three very prominent forwardly directed
spines, each of which bears 5 to 8 pointed spinelets. The middle
spine is directed forward and downward and, with the two
lateral spines, forms a deep, almost V-shaped gutter between
the supra-orbital borders, the opening of the V pointing forwards
‘(text-fig. 1). me :
The supra-orbital border bears a similar prominent spine with
accessory spinelets, which is directed forward and outward
(text-fig. 1,@). Behind this spine the border runs almost directly
backwards; there is a slight protuberance bearing a few spinelets
* Communicated by Dr. W. T. Catman, F.R.S., F.Z.S.
+ For the transference of this species from Cryptodromia to Petalomera, cf. li. A.
Borradaile, Ann. Mag. Nat. Hist. (7) x1. 1903, p. 3v0._
~ W. A. Haswell, Catalogue of Aust. Stalk- and Sessile-eyed Crustacea, 1882, p. 139.
§ Mary J. Rathbun, Proc. Zool. Soc. 1914, p. 653, pl. u. figs. 9, 10.
|| W. Stimpson, Smithsonian Mise. Coll. 1907, xlix. pl. xx. fig. 3.
»
Proc. Zoot. Soc.—1922, No. XIII. 13
194 MR. S. K. MONTGOMERY ON DIRECT
about the middle of its length (text-fig. 1, b), opposite which is
the eyestalk. The border then turns almost at right angles
iaterally and ends ina prominent tooth, covered with spinelets,
forming the post-ocular tooth of the young specimen (text-fig. 1, c).
On the lateral border, behind this, another strong tooth bearing
spinelets occurs before the cervical groove is reached. Directly
posterior to the groove is another similar spine, which is not so
strongly marked.
No subhepatie tooth can be discerned.
In the adult there are two teeth on the antero-lateral border ;
of these, the most anterior (text-fig. 2, c) must represent the
post-ocular tooth of the young. The adult post-ocular tooth
(text-fig. 2,6) may represent either the protuberance on the
supra-orbital border of the young, opposite the eyestalk; in this
case, the sub-hepatic tooth is a later development, and the prom-
inent anterior supra-orbital tooth of the young (text-fig. 1, a)
must become reduced to whe weakly developed supra-orbital tooth
of the adult (text-fig. 2, d); or, alternatively, the eye in the
adult may have passed even more anterior ly, so that the post-ocular
tooth of the adult would represent the anterior supra-orbital
tooth of the young, In this case, the adult supra-orbital tooth
would be a new structure, developed from the spines in the region,
and the sub-hepatic tooth may arise from the protuberance on the
supra-orbital border opposite the eyestalk.
The chele in the adult bear on the carpus a strongly marked
tooth (text-fig. 2, e) on the external angle, a small bluntly
rounded tooth (f) forming the extremity of the upper border,
and between them a very strong forwardly directed tooth (9)
forming the most anterior part of the carpus. Posteriorly, on
the outer border of the car pus are two ridgelike teeth (/) standing
close together. In the young (text-fig. 3), the first three teeth
are represented by weakly developed protuberances, but no sign
can be distinguished of the two ridgelike teeth.
The hand of the young is smooth, except for a few small spines
and hairs. The movable finger on its outer margin bears five
sharp teeth, excluding that at the tip (text-fig. 3); there 1s an
indication of a sixth tooth behind. The fixed finger bears four
sharp teeth, with a fifth poorly developed, again excluding the
apical tooth. In the adult, the teeth of the fingers are exactly
similar, with the exception that they are slightly blunter, and
that the posterior poorly developed teeth are slightly more
prominent. The hand of the adult is grooved on both sides towards
each finger to receive the thick tomentum which in life covers the
carapace and the greater part of the limbs; this grooving, though
typical in the Dr Beds as a whole, is not pr axon in NS young.
The only description of the development of a Dromiid which is
available is that of Dromia vulgaris by Cano* in 1894.
According to him, the crab is hatched from the egg in a zoea
stage, from which a metazoea develops; this is followed by a
* G. Cano, Atti del Accad. Sci. Soc. Reale Napoli, ser, ii, vol. vi. 1894, No. 2
pls. i. & ii.
DEVELOPMEN'’ IN A DROMIID CRAB. 19
megalopa stage 5mm. in length. Cano then deseribes a post-larval
stage, hardly differing from the adult, except for the supra-orbital
spines and the middle spine of the rostrum which are not yet
developed.
Text-figure J.
S. HPA. dod.
Petalomera lateralis Gray. Post-larval stage. X 25.
Text-figure 2.
P. lateralis; «dult female. x 3.
The specimens here described look most: like the megalopa of
Cano, but are considerably more advanced.
The antennules (internal antenne of Cano) are exactly similar
to those of the adult, consisting of a peduncle of three joints anda
196 ON DIRECT DEVELOPMENT IN A DROMIID CRAB.
flagellum, which appears to be double, but owing to the brittleness
of the dried specimen this could not be determined with certainty.
The antenne consist of four joints, so far as can be made out,
with a squame and a flagellum, the squame being rudimentary
and similar to that in Cano’s figure.
Text-figure 3.
y = =
ay
~ i
ios he
E.° la
\
at
I” S.KMSeL.
Cheliped of post-Jarval stage.
woe
The abdomen is not extended and folded at the tip as in the
megalopa figured by Cano, but is tucked beneath the body in
typically Brachyurous manner. It bears four pairs of pleopods
similar to those of the adult; the telson is also like that of the
adult, there being simply a trace of the sixth pleopod visible
between the 6th and 7th segments, and not, as in Cano’s megalopa,
a distinct ramus on either side.
The last two pairs of legs are distinctly cheliform, and not
weakly subchelate, as in the megalopa.
Sternal grooves cannot be distinguished in the young, nor can
it be seen whether there is an epipodite on the cheliped.
The young in this case are definitely post-larval, and the
growth-changes which occur before reaching maturity are (@) the
broadening of the carapace until it is broader than long, (6) the
forward movement of the eye, (c) the effacing of the grooves
between the regions of the carapace, (d) the thickening of the
legs and chelipeds, and (e) the appearance of ridges on them.
The thorns and spinules of the young are generally absent in the
adult, which, on the other hand, in life was covered with a short
tomentum, which is not seen in the young.
It is very extraordinary that in a well-known species such as
this, no record should hitherto have been made of the direct
development of a post-larval stage from the egg. The possibility
that the larvee have hatched from the egg, and passedthrough zoea,
metazoea and megalopa stages clinging to the pleopods under the
abdomen of the female, may be ruled out for the reason that it
would be impossible for an adequate food-supply to be maintained.
And, in fact, no record has been found of P. lateralis or
NV. serpulifera having been captured with eggs, though this is a
common occurrence with other Oxyrhynchs and Dromiids,
numerous examples being recorded in the collections both of the
‘Sealark’* and the Ceylon Pearl Oyster Commission 7.
* Mary J. Rathbun, Trans. Linn. Soc. London, (2) Zool. xiv. pt. 2, 1911,
p. 191 sqq.
+ R. Douglas Laurie, Herdman’s Rep. Ceylon Pearl Oyster Fish. (Royal Soc.,
London) y. 1906, p. 349 sqq.
ON THE LIFE-HISTORY OF CUCULUS CANORUS. 197
11. Notes on the Life-History of Cuculus canorus, with
exhibition of eggs. By A. H. Evans, F.Z.8.
| Received February 7, 1922: Read February 7, 1922. ]
A further paper on the Common Cuckoo might well be con-
sidered superfluous, if it were not for the fact that an examination
of the literature on the subject shews that this is by no means
the case; many of the details of the bird’s life-history are found
still to need proof, and are merely reiterated from the pages of
previous authors, who in place of such proof make assertions,
which though highly probable, require confirmation.
The eggs now exhibited have been collected during a period of
more than twenty years, and give sure proof of the facts
pointed out below. s
Those of us who saw Mr. Edgar Chance’s admirable film, which
was shown in this room some weeks ago, and redounded equally
to his credit and that of his photographer, were initiated into
the ways of an individual hen Cuckoo when depositing her eggs
and were given a clue to the probable number of them; while a
meeting of the British Ornithologists’ Club next month is to be
devoted to the Cuculide as a Family: but the present paper
deals neither with an individual and its idiosyneracies, nor with
the whole of this world-wide Family, but with a species—our
British Cuckoo—and its regular habits.
[t is by no means easy to examine into the bird’s life-history,
but the writer has had unusual good luck. In the comparatively
small grounds of Histon Manor, near Cambridge, he and the owner,
Mr. W. A. Harding, found that no fewer than five hen Cuckoos
would lay their eggs in a single year, and that in one case the egg
was so remarkable that recognition was instantaneous. Hardly
less remarkable were specimens from the Cam, most of which
are now exhibited by kind permission of Dr. Ticehurst and
Mr. Bonhote, and are used to strengthen the evidence for the
conclusions arrived at.
Incidentally the abundance of hens at Histon tends to discredit
the theory of polyandry.
The first series of clutches passed round shews the same hen
laying in the nests of the Greenfinch, Spotted Fiycatcher, and
Pied Wagtail, and proves that she does not always choose the same
foster parent. The egg, in this case peculiar and unmistakable,
bears no resemblance to the others inthe nest. Such resemblance
is the exception rather than the rule, except where, as in the
ease of the Meadow-Pipit, the typical Cuckoo’s egg is more or
less similar to that of its host, whose nests are plentiful and
easily found. Several examples are exhibited.
The second series affords strong corroboration ; the Cuckoo’s
egg, though less peculiar, is in each case undoubtedly that of
198 MR. A. H. EVANS ON THE
the same bird, which shews a partiality for Robins’ nests (3 out
of 4).
The third and fourth series are exhibited for the same purpose ;
in the third the egg is again remarkable, in the fourth Hedge-
Sparrows are the favourite hosts.
The fifth series includes reddish or greyish eggs, of a type not
uncommon along four or five miles of the Cam. ‘The preference
here is for the nests of Sedge- or Reed-Warblers, and the simi-
larity of coloration may shew the range in their case of kindred
birds. The specimens were taken by Messrs. Richmond, Mills
and myself: where two occurred in the same nest, they were
clearly the produce of different hens.
The single egg next to these was laid in a Pied-Wagtail’s nest
after the young had flown, the sole instance in my experience.
With regard to the choice ot foster parents, it should be noted
that in the latter part of May and in June there is often little
real choice, especially on the moors. The Cuckoo must take what
she can get, and individual hens seem to have a restricted range,
if we are to judge by the eggs. ‘They appear to be somewhat
lazy birds, which follow the “line of least resistance,’ choosing
Reed- and Sedge-Warblers for hosts on rivers and lakes, Hedge-
Sparrows, Robins, Wagtails and the like in lanes and gardens,
Pipits on moors and commons, with a decided preference for a
small, comfortably lined nest. No one would look for Cuckoo’s
eggs in a dense wood, or low down ina thick hedge or pile of
sticks, where Hedge-Sparrows often breed.
An examination of the dates on which the specimens exhibited
were taken shews a still more important fact, namely, that the
identical hen returns for several years in succession to the same
grounds or even to the same copse. Here we have definite proof,
as opposed to assumption. ‘This fact is not only made evident
by the eggs from Histon Manor, but by the two very remarkable
series from the collections of Dr. Norman Ticehurst and Mr.
J. L. Bonhote.
In the first series, where the Cuckoo’s egg resembles that of
1 Spotted Flycatcher, the foster parent was invariably a Pied-
Wace The nests were in pollard Willows between Grant-
chester and Waterbeach, a range of some eight miles. An egg,
less richly coloured, also taken near Waterbeach, about ten years
earlier, is sufficiently near the types to justify the conclusion
that kindred birds occurred there before the dates on the rest of
the series.
The second series of large grey eggs with uniform small stipples
came from a clay-pit near Wianerbeach. and, as in the last case,
were found annually for several years in Sedge- or Reed-Warblers’
nests. Here, again, I exhibit a similar, though greener, egg taken
many years later, which apparently shews kinship in the bird
which laid it. Be this as it may, the dates of the main series
prove the annual return of a hen bird to the very same spot.
The number of eggs of the foster parent allowed to remain
LIFE-HISTORY OF CUCULUS CANORUS. ; 199
in the nest varies greatly. As shown by Hancock (N.H. Trans.
Northumb. and Durham, viii.) almost the whole elutch may be
undisturbed.
Although this article has been written to accompany an exhi-
bition of eggs, and to bring out certain points which have
received insufficient attention, without the intention of criticizing
the work of others, it is impossible to end without mentioning
the standard paper on Cuculus canorus by Dr. Hugene Rey,
published at Leipzig in 1892 as No. 11 of Marshall’s ‘ Zoologische
Vortrage. This laborious and elaborate piece of work may be
taken to comprise all that was known up to that date of the
bird’s nesting-habits, and contains long dissertations on the
number of eggs laid in a year, the date of deposition, their size,
weight, texture, and the composition of the shell, with lists of the
foster parents, the number of their eegs suffered to remain in the
nest during inenbation, and so forth. Tables and even graphs are
given to elucidate the text, and, finally, full details of the immense
number of specimens accumulated.
Admirable, however, though this paper is, it fails in the points
upon which I am now insisting. Sixteen out of Rey’s seventeen
conclusions may be taken as correct, but when he states that most
hen Cuckoos lay in the nests of definite species of foster-parents,
except on rare occasions, his theory runs counter to experience.
Of course the objection raised is only to his use of the word
“most,” and would have failed if he had said “ not uncommonly.”
Again, he tells us that the bird generally lays her eggs in the
same locality or even spot, that is, that she does not wander far
in the nesting-season. If he had only been able to add the
expression ‘‘in successive years,” he would have anticipated my
observations, made over a longer period than his.
| i i a mi . Hie
a) yeti) V iA
' (
ine +l i vee
a 3 eh Ny Ha
awe Sari
Aste <4 Tiara
ede: By, Gy i} 4 hh
| nh i mips aye ay Ae
SAAN TEES Pa
ila mis ne
THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 201
EXHIBITIONS AND NOTICES.
February 7th, 1922.
Dr. A. Smiru Woopwarp, F.R.S., Vice-President,
in the Chair.
The Secretary read the following Report on the Additions to
the Society’s Menagerie during the months ,of November and
December, 1921 :—
NovemMBeErR 1921.
The registered additions to the Society’s Menagerie during the
month of November were 221 in number. Of these 104 were
acquired by presentation, 66 were deposited, 5 were received in
exchange, 44 were purchased, and 2 were born in the Menagerie.
The following may be specially mentioned :—
2 Brown Capuchins (Cebus fatuellus), 1 Squirrel-Monkey
(Saimiris sciureus), 1 Hairy-rumped Agouti (Dasyprocta prym-
nolopha), and 27 Rough-eyed Cayman (Caiman sclerops), from
British Guiana, presented by Dr. G. M. Vevers on November
12th.
4 Lions (Felis leo), born in India, presented by the Jam Sahib
of Nawanagar on November 7th.
1 Puma (Félis concolor), from Chaco in the Argentine, pre-
sented by Arthur R. T. Woods, on November 11th.
1 Yellow-bellied Phalanger (Petaurus australis), from New
South Wales, presented by William Huntsman, F.ZS., on
November 21st.
1 Squirrel-like Phalanger (Petaurus sciureus), from New South
Wales, presented by Mrs. Edward Goldsmith on November 28th.
1 Great Kangaroo (Jucropus giganteus), 2 Wombats (Phasco-
lomys mitchelli), 2 Long-nosed Bandicoots (Perameles nasuta),
from Australia, 1 Tasmanian Devil (Sarcophilus harrisi), from
Tasmania, 1 Tawny Frogmouth (Podargus strigoides), from
Australia, 1 Six-plumed Bird-of-Paradise (Parotia lawesi),
2 Magnificent Birds-of-Paradise (Diphyllodes hunsteini), from
S.E. New Guinea; also 2 Magnificent Fruit-Pigeons (Megaloprepia
magnifica), and 2 White-fronted Bronze-winged Pigeons (Henz-
cophaps albifrons), from New Guinea, new to the Collection.
Purchased on November 15th. )
1 Long-tailed Roller (Coracias caudatus), from South Africa,
presented by the Marquis ef Tavistock, F.Z.8., on November
16th. New to the Collection.
2 White-bellied Sea-Eagles (Haliaétus leucogaster), from
Australia, presented by Alfred Ezra, O.B.E., F.ZS., on
November 15th.
2 Australian Lung-Fish (Ceratodus forsteri), from Queensland,
purchased on November 15th.
Proc. Zoou. Soc.—1922, No. XIV. 14
202 THE SECRETARY ON ADDITIONS TO THE MENAGERIE.
Decemper 1921.
The registered additions to the Society’s Menagerie during the
month of December were 79 in number. Of these 30 were
acquired by presentation, 39 were deposited, and 10 were pur-
chased,
The following may be specially mentioned :—
A pair of Lions (felis leo), from Kathiawar, India, presented
by H.M. The King, on December 20th,
2 Northern Lynxes (Felis lyne isabellinus), from Dharmo
Valley, Almora District, 10,000 ft., presented by P. Wyndham,
CHC Bab ales:
2 very large Grooved Tortoises (Vestudo calcarata), from
N. Nigeria, presented by The Emir of Katsina.
Mr. A. H. Evans, F.Z.8., exhibited, and made remarks upon,
a series of Cuckoos’ eggs taken near Cambridge.
Lord Currrorp or CHaupieren, F.Z.S8., exhibited, and made
remarks upon, a series of photographs of Vototheriam mitchelli.
Miss L. E. Cuzpsman, F.E.S., gave an account of the position
and funetion of the Siphon in the Amphibious Molluse, Ampul-
laria vermiformis.
Mr. G. C. Rosson, F.Z.8., exhibited, and made remarks upon,
a series of models demonstrating the respiratory mechanism of
Ampullaria vermiformis.
February 21st, 1922.
Prof. EK. W. MacBripz, D.Sc., LL.D., F.B.S.,
Vice-President, in the Chair.
The Secrerary read the following Report on the Additions to
the Society’s Menagerie during the month of January, 1922 :—
The registered additions to the Society’s Menagerie during the
month of January were 151 in number. Of these 52 were
acquired by presentation, 76 were deposited, and 23 were
purchased.
The following may be specially mentioned :—
1 Macedonian Wolf (Canis lupus), from Maeedonia, presented
on January 14th by Lt.-Col. F. L. Giles, R.E., British Repre-
sentative Serbo-Bulgarian Boundary Commission, to whom it
was given by King Boris of Bulgaria.
ON THE INDIAN TORTOISE-BEETLE. 203
1 Dybowski’s Deer (Cervus hortulorum), from Manchuria, bred
at Horsham, purchased on January 2nd.
11 Plumbeous Quails (Synacus plumbeus), from S.E. New
Guinea, new to the Collection, deposited on J anuary 21st.
2 Angel-Fish (Petrophyllum scalare), new to the Collection,
from Brazil, purchased on January 30th.
The Srcrerary exhibited, and made remarks upon, a photo-
graph of the Society’s Gardens taken from an aeroplane.
Miss L. EK. Curzsman, F.E.S., exhibited, and made remarks
upon, living specimens of the Amphibious Mollusc, Ampullaria
vermiformis.
Mr. G. ©. Rozson, F.Z.8., described in fuller detail the series
of models demonstrating the respiratory mechanism of Ampul-
laria vermiformis exhibited at the previous Scientific Meeting,
and made remarks upon the respiratory mechanism of the
Ampullariide.
Mr. F. Martin Duncan, F.R.M.S., F.Z.S8., exhibited a series of
cinematograph films he had recently taken in the Society's
Gardens, showing the movements of Ampullaria vermiformis ; of
the Freshwater Crab, Cardisoma armatum; and of the Hawk's
bill Turtle, Chelonia imbricata.
Miss L. KE. Currsman, F.E.S., described the habits, in captivity,
of the Freshwater Crab, Cardisoma armatum.
March 7th, 1922.
Sir Stipvey F. Harmer, K.B.E., F.R.S., Vice-President,
in the Chair.
Morr Catan RnGan, FUR.S. RUZ.Ge exhibited, and made
remarks upon, some living specimens of the Indian Tortoise-
Beetle. Aspidomorpha sancte-crucis.
ROLES
ei
A ae a aut
OF THE
- PROCEEDINGS
|
OF THE
OF LONDON.
Cvrk
: WiNV
4 YU
7 wy
Oy
Pay Ee
1385 Texr-FIGURES.
‘JUNE 1922.
PRINTED FOR THE SOCIETY,
SOLD AT ITS HOUSE IN REGENT’S PARK,
; LONDON:
PATERNOSTER ROW.
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
ZOOLOGICAL SOCIBTY
Va ES:
ASOnian D
} ie =
CONTAINING Paces 205 to 481, wivH 10 PLatTEs aNnp
MESSRS. LONGMANS, GREEN, AND CO.
741922
?
ee [Price Twelve Shillings. }
=.
LIST OF. CONTENTS,
1922, Part II. (pp. 205-481).
EXHIBITIONS AND NOTICES.
The Secrerary. Report on Additions to the Society’s Menagerie during the month of
ebriany l922 arrestee.
My, ©. Tarr Reeay, M.A., F.R.S. Exhibition of a series of lantern-slides illustrating
specimens of various blind fresh-water Fishes
Ce er ee
Mr. R, H. Burns, M.A., F.Z.S. Exhibition of specimens demonstrating the recessus
orbitalis in Flat Fishes
ee eC ec Ca
Dr, L. Hocsny, M.A., F.Z.S. Exhibition of, and remarks upon, some small examples of
metamorphosed Mexican Salamanders (Amblystoma tigrinum)
ee ee eee ee tw we ce ae
The Szcrerary. Report on Additions to the Society’s Menagerie during the month of
March, 1922
a i eC er rr i er we
The Secretary. Exhibition of some dressed skins of a Tree-Hyrax (Procavia valida)
from Tanganyika Territory
eC ee eer ry
Prof. P. T. Fuyyy, Jxhibition of, and remarks upon, a cast of the skull of a Squalodont
Whale
i i
The Sucrerary. An account of Mr. Loveridge’s experiences while watching “ Lions at
their Kill”
ee te ree mee te PE OH ew Dee ew ee wwe mH eee OL er BOB eesereesoreseceenrene
Page
479
479
466
481
48]
Contents continued on page 3 of Wrapper.
THE NEMATODE PARASITES OF ELEPHANTS, 205
12. A Revision of the Nematode Parasites of Hlephants, with
a description of four new species*. By M. Kwatit,
SIDI, IO IP Lae SO OUEIES ie ISL)
(From the Helminthological Department of the London School of
Tropical Medicine.)
[Received March 2, 1922: Read April 4, 1922.
(Text-figures 1-71.)
Since Cobbold’s adinirable work on the parasites of the Indian
elephant in 1882, our knowledge of the Helminths harboured
by this species of elephant has been considerably enlarged
by Lieut.-Col. C. Lane, Railliet & Henry, and others. Little
advance has been made regarding the parasites of the African
elephant. ‘The new species recorded here, with one exception,
are parasites of the latter animal.
The material upon which this study is based was collected by
Prof. Leiper in Uganda as far back as 1909 from two elephants
shot by him. Im addition, specimens received from Dr. J. J.
Simpson, West Africa, the Raymond Laboratories in India, and
from Lieut.-Col. C. Lane, the lattex’s representing most of his type-
species from the Indian elephant, were available for study and
comparison. Leiperenia galebi was collected by Prof. Leiper from
an Indian elephant that died in the London Zoological Society’s
Gardens.
Besides all this unique material, Prof. Leiper placed at my
disposal his extremely valuable notes on the type-species in the
Vienna and Berlin Natural History Museums. These proved to
be of great assistance with regard to Belascaris lonchoptera
Diesing, and Murshidia linstowt, sp. n.
I feel deeply indebted to Prof. Leiper for his unrestricted
generosity and ever-ready help both as regards material and
information, without either of which this work could not have
been done.
The nematode parasites of the elephants are peculiar to these
hosts, and it is an interesting fact that the parasites found in the
African and Indian elephants never belong to the same species.
Grammocephalus clathratus, which was formerly supposed to occur
in botn species of elephants, has lately been shown by Lane to
be represented in the Indian elephant by a separate species.
Murshidia faleifera is another example. In this paper it is
shown that the African elephant harbours two species of Mir-
shidia, one of which apparently was mistaken for JMurshidia
falcifera.
* Communicated by Prof. R. T. Lerppr, D.Sc., M.D., F.Z.S,
7 In a preliminary note published in the Ann. & Mag, Nat. Hist. for Feb. 1922,
brief diagnoses were given of two new genera and seven new species which are here
fully described and illustrated.
Proc. Zoou. Soc.—1922, No. XV. 15
206 DR. M. KHALIL ON THE NEMATODE
There is some confusion regarding the correct zoological
terminology of the species of elephants. For this reason the
terms African and Indian are used throughout this work.
NEMATHELMINTHES.
Class NEMATODA.
Super-family Ra4BDIASOIDEA Railliet, 1916.
Family ATRACTID# Travassos, 1919.
This family name was given by Travassos to include the
following genera :—Atractis, Ozolaimus, Rodonia, Labiduris,
Crossocephalus, Macracis, Cobboldina, and Cyrtosomum, to which
IT add the genus Leiperenia.
Travassos’ diagnosis of the family runs as follows :—Cisophagus
with an anterior and a posterior bulb; viviparous, the female
genital system is generally single; vulva is placed posteriorly or
in common with the anus; parasites of vertebrates.
Genus LereErEeNtA Khalil, 1922.
Small nematodes just visible to the naked eye. The males are
slightly smaller than the femalés. Mouth is surrounded by more
than six lips. Cisophagus is divided into an anterior, short, and
strongly muscular portion, to which the name pharynx is given.
The posterior portion of the esophagus is about three times as
long as the pharynx. The anterior end of the body is provided
with a cuticular membranous expansion on either side. There
is a large excretory vesicle. The excretory pore is raised on a
apilla which is striated radially.
Male: The caudal extremity of the male is curved. There are
two unequal spicules and an accessory piece. There are four
papillz on the tail.
Female: Vivipareus, and the embryos reach an advanced
stage of development im wtero. The vulva is placed in the
posterior part of the body, separate, and a short distance in
front of the anus.
Type-species, Leiperenia leiper: (from the African elephant).
LEIPERENIA LEIPERI Khalil, 1922. (Text-figs. 1-4.)
Material.—The material consists of two males and three
females, taken from Prof. Leiper’s collection of Elephant
nematode parasites.
Shape of body.—These nematodes are very small and just
visible to the naked eye. The males are 3°8 mm. long. The
females are slightly longer, being 3°9 mm. in length. The
maximum diameter of the body is about the middle: it is*2 mm.
in the male and-:21 mm. in the female. The body narrows
slightly as if is traced towards the head end, which is truncated
PARASITES OF ELEPHANTS. 207
in appearance. Posterior to the head a cuticular expansion is
present on either side of the body. The female is practically
straight ; its tail is very long and is gradually attenuated to a
fine point. The caudal end of the male is bent towards the
ventral surface, and its end is more rounded than in the female.
Skin.—The cuticle is finely striated at intervals of -007 mm.
throughout the whole length of the body. The cuticular
expansions near the cephalic end of the body are placed on both
lateral sides, and extend from the level of the pharynx to prac-
tically the middle of the esophagus. They are *3 mm. in length
and ‘037 mm. maximum breadth, and are striated.
Text-figure 1.
Leiperenia leiperi Khalil. Cephalic end.
Mouth collar—Yhe mouth collar is very short and has a
rounded shape. It is ‘012 mm. long and -06 mm. in diameter.
The mouth-opening is practically circular and is surrounded by
ten small lips. The lip in the mid-ventral line and that in the
mid-dorsal line are the largest and broadest. The lips placed on
either side of these lines are smaller and rounded. There is no
mouth capsule.
Pharynx.—This is a short muscular canal. Its musculature is
more closely set and of a finer texture than that in the succeeding
esophagus. It is also distinctly demarcated from it by a deep
groove. For these reasons | named that specialized part of the
cesophagus the pharynx. The shape of the pharynx is cylindrical,
with a small bulging in the middle. It is ‘083 mm. long and
"06 mm. maximum diameter. From its cephalie end project
conical processes ; their apices are directed antero-laterally. The
number of these processes and their structure can only be
ascertained under the oil immersion lens. They are eight in
number surrounding the mouth-opening, and they are muscular
La
208 DR. M. KHALIL ON THE NEMATODE
processes from the pharynx. The lumen of the pharynx is
continuous with that of the esophagus.
(Hsophagus.—Vhe csophagus is long and thin. In some
specimens it pursues a wavy course. Its posterior end is a little
swollen. It is ‘38 mm. long and ‘073 mm. maximum diameter.
Text-figure 2.
Leiperenia leiperi Khalil. Anterior part of the body.
Chyle intestine.—‘vhe intestine is irreguiarly bent in its course.
Near its commencement its wall is surrounded by a clear
refractile band, the nature of which is unknown. The cellular
wall is slightly pigmented. The rectum is a short canal; in most
cases its structure is obscured by the greatly distended genital
organs of the female.
Hecretory system.—The large excretory vesicle receives two
excretory ducts, one from the lower part of the body and one
from the head end. The excretory pore is wide and is placed
"92 mm. from the cephalic end. It is placed on a raised papilla,
striated radially round the pore.
Nerve collar.—TVhe nerve collar surrounds the thinnest part of
the cesophagus *22 mm. from the head end.
Genital organs.—Male: The irregularly convoluted testis
reaches within -8 mm. from the head. The cement gland is
long, and the ejaculatory duct opens into the ventral aspect of the
cloaca.
Female: In the mature female the genital tract is so distended
with embryos that the details of this system are practically
impossible to make out. By the help of an immature specimen
the general features of the organs can be recognized. There
is only one ovary and one uterus, placed in the axis of the
body. The vagina is short, and opens in the posterior part of the
body +1 mm. cephalad of the anus.
Spicules.—There ave two unequal spicules and an accessory
PARASITES OF ELEPHANTS, 209
piece. The longer spicule is-3 mm.in length. It begins close to
the dorsal surface, and lies parallel to 1t for about two-thirds of
its course. Then it bends ventrally to pass into the cloacal canal
anterior to the accessory piece. he shorter spicule is*19 mim. in
length. - It lies by the side of the terminal portion of the longer
spicule. Its thick cephalic end is bent ventrally, and is thus
seen in the concavity of the longer spicule. The accessory piece
is ‘09 mm. long. Its cephalic extremity is large and rounded; its
end is attenuated to a sharp point.
Text-figure 3.
100
Leiperenia leiperi Khalil. Caudal end of male.
The tail.—The male tail is curved, forming a semicircle. It
is *38 mm. long. Its extremity is rounded; -16 mm. from the tip
of the tail there are four papille. Two of these are placed near
the mid-ventral line, one slightly caudal to the other. The other
two are placed one on either side of the tail. These are all simple
papille and project very little above the surface.
The female tail is very much longer and thinner than that of
the male, being *7 mm. in length; its termination is very fine.
There are four papille on either side of the anus. No other
papille could be detected on the female tail,
The Embryos.—The females are viviparous ; the embryos reach
an advanced stage of development while they are still in the
uterus. Six embryos can be seen in the uterus at one time.
They distend that organ enormously, and make identification of
the neighbouring structures difficult, 1f not impossible. Some of
these embryos measured °52 mm. in length, and -06 mm. in
210 DR. M. KHALIL ON THE NEMATODE
diameter. The pharynx, cesophagus, and a straight gut can be
easily recognized in them.
Habitat. The intestine of the African elephant (Uganda).
Text-figure 4.
Leiperenia leiperi Khalil. Tail end of female.
LEIPERENTA GALEBI Khalil, 1922. (Text-figs. 5-7.)
Material.—The material consisted of ten specimens collected
by Prof. Leiper from an Indian elephant that died in the London
Zoological Society’s Gardens. They were the only Helminth
parasites found in that elephant.
Shape of body.—These nematodes are just visible to the naked
eye. The male is smaller than the female. It is 3°25 mm.
long, while the female is 3°8 mm. The maximum diameter is
attained near the middle of the body. It is *17 mm. in the
male and ‘18 mm. in the female. The head end is truncated,
while the caudal is attenuated to a fine end, more delicate in
the female than in the male. The caudal end of the male is
curved ventrally. The head end of the body has a lateral
cuticular expansion on either side.
Skin.—The cuticle is striated throughout its length, except for
a short distance at the cephalic extremity. The striations are at
regular distances of °008 mm. apart. The cuticle is expanded in
a membranous fashion on either side of the cephalic end of the
body fora distance of -6 mm. It is striated in its greater extent,
the part corresponding in level with the pharynx is not striated.
The cuticular expansion is ‘023 mm. in breadth.
Mouth collar—The mouth collar is distinctly separated from
PARASITES OF ELEPHANTS. ANAL
the body bya groove. It is 0:18 mm. long and ‘08 mm. in diameter.
It has a rounded margin. The oral opening is circular and is
surrounded by ten rounded lips. The one in the mid-ventral
line and also that in the mid-dorsal line are broader and divided
into two by a superficial depression. There is no oral cavity.
Text-figure 5.
Leiperenia galebi Khalil. Cephalic end.
Pharynx.—The short cylindrical muscular pharynx is "14 mm.
in length and-] mm. maximum diameter. It is thus much
longer and broader than in J. leiperi. From its cephalic end
spring eight cone-shaped processes that surround the mouth-
opening. The fine apices of these processes correspond more or
less to the interlabial depressions.
(sophagus.—The thin muscular cesophagus is irregularly bent.
It is °39 mm. long in the male and -4 mm. in the female. Its
maximum diameter is at posterior end, and measures in the male
-06 mm. and in the female ‘07 mm.
Chyle intestine —The intestine pursues an almost straight
course. The cells of the intestinal wall are slightly pigmented.
The short rectum is indistinctly marked from the intestine.
Exeretory system.—There is a large excretory vesicle similar to
that described inthe type-species. The excretory pore is placed
on a large raised papilla 1-2 mm. from the head end. The
papilla is radially striated round the pore.
212 DR. M. KHALIL ON THE NEMATODE
Nerve collar.—The small nerve collar surrounds the thinnest
part of the cesophagus ‘25 mm. from the head end.
Genital organs.—Male: The convoluted testis gives rise to the
ejaculatory duct. This pierces the enormously elongated cement
gland to end in the cloaca.
Female: The female is viviparous; the embryos reach an
advanced stage of maturity, and so distend the uterus as to make
the examination of other structures difficult. The convoluted
ovary 1s single, and leads to the enormously dilated uterus. The
short vagina opens on the surface of the body :06 mm. in front of
the anus. The convolutions of the ovary reach within 52 mm.
of the head end.
Text-figure 6.
Leiperenia galebi Khalil. Tail end of male.
Spicules.—There are two unequal spicules and an accessory
piece. The longer spicule is -25 mm. long; thus it is shorter
than that of the type-species. It lies in the axis of the body,
midway between the ventral aud dorsal surfaces. Near its
lower end it bends boldly ventrally to pass into the cloaca. The
shorter spicule is °13 mm. jong. It is very slightly curved, lying
by the side of the terminal portion of the longer spicule. The
accessory piece is ‘13 mm. long. It has a thick bulbous end, and
is gradually attenuated to end in a sharp point buried in the
posterior lip of the cloacal opening. It lies practically across the
axis of the body.
The tail.—The male tail is gracefully curved and is *43 mm.
in length. itsendis biuntly rounded. ‘There area group of four
papille similar to those in ZL. lewperi. They are situated -17 mm.
from the tip of the tail. Two papille are in the middle line, one
PARASIVES OF ELEPHANTS. 213
slightly anterior to the other. Of the other two, one is placed
on either side of the body at the same level.
The female tail is short and conical, unlike that of Z. leipert.
Its tip is pointed. It is 53 mm. long. The anal opening is not
vaised or surrounded by prominent lips. On either side there are
four papille.
Text-figure 7.
Leiperenia galebi Khalil. Tail end of female.
Embryos.—The uterus is crowded with these giant embryos.
They attain -48 mm. in length. The various parts of the
digestive tract can be easily made out in them.
Habitat. Intestine of Indian elephant (India; died in the
London Zoological Gardens).
Discussion.
This genus has some points in common with the genus dA tractis.
The latter was made by Dujardin to include Ascaris dactyluris of
Rudolphi, 1819, but was not defined. The genus was defined by
yon Linstow in 1902. From von Linstow’s description, together
with a detailed account of the type-species by Railliet and Henry,
Leiperenia can be easily separated by the following characters :—
In Atractis there are six lips surrounding the mouth. There
are no cuticular membranous expansions in the anterior part of the
body. The esophagus is divided into two portions, of which the
anterior is the longer. ‘The number of the conical processes is
limited to six. There isa cuticular expansion along the posterior
two-thirds of the body. he species of Atractis are parasitic in
Land and Freshwater Tortoises and Reptilia.
214 DR. M. KHALIL ON THE NEMATODE
The two species of Leiperenia differ mainly in the length of the
spicules and accessory piece, and particularly in the shape of the
female tail, which is very long and narrow in JZ. leiperi and
stumpy and broad in Z. galebi.
Some of the genera included in the family Atractide are
credited with the ability of reproduction and maturation in the
same host. They are found in tremendous numbers in the host.
Super-family Ascarorpe4 Raill. & Henry, 1915.
Family AsScARID# Cobbold, 1864.
Sub-family AscArina Travassos, 1913.
Genus Brxascaris Leiper, 1907.
BELASCARIS LONCHOPTERA (Diesing, 1851) Leiper, 1911.
Strongylus elephanti Rud., 1819.
Ascaris lonchoptera Diesing, 1851.
Belascaris lonchoptera Leiper, 1911 (unpublished notes).
This was the first Helminth parasite recorded from the elephant.
It was listed by Rudolphi in his ‘Synopsis Entozoorum’ under
doubtful strong yles. The specimens were collected from the bile-
ducts of an Indian elephant and deposited in the Museum of
Vienna.
In 1847, Jackson recorded in the ‘ Descriptive Catalogue of
Boston Medical Improvement Society’ the presence of Ascaridz
and Flukes in the Indian elephant.
In 1851, Diesing described the doubtful specimens of Rudolphi,
and found them fa be female Ascaride and not Strongyles. He
named them Ascaris lonchoptera. He also referred to the
Ascaride of the Boston Catalogue as the same species.
In 1882, Cobbold expressed a doubt if such an illustrious man
as Rudolphi would have mistaken an Ascaris for a Strongyle.
In the same year Drasche redescribed and figured A. lonchoptera
from the Vienna collection, and left no doubt as to its Ascarid
nature, thus confirming Diesing’s observations.
Leiper, in unpublished notes to which I have had access, records
the results of his examination of the same specimens in the
Vienna Museum, and coneluded that they belonged to the genus
Belascaris (Leiper, 1907).
Although a good deal of attention has been paid lately to the
elephant parasites, especially in India, no Ascaridze have been
reported. So far, the male of Lelascaris lonchoptera remains
undescribed.
Habitat. Bile-ducts of the Indian elephant.
PARASITES OF ELEPHANTS. 215
Super-family Sprrororpea Raill. & Henry, 1915
(emend. Hall, 1916).
Family Sprrurip& Oerley, 1885.
Sub-family Sprrurin# Railliet, 1915,
Genus PArABroneMA Baylis, 1921.
Spiruride: Polymyarian worms, having the mouth bordered
by paired lateral lips, external to which there are a dorsal and a
ventral shield of cuticle. Each lip has one large median lateral
papilla and a pair of small sublateral papille. Of the dorsal and
ventral shields, each carries a pair of larger papille, situated
some distance behind the extremity of the head. The cuticle of
the head is thick, and folded in a complicated manner so as to
form a circlet of six horseshoe-shaped auricular appendages, of
which two are lateral, two subventral, and two subdorsal.
The body is rather slender, tapering rather more in front than
behind. The cuticle is transversely striated. The buccal cavity
a short distance from the oral aperture is transversely elongated,
then passes into a long cylindrical cuticular tube. The
cesophagus consists of a short, narrow anterior portion and a
long, somewhat wider posterior portion. The anterior portion
is surrounded by the nerve-ring.
The tail of the male is coiled ventrally into a spiral. The
spicules are markedly unequal. A somewhat triangular accessory
piece is present. ‘here are six pairs of papillee, four pre-anal
and two post-anal. There is also a large median double papilla
immediately in front of the cloacal opening.
The female is considerably larger than the male. The tail is
short and curved towards the dims sal side. The vulva is in the
region of the end of the cesophagus. The vagina is long and
narrow. It has a curious U-shaped bend in “Es course, a “short
distance from the vulva. The two uteri are parallel, running
backwards at first, one of them returning towards the anterior
end of the body. The female is viviparous. The uterus contains
immense numbers of embryos.
Type-species, Parabronema indicum.
PARABRONEMA INDICUM Baylis, 1921.
The male‘is 7°9 mm. and the female is 13 mm. in length.
The vulva is situated a little caudad of the end of the
esophagus. The longer spicule of the male is 0°93 mm. long
and the shorter is 0°39 mm. The tail of the male is 0°17 mm. and
that of the female is 0°32 mm. in length.
Habitat. Stomach-wall of the Indian elephant (India).
216 DR. M. KHALIL ON THE NEMATODE
PARABRONEMA AFRICANA Baylis, 1921.
This species was apparently mistaken by Baird for female
Grammocephalus clathratus. It was isolated and described from
the original material by Baylis in 1921, but Leiper in his
notes in 1911 recorded and drew this species from material in
the Berlin Museum, where it was kept under the name Ascaris
lonchoptera. He recognized this mistake in diagnosis, and added
that the same species is included wrongly by Baird in his type
of G. clathratus in the Natural History Museum of London.
The male is 40 mm. and the female is 57 mm. in length.
The lips have a median inward projection on their opposed edges.
Vulva is placed just caudad or cephalad of the end of the
esophagus. The left spicule is 3°15 mm. long and the right is
0°68 mm. The male tail is 0°85 mm. and the femalé tail is
0°52 mm. in length.
Habitat. Stomach-wall of the African elephant (died in
London).
PARABRONEMA SMIrHit Cobbeld, 1882. (Text-figs. 8-12.)
Filaria smithii Cobbold, 1882 (mot Filaria smitht Sambon,
1907, from the Grouse Lagopus scoticus).
Spiroptera smithi Raill., Henry & Bauche, 1914.
Parabronema smithi Baylis, 1921.
Source of Material—The material consisted of one male and
three females sent from the Raymond Laboratories, Calcutta.
They were collected from an Indian elephant.
Shape of body.—Very small nematodes. The females much
larger than the males The male is 4:1 mm. long and the
female 8 mm. The maximum diameter of the body is about
the middle of the worm, being 0-2 mm. in the male and -26 mm.
in the female. The female is always curved, the tail directed
dorsally. The tail of the male is spirally coiled ventrally.
Skin.—The cuticle is very finely striated throughout the
length of the body.
The head.—The cuticle surrounding the head end is raised from
the surface of the body, forming a complicated design. Its caudal
margin surrounds the circumference of the body 0-083 mm. from
the head end. It forms six auricular appendages, two lateral,
two subventral, and two subdorsal. Hach auricle is open towards
the head. ‘Their posterior margin is nearly flat, unlike the
rounded appearance in P. indicum or the angular appearance in
P. africana. Hach of the two lateral lips bears three papille, a
large one near the centre and two smaller ones near the margin
on either side. ‘The cuticular shields lower down bear four
papille. The terminal extremity of the head is 0°03 mm, in
diameter.
Cervical papilie.—The cervical papille are placed 0°209 mm.
from the head end.
Digestive system.—There is a buccal cavity just behind the oral
PARASITES OF ELEPHANTS. PATE
aperture. This leads into a long cuticular tube leading into the
esophagus. The length of that part of the digestive system is
‘limm. The wsophagus consists of two portions. The cephalic
part is thin and practically straight. It is 0-14 mm. long in the
male and 0:16 mm, in the female. This portion is surrounded
by the nerve collar. The caudal part of the esophagus is very
long and broad in diameter. It pursues a crooked course, being
acutely bent from side to side. It is 1-06 mm. long in the male
and 1-09 in the female. Its caudal end projects into the intestine
and is protected by a trilobed valve.
Text-figure 8. Text-figure 9.
OF
50p
Parabronema smithii Cobbold. Parabronema smithii Cobbold.
Ventral view of head. Lateral view of head,
Chyle intestine.—The intestine pursues a straight course and
is lightly pigmented The rectum is narrow and is constricted
from the rest of the intestine.
Exeretory pore.—This could only be made out in the female,
where it is 0°25 mm. from the head end.
Nerve collar.—Vhe thick nerve collar surrounds the cephalic
portion of the esophagus. It is placed 0°22 mm. from the head
end in the male and 0°23 mm. in the female.
Genital organs.—Male: The coils of the convoluted testes lie
more in the longitudinal axis of the body. The long cement
gland curves in the caudal end of the body to end in the
ventral aspect of the cloaca.
Female: The vulva lies a little cephalad of the end of the
cesophagus in the mid-ventral line. At a distance of 0-1 mm,
from the vulval opening, the caudally directed vagina produces a
distinct U-shaped kink, and then resumes its course caudally,
where it is directly joined by the two uteri. The uteri proceed
218 DR. M. KHALIL ON THE NEMATODE
caudally. They are packed with embryos and no signs of eggs.
The ovaries are confined mostly to the caudal half of the body.
Spicules.—There are two unequal spicules. The long spicule
is 0°56 mm. long. It lies to the left of the short spicule, which is
0:21 mm. long. The latter lies parallel with the lower end of
the long spicule, and curves ventrally towards the cloacal opening.
There is a small triangular accessory piece 0-03 mm. in length.
Text-figure 10.
Text-figure 11.
Parabronema smithii Cobbold.
Female tail.
Parabronema smithii Cobboid,
Anterior part of the body.
Caudal papille.—The male tail is provided with four pairs of
pre-anal papille lying on either side of the mid-ventral line.
Each is provided with a long core and a single termination.
Two pairs of post-anal papille could be made out, one close to the
cloaca and the other midway between the cloaca and the tip of
the tail.
Tail.—The male tail is spirally coiled ventrally. It.is shorter
than that of the female and its tip is rounded. It is 0:18 mm.
in length.
PARASITES OF ELEPHANTS. 219
The female tail is slightly bent dorsally. It narrows gradually
to its tip, which is slightly rounded. There is apparently no
papillz on the female tail. The length of the tail is 0-36 mm.
Text-figure 12.
Parabronema smithii Cobbold. Male taii-
Embryos.—The female is viviparous. The embryos are so
closely packed that it was impossible to arrive at an approximate
estimate of their length.
Habitat. Coats of the stomach of the Indian elephant (India).
Discussion.
Baylis described fully P. indiewm and P. africana. He was
not able to find material containing P. smithii, which has only
been alluded to once since it was described by Cobbold. There
has been some doubt as to the identity of P. indicum and
P. smithii, as Cobbold’s description was not full. For this
reason the latter species is described here in full. It is certainly
a different species from VP. indicum, although found in the
same host. It isa much smaller worm, and the measurements
of its various organs are likewise shorter than the corresponding
figures of P. indicum.
The shape of the auricular appendages is different from that
alluded to above. The African species is of very great length, and
its auricular appendages of the cuticular thickening of the head are
quite distinctive. All its other dimensions are correspondingly
longer than in the other species.
MIcROFILARIA.
Evans and Rennie record the presence of a microfilaria in blood
taken from an elephant during the daytime. It measured 1804
in length and 6 in breadth. It is possible that it was really
a microfilaria, but I suggest that it is more probable that the
220 DR. M. KHALIL ON THE NEMATODE
viviparous species of Parabronema living in the stomach-wall void
their embryos into the lymphatics or blood and not into the
intestinal canal.
Super-family SrroncyLorpea Wienland, 1858.
Family StRONGYLID# Cobbold, 1864.
Sub-family StroneyvLin# Railliet, 1893.
Genus DecrusiA Lane, 1914,
Fairly stout worms with a cup-shaped mouth capsule. There
is a marked duct of the dorsal cesophageal gland running along
the mid-dorsal line of the mouth capsule.
Male: The dorsal ray is undivided except at its extreme tip.
The spicules are equal and similar, fine-pointed, and without
accessory piece.
Female: The tail is blunt, the vulva is in the caudal third, the
uteri are divergent, the caudad-running uterus turning imme-
diately cephalad.
Type-species, Decrusia additiciia,
Decrusia ADDITICTIA Raill., Henry & Bauche, 1914.
Strongylus additictus Raill., Henry & Bauche, 1914.
Decrusia decrusi Lane, 1914.
Decrusia additicia Raill., Henry & Bauche, 1915.
Decrusia additicia Lane, 1915.
Strongylus additicius Ihle, 1919.
This species has the characteristics of the genus. The mouth
capsule is tilted dorsally. The external leaf-crown consists of
140-150 rays. Male is 14 mm. long and 1:0 mm. maximum
diameter. The spicules are 2'4mm. long. Female is 15mm.
long and 1-4mm. maximum diameter. The tail is 0:3 mm, long,
blunt at the tip. The vulva is 73mm. from the tip of the tail.
The ova are 75 long by 40 broad.
Thle thinks that this genus is an unjustifiable division of the
genus Strongylus. Although there are points of similarity
between the two genera, the dorsal tilt of the mouth-opening,
the long dorsal ray of the bursa, and the absence of an accessory
piece make this genus justifiable. In the genus Strongylus itself,
it was suggested by Looss that S. vulgaris, which is more allied
to S. equinus the type-species than Decrusia additictia, might
eventually prove to belong to another genus.
Habitat. Warge intestines of the Indian elephant (India).
Genus Mursuipra Lane, 1914.
Lane’s diagnosis of the genus runs as follows :—“ Fairly slender
worms tapering towards the head end. The male is widest just
PARASITES OF ELEPHANTS, 22h
cephalad of the bursa, the female tapers caudad to a fine point.
The head is discoid in shape. “The oral aperture is hounded by
two slightly marked lateral lips, each carrying a sessile lateral
and two prominent submedian papilla. The oral capsule is
roughly cylindrical, and circular in cross-section, while the wall
varies in shape in different parts. Since this is thicker caudad
and thinner cephalad on its dorsal and ventral than on its lateral
aspects, its cavity is, atits cephalad end, wider dorso-ventirad than
latervo-laterad. The cuticle bounding the mouth cavity is applied
to the mouth capsule only over its caudad portion. Along a cir-
cumferential line, which is further cephalad on the dorsal and
ventral than on the lateral walls, the cuticle turns axo-cephalad
and splits into rays having a similar direction ; and since further
these rays are shorter dorsally and ventrally than laterally, the
result is that their points which form the boundary of the oral
aperture, do not describe a circle but produce a dorso-ventral slit.
The cesophagus is fairly short, widening caudad of the nerve-
collar. Its caudal end is guarded by three intestinal valves. The
boundaries of the cells of the chyle intestine are more marked
than usual. The cephalic and the cervical glands are large and
well developed. The lateral cervical papille are long, slender,
and project somewhat cephalad.
“The male expands ventrad just cephalad of the bursa. The
number of bursal rays is as usual. The externo-dorsal rays lie
close to the lateral rays. The dorsal ray is deeply cleft in the
mid-line, and each half is composed of three subdivisions. There
appear to be no pre-bursal papillae. The spicules are equal and
similar, and the head of each is bent like the head of a golf-
driver, while the shaft may be straight or may be bent near the
oint. There is a colourless thickening of the cuticle in the
position of the accessory piece, which is §-shaped in lateral optical
section. The other male organs are as usual.
“The female has a long and pointed tail, the vulva lying shortly
cephalad of theanus. The vagina runs cephalad and divides into
two uteri, which have the same direction, are provided with
ovejectors, run parallel with one another, and end abruptly in the
two cephalad-running ovaries.”
I may add to this that an accessory piece is present, and
although the male is broad just above the bursa, this is not
always the greatest diameter of the body.
Type-species, Murshidia murshida.
MursHIDIA MURSHIDA Lane, 1914.
This nematode has the general characteristics of the genus.
The external leaf-crown consists of sixty leaflets. The vulva has
a cuticular prominence both cephalad and caudad of it. (For
measurements vide Table I., p. 228.) The lateral and dorsal rays
of the bursa have a rugged outline and a bulbous origin.
Habitat. Ceecum of Indian elephant (India).
Proc. Zoou. Soc.—1922, No. XVI. 16
Die, DR. M. KHALIL ON THE NEMATODE
MURSHIDIA FALCIFERA Cobbold, 1882.
Strongylus falcifer Cobbold, 1882.
Nematode No. 3. Evans & Rennie, 1910.
Strongylus falcifer Mitter, 1912.
Cylicostomum falciferum Raillet, Henry & Bauche, 1914.
Murshidia faleifera Lane, 1914.
The oral aperture is bounded by eighty rays. The bursal rays
have not the rugged outline seen in d/, murshida. (For measure-
ments see Table I.)
Mitter in 1912 gave a short description of this nematode with
figures. He represented the bursa with one ray missing, which
is either the externo-dorsal or one of the terminal branches
of the dorsal ray. In either case this may be attributed to
inaccuracy.
Habitat. Intestine of Indian elephant (India).
MURSHIDIA LINSTOWI, sp. n. (Text-figs. 13 & 14.)
Sclerostomum rectum von Linstow, 1907, from the African
elephant (not Strongylus rectus von Linstow, 1906, from Dolichotis
patagonica).
Cylicostomum rectum Gedoelst, 1916.
Murshidia recta Thle, 1919.
This species was first described by von Linstow in 1907 as
Sclerostomum rectum. The genus Sclerostomum Rudolphi, 1809,
Text-figure 13,
Text-figure 14.
Murshidia linstowit, sp. n.
Lateral view of male bursa,
Murshidia linstowi, sp. 0.
Cephalic end of body.
By kind permission of Prof. Leiper.
is a synonym of Strongylus Goeze, 1782, both having as their
type-species S. equinus Mueller, 1780, Thus Strongylus has
PARASITES OF ELEPHANTS. J23
precedence. Von Linstow described in 1906 Strongylus rectus
from Dolichotis patagonica.
According to Article 36 of the International Law of Zoological
Nomenciature, the specific name rectum, 1907, must be rejected
and can never be used again.
This is in complete agreement with the views expressed by
Lane. He rightly was unable to arrive at the conclusion that
the nematode described by von Linstow belongs to the genus
Murshidia, because the original description and figures are very
unsatisfactory. Only the figure of the male bursa.shows any
marked resemblance to Murshidia.
I have been fortunate in having access to a series of camera-
lucida drawings of von Linstow’s original material in Berlin
Natural History Museum, made by Prof. Leiper and very
generously placed at my disposal. Unfortunately, the exact
magnification was not noted, and therefore von Linstow’s measure-
ments are relied upon, as they correspond in proportion to the
camera-lucida drawings.
The mouth capsule is wide, surrounded by a chitinous ring of
the shape characteristic of the genus. The cesophagus is short
and thick. The excretory pore is placed a considerable distance
eaudad of the cesophagus. The vulva is surrounded by raised
cuticle, specially cephalad. The male bursa is short and the
lateral rays are not rugged in outline. The spicules are straight
and sharply bent near their terminations, which are bulbous in
shape. (For details of measurements see Table I.)
Habitat. From the African elephant (Cameroon).
Mursuipia HADIA Khalil, 1922. (Text-figs. 15-21.)
Material —The material consisted of one male and four females,
selected from Prof. Leiper’s collection of Elephant nematodes.
Shape of body.—The body of the male is straight, the female
tail is slightly bent ventrally. The male is 18-5 mm. long and
the female is 24mm. long. The maximum diameter of both sexes
is about the middle, being *67 mm. in the male and °82 mm. in
the female. The antero-posterior diameter of the male body just
cephalad of the bursa is 55mm. This is a little smaller than the
diameter at the middle of the body.
Skin.—The cuticle is finely striated at intervals of 009 mm.
Mouth collar.—This is well developed and rounded in out-
line. It is°05mm. long and ‘2mm. in diameter. It is distinctly
separated from the rest of the body by a deep groove.
External leaf-crown.—Arising from the inner surface of the
mouth collar is the external leaf-crown, consisting of forty rays.
These rays are longer laterally than ventrally or dorsally. Each
ray is narrow and gradually tapers to a point.
Head papille.—There are the usual four submedian head
papille. They project freely above the head. Each is sur-
mounted by a small knob which, as already mentioned, represents
16*
DR. M. KHALIL ON THE NEMATODE
bo
i)
=
Text-figure 15,
a aS
sete ==
——
SO
Murshidia hadia Khalil. Head.
Text-figure 16.
\\ /]
Ta.
3
v5,
Murshidia hadia Khalil. Cephalic end of body.
—
_——
PARASITES OF ELEPHANTS. 225
the outlet of the cephalic glands. The two lateral head papille
do not project above the surface, and are marked by a slight
depression. They are conical in shape with a wide base.
Cervical papille.—The cervical papille are very thin, long, and
point cephalad. They are placed 1-4 mm. in the female and
1:14 mm. in the male from the head end.
Mouth capsule.—The mouth cavity is very large and almost
globular in shape. It is ‘12mm. long and -15 mm. maximum
diameter. The oral opening is ‘07 in diameter. The mouth
capsule is a massive chitinous structure ‘(07 mm. in length and
‘025mm. in thickness. It is slightly curved inwards. A fine
chitinous process of the mouth capsule bounds the oral cavity in
the region of the mouth collar. The broad floor of the oral
cavity is free from any teeth.
Text-figure 17.
Text-figure 18.
Gan
Murshidia hadia Khalil.
Ovum.
Murshidia hadia Khalil.
Tail end of female.
(Hsophagus.-—There is a cone-shaped cesophageal funnel. It is
‘06 mm. in length and -1 mm. at its junction with the oral cavity.
The cesophagus is broad for its length. It is °85mm. long in the
male and ‘9mm. in the female. Its maximum diameter is
-26 mm. in the male and ‘34 mm. in the female. The caudal end
of the esophagus projects into the beginning of the chyle intestine,
forming three small lobules.
226 DR. M. KHALIL ON THE NEMATODE
Chyle intestine.—The intestine takes a practically straight
course in the axis of the body. The wall of the intestine is
annulated on the external surface by shallow depressions between
the cells. Except for a short distance at the beginning and end
the whole length of the gut-wall is deeply pigmented. 'The
rectum is an elongated and narrow canal indefinitely demarcated
from the intestine.
Exeretory pore.—The excretory pore is placed just caudad of
the nerve-ring in the mid-ventral line. It is ‘068 mm. from the
head end.
Nerve collar.—This is very thin in comparison with the size of
the worm. It is placed -6 mm. from the cephalic end.
Genital organs.—Male: The testis 1s irregularly coiled, reach-
ing within 5 mm. of the head. There is no distinct seminal
vesicle. ‘he cement gland is of enormous length.
Female: The wide opening of the vulva is surrounded by a
raised cuticular margin. The strongly muscular vagina runs
straight towards the head end. The two uteri are convergent,
ending in strong ovejectors lying side by side and joining the
cephalic end of the vagina. The ovaries are irregularly coiled,
reaching within 5 mm. of the head end.
Bursa.—The bursa is very short, and indistinctly divided into
three lobes of approximately the same length. It measures
‘22 min. in length and ‘7 mm. in breadth.
Text-figure 19.
Text-figure 20.
woe,
Murshidia hadia Khalil. Murshidia hadia Khalil.
Lateral view of bursa. Dorsal ray of bursa.
The ventral ray is bifid throughout its length, and is placed a
considerable distance from the ventral edge of the bursa. The
lateral rays arise by a common origin. At the site of their
bifurcation each ray has a bulb-like swelling. The externo-lateral
ray lies separate from the other two and close and parallel to the
ventral ray. It ends in a papilla on the external surface a short
distance from the edge of the bursa. The medio-lateral and
dorso-lateral vays lie close together. The externo-dorsal ray has
PARASITES OF ELEPHANTS. 227
a rugged outline. It arises in common with the dorsal ray, and
lies close to and parallel with the postero-lateral ray, ending a
short distance from the edge of the bursa on the external surface.
The dorsal ray arises by a very broad origin and bifurcates into
two branches, each ending in three rays. The outer two rays
arise by a common trunk and lie close together. The medial ray
is stouter and longer, and ends some distance from the corre-
sponding ray of the opposite side. The dorsal ray is ‘25 mm.
long.
Genital cone.—The genital cone ends in a sharp point. The
cloacal canal opens a little distance posterior to the apex.
Spicules.— The two similar spicules are very stout and practi-
cally straight. The club-shaped tip is bent sharply forwards.
They are 1-1 mm. in length. There is a small, curved accessory
piece ‘1mm. in length. The cephalic portion of the spicular
sheath is reticulated.
Text-figure 21.
Wurshidia hadia Khalil. Spicules and accessory piece.
Termination of the female.—The female tail gradually tapers to
a sharp point. It is 2°25mm. long. The vulva is placed ‘6 mm.
cephalad of the anus,
Ova.—The ova in the vagina are small: 40 long and 23 pu
broad. They are then in the morula stage of development.
Habitat. Intestine of the African elephant (Uganda).
228 DR. M. KHALIL ON THE NEMATODE
Discussion.
The main specific differences between the members of the genus
Murshidia ave to be found in the shape of the rays of the male
bursa, the length of the spicules, the length of the female tail, and
the distance between the vagina and the anus. These and other
differences are for convenience tabulated below.
Upon these data the species Murshidia linstowi was differen-
tiated.
Railliet, Henry and Bauche have recorded the presence of
Murshidia falcifera in the African elephant. This was probably
either Jf. hadia or linstowi, which bear a superficial resemblance
to MW. falcifera.
In the following table all measurements given are in millimetres.
The measurements of J. murshida and MW. falcifera are based
partly on Lane’s record and partly on material of both species
sent by him to Prof. Leiper. ‘These specimens are labelled by
Lane as type material of the respective species. The measure-
ments of I. linstowi are based on the original record of this
species by von Linstow. Where the sex is not stated, the
measurements are put in the centre of the column.
Taste [.
M.murshida. | M. faleifera. | M.linstowi,sp.n.| M. hadia.
3. India. aul a. India. Og. Africa. Q. | 3. Africa. Q.
min. mm. | mm. mm. | mm. mm. | mm, mm.
Total length ............... | 18-20 22-281/22-95 28-30] 25 29 185 24
Maximum diameter...... 6 “9 “74. TO |) Oe Te | By “82
CBOFOINEVAUE) sosdansaasaacen | 2 uss 5) || 497 8 | 85 ‘9
Nervelcollaneeeereeay see ale 31 | “48 BOT ecny ita aes | 6
Diameter of head ......... 17 BOW Say iil ee ie eceeaes | 2
Rays of leaf-crown ...... 60 80 Pore Hae abe | 40
Diam. of mouth capsule. ‘08 | “il |e eed toy “15
Length of spicules ...... | 1:28 155 | “96 relia
VEIN OH TEVAEMIE) dobadacdare 1G | 22 31 | 2°25
Vulva from tail end .... 23 | 29 | 4°3 | 2°85
ORES» Ghendaend heeael Nat 05 X03 052X024. | 04 023
Cuticular striation ...... 007 ‘007 ‘01 | ‘009
Cervical papille ......... "85 1:2 see 14 1-4
Hxcretory pore ............ 8 LOSE REN Rite 68
Genus Cuonrancium Raill., Henry & Bauche, 1914.
Asifia Lane, 1914.
Fairly stout, straight worms; the head is obliquely truncated
dorsad. The oral opening is directed towards the dorsal surtace.
PARASITES OF ELEPHANTS. 529
The oral aperture is surrounded by a circle of converging rays.
The mouth capsule is large and like a funnel in shape. Rounded
cuticular prominences protrude into the oral cavity.
Male: The most striking feature of the bursa is an accessory
ray projecting from the dorsal aspect of the main stem of the
lateral ray. The two spicules are similar and equal. There is
an accessory piece. Mailliet wrongly recorded that the dorsal
ray divides into four branches instead of three.
Female: The vulva is close to the anus, where the worm
suddenly narrows. The tail is bluntly conical.
Type-species, Choniangium epistomum.
CHONIANGIUM EPISTOMUM Piana & Stazzi, 1900.
Sclerostomum epistonum Piana & Stazzi, 1900.
Choniangium epistomum Raill., Henry & Bauche, 1914.
Asifia vasifa Lane, 1914.
The male measures 14 mm.in length, with a maximum diameter
near the head of 0°75 mm. ‘The spicules ave 20 mm. in length.
The female is 19 mm. in length, with a maximum diameter of
1-'Omm. The vagina runs cephalad, joining the two parallel uteri.
The ova measure 50 w by 25 p.
Habitat. Cecum of the Indian elephant (India).
Genus QurtontA Lane, 1914.
Hvoansia Railhet, Henry & Joyeux, 1913 (not Hvansia Scott,
1906, for a copepod).
@uilonia Lane, 1914.
Nematevansia Thle, 1919.
Quilonia Ihle, 1919.
Fairly slender worms; the head discoid, the mouth terminal
and surrounded by two sessile lateral and four prominent sub-
median papille. The rays of the external leaf-crown are few
and characteristically curved. The cuticular lining of the oral
cavity does not lie directly in contact with the chitinous mouth
capsule. The mouth-opening is narrower than the diameter of
the oral cavity. The esophagus is nearly cylindrical in shape.
Two or more teeth project into the oral cavity.
Male: The bursa is more or less distinctly divided into three
lobes. The dorsal lobe is longer than the lateral lobes. The
bursal rays are the usual number. The dorsal ray is bifurcate,
each branch having three subdivisions. The spicules are equal
and similar, and each has a sickle-shaped point. The accessory
piece 1s curved from side to side, the concavity being ventrad.
Seen from the dorsum it is wider cephalad than caudad.
Female: The vulva is in the caudal third of the body. The
uteri are opposed, divergent and furnished with marked ovejectors.
The caudad uterus turns cephalad immediately beyond its
ovejector. ‘The two uteri run cephalad side by side. Brown
a ‘
230 DR. M. KHALIL ON THE NEMATODE
cement on the vulva is fairly marked. The ova are colourless
and thin-shelled and have a granular yolk.
Type-species, Quilonia renniei.
QUILONIA RENNIEI Raill., Henry & Joy., 1913.
Nematode No. 2 from Indian elephant. Evans & Rennie, 1910.
Hvansia renniet Ruill., Henry & Joy., 1913.
Vuilonia quilong Lane, 1914.
Kvansia rennier Lane, 1915.
Nematevansia rennier hile, 1919.
Vulonia renniet Ihle, 1919.
The main features of this species, as described by Evans and
Rennie and also by Lane, are: («) the leaf-crown projects freely
above the head, (6) the leaflets are thim and long, (c) the dorsal
vay bifurcates low down in the bursa, and (d) its tri-radiate final
branches are very thin and arise at different levels. (For
measurements see Table IT.)
Habitat. Cecum of the Indian elephant (India).
(UILONIA TRANVACRA Lane, 1914.
Quilonia tranvacra Lane, 1914.
Hvuansia tranvacra Raill., Henry & Bauche, 1915
Nematevansia tranvacra Ihle, 1919.
Quilonia tranvacra Lane, 1921.
The leaf-crown does not project above the head. It consists
of ten rays. The branches of the dorsal ray are stouter and
longer than in Q. rennieit. (For measurements see Table II.,
p. 249.)
Habitat. Cecum ? of the Indian elephant (India).
QUILONIA APIENSIS Gedoelst, 1916. (Text-figs. 22-28.)
Hvansia apiensis Gedoelst, 1916.
QYuilonia apiensis Lane, 1921.
This parasite was recorded by Gedoelst unaccompanied by figures.
In the genus Quilonia it is extremely difficult to differentiate
species on description only. Lane, in describing YY. africana,
expressed his regret that Q. apiensis was recorded without figures.
The same difficulty was experienced in examining the material
at my disposal without a full knowledge of @. apiensis. Later,
Prof. Gedoelst very generously sent co-type material of his species
to be added to Prof. Leiper’s collection. This made possible a
complete description of the species with camera-lucida drawings.
Material—The material consisted of 25 specimens labelled
Quilonia apiensis (co-types) from Prof. Gedoelst. Of these, eight
specimens are kept in the Helminthological collection of the
London School of Tropical Medicine. In addition, three speci-
mens were identified by me in Prof. Leiper’s s material,
Shape of body.—The body is straight, slender, and tapering
towards either end. The male is 16 mm. long and the female is
PARASITES OF ELEPHANTS. sit
19-21 mm. long. The maximum diameter of the body is about
the middle. It is -62 mm. in the male and -78 in the female.
The average length of this species given here is greater than
that origine ally recorded. Gedoelst stated that the length of the
male 1s 12: 8mm.and the female 17°83mm. That shes: measure-
nents were based on a single individual male and one female is
evident from his remarks: “ LD’ unique exernplaive que nous avons
étudié mesurait, etc.” The measurements recorded here are the
average of all the specimens available.
Shin. —The cuticle is striated at regular intervals of ‘02 mm.
Mouth collar.—Vhe mouth collar is distinctly separated from
the rest of the body by a groove. There is a curious cuticular
prolongation in the form bi a spine at the site of this groove
Text-figure 22.
Quilonia apiensis Gedoelst. Lateral view of head.
The mouth collar is irregularly rounded in outline, being -04 mm.
in length in the male and -08 in the female. It is -16 in
diameter.
Kxternal leaf-crown.—This consists of twelve leaflets. These
are thin and curved at the top, ending in a fine point. They
project freely above the head.
Head papille.—The four submedian head papille are thick.
Hach is surmounted by a knob-like structure. The two lateral
papille are thin and do not project above the surface. Their
termination 1s marked by a depression.
Cervical papille.—The two cervical papille are thin and point
slightly cephalad. | They are placed on the lateral lines ‘98 mm.
from the head end in the male and 1:04 mm. in the female.
Mouth capsule.—The chitinous mouth capsule is a short ring
"04 mm. in length and -16 mm. in diameter. It is not in
932 DR. M. KHALIL ON THE NEMATODE
contact with the wall of the oral cavity. The oral cavity is
wide laterally and contracted from before backwards. It is
cone-shaped.
Internal leaf-crown.---This consists of twelve leaflets corre-
sponding to the external Jeaf-crown. Each is broad, with
a rounded blunt end and does not project above the mouth
collar.
(Hsophagus.—There is a small but distinct cesophageal funnel
having a chitinous wall. From the top of the cesophageal columns
two spinous processes project into the mouth capsule. These are
not so prominent as in YY. africana. The csophagus is ‘73 mm.
long in the male and ‘8 mm. in the female. [t is °25 mm. in
maximum diameter in the male and :28 mm. in the female.
Text-figure 23.
SOW
Quilonia apiensis Gedoelst. Dorsal view of head.
The cesophagus is slightly narrowed at the site of the nerve collar.
The caudal end of the cesophagus projects into the beginning of
the intestine, forming a tri-lobed valve.
Chyle intestine.—The intestine is straight and thick-walled.
It is deeply pigmented in its middle two-thirds. In most
specimens there is a characteristic constriction in the outline of
the intestine 2 mm. from its caudal end.
Rectum.—A marked groove indicates the beginning of the
rectum. Its walls are thinand chitinous. The lumen is spindle-
shaped, -42 mm. in length.
Excretory system.—The excretory vesicle is small and has a
delicate wall. The excretory pore is placed ‘6 mm. from the head
end.
Nervous systenu.—The nerve collar surrounds the narrowest
PARASITES OF ELEPHANTS. 933
part of the cesophagus and is ‘24 mm. from the head end. ‘Iwo
long bands stretch from the nerve collar caudally.
Geiital organs.—Male: The fine tubules of the testis are bent
irregularly, reaching to within 4mm. of the head end, There is
no distinct vesicula seminalis. The cement gland is of enormous
length, ending in the anterior part of the genital cone.
Female: The two convoluted ovaries reach within 5 mm, of
the head end. The two uteri lie side by side in the longitudinal
axis of the body. Each ends in a strong ovejector—that of the
anterior uterus is cephalad and that of the posterior uterus is
caudad of the vagina. The two ducts from the muscular part of
Text-figure 25.
Text-figure 24.
Quilonia apiensis Godoelst.
Ventral view of anterior portion of body.
Quilonia apiensis Gedoelst.
Female tail.
the ovejector join a very short vagina placed transversely to the
longitudinal axis of the body. The vulva is not surrounded by
raised lips. It is situated 6°2 mm. from the tip of the tail.
brownish cement surrounds the vulval opening.
Bursa.—The bursa is more or less divided into three lobes.
The posterior lobe is slightly longer than the lateral lobes. Seen
from the lateral aspect, it resembles somewhat the bursa of
Q. uganda in shape, although the dorsal ray is totally different.
The bifid ventral ray lies close to the lateral rays. ‘he fissure
234 DR. M. KHALIL ON THE NEMATODE
between the medio-lateral and the postero-lateral rays is deeper
than that between the medio-lateral and the ventro-lateral
rays. The externo-dorsal ray arises from the stem of the
dorsal ray. he dorsal ray is ‘7 mm. long. It bifurecates in
its lower third, Each branch after a short course divides
Text-figure 26.
Quilonia apiensis Gedoelst. Ovum.
into three rays at the same level. In many specimens, however,
the lateral two branches tend to lie close together, and the
fissure between them is not so deep as that which separates the
medial branch,
Text-figure 27.
Text-figure 28.
/00p
Quilonia apiensis Gedoelst.
Dorsal ray of bursa.
Quilonia apiensis Gedoelst.
Lateral view of bursa.
Genital cone.—The genital cone is blunt at the apex, which lies
in line with the ventral surface of the body. The cloacal
canal opens posterior to the apex and is surrounded by four
rounded lips.
PARASITES OF ELEPHANTS. 935
Spicules.—The two similar spicules are gracefully curved
ventrally. Their tips are fine and bent dorsally. ‘hey are
‘91 mm. in length. The accessory piece is*19 mm. in length.
It is concave ventrally, providing a canal for the spicules. It
is thicker at its cephalic than at its caudal end.
Termination of female.—The tail of the female is rather broad. —
The tip in most specimens is diseased and deeply pigmented,
but a few retain the tip, which is fine. The tail is 2°66 mim.
in length. The anal opening is slightly raised above the surface.
Ova.—The ova in the vagina ave thin-shelled, measuring 88 ju
by 48. They contain a morula.
Habitat. Intestine of African elephant (Belgian Congo and
Uganda).
(JUILONIA AFRICANA Lane, 1921.
This species is represented by more specimens than any
other in the Uganda material. It agrees in all characters
with Lane’s description, except in the length of the female
tail and the distance of the vulva from the anus. These were
given by Lane as 4 mm. the length of the tail, and 4 mm. the
distance of the vulva from the anus. On re-examining Lane’s
type-species, kept in the Natural History Museum, I could not
find a single specimen with these measurements. All corre-
spond to my previously taken measurements, being 3°6 mm.
length of tail, and 3:25 mm. distance of vulva from the anus.
Lieut.-Col. Lane ascribes this to the fact that the proofs of his
paper were not corrected by him. (For detailed measurements
based on my observation, see ‘lable IT.)
Habitat. Stomach of African elephant (South Africa and
Uganda). “e
QuiLontA UGANDA Khalil, 1922. (Text-figs. 29-34.)
Material.—The material consisted of fifteen specimens sorted
from the Uganda collection.
Shape of the body.—The body is short and straight. The
female is slightly longer and thicker than the male. The male
is 11°5-12 mm. long and the female is 16 mm. long. The
maximum diameter of the body is about the middle. It is
.OD mm. in the male and ‘64 mm. in the female.
Skin.—The cuticle is striated more closely near the head end
than in the middle of the body. Near the head the intervals
between each two striations is ‘01 mm., near the middle of the
body it is °02 mm.
Mouth collar.—The mouth collar is distinctly separated from the
body by a groove. It is rounded in outline, -04 mm. in length
and *19 mm. in diameter.
Anterior leaf-crown.__From the inner surface of the mouth
collar projects the anterier leaf-crown, It consists of twelve
236 DR. M. KHALIL ON THE NEMATODE
leaflets, fairly broad and bent characteristically at their tips.
They project a little distance above the head.
Head papille.——The four submedian head papille project a little
above the mouth collar. They point straight forwards. Their
bases are broad and each is surmounted by a long knob, the
function of which is pointed out elsewhere. The two lateral
head papille do not project, and their termination is marked by
a depression.
Cervical papille.—The two delicate cervical papille are placed
a little posterior to the level of the caudal end of the esophagus.
They are °78 mm. from the head in the female and -6]1 mm. in
the male.
Text-figure 29.
Quilonia uganda Khalil. Head.
Mouth capsule.—The chitinous mouth capsule is -15 mm. in
diameter. It is oval in optical cross-section and is not in contact
with the wall of the oral cavity. The oral cavity is shallow and
funnel-shaped. It is 06 mm. deep and *11 mm. diameter at its
floor. The mouth-opening is 07 mm. in diameter, From the floor
of the oral cavity project two large chitinous oval teeth, each
surmounted by a delicate sharp spine. These are placed on the
cephalic end of the two sub-ventral cesophageal columns.
Internal leaf-crown.——The short stumpy leaflets of the internal
leaf-crown correspond in number to the external crown. They
have a rounded end and do not project above the surface.
(Hsophagus.— There is a shallow cesophageal funnel surrounded
by a chitinous rim. The cesophagus is short and thick, being
-52 mm. in length in the male and 62 mm. in the female. Its
maximum diameter is °2 mm. and :21 mm. in the respective
sexes. The end of the cesophagus projects into the chyle intestine,
its opening being guarded by a trilobed valve.
PARASITES OF ELEPHANTS. 237
Chyle intestine.—Vhe course of the chyle intestine is straight in
The intestinal wall is thick and deeply
the axis of the body.
The rectum is a narrow, long canal
pigmented in its caudal half.
"28 mm. in length, demarcated from the intestine by a groove
round the circumference of the gut.
Text-figure 31.
Text-figure 30.
=
ao
—
ARTA RAS
See
H i
——
saci din on a on er LL rel ene
ala ile
100 7
Quilonia-uganda Khalil.
Cephalic end of body.
Quilonia uganda Khalil.
Female tail.
Text-figure 32.
Quilonia uganda Khalil. Ovum,
Nerve collar.—The thick nerve collar is placed °25 mm. from
It surrounds the narrowest part of the
the cephalic end.
cesophagus.
Proc, Zoou. Soc.-—1922, No. XVII. ‘lis
238 DR. M. KHALIL ON THE NEMATODE
Excretory pore.—The excretory pore is ‘47 mm. from the head
end.
Genital organs.—Male: The convoluted testis reaches within
3mm. of the head end and is irregularly coiled. The vesicula
seminalis is spindle-shaped, lying in the long axis of the body.
The cement gland is very long, measuring 3:5 mm.
Female: The irregularly convoluted ovaries reach within 3 mm.
of the head. The divergent two uteri lie side by side for most
ef their course. Hach ends in a strong muscular ovejector
which lies, in the case of the anterior uterus, cephalad and, in
the case of the posterior uterus, caudad of the vagina. The latter
is a short horizontal duct joining both uteri to the vulva. The
opening of the vulva is not surrounded by raised lips, and is
placed 2:17 mm. cephalad of the anus.
Text-figure 33.
Quilonia uganda Khalil. Lateral view of bursa.
Bursa.—Seen from the lateral aspect the bursa has a spindle
shape, lying obliquely to the longitudinal axis of the body. The
dorsal lobe is slightly longer than the lateral lobes and has a
rounded end. The ventral, lateral and externo-dorsal rays con-
form to the general design in the genus, but are all more slender.
The dorsal ray is long and thin, being °5 mm. in length. It
bifurcates in its lower third into two long and slender branches.
These in their turn branch nearer their ends. The most lateral
ray arises first, and the two medial rays arise by a common trunk
branching lower still. This arrangement is exactly the opposite
of that in Y. apiensis.
‘; The pre-bursal papille are placed °52 mm. from the tip of the
bursa.
Genital cone-—This cone is placed nearer the ventral than the
dorsal surface. Its tip is rounded. The cloacal opening is
PARASITES OF ELEPHANTS. 239
marked by a depression in its outline a little distance dorsal to
the apex.
Spicules.—The two similar spicules are ‘72 mm. long, the
shortest in all the members of the genus. The tips of the
spicules are sharp and bent ventrally. The elongated accessory
piece isl mm.in length. It is curved ventrally to lodge the
spicules,
Text-figure 34,
Quilonia uganda Khalil. Wentyral view of bursa and spicules.
Termination of female.—The female tail is 2°43 mim. in length.
It ends in a fine point. The anal opening is marked by a
round depression from the surface.
Ova.—The ova seen in the terminal part of the uterus are
thin-shelled. Their contents are in the morula stage. The
dimensions of the ova are 60 p in length by 30, in breadth.
Haoitat. Intestine of African elephant (Uganda).
QUILONIA BREVICAUDA, sp. n. (Text-figs. 35-39.) -
Material.—The material consisted of two specimens sorted
out from the Uganda collection of nematode parasites of the
elephant.
Shape of the body.—The female is much stouter and longer
than the male. The outstanding feature of this species, as its
name implies, is the short tail of the female ending bluntly,
as can be seen with the naked eye. The male bursa is very
much smaller than in the other species. The male is 13mm.
long. The female is 21 mm. long. Tbe maximum diameter
Was
240 DR. M. KHALIL ON SHE NEMATODE
of the body in either sex is about the middle. It is ‘45 mm. in
the male and ‘85mm. in the female.
Skin.—The cuticle is striated at intervals of ‘02 mm. through-
out the length of the body.
Mouth collar.—The mouth collar is rounded in outline. It
is ‘06 mm. long in the male and ‘08 mm. in the female.
Its diameter is -18 mm. in the male and -22 mm. in the
female.
Eeternal leaf-crown.—This consists of ten very slender leaflets,
bent near them termination. They do not project freely above
the head.
Text-figure 35.
Quilonia brevicauda, sp.n. Head.
Head papille.—Vhere are the usual four submedian head
papillae. These are stumpy and surmounted by a knob each.
The lateral head papille do not project above the surface and
are marked with a depression.
Cervical papille—The two cervical papille are very thin and
directed slightly cephalad. They are *95 mm. from the head end
in the male and 1:1 mm. in the female.
Mouth capsule.—The chitinous mouth capsule is very short.
It has a diameter of -13 mm. in the male and *14 mm. in the
female. It lies closer to the oral cavity than in the other
species. The oral cavity is funnel-shaped, narrowing tcwards the
mouth-opening. Into the mouth capsule project three formid-
able Jong teeth; each is placed on the top of one of the three
cesophageal columns. This peculiarity is apparently limited to
this species.
Internal leaf-crown.—This consists of biunt and slightly pro-
jecting processes placed deep in the oral capsule.
PARASITES OF ELEPHANTS. 241
Hsophagus.—There is a small esophageal funnel. The ceso-
phagus is elongated, being -52mm.in length in the male and
‘7 mm.in the female. It is slightly constricted in the region of
the nerve collar. From its posterior end project the three lobes
of the cesophageal valves.
Chyle intestiie.—The chyle intestine pursues a straight course.
It is pigmented in its middle. The rectum is sharply demar-
cated from the end of theintestine. It is narrow and thin-walled,
and measures ‘24 mm. in length.
Text-figure 36.
Quilonia brevicauda, sp.n. Cephalic end of body.
Execretory pore.—This ig situated in the mid-ventral line
-73 mm. from the head end in the male and °88 in the female.
Nerve collar—The nerve collar surrounds the cesophagus
-29 mm. from the head end in the male and °39 mm. in the
female.
Genital organs.—Male: The convoluted testis reaches within
3 mm. of the head end. There is no distinct seminal vesicle.
The cement gland is very long.
Female: The convoluted ovaries reach within 6 mm. of the head
end. The two divergent uteri lie side by side for the most of
their course. Each ends ina strong muscular ovejector. That
242 DR. M. KHALIL ON THE NEMATODE
of the posterior uterus makes a loop caudad of the vulval cpening
before joining the ovejector of the anterior uterus to form the
very short horizontal vagina. The vulva is covered with dark
brown cement. It is 5°l mm. from the tail end and 3:35 mm.
from the anus.
Text-figure 37.
Text-figure 38.
SO
Quilonia brevicauda, sp. n.
Ovum.
Quilonia brevicauda, sp, n.
Female tail.
Bursa.—Vhe male bursa is short and broad. It is 63mm. in
length and 52 mm.in breadth. The ventral, lateral and externo-
dorsal rays are similarly arranged as in other members of the
genus. The dorsal ray is shortand broad. It is°5 mm. in length.
It divides in its lower third into two thick branches. Each in
turn ends in three small and thick rays. The most lateral
arises separately and at higher level. The middle ray is the
shortest.
Genital cone—The blunt genital cone hardly protrudes into
the bursa. ‘The cloacal opening lies a little dorsal to its blunt
apex.
Spicules—The two similar spicules are ‘81mm. in length.
Their termination is very fine and directed ventrally. The
accessory piece is "19 mm. in length. Its lateral margins curve
forwards, forming the dorsal wall of the cloacal duct.
PARASITES OF ELEPHANTS, 243
Termination of the female.—This forms the most. charac-
teristic feature of this species. The female tail is broad and
short. It is 1:75 mm. in length. It is of practically uniform
diameter till near the tip, where it is cone-shaped. The tip is
blunt.
Text-figure 39,
1
Quilonia brevicauda, sp. n. Bursa and spicules.
Ova.—The ova seen in the terminal portion of the uterus
contain a morula, They measure 56 in length and 33, in
breadth.
Habitat. Intestine of African elephant (Uganda).
QUILONIA ETHIOPICA, sp.n. (Text-figs. 40-45.)
Material.—The material consisted of eight specimens sorted
from the Uganda collection.
Shape of body.—The body is straight, broader near the middle,
and gradually tapering towards either end. Male is 13°5 mm.
long and °55 mm. maximum diameter The female is 19 mm. in
length and -7 mm. maximum diameter.
Skin.—The cuticle is finely striated through the whole body
except for a short distance caudad of the head. The striations
are at intervals of :(029 mm.
Mouth collar—The mouth collar is rounded in outline. It is
944 DR. -M. KHALIL ON THE NEMATODE
‘06mm. in length in the male, ‘08mm. in the female. The
diameter is ‘14 mm. in both sexes.
Keternal leaf-crown.—The external leaf-crown consists of
twelve leaflets. These are the most prominent characteristic of
the species in their appearance and arrangement. They are broad,
leaf-like, and slightly bent at the tip. They do not protrude
above the head.
Head papille.—The four submedian head papille are displaced
laterally by the external leaf-crown. They project directly
forwards, and each is surmounted by a small papilla, the nature
of which is referred to elsewhere. The lateral papille do not
project, and their termination is marked by a depression.
Cervical papille.—The two laterally placed cervical papille are
directed slightly cephalad. They lie ‘7mm. from the head end
in the male and ‘86 mm. in the female.
mele 40,
50
Quilonia ethiopica, sp.n. Head.
Mouth capsule—The chitinous mouth capsule is extremely
shallow. In optical cross-section it lies oblique to the axis of
the body. Itis -018 mm. in depthand-12 mm. in diameter. The
chitinous mouth capsule does not come in contact with the wall
of the oral cavity. The latter is practically cylindrical in shape
and slightly contracted in the middle. There are apparently no
teeth projecting into the mouth cavity.
Internal leaf-crown.—The twelve leaflets of the internal leaf-
crown can be traced from the floor of the mouth cavity to about
its middle. Hach leaflet is broad at its base and tapers toa
point. It has a distinct cleft in the middle, and thus each is
really two leaflets side by side.
@sophagus.—There is a shallow cesophageal funnel surrounded
by a chitinous rim lying on the three cesophageal columns.
The cesophagus is shortand thick. It is ‘55 mm. in length in the
male and ‘07 mm. in the female. It is slightly narrowed at the
PARASITES OF ELEPHANTS. QA5
site of the nerve collar. Its maximum diameter is ‘21 mm. 1n
the male and ‘26mm. in the female. The posterior end of the
cesophagus projects into the intestine, and its opening is guarded
by a trilobed valve. -
Chyle intestine. —The straight chyle intestine is pigmented in
Text-figure 41,
420 Ms
—
Quilonia ethiopica, sp.n. Cephalic portion of body.
its middle third. The narrow rectum is -23 mm. in length and
sharply demarcated from the rest of the intestine.
Excretory pore.—The excretory pore lies in the mid-ventral line
°6 mm. from the head end.
‘Text-figure 42.
Quilonia ethiopica, sp. ii. Ovum.
Nerve collar.—The nerve collar is °25 mm. from the head end.
Genital organs. Male: The testis is convoluted irregularly,
reaching within 3°5 mm. of the head. There is a peculiarly
246 DR. M. KHALIL ON THE NEMATODE
shaped seminal vesicle, having the form of two spindles con-
nected with a narrow duct. The cement gland is long and
pierced by the long ejaculatory duct, which opens into the ventral
aspect of the cloaca. °
Female: The convoluted ovaries reach within 4 mm. of the
head end. The divergent uteri lie side by side for most of
their course. Each is provided with a well-developed muscular
ovejector. The posterior ovejector loops caudad of the vulvar
opening before joining the anterior ovejector to form the short
horizontally placed vagina. The vulva is covered with brown
cement and lies 4°97 mm. from the tail end.
Text-figure 43.
Quilonia ethiopica, sp.n. Male bursa and spicules.
Bursa. The male bursa conforms to the usual type. The
dorsal lobe is slightly longer than the lateral lobes. The bursa is
‘8mm. inlengthand*6mm.in breadth. The ventral, lateral and
externo-dorsal rays are arranged in a similar fashion, as seen in
the other species. The dorsal ray is ‘67mm. in length. It
bifureates in its lower third, The terminal branches are slender
and long. ‘The lateral branch separates first.
Genital cone.—The genital cone is fairly long and pointed.
The cloacal opening lies near its apex.
Pre-bursal papille.—These lie on the lateral lines ‘75 mm. from
the end of the bursa.
PARASIVES OF ELEPHANTS. 247
Spicules.—The two similar spicules are ‘83 mm. in length.
They end in a fine termination, bent forwards. ‘The accessory
Text-figure 44.
fl
100 p
Quilonia ethiopica, sp.n. Ventral view of bursa.
Text-figure 45.
tone
Quilonia ethiopica, sp.n. Female tail.
piece is *l1 mm. long. It has a bulbous cephalic extremity.
grooved ventrally to lodge the spicules.
Itis
248 THE NEMATODE PARASITES OF ELEPHANTS.
Termination of the female.—The female tail is 2°3 mm. long. It
tapers gradually. Its tip is rounded.
Ova.—The ova in the terminal part of the uterus are oval in
shape, thin-shelled, and contain a morula. ‘They measure 63 pu
long and 35 pw broad.
Habitat. Intestine of African elephant (Uganda).
Text-figure 46.
Quilonia africana Lane. Abnormality in the rays of the bursa.
In a male specimen of Q. africana which was otherwise normal,
the externo-dorsal ray was completely missing on one side, as
is shown in the accompanying camera-lucida drawing. This
abnormality must be very rare, and has apparently not been
recorded before.
Discussion.
As was mentioned before, the identification of any particular
species of the genus Qwéilonta from descriptions only is very
difficult. Diagrams representing the head and tail ends are
necessary. ©. brevicauda is easily distinguished on account of
its short stumpy tail. For the rest, careful comparison is neces-
sary for diagnosis. ‘he important features in this connection
are the shape of the head, the external leaf-crown, the dorsal
ray of the male bursa, the length of the spicules, the length of
female tail, and the distance between the vulva and anus.
The fact that the external leaf-crown may or may not project
above the head divides the genus into two groups. In Q. renniei,
i ce <q Weg
660.
L6-1
6-6
Tis
€8.
SG.
9.
98. yi
9G. 1G.
Lg. ce.
GI-
10.
‘sICU ZT
Ut.
L. ee.
61 ¢.€1
“One
*pardory79 *2
neg hq 119g
G0:
L.¢
GL.L
61.
18.
“PpNYIINANG *@
og Sq gg
G0:
6-G
€V-6
ile
GL.
GG.
LY.
8L. T9.
Ge
GQ. GG:
CT.
90.
"SAB ZT
6I1-
79- SS
9T GIT
coe Se
“ppunbr
ile
ST.
1 gp Aq 71 €8
80. 90.
*sAUL ZT
LG. 6.
BL. Gd.
16-61 9
‘sasuaidy *%
nop Sq OL
10.
8.9
6.3
SG.
SI.
da.fe Sq I G.18-SL
Se
TEO.
0.9
0-6
PIG] ‘ouvry vruopngy snuas ey4 Jo saroads jo syueutetnsve|{— JT] WTAV],
as aqusdvo yyuow jo “UBT |
verses" 19F 9 UBIP WU UIX BIL |
WOIFRLTGS IBMT NY)
“puta [IB} TWOAF BATA |
|
rep apBUIAy |
aoatd A.10S88900
coer nen gemnordg |
soe seeeerere esses IRITOD AATONT
reer 919 ALOJOLON GL
ess carded [BOLAAD |
ef “UURIP “XVIAT
snSvydosa Jo yy.ouey
|
|
K41AvO [RIO JO YSU |
“UMO.1)- PROT [BULTAFXOT |
AB] ]O9 qynout jo “WleT( |
ses HASTA [RIOT
250 DR. M. KHALIL ON THE NEMATODE
apiensis, uganda, and ethiopica the external leaf-crown projects.
In Q. ethiopica particularly the shape of its leaflets is so distinctive
as to form a specific difference. In Q. tranvacra, africana, and
brevicauda the external leaf-crown is more or less hidden by the
mouth collar.
Table II. gives in detail the various measurements useful in
differentiating any species of Quilonia. The data for Qwilonia
renniet and tranvacra ave taken from Lane. ‘The data for the
rest of the species are based on my personal examinations. The
numbers are given in millimetres.
Genus PrertIpoPHARYNX Lane, 1921.
Body is slender, with discoidal head. The rays of the external
leaf-crown vary in length. The oral cavity has the shape of an
inverted funnel, its cuticular lining coming into contact with the
circular chitinous ring which forms the oral capsule. The
cesophagus is short and wide. The internal aspect of the three
cesophageal columus may or may not be sculptured in a plumose
fashion. When this is present, the free internal surface of the
cesophagus is moulded into a number of freely projecting ridges
which run obliquely outwards and posteriorly, giving it a
feathered appearance.
In the male bursa the lateral ray has a prominence or acces-
sory ray on its posterior border, while of the three branches of
each dorsal ray the two outer are fused partly or to nearly their
tips. The spicules are equal and similar, and an accessory piece
is present.
‘The vulva lies closely anterior to the anus, the vagina running
anteriorly and dividing into two parallel anteriorly directed uteri,
each provided with an ovejector.
Type-species, Pteridopharynx africana Lane, 1921,
PLrERIDOPHARYNX AFRICANA Lane, 1921.
Very little needs to be added to the description given by Lane.
The leaflets of the external leaf-crown are sixteen in number.
There are two teeth projecting into the floor of the mouth capsule,
apparently regarded by Lane as the beginning of an esophageal
funnel. The dorsal ray of the bursa before its final division
has an irregular outline. In some specimens this has the shape
of one or two processes, but this feature is not constant. Lane
did not see ova in any of his female specimens. I am able to
record their measurements as 62 « long by 32 « broad.
Habitat. Stomach of the African elephant (South Africa and
Uganda).
PYrERIDOPHARYNX ANISA Khalil, 1922. (Text-figs. 47-51.)
Source of material.—The material consists of more than thirty
specimens. These I sorted out from the large stock of parasites
collected by Prof, Leiper in Uganda.
PARASITES OF ELEPHANIS. Zl
Shape of body.—In the preserved condition the body of both
males and females is practically straight. The male is 14-15 mm.
long. ‘The female is slightly longer, being on an average
16:5-17mm. The greatest diameter of the body is about the
middle, being ‘46mm. in the male and °65 mm. in the female.
The body tapers slightly towards the head end and the caudal
end. In the female the tail becomes suddenly constricted from the
body, appearing to the naked eye as a spinous process projecting
from the posterior end of the body.
Skin.—The skin is finely striated throughout the whole length
of the body, including the tail of the female. The striations are
recularly placed at distances of 10 between. There is no
cuticular expansion.
Mouth collar.—The mouth collar has a rounded outline. It is
separated from the rest of the body by a distinct groove. The
diameter of this collar is shorter than that of the body in its
immediate neighbourhood. It is *125 mm., while its length
is ‘02 mm.
Text-figure 47.
Pleridopharyne anisa Khalil. Head.
External leaf-crowi.——This consists of twenty leaflets, broad at
their base and sharp at their termination. ‘The outline of the
tips of these leaflets, seen from the ventral aspect, forms a crescent,
being longer in the centre than at either side.
Head papille.—The four submedian head papille are long, and
each surmounted with a small knob. The latter is apparently
continuous with a core inside the papilla, and presumably
represents the duct of a head gland. The two lateral papille
do not project above the surface; their terminations are marked
by a depression in the cuticle.
252, DR. M, KHALIL ON THE NEMATODE
Cervical papille.—These are long and thin, They are placed
‘92 mm. from the cephalic end of the body.
Mouth capsule.—The mouth capsule is pear-shaped. It is longer
than it is broad. It is ‘093mm. long and -065 mm. in maximum
ciameter. The chitinous plate forms the wall of the posterior
half of the mouth capsule, and a funael-like extension of it forms
the wall of the anterior portion. The chitinous plate is ‘06 mm.
long; its upper endis sharply cut in an oblique direction. From
the posterior end of this chitinous ring project a series of sharp
teeth-like processes overhanging the opening of the cesophagus,
Csophagus.—There is a broad, shallow cesophageal funnel
bordered by a thick cuticular edge. The length of the esophagus
is ‘45mm. in the male and ‘57 mm. in the female. Its maximum
Text-figure 48.
Pteridopharynx anisa Khalil. Ventral view of cephalic end of body,
diameter is:l14mm in either sex. There is hardly any constric-
tion at the site of the nerve collar.
Chyle intestine—The beginning of the chyle intestine overlaps
the wsophagus. It pursues a straight course to the rectum. The
cellular lining of its walls is pigmented in its middle. The lumen
narrows befure joining the rectum.
Rectum.—This has a wide funnel-shaped cavity in the female.
It has a thin cuticular lining in contradistinction to the thick
cellular wall of the intestine. The caudal end of the rectum
narrows to a short narrow anal canal, the opening of which is
surrounded by a slightly raised lip.
Ewcretory pore.—Lhe excretory vesicle is a small fusiform sac,
with a short duct opening ‘87mm. from the cephalic end on the
ventral surface.
PARASITES OF ELEPIIANTS. 253
Head glands.—There are four elongated, deeply pigmented
bodies surrounding the beginning of the cesophagus. They con-
tain orange-coloured granules, suggesting their being glandular.
Their anterior end is elongated in the form of a duct lying to the
outer side of the chitinous ring of the mouth capsule. Their
exact termination could not be made out, but probably they
traverse the submedian head papille and open at their tip. The
presence of ducts opening on the tip of the head papille was first
recorded by Cooper Curtice and confirmed by Giles in sophago-
stomum columbianun.
Both these observers, however, interpreted them to be the ducts
of the lateral canals. Looss’ comment was: “I do not entertain
the slightest doubt that they have seen the outlet of the head-
glands.”
Text-figure 50.
Text-figure 49.
Pteridopharynx anisa Khalil.
Ovum.
Pteridopharyne anisa Khalil.
Female tail.
Nerve collar.—The thick nerve collar is -25 mm. from the head
end. ;
Genital organs.—Vhe male testes are irregularly convoluted,
reaching to within 3mm. of the head. The cement gland is long,
terminating at the base of the genital cone, where the ejaculatory
duct opens into the cloaca.
The female vagina is a long, straight canal lying in the axis of
the body. At its cephalic end the two strongly muscular ovejectors
open into it, connecting it with the corresponding uterus. The
conyolutions of the ovaries reach to within 4:5 mm. of the cephalic
end of the body.
Proc. Zoou. Soc.—1922, No. XVIII. 18
254 DR. M. KHALIL ON THE NEMATODE
Bursa.—The bursa is long, and is indistinctly separated into
three lobes, especially when seen in a side view. ‘The elongation
is due mainly to the dorsal lobe. The bifid ventral ray is short
and placed close to the cephalic end of the bursa, where it may be
overlooked on casual examination.
The lateral group of rays arises by a conmon trunk, and, in
addition to the ordinary three branches, gives a fourth stumpy-
looking ray. This is directed caudally. The externo-dorsal ray
is stout and arises close to the lateral rays. It does not reach the
edge of the bursa, The dorsal ray is ‘7 mm. long. It gives a
Text-figure 51,
M0
Pteridopharyne anisa Khalil. Lateral view of male bursa and spicules.
branch on either side after °18mm. course. The ending of each
of these branches is bifid. This bifureation can be traced for a
longer distance than in P. africana. The main stem of the dorsal
vay after another course of -2mm. divides into two branches
which end at the posterior limit of the bursa. The bursa is
*82 mm. long and -47 mm. broad.
Genital cone.—This has a truncated apex. The cloacal opening
is surrounded by a protruding lip, and is directed ventrally and
caudally.
Sprcules.—The two similar spicules are °82 mm. in length,
PARASITES OF ELEPHANTS, 255
Their cephalic ends are thick and surrounded by an everted lip.
In their course they apparently entwine each other. Their tips
are sharp and curved backwards. The accessory piece is +13 mm.
long and curved dorso-ventrally.
Ternination of the female.—At the level of the vulva the
ventral surface curves towards the dorsum, forming a conical-
shaped tail. The opening of the vulva is directed caudally. The
anus opens "12mm. posterior to the vulva, and is also directed
caudally. The tail is °52 mm. in length; its end is rounded.
There are two caudal papille marked on the surface by a depres-
sion *33 mm. from the extremity of the tail.
Ova.—The ova in the terminal portion of the uterus measure
43 u by 30. ‘They are then in the morula stage.
Habitat. Intestine of African elephant (Uganda).
Discussion.
This species differs from P. africana in the shape of the mouth
capsule, which is more elongated ; the absence of the plumose
appearance of the esophagus; the greater distance between the
vulva and anus; and the general appearance of the tail in the
female. Inthe male the spicules are shorter, and the bifid end
of the branch of the dorsal ray is also much longer. The total
length of the body and of the various structures are also different.
As the plumose corrugation of the asophagus for which Lane
suggested the generic name Pteridopharyna is absent, this can
only be considered a specific character. The same plumose
appearance is present in Memphisia aziza, which undoubtedly
belongs to a different genus on account of the configuration of
the head and the presence of an additional ray in the bursa, both
of which are important morphological characteristics.
Genus Mempuista Khalil, 1922.
Body is slender. The rays of the external leaf-crown do not
project freely above the head. The oral cavity is deep, and
is wider from side to side than from before backwards. There
ave chitinous teeth in the floor of the mouth capsule. A shallow
cesophageal funnel is present. ‘The cesophagus is short and thick.
Plumose appearance of the cephalic end of the lumen of the
cesophagus may or may not be present.
Just caudad to the mouth collar the cuticle is raised to form
a collar surrounding the circumference of the body. The degree
of projection of this cuticular collar varies in the two species.
Male. The lateral ray has an accessory ray on its posterior
border. The externo-dorsal ray has a branch directed caudally.
Of the three branches of the dorsal ray the outer two are fused
to nearly their tips. Their dual nature is indicated by ending in
two papille. The two spicules are equal and similar. An
accessory plece is present. Ls
256 DR. M. KHALIL ON THE NEMATODE
Female: The vulva lies closely anterior to the anus. The
distance between the two openings varies in the different species.
The vagina runs anteriorly in the axis of the body. The two
parallel uteri are provided with strong ovejectors.
Type-species, Wemphisia memphisia Khalil, 1922.
MempuistA MEMPHISIA Khalil, 1922. (Text-figs. 52-58.)
Material.—The material consists of more than sixty specimens
sorted out of Prof. Leiper’s Uganda collection.
Shape of body.—Vhe body is slender, and in the preserved state
straight. The males vary between 13°5 and 14mm. in length.
The females are slightly longer, 14°5-16 mm. in length. The
maximum diameter is about the middle of the parasite, being
Text-figure 52.
Memphisia memphisia Khalil. Ventral view of head.
-55 mm in either sex. The body is slightly attenuated at the
cephalic end. Just posterior to the mouth the body is encircled
with a thick cuticular collar. The tail of the female is suddenly
constricted from the body, appearing to the naked eye asa large
posteriorly placed spine.
Skin.—The cuticle is closely striated at intervals of about 10 ph.
Towards the anterior end of the body the cuticle becomes inflated,
forming in optical section two alze which surround the whole cir-
eumference of the body. It is ‘2 mm. long, and extends ‘03 mm.
beyond the circumference of the body. It is placed at the level
of the anterior part of the cesophagus.
Anterior to this is a definite collar-like expansion of the
PARASITES OF ELEPHANTS. 257
cuticle, sharply demarcated from the rest of the body. It is
"05 mm. long, and projects by the same amount from the
body. The anterior cuticular expansion is not striated, but.
the posterior is. ;
Mouth collar.—This is very short and indistinctly separated
from the rest of the body. It is ‘02 mm. in length and ‘12 mm.
in diameter.
Anterior leaf-crown.—This consists of 36 incompletely sepa-
rated leaflets. They do not project freely above the mouth
collar.
Head papille.—There are four submedian head papille. They
are stumpy and surmounted with knobs continuous with,
apparently, a duct traversing the papilla. The two lateral head
papille do not project on the surface.
Text-figure 53.
Memphisia memphisia Khalil. Lateral view of head.
Cervical papille.—These are very long and delicate. They are
placed -92 mm. from the cephalic extremity.
_ Mouth capsule—Seen from the ventral surface the mouth
capsule is practically quadrangular, being slightly broader at the
outlet. A side view shows the same structure as funnel-shaped,
narrowing at the outlet. The length of the oral cavity is -08 mm. :
its maximum breadth is ‘(09 mm., and from before backwards it
measures ‘OOmm. The chitinous ring surrounding the oral
cavity is thickand nearly vertical. Its upper extremity is slightly
bent peripherally, as seen in ventral view. There are two teeth
projecting into the mouth cavity. :
- Bsophagus.—There is a shallow cesophageal funnel lined with
cuticle. The cesophagus is short and broad. It is -48 mm. in
958 DR. M. KHALIL ON THE NEMATODE
length and *18 mm. maximum diameter, and is very slightly
constricted at the site of the nerve collar. The caudal end of
the cesophagus projects into the beginning of the chyle intestine
in the form of three lobes. }
Ohyle intestine.—This pursues an almost straight course. Its
cellular wall is pigmented in its middle portion.
Rectum.—The intestine is sharply demarcated from the rectum.
The thick cellular wall of the intestine stops ebruptly. The wall
of the rectum is very thin and chitinous. Its cavity is pear-
shaped, ‘18mm. long and ‘1mm. maximum diameter. There 1s
hardly any anal canal. The opening of the anus is marked by a
depression. here is a lip-like projection between the anus and
the vulva.
Text-figure 54,
100 ys
Memphisia memphisia Khalil. Lateral view of cephalic portion of body.
Nerve collar.—The thick nerve collar is placed round the
cesophagus *25 mm. trom the cephalic end.
Genital organs.—-Male: The testes are convoluted irregularly,
reaching to within 3mm. of the head. There is a dilated,
elongated vesicula seminalis. he ejaculatory ducts penetrate
the cement gland. This is of enormons length, and is apparently
divided into three parts by deep grooves. The ejaculatory ducts
open into the cloaca.
Female: The thin-walled vagina proceeds cephalad in the
long axis of the body. The two strongly muscular ovejectors
open into its apex side by side, connecting it with the two dilated
PARASITES OF ELEPHANTS. 959
uteri, which are packed with eggs. The ovaries are irregularly
convoluted, reaching within 4-3 mm. of the head. The vagina is
1:7 mm. long.
Bursa.—The bursa is elongated, and distinctly divided into
three lobes. The dorsal lobe is nearly double as long as the
lateral lobes. The whole bursa is -75 mm. Jong and °55 mm.
broad. The ventral, lateral and externo-dorsal rays are crowded
together, while the branches of the dorsal ray are wide apart.
The bifid ventral ray is placed near the anterior edge of the
bursa. The lateral, in addition to the usual three branches, gives
a fourth stumpy ray, directed caudally. The externo-dorsal ray
Text-figure 55.
Text-figure 56.
Memphisia memphisia Khalil.
Ovum.
Memphisia memphisia Khalil,
Female tail.
does not reach the edge of the bursa. Near its origin it gives a
long branch, directed caudally and kaving a rounded extremity.
Tt is ‘1 mm. in length, and is constantly present in all the males
examined.
The dorsal ray is °7 mm. long. It gives a branch on either
side before ultimately dividing into two rays. ‘The first two
branches have a double papilla at their end. The terminal
branches end a long distance apart at the caudal extremity of the
bursa.
Genital cone.—The genital cone is placed nearer the ventral
than the dorsal surface of the body. The cloacal opening is
placed a little posterior to its apex, and is surrounded with four
ill-defined cuticular knobs.
260 DR. M. KHALIL ON THE NEMATODE
Spicules.—The two similar spicules are ‘9 mm. long. They
are thin, with a fine termination bent dorsally. There is
i Short accessory piece ‘1 mm. long, curved on itself, form-
ing a canal for the spicules, and also arched in the longi-
tudinal axis; the concavity produced is directed dorsally. The
cephalic third of the spicular sheath is finely striated trans-
versely.
Termination of female.—The vulval opening is surrounded
by a prominent lip, especially on the caudal side. This latter
Text-figure 57.
Text-figure 58
Memphisia memphisia Khalil.
Male bursa, ventral view.
Memphisia memphisia Whalil.
Male bursa, lateral view.
separates the vulva from the anus. Both openings are directed
practically caudally. The distance between the two openings is
‘08 mm. At this level the diameter of the body is suddenly
constricted, changing from *26 at the level of the vagina to
‘Lmm. a little below the anus. The tail is 58 mm. long, and its
end is pointed.
Ova.—Ova found in the vagina are in the morula stage, and
measure 55 pw by 35 p.
Habitat. Intestine of African elephant (Uganda).
PARASITES OF ELEPHANTS. 261
Mempuista AzizA Khalil, 1922. (Text-figs. 59-64).
Material.—The material consists of more than sixty specimens
sorted from Prof. Leiper’s collection.
Shape of body.—The body is slender. It is slightly shorter
than I. memphisia. The males are 12 mm. long and the females
are 15 mm. The maximum diameter is about the middle of the
body, being *5 mm. iv both sexes. The head end is very narrow,
being ‘1 mm in diameter. There is a cuticular collar, less pro-
jecting than in Jf. memphisia. There is no cuticular expansion.
The female tail is less abruptly constricted than is the type-
species.
Text-figure 59.
$04
Memphisia aziza Khalil. Wentral view of head.
Skin.—The cuticle is very finely striated. The striations are
at intervals of ‘Ol mm. The cuticular collar is demarcated from
the body by a shallow groove.
Mouth collar.—The mouth collar is distinctly separated from
the rest of the body by a groove. It is irregularly rounded
when seen laterally, and almost quadrangular when seen from
the ventral or dorsal surface. The latter appearance is due to
the presence of the lateral head papille being on the extreme
262 DR. M. KHALIL ON THE NEMATODE
edge of the mouth collar, The mouth collar is ‘02 mm. long and
‘] mm. in diameter. .
Head papille.—The four submedian head papille are fairly
Jong, with a knob-like mass on their top. This is continuous
with a duct-like canal passing in the substance of the papille and
previously alluded to as possibly the duct of a corresponding
head gland. The two lateral papille are small and delicate, and
directed mainly laterally, giving the square-like appearance to
the head.
External leaf-crown.—This is composed of 24 thin leaflets
surrounding the oval-shaped mouth-opening. Seen from the
Text-figure 60.
50 ps
Memphisia aziza Khalil. Ventral view of head.
ventral aspect the rays in the centre project more than those
on either side. :
Cervical papille.—The long and thin cervical papille are placed
82 mm from the cephalic extremity.
Mouth capsule.—The oral cavity is longer than it is broad. It
is pear-shaped, being narrow at its outlet. The chitinous ring
surrounding the oral cavity is very thick, and almost rectangular
in optical section. It is -045 mm. in length and -072 mm. in
diameter. A thin chitinous offshoot surrounds the oral cavity
in its terminal half. At the floor of the mouth capsule there is
a ring of 24 sharp teeth, probably representing an internal leaf-
crown. ;
Cephalic glands.—There are four cephalic glands surrounding
PARASITES OF ELEPHANTS, 263
the anterior part of the cesophagus from the level of the nerve
collar to the mouth collar. They are placed in the submedian
planes. Hach has a long, thick duct, which can be traced easily
to the level of the groove separating the mouth collar from the
body. As has been mentioned already, probably these ducts
open on the top of the submedian head papille. Their contents
are big granules in the centre, surrounded by fine granules which
also fill the ducts.
(Hsophagus.—There is a shallow cesophageal funnel sur-
rounded by a thick chitinous wall. The lumen of the
anterior ‘O07 mm. of the cesophagus is ridged, having a
plumose appearance similar to that seen in P. africana. The
rest of the lumen of the esophagus is -4 mm. in length and
Text-figure 61.
Memphisia aziza Khalil. Lateral view of cephalic portion of body.
‘14 mm. maximum diameter. It is slightly constricted at
the site of the nerve collar. The caudal termination of the
csophagus projects into the lumen of the chyle intestine,
forming three small lobes.
Chyle intestine.—The cellular wall of the chyle intestine is
pigmented throughout its entire course. It pursues a nearly
straight course.
Rectum.—This is a short tubular structure, with chitinous
wall sharply demarcated from the rest of the intestine.
Hacretory system.—The excretory vesicle is large and oblong.
The short excretory ducts open in the median ventral line ~
*72 mm. from the cephalic extremity.
964 DR. M. KHALIL ON THE NEMATODE
Nerve collar.—This is placed *2 mim. from the head surrounding
the cesophagus.
Genital organs.—Male: The irregularly convoluted testes reach
within 2°5 mm. of the head extremity. There is a dilated
seminal vesicle. The ejaculatory duct traverses the long cement
gland to terminate in the cloaca.
Female: The vagina pursues a vertical course in the long
axis of the body. It is 52mm. long. Into its cephalic end the
two strongly muscular ovejectors open, each connected with the
corresponding uterus. The two uteri lie side by side, lying
mainly in the axis of the body. The irregularly convoluted ovary
reaches within 2°75 mm. of the head end.
Text-figure 62.
Text-figure 63.
Memphisia aziza Khalil.
Ovum.
Memphisia aziza Khalil.
nl 1 I
Female tail.
Bursa.—The bursa is distinctly divided into three lobes.
The dorsal lobe is much longer than the lateral. The bursa
is smaller than that of J/. memphisia, but corresponds to it in
shape and in the arrangement of the rays. It is 5 mm. in length
and:45 mm.in breadth. The ventral and lateral rays are similar
to the corresponding rays in the type-species. The fourth division
of the lateral ray is shorter and knob-like. The externo-dorsal
ray does not reach the edge of the bursa. It gives off near its
origin a small ray, which is characteristic of the genus. It is
shorter than that in J. memphisia, being ‘05 mm. in length,
and its end is pointed.
PARASITES OF ELEPHANTS. 265
The dorsal ray is ‘47 mm. long. Each of its first two branches
terminate in two papille. The terminal two branches terminate
close to each other at the caudal end of the bursa. .
Genital cone.—This blunt cone projects from the roof of the
bursa nearer the ventral than the-dorsal surface. The cloacal
opening is a little dorsal to its apex. It is surrounded by
cuticular bosses.
Pre-bursal papille.—The pre-bursal papille are fairly long, and
are placed *7 mm. from the caudal end of the bursa.
Text-figure 64.
Memphisia aziza Khalil. Male bursa.
Spicules.—The two similar spicules are -75 mm. long. Their
tips are curved, pointing dorsally. The spicular sheath is trans-
versely striated in its upper third. The accessory piece is *] mm.
long, and is curved from side to side, enclosing the spicules. It
is also arched in the longitudinal axis; the concavity is directed
dorsally and caudally.
Termination of the female.—The tail of the female tapers
gradually to a point, unlike the suddenly constricted tail of J
memphisia. The vulva is placed -25 mm. cephalad of the anus.
The tail is ‘66 mm. long. There are two caudal papille, marked
by a slight depression in the lateral line. These are placed
“14mm. from the tip of the tail.
966 DR. M. KHALIL ON THE NEMATODE
Ova.—The ova, as seen in the vagina, are thin-shelled. They
are 60 » in length and 35, in breadth. The contents are in the
morula stage of development.
Habitat. Intestine of African elephant (Uganda).
Discussion.
This genus is in some respects allied to Pteridopharyna. The
presence, however, of the cuticular collar round the anterior end
of the body, and Ane peculiarity of having a branched externo-
dorsal ray in the bursa, clearly differentiate the members of the
genus Memphisia.
The two species here described are easily sorted out, even with
a hand lens. The very marked and projecting pu sitolare collar
of M. memphisia, together with the spike-like female tail, suffice
to differentiate them. The spicules in J/. aziza are shorter;
the distance between the anus and vulva is longer than in
M. memphisia. In addition, the total length and the diameter
of the various structures of the body and the position of
the cervical papillae and excretory pore are different in both
species.
No member of this genus has been recorded from the Indian
elephant.
Genus Amira Lane, 1914 (amended).
Fairly small worms with thick cuticle. Cuticular bosses may
be found anterior to the male bursa. The mouth is terminal,
and is surrounded by an external and an internal leaf-crown.
There are six head papille, none of which are prominent.
The mouth capsule is very short; the cesophagus is hour-glass
in shape, the nerve collar surrounding the constriction. There
is a large esophageal funnel. The cesophagus has a cuticular
lining.
The male: The dorsal lobe of the bursa may or may not
be enormously elongated. The pre-bursal papille are very much
elongated. ‘The two equal spicules are of enormous length and
very fine. There is an accessory piece.
The female: The vulva lies close cephalad of the anus. The
long vagina runs cephalad, and divides into two parallel cephalad-
running uteri provided with ovejectors. The colourless thin-
shelled ova are seemented.
Type, Amira pileata Railliet, Henry & Bauche, 1914.
Amira PILEATA Raill., Henry & Bauche, 1914. (Text-figs. 65-68.)
Cylicostomum pileatum Raill., Henry & Bauche, March 1914.
Amira omra Lane, July 1914.
Source of Material.—The material consists of one male and five
females, which were sent to Prof. Leiper by Lieut.-Col. Clayton
Lane. They are the co-types of Amira omra.
PARASITES OF ELEPHANTS, 267
Historical Review.—This species was independently described
by Railliet and his co-workers on the one hand, and by Lane on
the other. The descriptions in some respects are complementary.
Lane definitely mentions that an accessory piece is absent.
Railliet and his colleagues do not mention it at all. There is
important discrepancy in the bursal rays as described and drawn
Lane records the presence of a long pre-bursal papilla assuming
the appearance of a ray and lying close in front of the double
ventral ray. This is completely missed by the other observers.
On the other hand, Railliet figures two small branches to the
dorsal ray, while Lane records one only. The head end is well
Text-figure 65.
Amira pileata Raill., Henry & Bauche. Cephalie portion of body.
represented in the diagram of Railliet. It is thought advisable
to re-describe the bursa.
The Bursa is greatly elongated. This is due mainly to the
length of the dorsal lobe. The bursa is 1:9 mm. in length
and 0-55 mm. in breadth. ‘The cuticle in front of the bursa on
the ventral surface is very much thickened. The free edges of
the lateral lobes of the bursa are folded inwards, obscuring the
exact mode of ending of the lateral rays.
The pre-bursal papilla is very long and thin. It pursues a
wavy course, and ends close to the termination of the ventral
ray. It is 0°25 mm. in length.
268 DR. M. KHALIL ON THE NEMATODE
The ventral ray is bifurcated in its terminal half. The two
branches lie close together. The anterior branch is thinner than
the posterior.
The three lateral rays have a common origin. The postero-
lateral ray branches before the other two rays. All three rays
lie close to each other throughout their entire course. Their
terminal third is folded inside the bursa.
The externo-dorsal ray is slender, and pursues a curved course.
It does not reach the edge of the bursa, but ends by a papilla on
the external surface.
The dorsal ray is unique in virtue of its enormous length. It
measures 1°67 mm., and divides high up within -15 mm. of
its origin. After a short course each primary division gives off
Text-figure 66.
Amira pileata Raill., Henry & Bauche. Male bursa.
a long and a short branch lying close to each other. The
longer branch is "12 mm. and the shorter ‘05 mm. in length.
These branches appear as offshoots from the main stem, which
continues a straight course, parallel and close to the corre-
sponding ray of the other branch. Both end in a sharp point.
Genital cone.-—The cloacal opening is placed near the apex of
an elongated genital cone. ‘The cone projects from the roof
of the bursa between the pre-bursal papille. It is placed
ventrally and caudally, and is °18 mm. in length and ‘04 mm. in
breadth. The cloacal canal traverses its centre. Its opening
is overlapped by its projecting anterior lip, which forms the
apex of the cone.
PARASITES OF ELEPHANTS. 269
Along the dorsal wall of the cloacal canal is placed the accessory
piece. It is grooved on its ventral surface with curved edges.
The cephalic extremity is slightly broad. The caudal lies a
Text-figure 67.
Text-figure 68.
Amira pileata Raill., Henry & Bauche.
Ovum.
Amira pileata Raill., Henry & Bauche.
Female tail.
short distance cephalad of the apex of the genital cone. Its total
length 1s °165 mm.
Habitat. Cxecum of the Indian elephant (India).
AMIRA SAMEERA, sp. n. (Text-figs. 69-71.)
Source of Material.The material consists of a single male
discovered accidentally amongst the type material of Pterido-
pharynex africanw Lane, deposited in the Natural History
Museum. I was enabled to examine this material through the
kindness of Dr. Baylis. It was collected at Lawrence Hill,
Addo Bush, Cape Province, Africa, and was presented to
the Natural History Museum by the Imperial Bureau of
Entomology in August 1919. This collection had been examined
by Lieut.-Col. Clayton Lane, and had been separated by him
into two hitherto undescribed species. Careful search through
the whole material failed to discover further specimens of
A. sameera.
Shape of body.—Body is stout in comparison with its length.
It is 10°5 mm. long. The greatest diameter of the body is
about its middle, where it is -45 mm. It tapers slightly
towards the head, and also towards the caudal end, just
anterior to the bursa, which, however, gives a truncated
appearance to the posterior extremity of the male.
Proc. Zoou. Soc.—1922, No. XIX. 19
270 DR. M. KHALIL ON THE NEMATODE
Skin.—The skin is striated to within a short distance of the
head. The striations are crowded near the cephalic extremity,
where they are ‘02 mm. apart; at the middle of the body they
are ‘025 mm. apart, while near the bursa they are far between,
being on an average ‘06 mm. apart.
The thickness of the cuticle is ‘02 mm. in the cephalic two-
thirds of the body; caudally it becomes thicker. Near the bursa
the cuticle reaches its maximum thickness, ‘08 mm.
Mouth collar..—This is distinctly separated from the rest of the
body by a shallow groove. It is regularly rounded in outline.
Its length is (048 mm. and its diameter -25 mm.
Text-figure 69.
Amira sameera, sp.n. Cephalic portion of body.
External leaf-crown.—The inner surface of the mouth collar
is occupied by a circular crown consisting of 36 leaflets. These
leaflets are slender, -065 mm. in length, and tapering at their
free end.
Head papille.—Vhe four submedian head papillee are short and
conical. ‘The two lateral papille have a rounded top.
Cervical papille could not be made out in this specimen.
Mouth capsule.—The shallow mouth capsule is ‘05 mm. deep
and -165 mm. in diameter. The chitinous plate surrounding the
mouth is thick and curved. Its concavity faces the lumen of the
mouth,
PARASITES OF ELEPHANTS. 271
Internal leaf-crown.—This is placed over the top of the
chitinous ring of the mouth capsule. It consists of 36 very short
leaflets corresponding to those of the external crown.
(Hsophageal funnel.—This is large and characteristic of the
genus. Itendsat the level of the nerve-ring where the esophagus
is constricted. There are three chitinous plates lining this
funnel as well as the rest of the esophagus. The cavity of
the funnel is 21 mm.in length and :175 mm. maximum diameter.
(Gsophagus is short and thick. It is bulbous at either end and
constricted where the nerve collar is placed. The total length of
the esophagus 1s ‘6 mm.; the anterior portion is °21 mm. and
the posterior 39 mm. The diameter of the anterior portion is
-25 mm., while the posterior is °22 mm. The posterior end of the
Text-figure 70.
100 ft
Amira sameera, sp.n. Dorsal view of bursa.
cegophagus projects into the beginning of the chyle intestine,
forming three small lobes. wilt 8
Chyle intestine—The course taken by the chyle intestine is
practically straight. Its wall is thick and the cells are not
pigmented.
Excretory system.—The excretory pore is placed *7 mm. from
the head end. The thin-walled excretory -vesicle lies close by
and has a thin wall.
Nervous system.—The nerve collar is 33 mm. from the head
end. It lies in the deep constriction of the cesophagus.
Genital organs.—The convolutions of the testes reach to within
1-3 mm. of the head. They run for the most part in the axis of
the body. The cement gland is nearly half the length of the
noe
sida 19%
272 DR. M. KHALIL ON THE NEMATODE
Bursa.—The bursa has a wrinkled appearance. Its edges are
curled inside. The dorsal is indistinctly separated from the
lateral lobes, It is not elongated as in A. pileata. The general
arrangement of the rays, however, is common to both species.
The bursa is °67 mm. long and ‘54 mm. broad.
The pre-bursal papilla is thin and wavy, and is ‘3 mm, in length.
It is easily mistaken for a ray.
The ventral ray is bifid in its terminal half. The two branches
lie close together. |
The lateral] rays lie close together, and their terminations are
bent inwards exactly in the same manner as in A, pileata.
The externo-dorsal ray is long and ends very near the edge of
the bursa.
Text-figure 71.
Amira sameera, sp.n. Lateral view of bursa.
The dorsal ray is massive. It divides at the level of the origin
of the externo-dorsal ray. Each of these primary divisions gives
a thick, short lateral branch, which almost immediately divides
into two. These are very short and do not reach the edge of the
bursa. The main stem is longer and ends near the corresponding
ray of the other side.
The length of the dorsal ray is °25 mm.
Genital cone.—Vhe genital cone is sharply pointed. It is,
however, a more massive structure than that of A. pileata. The
apex of the cone lies nearly in a line with the ventral surface of
the body. It is not marked with any cuticular thickenings.
Genital opening.—-The cloacal opening is placed practically at
the apex of the genital cone.
PARASITES OF ELEPHANTS. 573
The spicules.—The elongated spicules pursue a wavy course.
They are about 2°9 mm. in length. Their tips are thickened.
The accessory piece is short, 12mm. in length. It is strongly
curved from side to side, and less so in the longitudinal direction.
The concavity of the latter curvature is directed caudally and
dorsall y.
Habitat. The stomach of the African elephant (South Africa,
Addo Bush).
Discussion.
The striking difference between the two species of the genus
Amira is the shape of the bursa. On careful study, however,
this difference is minimized by the great similarity between the
arrangement of all the rays and the pre-bursal papille. The
dorsal rays in both species have six branches each. Four of these
branches are little more than bud-like processes. The remaining
two branches are enormously elongated in A. pileata and short in
A. sameera. This I considered as a specific difference. It is not
justifiable to regard it as generic. For this reason I have had
to amend Lane’s generic diagnosis. The configuration of the
mouth, the csophagus, and spicules are strikingly similar, with
differences as regards size only.
Genus Equinurpia Lane, 1914.
Fairly large and stout bursate nematodes. There are an
external and an internal leaf-crown. The mouth capsule and the
oral cavity are more or less globular. The dorsal esophageal
gland discharges through a dorsal gutter.
Male: There are 11 pairs of rays supporting the male bursa,
of which two are ventral, three are lateral, three are externo-
dorsal, and three are dorsal. The spicules are equal and similar.
There is ne accessory piece.
Female: The vulva opens on the summit of a prominence
adjoining the anus. The uteri are convergent and lie parallel to
each other, and are furnished with weak ovejectors.
Type-species, Lquinurbia sipunculiformis.
EQUINURBIA SIPUNCULIFORMIS Baird, 1859.
Sclerostoma sipunculiforme Baird, 1859.
Nematode No, 4. Evans & Rennie, 1910.
Cylicostomum sipunculiforme Raill., Henry & Bauche, 1914.
Hquinurbia sipunculiforme Lane, 1914.
This species has the characteristics of the genus. Males are
15 mm. long and 1:3 maximum diameter. The female is 27°5 mm.
long and 15mm. maximum diameter. The spicules are 1°5 mm.
long. The tail of the female is 0°8 mm. long. The ova are 60 p
in length and 80-35 p» in breadth.
Habitat. Cecum of the Indian elephant (India).
274 DR. M. KHALIL ON THE NEMATODE
Family ANcYLOStomiDbD# Looss, 1911.
Sub-family AncyLosromin# Raill. & Henry, 1909.
Genus GRrAMMOCEPHALUS Raill. & Henry, 1910.
The head is curved towards the dorsum. Mouth capsule is
wide anteriorly and narrowed posteriorly. There are three teeth
projecting from the dorsal and lateral walls of the mouth capsule.
(Esophagus is long and simple. rom the dorsal wall of the
intestine, close to its union with the cesophagus, springs a long
diverticulum, running cephalad. The dorsal rays are bifid near
their tips. There are two similar spicules and no accessory piece.
Vulva lies close to the middle of the body. The short vagina is
joined by two divergent uteri.
Type-species, Granmmocephalus clathratus.
GRAMMOCEPHALUS CLATHRATUS Baird, 1868.
Sclerostoma clathratum Baird, 1868.
Strongylus clathratum Cobbold, 1882.
Grammocephalus clathratum Rail. & Henry, 1910.
Male is 45°52 mm. in length and 1:15 mm. in maximum
diameter.
Female is 36 mm. long and 1:03 mm. in maximum diameter.
The median dorsal tooth is not particularly prominent. The
intestinal diverticulum is 2°3mm. long. The spicules of the male
are 1'2mm. in length. The female tail is 0°67mm. long. The
vulva is 17 mm. cephalad of the anus. ‘The main stem of the
dorsal ray is longer than that of the next species.
Halitat. Bile-ducts of the African elephant (South Africa).
GRAMMOCEPHALUS VAREDATUS Lane, 1921.
Probably the supposed G*. clathratus reported from the Indian
elephant were really this species. ‘These were :—
Strongylus clathratum Cobbold, 1882.
Sclerostomum clathratum Piana & Stazzi, 1900.
Nematode No. 1. Evans & Rennie, 1910.
Grammocephalus varedatus Lane, 1921.
This is a somewhat longer species. The males are 55 mm. long
and the females are 47 mm. ‘he dorsal ray divides close to its
origin. The spicules are 1°35 mm. in length. The female tail
is O'8mm. in length, and the vulva lies 22mm. cephalad of
the anus.
Habitat. The bile-ducts of the Indian elephant (India and
Burma).
bo
a
Or
PARASITES OF ELEPHANTS.
Genus Baramostomum Raill. & Henry, 1909.
Small nematodes having the head curved towards the dorsum,
the oral aperture facing anteriorly and dorsally. The oral
capsule is fissured on its dorsal and lateral aspects. Its internal
surface is raised mainly ventrally and laterally into shelf-like
projections encroaching on the oral cavity. A small dorsal tooth
surmounts the dorsal column of the esophagus, which is simple
and cup-shaped.
Male: The dorsal rays are Separate for nearly their whole
extent, the externo-dorsal rays springing from the individual
dorsal rays. The lateral rays turn ventrally. The spicules are
stout. There is no accessory piece.
Female: The vulva lies near the middle of the bedy. The
uteri are divergent, furnished with str ong ovejectors.
Type-species, Bathmostomum sangert.
BATHMOSTOMUM SANGERI Cobbold, 1879.
Dochmius sangert Cobbold, 1879.
Uncemaria sangert Railliet, 1896 (not Alessandrini, 1905).
Uneinaria os-papillatum Piana & Stazzi, 1900.
Baithmostomun sangeri Raill. & Henry, 1909.
The oral cavity is wider than long. There are two sub-ventral
teeth. The male is 15-16mm. long and the female is 20mm.
long. The spicules 0-47 mm. in length. Vulva lies near the
middle of the body
Hatitat, Cecum of Indian elephant (India).
Genus Bunosromum Raill., 1902.
Capsuled bursate nematodes having the mouth guarded by
ventral semilunes. The dorsal and externo-dorsal rays ave asym-
metrical. The mouth cavity contains a dorsal tooth, being the
freely projecting ducts of the dorsal cesophageal gland. There is
a pair of sub-ventral lancets at its base.
Bunostomum rouiatum Cobbold, 1882.
Strongylus foliatus Cobbold, 1882.
Unemaria sangert ‘Messandrini) 1905 (not Railliet, 1896).
Bunostonvum foliatuwm Raill., Henry & Bauche, 1914.
This nematode was imperfectly described by Cobbold in 1879,
and has never been recorded since. Its inclusion in the genus
Bunostomum is regarded by Railhet as a provisional matter.
Probably it will prove to be a representative of a new genus.
"GQBL “Plogqon wngn270f gq ‘OGL “Tey WaWorLsoNnag
a ————__- —__ ——+ ee —. “TI6L ‘ssoory
“GLBT ‘PlOqqo,) tlabuns “gq | ‘6061 “H ¥ Wf WaWworLsownrveg
—_— wa. SS SS me a Oo el ae al ei ae WAINOLSOTAONV
"BORL “parece si2pp.0y70)0 *4D ‘TSG “OUR'T sip patma “H ‘OI6L “H YY SATVHamOoWN VAY
| “OOGBT “12Z04g I BURT, wnumojzsida “D TIGL “A YH “YW WaAINvVINoHD |
“OSRT ‘pag, sywcofipnowndis ap PIGL ‘OUBY] VIddONIOO Hy
PPLGL Sl PCE OM PlpeUIppY visn.woaq “PI6L ‘OUBTT VISOYO A]
‘ucds “puaauns “FF | PIG ‘oyonng x pT “py wpvaqid py ‘PIGL ‘ouvry vary
AA SUES ERAS CNS To Se ne genres | osmer Gavia Srp
‘COBL “[LRUM peseyduau “py | 666L [YUM VIsthawa ji | “FORT “ploqqog
ESTE UCT LD ag, ae
“1G6L ‘ouLy vuvowt fd *
‘TS6L ‘ouvTy XNXUVHdoaruaLg | WALTADNOULS
‘uds ‘pordoryja *
“ucds “ppnrvoinetg “QO |
“GOST ‘TBI vpunbn °% |
“1ZGL ‘ouvry puvortf[p °& | “S16L “Ap H LiuazPy “yy eazeewaw
OLBL “Jspooper) sesuardy °% |
“PIGL ouvyT VtovaUy.t7 “
“PIGL “ouLury VINOTIA?)
“B88 “PIO Mlaliomf “TL |
"PIGBL OUT VpLysanrue “Pr
“ud amopswue) “Tir
| ‘tT eT & Na Tr
‘BGT MMM Pepe Te De ae SEN
‘GSST SPlOgqoD wwiygeus *q *EQgT ‘AdpteG
“LOGL ‘suydegr punorefp «gq | “1LZGBL “SAU, wnowpur *gT | “TEG6L ‘St[Avgr VWGNOMEVUV WaIMAWIds
| "FOBT “PIOdqoD
“TEST “ousony wtazdoyouo) “gg | “LOGL ‘lodiaryT sTuyosviag WAIN VOSV
| | "GIGI ‘sossvawiy,
“GAG “YM Blade) 7 | “GCGL “TRY 199196 | “GEG “WIVUM VINGuad Ia] W CI.LOV aL
‘yuvydapy Uvonyy ur soedg | ‘quuydapgy uurpuy ur sermadg | *snulay) “ATU IP
‘queydery og wor pepz0oa.t SIJISBILA O9popwMEN [[v JO uorqtsod orp rmeysMQ—' PTT ATAvy,
THE NEMATODE PARASITES OF ELEPHAN'S. Qe4
ComPiuTE BiBiioGrapuy of the Parasites of Elephants.
ALESSANDRINI, G. 1905. Si di aleuni Uncinariw parassite dell’ uomo e di altri
vertebrati. Boll. Soe. Zoo}. Ital. ser. 1. pts. 1-8.
Barep, W. 1859. Description ofa new species of Entozoon (Selerostoma
sipunculiforme) trom the intestine of the Elephant.
Proc. Zool. Soe. London (410), pt. 27, pp. 425-427.
Bayuts, If. 1521. A new genus of Nematodes parasitic in Elephants.
Parasitology, vol. xiii. No.1. Cambridge.
Necropsia di una elephantessa. Cisti d’ echinococco
nel egato e nei polmoni. Bologna.
CoxnBo.xp, T. 8. 1869. Description of species of Trematodes from the Indian
Klephant. Quarterly Journ. of Microscop. Sci.,
January.
*BoOnvicini, A. 1897.
5 * 1869. Ibid. in Supplement to Entozoa, p. 80.
* 3 1875. Further remarks on Parasites from the Horse and
Klephant, with a notice of new Amphistome from
the Ox. The Veterinarian, November.
a 1879. Parasites of Pachydermata in ‘A General Treatise
on Parasites,’ Part X., sect. J., Book II., p. 353.
London,
os %. 1882. The Parasites of the Elephants. Trans. Linn. Soc.
London, 2nd Ser., Zoology, vol. ii. pt. 4, pp. 223-
288, pl. xxili.-xxiv.
Dirsine, K. M. 1851. ‘Systema Helminthum,’ vol. ii. pp. 176 et 560.
* 45 1857. Sechzehn Arten von Nematoiden. Denkschr. d.
math.-nat. Cl. d.k. Akad. d, Wissensch. Bd. x., 111.
p. 18, Taf. 1. figs. 14-17.
e is 1858. Revision der Myzelminthen. Vienna-Academy
teports, Bd. xxxu. p. 35.
2p 5 1861. Kleine helminthologische Mittheilungen. Sitzungsb.
d. k. Akad. d. Wissensch. Wien, math.-naturw. Cl.
Bd. xli., i. Abt. (4), pp. 269-282.
Drascue, R. 1882. Revision der in der Nematoden-Sammlung des k.-k.
Zoologischen Hofcabinetes hefindlichen Original
exemplare Diesing’s und Molin’s. Verhandl. d.
k.-k. zool.-bot. Gesellsch. in Wien, Bd. xxxii.
p. 193, Tat. xi—xiv. fig. 21.
Evans, G. H., & 1908. Notes on some Parasites in Burmah.—Il. Journ.
Rennie, T. Trop. Vet. Sci. Calcutta, vol. iv. pp. 184-148,
pls. v.-vul.
" es 1910. Notes on some Parasites in Burmah.—III. A few
common parasites of Klephants. Journ. Trop.
Vet. Sci. Calcutta, vol. v. pp. 240-250.
*Fitg, R. H. 1876. “ Anatomy of Fasciola jacksoni.” Reported from
Boston See. of Med. Scien. in the New York Med.
Journ., November.
GEDOoELST, L. 1911. Synopsis de parasitologie de Vhomme et des animaux
domestiques, xx+832 pp.; p. 327, fig. 1. Lierre
& Bruxelles.
1916. Notes sur la faune Parasitaire du Congo Belge. Rev.
ae if Zool. Africaine, vol. vy. Fasc. i. December.
*Horyra, F., & 1918. Spezielle Pathologie und Therapie der Haustiere-
Marek, I. Kvankheiten der Atmungsorgane der Verdauungs-
organe des Nervensystems der Bewegungsorgane
und der Haut, xiv+1088 pp.; 207 figs., 7 pls.
Jena.
* Not available for reference.
278 DR. M. KHALIL ON THE NEMATODE
Inte, J. E. W.
EP) 35
*Jackson, J. B.S.
Kwai, M.
Lane, C.
von Linstow.
Lorenz, L.
Mirtrer, 8S. N.
Prana, J. P., &
Srazzi, P.
PorriER, J.
Rarcurer, A., &
Henry, A.
oP) oP)
RaiLuieT, A., Henry,
A., & JovEux, C.
bb 39
Ratiurer, A., Henry,
A., & Baucus, J.
bP 39
Rupowpaut, C. A.
*Srazzi, P.
Strives, W., &
GOLDBERGER, J.
Stossicu, M.
1919. Ueber Ancylostoma perniciosum vy. Linstow und die
1919.
1847.
1922.
1907.
1880.
Strongyliden des Hlefanten. Bijdr. tot de Dierk.
Afl. xxi. p. 97.
Notiz zu meinem Aufsatz Ueber Ancylostoma ete.
Centralb. f. Bakt. etc., orig. Bd. viii. Heft 7.
Descriptive Catalogue of the Medical Improvement
Society’s Cabinet, Boston, U.S.
A preliminary Note on some new Nematode Parasites
from the Elephant. Ann. & Mag. Nat. Hist.,
February.
. Bursate Nematodes from the Indian Elephant.
Indian Journ. of Med. Research, vol. 11. No. 1,
rp. 380.
. A further Note on Bursate Nematodes of the Indian
Elephant. Indian Journ. of Med. Research,
vol. 111. No.1, p. 105.
. Some Bursate Nematodes from the Indian and A frican
Elephant. Indian Journ. of Med. Research, vol. ix.
No. 1, pp. 163-172.
. Entozoa des zoologischen Museums der Kaiserlichen
Akademie der Wissenschaften zu St. Petersburg.
Ann. Mus. Zool. Acad. d. Sc. de St. Petersburg,
vol. viii. (3-4), pp. 265-295, pls. 17-19.
Nematoden aus dem Koeniglichen Museum in
Berlin. Mitteil. aus dem Zoolog. Mus., Berlin,
vol. 111. No. 8, p. 255.
Ueber Distomum robustum, sp. n., aus dem afrikan-
ischen HElephanten. Verhandl. d. k.-k. zool.-bot.
Gesellsch. in Wien, vol. xxx. p. 5838, figs. 1-6.
. Some Entozoa of Indian Elephants. Journ. of
Comparative Path. and Therap. vol. xxv. part 2.
pp. 111-115, 1 pl.
. Elminti intestinale di une elefantissa. Archiv.
Parasitolog. vol. ili. pp. 5389-529.
. Tvématodes parasites de l’éléphant d’ Afrique. Paris
C. R. Assoc. Frang. Avane. Sci. xxxvii. (Clermont-
Ferrand, 1908), Notes et Mém. pp. 580-596.
. Sur la classification des Strongylides. C. R. Soc.
Biolog., Paris, vol. Ixvi.
. Quelques Helminthes nouveaux ou peu connus du
groupe des Bunostomiens. Bull. Soc. Path. Exot.
vol. 111.
. Sur deux Trématodes des Primates. Bull. Soc. Path.
Exot. vol. v. No. 10.
. Un nouveau Strongylides des Singes. Bull. Soe.
Path. Exot. vol. vi.
. Sur les Helminthes de ’éléphant d’Asie. Bull. Soe.
Path. Exot. vol. vii. Nos. 1, 2, &3.
. Sur les Helminthes de l’éléphant d’Asie. Note
complémentaire. Bull. Soc. Path. Exot. vol. viii.
pp. 117-119.
. Synopsis Entozoorum, p. 36. Berlin.
. Lelmintiasi nell’ elefante. Mod. Zooiatro Torino,
vol. 11. (23), 10 Dic. pp. 451-454 (24); 25 Dic.
pp. 466-69.
. A Study of the Anatomy of Watsonius (n. x.)
watsoni of Man and of nineteen allied species, etc.
Hygienic Laboratory Bull. No. 60. Washington.
. Strongylidze Lavoro monografico. Bull. Soc. Adriat.
di Sc. nat. in Trieste, vol. xix. pp. 55-162.
* Not available for reference.
PARASILVES OF ELEPHANTS, 279
Vevers, G. M. 1920. Report on Entozoa collected from animals which died in the
Zoological Gardens of London during 8 months, 1919-1920.
Proc. Zool. Soc. London, Sept., pp. 405-410.
BrBuioGRAPeHy referred to in the text in connection with the
anatomy or systematic position of the parasites.
GILES. 1892. On the nodular disease of the intestine in Sheep. Scient.
Mem. of Med. Offic., India. Part vil. p. 36, fig. 1.
Caleutta.
von Linstow. 1906. Nematoden des Zoologischen Museums in KGnigsberg. Arch.
ftir Naturgeschichte, Band 1. Heft 3.
Looss, A. 1891. The Sclerostomide of Horses and Donkeys in Egypt.
Records of the Egyptian Government School of Medicine,
Cairo, p. 94.
1899. Weiterer Beitrag zur Kenntniss der Trematoden-Fauna
Aegyptens, zugleich versuch einer natirlichen Gliederung
des Genus Distomum Retzius. Zool. Jahrb., Jena, Abt. f.
Syst. vol. xii. (6-6), pp. 521-784.
Sampon, W. 1907. Note on a Filaria of the Red Grouse. Journ. Trop. Med. &
Hye., London, vol. x. p. 304, 1 fig.
Scort, T. 1906. Notes on British Copepoda: change of names. Ann. & Mag.
Nat. Hist. (7) xv. p. 460.
TRAVASSOS. 1919 (2). Espaco de uma chave ceral dis nemotodes parasitos.
Revistade Vetrinaria E Zeotechnia, Rio de Janeiro.
ADDENDUM.
Since the manuscript was sent to the press, a paper by
Gedoelst, entitled ‘ Quelques nématodes parasites de l’Hléphant
africain,”* has come to hand. The author describes a new species,
Ascaridia rodhaini Gedoelst, 1922, from the African elephant.
He also records a single female, Amira pileata, from the same host.
It is very probable that the latter is the female of Amira
sameerd, sp. n., described above. The author redeseribes dur
shidia recta von Linstow, which is renamed above I. linstowr,
sp. n. Unfortunately, however, Gedoelst’s paper is devoid of
illustrations.
* Bull. Soc. Path. Exot. t. xv. Séance, 8 Févr., 1922, No. 2.
4 itd ui f : oi
waft, (ch y, f
AM QE. ot
wear, we
a ey ut Dwi af
aE ae gs We lait -
aie Paap tity th ies ‘
shes nd fobviae fh tinted alan ifthed: A
ca 77 a ie nasal uefa tb: ae |
aA Ot pa we
CRIED. ATS
hot” Bebe ohne ir,
EY neich, pay)
ON DEATHS IN THE GARDENS IN 1921, 281
13, Report on the Deaths which occurred in the Society’s
Gardens during 1921. By N.S. Lucas, M.B., F.ZS.,
Pathologist to the Society.
[Received February 10, 1922: Read March 7, 1922.]
On January Ist, 1921, there were in the Gardens 563
mammals, 1618 birds, and 549 reptiles.
During the year 409 mammals, 924 birds, and 594 reptiles
were added, making a total for the year of 972 mammals, 2542
birds, and 1143 reptiles.
During the year 208 mammals, 519 birds, and 281 reptiles
have died. This gives a percentage of 21-4 for mammals, 20°4
for birds, and 24°5 for reptiles.
If, however, only those animals are taken which have lived
in the Gardens for six months or more, it is found that only
Table of Causes of Death.
| | | Numbers of
Causes of Death. Mammals.) Birds. Reptiles. Explanatory
|
| Notes.
Pulmonary disease. | |
TPMGOIENG TID Goooosare bos ong 990 soo nDA cee | Dal 33 5
Congestion of lungs ............... 16 44 | 25
Gidemayoflumigsiseesss.-heee cea ace il
JN OSGI Ort UBTREP G55 06005 sen coence ese Aa 1
Acariasis of lung 1 5 |
Kmpyema eyues lereceaseat cal 1 eo mebliteaiias 1
I BHROAEIANIGTS) ° Go0\3o5 Heaodd oboe. soe S00 cee 13 ne) ale
Cardiovascular. | | |
TERS DICRSHODIOTS! su52a33hnapnogris Soabee a2! ie 3 a
ATES TTI Bae Beedod Renee Goto saeco He 1 | 1
Rupture of mesenteric artery .. Don SI et
Rupture of splemic artery ......... | a eae ht H
Rupture of pancreatic artery ...| os | 1 | ae |
Renal disease. | |
INephiatist ec atedrm ern aceecene 13 16 2 2; |
Granuloma of kidney ....... .......) Me | 2 il 3
|
Alimentary Tract. |
Shommealeihlsur cesses sceecee tents 03 a peers | 4.
Acute dilatation of stomach ..... | a | |
Gastiiiisaamnet cant a: cin ndrsteee creel 1 | 1 1
| Gastro-enteritis! -...-4-.0--4e0+-- 2: 13 (0) 10
Perforation of gizzard ............| 6 | 2 0 5
A IGETIGISIR BREN. see eee sean en cate 41 183 a | 6
Constipation) (e.--s5-s--e-eee ese 2 1 @} Wf
IiintuSstise ephion ey eee eee eeel 4, 1 OOS 8
Gastric ulcer Ae Cire eee Neal
Ulceration of intestine ............ |
JEHOVDSUS AO Goo<c vn beoseose6sea coe! il a nae
IBEHO MIDIS, “agsancsnoabesbecbesonaesel 2 1 1 |
bo
fe 2)
bo
DR. N. 8S. LUCAS ON
Table of Causes of Death (cont.).
: | Numbers of
| Causes of Death. Mammals. Birds. Reptiles.) Explanatory
| | Notes.
Hepatic.
Fatty degeneration of liver......... aa 6 0) |
Myeloid metaplasia ............... an il 0 9 |
INGCTOSIS On UIVel eee. eteneeeieeeen- al 0)
Portal pysemia ita 0)
Pancreatic.
li mee anicreatitisi. ceric eccsaee ges 3 0) (0)
| Genital organs. |
Wheders]ombeelUaYes 00008 ang anc nen opo tse Sap @) g 1
Tnflammation of oviduct ......... 0 a 9
Postpartum sepsis ..............0-.. 1 0 0 10
Nervous System.
Congestion of brain Silhedan 1
Abscess of frontal sinus............ 1
Acute Infections. |
TINS ROWI@SGS “pages ton donna eanasce. 17 9 (0) 11
INGO SScc2 aan senene ondoncsd adanne seance 23
Septiczemia .........-0--00+.- 22.2 = 9 vf 2
Pyzemia Seen ai Bh 1 ae
[Efelinm bhitasis ereees see -ee er seeee 2 6 8
(CRATES OH PAW. copcenog0v00000 b30.208 cc 2 12 |
Malignant disease.
Carcinoma of thyroid 1 a 13
Sarcoma sas 1 2 14
| Miscellaneous Causes.
| Osteomalacea ........05--c sere Se 1 Bt 15
| TOYS OMI | sossaago2che sesso 8dr 220 apenas 3 5 3
| LO TENON, as cocoong0s coh vod so nop becca aa nee 36
TBS AROSWOUS)’ cas cognpa nog 390000097200 088boE 1 igh sur |
Hemorrhage, fractured femur ... il |
IDO wn CONeeree eee eee ee caee se 2 zis
ells ee nm ie iia cian Gah Ula 60 3 16 |
Destroyed by rats .............-.... il 13 0) |
INO GUE EANTTLEl son) s90 op oa8 do0 soa jdm aoe 3 16 14 |
INOW GATTM ONGC! 455000 son aosons oac44e 9 1 11
ID Yeo AA HOEOE! bso aps noo cma hee ext angnon sos 5 24. Toy)
98 mammals and 93 reptiles have died. This gives a percentage
of 10:0 for mammals and 8-1 for reptiles.
It is not possible to give an accurate figure for birds, since so
many of the smaller birds of the same species are kept together,
which makes it impossible to distinguish a bird which has been
in the Gardens a month from one which has been there a year.
Explanatory Notes.
1. Empyema.—This occurred in a black bear.
2. Nephritis.—The small incidence of this disease referred to
last year has continued. It is noted to increase among
the mammals in cold weather.
’
3.
4.
5.
6.
15.
16.
DEATHS IN THE GARDENS IN 1921. 283
Granuloma of kidney.-—These two cases were in a waxwing
and a sheldrake. The latter had enormous masses of
granulomatous tissue among which were small islands of
renal tissue, yet the bird was in good plumage and fat
and had lived for 18 years in the Gardens.
Stomatitis.—This shows a decrease.
Perforation of gizzard. Both cases were due to pieces of
wire and were found in ground-feedin g birds.
Knteritis.—This is really a group of diseases due to various
causes, all resulting in inflammation of the intestines.
The causes are at present unknown.
Constipation.—A remarkable case occurred in a_peccary.
The coils of intestine were adherent to one another,
thereby causing a vicious circle. There was nothing to
indicate which was the primary condition.
Intussusception.—Three of the four cases occurred in
spotted cavies.
Myeloid degeneration of liver.—In a double-striped Thicknee.
Postpartum sepsis.—In a Dingo. Only four puppies out of
eight were born alive, and of these three died early of
pneumonia and ulceration of intestine.
Tuberculosis.— Both this disease and mycosis show a low
incidence this year.
Caries of jaw.—This occurred in a hedgehog and a Malabar
squirrel.
Carcinoma of thyroid.—This occurred in a fox. There were
large secondary growths in the lungs.
Sarcoma.—These three cases were as follows :—-In the breast
muscles of a red-vented bulbul. In the neck muscles of a
Carolina duck. A melanotic sarcoma starting in the
suprarenal of a Lechee antelope.
Osteomalacea.—This occurred in a cockatoo.
Killed.—This heading includes those killed by order for
injury, or by their companions. It does not include those
killed for disease. These are entered under the diseases
found post-mortem.
ae
yalngatr ante
Totieaar | ere
ioe We he ed eae
| apes ecuen he ee
: CTs oe Dell chips
mer went at, ry hed jena cist
ao fo Pea ime
| Sp dewemitebeivelle Pr sth eae Be eRe praise fo
Ae palsanl ise shepoce Set ih. ldtent hodixee-fee
es ekg abs, atage ae -phlondiog A
+ ee ih ego batie waiload:
Shia vata helienecat el a: trvsasigy Aid
var ahod.cd Botlhe somal pales fog:, eotinod
pease iia aaeaty, #1 tetany ee tein: xa *
—— faired is e angi
a Hes Miss: “Was ae " vote
fi ee tice UPS: aie of ae
Se. SR AY aan a iin: 3 ety? Satis" fer
cave byt a Ee male pm
Bo hid racy re nao ac)
te Sita acim i bb ay
ON SOME SPIDERS FROM SOUTH ANNAM, 285
14. Some Spiders from South Annam.
By H. R. Hoae, M.A., F.Z.8.
[Received February 17, 1922: Read March 21, 1922. |
(Text-figures 1-10.)
The spiders described below were collected by Mr. C. Boden
Kloss in the course of a journey among the hills in 8. Annam,
in the spring of 1918. He has very kindly sent them to me for
identification.
Family AVICULARIIDA.
Genus MacrorHe.e Auss.
MACROTHELE MACULATA Thor., var. ANNAMENSIS, var. nov.
Dr. Thorell described as a new species, named as above, a non-
adult female from Java, placing it in C. Koch’s genus Diplura.
There is no doubt about it belonging to the present genus. Of
the two specimens here recorded, one is probably adult and the
other certainly not. They agree closely with Thorell’s description,
with the exception that his specimen had two rows of white spots
on the back on a black ground, while in these the spots are
wanting.
He further states that the eye area is 25 times as broad as
long. In these it is barely twice as broad, meaning that the
space between the front and rear rows has slightly widened. As
I believe this may be caused during growth to maturity, I have
merely deemed this a variety, due to the want of spots, instead
of, as it would otherwise have to be, a new species. There are
two females from Langbian Peaks, 5. Annam, 6500 ft.
Family ZODARID &.
Subfamily StoRENOMORPHIN2.
Genus STORENOMORPHA Sim.
STORENOMORPHA COMOTTOI Sim.
I have little doubt that a male and female from the same
locality as the foregoing belong to M. Simon’s above-named
species, which he records from Burma.
The minute third claw is little more than a plate with a small
point. It is close up to the scopula and very difficult to see at
all. All three claws spring from a hollow at the end of the
joint.
Proc. Zoou. Soc.—1922, No. XX. 20
286 MR. H. R. HOGG ON SOME
Subfamily ZoDARTIIN &.
LANGBIANA, gen. nov.
This genus differs from Diores Sim. in having the median eye
area as broad as or slightly longer than broad. The row of
spinous bristles on the tibia and distal joint of the female palp is
lacking. ‘The claw pectinated. There are short weak spines on
all joints of the legs except the tarsal.
LANGBIANA KLOSSI, sp. 0.
The cephalic part of the cephalothorax and mandibles are dark
red. The thoracic part, lip, maxille, sternum, legs, and palpi
bright orange. The abdomen above is black-brown, with two
longitudinal lines of yellow-grey spots, five in each, to near the
posterior end, where are a few transverse lines of similar colour,
On the under side it is yellow at the base as far as the genital
fovea, below which are alternate black and rather wide yellow-
grey longitudinal lines, some seven in number, reaching to the
spinnerets, the latter being yellow.
Text-figure 1.
Langbiana klossi, gen. et sp. n.
a, eyes; 6, lip and maxillz; c, epigyne: d, spinnerets from under side. In a the
front laterals should be lower down and rather nearer the median than drawn.
Both rows of eyes are procurved, the laterals of the posterior
row having their uppermost points on a line with the bottom of
the median, and the upper points of the front laterals in a line
with the centres of their median.
The eyes of the rear row and the laterals of the front row are
each about two-thirds the diameter of the front median. The
latter are half their diameter apart and twice that distance from
the rear median, which are their own diameter apart and twice
their diameter from their laterals.
SPIDERS FROM SOUTH ANNAM, 287
The front laterals are their diameter distant from their
median.
The clypeus is one and a half times longer than the median
eye-area and perpendicular.
The mandibles are conical, one and a half times as long as they
are broad at the base; the fangs short and stout.
The lip is longer than broad, rounded anteriorly, three-fourths
the height of the maxille. The latter bend over the lip and are
convex, not at all impressed.
The sternum is shield-shaped, truncate anteriorly, 14 times as
long as broad, rounded at the posterior end, but not protruding
between the rear cox, which are rather wide apart.
The abdomen is ovate, broader in front than at the rear,
rounded at the sides. At the posterior end on the under side is a
transverse row of short spines in front of the spinnerets. Of
these the inferior are cylindrical, raised on a membranous
base, with a short 2nd joint. They stand close together.
The superior are similar, but half as long and only half the
diameter, standing farther apart. The epigyne consists of two
oval depressions, their length apart, with another shallow hollow
above each.
The legs are long and fine, the joints cylindrical, with a few
seattered hairs along the outer sides, and short weak spines on
ll joints except the tarsal. Near the distal end of the metatarsal
is a thick bunch of spinous bristles. The tarsal joints of the 2nd
and 3rd pairs (the others broken) are three-fourths the length of
the metatarsal. The claws are two, with about six pectinations,
and spring from a short onychium.
The measurements in millimetres are as follows :—
Long. Broad.
as fi? im front.
Cephalothorax... 38 { 4
2
Abdomen......... 32 24
Mandibles ...... le
Trochanter. Patella Metatarsus
Coxa. & femur. & tibia. only.
Tees es ale Dar yl 24 32 Be ee JL SY
Metat. & tars.
9 3 QL at oars 3
ine 22 24 no:
‘ 3 91 93 Sn So
3 ZA “7 “4 OFT — 94
Metat. only.
1 93 L 43 Ban. 1 f
As iy dle 23 33 42 = 12242
, iL ll il pate 3
Palpi Rohe tieretelehey ater o 3 ] Fay il 8 il = 37
A single female from Dalat, Langbian Mountains, 8. Annam,
5000 ft.
20%
288 ' MR. H. R. HOGG ON SOME
Family THERIDIID.
ARGYROELOS, gen. nov.
Differs from Argyrodes Sim. in having the front median eyes
smaller than the laterals and the median eye-square narrower
anteriorly than at the rear. The maxille bending over the lip
nearly meet one another.
ARGYROELOS MICANS, sp. n.
The cephalothorax, mandibles, lip, maxille, and cox are all
bright orange. The sternum somewhat yellower, legs and palpi
paler orange, with yellow-grey hairs and brown bristles. The
abdomen is jet-black, with a mottled white streak across the
base, another across the middle, a third at the rear end with a
diamond-shaped spot between the latter two; the under side is
similarly black, with a short white transverse streak behind the
genital fold and a fainter white fillet at the sides. The epigyne
orange and spinnerets dull brown.
Text-figure 2.
Argyroelos micans, gen. et sp. n.
a, eyes; b, epigyne; c, lip and maxille; d, tarsus iv.
The cephalothorax is 14 millimetres longer than broad, rounded
and narrowed in front and to a less degree at the rear, rounded
at the sides. ‘he cephalic part is marked by depressions from
behind the side eyes tu a broad but shallow transverse depression,
from the ends of which two turn rearwards.
The rear row eyes are very slightly recurved, equal in diameter,
the median pair being 13 of their diameter apart and 12 of the
same from their laterals. The latter are contiguous with the
SPIDERS FROM SOUTH ANNAM., 289
front laterals, on a joint tubercle. The front median eyes are
distinctly smaller than the others, their diameter distant from
the rear median, slightly more from one another, and 1} diameter
from their side eyes.
The surface of the cephalothorax is smooth, with the exception
of very fine bristles on roots scattered at intervals on the cephalic
part.
The clypeus is convex, perpendicular, twice as deep as the
length of the median eye square.
The mandibles are straight along the outer edges, divergent on
the inner; on the inner margin of the falx-sheath at the far
corner is a triangular tooth, on the outer margin two smaller
near the middle. The fangs are long and slender.
The lip is about 23 times as broad -as long, curved in front,
straight at the sides, ‘and immovable. The maxille are nearly
three times as high as the lip, over which they bend, nearly
meeting one another, with straight edges on the upper part of
the inside, truncate anteriorly, and with a hollow about the
middle of the outer side.
The sternum is a long shield-shape, curved in front round the
bases of the lip and maxille, and passing in a narrow point
between the rear coxe.
The abdomen is oval, smooth, slightly protruding over the
cephalothorax. On the upper side it is sparsely furnished with
rather long, fine bristles, on the under side similar but shorter.
The epigyne consists of a lunate flat shelf protruding from a
kidney-shaped convex area, which fills the median part of the
former. ‘The spinnerets are not quite terminal, on chitinous
bases, conical, with a short 2nd joint.
The legs are moderately long and fine, without spines, the
metatarsal joints longer than the tibial. The two upper tarsal
claws are bent at the anterior end, with about five pectinations
on the basal half. The third claw smooth. The palpal claw is
straighter, with about five pectinations also.
The measurements in millimetres are as follows :—
Long. Broad.
do Bo
Cephalothorax... 43 { cs Tay ate
£
Abdomen......... 54 43
Mandibles ...... 1
Tib. Metat
Coxa. Tr. &fem. & pat. & tars.
WEBS) aces sack il. le 6 6 foi == 202
Dy 1 4s = ae eels,
3 2 a5 34 44 = 123
4, ls DS 54 6s = 182
AOI cas dae 4 le 1 ey on
290 MR. H. R. HOGG ON SOME
One female from Dalat, Langbian Mountains, 8. Annam,
5000 ft.
The cephalothorax, broader at the rear, with a shallower
thoracic fovea, front median eyes smaller than the others, and
maxille bending more over the lip, are sufficient to distinguish
this from Argyr odes, which it resembles in mandibles, lip, long
metatarsi, and other points.
Family ARGIOPIDS.
Genus NepuintaA Leach.
NEPHILA MACULATA.
Aranea maculata Fabr. Ent. Syst. ii. p. 425 (1793).
NV. maculata, forma principalis, Thorell, Rag. Austro-Mal. iii.
p. 145 (1881).
Two females from Phanrang Province, S. Annam.
Genus ArGiops Aud. in Sav.
ARGIOPE ZTHEREA Walck. Ins. Apt. ii. p. 112 (1841).
One non-adult female. Dran, Langbian Mountains, S. Annam,
3000 ft.
Genus GASTERACANTHA Sund.
GASTERACANTHA PREPEXTATA Walck.
Two females. Dran, Langbian Mountains, 8. Annam, 3000 ft.
Family THOMISID&.
DL@A SHIRLEYI, sp. n.
The cephalothorax is dark red, the mandibles, lip, and maxillee
rather paler, the former with nearly white fillets on the outer
margin of the falx-sheath, the sternum darker red again. The
femora and patelle of the front two pairs of legs are dark brown
tibia red at the base, dark at the distal end, metatarsus and
tarsus pale reddish brown, al) with short, fine brown hairs or
bristles. The coxee of the latter also darker red than the rear
two pairs. The other joints of these and the palpi are pale
yellow-brown.
The abdomen above is dark brown, smooth, and shiny, with
faint yellowish markings, one rather large spot at the base, a
pair of large round spots about the middle, with a pair of
smaller below them; nearer the posterior end a pair of strokes
sloping outwards from the centre line. Across the posterior
end are three transverse dark brown corrugations, slightly
paler along their anterior sides. Along the sides of the
abdomen are similar longitudinal corrugations, which on the
under side between the spinnerets and the genital groove sur
round, in a shield-shaped area, another transverse series. A pair
SPIDERS FROM SOUTH ANNAM. 291
of chitinous pale red-brown spots lie one each side above the
pulmonary openings, the spinnerets of a similar colour are
enclosed in a circular corrugation. The shape is about oval.
The cephalothorax is as broad as long, truncate in front, where
it is less than one-half its greatest width, rounded at the sides.
Its surface is roughly coriaceous, with a few bristles on roots
scattered over it.
‘Text-figure 3.
Diea shirleyi, sp.n., 3.
a, eyes; 6, male palp.
The mandibles are short, straight, and about twice as long as
broad.
The lip is as long as broad, rather broadly rounded in front,
half the length of the maxille. The sternum broad, heart-
shaped, passing between the rear coxee, which are wide apart.
The palp is of a normal type, an oval cushion with a flagellum
running round it.
The measurements in millimetres are as follows :—
Long. Broad.
y = in front.
Cephalothorax... 12 | Ac:
8
Abdomen......... 24 13
Mandibles ...... a
Pat. Metat.
Coxa. . Tr.&fem. &tib. & tars.
OSS scypatiesicr ile 4 24 22 72 eae
2. 4 22 3 2 ce
1 1 il a a
I 3 po 1
4. 2 li 12 a 47
PN Olle Sheri cee 2 3 4 mind a
292 MR. H. R. HOGG ON SOME
One male from Dalat, Langbian Mountains, $8. Annam, 5000 ft.
This in many respects resembles Thorell’s D. zonura, described
from three females collected in Java and Sumatra, and may
possibly be the male of them ; the legs are not quite in the same
proportion, neither does he mention the numerous corrugations
on the sides and underneath, nor the hard shiny back of the
abdomen.
Family CLUBIONID.
Genus PANnbDERcEYTES L. Koch.
PANDERCETES OCHREA, Sp. 0.
The cephalothorax and mandibles are bright orange-yellow,
with white and red hair on the former and white bristles on the
inner sides of the latter. The fangs are yellow-brown. On the
thoracic part of the cephalothorax are two transverse rows of pale
brown hair. The eye-space is almost wholly dark brown, but
interspersed with red and white hair.
Text-figure 4.
Pandercetes ochrea, sp.n. , eyes.
The lip, maxille, and sternum are bright pale yellow, the coxe
orange.
Tne legs are pale yellow, with smooth white hair and upstand-
ing brown bristles, very long brown spines, and grey scopule on
tarsi and metatarsi.
SPIDERS FROM SOUTH ANNAM, 293
The upper side of all the femoral joints are embellished with
round brown spots in more or less longitudinal rows.
The upper side of the pale yellow abdomen is thickly covered
with smooth white hair, the underlying surface being dotted like
the femora with numerous rather large round brown spots. The
underside is somewhat similar, but the hair-covering is shorter
and finer and there are no brown spots. The median area is
mottled with smaller white spots. The spinnerets are bright
yellow.
The cephalothorax and eyes are quite typical of IL. Koch’s
genus.
The lip, broader than long, is not quite half the length of the
maxille. The four teeth on the inner margin of the falx-sheath
are equal in size. Two on the outer margin are smaller, the
lower one smallest of all. Besides the tarsal scopula, a rather thin
one runs about half-way up the metatarsus of all legs.
Underneath the tibia and metatarsus of each leg are two pairs
of powerful spines 23 millimetres long besides others shorter.
The claws are stout and well curved, with four pectinations.
The abdomen is truncate in front and rather pointed pos-
teriorly.
The measurements in millimetres are as follows :—
Long. Broad.
Cephalothorax... 53 { 2 hin IER
Abdomen ......... 63 4
Mandibles ...... Hat
4 Pat. Metat.
Coxa. Tr. & fem. & tib. & tars.
Wess tia: 1, 2 163 12 95 = 34
2. or Il ls 10 2 = Se
Be 2a 9 92 9 2S
4, in DF 12 12 = dos
Palio} gare! il 3 3 3 = 10)
Tt will be noted that the second and fourth pairs of legs are
equal and longest, 64 times the length of the cephalothorax.
In P. gracilis L. K. the first pair of legs are longest, 7; times as
long as the cephalothorax, and there is a pattern on the abdomen.
In P. plumosus Dol. the legs are covered with thick bunches of
hair and first pair longest. In P. longipes Thor. the second pair
are 9 times as long as the cephalothorax. In P. isopus Thor. the
first, second, and fourth pairs of legs are equal in length, only
4 times as long as the cephalothorax, and coloration much darker,
especially the sides of the latter.
This must certainly be a new species, and J have named it
accordingly.
294. MR. HU. R. HOGG ON SOME
There is one not quite adult female and one smaller from Dran
in the Langbian Mountains of 8S. Annam, taken at an elevation
of 3000 ft.
Genus ADRASTIS Sim.
ADRASTIS LASHBROOKI, sp. 0.
Male. The cephalothorax is bright orange-yellow, with a
narrow brown stripe passing from the eye-space to the rear, from
which protrude forward at the rear of the cephalic part two
right-angled prongs, the whole forming a trident-shaped pattern.
Behind these are two similarly coloured forward curving lines.
The rear slope is brown.
The mandibles are orange. The lip, maxille, sternum, coxe,
and under side of abdomen paler yellow. The upper side of the
latter dark grey, with a round pale spot at the base. The legs
Text-figure 5.
Adrastis lashbrooki, sp. u., 6.
a, eyes; 6, male palp; ec, epigyne.
pale yellow, with brown spines and nearly white scopule. The
palp is dark grey on the back of the distal joint.
The Female is darker on the main part of the cephalothorax,
but a pale band runs round the margin, and the brown pattern-
markings are fainter. The abdomen on the back is dark grey,
with three pale spots at the base in a transverse line and a pale
grey transverse line at rear, also many grey hairs among the
darker. The under side of the abdomen is yellow-grey, with dark
grey hairs mixed with white; the scopula and claw-tufts a trifle
darker than in the males. The epigyne brown.
SPIDERS FROM SOUTH ANNAM. 295
The cephalothorax is half a millimetre longer than broad,
rounded at the sides, truncate in front, curving inwards at
the rear. The eye-space is raised up above the adjacent area
at the sides and in front, but at the rear is on a level with
the dorsal surface.
The rear row of eyes is recurved, the median sessile 1} diameters
apart. The laterals on prominences are 14 times as large and
14 times as far away as the median are from one another, The
front row are only slightly recurved, the laterals as large as the
rear laterals, The front median are rather smaller than the rear
median, 14 diameters apart, half that distance from the laterals,
and 2 diameters from the rear median. The clypeus is about the
depth of the diameter of the rear median eyes.
The mandibles are conical, slightly kneed at the base, with
three equal teeth on the inner falx-sheath margin and one quite
small below them.
The lip is convex, as broad as long, rounded at the sides, trun-
cate in front, less than half as long as the maxille, which are
widest in front and narrowed at the base to one-half the width
in front.
The sternum is as broad as long, rounded at the sides,
truncate in front, and thickly covered with upstanding bristly
hair.
The legs are moderately stout, with fine downlying hair, three
pairs of spines under each tibia, and three long ones on the outer
side of i. Scopule on all tarsi and on metatarsus of i. and il.
Two claws with about 7 or 8 pectinations. On the tibial and
distal joint of the palp are three spines each.
The epigyne consists of a pair of convex bean-shaped promi-
nences, placed longitudinally a short distance apart, narrowed at
the upper ends to one-half the width at the base. Above each of
them is an oval hollow. The whole lies on a semi-oval promi-
nence covered with short rough hair.
In the Male the rear row of eyes is less recurved and the
abdomen narrower in comparison with its length.
On the outer side of the tibial joint of the palp is a rather long,
hollowed, pointed apophysis, with three curved spines on the
inner side. A fine pointed flagellum passes about half-way round
the bulb, and a stouter doubly- -curved apophysis projects from
the opposite side.
The measurements in millimetres are as follows :—
Female. Male.
Long. Broad. Long. Broad.
24 in front. 12 in front.
Cephalothorax... 43 ee Par 32 i ata Gt
4 2
Abdomen ......... 6 4 5 24
Mandibles ...... 2 lt
i)
we)
(or)
MR. H. R. HOGG ON SOME
Female.
Pat. Metat.
Coxa. Tr. & fem. &tib. &tars.
ews ee recee ie 13 4} 5 LN eS
oN la 5 5 Aer ee le
Be ae) 4 4 32 = 123
3 12 44 4s Uys TY 105)
PAIL eessh Bareaeeee 4 2 2 DIO) a ieeet gO)
Mate.
essie. Se. if It 4 5h ave) ioe
2 1 AS 6 De ele
3. 11 32 44 a ke
A, 14 4 5 Of oe 1s
alot Ween ens. 5 ils les —— 5
Three males and one female from Langbian Peaks, 8. Annam,
6500 ft.
This species is distinguishable from A. atomaria Sim. by its
more uniform colour, freedom from spots on the cephalothorax
and legs, as well as the more even length of the latter, of which
the second pair is the longest.
Genus Patysres L. Koch.
PALYSTES LEDLEYI, sp. n.
Female. The cephalothorax is yellow-brown, with fine down-
lying pale yellow-brown hair scolloped against a paler area at
the rear; on the clypeus whitish yellow. The mandibles are
brown with brown hair. The fangs dark red-brown. The lip,
maxille, sternum, and cox pale yellow-brown. The remaining
joints of the legs and palpi are darker, the hairing ashy grey
under the femora and on the scopule, other wise yellow- brown.
The abdomen is pale yellow, with fine, short, smooth, yellow-
brown hair above and four muscle-spots rather darker in a
trapezium, narrowest in front.
The under side is similar in colour, with a dark grey triangular
pattern between the genital fold and the spinnerets, and three
longitudinal lines in the middle.
The cephalothorax is highest just in front of the rear slope
and declines to the eye-space; it is rounded at the sides, and
narrows in front to one-half its greatest width. The mandibles
are as long as the front is wide, perpendicular, convex. On the
inner margin of the falx- sheath are three lar ge teeth followed by
one smaller, and low down on the outer one large between two
smaller.
The rear row of eyes is slightly recurved, the median their
diameter apart, and 14 of same from the laterals, which are also
SPIDERS FROM SOUTH ANNAM. 297
14 times wider in diameter; the front row is straight, the
median rather smaller than the rear median, two-thirds of their
diameter apart, and one-fourth of the same from their laterals,
which are as large as the rear laterals. They are 14 their
diameter from the rear median. The whole eye-space and the
elypeus is furnished with long, upstanding brown bristles.
The lip is broader than long, rounded in front, and less than
one-half the length of the maxille, which are upright, convex,
truncate in front, and slightly hollowed on the outer margin.
The legs are moderately stout and long. The front femora
have i., i1., 1. spines on the upper side, none on the under. On the
fourth pair li.,1., 11, i, above. The patella are unarmed. Under
Text-figure 6.
Yy,
Yy,
YY,
My},
Y
i
Palystes ledleyi, sp. un., &.
a, eyes; b, male palp; c, epigyne.
the tibia of all legs three pairs of spines. On the front two pairs
the scopula reaches two-thirds the length of the metatarsus, on
the third and fourth pairs the whole distance. On the patella
joint of the palp is a pair of spines, one each side; on the tibial,
three long on the outer side, two on the inner, and on the distal
joint three long on each side. :
The epigyne consists of two kidney-shaped prominences, placed
diagonally in the sides of a circular hollow, and meeting over a
pair of small chitinous lobes at the lower end.
The Male is darker on the cephalothorax, the centre of the
thoracic part black-brown, with the lower edge scolloped, and a
nearly white fillet round the margin.
298 MR. H. R. HOGG ON SOME
On the tibial joint of the palpi is an apophysis, hollowed out
in the middle, with a blunt point on the outside corner and a
square-topped projection on the side nearest the joint. On the
opposite side of the same joint are three long spines springing
from near the base and reaching past the distal end.
The measurements in millimetr es are as follows :—
Female. Male.
Long. Broad. Long. Broad.
Cephalothorax ... 12 ' e vie GID 9 | ; 1
Abdomen ~. 2.221: 18 ON 143 93
Mandibles ...... i5) 4
Female.
Pat. Metat.
Coxa. Tr. & fem. & tib. & tars.
WES hee hen Ws 4 14 18 15 = il
Dy 4s 15 20 Ug == BSS
ae 4 13 143 5 —
A, 43 14 15) 16 = 49i
Ballou ect eae 2 54 5 Go = 134
Male.
eS She .2 ee i 33 Its 14 1 es aul
2 4 134 15 134 = 6
B} 34 10 114 NO) =. BS
4 4 11 124 2)
Palllot Miche see tere 2 4 4 oil!
There are two males and eight females from Dre an, in the Lang-
bian Province of 8 . Annam, from an altitude of °3000 ft., Sail
Mr. Kloss has also cone me one male and one female from ire ls
Lumpur, Malay Peninsula.
The measurements, and many other points, are very close to
those of Dr. Thorell’ s P. incanus from Borneo, but in that
species the first pair of legs is longest; the remainder are shorter
in proportion in this species ; the side- -eyes of the rear row
are as large as the side-eyes of the front row instead of having
the rear row all nearly equal, and the rear laterals are on lar ge
tubercles instead of being sessile.
Genus SERAMBA Thor,
SERAMBA PICTA? Thor.
One male, not fully adult. Dran, Langbian Mountains, 3000 ft.
SPIDERS FROM SOUTH ANNAM. 299
Genus Crmnus Walck. ,
CYENUS FLAVIDUS, sp. n.
The cephalothorax is bright orange-yellow, with dark grey
haivs at the sides and scattered white hairs along the margin.
The thoracic fovea is brown.
The eyes topaz-yellow, the rear median on black rims. he
mandibles, slightly darker, have long brown bristles and short
downlying white hair, chiefly near the base; fangs red, fringes
grey.
The lip, maxille, sternum, and coxe are all about the same,
pale yellow with brown bristles and scattered dark grey hairs,
rather lighter on the fringes of the maxille. The legs are orange-
Text-figure 7.
Ctenus flavidus, sp.n., 2. a, eyes; b, tarsus of iv leg.
Pardosa dranensis, sp.n., Y. c, eyes; d, epigyne.
coloured, with brown spines and grey tarsal claw-tufts. The
abdomen is pale yellow on the upper side, paler still on the
under, with short white hairs scattered about.
The cephalothorax is truncate and narrowed in front, gradually
widening at the sides of the cephalic part and much rounded at
the sides 6f the thoracic, hollowed at the rear. It is straight
along the median line, but slopes steeply to the side near the
margin. The mandibles are kneed at the base, the fangs long
and slightly curved. On the inner falx-margin are three large
teeth followed by one smaller, and on the outer three.
The front median eyes are two-thirds the diameter of the rear
median, and the small oval laterals of the front row have their
upper edges on a line with the upper edge of the latter. The
rear laterals are intermediate in size between the front and rear
median, and their lower edges are on a line half their diameter
300 MR. H. R. HOGG ON SOME
above that touching the tops of the rear median and front
laterals.
The lip is as broad as long, slightly hollowed in front, rounded
at the sides, and narrowed at the base. It hardly reaches to one-
half the height of the maxille. The latter are convex, upright,
rounded on the outer side, the anterior margin furnished with
thick heavy fringes.
The sternum is a broad shield-shape, convex, narrowed in front,
rounded at the sides, and, rather finely pointed at the rear, it
terminates in front of the rear coxe. The legs are moderately
stout, with five pairs of spines under tibia 1. and i. and three
pairs under metatarsus 1. and i.
The tarsi have thick claw-tufts, no scopule, but rather thick
bristles, along tarsus iv. The claws are well curved, stout, with
three teeth near the base.
The abdomen is ovate, broadest posteriorly, with a broad
pedicule, visible from above; the surface is smooth, with only
scattered hairs and bristles. The inferior spinnerets are short
and conical, broad at the base, with a hemispherical 2nd joint.
The superior are similar, but much smaller still, rather wide
apart, standing behind the outer margin of the inferior.
The measurements in millimetres are as follows :—
Long. Broad.
Cephalothorax ... As | ie iio roa
INibpdomiene eee re: D3 4
Mandibles ...... 12
Pat. Metat.
Coxa. Tr. & fem. & tib. & tars.
egy ejeceeoe Ie = + 4 See 2 ols
2: 5 At 4 Be 1G)
: Wa 3 3 3 = llQz
A, es 5 45 5 = ls
BATE eames 1 1s ly Anh Tiny a5
I have described this non-adult female from Dran, Langbian
Mountains as a new species only. The position of the lateral
eyes of the front row, as far to the rear as the tops of the rear
median, was L. Koch’s grounds for constituting his genus
Pycnoctenus and removing a single species from Walckenaer’s
genus of Ctenus. In the narrowness of the clypeus, however,
the above-described differs from both genera. The shape of the
cephalothorax and abdomen agrees with L. Koch’s genus, but I
am unable to distinguish any third claw, and in his the lip is
manifestly more than half as long as the maxille, while here it
is the opposite.
I therefore leave it as a Ctenus, in spite of the narrow clypeus
and the front laterals, because it follows that genus in too many
other points.
SPIDERS FROM SOUTH ANNAM. 301
Family AGALENID4#.
Genus AGALENA Walck.
AGALENA DORIS, Sp. n.
The cephalothorax dark brown at sides, with paler brown
median stripe reaching from the rear row of eyes to the rear
slope. ‘The hairy covering is plumose, smooth brown at the sides
and white on the median area. The mandibles, lip, aud maxille
are pale yellow-brown, with similar white hair to that on the
cephalothorax, the sternum darker, the hair longer and pointing
forwards, the coxe pale, with short plumose hair and longer
upstanding bristles. he legs and palpi pale brown, banded
with white hairs, but darker on the under side of thefemur. The
abdomen is yellowish grey all over, with scattered plumose white
hair. The eyes are topaz-yellow.
Text-figure 8.
1. Agalena doris, sp. n., d- ; €YeS
2. Agalena tenuis, Sp. 0.5 de
b, spinnerets of both species; ¢, lips and maxille of both species ;
d, male palp of A. tenuis; e, eyes of A. tenuis.
The cephalic part of the cephalothorax is straight and narrow,
truncate in front, the thoracic part sloping rather steeply there-
from to the side-margins. The rear row of eyes are equal and
equidistant their diameter apart, and so far procurved that a
line across the upper points of the laterals is wholly below the
lower edge of the median. The front row is shorter and like-
wise procurved, the eyes about the same size as the rear row,
being about two-thirds of their diameter apart, and only somuch |
procurved that the upper points of the laterals reach as far back
as the centres cf the median.
Proc. Zoo. Soc.—1922, No. XXI. 21
302 MR. H. R. HOGG ON SOME
The median quadrilateral is longer than broad and narrower
in front than at the rear. The clypeus is as deep as the median
eye-area, perpendicular, and slightly hollowed below the front
vow of eyes. The mandibles are conical, with three small teeth
on the inner margin of the falx-sheath.
The lip is straight in front, broader than long, with the maxilla
slightly inclined over it and barely twice as high as the lip.
The sternum is as broad as long, truncate in front, rounded at
the sides, the blunt rear end passing between the rear coxe.
The whole area 1s distinctly convex.
The joints of the legs are cylindrical and fine. There area few
short spines on the tibial and metatarsal joints, with a bunch at
the distal end of the latter. The superior tarsal claws have
about six pectinations at the basal end.
The abdomen is ovate, broadest anteriorly and truncate
posteriorly, and is furnished with a thin coating of downlying
plumose hair and scattered upright plain hair, The inferior
spinnerets are cylindrical, their diameter apart, covered both
with the plumose hair and longer fine bristles. A very short
hemispherical 2nd joint inside the circular termination of the
Ist. ‘The Ist joint of the superior is about the same length
and shape as that of the inferior, with a 2nd joint longer and
tapering to a fine point.
The measurements in millimetres are as follows :—
Long. Broad.
{ 14 in front.
Cephalothorax ... 23 5
Nbdomenpeneese 33 24
Mandibles ...... le
Pat. Metat.
Coxa. Tr. &fem. &tib. & tars.
eos aH ya: il 2 23 3 ye OS
2. 2 Din 3 on a ee
Bh 2 De 2 HE 2
4, 3 22 oe A — alt
MPA | cess ee = Ve 2 [ke eau oe
These are two males, neither quite adult, from the Langbian
Peaks, 8. Annan, 6500 ft. The genus seems to be very poorly
represented in any of the surrounding areas, and I have little
doubt that the species is hitherto undescribed, as also the
following from the same part.
AGALENA TENUIS, Sp. 0.
A non-adult female and fully adult male from the same locality,
while resembling the above in the more important respects,
can readily be distinguished therefrom.
SPIDERS FROM SOUTH ANNAM. 303
The colouring of the cephalothorax and abdomen both above
and below is a uniform deep black-brown, with the exception of
a lozenge-shaped wliutish-haired patch round the thoracic fovea
of the former. The eyes are black.
The mandibles, lip, maxille, and sternum are rather dark
yellow-brown.
The legs are brown, with the exception of the tarsal joints,
which are yellow.
The shape of the cephalothorax, eyes, mandibles, lip, maxillee,
and sternum are as in the above-described, but the legs are finer
and more heavily bespined—a single spine on the inner side of
each femur, two single on the upper side of femur 1 and 2.
The spines on the tibia and metatarsus are longer and stouter,
and the tarsal joint of the third and fourth pair of legs is more
than half as long as the metatarsus of the same. The abdomen
is narrower in proportion to its length. The sternum projects
rather farther between the rear coxe, and the whole specimen
is distinctly smaller and more lightly formed.
The measurements in millimetres are as follows :—
Long. Broad.
21 5 ‘
2 in front.
‘Cephalothorax . . 23 J “
ple
/NJOGICASIN, Gaonbeooe Mee 1}
: Pat. Metat.
Coxa. Tr.& fem. & tib. & tars.
ITER ae cael il, 3 2 25 35 = | 82
em is Voce a7.
a 2 2? ae Aaa
Me cites 23 23 Sp ss ls
Family Lycosi1p.
Genus Parposa C. Koch.
PARDOSA DRANESSIS, Sp. n. (See text-fig. 7, p. 299.)
The cephalothorax has a pale yellow-brown median streak
reaching from the median eyes to the top of the rear slope; the
sides darker yellow-brown; scanty downlying grey hair over
the whole, thicker round the marginal streak. The eyes are
yellow.
The mandibles are rather dark brown; fangs red; the lip,
maxilla, sternum, and coxe yellow; the legs yellow, with brown
rings.
The abdomen above is yellow-brown, mottled all over with
small round yellow spots; scattered grey hairs smooth and down-
lying, thicker on the sides. On the under side it is paler yellow-
brown, uniformly covered with fine, smooth grey hair. The
epigyne is yellow.
Ail
304 MR. H. R. HOGG ON SOME
The cephalothorax is narrow in front, akout one-third of its
greatest width. The mandibles are longer than the clypeus.
is wide; fangs short and stout; two equal-sized teeth and one
very minute on inner falx-sheath.
The eyes of the front row are procurved, the laterals two-thirds
the diameter of the median and that distance apart. The eyes.
of the second row have a diameter three times that of the front
median, and stand out rather prominently on black rims; they
are as far apart as the diameter inside the black rims. The
rear eyes are two-thirds in diameter those of the second row,
two of their diameters apart and the same distance from the
second row.
The lip is much brcader than long, about one-third the height
of the maxille.
The sternum is a broad oval, smooth and slightly convex.
The abdomen is oval; the upper surface smooth, with only a
few scattered fine hairs, but they cover entirely the under side.
The epigyne consists of an oval transverse base, from whieh
springs the median rib. This dividing at the apex encircles.
hollows—one on each side—and returns to the middle in a small
shiny protuberance. Upright oval bosses stand one at each side
of the base. ~
The measurements in millimetres are as follows :—
Long. Wide.
a2
3 in front.
Cephalothorax ... 3 f a
a4
Abdomen ......... 3t 24
Mandibles ...... it
Pat. Metat.
Coxa. Tr. &fem. & tib. & tars.
Weesen eas Te 3 21 21 ee ok
9) 3 = ne sae Lae
ery 4 —
‘ 3 1 1 3 =o all
3, q 24 24 oF = 94
3 3 Ab) eS 3
4. 1 23 28 45 = 103
Pal pl GeJoordodoud = ] t I 2 = 34
A single female from Dran, Langbian Mcuntains, 8S. Annam,
3000 ft., is very similar in appearance to the Australian Lycosa
albosparsa L. Koch, but has the front row of eyes shorter than
the second instead of longer as well as the slight generic
differences.
Family PIsAURID2.
Genus Hyeroropa Thor.
Hycroropa tonairarsis Thor.
Three females from Deban, Pharang Province, South Annam,
650 ft.
SPIDERS FROM SOUTH ANNAM. 305
The measurements in millimetres are as follows :—
Long. nee
Cephalothorax ... 34
D Bes
ANodomens........ 5 24
Mandibles ...... I
Pat. Metat.
Coxa. Tr. & fem. & tib. & tars.
1 bracers Il. ] 6 a 9l = 231
My il 6 6 ee = UE
as 2 3 3 B29 es Oe
q la 5 D 7 = lies
sahaS 1 l 1 Te tee
EEO eit a Re z le 1+ We = 4S
With the exception of being smaller, this agrees closely with
Thorell’s description of his ie (Dendrolycosa) longitarsis from
Kandari, Celebes Is.
Family SALTICID.
Genus MErNrEMERUS Sim.
MENEMERUS FELIX, sp. n.
The ground-colour of the cephalothorax is black-brown round
the edges of the cephalic part, dark yellow-brown in the middle
and on the thoracic part, with white and orange hairs in front
between the eyes, and white hair-patches on the median streak.
At the sides are broad black streaks, with a white hair-fillet
round the sides; a white hair-fillet surrounds the whole upper
side of the abdomen, with a longitudinal median streak of white
hair intermixed with orange in the posterior half ; between this
and the side fillet is a black area on each side. A white hair-
spot lies on this on each side near the posterior end. ‘The under
side of the abdomen is greyish yellow in front of the genital
groove, from which a darker longitudinal stripe runs to the
spinnerets, each side of which are white and orange hair mark-
ings covered with rather large irregular brown ‘patches. The
mandibles are black-brown, with scattered white hairs on the
inner side. Fangs reddish.
The lip, maxille, and sternum are bright yellow, with short.
brown hair on the former and long upstanding white hair on the
latter.
The palpi are bright yellow, with short brown and long white
hair.
The coxe of the legs are bright yellow, but the femoral, patellar,
tibial, and metatarsal joints are brown, with white and orange
hairing and long brown bristles; the tarsal joints yellow, with
grey claw-tufts.
306 MR. H. R. HOGG ON SOME
There are three pairs of short spines under tibia i. and ii., two:
pairs under metatarsi i. and ii.; on metatarsus iv. are spines near
the base as well as the bunch ah the distal end.
The measurements in millimetres are as follows :—
Long. Broad.
ie ee
Cephalothorax ... 33 { ak iv ior
No donveneenn ate 33 2
Pat. Metat.
Coxa. Tr. & fem. & tib. & tars.
a a 93 23 valle tet in
Legs weet ene 1 Ie “aq O44 “4 = 93
Dig si 22 3i 2 = BR
3. ls 2% 24 2 eos
1 93 : =
TS tie 23 3 3 = 10
There are a male and two females from Dran, Langbian
Mountains, Province of 8. Annam, taken at an altitude of
3000 ft. None of them quite adult.
Text-figure 9.
Menemerus felix, sp. n., 6.
a, Menemerus crassus, sp. n., 9 ; b, Menemerus felix, profile; c, eyes from
in front, g ; d, eyes from above, 2 ; e, epigyne of M. crassus.
These specimens certainly appertain to the above genus, and
are very close to M. Lucas’s VW. (Attus) melanognathus, a very
widely-spread species which I have not seen; but judging from
the list of allied species given by Dr. Thorell (‘ Ragni Indo-
Malesi,’ p. 307), it would seem +0 differ only in the pattern of
the abdomen, generally lighter colouring, and longer legs.
SPIDERS FROM SOUTH ANNAM. 307
MENEMERUS CRASSUS, Sp. n.
Female. Cephalothorax black-brown; a rather large yellow-
brown spot surrounds the fovea just behind the line of the rear
eyes, and is continued as a narrow streak to the rear slope. Down-
lying lanceolate red and white hairs cover the upper part of the
cephalothorax; a white fillet round the margin and only red
hairs round the front eyes and on the clypeus; above the front
row of eyes is a row of long brown bristles, and a thick bush of
long yellow-grey bristles protrudes from the lower part of the
clypeus and base of the mandibles, the latter being dark red-
brown. The lip and maxille are yellow-brown with yellowish
grey fringes. The sternum is brighter, with long upstanding
bristles round the margin. It is twice as long as broad, and
does not extend between the rear coxe.
The legs are bright yellow-brown on the under side, nearly
black-brown on the upper, with the exception of the tarsi which
are yellow-brown. The claw-tufts are grey. Both they and the
palpi are closely covered with short red and white lanceolate
hairs, and on the tibial and distal joints of the latter are long
outstanding yeilow-grey bristles.
The abdomen above is pale yellow-brown, with similar hair to
that on the cephalothoiax but white only round the base, where
there is also a band of long white bristles.
The under side is similarly coloured, but the hair-covering is
yellowish grey intermixed with brown bristles on roots. The
epigyne is brown, and the spinnerets on the upper side are
covered with red and white hair.
The measurements in millimetres are as follows :—
Long. Broad.
alee
Cephalothorax... 5 [ar oe ont:
| 33
Abdomen ......... 7 4i
Mandible ......... ig
Pat. Metat.
Coxa. Tr. & fem. & tib. & tars.
eS ee ss sic: ike 13 34 + 2 i=) elle
2. ile 33 3t ee — atl)
Je 12 3 3f Seer =e Oe
4, is 32 + Ape ot
BEY Adie deus hong -cldentieadd dh a 2 = 3
There is one female from the same locality as M. felix, but it
is a good deal larger, the abdomen is more elliptical, and the
long bunches of bristles on the palpi and fore part of the cephalo-
thorax form a special feature.
MENEMERUS DABANIS, Sp. 0.
Male. Cephalothorax black-brown in front and at the sides of
the cephalic part, red-brown in the middle and on the thoracic.
308 MR. H. R. HOGG ON SOME
A fillet of white hair round the margin: a white hair-patch lies
round the thoracic fovea. Long white bristles behind the front
eyes and on the clypeus. Red hair round the front eyes.
The mandibles are black-brown, with long white bristles at
the base; fangs paler red. The maxille are dark yellow-brown,
the lip darker brown with dark grey hair. The sternum is dark
yellow-brown with white hair.
The abdomen is dark yellow-brown above, with a bread median
streak, narrowing posteriorly, of white hair on pale yellow-brown
and iene upstanding white bristles. On the under side it is white
at the sides, with a yellow-brown median area reaching from the
base to the spinnerets. The latter are dark brown, w ith a white
hair-streak on the upper side of the superior pair.
Text-figure 10.
Menemerus dabanis, sp. n.
a, eyes; b, profile; ec, under side of abdomen; d, left palp.
The palps are dark brown on the femur, paler yellow-brown
on the other joints, with long upstanding white bristles.
The first and second pairs of legs are dark brown from femur
to metatarsus, with long white bristly hair mixed with black
forming a fringe on the under side of the femur, tibia, and
metatarsus. The tarsal joint is paler yellow-brown with a dark
grey claw-tuft.
The third and fourth pairs of legs are pale yellow-brown on the
femoral joints, darker on the tibial and metatarsal, with long
white bristles and downlying white hair.
The cephalothorax is flat on the cephalic part, highest at the
rear, and sloping anteriorly. The thoracic part slopes at first
slightly and then steeply to the rear, with a short fovea behind
SPIDERS FROM SOUTH ANNAM. 309
the rear rows of eyes. The front median eyes are twice the
diameter of the front side, which are well separated from the
former, situated farther back and higher up, making the whole
row recurved. All these eyes have wide black rims. The eyes
of the third row are of the same diameter as the front laterals,
and the row is only very slightly broader than the front. The
small eyes of the second row are about equally distant between
the front laterals and the rear eyes.
The clypeus is half the width of the front median eyes, and
wholly covered with long bristly hair pointing inwards from
each side.
The mandibles have long bristles at the base, with long but
fine hair over the remainder. The single tooth on the inner
side is conical and about half-way down the margin of the falx-
sheath. On the outer margin are two smaller teeth at the
lower end.
The sternum is pear-shaped, narrowed anteriorly, the front
lying between the coxee of the first pair of legs, which are the
width of the base of the lip apart.
The abdomen is twice as long as broad, widest near the base
and narrowing to the posterior end. The downlying hair at the
sides of this as well as on the legs and the cephalothorax is
lanceolate.
The front pair of legs is longer and stouter than the others,
the femur being club-shaped. ‘The second pair similarly shaped
but smaller. On the patella and femur of each of these pairs is
a long thick fringe of bristles—under tibia i. and ii. are three
pairs of spines; under the metatarsus of i. and 11. are two pairs
of stout spines and one smaller spine; under metatarsus ill.
and iv. are two pairs of spines, one pair at the sides and a
bunch at the distal end.
The measurements in millimetres are as follows :—
Long. Broad.
95 4 D
Cephalothorax ... 32 i 31 tpazOne:
Abdomen ......... 5 24
Mandibles ...... 12
Pat. Metat.
Coxa. Tr.&fem. & tib. & tars.
MEGS nares. Ie la 4 43 ee
2. IE 3 3 22 =e O86
3. = 22 22 Fa Liesl
4, 1} 34 32 3 eee AOE
Ballplenieasctts< dec 3 le il Weg ie Ae
One male from Daban, Pharang Province, 8. Annam, from an
altitude of about 650 ft.
310 MR. H. R. HOGG ON SOME
4
TACHYSKARTHMOS, gen. nov.
Differs from Varne EK. Simon, of the group Hasariex, in
having flat lanceolate bristles and sumple hairs in place of
shining squamules, in having the rear pair of eyes dis-
tinctly. large instead of rather small, in having the bunches
of spines on the four rear metatarsi larger and more numerous
and the third pair of legs shorter and not thicker than the
fourth pair.
TACHYSKARTHMOS ANNAMENSIS, Sp. n.
Male. The cephalothorax is black-brown, sparsely covered with
white and orange lanceolate hair, and brown bristles behind the
front row of eyes; the mandibles, lip,and maxille yellow-brown 3.
sternum and coxe brighter yellow-brown, with pale brown up-
standing hair. The legs darker yellow-brown on the femoral,
patellar, and tibial joints, paler on the metatarsal and tarsal,
sparsely furnished with fine upstanding brown bristly hair and
grey claw-tufts. The palpi yellow- brown on the femoral and
under side of the patella and tibial, but on the upper side of these:
a thick mat of white lanceolate hairs ; the distal joint yellow-
brown.
The abdomen above is mottled with small yellow and brown
spots, with a patch of brown bristly hair at the base; behind this.
a horseshoe-shaped fillet of white lanceolate hair reaching half-
way down each side of the abdomen. On the lower half ‘of the
back is another smaller similar fillet of white hair. The under-
side is smooth yellow-grey with minute brown simple hairs;
spinnerets brown.
The cephalic part of the cephalothorax slopes downward from
the rear row of eyes to the front and from about the same point
to the rear, the thoracic part being in length as eight to five
of the cephalic. The surface is smooth and shiny. The eye
quadrangle is one-third wider than long. The eyes of the rear
row are very convex and as large as the front laterals, shghtly
narrower than the front row. The small second row eyes are
situated midway between the rear and front laterals. The eyes
of the front row are slightly recurved, the laterals half the
diameter of the median, and the clypeus half the width of the
latter. A short thoracic fovea lies just behind the rear row of
eyes.
The mandibles are conical, straight, and perpendicular, with
rather long fangs.
On the inner margin of the falx-sheath is a bifurcated tooth,
with two single ones on the outer margin.
The lip is longer than broad, rounded anteriorly, and more
than half the lent eth of the mioecilles which are upright, straight.
at the anterior end, rounded on the outside of their upper end,
and tapering to the base.
SPIDERS FROM SOUTH ANNAM. 31}
The sternum is ovate, widest in the middle, longer than broad,
tapering to rear and front, where it is truncate the width of the
ase of the lip.
The coxe on each side are contiguous, as are also the rear
pair.
The tibial joint of the palpi is longer than the patella. There
is no apophysis at the base of the distal joint, which holds a
simple oval convex bulb with only a short flagellum.
The legs are moderately long, the femoral joint of the first
pair thicker and more club-shaped than the others.
There are three pairs of spines under tibia i., one long pair at
the base of metatarsus i., and a single spine on the inner side
near the distal end.
The two tarsal claws have a few teeth at the basal end, covered
by claw-tufts of thick flat bristles. On metatarsi ii. and iv.
are one single and two pairs of Jong spines with a bunch at the
distal end. A pair of spines on the patella of the same, none on
the femur.
The abdomen is ovate, rounded in front, rather pointed at the
rear. The texture is soft, with flat lanceolate bristles on the
upper side and fine short simple hairs on the under side and a
bunch of bristles at the base.
The measurements in millimetres are as follows :—
Long. Broad.
2 in front.
Cephalothorax ... ot { .
‘ 91
“95
FAocloment see ena. 3 OF
Mandibles ...... i
Pat. Metat.
Coxa. Tr.&fem. & tib. & tars.
Wecsi te a i ae an mC Rp he Fhe Ry,
2. 1 : eee hr ae
Brady ind 22 12 Mua 62
Be om <3 oe ee. ee ee
alpine een ne = 1 1 pe beet 833
A single male from Dran, Langbian Mountains, 8. Annam,
3000 ft., would appear to come nearest to M. Simon’s African
genus of Larne, but it is clearly necessary to place it in a new
genus, on account of the difference given above.
I would like to record an observation made by an esteemed
correspondent, Mr. H. C. Abraham, an officer of the Govern-
ment Survey Department at Taiping, F.M.S.
Mr. Abraham, who is an enthusiastic arachnologist, was
watching a female of the Giant Orb Weaver, Vephila maculata,
on her web, running about on which were three of the minute
Slee ON SOME SPIDERS FROM SOUTH ANNAM.
males. While he was watching, one of the males ran up to her,
clasped her, and served her, followed at short intervals by each
of the others.
The female and her three suitors are now in Mi. Abraham’s
collection.
It might well be that, owing to the great difference in size
between the sexes in this genus, such a procedure is necessary in
order to fully fertilize her eggs, but I do not think it has been
previously noted that to do so the services of more than one
of the other sex has been required.
ON NEW REPTILES FROM tANGANYIKA TERRITORY. 313
15. New Reptiles from Tanganyika Territory.
By Artuur LoveripcE, C.M.Z.S.
[Received March 24, 1922: Read April 25, 1922.
The opisthoglyphous snake and lacertid lizard here described
occurred in collections of reptiles made in East Africa during
1920.
Tn naming this new snake after Miss J. Procter, #.Z.8., I wish
to testify to her kindness in examining the dentition of this and
other specimens and in reading over the proofs. My thanks are
also due to Mr. G. A. Boulenger, F.R.S., for making a correction
in the description of the new lizard.
GEODIPSAS PROCTER, Sp. Nn.
Tf included in this genus (Blgr. Cat. Snakes, ili. p. 32), to which
it is closely allied, the generic description will have to be
enlarged to include snakes with single as well as double sub-
caudals.
Dentition.—16 or 17 maxillary teeth (including the fangs).
Solid teeth followed after a slight interspace by two enlarged
grooved fangs. Mandibular teeth subequal.
Description.—Rostval broader than deep, just visible from
above ; internasals as long as broad, shorter than preefrontals ;
frontal once and a half as long as broad, shorter than its
distance from end of snout, shorter than the parietals; loreal as
long as deep; three pre- and two postoculars; temporals one
plus two; eight upper labials, fourth and fifth entering the eye ;
four lower labials in contact with the anterior chin-shields,
which are a little shorter than the posterior. Scales in seventeen
rows. Ventrals 148; anal single; subcaudals 49, single.
Ooloration.—Purplish iridescent black above, slightly paler
beneath. Throat white. Head brownish ; labials white, with a
brownish mark on the posterior border of each; an ill-defined
black stripe from posterior border of eye to commissure of mouth,
erossing 7th and 8th labials. A distinct black dorsal stripe on
median row of scales from frontal to end of body; indistinct
on tail.
Sex ¢
Length of head and body 310mm.
<3 bog CEO tema,
Afinities—Very similar in all respects to Geodipsas vauero-
cege Tornier, 1902, from which it differs in having 3 instead
of 2 preoculars, smaller parietal shields, and single subcaudals.
Locality.—On path in thick bush-country, Uluguru Mts.,
3 miles from Morogoro, T.T., at an altitude of about 3000 ft. ;
Loruttis IEG
314 MR. A. LOVERIDGE ON NEW
Type-specimen given to the British Museum. Only the type
was collected, and though I sent a native collector to the
same locality for a month for the express purpose of obtaining
more specimens, he failed to get any.
NUCRAS KILOS&, sp. n.
Some lizards collected at Tindiga, Kilosa District, T. T., belong
to an undeseribed species. ‘The following description is based on
a series of seven individuals, six of which were males. These
have been indicated by letters @ to /, males; g, female :—
Description.—Head moderate; snout obtuse; nostril pierced
between three shields; nasals not or but slightly swollen, lower
separated from the rostral; two prefrontals ; frontal feebly (a, d)
or strongly (0, c) grooved ; four to six supraoculars (four in a, @;
five in 6; six inc), the two middle ones being much the largest
in a, the anterior two largest in 6, ¢; interparietal at least twice
as long as broad, longer than the suture between the fronto-
parietals; an enlarged band-like temporal shield and a curved
tympanic ; temporal scales granular, smooth, no auricular denticu-
lation ; lower eyelid scaly, subocular narrower below. than above,
bordering the lip between the fourth and fifth upper labials ; two
or three anterior pairs of chin-shields in contact. Gular seales
gradually increasing in size towards the collar; latter slightly
curved, perfectly free, formed of nine (rarely seven as in @) plates.
Dorsal scales rhomboidal, juxtaposed, keeled, 42 to 56 across the
middle of the body. Ventral plates in 6 or 8 longitudinal, and
97 to 31 transverse series. ‘The hind-limb reaches the wrist
(b, c,d), elbow (e), or axilla (@) in the male, and the elbow in the
female type. Foot as long as, or a little longer than, the head.
A single (occasionally double) series of large subtibial plates.
Eleven to thirteen femoral pores(11 in ga, Qg;12 in 3 3 b,d;
13 inc). Tail nearly (4, e) or more than twice (6) the length of
head and body in male, much more than twice the length in
female type; caudal scales strongly keeled except basal sub-
caudals, which are obtusely keeled.
Coloration in life— 3. Upper surface of head, fore-limbs, and
body nearly to hind-limbs brown. Head, limbs, and upper sur-
face of tail a bright yet translucent red, which entirely disappears
on preservation. A narrow, yellow, vertebra! line commences at
interparietal ; it lies between a pair of dorso-lateral lines which
start from the band-like temporal scale; these three lines all
vanish on the base of tail; the space between these dorso-laterals
and the vertebral line is dotted and marbled with black spots,
which are sometimes almost absent in parts and in others
confluent in black patches, or form a black border to the vertebral
line. A single (sometimes double) series of yellow dots and
dashes on flanks; below these a white or yellowish line com-
mencing at commissure of mouth: passes through the ear-openin g
and may be continuous to hind-limb, but is more frequently
REPLILES FROM TANGANYIKA TERRITORY. 315
broken up into a seriesof dashes. In the type a the whole of
the upper surface is spotted with yellow and white (probably
breeding marks) ; this isabsent in all the others. Under surfaces
china-white except distal portion of tail, which is rufous.
2. Upper surfaces all brown and much darker than male ;
only one series of dots on flank. Lower surface all white; no
rufous colour at all. Measurements :
Males (a). Head and body 65mm. ‘Tail127mm. British Museum.
(b). = > 62mm. » 128mm. Loveridge Coll.
(e). 5 >» 66mm. > impaired. ‘s 5)
(d). 3 » 65mm. 3 a A.M.N.H.
(e). e ; 60mm. , 116mm. Skull, Loveridge Coll.
(f). 5 » 65mm. », impaired. Dr. Barbour Coll.
Female (g). _,, By YAMIN, » 127mm. British Museum.
Affinities.—Closely allied to WV. emini Blgr., from which it
differs in its keeled scales and different markings.
Locality —Running on paths at Tindiga, 5 miles south of
Kilosa, Morogoro District, T. IT. Has since been found at Kilosa
also.
Type-specimens.— 3 a, 9g, donated to the British Museum.
Dates.—(a, g) 19. iv. 21; (f) 20. vi. 21; (6, 6, d,e) 19. xu. 21.
Notes.—Not uncommon but active, running on paths in hot
‘sunshine.
Female retired into hole, and on being captured was found to
have four eggs in ovary measuring 5 mm. in diameter. Stomach
contents included a parasitic nematode, a field cricket well
chewed but swallowed whole, limb of Hymenopterous insect, and
the lizard’s own sloughed ventral scutes.
LP agi feces vitae bien: sn
Ra navel
eas
P, Z. S. 1922, BEAINE, Elods
232°
a
a
Fi
HIPPOTRAGUS VARIANI, al
770
ye een
EAR aN
(iyi
(P YASIIN SNOVALOddIH
limtdSaNivVad ece6l=S Zed
Re ee eee ee A, ee ae
i ‘SANINOA SNOVYLOddIH
Uitte aaNet cdc bleeS=Z-id
IV.
Pl.
1922, BLAINE,
Ln See
B. H.VARIANI. Cy He BQUINUS:?
A. HIPPOTRAGUS NIGER.
Py Zo SS, IO22, IBE/ANUINVE, ell. W.
A. Angolan Sable Antelope. ¢ B. Angolan Springbuck. ? 3¢
Pa ZS 1922) BEATINIE: Sela
A. Old Stallion Hartmann’s Zebra. B. Dorsal view.
RP. Z-'S. 1922, BLAINE, PIowile
A. ANTIDORCAS ANGOLENSIS, sp.no. Bee EUCHORE.d.
ai! basioccipital. b basioccipital.
PZ Sel Z a SAIN Eee ante
A. ANTIDORCAS ANGOLENSIS, sp.n.Q. B. A. BNIB Gs
ON THE ZEBRAS AND SOME ANTELOPES OF ANGOLA, aN
16. Notes on the Zebras and some Antelopes of Angola.
By Ginpert Buarne, F.Z.S.
[Received February 7, 1922: Read March 21, 1922.
(Plates I.—VIII.*)
Part I.
In the summer of 1918,Mr. P. Van der Byl and I, who had
both for a long time been contemplating a trip to Angola to ob-
tain specimens of the Great Sable Antelope, on the subject of
which letters had from time to time been appearing in the ‘ Field’
from the pen of Capt. H. F. Varian, were able to fulfil our
intention.
We set out under very favourable auspices with the kind
assistance of the British Museum (Nat. Hist.), and of Mr.
Varian in Angola, to whom are due our grateful thanks for
the attention he gave us and the unremitting care he took to
insure the success of our enterprise. Unfortunately, Mr. Van
der Byl, after spending six weeks up country, contracted malaria,
which left him in such a condition that he found it necessary to
return to the Kapango Mission Station in the Bihe District,
whence, after a few weeks’ convalescence, he reached the coast and
went home.
Mr. Oldfield\Thomas (P. Z.S. 1916, p. 298) had already
described the Angolan Sable Antelope as a new subspecies
from material provided by Capt. Varian, taking as his type a
face-skin and a frontlet with horns. He was able to obtain the
loan of a single skull for comparison, but owing to insufficient
evidence did not feel justified at the time in making a new species.
I was able to bring home complete specimens of each sex, including
some sub-adult examples, and with this material to work upon, I
feel less diffident in establishing its claim to specitic rank, and
T have the concurrence of Mr. Thomas in so doing.
There are certain remarkable modifications to be found in the
skull, and these, coupled with the large size and immense horns
of the males, the entirely black face, and the distinctive body-
colour of the females, combine to make a very strong case.
The Angolan Sable also, although as far as present knowledge
goes confined within narrow limits, appears to be completely and
widely isolated by natural boundaries from true Sable.
The name variant that Mr. Thomas has chosen fits it well, for
it is through the active interest of the distinguished Engineer of
the Benguela Railway that this splendid Antelope, so far the finest
in all Africa, has been brought to light.
* For explanation of the Plates, see p. 339.
Proc. Zoou. Soc.—1922, No. XXII. 22
318
MR. G. BLAINE ON THE ZEBRAS
In the following table the differences between the skulls of
THippotragus niger and Hippotragus variani are set forth :—
A. niger.
Face shorter and broader, the frontal
field which gives rise to the horns
being more prominently developed,
with the bases of the horns directed
more obliquely backwards.
Nasals shorter, their frontal extremities
not reaching the plane of the ante-
orbital processes.
Premaxille less spatulate.
Lacrymal depression scarcely defined.
Orbits round.
Maxille showing a distinct groove
running parallel to the inferior border
of the nasals,and continued across the
upper ends of the premaxille.
Tympanic bulle small, with styliform
process flattened and blade-like.
Paroccipital processes wide, flattened,with
posterior margins convex. Viewed
from behind they curve slightly out-
wards, then downwards and inwards
at their apices, a large portion of their
posterior borders being on a level with
the posterior face of the condyles.
Foraimen ovale large and oval.
Mesopterygoid fossa wide, triangular in
shape, with its containing walls
diverging from the basisphenoid down-
wards, the post-hamular margins
forming everted flanges.
Lateral fossee wide.
Ascending wall of maxilla behind the
tooth-row nearly vertical, and strongly
carinated.
Posterior palatine notches in line with,
or in front of the plane of the posterior
edges of the last molars.
Teeth smaller.
H, variant.
Face longer and narrower, with less ex-
tensive frontal field and the bases of
the horns rising nearly vertically and
directly over the orbits.
Nasals longer, their frontal extremities
reaching the plane of the ante-orbital
processes,
Premaxille spatulate, with
flanged anterior margins.
Lacrymal depression clearly defined.
Orbits oval, the vertical axis being longest.
Maxillz showing only faint traces of
groove.
broadly
Tympanic bulla large, with styliform
process round, very small and spike-
like.
Paroccipital processes narrow, with
straight posterior margins and no
lateral curvature, but having a strong
forward and shght inward inclination.
They are more forward in position,
being almost wholly in advance of the
condyles.
Foramen ovale small and round.
Mesopterygoid fossa narrow, with
parallel walls.
Lateral fossee narrower, being constricted
by the inflation of the posterior ascend-
ing walls of the maxille.
Ascending wall of maxilla behind the
tooth-row obliquely directed with its
suface inflated, and less carmated.
Posterior palatine notches behind the
plane of the posterior edges of the
last molars.
Teeth larger.
The skull of H. equinus resembles that of HW. niger, and differs
from H. variant in the form and position of its paroccipital pro-
cesses, in the open triangular outline of the mesopterygoid fossa,
and in the vertical direction of the ascending walls of the maxille,
which are laterally compressed and prominently carinated.
AND SOME ANTELOPES OF ANGOLA. 319
Comparative measurements in millimetres of skulls of the
three species taken from the B.M. Coll. (N.H.).
niger. variant. equinus.
Greatest’ length..:.................. 4388 (= 17% 475 (= 182 470 (= 183
inches). inches). inches).
5 GareEAGI EIN Waemasece neon ade 162 162 179
Sumnut of orbits to base of
NONI AGNI LS 5 cAsasnvenmesssonse | Coll 4d 43
Tengthy of masals.................. 164 203 178
a <9 WORDT ED cA Geoonaond LUZA0) 164 132
Width across maxillary flanges. 60 71 65
>» Of mesopterygoid fossa. 24 16 29
Length from anterior angle of
mesopterygoid fossa to end of
premaxille .. Hee ce recht (OO) 275 257
Length of upper hove TOW Eenoce 112 122 132
In comparing skins of ziger with those of variant, the most
striking differences are found in the females, for whereas in the
males of the latter a greater intensification of colour is shown, in
the females the colour is quite distinct.
In true Sable the prevailing colour of the females is a sombre
brown, very old examples becoming almost as black as the males,
but of a more rusty hue. In variani the females are of a brilliant
golden-chestnut colour, and never assume the black livery of the
males. A very old female may develop a deeper chestnut hue,
but there is no suggestion of black, as is seen in young males
while still in an adolescent stage.
General Description of H. variani.
Male. Amassive upstanding Antelope,almost as large as equinus
but not so high in the leg. ‘he foreface is long and narrow, the
forehead not so high and prominent as in niger, with immense
horns rising perpendicularly from above the eyes and curving back-
wards in an individually variable are. here is usually a wide
outward sweep towards the tips; more rarely the horns are
parallel throughout their length. They are much compressed
laterally, with very strong annulations. A section through them
taken 8’’ above the base measures roughly 33x 2". The annu-
lations have a tendency to become obliterated on their lateral
surfaces centrally, especially on the inside. Good horns may
measure 58, with girth 103’’; but a length of 64” has been
recorded, The unannulated terminal portion i is generally straight.
For this reason, coupled with their perpendicular rise from ‘the
frontals, the enals of the horns, when the animal is standing erect,
do not as a rule curve dancers, but are carried more or ese
parallel to the line of the back.
The ears are comparatively small in fully adult males. In
young males and in the females the ears are larger.
The neck is full, short, massive, oval in seceom, and wedge-
shaped fiom alors Jlommnvantile The mane, rnd is full and
29%
a
320 MR. G. BLAINE ON THE ZEBRAS
composed of rather soft crimped hairs, unlike those of eguinis
which are straight and brush-like, reaches to the middle of the
back. The back itself is level and short, with immense breadth
across the loins, and quarters wide, full, and round like those of a
cobby horse,
The tail, which is crested along its dorsal surface, terminates
at about the level of the hocks in a thin tuft.
The testes in fully adult males are very large, the scrotum
being pear-shaped as in bovines.
Colour above intense glossy black, under parts and inside of
thighs white. Forehead and foreface w holly black, having in
some specimens a faint trace of the cheek- -stripes indicated by a
few buttish hairs. Lacrymal tufts with extension upwards in the
form of a frontally converging whorl of hair to the level of the
eyebrows, buff. Lower part of cheeks, chin, and lips creamy buff.
Hair of inside of ears white with buff edges; back of ears tan.
Mane black, with a few reddish-tan hairs on crown. ‘Tail black,
with reddish-tan hairs s along its dorsal crest; tuft wholly black.
A deep tan patch covering side and back of hocks, and a stripe of
the same colour extending down back of forelegs to fetlocks.
Legs otherwise black. Jy ‘ides in old males light taw ny or light
brown, giving a goat-like expression to the eye.
Young males resemble females. but the prevailing body-colour
is duller, being tawny and black in almost equal proportions.
As the animal advances in age, the black predominates, spreading
forwards from the neck, chest, and shoulders, along the flanks and
on the quarters, the last part to change colour being the lumbar
region of the back. Face black as in old males.
Females. Bright golden chestnut, lightly suffused with black on
the flanks, due to a percentage of the hairs being tipped with biack,
and deepening to a dusky brown with a sprinkling of long tawny
hairs along the under side of the neck and on the chest. Under
parts, including inside of hams, white. Legs golden chestnut, with
a broad dusky streak down centre of knees. Back of pasterns and
coronary band black. A blackish-brown blaze down centre of
face flanked by narrow indistinct cheek-stripes. Lacrymal tufts,
inside ears, and sides of jaws buffy white, paling to white on lips
and chin. Back of ears, crown,and forehead golden chestnut.
Mane blackish brown, with an admixture of golden-chestnut hairs
which replace the black on the crown and in rear of the withers.
Tail-tuft deep brown. Irides brown.
Habitat. The Angolan Sable Antelope is found in the strip of
country enclosed by the upper waters of the Quanza River and its
eastern tributary the Luando, and itis also said on good authority
to occur between that river and its western tributary the Kutatu,
all three rising in the great central plateau that divides the Congo
basin from that of the Zambesi. East of the Luande the country
is dry, barren, and sandy, being known locally as the “* hunger
country,” and all reports, both native and European, tend to
prove that the range of the Sable does not spread beyond the
AND SOME ANTELOPES OF ANGOLA. 321
confines of these rivers and certainly not across the watershed.
It is, therefore, very restricted as to locality, and this cireumstance
may account in part for its highly specialized development. Col.
J.C. B. Statham, who in 1920 penetrated into this country from
Malange in the north in quest of this Antelope, did not e neounter
it in any large numbers until he reached the region in which, in
the autumn of 1918, I procured all my specimens: viz., some
70 miles south-east beyond the junction of the Quanza and
Luando. The distance between these two rivers is about 30 miles,
with a low, flat, ironstone ridge forming the divide, being steepest
towards the Quanza, from which it is distant about 10 miles.
The elevation of this country is roughly between 3000 feet and
4000 feet. ‘The Sable here are mostly found on the Luando side
of the watershed where several streams rise, such as the Luce,
the Kaluando, the Dunde,and the Lusinge, the latter being sat
and forming a group of salt-pans known to the natives as “ Ochi-
Songwe.” ‘This area is covered with an extensive bush-forest,
having narrow plains bordering the rivers and strung out along
the headwaters of the streams, and interv ening here and there
as round or oval openings, ter med “dambos.” The trees vary in
density, but nowhere is it possible to see more thau 300 yards
ahead between their massed trunks. The undergrowth is light,
consisting of little low seedlings of bush a few feet high. and a
fine, soft, sparsely-growing grass, which is the principal food of
the Sable. There are also extensive beds of a low legumincus
plant with a dull pink flower on which the Sable occasionally
browses, and numerous bulbs and tubers, with some very beautiful
flowers. The ground is thickly carpeted with dead leaves, and
studded at intervals with enormous termite mounds upon which
grow trees and bushes. The soil is a sandy loam enriched with
leaf-mould, giving place on the dambos to the usual sun-haked
knobbly grey clay, where a hard, coarse grass grows which the
Sable never seems to eat. They were partial to the denser parts
of the forest, and especially where certain trees are abundant,
probably several varieties of Cassia, including the graceful Cassia
occidentalis and Huapaca gossweileri, a tree of stiff-set habit,
having large, expanded, racket-shaped leaflets radiating from one
stem like those of the horse-chestnut.
The numbers in a herd vary from about eight to twenty indi-
viduals, about half of them being bulls. Two young bulls are
frequently to be met consorting together by themselves, always
very shy and wary. They have probably been driven out of the
herd by a jealous old bull. The sentinel of a herd was nearly
always a young bull. They no doubt make the best guards from
experience gained when running singly or in pairs in the forest.
I never encountered an old bull by himself. In a small herd
there is usually only one big black bull. In large herdsthere may
be two and quite a number of young bulls ranging from sub-adult
nearly black ones with half-developed upright horns to younger
ones of a dusky-tawny colour.
322 MR. G. BLAINE ON THE ZEBRAS
The herds of Sable at this period of the year, viz. September
to December, do not move about a great deal, and it was possible
to walk a long distance of 8 or 10 miles through the forest
without crossing a single fresh track, and yet within the circle so
described two or three herds might be harbouring. As referred
to above, they confined themselves to certain sections of forest
which could always be recognized by the kind of trees of which it
was composed, and the art of finding the Sable was a question of
keeping within the limits of these trees.
Unless disturbed a herd remains in that section in which it has
established its feeding and resting quarters. Grazing through
the early morning till about 9 a.m., it wanders off to a denser
patch, where it rests till about 3 p.m., the whole herd lying down
rather scattered and often without a sentry.
No concise information relating to the breeding habits was ob-
tainable, but no small calves were noticed running in the herds,
and as the big bulls were all with the cows, it might be inferred
that the autumn is the rutting season. One afternoon J had a herd
under observation in some thick forest, and noticed the old bull
apparently taking stock of his cows. He rounded them up one
by one, driving each to the front, and then turned back to fetch
another. His methods were rather autocratic, for he ran at the
cow and butted her behind with the front of his horns if she was
at all inclined to loiter. Having passed them all in review, he
followed sedately in rear of his protégées.
The big bull is generally somewhere in the background, that is
to windward, and it is difficult to see him—black among black
shadows and the lattice of black tree-trunks. He spends a great
deal of his time lying down, but often a great pair of curving horns
rising directly above the undergrowth discloses his position. The
brilliant chestnut cows, on the other hand, easily catch the eye.
At intervals one or another, generally a young one, gets on its
legs and begins to graze, and then lies down again on the same
spot. At length all rise together as by a preconcerted signal, and
wander off slowly grazing.
Their drinking habits were not noted, but as sufficient water
was always obtainable from pools in the stream-beds in the high
forest, it is probable that they drink regularly. They did not
appear to go down to the Luando to drink.
The bulls have a habit of rubbing their great horns on the bark
of saplings and small trees, the branches of which they break off.
This action imparts to the horns a red tan colour, very beautiful
in contrast with the velvety blackness of the skin. They also,
after soiling, invariably scratch with their hind feet, more canis,
cutting long grooves in the earth. The rubbed bark, broken
branches, and sapling tops hanging head downwards from the
stem by a strip of bark, together with furrows scratched in the
soil, are a certain indication establishing the presence of Sable in
distinction from Roan Antelope. The latter occasionally go
through the same performance of scratching, but not so regularly
AND SOME ANTELOPES OF ANGOLA. 323
as the former, and do not break down branches with their horns.
It takes some experience to distinguish between the spoor of the
two species. Roan, however, have rather a longer hoof, with the
toes sharply pointed and separated at the tips. It is also larger,
if that of an old bull. Sable have a hoof wider at the heels
with blunter toes, the outer toe in bulls being usully worn
square at the tip. The spoor of herds is easy to distinguish,
cow Sables having a neat triangular hoof resembling that of
Waterbuck.
The Roan also do not frequent the same parts of the forest in
which the Sable make their home, preferring the smaller, thinner,
and more open tree-bush in the vicinity of the dambos, which are
their feeding-grounds, Sable do not seem to care for the dambos,
and when moving through the bush, usually make a detour inside
the edge of the cover rather than face the open space, especially
when alarmed. On first being disturbed they do not run far, bu
stop to look back from the shelter of any convenient thick cover,
and if then not followed too hastily, will generally settle down, and
may be disturbed several times without going far, leaving a sentry
posted. But if one of their number has been shot, the herd will
keep going a long distance, and clear out to another part of the
forest. When wounded, a Sable is extremely wary and diffi-
cult to approach, and before lying down is careful to select a
good position from which to guard its back tracks. In this
country they have few natural enemies, such as lions or wild
dogs.
The country is thinly populated, the villages of the natives, the
Luimbe, being few and far between. They are a poor race,
cultivate scarcely at all,and at the time of our visit were sub-
sisting mainly upon wild fruits and honey. They had no cattle
and very few sheepand goats, but a recent survey has pronounced
the country suitable for ranching purposes, the fine quality of the
herbage, coupled with the salt-licks, making ideal conditions for
cattle raising. All these causes no doubt have combined to favour
the development on special lines of the Great Sable.
The only other Ungulates observed in the country were Roan
Antelopes, Reedbucks, Oribi, on the small plains, and a Grimm’s
Duiker, identified as Sylvicapra grimmi leocoprosopus, which were
numerous in the forest. The fresh tracks of a troop of four Eland
appeared for several days, but these were probably visitors to the
salt-pans.
Adult Sable bulls have a peculiar pungent smell, which pervades
the whole animal and clings to the skin for weeks after death, in
spite of daily exposure to the sun and wind. It resembles a
vegetable rather than an animal smell, and suggests the aroma
of the bush itself, although not of any particular plant. They
are also covered with several kinds of ticks, which are very
numerous on the neck and shoulders, where the hair is worn thin
in consequence. The younger bulls had fewer ticks and the cows
fewer still,
324 _ MR. G. BLAINE ON THE ZEBRAS
Some measurements in inches of two adult bulls and one cow
are given below :— Rin tf
(1) (2) (3)
Oldad. g. Ad. ¢g. Ad. 2.
iEeadsamd! bodiyop cer nceeaa senate ecre 80 88 15
Wy ee ta iene ties Some Weate dak ome loers 21 212 15)
* Mean height at shoulder ............ 53 554 AT
Girth behind shoulder ............... 663 6435 58
Length of head from crown to lip... 163 = —
Girth of neck at throat ............... 294 29 25
be Bu eheSburaas ae eae AQ 48 393
Length of ear from notch to tip... 9% 10 9
(1) A splendid old bull with very massive horns, 58” long and
104” round their base.
(2) A very big adult bull, with horns still growing and traces
of tan on the top of the loins.
(3) An adult cow that had probably given birth to several
calves.
In conclusion, one is glad to remark that there is no reason
why this splendid Antelope should not continue long to exist in
its present environment. During three months spent in the
locality I must have seen at least 100 individuals, forming
several strong and flourishing herds, and measures have been
taken by the Portuguese Government, at the instigation of
Mr. Varian, to protect them.
The local native name is ‘‘ Sumbakoloko.”
REDUNCA ARUNDINUMYT Bodd.
On comparing a number of skins of this Antelope from various
localities as far as Lat. 10° S., which is approximately the limit of
its known range, it is difficult to arrive at any positive con-
clusions. In the B. M. collection the skins from South Africa,
Zululand, the Transvaal, and Southern Mozambique are greyer.
A skin from N.E. Rhodesia and a series collected in Angola are
more fulvous. The most pronounced fulvous skins are two
collected by the Rudd Expedition—one from near Beira, and the
other from Gorongoza District in P. E. Afriea. A paler and
greyer race has been described by Lord Rothschild (P. Z. 5. 1907,
p- 237) from Fort Jackson in N. Rhodesia. The Nyasaland race
* This measurement is taken by holding the foreleg perpendicularly under the
animal, forcing it into the shoulder and then measuring from heel to top of withers.
This method would give a mean height. The shoulder height of the animal when
alive would probably give 1’’ more.
+ R. arundinum occur in the Bahr-el-Ghazal Province of the Sudan in the
neighbourhood of Dud Majok, N.E. of Wau. Along the Meshra-er-Rek-Wau road
and westwards along all the tributaries of the Bahr-el-Ghazal River a small Bohor
Reedbuck is common.
AND SOME AN'TELOPES OF ANGOLA. 325
appears also to be greyer. ‘There does not, therefore, seem to be
a line of geographical progression from grey to fulvous in any
given direction; but material on which to work is sadly deficient.
There are no differences to be detected in skulls from any of
these places.
Body measurements in inches of an adult maie Reedbuck from
the Luando, Angola:—
elleardtania! lb odivyaveeeeee eee BARA is sol fns 62 .
{LAWL Sevan Cre ROE Ae CRIES Ms cetird aie ee we
Mean height at shoulder ................... 38
Girth behind shoulder ..................... 4]
Length of ear from notch to tip ......... 62
A new Orabi.
OUREBIA RUTILUS, sp. Nn.
A beautiful Oribi, distinguished by its brilliant colour, wavy
or curly hair, and the obsolescence of the bare patch below the
ear.
Skin.—Colour of upper parts uniform tawny orange, changing
abruptly to white underneath. The white area extends higher
on the flanks than usual. Inside of ears, superciliary stripe,
lips, chin, and throat white. Back of forelegs, hoofs, and of
hindlegs to hocks white. There remains a slight trace only of
the usual round subauricular patch, this area being almost com-
pletely covered with fine, short, buffish hairs. Tail black and very
short, about 30 mm. Skull rather long and narrow, with long
nasals, flat straight profile,and comparatively shallow lacrymal
pits. Dimensions of typical skull: length 177 mm., breadth
75 mm., nasals 67 mm., upper tooth-row 52 mm.
Hab. The country between the Quanza and Luando Rivers,
Angola.
Type. Ad. 3, skin and skull, B.M. no. 20.4.27.40. Collected
and presented by G. Blaine.
There are six specimens of this Oribi in the B. M. collection
from the above locality—two males and four females, all adult,
shot by the writer in the autumn of 1919. They are remarkably
uniform in colour, one specimen only, a female, differing from
the rest in having a blackish patch on the crown. In one char-
acter, viz. the suppression of the subauricular patch, they show
some affinity to the Cape Oribi, but have much shorter tails,
so short, in fact, as to be almost rudimentary. O. hastata from
Nyasaland has a pronounced bare subauricular patch. Neither
has the long curly hair and brilliant colouring of this species.
In comparing skulls, those of Oribis from the Cape ave rela-
tively shorter and broader, with short nasals; those from Rhodesia
and Nyasaland are longer and narrower, with long nasals. The
skulls of O. rutilus approximate the latter, but are straighter in
profile than either and have shallower lacrymat pits.
326 MR. G. BLAINE ON THE ZEBRAS
Comparative skull measurements in millimetres :—
B.M. No. Length. Breadth. Nasals. Tooth-row.
Oribi (Cape)...... 135 a 165 ili D3 50
(approx.).
Oribi (Rhodesia). 12.5.10.2. 168 72 60 51
HOSTAL arash lee 13.9.5.4. 176 (fl 65 53
TUES aise e ctte 20.4.27.40. 177 75 67 52
Ovibis have peculiar inguinal glands, and in these Angolan
Oribis they were very noticeable. They are covered at the sides
by a modified pouch, consisting of a pair of flaps of tough parch-
ment-like granular skin with serrated edges, sparsely covered
with coarse white hairs and enveloping the genital organs of the
male and the mammee of the female. The female seems to shew
a greater development of this curious growth than the male.
This peculiarity is briefly referred to by Roosevelt in * African
Game Animals,’ il. p. 555, as follows :—“ At the groin are a pair
of deep inguinal sacs marked by a growth of long peculiar pithy
hairs.”
Pocock*, in describing these inguinal glands in Ourebia, gives
examples of their variation in the different races of that genus,
notably in nigricaudate, montana, and owrebi.
Since my observations of these glands in the case of rudtilus
differ in some respects from his descriptions, it is probable that
this variability is characteristic of the genus.
SYLVICAPRA GRIMMI LEUCOPROSOPUS.
OQ. Neumann, Sitz. Ges. Nat. Freunde, 1899, p. 18.
In my collection of Antelopes from Angola I brought back
seven specimens of a Grimm ’s Duiker from the country between
the Quanza and Luando Rivers. These have been found to agree
best with Herr O. Neumann’s description of his lewcoprosopus,
which he thought might have come from the interior of Angola,and
in this supposition he was probably correct. He describes as his
type a male living with a female in the Berlin Zoological Gardens.
The female he referred to Cephalophus altifrons (=ocularis) Peters
from Mozambique or to flavescens Lovenz from Matabeleland as
being greyish brown in colour with less white on the under parts.
But he gives a very full and careful description of the male, to
which the above specimens conform most accurately, and especially
touching the masked or spectacled appearance of the face viewed
frontally, due to the white rings encircling the eyes.
There are two skins of S. splendidulus Gray and also a stufted
specimen in the B. M. collection, allfrom Angola. They represent
* P.Z.S. 1910, pp. 882-884; also A.M. N. H. 1918, p. 430.
AND SOME ANLELOPES OF ANGOLA. Bei
a Duiker with the usual characters of grimmi. In one detail
only are the above at variance with Neumann’s description. He
states that his Antelope was smaller than average grimmi,
whereas the writer, who saw many of them in the open forest
between the Quanza and Luando, considers that they were above
the average size for this species. But the animal which he
described was a menagerie specimen, and therefore unlikely to
have attained a perfect development.
They are as richly and strikingly coloured as are the Oribi and
the Great Sable Antelope that inhabit the same country.
A fetus and a young female a few weeks old are greyer and
more grizzled in colour, resembling the northern race abyssinicus,
and show no traces of the white markings of the adults.
Parr Il.
Between Benguela and Mossamedes lies a wild desert region of
rock, sand, and thorn-scrub, almost waterless and uninhabited
save for the presence of nomad Baquando, half negroid, half bush-
men, who roam over the country with their herds of goats, and a
few Portuguese and native fishermen, existing from hand to
mouth along the beach.
There is a deserted sugar factory at Equimina, its spacious and
not unimposing facade rising above a close-set tangle of tropical
bush and old overgrown gardens, near the middle of a wide and
pleasant bay, one of the few places along this coast possessing a
supply of good fresh water. At one time the site of a prosperous
plantation employing considerable native labour, the place has
now shrunk to a small untidy village harbouring an uncouth
assortment of beach-combing blacks and two or three Portuguese
fishermen.
In the next bay to the south, called Elephant Bay, distant
60 miles from Benguela, a whaling station has been established
by a Norwegian company.
Mr. Tyler Thompson, an Englishman well known in Angola,
has been in sole charge of this whaling station during the war,
and it was here that I landed at midnight on December 20th,
1918, after four consecutive days and nights spent in an open
fishing boat, tacking up against contrary winds from Benguela. It
was some relief to disembark on the threshold of an Englishman
in this otherwise inhospitable wilderness.
Mr. Thompson, being an old elephant hunter with an intimate
knowledge of the game of the country, was able to give me much
valuable help and information, and I am indebted to him for the
success of my hunting excursions in this difficult country. A
supply of good water in small barrels, and fresh frait and
328 MR. G. BLAINE ON THE ZEBRAS
vegetables from his carefully tended garden at Equimina, sent up
to my camps by relays of carriers, helped to smooth over many
minor discomforts, and enabled me to look back upon this trip
as one of the most interesting and delightful of my African
adventures.
Without some elementary knowledge of geology, it is difficult
to give a deseription of a country whose outstanding features are
bare rock and sand. Having very little, | must, therefore, be
excused for a tenative and sketchy account of its outlines, which
are so strange, wild, and rugged as to merit a description.
After passing the mouth of the Coporolla River below Dombe
Grande, the littoral plain disappears, and gives place to high lime-
stone cliffs alternating, as at Elephant Bay, with red sandstone.
The coast-line here is indented with small bays bordering bush-
covered sandy flats, which run inland for a space of 2 or 3
miles and are bounded by cliffs. Numerous dambas, or dry
water-courses, debouch into these bays through steep defiles worn
in the encircling walls of rock. Access to the country behind is
only practicable up one of these dambas, as any attempt to climb
to the top of the cliffs and thence proceed across country involves
the surmounting of an appalling series of obstructions; but by
plodding patiently along a damba through all its intricate
windings, one is eventually rewarded by reaching a country
where it is possible to follow the direction that one fancies.
The lower reaches of these dambas are contained by perpen-
dicular walls of ‘‘ pudding-stone” conglomerate. As they continue
upwards, the cliffs give way to precipitous slopes of cretaceous
rocks. Finally becoming shallower, they pass between unduiating
ridges covered with loose stone and shale, merging on to sandy
flats, whence they break out into a aehronle ag dry water-
courses.
At this point the terrain expands into many little plains which
are more or less confluent, baving a central nucleus traversed by
an uninterrupted level stretch from 12 to 15 miles long. ‘These
plains spread laterally into irregular bays and gulfs, and send out
long corridors through rocky defiles to end in “dambas which lose
themselves among one hills. Kneompassing them on every side
are steep stony ridges and conical kopjes built up of gneisses and
mica-schists, with huge outcrops of white quartz occasionally
crowning their summits.
A moraine of loose fragments of glistening white quartz litters
the lower slopes between! the hills. Elsewhere large surfaces of
undulating ground are thickly spread with the same débris,
which reflects a blinding glare from the pale metallic blue of
the sky.
There are no trees to cover the nakedness of the land, but pale
green thorn-bushes of the wait-a-bit variety, nearly all having
ord hooked thorns, with some cactus and euphorbia, are dotted
more or less evenly about this country, the peculiar features of
AND SOME ANTELOPES OF ANGOLA. 329
which in their wild disarray appeal strongly to the imagination.
Here Pelion has been piled upon Ossa, there Ossa has heaved up
and overthrown Pelion, so fantastically and in such confused
masses do the hills range themselves or fall asunder into island
groups and solitary kopjes in sandy wastes.
Some 57) miles inland from the coast and dominating all this
region is a vast mountain rising in two great blocks from a sunken
plain to a height of 5000 ft. or more. It is crowned with tower-
ing precipices of bare rock fluted with perpendicular fissures,
while its flanks and base are covered with a downward-spreading
forest of thick bush.
Owing to the restraint put upon my mobility by a crew of
untrustworthy carriers who terminated their contract by running
away, | was unable to reach this mountain, which would well
have repaid.a visit. It probably stood at the inland limit of the
desert region.
There is no surface water, but by digging in some of the
dambas it is possible to find water in certain places. Most of it
is brackish and unfit to drink, and in the lower reaches of the
dambas a brackish water often oozes out of the sand and trickles
for a few hundred yards to be again absorbed. Water was,
however, available from used water-holes near the two camps
J made in this locality, the farthest one being in rock and con-
taminated by the Baquando goats and baboons that frequented it.
This is a hunter’s paradise. To climb in the early morning
to the topmost pinnacle of some hill selected for a wide
range of view, to sit in the cool 8.W. breeze as it streamed
inland from the Atlantic and watch the sun rise over a
shoulder of the great mountain, was an unforgettable expe-
rience. As the clear white light came slanting across the
crests of the hills and began to radiate downwards on to the
plains, every detail below, becoming illuminated, sprang clearly
into view. At this time the game was all on the move, and
could be detected with glasses 3 or 4 miles away. Below, in the
foreground, delicate clean-cut Springbuck in open herds moved
briskly between the bushes from tuft to tuft of grass, throwing
long blue shadows across the cold, pale sand. Beyond, a solitary
bull Gemsbuck, standing motionless as if carved out of stone.
would suddenly spring into life, and with characteristic pendulum-
like nodding of the head and smooth, rapid walk, join a herd of
his kind already grazing on the edge of a dry water-course. Or
the more bulky and indolent forms of a herd of large Mountain
Zebra would be grouped in tones of pale gray and pinkish white
among the thicker bushes clustered about some stony ravine.
The sun, meanwhile soaring perceptibly higher above the crests of
the lovely purple mass of the mountain, swept back the shadows
into the farthest recesses of the landscape, changing the cold
blue light of dawn into the sunny brightness of an African
morning.
330 MR. G. BLAINE ON THE ZEBRAS
Equus HARTMANN Matsch.
Equus hartmanne Matschie Sitzber. Ges, Nat. Freunde, 1898,
p. 174.
Equus penricet Thos. A. M. N. H. ser. 7, vol. vi. p. 465 (1900).
Most characteristic of this land of stones 1s this big Mountain
Zebra, a fair cousin of the true Zebra of the Cape, now almost
extinct.
Originally described by Matschie from Huanib and Uniab on
the coast N. of Walfisch Bay, and later by Thomas fiom a speci-
men presented by Mr. G. W. Penrice from the Moninha River
N.E. of Mossamedes, there can be little doubt that the two are
one and the same animal. My specimens were collected near the
coast .at Elephant Bay, 100 miles N. of Mossamedes, and corre-
spond accurately with Matschie’s original description.
Habitat.
The home of this Zebra appears to be in the coastal belt of
S.W. Africa where the country 1s rocky and precipitous. I have
seen the spoor on the wet sand of the sea-shore at low tide, and
have often watched them grazing on the low-lying flats and on
the plateau-like tops of the adjacent cliffs within a mile of the
sea. According to Mr. Tyler Thompson’s observations, with
which my own coincide, they do not penetrate further than
30 miles inland, and are strictly confined to the waterless desert
region.
Description.
Skull.—A. comparison of the skulls of zebra with hartmanne
shows slight differences only, the latter being more massive.
Zebra shows modifications of the frontal aspect between and
below the orbits. There is a depression along the naso-frontal
suture. The nasals also are depressed centrally, and the maxille
are compressed laterally above the infra-orbital foramen.
Hartmanne has the frontal profile straight, the nasals straight,
and no lateral compression of the maxille in the region of the
nasals.
Tn zebra in the flesh, the profile is dished or dented in between
the rather prominent field of the forehead and the muzzle, as is
seen in an Arab horse.
In hartmanne the whole profile is flat with a descending
muzzle, giving the head a Roman-nosed appearance.
There is also noticeable at the anterior end of the malar, near
its junction with the maxilla, on the inferior surface of the malar-
maxillary ridge and in line with the last molar, a roughened bony
spur. This is not noticeable either in zebra, quagga, or grevyt,
and has probably been developed for the attachment of a more
powerful muscle to meet some special requirement of mastication.
Skin.— Direction of hair asin zebra. Resembles zebra in colour-
pattern, except that the relative areas occupied by dark stripe and
AND SOME ANTELOPES OF ANGOLA. Boll
light ground are reversed. Colour of stripes rusty liver-brown,
darkening to sepia on neck and down legs to hoofs, where it forms
a broad coronary band. On the top of the back, and notably in
the lumbar region, the stripes are speckled with ochraceous hairs.
Gridiron pattern fades into indistinct lines and spots as it
approaches the tail; also down its basal third. Central dorsal
stripe very narrow. Stripes down flanks taper to a point in line
with elbow, leaving the chest, belly, and inside of thighs white.
A broad band of sepia runs down centre of belly to chest.
Ground-colour above a warm tone of ochraceous or sandy buff,
paling below to buffy white. Centre of tail ochraceous buff,
fading to white at sides. ‘Tail tuft ragged and thin, buffy white
above, changing to bistre at the end, the under hairs being black.
Head.—Stripes on forehead liver-brown, close and narrow with
blurred edges, paling to warm sienna down face and on cheeks,
Muzzle sienna. Ears as in zebra but more lightly marked, with
brown tips. Mane striped dark brown and white, the dark
stripes fading to bistre at the ends of the hairs, which are long
and ragged, the longest measuring 7 inches.
The skin has a faded rusty appearance throughout. Chest-
nuts on forelegs large, smooth, and oval, measuring 43 inches by
33 inches. Hoof measuring 43 inches by 24 inches, both fore
and hind bemg very similar. They are very hard. ‘The frog
shows only an outline, being worn quite smooth. The heels are
long.
This Zebra is a massively built animal, considerably larger than
the Bonte Quagga and approximating to a Grevy Zebra in size.
The head is long and heavy, with a large muzzle. That of one
old stallion I shot resembled a cart-horse, with deep pits above the
eyes, Roman nose, and pendulous lower lip. The ears are large.
There is in both sexes a dewlap 3 inches deep and commencing
at about the same distance from the throttle. The loose skin
of the dewlap is much thickened towards its apex, the whole
of the skin of the neck increasing in thickness from above down-
wards. The end of the dewlap contains au indurated ball of
flesh, about the size of a large walnut, lodged in fatty tissue which
is adhesive to the skin. The neck is short and deep, the withers
low, and the back level and rather long. The Jimbs are massive,
with big bony knees and hocks and big cannon-bones.
In action this Zebra moves more freely than the Bonte Quagea,
whose paces are short and lumbering. When suddenly startled,
he gallops off with head carried high and nearly horizontal and
neck arched backwards, but soon settles down to a slinging trot.
At this pace the knees and hocks are well flexed. A herd in
retreat always makes for the hills, up which they clamber with
marvellous ease. and surety over the roughest, stoniest ground.
The hills of this country are traced all over by their paths, so
worn as to resemble native foot-tracks. It is only possible to get
over some parts of the country by using the Zebra paths, and
the main footpath along the coast between Dombe Grande and
332 MR. G. BLAINE ON THE ZEBRAS
Mossamedes is composed of their joined-up tracks. These Zebras
do not run in large herds, eight being an average number in a
herd. ‘They graze in the mornings and afternoons on the tufted
grass that grows on the plains and on the lower slopes of the hills,
sheltering from the sun during the heat of the day under thorn-
bushes, where they doze away “the mid-day hours. ‘They are not
very watchful, nor does there appear to be a sentry in a herd, but
when alarmed, they make off at once, and, unlike the Bonte
Quagga, do not stop to look back until they have gone a long
distance. They then retire up one of their numerous paths in
single file, and then disappear over the sky-line into another part
of the country. During the night they make their way towards
the coast to drink at one of the rock-pools of brackish water in
the lower reaches of some damba.
Mr. ‘Tyler Thompson has described to me how, when first he
went to Elephant Bay, the Zebras used nightly to drink at the
fresh-water pool under the cliff. Led by an old stallion, the herd
would come down at a gallop and halt just short of the water on
the plain, while he would continue his career up the side of the
cliff to take his stand upon some point of vantage overlooking the
water. Having satisfied himself that no danger was present, he
would signal “all clear” with a low neigh, when the herd would
go down to the water. While they were drinking he would
remain at his post, and descend to drink himself when they had
all retired. No doubt this precaution was necessary, as lions had
often been known to visit the bay at night. It is curious that
most of the fresh water in this country is found at or near
sea-level, and the water in the dambas, where it approaches
the surface, may be fresh on one side and brackish on the
other.
During the season of light rains when I was hunting in these
parts, herds of the local race of Bonte Quagga (Hquus quaygga
antiquorwm) came down from the interior for the fine grass, and
were often seen on the same plain with herds of Hquus zebra
hartmanne, but never associating together. The latter do not
go about in large herds, from six to twelve being the ususl number,
though as many as twenty have been seen together, which is
unusual. Old stallions are oftien solitary.
Even at a distance the two species appear quite different: for
whereas the Bonte Quagga is a conspicuous and by no means
harmonious object, looking black or smoky grey against the
prevailing tone of the country, this Zebra * never appears dark in
any position in relation to the light, but always either white, pale
erey, or reddish-sand colour. When standing against a back-
ground of rocks of which the prevailing tones are various shades
of warm grey, the outline melts away and the whole animal looks
transparent, the stripes dissolving into pale shadows.
* In a shot specimen of Hquus hartmanne the ground-colour of the dorsal surface
was noticed to be exactly the same shade of pale reddish ochre as the sand on which
it was lying.
AND SOME ANTELOPES OF ANGOLA. 333
The call of this Zebra is a low, snuffiing neigh or whinny,
quite unlike the oft-repeated hysterical ‘ bweha-bweha” of Bonte
Quagea. It also makes a loud squeal.
The flesh is fine-grained, with white fat and sinews, and by no
means bad eating with a slightly sweet taste. That of the Bonte
Quagega is coarse, rank, and unpalatable, in colour dark red with
yellow fat and sinews.
Below are the body measurements in inches of two adult
stallions :—
Telerncl Bin loleXeha Medel desnon oademeen howe co oo Ole. 92
LETT Li ole eg Ne Oe a PEI SU ae 194 21
Mean height at shoulder ............ 52 D445
Girth behind shoulder .........:..... DOS 604
Length of head from crown tolip... 28 27
Gurthyot neekiatiubrosiess epee Bx) 30
ms oe ehest, Seana: 46 47
lemoGh 3780) Res ee Beare f 9 )
ie { JOVREPNG US) CAAA hobp ce ccistrha’ 4 4
ANTIDORGAS.
On preparing to work out the Angolan Springbucks, the first
difficulty to be encountered was the usual lack of material. Apart
from the Angolan series, only two complete skins with skulls
could be found in the B. M. collection, and one of these was a
stuffed specimen. The remainder consisted of stuffed heads from
the Selous collection and a few ancient relics without histories.
There is, moreover, very little literature giving any detailed
description of this Antelope for the reason that its highly-
specialized character so readily distinguishes it from all others.
Harris f gives a careful description, and Brooke ¢ follows with
another careful and accurate description, upon which I base the
differences found in the Angolan series.
Even from the scrappy material available it is evident that
two distinct races exist, the most obvious character of the one
being the imperfect development of the horns in the female,
which are small, thin, and indistinctly annulated, and of the
other the fully-developed horns in the female, which are long,
lyrate, and strongly annulated.
Other differences appear in the skulls, to which reference will
be made later, but it is first necessary accurately to define typical
euchore.
The best examples to be found representing anything ap-
proaching a series are four specimens from the Orange River
* Very old stallion. } f ,
+ ‘Wild Animals of Southern Africa,’ 1840, pl. ii. Does with very slender
horns either straight or capriciously bent, exhibiting a few indistinct annuli at their
base.
+ P.Z.S. 1872, p. 550. Horns in both sexes, but very small im female. Had.
S. Africa.
Proc. Zoou. Soc.—1922, No. XXIII. 23
334 MR. G, BLAINE ON THE ZEBRAS
Colony, all shot by Mr. Selous—one an old male, stuffed, in the
Public Gallery, and the other three stuffed heads of females from
which the skulls have recently been removed.
A description of the heads will suffice, as the colour and its
distribution on the bodies of Springbucks does not appear to vary.
The male has no trace of a triangular reddish-fawn patch on
the forehead, the face being entirely white. The lateral cheek-
stripes are distinct, but narrow and continued above the eyes to
the bases of the horns. ‘The ears are comparatively small,
146 mm. from notch to tip. The horns are basally stout, lyrate,
and strongly annulated, having a perceptibl2 spiral tendency
which might be termed an interrupted spiral. They rise at an
obtuse angle with the frontal plane,so that a line drawn per-
pendicularly through the centre of the orbits falls within their
axis. The tips are hooked backwards and inwards.
Of the females, one is without the triangular fawn frontal
patch. The other two show it pale and indistinct. Har from
notch to tip 133 mm. The horns are short, thin, nearly straight,
and hooked either forward or forward and inward at the tips,
with indistinct annulations.
In the male the skull is relatively short, wide, and deep, with
horn-pedicles upright. The frontal profile is concave, with prom1-
nent ends to the nasals. The lacrymal pits are very deep.
Basally the palatal surface is convex, and the basioccipital has
greater breadth and length. The skulls of the females are
similar in character.
Comparative measurements in millimetres of the skulls of male
and female Springbucks from the Orange River Colony and from
Angola :—
O.R.C. Angola O.R.C. a.
3. 3. 4 2
B.M. no. B.M.no. B.M.no. B.M.no.
98.1]. 20.4.27. NOUS e202 Te
28.8. 32. 336. 34.
Grea testrlemetin /aynmeeny- meee 220 240 220 223
“4 breadth eens 95 105 99 95
Vertical depth. dct: 8. fyi ke-weee 98 98 90 87
TN eisai 1S:. Mande dies ul eediune neuen (6) 87 73 83
Premanalileotsh:(olee sea) e aa eee da a 76 7
Basioccipital surface ......:..... 22x27 25x25 27 x 23
Diowermcoth=nows:- <5 panes 65 70 63 65
Length of horns in inches... 123 13 (es ils
A stuffed head of a male from N. Kalahari in the Selous col-
lection resembles the Angolan Springbuck in size, length of face,
and in having the frontal patch and the horns more nearly in line
with the frontal plane. B.M. no. 19.7.15.333.
A female from N. Bechuanaland in the same collection shows
the same characters as the above, and has long, straight horns,
hooked slightly inwards at the tips, but not so perfectly annulated
as in the Angolan females. B.M. no. 19.7.15.334.
AND SOME ANTELOPES OF ANGOLA. 335
Description of Angolan Springbuck.
ANTIDORCAS ANGOLENSIS, Sp. n.
Larger than euchore, with longer, narrower head and axis
of horns in hne with frontal plane. In males, horns not so stout
ab base and, viewed laterally, less sinuous than in ewchore.
In females, horns long, perfectly developed and annulated, lyrate
in form, with tips hooked inwards and backwards, and in all
respects exact models of the male horns. ars very large: of a
male 165 mim., female 184 mm.
Colour as in euchore, but stronger and brighter throughout;
the triangular frontal patch always present and edged below with
bright mahogany- brown.
Skull relative ely longer, narrower, and verticaily less deep than
in eewchore, with straight frontal profile and shallower lacrymal
pits. Horn-pedicles in line with frontal plane, so that a line
drawn vertically through centre of orbits falls outside and in
front of theiraxis. Basally palate not convex, basioccipital having
equal or greater length than breadth.
Hab. Vhe coastal region of Angola from Benguela southwards
towards Damaraland.
Type. Adult female skin and skull: B.M. no. 20.4.27.34, col-
lected by G. Blaine.
The Angulan Springbucks herein deseribed were obtained from
the coastal belt between Benguela and Mossamedes. The Copo-
rollo River appears to be the approximate northern limit of their
range, though a few stragglers have been seen, in some years,
across the Benguela Railway. They roam in herds over the
sandy flats, though never, I believe, in great concourses, grazing
on the soft feather y grass that grows on the open flats, and nibbling
the foliage of the thorn-bushes that line the courses of the sand-
rivers, and are scattered on the stony approaches to the hills.
They are also fond of frequenting the quartz-strewn wastes,
where their bright, clear appearance and elegant outline is in
lovely harmony with the glistening white stone and pale green
vegetation.
‘The habits of Springbuck have been so often and so well
described by naturalists and hunters, that it is unnecessary to
add my observations to theirs. J am able, however, to record
one fresh scrap of information which I believe to have escaped
notice. In Angola the fawns are dropped in January. During
the early part of that month, in 1919, 1 saw many Springbuck,
the does being then heavy with young, but I did not see any fawns
about. On the 12th of January, I went down to the coast
returning to my camp inland three weeks later. During that
interval the does had dropped their fawns, and they were to be
found lying about on stony ground in the open or under thorn-
bushes, where their mothers had left them during the day.
I once very nearly trod on one of these little creatures, huddled
between some pieces of quartzite, for they lay very close until
93%
336 MR. G. BLAINE ON THE ZEBRAS
detected. Although probably not more than ten days old, it
jumped up and trotted off with characteristic stilty action, head
between forelegs, arched back, and with the white dorsal crest
erected and spread like an open fan. I followed and it cantered
away, making miniature bounds in the air, in perfect mimicry of
an adult Springbuck. JI encountered several move during that
day, and they all went through the same acrobatic performance,
nor were any of the natives with me fleet enough to run one
down.
Measurements in inches of an adult ¢ Springbuck are here
CIV CN
ieaid and Mbodiyns0s Bae i onry cn totes! a D143
Mavi (iets Seis ates fan hehe ete mR ASE 94
Eleio tb ait snoul der Deareneece as eee eer ee 32
Garth sbehimdsshoullider: te see peers or
Fi SORMECK elt urOAb lee mae eee meee: 143
ins Ne Chestiwassate.: se aeeee nee a 214
henge bhset jean yi. (et aye eck ALB a ee tke 6i*
Besides Springbuck, the following Antelopes were seen within
20 miles of the coast :—Hland, Kudu, Gemsbuck (Orya gazella),.
Steinbuck, Klipspringer, and Dik-Dik. Also Chapman’s Zebra..
Of these, the Hland and Chapman’s Zebra were visitors that had
wandered down from the interior, atiracted by the fine green
grass which had sprung up during the season of hight intermittent
vains that fall between the months of December and April. The
remainder are regular denizens of this barren region.
OryX GAZELLA.
Above all other Antelopes the Gemsbuck seems to embody the
spirit of the African veldt. He is at home in vast shadeless.
spaces under a fiery sun, reared on the pale desert: grass and
sheltered by the scanty wait-a-bit thorn. The Kudu is mere
graceful, the Sable more magnificent, the Eland more stately, but
the Gemsbuck has a character all his own. He is thoroughbred
of the desert. His classic outline suggests some hunting scene
depicted on an Assyrian bas-relief. HKnergy, strength, endurance
are the key-notes of his conformation. His frame is massive
without being heavy, and, although devoid of hollows and angles,.
does not carry an ounce of superfluous flesh and is supported upon
light clean-boned limbs. The straight rapier-like horns, dark
brilliant eye, and clean hard colour of the pelage, handsomely
varied with black and white, go to form, in my estimation, one of
the finest game animals in existence. This opinion I know is not
generally shared by others who have seen this Antelope in its
wild state. To me there is completeness both of character and
appearance and such perfect adaptation to environment as always.
to draw my admiration.
* Kar of 2 7 inches.
99
AND SOME ANTELOPES OF ANGOLA. 330
The Gemsbuck has a small pouch ef loose skin just below the
junction of head and neck at the throat, which, like its neighbour
the Mountain Zebra, contains a hard, round lump of fleshy
matter. In the bulls the skin on the top of the neck is im-
mensely thick, being as much as 14 to 2 inches through, thinning
to about 4 inch on the under side. The hair is short, hard,
brittle, and shining—about the most perfect form of coat Nature
could devise to resist the aggressive clinging wait-a-bit thorns,
while the skin all over the upper parts of the body is tough and
thick.
T see that Lord Rothschild has made this Gemsbuck into a
new subspecies, and has been kind enough to attach my name
thereto. His reasons for doing so are based upon its paler and
greyer colour, and the reduced and partially interrupted areas
occupied bythe black markings in comparison with Oryx gazella.
The discovery of this new form tending in coloration towards
beisa has caused him to consider that the straight-horned Oryx
may now all be classified as local forms of one species. This is
very interesting, and led me recently to visit the Zoological
Gardens to get a close view of the Arabian Oryx deposited there
by H.M. the King in 1920. I noticed that this Oryx appears to
have the same pouch on the underside of the throat as Oryx
gazella, only in a lesser degree. Neither beisa, callotis, nor
algazel, the scimitar-horned Oryx, has this appendage.
The range of the Gemsbuck in Angola stops at the Coporollo
River, north of which none have ever been seen. They are
never, like the Kudu and the Mountain Zebra, found close to
the sea, nor did I see their tracks in the dambas leading coast-
wards from the interior plains. They were not in large numbers
in the neighbourhood of Elephant Bay during my visit, where
I only saw a few herds and some solitary bulls.
All Oryx are naturally very wild, and these were no exception to
the rule, They are great wanderers, and any suspicion of human
presence causes them to leave the neighbourhood.
Although generally found on the flats where the best pasture
was obtainable, they had no hesitation in climbing the hills, with
which they were obviously well acquainted, for their tracks were
to be found all over them along the Zebra paths.
One frequently came across little secluded flat basins high up
among the rocks at a place where several dambas joined, where
‘a herd of Gemsbuck had been resting, some lying down, and,
judging by the quantity of droppings both old and fresh, these
were favourite spots. In the heat of the day a whole herd would
often lie down under a single thorn-bush. I have seen as many as
ten so grouped. Solitary bulls are careful to select a bush in the
open, and two or three may often be seen in the same plain lying
each under his favourite bush, widely segregated from each other.
Under certain isolated bushes the sand was always much trodden
and scooped out, showing where Gemsbuck were in the habit of
resting. Gemsbuck keep generally so far in the open away from
338 MR. G. BLAINE ON THE ZEBRAS
cover that they are difficult to stalk. Attempts to descend upon:
them from above down a hill-side generally resulted in failure,
as they were very quick in spotting any movement above. When
alarmed they get off the mark more rapidly than any other
Antelope [ know. Even when lying down they are up and away
with the quickness of thought, but always stop to look round
after going two or three hundred yards. The action is smooth,
both in the trot and gallop, and close to the ground, with the
nose stretched out, the horns thrown back, and the neck not
appreciably raised. The heavily tufted tail streams out behind
at the gallop, but when trotting is swished from side to side.
Gemsbuck are very keen- -sighted, and when their suspicions.
ave aroused will continue to stare steadily at the spot from which
they sense danger with great persistency, and will detect the
slightest movement behind any but dense cover. 1 remember
one raorning watching a herd, after grazing on some distant
stony flats, wander off and take up a resting position on a
shoulder connecting two quartz-topped kopjes which appeared to
be very favourably situated for a stalk. After making a detour
of several miles to approach one of the kopjes from a flank,
I climbed to the top of the great white bouiders and peered over
into the pass immediately beneath. Fer the first moment I saw
nothing but a stretch of glaring sand and some thorn-bushes.
The next the whole herd magically leapt into view. Hach member
of it was standing perfectly motionless beneath me—most of them
in the open, a few under bushes. They looked absolutely trans-
parent, their outlines fusing into the reddish sand. Yet every
detail was there, for they were very near. I have often before
been struck by this curious effect, especially with Oryx. .Jé appears
to be due to the perfect toning of the body- Shite with a back-
ground of sand or light stone, and to the fact that the black
markings are exactly the same tone ag the shadows thrown by
any solid object under intensely bright shimmering light. For
instance, the black markings of an Oryx standing “under a thin
thorn-bush through which the light penetrates, become so per-
fectly fused with the shadows thrown by the branches that it is.
impossible to detect, even with powerful field-glasses, where the
solid marking leaves off and the shadow begins. This I have
often tested.
The Gemsbuck cows in this country drop their calves in
January. They are curious little creatures with large ears, short
necks, and of a grizzled fawn-colour, and I remember once mis-
taking one that I saw in the distance cantering after its mother
for a hunting-dog. The dark markings first appear in indistinct
tones of greyish ‘brown diffused about the areas, Which are even-
tually occupied by the black bands in more concentrated form.
The white markings on face and legs have not yet been assumed.
The bulls at this time are generally away from the herds.
t counted two herds composed entirely of cows, one of sixteen
and the other of ten.
AND SOME ANTELOPES OF ANGOLA, 339
Measurements in inches of adult bull and cow Gemsbuck are
here given :—
Ge OF
Jeleeval eiavel looted ccoccdosaudoarncoence 79 iG
SRE erie ek ee Os Ra a 8) 18
eishinatyshoulder) =ss..ss.2ese 464 DO
Girth behind shoulder ............ 61 60
> Of neck at throat ......... Paste do) 25s
5 nh Chest eee 50 39
ema ciro tears. 5 ai sie eee lie =
s
A Note on the Blue Buck (Cephalophus melanorheus).
At the Loanda Zoological Gardens were a male and female
Blue Buck in a large paddock. The general colour of this pair
was ash-grey with patches of arusty tinge, The latter may have
been due to a seasonal change of coat or to conditions of
captivity.
Description.— Head darker than body. Rump not noticeably
darker than body. Legs ash-brown. Tail fringed with white
hairs, with the base and centre black.
This pair, which appeared to be rutting, were following one
another about the enclosure, and were seen repeatedly to face one
another and rub the facial suborbital glands together, with a
stropping action, continued repeatedly, first on one side and then
on the other.
EXPLANATION OF THE PLATES.
Poate I. Hippotragus variani: ad. f skull in B.M. Coll. Length from occiput
to end of gnathion 475 mm.
Puate Il. Aippotragus niger: ad. 8 skull in B.M. Coll. Length from occiput
to end of gnathion 488 mim.
Pratr iil. Hippotragus equinus: ad. § skullin B.M. Coll. Length from occiput
to end of gnathion 470 mm.
Pratt iV. Basal aspect of skulls of A niger, B variant, and C equinus, showing
relative positions of ¢, c, condyles and , p, paroccipital processes.
Pratt V. A. The Angolan Sable Antelope.
B. Specimens of g and 2 Angolan Springbuck (stuffed) in the B.M.
Coll.
Pratr VI. A. Old stallion of Hartmann’s Zebra, showing dewlap.
B. Dorsal view of Hartmann’s Zebra, showing gridiron pattern.
Pratt VII. A. Antidorcas angolensis: ad. 8 skull in B.M. Coll.
B. Antidorcas euchore: ad. 8 skull from the O.R.C. in B.M. Coll.
a’ and 6’, basi-occipitals of above.
Prats VIII. A. Antidorcas angolensis: ad. 2 skull in B.M. Coll.
B. Antidorcas euchore: ad. 9 skull from the O.R.C. in B.M. Coll.
The above skulls (Pl. VIII.) are shown to too large a scale.
They should appear smaller than those of the males shown on
Tels WD
ri
i ) 4
¢ fares
hy
1
Me
Ona |
u
WT
Way
ey a
i Ln a hae
it (ics ay ABH, 20
4 ees Am F
RESPIRATORY MECHANISM OF THE AMPULLARIIDA. 341
17. Notes on the Respiratory Mechanism of the Ampullariidee.
By Guy C. Rosson, M.A., F.Z.S.
(Offered for publication by permission of the Trustees of the British Museum.)
| Received February 21, 1922: Read February 21, 1922
During the latter part of the last century a good deal of work
was done upon the respiration of the amphibious gastropod
Ampullaria. The studies of Bavay, Sabatier, Bouvier and Fischer
together with the more recent work of Ramanan yielded a
satisfactory general account of the process. The method by
which the gill and lung are alternatively employed, the con-
ditions under which one or the other becomes functional and
the relation of the circulation to the respiratory organs are there-
fore fairly well known. There are, however, several details upon
which the older accounts are vague, and certain features seemed
so inexplicable that the donation by Colonel Knight and Colonel
Porteous of living examples of A. vermiformis to the Society,
afforded a very welcome opportunity for fresh study.
The general phenomena of pulmonary and branchial respiration
recorded by Jourdain (6), Bouvier and Fischer (3); ), and Bavay (1)
for various Neotropical forms are found to take place in A. vermi-
Jormis. ‘There is no need, therefore, to describe the details which
may be found in the two latter papers. Bavay described three
types of respiration, and I propose to use the term ‘ siphonal’ for
the one in which air is drawn into the lung through the siphon
while the animal is still under water. It is to be distinguished
from ordinary branchial respiration on the one hand and from
direct pulmonary respiration on the other.
A. The first point upon which fresh evidence may be offered
is the function of the longitudinal ridge (‘repli palléal’ ) which
sub-divides the pallial cavity. Jourdain, Bavay and others speak
as though the pallial cavity were divided into two separate parts
by the ridge in question which runs up the floor of the cavity
parallel to and on the left-hand side of the gill and rectum. The
precise role of sucha division is not specified by previous workers ;
but one is left with the distinct impression that this apparent
division of the mantle-cavity into right (branchial) and left
(pulmonary) areas is of importance in respiration. The available
facts, however, a uppeet to indicate that this is not the case. To
seal effectually the ‘pulmonary’ pallial chamber from water or the
branchial chamber from air, the separation of the two chambers
should be complete. But in over twenty examples of various
species and genera I have never found the ridge nearly high
enough along its entire length to touch the pallial roof. Bavay,
342, MR. G. C. ROBSON ON THE RESPIRATORY
on the other hand, says that it fits into a groove in the latter.
It is possible that, in the spirit specimens | have examined, the
ridge is contracted and drawn away from the roof of the pallial
cavity.
In the first place it is unfortunate that we can have no direct
evidence on this subject. In a living animal we are obviously
unable to see whether the pallial roof is in contact with the entire
length of the ridge. We have, therefore, to fall back upon in-
direct evidence. ‘Lhe following facts appear to indicate that the
separation is not complete. The ridge is sometimes quite high
enough anteriorly to touch the roof ae the cavity; but it Alaa:
invar siably decreases in height postericrly so that it is difficult to
see how complete insulation of the two chambers can be ther eby
effected. Secondly, 1 have never found a groove deep enough to
hold the top of the ridge steady against the pressure of water.
Thirdly, I have freque ently found the albumen gland so much
distended that it presses down a portion of the ridge and keeps
the pallial roof away from it, thus opening a more complete
communication between the two cavities. Finally, there can be
no actual need to prevent water from entering the so-called pul-
monary pallial cavity from the side of the gill, as it may be seen
frequently entering it from the left-hand side. It is, perhaps,
from this latter fact that we may derive an explanation of the
function of the ridge. Water apparently enters the pallial cavity
from the left-hand side, passes backwards, and then is passed
forwards over the gill to be finally discharged from the right-hand
side. It seems likely that the ridge functions in euiding the
current of water, and its structure (usually high sauna only and
diminishing in size posteriorly) is in accordance with such a
function. “The criticism will at once suggest itself that similar
arrangements for directing the current “of water in branchial
respiration are not found in the large assemblage of other Proso-
branchs which employ branchial respiration. Difficult as this.
argument may be, it appears to offer less obstacles to a solution
than the fact that a similar ridge is found in Vivipara in which
no lung is found!
The function of the ridge in that genus has been discussed by
Sewell (9), who regards it as ‘respiratory, though he does not.
make its precise role clear. The possibility that ene ridge may
form part of a gutter down which the renal excretions pass to
the exterior should not be overlooked. Bouvier (2) suggests that
it acts in this capacity in Ampullaria, as a result of the displace-
ment of the gill into the area of renal excretion. It should not
be overlooked, however, that the ridge is found in Vivipare in
which there is a fully-formed ureter as well !
B. The next point of interest is whether during siphonal
respiration there is a direct connection between the base of the
siphon on its inner side and the orifice of the lung. It must be
admitted in advance that the evidence to be adduced is mainly
MECHANISM OF THE AMPULLARIID®. 343:
negative ; but as the question is very obscure and at the same:
time connected with the whole question of the respiratory
mechanism, we may attempt to make the issue a little clearer.
Bavay and Bouvier (2 @) imply that the siphon and respiratory
orifice form a continuous tube. If this is the case, the ai
from the exterior passes down the siphon straight into ‘the lung
and none of it is wasted in the pallial cavity. As to the details,
Bouvier is the more explicit. After admitting that during the
act of siphonal breathing it is impossible to see what is going on
in the pallial cavity, he says that afterwards the “plancher pul-
monaire’ becomes visible (i. e. by the modification of the shape of
the siphon and its base) and that it is “susceptible de faire saillie
en avant jusquau voisinage de Ja base du siphon.” The chief
thing to notice is that during the actual process of siphonal
bre athing i it 1s impossible to see » what is going on inside the pallial
cavity. It is a controversial point whether one is entitied to
assume that siphon and vulmonary orifice are continuous at a
given moment because a little while afterwards the latter is fuund
projecting towards the former! It is thus dificult to see on what
critical facts Bouvier and Bavay based their statements. As far
as A. vermifornisis concerned, even if it is seized during siphonal
breathing and the pallial cavity forced-open, the siphon resumes
its normal shape before the base and respiratory orifice come
into view.
A priori one would be tempted to assume that the connection
in question is necessary. It would be physiologically advanta-
geous to have a means whereby the air is conveved straight into.
the lung and not wasted in the pallial cavity. It might also be
desirable to keep air away from the gill. But we may point out
(a) that the siphon is not used at all in direct pulmonary breathing,
so that the question of physiological economy does not seem to be
involved, and (6) that during siphonal respiration the mantle
cavity is kept tightly closed by the down-drawing of the mantle
folds, so that there is no risk of water entering the cavity and
finding its way into the lung.
C. A comparison of the observations upon Neotropical forms
with those published by Ramanan(7) upon the Oriental Pachylabra
globosa and by Bouvier and Fischer upon the Hgyptian Lanistes
bolteniana, suggests that the structural differences upon which
Fischer (5) separated the American, Indian, and African forms
are correlated with differences in the mode of respiration.
Fischer divides the Ampullariidz into the following genera :—
1. Ampullaria s.s. including Marisa (=Ceratodes): siphon
very long. America.
. Pachylabra: siphon rudimentary. Africa, Asia, Oceania.
. Asolene: siphon absent. South America.
. Lanistes: (siphon ?). Africa, Madagascar.
. Meladomus: (siphon ?). Africa.
Ore co bo
344 MR. G. C. ROBSON ON THE RESPIRATORY
It is desirable to know to what extent the differences illustrated
by his classification are to be found in other features of the
respiratory system.
In order to test this I have examined the respiratory organs
of the following forms, all of which are represented in “the
material available in the Zoological Department of the British
Museum :—
% Marisa corinu-arietis (Trinidad).
2. Ampullaria vermiformis (Pernambuco).
: BA glauca (Demerara).
4, 5 insularum (South America, ? loc.).
5. Lauistes afinis (Victoria Falls).
6. » oliviert (Kgypt).
7. Pachylabra sp. (Sikkim).
=) an sp. (Perak).
% gordont (Victoria Nyanza).
According to Bouvier and Fischer (3) Lanistes is less specialized
for siphonal respiration than Ampullaria. The siphon is short
and does not appear to form a complete tube. On coming to the
surface of the water it apparently places its respiratory orifice
in direct communication with the air, and the siphon is not an
indispensable part of the mechanism. A. (=Pachylabra) glauca,to
judge from Ramanan’s account, represents a more advanced con-
dition, but not yet approximating to Ampullariu s.s. There
is true siphonal respiration and apparently, as in the South
American verniformis, the base of the siphon assists in closing
the left-hand side of the pallial aperture. But the siphon is very
short, its orifice very broad, and there is no pumping-movement
of the head such as Bavay and others have described for South
American forms and as Miss Cheesman and I have witnessed in
A, vermiformis. We finally have complete siphonal breathing in
Ampullaria s.s. achieved by a very long flexible siphon.
In Ampullaria and Marisa there is a rather small and sub-
circular or ovoid respiratory orifice situated on the anterior end
of the lung-cavity rather towards the left. Its edges are some-
times thickened slightly and tend to overlap each other. It is
difficult to see whether closure is effected by a sphincter or by the
overlapping edges.
In the two Asiatic species examined and in P. gordoni from
Kast Africa a very different condition is found. The respiratory
aperture is large and gaping, and is not a simple orifice in the
floor of the lung. Generally found more in the middle of the
latter, it extends backwards and to the right—as far as the right-
hand wall of the: pulmonary cavity—as a wide longitudinal slit
between two markedly overlapping edges, the inferior of which
(left-hand) is inserted into the floor of tne! pallial cavity.
In Lanistes the orifice occupies more or less the same position
as in Ampullaria, i.e. anterior, to the left and close to the osphra-
dium, It shows, however, the same general structure as in
MECHANISM OF THE AMPULLARIIDA. 345
Pachylabra, though an individual feature is seen in the develop-
ment of an upraised ridge on the right-hand edge. The orifice is.
almost as long as in Fecha a, but i in the two species examined
it does not extend so far backwards.
It is impossible as yet to discuss the course of evolution in the
Ampullarude. It would seem, however, that the method of
respiration and the structure of the siphon and the respiratory
orifice exhibit a correlated and progressive modification. first
the siphon is short and almost functionless, the respiratory orifice.
elongate. Then the siphon acquires an intimate association with
the left-hand side of the pallial aperture which it assists to close
during siphonal respiration. It increases in length and can
become completely tubular. The respiratory orifice becomes
restricted in size and anterior in position and possibly more.
directly approximated to the base of the siphon.
D. The development of the lung-cavity * from the branchial
chamber in the Ampullartide is comparable to the similar
phenomenon in Lirgus latro and the terrestrial Brachyura and in
certain gobiid fishes, and the amphibious habits of these animals.
are in general alike. Semper (8) and others have dealt with the
causes which may lead to the development of the amphibious
habit, so that there is no need to discuss this matter in detail.
It might be pointed out that in the case of Ampullaria the
development of a lung is apparently not necessarily connected
with the periodie diying-up of the streams and the sheets of water
in which the molluses live. Ampullaria, to judge from numerous
observations, is capable of spending a long period in a state of
reduced metabolism familiar to all students of molluscan esti-
vation. This process is of course facilitated in operculate forms
by the retention of moisture in the tissues owing to the complete
insulation of the animal by the operculum. The methods by
which the Ampullaridze are ‘able to resist the annual periods of
desiccation are, therefore, probably the same as that of other tropical
molluscs. The significance of the development of the siphon is
quite a different matter. The development of the siphon is
probably correlated with an increased tendency to live in streams
covered with a dense mat of surface vegetation in which the
short siphonal forms couid not exist. In drawing attention to a
possible adaptationai value of the siphon, I do not wish to assert
that it was necessarily adaptive in its origin,
I am indebted to my friend Dr. Baini Prashad of the Zoological
Survey of India for permission to state that, so far as Pachylabra
is concerned, his views (arrived at independently) with regard to.
the adaptational value of the long siphon are the same as those
advanced above.
* It is interesting to note that Brooks and McGlone (4), who have studied the
development of the North American A. depressa,iind that the lung-cavity is de-
veloped as an infolding between the ridges from which the gill and osphradium
are developed, all three structures arising from practically the same area in the:
embryonic mantle-cavity.
346
RESPIRATORY MECHANISM OF THE AMPULLARIIDA,
T have also to express my obligation to Miss EK. Cheesman for
facilitating my observations on the animals placed under her
charge and for several valuable suggestions.
i
2.
Works quoted.
Bavay. 1873. Rev. Sci. Nat. ii. (1).
Bouvier. 1888. Soc. Philom. Paris: Mémoire Centénaire.
2igs lb: 1891. ‘Le Naturaliste,’ (2) xcii. p. 143.
CO OG MED Cr
(ml
Bouvier & Fiscuer. 1890. Comptes Rendus Ac. Sci.
Paris, exi. 2.
Brookes & MoGtonsz. 1908. Carnegie Instn. Washington.
Publication 102, p. 95.
Fiscuer. 1887. Manuel de Conchyliologie, Paris, p. 737.
JouRDAIN. 1879. Comptes Rendus Ac. Sei. Ixxxviil. p. 981.
Ramanan. 1903. Journ. Malac. x. 4. p. 107.
Semper. 1883. “Animal Life.” Internat. Sci. Ser. London.
SEWELL. 1921. Records Ind. Mus. xxii. 3. p. 221.
MONKEYS AND THE FEAR OF SNAKES. 347
18. Monkeys and the Fear of Snakes. By P. CHanmurs
Mrrenext, D.Se., LL.D., F.R.S., C.B.H., Secretary of
the Society.
[Received March 21, 1920: Read March 21, 1922.}
When I became Secretary of the Society, tlhe practice of
feeding live animals to snakes was violently attacked, on the
ground of cruelty, and vigorously defended as naturat and
necessary. J enquired into the attack and the defence. ‘The
latter was easily disposed of. JI could not agree that even if
cruelty were part of the order of Nature we were justified in
continuing it under artificial conditions. Nor could I accept the
odd opinion urged on me in particular by the late Dr. Plimmer
that there was merit in the flesh of prey killed by the serpent
itself, absent from prey killed by other means. But the question
was answered by practice. Mr. HE. G. Boulenger, our Curator of
Reptiles, to whom I am much indebted for his patient and
careful observation of my suggestions, is now in agreement
with me that snakes do not vequire a living prey.
The cruelty alleged by the attack is a more difficult, and from
the scientific point of view, more interesting question. So far as
I could see, frogs, rats and mice, rabbits, guinea-pigs, fowls and
‘ducks, introduced to snakes, had no sense of impending fate, and
treated the creatures which were going to destroy them as
indifferent objects. The sense of terror was in the spectators,
notin the victims. But I wished to carry the matter further, and,
with the assistance of Mr. R. I. Pocock, the Society's Curator of
Mammals, made a large series of experiments by introducing
live snakes to different kinds of animals. ‘The results of these
observations have already been communicated to the Society
(P. Z.S. 1907, p. 785), and I have also repeated some of the
actual experiments at one of our Scientific Meetings. Except
in the cases of a few of the higher birds, notably parrots and
Passerines, and the Primates excluding Lemurs, animals show no
special relation of fear in the presence of living snakes, even
when the latter are allowed to approach them closely, protruding
their quivering tongues. But the reaction of fear is immediate
and painful to see when a snake is introduced to a parrot or
starling in a cage, or to any of the true monkeys from the South
American marmosets to the Great Apes.
The instant recognition and betrayal of fear by special sounds
and rapid retreat, even in the case of a lethargic old Orang-utan,
were so plain that I was disposed to treat them as instinctive,
and even went the length of suggesting that the human dread of
snakes might be an instinct inherited from our ape-like ancestors.
348 MONKEYS AND THE FEAR OF SNAKES. ;
But “instinct” is a difficult conception, especially in the mental
field of higher animals where experience plays a dominating part.
For some years I have been waiting an opportunity to see if a
monkey born here and reared without any individual experience
of snakes would display an instinctive dread of them. That
opportunity has not yet occurred. But last year, in January, a
young male chimpanzee was sent to us, the youngest I have ever
seen, and so small a baby that it seemed not much more than
weaned. It thrived in the Ape House, soon lost its shyness,
and became quite lively and intelligent. In the course of the
summer, with Mr. Boulenger, qT took: a very active tree-boa to it.
Mr. Boulenger held the chimpanzee and J had the snake twisted
round my arm, with its head and about 2 feet of neck pro-
truding. I put it cautiously near the chimpanzee, who leaned
out towards it; I allowed the flickering tongue of the snake
actually to touch the chimpanzee’s face; the latter showed no
sign of alarm and was quite ready to kiss the snake’s mouth. It
was clear that, in this case, no particle of dread of the serpent was
present. Since then, the chimpanzee has on several occasions
been shown snakes, ine luding a large King snake. The latter he
handles quite freely, plays with it and pulls it about.
It is possible that this case is an example of failure of the
development of a normal instinct in an individual. This may be
true, and I do not wish to found a-general conclusion on a single
example. But so far as the evidence goes, it would appear that
the dread of snakes is not instinctive in chimpanzees. As it 1s
certainly present in the older chimpanzees on which J have made
the experiment, the alternative 1s to suppose 1t an acquisition due
to experience or the imitation of other chimpanzees which have
had individual experience.
P. Z. S. 1922, CUNNINGHAM, PI. I.
“ Pile” Coloration in Fowls, ¢ and @ L.
a ie ; x
ne
Nuh
P. Z. S. 1922, CUNNINGHAM, PI. II.
Recessive colour in hens from cross of White Leghorn 2
x Black-red ¢
MBENDELIAN EXPERIMENTS ON FOWLS. 349
19. Mendelian Experiments on Fowls. III. Production of
Dominant Pile Colour. By J.T. Cunnincuam, M.A.,
F.Z.S8.
[Received March 1, 1922: Read April 4, 1922. |
(Plates I. & IT.)
Two previous papers by me on Mendelian experiments in Fowl
breeding have been published in the ‘ Proceedings of the Zoological
Society.’ The last, in the volume for 1919, described the pro-
duction of a recessive pile from a cross between a Silky hen
and a bankiva cock. The present paper records the results of a
cross between a dominant white, namely White Leghorn, and
Black-red Game, made with the object of finding whether a
dominant pile would result.
In March 1919 I purchased from a Yorkshire breeder a Black-
red Game cock, which was stated to be pure-bred, and mated it
with two white Leghorn hens kindly supplied by Mr. Seth
Snith from poultry in the Society’s possession. From these
parents two broods of chickens were hatched—the first on April
27—28th, the second about May 10th. There were eight chicks
in each brood.
The colour of the down of these chicks was canary yellow,
darker round the neck, and diminishing to white behind and
below, but none were pure white all over. There was no sign of
stripes. Bateson (‘ Principles of Heredity,’ 1909, p. 102) states
that the F, offspring of White Leghorn by Indian Game or
Brown Leghorn is, when newly hatched, white with a few specks
of black. There may be some breeds of fowls which are pure
white in the down, but all that I have studied are not white but
yellow or fawn colour. It is well known that young ducklings of
white breeds are yellow in the down. Several of the chicks of
my cross had one or more black spots and others had black and
red specks on the head, but I was not able at the first examination
to take them up and examine them closely. The absence of
striping is Interesting. The chicks in the down of the Japanese
Long-tails which I bred some years ago, and, I believe, the chicks
of Black-red Game and of Game fowls, generally have a median
dorsal dark brown stripe and two lateral stripes of the same
colour, the rest of the down being fawn colour or buff. The
recessive or non-coloured chicks from the cross of Silky by G.
bankiva had yellow down on the body generally with white
stripes in place of the dark stripes of the coloured chicks.
According to Bateson the chicks of pure-bred Silkies have the
same marking as this. It would seem, therefore, that in the
cross White Leghorn by Black-red the dominance of the white
Proc. Zoou. Soc.—1922, No. XXIV. 24
350 MR. J. 1. CUNNINGHAM ON
prevents the dark striping of the chicks, though it does not
prevent the occurrence of spots and sp ecks of black or red. On
the other hand, Bateson (loc. cit. p. 120) states that in the down
the chickens os Pile fowls have longitudinal stripes of light
chestnut.
On June 4 I made a more thorough examination, but the
chickens were wild and difficult to catch, and on reading my notes
J found I had only handled 13 out of 16. The first brood were
now almost fledged, the second about half-fledged, so that the
colour recorded now is that of the first plumage “of true feathers.
The condition observed was as follows :—
Tn three no colour was seen: feathers all white.
Four had much brown colour on back, wings, and breast.
One had numerous scattered black feathers.
Five were white with one or more black specks.
The birds were examined thoroughly about six weeks later,
namely on July 16 when they were completely fledged, and the
sex was recognisable from the size of the combs, althou gh the tails
of the cocks were not fully developed.
Cocks, total number 8.
Four of these may be described as imperfect piles; their
coloration was :—
Numerous red patches on the saddle and backs of the wings,
slight yellow on neck hackles, and very slight on loin hackles ;
the rest of the plumage white. Legs white with yellowish
tinge.
The other four had much less colour, and some of them had
more black :—
5, Same marking, but golden yellow instead of red; only a
few red feathers on the wings.
6. Black spots on saddle and back of wings, one black spot on
the breast, and oneon neck. Slight golden yellow on loin hackles.
7. Only a few small red patches on back of wings, but three or
four single black feathers; also a tinge of gold on saddle.
8. Black and red ticks on back of wings and back, more black
on the right side.
Hens, total number 7.
Four of these had a great deal of reddish-brown colour, different
from that of the cocks, which was a brighter red. This colour
was ‘laced ” all over back, and saddle hackles, more continuous and
uniform on back of wings. There was uniform brown on throat,
breast, and abdomen, but pale, ¢.e. diluted with white. Only the
end of the tail was quite white. (PI. I. fig. 2.)
This coloration agrees fairly well with the description by
Mr. Douglas of hen piles in Wright’s ‘ Book of Poultry, 1885 :—
“Head ieee golden chestnut; hackle white, laced with yellow
chestnut; back a ne white ground-colour, slightly laced, a
shade of | gold prevailing; salmon-coloured wing-coverts almost
MENDELIAN EXPERIMENTS ON FOWLS. 351
similar to back, but just a little heavier in the dark colour; breast
a rich chestnut right up to the throat, running off to a white, but
not pure on the thighs; tail almost a pure white.”
& and 6. No brown but scattered dark specks, many of the
feathers in these with minute points of black all over,
appearing grey. No. 5 with more black than No. 6 but of
same type.
7. Deep black isolated feathers on wings and back, with slight
trace of red, uniform over back.
One chick had been lost.
It will be seen, therefore, that on July 16th in these I, birds
there were two groups—one of four cocks and four hens of the
pile type, and the other of four cocks and three hens with small
scattered spots of black and red. No. 5 among the cocks belonging
to the second group is more similar to the pile, but with colour
much less developed.
Subsequently the red colour in the cocks increased considerably,
and the pile coloration was more completely developed. Thus on
Sept. 27 the birds were nearly full grown and with fully-developed
plumage. There were six cocks that looked like fairly typical
piles, 7. e. white birds with rich red backs. In addition there
was one with similar marking, but straw colour on the back
instead of red, and another with rather smaller comb and wattles
and no broad area of colour on back, but separate spots, and a lean
and game-like shape. The four pile hens were much the same as
before, and the other three hens still had only scattered specks of
black and yellow.
In December one of the pile cocks was accidentally killed. The
last of the cocks described above, which was more backward in
development than the others, had by this time developed the pile
coloration like the rest, but with pale yellow colour in the back
and saddle, not red.
Thus the F, generation in this experiment shows that the pile
coloration in the cocks is a heterozygote. It may be said that
the white of the Leghorn is dominant to the black in the black-
red, but not to the red: hence the heterozygote is white-red
instead of black-red. The dominance to black is not, however,
quite complete, as black specks or occasional black feathers occur.
In the hens the pile coloration takes a different distribution,
corresponding to the different distribution of black in the hen of
the black-red.
F, Generation. Year 1920.
In Feb. 1920 I mated the F, fowls in two separate pens for
breeding. In one pen were put one of the most deeply coloured
pile cocks with the four typical pile hens. One of the latter died
before breeding. In the other pen were one of the palest pile
cocks with three hens which were white with spots and specks of
colour. A number of eggs, about 25 from each pen, were obtained,
94%
Pai
352 MR. J. T. CUNNINGHAM ON
and hatched in incubators about April 14th. The chicks were
examined on April 17th, with the following results :—
A. From the pile parents there were 16 chicks, 13 of which
were yellow without spots. The other three were coloured.
(1) Bluish primaries and rump, reddish back, sides and belly
yellow.
(2) Smoky biack on head and back, and on wing-primaries.
(3) Smoky black on back and head.
B. From the pale pile cock with white hens there were 19 chicks :
of these 11 had yellow down apparently without colour,
and 8, including one that died after hatching, had some
colour. Of these 8, four had one or two black spots and
four had more contimuous colour, the darkest being the
dead one, which was dark smoky black over the head and
the whole of the back and wings.
The other three were as follows :—
(1) Slightly greenish on rump.
(2) Slightly bluish rump, head ditto, otherwise yellow.
(3) Back dark reddish combined with blackish colour, slight
black spot on head.
If we consider these lots separately, the theoretical expectation,
on the supposition that the black-red colour of the original male
parent was a single character, would be one-fourth black-red in F,,
the black-red being recessive. Thus in the chicks from Pen A,
16 in number, there should have been 4 coloured: the actual
number was 3. Of the chicks from Pen B, 19 in number, one-
fourth would be between 4 and 5, or very nearly 5. The actual
number was 4 if we include only those which had more or less
continuous colour. If we take the two lots together’, all the F,’s
being heterozygote, the number of coloured recessives was 7 out
of 35. It would be expected, however, that black-red recessive
chicks in the down would be brown with black stripes, like the
black-red dominants in my cross between bankiva and Silky.
There were none at all like this.
On October 9th I made a careful examination of these birds of
the F, generation, which were then in mature plumage, and their
characters were :—
A. From pile parents.
7 Cocks, all showing the pile coloration, but with different
degrees of intensity in the colour on the back, ranging from pale
yellow to red.
1. Deeply coloured pile; legs yellow. A photograph of the
skin of this bird is shown in PI. I. fig. 1.
2. Similar but not so deep a colour ; legs yellow.
MENDELIAN EXPERIMENTS ON FOWLS. 353
3. Pale orange on back and hackles, with a few black specks ;
legs pink.
4, Pale pile; legs yellow.
5. Pale pile ; legs pink.
6 and 7. Very pale pile; both with legs pink.
6 Hens. Two of these were coloured all over; three had the
pile coloration, 7.e. feathers laced with fawn colour on the back,
the breast and abdomen uniform pale brown ; one was white, with
trace of reddish lacing on the neck.
1. Almost entirely black, with white edging to feathers on
wings and breast and very slight yellowish lacing.
@EIEE eters] 22)
. Grey all over, with slight orange lacing: legs not noted.
(Bl’ DS figs 1.)
3. Strong dark pile coloration of hen type; legs pink.
4, Good hen pile, well laced ; legs pink.
5
6
bo
. Pile lacing rather pale; legs pink.
. White with trace of reddish lacing on neck; legs pink.
B. Offspring of cock with slightly yellow back, and hens white
with black specks. .
8 Cocks. One of these was a moderately coloured pile ; six were
pale or very pale piles; and one was not a proper pile, but had
spots of orange and black on wings, hackles, and tail. Four of
them had yellow legs and four had pink legs.
9 Hens. Two of these were typical piles, with pale back laced
with fawn colour and brown breast, one rather more deeply
coloured than the other. One was reddish brown all over with
black specks. The rest, six in number, were white with scattered
specks of colour, and two of them hada yellowish tinge on head
and neck-hackles. I give below the notes for each of these
individually :—
4. White with minute specks of black.
5. Yellow on headand hackle ; back and tail with many black
spots.
: Galen, on head and hackle; black specks on back.
. White with a few grey marks on feathers.
. Very slight specks of black here and there; otherwise
white.
9. White with few black spots.
@~Ilo
Five of these hens had pink legs and four had yellow legs.
I have not paid special attention to the inheritance of the
colour of the legs in this experiment, but will merely point out
the leg colours noted in the F, generation. Yellow leg means
yellow skin, and what I have called pink leg means white skin.
The yellow skin was a character of the White Leghorn hens of
the parents in the cross, the white skin a character of the Black-
354 MR. J. T. CUNNINGHAM ON
Red Game Cock. According to Bateson, white skin is dominant
to yellow, and I believe this was the case in the F, generation in
this experiment. In the F, generation, omitting ones specimen in
which the colour of the legs was not noted, there were 19 pink
legs to 11 yellow. The “number of recessives was therefore
considerably in excess of the expected number, namely 11 yellow
to 33 pink, or 1 to 3. ‘This is, of course, in a total of 30
individuals of no special significance.
It is evident from these results that the Pile type of coloration
is, like the Andalusian, a heterozygote and not a pure character
which breeds true, and that it results from crossing the dominant
white of the White Leghorn with the Black-red of the Game.
Fanciers have stated this before, but they have not recorded the
individual results nor the segregation of the F, generation. Thus
Mr. John Douglas in Wright's < Book of Poultry,’ 1885, states :
“You can also get a very rich Pile by putting a Wheaten hen to
a White or Pile cock.” The Wheaten hen is one of the types of
the hen of the Black-red Game. Mr. Douglas also says that Piles
breed true to colour, but that now and then a cross of the Black-
red is thrown in to give hardness of feather. My results are not
in agreement with thisstatement. Mr. Fred Smalley, of Silverdale,
Lancashire, who bred Pile Game for many years, kindly answering
enquiries from me in 1913, wrote that Pile was dominant to
Black-red, and that when bred together, Piles worked out to too
much washed-out colour, but could never produce pure white.
For this reason they were crossed with Black-red once in seven
years. He also informed me that a cross of White Leghorn hens
with a Brown Leghorn cock, or White Wyandotte hens with
Partridge Wyandotte cock, would produce birds of the pile
coloration, though they might not be up to standard Pile colour.
In the F, generation, however, the heterozygotes were not all
alike, but varied considerably in the amount of colour shown—in
other words, in the degree of dominance of the white character of
the maternal parent. The 8 cocks all ultimately developed the
pile marking, six of them having red colour on the back of
varying intensity, and two having only yellow instead of red, one
having straw-colour and the other pale yellow. Of the 7 hens,
four were fairly typical Piles, the other three were white with
scattered black or grey feathers.
The two groups of F,’s mated to produce the second generation
both produced some piles, as might be expected since the birds of
EF, were all heterozygotes. The birds of F, would be expected to
produce pure dominants (DD), 7. e. white, piles (DR) and black-
red, 7. €. recessives (RR) in the proportion of1:2:1. The deeply
coloured hens in the F, generation are the recessives: unfortunately
there were no deeply coloured, 7. e. black-red, cocks. The three
coloured hens are all very different from each other, and this is an
interesting fact. If we regard the black-red coloration as a single
character, a recessive allelomorph to the dominant white, we should
naturally expect recessive individuals to be closely alike, whereas
MENDELIAN EXPERIMENTS ON FOWLS. 305
here we have one black, one grey, and one reddish brown. The
recessive character therefore appears in different degrees in
different individuals, and the fact that the black hen has some
white on its feathers shows that the segregation is not complete
and the recessive character not perfectly pure.
The pile birds, both cocks and hens, were soon after Oct. 9th
separated from the rest, and on Nov. 20th I examined the hens
which were not pile, in order to satisfy myself whether there
were any pure dominants, 7. e. white without specks of colour.
Of Lot A there were none without some lacing to the feathers, as
seen from the observations of Oct. 9th, given above. Of the six in
Lot B which were not deeply coloured nor pile, there was not one
without some trace of colour. Three of them showed slight
lacing of colour on the neck-feathers, and all except one of these
had a few black or grey specks.
According to these results, then, there were no pure dominants
either in Lot A or Lot B of the F, generation. Two hens in
Lot A had very pale lacing. In the cocks of this lot there were
all degrees of intensity in the red colour of the pile. In Lot B
there was one cock which was not a proper pile, but had spots of
orange and black on its back, and six hens with either a trace of
lacing or some specks of colour. If these six hens in Lot B are
to be regarded as pure dominants, we have six out of 17 or nearly
one in three instead of one in four. This may seem a good
approximation to the theoretical proportion, but it is unlikely
that all the dominants should be female. The fact, however, is
that none of these birds were without evidence of colour, and
therefore it cannot be said that there was complete and clear-cut
segregation.
As I was unable to make many further experiments with these
birds of the F, generation, I decided to mate some of the hens
with Jeast colour with another black-red cock to see if they
behaved as heterozygotes or as pure dominants. Accordingly
two of the F, hens above described with least traces of colour
were mated in March 1921 with a bankiva (black-red) cock which
was among the birds at that time alive in the Zoological Gardens.
From this mating nine offspring were produced and reared. As
there were traces of colour in the female parents, and some of the
F, hens were also white with black ticks, I thought it possible
that these F, hens would behave as heterozygotes, and not as pure
dominants. I therefore rather expected that half the chicks
would develop into piles and half into black-reds. All the nine
chicks obtained, however, developed in mature plumage into fairly
typical piles. There were five cocks and four hens, only one
cock and one hen having somewhat paler colour than the rest.
Although the number of chicks obtained is small, I am of
opinion that the F, hens behaved genetically as dominant: if they
had behaved as heterozygotes, I think there would have been at
least one, and probably more than one, black-red among the
offspring, the theoretical explanation being, of course, half the
356 MR, J. Tt. CUNNINGHAM ON
number piles and half black-reds. At the same time it is to be
noted that the pile coloration in these birds was more pronounced
and more uniform than in the heterozygotes of F,, whereas if
the F, mothers had been pure dominants, there is no reason why
the offspring of F, DD’s with the RR black-red should have any
more colour than the DR’s of F,.
If we regard the dominant white as one character and the
black-red colour as its allelomorph, the evidence of these
experiments seems to show both imperfect dominance and
incomplete segregation. The pile produced by my cross, and
probably the pile of fanciers also, is a heterozygote, like the blue
Andalusian, and it should produce in the F, generation whites,
piles, and black-reds in the proportion 1: 2:1. In F, the white
is not acomplete dominant; it suppresses the black colour almost
but not quite completely, but the red colour was developed in
varying degrees in the piles, eight cocks and four hens, while the
other three hens showed only scattered specks of black and red.
In the F, generation both groups A and B were the offspring
of heterozygotes, the parents of A being of the pile coloration,
those of B showing greater dominance of white. ‘The two groups
may be compared thus :—
F, Group A. LI, Group B.
Recessives. Recessives.
2, one black and one grey, each with 1 9 reddish brown with black specks.
some white feathers.
Heterozygotes. Heterozygotes.
5 & deep pile to pale pile. 3 g: one moderate pile with a few black
3 © dark pile to rather pale lacing. marks, two pale piles with spots of
black, or orange and black.
4.92: two typical piles, two yellow on
head and hackle, with many black
spots.
? Dominants. * Dominants.
5 go very pale piles, two with only a
slight tinge of yellow on back of
wings, perhaps not more than occurs
occasionally in Leghorn cocks.
4.9 white with scattered specks of
| black or grey.
2 very pale pile.
i % white with trace of reddish lacing in
‘neck.
|
But while, as this table shows, the recessives can be definitely
distinguished from the heterozygotes, it is not possible to separate
the pure dominants from the heterozygotes. In the whitest birds
there are some traces of colour, and the amount of colour forms a
continuous series from the typical pile cock and hen to the birds
with least colour. In the recessive hens also there are distinct
traces of white, The segregation therefore is imperfect.
MENDELIAN EXPERIMENTS ON FOWLS,. By!
In the second cross with G. bankiva g the F, hens with least
colour were used, and, as mentioned above, the offspring were all
pile, i.e. showed more pronounced coloration to a more uniform
degree in the different individuals. It may be suggested that this
was due to mating with a black-red of different race, but [ think
it shows that the F, white hens had more colour in their heredity
vhan the original White Leghorn hens. On the other hand, the
genetics may be more complicated than merely the meeting of a
single pair of allelomorphs, dominant white and black-red.
Bateson in 1909 concluded from his own breeding experiments
that the black-red colour was due to two complementary factors
X and Y, and that the dominant white of White Leghorn was
due to a suppressing factor 8 for which birds of that breed are
homozygous. One or both colour factors may be present in the
White Leghorn, but even when both are present, colour is
suppressed by the factor 8.
Some dominant white are of the genetic constitution Xx Yy
SS: the factors of the Black-red Game may be assumed to be
XXYY. My cross will then be :—
XG Vy SSX OY:
Gamletes|j senate WOXCS YES WoNCO VXI OG NO NA.
By dened sect ce betes XYXYS, XYYxS, XYyXS, XYyxs.
All these combinations contain both factors for colour, t.e. XY,
and all contain only one suppressing factor 8. This would help
to explain why in F, colour is not entirely suppressed, but it
does not explain w hy the black is suppressed more than the
red. It would also help to explain why the degree of coloration
is different in the F, birds, for in one combraeen both X and Y
are double, while in the other three XY together are present only
once, nigh either an additional X, an saglGbiis mel. NZ or neither.
The birds of constitution XYXYS then may be the typical piles,
and the others the palest cocks and white hens of F,. The
difference, however, was not sharply marked.
Again, one or both of the White Leghorn hens in my first cross
may have been XxyyS. In this case the cross could be analysed
thus :-—
XxyySS x XXYY.
Gamietes 458.2. 65: XyS, xyS xX XY.
PSR UNO RAW. oP XYXyS, XYxy8.
There would be only two combinations, and each would have
the compound XY only once, or in the simplex condition as in the
two last combinations in the former case. There would probably
then be no typical or deeply coloured piles. It is more probable,
therefore, that in my cross the constitution of the hens was
XxYy8S8. Ido not know whether a White Leghorn of consti-
tution XYXYSS could occur, and therefore wlll not consider it
here,
358 MR. J. T. CUNNINGHAM ON
We may suppose that the mating of the F,’s may be represented
thus :—
OROUP eA en ernie cre XY OVS xX OS,
Gigovbl oad o Ph ee MRE KY VS xX Vy Xs.
Group A, gametes... XYS+XY x XYS + XY.
Hd, . anata tay ee XYXYSS+ XYXYS+XYXYS+XYXY.
The first should be pure white dominants, the next two typical
piles, and the last recessive black-red, the suppressing factor
being absent.
Group B, gametes... XYS+xYS+ XY+xY x XYS+ XyS+
XY + Xy.
1 UR aad tera PA le cg XEXTYAVISS; KON Vy Sos MOVIN S| KONG Woo:
2XXYYS, 2XXYVyS8, 2XxVYS, 2xXyYS8.
XX VIXUYG SNORING, BROW OXGye aXayV.
Thus there would be four combinations of Xx and Yy, namely
OWN PONCE DCN PING SONG 5
and these would be combined with SS, 8, or absence of S. There
would be, therefore, in 16 individuals four coloured recessives of
varying composition, four pure dominants, and eight heterozygotes
or piles. This agrees fairly well with my results, if we assume
that the different combinations of the colour components affect
the degree of visible colour.
Tt is evident, however, that the combinations which are
recessive for S ought to show no more white than did the
original Black red, whereas in my specimens the three recessive
hens show distinct evidence of white: this is more in accordance
with the assumption of imperfect segregation.
In the second cross the hens with most white and least colour
were mated with another black-red. If we suppose that the
white hens had more of the colour factors than the original
Whites and were of constitution XY XYSS, then the cross would
be
ONOSOMISIS) SS DOVE.
Gametess. gee. eee nae XGVIS Anni XN
I EAROASLID A Nay gunnadgee 5 XGVOXONES:
Thus all the piles in this case will be homozygous for the
combination X Y, whereas in the first cross many of the piles were
heterozygotes for this combination.
Lastly we may consider the possible genetic combinations in
the fancier’s process of crossing pile with black-red to improve the
colour of the former. The cross might be
XYXYS x XYXY.
Gametes .......1, YS OX WIS EXO;
LOWRIE cance XYXYS + XYXY.
MENDELIAN EXPERIMENTS ON FOWLS. 359
In this case the piles would be as before, and there would be no
improvement.
But the pile might be
MOYES >= DOYS
Gametes...... XOVS = eViSi == EXO =a 5 OXOVE
leony Ga, | NOOSE DONG AVS) 2b OD ONE ROE
Therefore half the Piles would be improved by being now
homozygous for the combination XY instead of heterozygous.
At the same time the effect on the Black-red recessives from the
cross agrees with what fanciers say of the inferiority of such birds
to the pure-bred Black-red, for half of these recessives are
heterozygous for the XY combination.
EXPLANATION OF THE PLATES.
Prater I.
Fig. 1. Fo g, Pile cock from cross of White Leghorn hens with Black-red Game
cock. Killed March 1921, hatched April 14th, 1920.
Vig. 2. Fy) 9, Pile hen, rather deeply coloured: back light brown or fawn colour,
“laced” but not quite regularly, some white at base of each feather. Neck
hackles white, with orange edges to feathers (“lacing”) ; abdomen uniform
pale brown.. Killed Dec. 1920, hatched April 28th, 1919.
Prats IT.
Wig. 1. F2 2, coloured recessive. General colour diluted black, i.e. grey. » Neck
hackles laced with pale yellow ; feathers of back dark grey, laced with pale
fawn; tail dark grey; abdomen like the back, but much paler. Killed
March 1921, hatched April 14th, 1920.
2. F, 2, coloured recessive. General colour sooty black, with no red. Neck
hackles black, with streaks of light brown; back mostly black, with
slight lacing of pale brown or drab; breast and abdomen pale grey; some
white on wings. Killed March 1921, hatched April 14th, 1920.
eel
ug
&
ON THE LAND CRAB, CARDISOMA ARMATUM. 361
20. Observations on the Land Crab, Cardisoma armatum,
with especial regard to the Sense Organs. By Miss L.
HK. Curssmayn, F.E.S., Curator of Insects to the Society.
[ Received March 18, 1922: Read April 25, 1922. |
As little has been written on the habits of this group of
Crustaceans, the following observations on nine specimens from
the River Gambia, although made under the artificial conditions
of the Caird House for Insects, appeared worth recording.
Sight.—The eyes on their pedicels only partly fill the large
sockets. They function ineffectually in daylight, when the crab
appears to rely for guidance solely on the sete with which the
whole body is amply provided ; but in twilight the crab is able to
focus on a point with tolerable accuracy.
The third pair of maxillipeds are lined at the apex with very
soft hairs which are kept moistened; this organ is supposed to
cleanse the eyes from particles of grit etc., but is also used directly
they are exposed to a strong light. Miss Rathbun * mentions, in
reference to this species, that direct exposure to the sun is fatal to
them. This, of course, refers to the tropical sun of West Africa.
When taken out into the sunlight from the subdued light of the
Caird House in July the crabs used their maxillipeds with great
energy during the first quarter of an hour, they then submerged
in a tank of water—doubtless to moisten the respiratory surface
of the branchial cavity—and afterwards seemed indifferent to the
light, although exposed to it for move than an hour. Nor could
they be induced to use their maxillipeds again on that occasion,
even when the eyes were purposely sprinkled with water and with
dust.
An experiment with flashlight was carried out in the Caird
House. The crab which was exposed to the sudden strong light
was sitting in the water in front of the bay, but left the tank
with unusual speed and precipitated himself to the far end of the
bay, where he assiduously brushed his eyes with both maxillipeds
at once for several minutes. Later he was brought back for a
repetition of the same experiment ; he submerged at once in the
tank, and remained below water with only the eyes showing until
the flash, when he scrambled out and repeated the former
maneeuvres precisely. ‘The sudden transition from a dim to a
very strong light appeared to occasion him real distress ; but the
switching on of the electric lights in the bay after dark, though
it will arrest the crabs’ movements temporarily, does not
necessitate the use of the maxillipeds.
* Bull. Amer. Mus. Nat. Hist. xli, 1921, p. 459.
362 MISS L. E. CHEESMAN : OBSERVATIONS ON
Sense of Hearing.—No sign of response to any sound could be
obtained. During the experiment with flashlight mentioned
above, it seemed unlikely that the report of the concussion was
noticed ; it was the light alone which had a disturbing effect,
because two of the crabs were at an equal distance from the
light as that specimen which was being experimented upon, but
the former were shaded from the glare by a piece of cork-
bark. The two which were exposed to the sound but not to the
light were carefully observed during the reports but made no
movement,
Sense of Taste and Smell.—The seat of this sense must be inside
the mouth, possibly on the mandibular palp; stones, morsels of
brick, and upon one oceasion seaweed, were taken into the mouth,
and it was fully two minutes before they were rejected as
inedible. The former could be heard distinctly to grate against
the mandibles. The antennules are not used in connexion with
food; they are principally used when the crab is under water,
and may test the salinity of the water.
Sense of Touch.—Sete are scattered over the limbs, and as the
crab moves sideways some of these are continually in contact
with the ground and reveal the nature of the surface and the
presence of food. Although the crab is undisturbed by any
vibration of the air, there is immediate reaction to a vibration of
the ground. If a smart tap be given to the wooden partition
of the bay when the crabs have their backs against it, they will
respond to each tap by swinging the eyes into the sockets; or
will move away as though alarmed. he tufts of sete at the base
of the claw of the first pair of walking legs are in constant
requisition while the crabs are feeding, to touch the food before
it is conveyed to the mouth.
Further Observations on the Habits.
Food.—Dead leaves and twigs form the main food of this species
in captivity. They also eat fresh fish and mice. They refused
stale meat, but one night a mouse which had broken into the bay
was caught by a crab, and by morning was half devoured. They
were then supplied regularly with dead mice, and averaged about
one in every three days. After nearly eight weeks they refused
this diet and returned to vegetarianism, broken by an occasional
meal of fresh fish or an occasional mouse; but they prefer all
flesh food very fresh. It was during this surfeit of mice that
their attitude of toleration towards one another was renounced
for combativeness, and five specimens died of injuries caused by
entire limbs being severed from the body. With the change of
diet this pugnacious mood subsided, and the chief aggressor now
lives peaceably with two smaller specimens.
Method of Catching and Eating Mice.—The actual capture of a
live mouse has not been witnessed. When two live mice were
put into the bay one evening they showed a contemptuous
THE LAND CRAB, CARDISOMA ARMATUM. 363
disregard for the crabs. But though their agility afforded them
an easy escape when a lunge was made in their direction, they ran
over and under the crabs with a recklessness which brought them
constantly into position to be seized by the claws, and this is
doubtless how their capture is effected.
When devouring a mouse the crab begins with a leg or the
tail, touching it with the setee already alluded to and putting it
into position with the small claw. The whole mouse is gradually
drawn into the mouth and no part of it afterwards ejected.
Water.—Fresh water for immersion was preferred to artificial
sea-water, even when this was diluted until only slightly brackish.
A tank of natural sea-water was provided, and they bathed in this
with evident pleasure, but by the next day they showed a
preference for fresh water. When a week later a tank of sea-
water was again introduced they treated it with indifference
“Tae ah hag Ra a Ute oats chet
Sil ghae ged ade ri
Syd QonPR Amd Ob eh i.
ti,
\\
4 tt ,
eas, ths Shae ici pines
May: eran Wes | PNR or bai
Ca) KONAE ae Oe. peer eg ny ass wii’ Ne
ene TA Biota | pete wa Pile i eae fee Sivan
D iy aanaie eis wk Led apa
Oe ie yt
i J sala’ (ie: veah
% ne. MER? at Bie ie om si abi eae Abbie a wy Pat pike
ot GY 1 ee 8 ch pe / Diy A eS ; ot
- saan VAh SAN Sachs ete TH Tn rt tie 4 in Mas ‘5 fiyritinine i ie ta
Tyee ts a ae hale heb hacigh ie oe. aa Birla dak yes a,
Wis RETA WAY 0 thane tab: IRE EG A ih | Se a weit i, tan lal
Ti aie tei ye ea Lh iat m4 weihiy Wa ies Ts ‘xt a ee
Fiery Pie SDs pba Fea Ee ine i pals i ft i i i
aa ht See ss aa a 7 mia a
Si eo ee ae “Vipdans AN iy nwdoas he AR Readies,
nid, Coes halo 8 bs AY me jie adel ta eae ee A
Dante, el eit ivi Hal Hi ee fly ela
j a Wea an & eaten alah ey er Maine your mi
; tek WR ph i
yey Hho) eee .
ako herp liens PN
* et bind fae
4,
bey il mia bhi rete me has ey em 91% Liisa Fe ;
hs ah Se Saha bet fii te Ph Med i ty ae .
Mise Nile eecaitret ie
2 : r any barivoiead 4A) ee H
ee 2 De re itt pay
aha iM yaa bod a, aang Ne ry
wt
EXTERNAL CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 365
21. On the External Characters of some Hystricomorph
Rodents. By R. I. Pocock, F.R.S.. ¥.Z.8.
[Received February 7, 1922: Read March 21, 1922. ]
(Text-figures 1-28.)
, ConTENTS. Page
Inrtrrodie tions: ue one datee wee es I eee 3G
Eubtverrannh oui). £ eet ed ae ate Best
MacialhVaibrissea: Oi bs. 0 ee ee ee a ee eT
Mouth and Cheek-pouches..............0...-.....0s0...... 374
We ete mss. Pests os ed h on cack: ee le IRS
Genital @Oreansyots alee ene ee een)
GenitallOrsansrotslem~ale spe assent meneANTICS
Notes on Families and Subfamilies ..................... 422
Introduction.
The classifications of the Hystricomorpha in English text-books
of Zoology are based upon the one proposed by Alston in 1876
(P. Z.8. 1876, pp. 90-97), which was itself an amplification and
in some particulars a modification of the arrangement suggested
by Waterhouse in 1848. Alston added to the group the family
Dinomyide, which, following Peters, he placed between the
Dasyproctide and Caviide; and the Ctenodactyline, which he
ranked as a subfamily of Octodontide. He also transferred
Petromys from the Echymyina (Echinomyine), where it was
placed by Waterhouse, to the Octodontine.
His system, adopted to the letter by Flower and Lydekker
(‘Mammals Living and Extinct,’ pp. 484-491), was as follows:—
HyYsTRICOMORPHA.
1. Octodontidee.
Ctenodactyline (Ctenodactylus, Pectinator).
Octodontine: (Petromys, Ctenomys, Schizodon, Spalacopus,
Octodon, Abrocom«a).
Echinomyine (Carterodon, Myopotamus, Cercomys, Lon-
cheres, Mesomys, Hchinomys, Dactylomys, Plagiodontia,
Capromys, Aulacodus).
2. Hystricide.
Sphingurine (Chetomys, Sphingurus, Eirythizon).
Hystricine (Atherura, Hystriz).
3. Chinchillidee (Chinchilla, Lagidium, Lagostomus).
4. Dasyproctide (Dasyprocta, Calogenys).
5. Dinomyidee (Dinomys).
6. Caviide (Cavia, Polichotis, Hydrocherus)
Proc. Zoou. Soc.—1922, No. XXV.
bo
Or
366 MR. R. I. POCOCK ON THE EXTERNAL
Winge (E Museo Lundi, i. pt. ii. pp. 126-135, 1888) differed
from Alston in some particulars. He referred all the Hystri-
comorphs to one family Hy which was divided into the
following subfamilies :—
(1) Octodontini, equivalent to Alston’s Octodontine + Hehino-
myine ; (2) Capromyini, for Aulacodus, Capromys, Plagio-
dontia, and Myopotamus ; (3) Ctenodactylini, tor Cteno-
dactylus, Pectinator, and Petromys; (4) Hystricini and
(5) Eriomyini, equivalent to Alston’s Hystricidee and
Chinchillide respectively; and (6) Dasyproctini, embracing
all the genera referred by Alston to the Dasyproctide,
Dinomyidee, and Caviide.
Thomas (P. Z. 8. 1896, pp. 1024-1025) made the following
changes in Alston’s system. Following Winge, he transferred
Petromys from the Octodontine to the Ctenodactyline, and
removed Capromys, Myocastor (Myopotamus), and Thrynomys
(Aulacodus) from the Kchinomyine to form the subfamily
Capromyine. He also altered the name Echinomyine to Lon-
cherine, gave full family-rank to the Old World and New World
Poreupines respectively, calling the former Hystricide and the
latter Erethizontide and dividing this family into two subfamilies
—Cheetomyine for Chetomys and Erethizontine for Hrethizon and
Coendu (Sphingurus).
Tullberg (Nova Acta Sci. Upsala, (3) xviii. pp. 82-149, 1900)
introduced some innovations, notably the institution of two new
families—the Aulacodide for Aulacodus (Thrynomys), previously
associated with Capromys and Myocastor, and the Petromyide
for Petromys alone, severing the latter both from the Kehino-
myide and the Ctenodactylide. He refused, moreover, to admit
the affinities of the Ctenodactyline with the Hystricomorphs,
classifying them with his Myomorph + Sciuromorph division of
Rodentia. He fused the two families Dasyproctide and Cavide
of Alston into one called Caviide, keeping Dinomys provisionally
as the type of a special family till more is known about it. The
family-name Octodontide he changed to Echinomyide, and
altered Capromyine to Myopotamini.
Beddard’s_ classification (‘ Mammalia,’ pp. 487-502, 1902)
differed from Thomas’s in restoring Petromys to a place in the
Octodontinz, where Alston put it, and in granting family-rank
to the Ctenodactyline genera, Ctenodactylus, Pectinator, and
Massouteria.
Weber (Die Siiug. pp. 505-507, 1904) admitted no subfamilies,
placing the genera in seven families as follows :—(1) Ctenodacty-
lidees (Ctenodactylus, Petromys, Pectinator); (2) Octodontide
(Ctenomys, Octodon, Abrocoma, Loncheres, Echinomys); (3)
Capromyide (Capromys, Myocastor, * Thrynomys); (4) Chin-
chillidee (Chinchilla, Lagidiwm, Lagostomus); (5) Caviidee (Calo-
genys, Dasyprocta, Cavia, Dolichotis, Hydrocherus) ; (6) Erethi-
CHARACTERS OF SOME HYSTRICOMORPH RODENTS, 367
zoutide (Hrethizon, Coendu, Chetomys); (7) Hystricide (Aystria,
Atherura, Trichys).
Of the above-quoted papers Tullberg’s is by far the most
comprehensive. He dealt as exhaustively as the material at his
disposal permitted with the skull, skeleton, and internal anatomy,
and, in addition, described and figured the feet of several of the
genera, but dismissed very briefly such external organs as the
ears and rhinarium. I have been able to supplement his account
so far as the organs investigated are concerned, by the exami-
nation of some genera he did not see; and in a few particulars
connected with the male external genitalia my results differ from
his. Winge also figured the feet and the heads in profile view,
showing the ears and vibrisse of some genera of Loncherine.
Boas also (‘Ohrknorpel der Siiugthiere,’ p. 119 ef seg., 1912)
described the ears of some genera, figuring those of Cavia,
Dasyprocta, Celogenys, Hydrocherus, Lagostomus, and Hystrix.
The observations set forth in the following pages have been
taken mainly from fresh material examined immediately after
death in the Society’s Prosectorium. This has been supplemented
by specimens preserved in alcohol in the Society’s collection, and
by dried skins where soft material was unavailable.
7 _ The Rhinarium.
In Hystrix and Atherura the rhinarium is ill-defined. In
Hystrix it is hairy to the edge of the nostrils. In Atherura
there is an area of naked skin both above and below these
orifices. In both genera the nostrils are transversely elongated
slits, expanding anteriorly and internally; and there is a very
well-defined smooth philtrum, completely dividing the upper lip
into its right and left moieties, which are independently movable.
(Text-fig. 1, A-C.)
The rhinarium of Thrynomys is well defined, although the
hair encroaches upon it above and beneath laterally, leaving only
a narrow naked rim above and below the nostrils, which are
tolerably widely separated. It extends, as in Hystrix, to the
edge of the upper lip, forming a philtrum, wider above than
below, and completely dividing the lip into a right and left
portion. (Text-fig. 1, D.)
Jn Dinomys, according to Peters, the upper lip is deeply cleft —
apparently very much as in Hystrix and Atherura; and above
it there is a distinct triangular rhinarium, naked in front and
round the nostrils, but hairy above owing to the forward extension
of the hair of the muzzle well in advance of the posterior ends
of the nostrils, which are described as S-shaped, a form these
orifices assume in many of the Hystricomorphs *.
Dinomys is the only American genus of Hystricomorphs, so far
* Peters seems to have regarded Dinomys as akin to Celogenys. He appears to
have been misled in this matter by the similarity in colour between the two genera
There is, however, no obvious evidence of kinship between them.
25*
368 MR, R. I. POCOCK ON THE EXTERNAL
as my observations go, that has a complete philtrum continuous
with tle rhinarium above and cleaving the upper lip. The
nearest approach to it is seen in such forms as Ciwlogenys and
Text-figure 1.
SY
<=
=
GC Pus
Yi
A. Side view of head of Atherura africana to, show the ear, rhinarium, and
facial vibrissze, the latter drawn relatively shorter than in the animal.
B. Rhinarium and upper lip of the same. X 2.
C. > a m9 Hystrix africe-australis. X .
D. % op 4s Thrynomys swinderianus, from a dried skin.
<
KE, Ear of Hystrix africe-australis. X +
3°
CHARACTERS OF SOME HYSTRICOMORPH RODEN'S. 369
Dolichotis*, for example, in which there is a parting in the hairs
in the middle line of the upper lip which, especially in dried
skins, sometimes simulates a small philtrum ; but, although this
may be the remains of a true philtrum, it is very differ ent from
that structure in Zystrix and Thrynomys.
The rhinaria of Coendw and Frethizon differ greatly from
that of Hystrix. In Coendu it is hairy and the nostrils are
quite small and vertical rather than transverse in direction, the
postero-lateral sht being absent. In Hrethizon the rhinarium is
marked by shorter hairs than that of the area round it. The
nostrils are larger and more transverse than in Coendwu and are
very close together, the space between them and their very
narrow upper edge being smooth. In neither genus is there a
philtrum. (ext: fig. 2, 10) E.)
In Capromys the rhinarium is large and naked, but not very
well defined laterally, where it passes into the short-haired area,
surrounding it above as well as laterally and below. It is marked
by a median groove in front. The nostrils are elongated and
expanded anteriorly. (Text-fig. 2, A, B.) In Jyocastor the
rhinarilum is wider as compared with its depth than in
Capromys, the areas above and below the nostrils being much
narrower; there is no median groove, and the nostrils them-
selves are much smaller, forming crescentrically valvular slits.
(Text-fig. 3, A.)
In Ctenomys, Octodon, and Cavia the rhinavium forms a naked
area round the nostrils, the areas above and below these orifices
being narrow. (Text-fig. 4, C.) In Octodon it is mesially grooved
and angled below. In Ctenomys it has the upper edge biconvex
and mesially angled, the inferior edge transverse in the middle.
(Text-fig. 4, A, B.) In Cavia the nostrils are wider than in the
two Octodonts above described, the upper edge of the rhinarium
is concave, the lower convex from side to side, and there is
sometimes hair between the nostrils. (Text-fig. 4, F.)
In Dolichotis patagonica the entire nose is large, with a wide
rhinarium concave in the middle and convex at the side above
and below, the finely hairy and tolerably deep areas bordering
the large transversely extended nostrils above and below being
thickened and muscular. In one example of this species the
internarial area was hairy in the middle line, whereas in an
example of D. salinicola it was naked; but this character is
possibly not constant. (Text-fig. 3, D.)
In Celogenys the upper rim of the nostril is swollen and
muscular, but the rhinarium itself is scarcely defined, being
covered with very fine short hairs, leaving only a narrow hairless
area above and below the elongated narrow nostrils. Judging
from dried skins the rhinarium of Dasyprocta is very similar.
(Text-fig. 3, B.)
* T am not sure about the structure of the upper lip in Dasyprocta. The only
spirit example available had the rhinarium destroyed; but dried skins suggest the
absence of a complete philtrum. In living exampies there appears to be a median
eroove formed by a vertical fold in the skin of the upper lip.
370 MR. R. i. POCOCK ON ‘iH EXTERNAL
In Chinchilla, Lagidiwm, and Lagostonus the rhinarium is also
covered with fine short hairs almost up to the level of the nostrils,
but whereas in Lagostomus the nostrils are elongated and the
short-haired area around them sharply defined by the coarse
Text-figure 2.
WZ Ay
GE,
KEG |
. Side view of head of Capromys pilorides, showing the ear, rhinarium, and
facial vibrissxe, the mystacials shortened in the drawing.
B. Muzzle and mouth of the same from the front, the divided palatal flaps (p)
shown above the tongue (¢). X 2.
C, Rhinarium and upper lip of Hrethizon dorsatum. X +.
D. Muzzle and mouth of Coendu prehensilis, from the front. X +.
hairs of the rest of the muzzle, in Chinchilla and Lagidium ihe
nostrils are much smaller without any sharply defined short-
haired area around them. (Text-fig. 3, C; 4, D.)
>
GHARACTERS OF SOME HYSTRICOMORPH RODENTS. 371
In Hydrocherus the vhinarium is scarcely defined, although
the skin between the nostrils is naked. The nostrils are very
widely separated and small, without any postero-lateral slit
The upper lip is exceedingly deep and naked or scantily hairy
in the middle.
The rhinarium of Ctenodactylus is continued to the edge of the
upper lip by a naked area of skin, which, like the rhinarium
itself, is capable of lateral compression, heing broad or narrow
according to the degree of contraction of the muzzle. The upper
end of this labial tract is continued on each side asa naked rim
beneath the nostrils. The upper edge of the xvhinarium is’
convex with a median angular emargination. The area above
the nostrils is deep; the nostrils themselves when expanded are
large orifices with very short posterior slits, and the narrow space
between them is marked by a deep median groove extending
from the top of the labial tract to the summit of the rhinarium.
(Text-fig. 4, G.)
In view of the former association of Ctenodactylus with the
Jerboas (Jaculide) it is interesting to note the close similarity
between that genus and /aculus in the rhinarium, not only in
structural details but in its compressibility.
Facial Vibrisse.
In P. Z. 8. 1914, pp. 903-905, I deseribed briefly the facial
vibrisse of several species of Hystricomorpha belonging to the
genera Atherura, Coendu, Hrethizon, Octodon, Chinchilla, Lago-
stomus, Dolichotis, Cawlogenys, Dasyprocta, Cavia, anl Hydro-
cherus, and showed the general constancy of the presence of
well-developed mystacial, superciliary, and genal tufts, the latter
being usually set high up the face near or even behind the
posterior angle of the eye. Since then J have extended my
observations, and can add several more species to the list.
In Hystrix, Acanthion, Atherura, and Trichys the vibrisse are
as originally described in Atherwra, namely, exceedingly long
and well developed, and there is always an interramal tuft. In
Thrynomys, on the contrary, although its mode of life is tolerably
similar to that of the Old World Porcupines, the mystacials are
comparatively slender and short, and in two dried skins I can
find no trace either of superciliary, genal, or interramal tufts in
the coarse hairs clothing the head. Peters described the mysta-
clals as long in Dinomys and the genals as set behind the eye,
but says nothing about the interramal tuft.
In the genera of Loncherine figured by Winge, namely,
Loncheres, Echimys, Cannabateomys, Trichomys (Nelomys),
Carterodon, and Mesomys, well-developed mystacial, superciliary,
‘and genal vibrisse, the latter behind the eye, are shown, but no
interramal tuft is indicated. The omission of the interramal tuft
can hardly be taken as proof of its absence in these genera.
Nevertheless, I failed to find it in examples of Ctenomys and
ae MR. R. I. POCOCK ON THE EXTERNAL
~_
Octodon, which have the other vibrissz well developed, although ©
in Ctenomys the mystacials are shorter than in the other genera
of Octodontide above quoted. In the examples of Capromys,
Text-figure 3.
aaa tin
et NTR
A\ Ah
7
(th
ORO
s
iy
Wy
Yp AY AAT
7
SS
SS
=
tm
LW
ANS
Be A
OO ‘
MN
i S
D
\
EE
A. Muzzle and mouth of Myocastor coypu, from the front: p, palatal flaps
fused behind upper incisor teeth; 7, lmgual flap. X 4. :
B. The same of Ceelogenys paca. Lettering as in A, with 0, orifice of external
cheek-pouch. > +.
C. Rhinarium and upper lip of Lagostomus trichodactylus, from dried skin. X +.
D. The same of Dolichotis patagonica. X 4.
Coendu*, and Hrethizon examined the interramal is also absent,
and the same is true of all the adult specimens of JM/yocastor.
* In C,-prehensilis the fore and hind limbs and the lower portion of the sides of
the body are provided with a few long scattered tactile vibrissz recalling those
on the body of Hyrax (Procavia).
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. ole
Text-figure 4.
NU
(
Wy
A. Side view of head of Ctenomys mendocinus *.
B. Rhinarium and upper lip of the same.
¢. 8 5 55 Octodon degus *.
D. 3 +5 38 Chinchilla lanigera.
E. Muzzle and mouth of the same, with the fused palatal flaps and the
small cheek-pouches.
F. Rhinarium and upper lip of Cavia porcellus.
ot be 35 A Ctenodactylus gundi.
H. Side view of head of the same.
* Jn this and other illustrations in this paper, the figures labelled Cfenomys
mendocinus and Octodon degus were taken from specimens that came respectively
from Cordova in the Argentine (W. A. Sinithers) and Valparaiso (W. Goodfellow).
374 MR. kt. f. POCOCK ON THE EXTERNAL,
The genal tuft, too, is generally indistinguishable in the latter
genus, although in a young specimen I have detected both genal
and interramal tufts, the former consisting of two vibrisse close
to the hinder corner of the eye. |
In Calogenys and Dasyprocta the ordinary vibrissee are long
and numerous, especially those of the genal tuft in Calogenys,
and the interramal tuft is present. In Dolichotis the vibrisse
generally resemble those of Dasyprocta; but in an example of
D. salinicola the interramal vibrisse, present in a specimen of
D. patagonica, were absent. In Hydrocherus all the vibrissee are
short and slender, the interramal are absent and the genal are
set below the level of the eye. In the typical species of Cavia
(C. porcellus), the common guinea-pig, and in ©. aperea, the
mystacials, superciliaries, and genals are as in the Octodontide,
Dolichotis, and others, although relatively somewhat shorter; and
the interramals appear to be absent. But in an example of
Galea littoralis Thos.*, the interramals are represented by four
long bristles arranged along the posterior border of a nearly
naked area behind the chin, an arrangemert recalling that of the
ruminant ungulate 7’ragulus.
In Chinchilla and Lagidium the mystacial vibrisse are exceed-
ingly long and coarse, but the superciliaries are much finer and
softer. The genals also, when present, are fine and soft, but I
found them in only one example of Chinchilla. In another
example of that genus and in a specimen of Lagidiwm I could
not detect them. In Lagostomus the mystacials, superciliaries,
and genals are all long and coarse, and this genus in addition
has a long thick mat of bristles on the cheek below the eye.
The interramals appear to be undeveloped in these three genera.
Ctenodactylus has. long mystacials, superciliaries of medium
length, but no discernible genals or interramals.
The Mouth and Cheek-pouches.
The mouth of the Hystricomorphs, as in other groups of
Rodents, is provided with lobes of skin jutting into it from the
cheek on each side and serving to shut off the anterior from the
posterior part of the buccal cavity so as to prevent gnawed
fragments of wood from passing into the throat. A pair of
these, one on each side of the tongue, capable of meeting behind
the lower incisor, may be called the lingual lobes, while an upper
pair, capable of meeting across the palate behind the upper
incisors, may be called the palatal lobes.
The lingual lobes, so far as my observations extend, always
retain their distinctness and can be separated or brought together;
but the palatal lobes are more variable. Typically and in most
* For the genera of this group, see Thomas’s paper, Ann. & Mag. Nat. Hist. (8)
Xvill. pp. 301-303 (1916). Iam indebted to Mr. Thomas for the correct names of
the species of Cavies recorded in this paper. The specimen above referred to as
Galea littoralis is the one I erroneously identified as Cavia rufescens when I
described its facial vibrisse in P. Z. S. 1914, pp. 900 and 905.
CHARACTERS OF SOME HYSTRICOMORPH RODENTS 375
genera, such as Hystrix, Atherura, Hrethizon, Coendu, Capromys,
Ceelogenys, Cavia, Dolichotis, and Ctenodactylus, they ave sepa-
rated, although capable of being made to meet in the middle
Text-figure 5.
i!
A. Side view of head of Celogenys paca, the outline of the external cheek-
pouch dotted in ; 9, its orifice.
B. Transverse section of the head of the same anterior to the molar teeth, the
internal cheek-pouch on the left of the figure distended; 0, orifice of
external cheek-pouch; ¢, tongue; bony tissue dotted.
line, but in Ctenomys, Myocastor, Chinchilla, and Lagostomvus they
are fused across the palate anteriorly, although partially separated
posteriorly.
376 MR. R. I. POCOCK ON THE EXTERNAL
The only genus of the group which has genuine cheek-pouches
is Cologenys; but near the edge of the cheek, just inside the
mouth, in Chinchilia I find a small integumental pocket on each
side. These do not seem large enough for the storage of food,
and I am unable to surmise what their function may be. (Text-
fig. 4, E.)
The cheek-pouches in Calogenys lie on each side of the maxillary
portions of the palate, mainly in front of the molar teeth, and are
associated with the well-known and unique structural peculiarity
of the skull from which the genus takes its name. The outer
wall of each pouch is the original skin of the cheek, which
inferiorly passes downwards on to the face behind the corner of
the mouth. But this wall is entirely concealed from view
externally by the skin-covered, arched, laminate outgrowth of
the maxillary portion of the zygoma, which curves downwards
below the level of the palate and the molar teeth; and this
lamina itself forms the outer wall of a hair-lined external cheek-
pouch the orifice of which lies on the face above and behind the
angle of the mouth. Thus the original wall of the cheek,
covered with hair externally and with moist mucous membrane
internally, forms a partition between the internal and external
cheek-pouches; and this partition, being pliable, can be thrust
by pressure from inside the mouth so as to bulge into the cavity
of the outer cheek-pouch so as to diminish its size and increase
to a corresponding degree the size of the internal cheek-pouch.
Tf the latter were packed full of food, the external pouch would
be practically obliterated ; but there would be no visible swelling
on the face, such as is seen in the case of monkeys, hamsters, and
other mammals provided with these food-receptacles, because of
the presence of the secondary cheek formed by the bony out-
growth of the zygomatic arch. (Text-fig. 5, A, B.)
There can, [ think, be no doubt that the internal cheek-pouches
are used for storing food, although I have never found any food
in them in dead specimens of Celogenys. I attach no import-
ance, however, to this negative piece of evidence, because the
pouches of monkeys are usually empty when they die. That the
external cheek-pouches are not used for storing food is also, in
my opinion, beyond doubt, because the fore feet of Cwlogenys are,
judging from their structure, altogether unsuitable for thrusting
food into them.
From the description above given, it will be evident that the
laminate expansion of the maxilla acts as an external shield to
the cheek-pouch when distended. But this affords no expla-
nation of the equally remarkable expansion of the jugal portion
of the zygomatic arch behind the cheek-pouch.
The Har.
The ear in Hystriv and Atherura is comparatively small, but
stands well up from the head with a tolerably evenly rounded
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 377
upper border. In both genera the tragus is distinctly developed,
and there is a large antitragal thickening. In Hystrix the
antero-internal ridge descends internally to the tragus ; in Athe-
rura it is a rounded lobate thickening above the tragus. The
supratragus is a narrow ridge concealed in front by the antero-
internal ridge and ending posteriorly in a ridge which descends
into the cavity of the ear. Behind the antitragus there is a long
deepish groove, and behind this the posterior edge of the ear
forms a flap capable of being folded forwards. (Text-fig. 1, A, E.)
The ear of Dinomys is simple, with a ridge-like supratragus
and a thick, but not prominent antitragus. (Text-fig. 6, E.)
The ear in Hrethizon is covered thickly with woolly hair and
surrounded with spines. It is quite simple in structure and has
a tolerably evenly convex edge, which anteriorly above is curled
over and overlaps the anterior end of the simple supratragal
ridge, which posteriorly curves downwards and runs into the
tolerably large antitragal thickening. The lower margin of the
ear beneath the “aditus inferior” is a simple shallow ridge, and
the auditory orifice is not concealed by a tragal lobe, and there is
no definitely developed antero-internal ridge. (Text-fig. 6, C.)
In Coendu the ear is of a much more specialised type. Its
outline is irregularly angular. The antitragal portion is exces-
sively developed into a high valvular thickening, and the con-
sider ‘ably overturned antero-superior edge is jointed at the level
of the supratragal ridge, so that the upper portion of the ear can
be folded down upon the lower portion, reducing the size of the
cavity, which can be similarly compressed from behind by the
forward movement of the posterior portion. Modified as the ear
is, 1t is clearly derivable from the type seen in Hrethizon; and, as
in the latter, the auditory orifice is not protected by a tragal
lobe. (Text-fig. 6, D.)
The ear of Capromys is tolerably similar to that of Hystria,
but not so expanded at the summit. The antero-internal ridge
descends to the auditory orifice and is continued superiorly a little
above the point of attachment of the pinna, but the overfolded
portion of the edge of the ear is here much narrower than in
Hystrix. There is a small tragus concealing the orifice, and a
narrow notch betwen it and the antitragal ridge which is also
less well developed than in Hystrix, and above and behind the
antitragus there is a small depression much shorter and shallower
than in the latter genus. As in Hystrix the simple supratragus
is overlapped in front by the antero-internal ridge. (Text-
fig. 2, A.)
In Myocastor the ear is lower and rounder than in Capromys,
and has a laminate bulge on its posterior border above and
behind the antitragus. The antitragus is better developed than
in Capromys, and is provided with a large tuft of hairs which
serves to keep water out of the auditory ‘orifice, which similarly
has a crown of short hairs above it to serve the same purpose.
The antero-internal ridge, instead of descending nearly vertically
L
MR. R. I. POCOCK ON THE EXTERNAL
Text-figure 6.
A. Ear of Myocastor coypus, with hairs removed.
B. Lower portion of the same with vertical cut through the intertragal notch,
and the tragus turned forwards to show the fringe of hairs above the
orifice and the tuft of hairs on the antitragus (a).
C. Ear of Wrethizon dorsatum, with the hairs removed.
D.
K.
F.
G.
H.
Coendu prehensilis.
Dinomys branickii, copied from Peters.
Dasyprocta sp.?
Oavia aperea, with valvular fiap upturned below supratragus.
Galea littoralis, without valvular flap below supratragus.
Octodon degus.
Loncheres armatus, copied from Winge.
Hehimys cajennensis, copied from Winge.
Ctenomys mendocinus.
Ctenodactylus gundi.
a, antitragus; s, supratragus.
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 379
to the auditory orifice as in Capromys and Hystria, curves some-
what backwards beneath the supratragus and loses itself in a
thickening occupying the cavity of the ear above the orifice.
(Text-fig. 6, A, B.)
In the ear of Octodon the laminate portion is widely expanded,
especially below behind the antitragal area, but the edge above
this is distinctly emarginate. The cavity is, however, capacious,
although the supratragus which borders it above and in front does
not stand out as a definite shelf-like ridge. The extension of
the antero-internal ridge curves backwards and then upwards,
detining a deep pit as in Carlogenys and Lagosiomus. The tragus
is reduced to a little excrescence, not concealing the orifice which
lies behind the ridge forming the anterior border of the unusually
elongated notch (aditus inferior). The antitragus is very large
and fleshy, but has no trace of pouch. (Text-fig. 6, I.)
In the six genera of Loncherine Octodontide figured by
Winge, namely, Loncheres, Hchimys, Cannabateomys, Trichomys
(Nelomys), Carterodon, and Mesomys, the ears are apparently
simple in type, moderately large or small in size, and stand away
from the head inferiorly from a point beneath the antitragus or
the notch in front of it, which is distinct in all of them. The
anterior edge is folded over from a point below the anterior end
of the supratragus, its inferior end curving backwards and down-
wards into the cavity of the ear, this curvature being especially
strongly marked and high up in Loncheres, where the ridge is
curled so as to cireumscribe a definite pit as in Octodon. This
peculiarity is not so well marked in the other genera. In
Loncheres, too, the pinna is relatively smaller and the antitragus
larger. Carterodon has relatively the smallest antitragus of all.
The tragus is small in Lchimys, Trichomys, and Cannabateomys,
and apparently undeveloped in the others. The supratragus
forms a simple, scarcely a shelf-like, ridge roofing the cavity of
the ear above anteriorly. The posterior border of the ear is
slightly emarginate and angled above in Loncheres, Cannabate-
omys, Trichomys, and Mesomys, convex and angled in Carterodon,
nearly straight and quite unangled in Hchumys. The inferior
portion behind the antitragus is well developed in all, but shows
no trace of a pouch. (Test-fig. 6, K, L.)
Judging from the figures from which this description is taken,
the ears of ail these genera are of a simpler, more primitive type
than those of Octodon degus.
Judging from the illustration of Petromys, published by
A. Smith (Illustr. Zool. 8. Africa, Mammalia, 1849) the ear
resembles that of the typical Octodontide in a general way, but
details are not described or figured.
The ear of Ctenomys is greatly reduced and simplified im
adaptation to subterranean life. The apex is pointed; the pos-
terior border is lightly concave above, but only stands freely away
.from the head from a point a little below the supratragus, which
is not defined as a definite ridge, but is merely represented by the
380 MR. R. I. POCOCK ON THE EXTERNAL
anterior part of the wall of the subcircular cavity of the ear con-
taining the large exposed auditory orifice, the tragus, antitragus,
and the intervening notch being obliterated. ‘The antero-internal
ridge, forming the continuation of the strongly overfolded an-
terior rim of the ear, gradually disappears inferiorly at about the
level of the orifice and does not curve sharply backwards beneath
the supratragus. (Text-fig. 6, M.)
In Celogenys this ridge beneath the supratragus is much more
sharply defined and forms the lower edge of a well-defined depres-
sion, of wnich the supratragus, which has a definite thickening,
is the upper border, and above the supratragus there is another
weaker ridge. Mitevel is a small tragus, but the antitragus is very
large and thick, and is marked above and _ posteriorly “by a small]
shallow pouch. Above this pouch the antitragus is continuous
with a ridge curving forwards above and forming the posterior
rim of the cavity of the ear. The laminate portion of the ear is
small, but erect, with a slight emargination near the middle of its
posterior border, a rounded summit, and an overturned anterior
margin continuous below with the antero-internal ridge. (‘Text-
1235, (fs, 1D),))
The ear of Dasyprocta is wider as compared with its height
than in Celogenys. As in that genus the inferier end of the
overturned anterior rim juts into the cavity, forming a horizontal
shelf-like ridge beneath the supratragus, which is itself a long
narrow ridge with a well-defined depression above it. There is a
smail tractus as in Celogenys, but the antitragus is much less
developed than in that genus and has no pouch above it. Above
the tragus there is a little curled thickening. (Text-fig. 6, F.)
In Cavia and allied genera the ear resembles tolerably closely
in shape that of Dasyprocta, although varying within the limits
of the genus. But it always differs in having the supratragus
converted into a laminate valvular flap. In Galea littoralis the
ridge beneath the supratragus, which is also present in Dasyprocta
and Oelogenys, is quite thin and of uniform width throughout ;
the tragus is distinetly defined, and above it there is a thickening
recalling that seen in Dasyprocta but larger; the antitragus, too,
is well developed, and the portion of the pinna below and behind
it is comparatively wide, wider than in Dasyprocta. But in Cavia
aperea and the Common Guinea-pig (CU. porcellus) the ridge
beneath the supratragus is large and valvular, like the supra-
tragus itself, there is scarcely a trace of tragal and antitragal
thickenings, and the pinna is narrower behind the antitragal
ridge. When the ear of the Guinea-pig is folded the two valves
close down over the orifice, the supratragal valve lying upper-
most. The ear of Kerodon resembles that of the two last-
mentioned species of Cavia, although the two valves are relatively
a little smaller and the tragus and antitragus are a little better
developed. (Text-fig. 6,G, H.)
The ear of Zhrynomys, like that of Cavia, has the supratragus
valvular and the antero-internal ridge passing beneath it. There
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 381
is a small tragus and a moderately large antitragus. (Text-
fig. 7, A.)
“The ear of Dolichotis is much longer than in Cavia or
Celogenys, and is somewhat narrowed ‘above by the tolerably
deep and long emargination of its supero-posterior edge. The
supratragus is a simple ridge without the thickening seen in
Celogenys. The continuation of the antero-internal ridge i 1s very
distinct and long, longer than in Celogenys, but equally simple
in structure. Beneath it there is a similar ridge curving abruptly
downwards to end in a thickening above the auditory orifice.
The tragus is quite well developed cmd high, but the antitragus
is not better developed than in Cavia. (Text-fig. 7, C.)
In Hydrocherus the ear is very much simplified, the only
strongly developed ridge being the supratragus, which terminates
somewhat abruptly near the middle of the cavity. The antero-
internal ridge is not turned backwards beneath the supratragus,
but gradually fades away inferiorly. There is no definite tragus
or antitragal thickening, and the ridge representing the latter is
not continued so high as the posterior end of the supratragus.
The depression in which the orifice lies is thickly overgrown with
hairs covered with waxy secretion, the two combining to exclude
water. (Text-fig. 7, C.)
The ear of Layostomus trichodactylus is very specialised, but
seems to be an extreme modification of some such type as that
seen in Calogenys. Its laminate portion is very much larger
and is tolerably evenly oval. There is a well-developed tragus
separated by a large notch from the antitragal portion, which,
however, does not form a thickened bulge as in Celogenys, but is
continued obliquely upwards and backwards as a long straightish
sharp ridge defined behind by a shallow depression ‘running in-
feriorly into a shallow pouch, doubtless homologous to the
similarly placed pouch above and behind the antitragus in
Celogenys. The backwardly curved continuation of the antero-
internal ridge is strong and forms the lower border of a deep
depression, bordered above by the thickening representing the
supratragus, but this does not form a definite ridge as in most of
the ears hitherto described. (Text-fig. 7, F.)
The ear of Chinchilla is tolerably similar in type to that of
Lagostomus, but is much more expanded, except at the base
where it is narrower and more tubular, so that the orifice lies
deeper in the ear at the bottom of the depression bordered above
by the continuation of the antero-interna] ridge. No trace of
the supratragus remains. The tragus, too, has disappeared ; but
the antitragus is developed into a prominence relatively as large
as in Ceelogenys ; and this is hollowed into a deep pocket, from
the centre of which arises a straight ridge corresponding to the
similar straight ridge in Lagostomus and to the curved ridge
defining the cavity of the ear posteriorly in Celogenys. (Text-
fig. 7, E.)
Proc. Zoou, Soc.—1922, No. XX VI 26
382 MR. R. I. POCOCK ON THE EXTERNAL
The ear of Ctenodactylus is tolerably small, projects but little,
is sloped backwards, and is covered with hair behind and on the
Text-figure 7,
WE
)
ate %
S\ WW Win Wy
A. Ear of Thrynomys swinderianus, from dried skin.
», Hydrocherus capybara.
» Dolichotis patagonica.
, Ceelogenys paca.
» Chinchilla lanigera, the position of the orifice dotted in.
5 Lagostomus trichodactylus,
eo On
CHARACTERS OF SOME HYSTRICOMORPH RODENTS, 383
edge of the cavity. The edge or rim is raised all round, defining
a deep hollow divided into two by a large valvular supratragus,
which separates the upper third from the lower two-thirds of the
cavity. There is no tragus; but the antitragus forms a long,
narrow, infolded, naked ridge, from the upper end of which a
short ridge runs obliquely forwards, wpwards, and inwards to
terminate in the middle of the cavity beneath the supratragus.
A corresponding ridge occupies a similar position in the anterior
half of the lower two-thirds of the cavity. The oritice of the
auditory meatus is a large hole situated just within the lower
half of the antitragal ridge. (Text-fig. 6, N.)
According to Dobson, the ear of Pectinator vesembles that of
Ctenodactylus except that it retains the antitragal thickening
which the latter, he alleges, has lost. Since, however, there is a
long low antitragus in the examples of Ctenodactylus | examined,
this distinction does not hold. Dobson probably overlooked this
structure in Ctenodactylus owing to its being normally folded
inwards. Assuming that their ears are similar, these two
genera differ markedly not only from all the Hystricomorphs,
but from all other Rodents known to me in the structure of
this organ.
The Feet.
In Hystrix, Acanthion, and Atherura the fore foot is short and
broad. The four main digits are short, subequal, moderately widely
separable, and united by naked integument up to the proximal end
of the large smooth digital pads. The claws are tolerably long
strong, and fossorial. The pollex, on the other hand, is éreatly
reduced and represented externally merely by its nail and small
pad. It arises alongside the outer distal extremity of the radial or
inner carpal pad, behind or above the inner lobe of the plantar
pad. ‘The plantar pad is smooth, about twice as wide as long, and
three-lobed. The median lobe projects in the middle line and
is larger than the lateral lobes, of which the inner is smaller
than the outer. The posterior border of the pad is almost
straight from side to side. Immediately behind it are two large
smooth carpal pads, separated by a naked space, or nearly in
contact, and behind the pads there is a short area of naked skin
overlapped by the bristly hairs above it.
The digits and plantar pad of the hind foot are, in general
features, tolerably similar to those of the fore foot, except that
the hallux is better developed than the pollex and arises just in
front of the inner lateral lobe of the plantar pad. The sole of
the foot behind the plantar pad is smooth and naked back to the
heel, and is occupied for the most part by two large, but not well-
defined, metatarsal pads. (Text-fig. 8 )
The feet of Dinomys, judging from the description and figures
published by Peters*, appear to be more like those of Hy ystrva
* Festschr. Ges. Nat. Berlin, 1873, p. 228, pl. iii. figs. 2, 3
DA
384 MR. R. I. POCOCK ON THE EXTERNAL
than of any other genus of Hystricomorpha, and are unlike the
feet of Cologenys with which Peters compared them.
The pollex and hallux, which are quite short and practically
functionless in Hystriv, have disappeared; but, as in Hystrix,
Text-figure 8.
A. Right fore foot of Atherura africana.
B. ,, hind ,, ., thesame.
C
1
, tore foot of Hystrix africe-australis.
D. .,, hind ,, ,, the same.
digits 2-5 are short, thick, and strongly developed, 3 and 4 being
subequal and a little longer than 2 and 9, which are themselves
CHARACTERS OF SOME HYSTRICOMORPIH RODENTS. 385
subequal, 2 being set a little more forwards than 5. ‘The inter-
digital webbing, however, does not perhaps extend so far distally
as in Hystriv, and the soles are—in part, at all events—covered
with squamiform papille instead of being smooth. The plantar
pad of the fore foot seems to be of much the same form as in
Hystrivz, and just behind it and in contact with +t there is an
immense carpal pad with a median depression indicating its
division into the two normal elements. The plantar pad of the
hind foot is reduced to three smooth isolated prominences rising
from the papillate integument; and the metatarsal area shows no
trace of the two pads, but is uniformly papillate in its distal
portion, where it is scarcely defined from the plantar portion,
and smooth and swollen in its proximal portion up to the heel.
(Text-fig. 19, A, B, p. 402.)
In Octodon degus the fore foot is artiodactyle, the third and
fourth digits being paired and subequal and a little longer than
the second and fifth, which are also subequal, the fifth, however,
being a little the shorter of the two. These four digits are stout,
shortish, with sharp moderately long claws; the pollex is quite
short and represented externally by little more than its claw.
The plantar pad is large, as wide as the foot, considerably wider
than long and markedly three-lobed, the median lobe being larger
than either of the others. It is covered with papille, which
spread on to the base of the digits, and each lobe is provided with
a single large papilla opposite the three interdigital spaces. The
double carpal pad of which the two elements are mesially in con-
tact, the outer being a little the larger, is a little larger than the
plantar pad and, like it, covered with papille, with a single
enlarged papilla at the antero-external corner of each. Behind
this pad there is a naked triangular area of skin. (Text-fig. 9, B.)
In general features the hind foot 1s like the fore foot, but the
digits are much longer and the plantar pad narrower and the
hallux is relatively much longer than the pollex. There is a
single enlarged circular papilla opposite the four interdigital
spaces, and two others on the metatarsal area, one set forwards
on the ulnar side just behind the outer moiety of the plantar pad
and the other farther back on the radial side some distance
behind the papilla opposite the space in front of the hallux. The
whole plantar surface is covered with small. papille, except the
heel, which is naked; and there is a median groove defining the
two elements of the metatarsal pads. I regard the feet of
Octodon as the most generalised type found in the Hystricomorph
Rodents. (Text-fig. 9, A.)
Winges figures of the feet of several genera of Loncherine
(Loncheres, Echimys, Trichomys (Nelomys), Carterodon, Mesomys)
show that, with variations in detail, they conform tolerably closely
with those of Octodon degus. They are pentadactyle with greatly
reduced pollex and short hallux; the primary interdigital ele-
ments of the plantar pads are separated, but they are rela-
tively much larger than in Octodon, and the papillate areas of
386 MR. R. I. POCOCK ON THE EXTERNAL
integument between them are correspondingly reduced ; and the
same applies to the carpal and metatarsal elements. Also the
fifth digit both inthe fore and hind foot is relatively shorter and
set farther away from the fourth in Lchimys, Carterodon, and
Vext-figure 9.
. Right hind foot of Octodon degus.
ONC HP a aeeet eyscmnes
» hind foot of Capromys pilovides.
» fore ,, ,, the same, showing the smooth area of the carpal pad
behind the pollex.
Sone
Mesomys. In Mesomys the feet are more fossorial in type, being
broader, and supplied with shorter digits and longer stronger
claws. Moreover, the fourth digit of the exceptionally wide fore
foot is considerably shorter than the second and third.
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 387
The fore foot of Capromys is very like that of Octodon, except
that the four main digits are relatively considerably longer,
Text-figure 10.
y
Lf
LAU aS
SS
A. Right fore foot of Hrethizon dorsatum.
B. ,, hind ,, ,, the same (the sole is sometimes more expanded
Cc
D
behind the hallux).
fore foot of Coendu prehensilis.
hind ,, ,, the same.
h, hallux; e, prehensile expansion of sole.
39
3”
there are no enlarged papille on the plantar or carpal pads, and
the external moiety of the carpal pad is relatively larger and
388 MR. R. I. POCOCK ON THE EXTERNAL
quite smooth on its postpollical portion, The hind foot is aiso
like that of Octodon, but is relatively broader and shorter, being
actually broader, instead of narrower, than the fore foot. More-
over, the hallucal or inner element of the plantar pad is not
fused with the metatarsal pad, but forms part of the plantar pad
itself, which is thus four-lobed instead of three-lobed as in
Octodon, and there is no trace of divisional line on the meta-
tarsal pads, and there are no enlarged papille either on the
plantar or metatarsal pads. (Text-fig. 9, C, D.)
Text-figure 11.
A. Right hind foot of Hrethizon dorsatum.
B. Right fore foot of Coendu prehensilis.
C. Right hind ,, ., the same.
h, hallux; c, heel; e, prehensile expansicn of sole.
Except that they are broader and shorter, with shorter digits,
partially webbed basally, much longer claws, and the component
elements of the plantar and carpal pads less clearly defined,
the feet of Hrethizon are like those of Capromys. The above-
mentioned differences, together with the suppression of the
pollex, externally attest higher specialisation of the feet, in
accordance probably with greater adaptation to arboreal life.
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 389
Specialisation of the feet on the lines indicated in Hrethizon
is carried a stage further in Co end, where an excrescence on the
inner side of the plantar area, giving greater width to the sole,
takes the place of the poliex, while a very much larger, nearly
semicircular expansion, occupying the whole length of the sole
of the hind foot, takes the functional place of the hallux*. This
expansion is movable up and down to a certain extent, and
enormously increases the supporting area and gripping power
of the foot. As in Hrethizon, the plantar, carpal and metatarsal,
and the digital pads are covered with squamous papille. (Text-
figs. 10, 11.)
In the disposition and proportionate length of its digits the
fore foot of Myocastor resembles that of Capromys, but the four
main digits are relatively shorter and stouter, and have thicker
blunter claws. The pads are better developed, being relatively
larger and more cushion-like. The median lobe of the plantar is
approximately twice the size of either of the laterals, and there
is a deep or shallow division between the two elements of the
carpal pad. ‘The radial moiety of the latter is also covered, like
the rest of the carpal pad, the plantar, and the digits, with squami-
form papille. I found no trace of carpal vibrisse. (Text-
figee lanes)
The hind foot is very much larger than the fore foot 7, mainly
owing to the elongation of the digits, of which the third is the
longest, the second and fourth a litle shorter and subequal,
the fifth considerably shorter than the fourth and the first than
the fifth. The first, second, third, and fourth are united by
swimming-webs extending up to the digital pads; but the only
trace of web between the fourth and fifth is a flap of skin
running along the inner edge of the fifth digit, which is thus
left free for the purpose, I believe, of acting more efticaciously
as a skin-scraper. The plantar pad is indistinctly defined, being
represented by lobes of skin at the base of the first, second,
third, and fifth digits. The comparatively short metatarsus is
naked back to the heel. Its anterior portion, like the plantar
area and the digits, is covered with squamous papille, but a
varying amount of the heel is smooth, especially on the outer
side, (Text-fig. 12, A.)
The feet of Dactylomys and Cannabateomys are very different
from those of the typical genera of Loncherine. In C. amblyonye,
as figured by Winge and Tullberg, the lower sides of the digits
and soles are miaiaerunlly and closely covered, except on the thea
with squamous papille, lineally arranged on the digits, but show
no trace of definite pads apart from. the inteoumental grooves
vesulting from the folding of the digits on the soles, The fore
foot has a minute functionless poles close to the base of the
second digit, which rises nearly on the same level as the fifth,
* Tn a young example of C. prehensilis there was a larger external hallux occupy-
ing the same position as the hallux in Erethizon.
+ Precisely as in the Otters and for similar reasons.
390 MR. R. I. POCOCK ON THE EXTERNAL
but exceeds it in leneth, although much shorter than the third,
The third and fourth are long, thick, and subequal. The fifth
only reaches the distal end of the first phalange of the fourth.
The ends of the digits are thick and bluntly rounded, and
furnished with nail-like claws, which are concealed from the
inferior aspect. The sole is marked with a Y-shaped groove,
the branches of which run to the margin at the spaces between
the second and third and fourth and fifth digits, the upright
Text-figure 12.
A. Right hind foot of Myocastor coypus.
Dts LONE - D .
presumably marking the original line between the two elements
of the carpal pad. ‘The hind foot has the sole short, only a little
longer than in Céenomys, and, as in that genus, broad in front
and narrowed behind. The second, third, fourth, and fifth digits
bear much the same relation to each other in length as those of
the fore foot, but the hallux is well developed as compared with
the pollex and stands well away from the sole. ‘The nails are as
in the fore foot, except that on the second digit the nail is claw-
like, according to Winge, and projects beyond the tip of the
digit, acting, no doubt, as a seratcher.
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 391
In Celogenys the fore foot is five-toed and artiodactyle. The
pollex is greatly reduced and represented externally merely by
its small nail, which is set some distance above the second digit.
The third and fourth digits are thick and subequal, and united
by webbing halfway between the plantar and digital pads. he
second is as thick as the third and considerably shorter ; but it
is stouter and longer than the fifth, and set a little more forwards.
The web between the second and third and fourth and fifth is
less extensive than that between the third and fourth. These
four digits have powerful claws and well-developed smooth pads.
The carpal pad is large, three-lobed, and projects in the middle
in front, this portion having a truncated anterior margin. It is
roughened with papille, if not all over, at least in the middle
line and laterally at the base of the second and fifth digits.
Separated from the plantar pad by aspace of naked wrinkled skin
are the two well-developed nearly smooth carpal pads which are
in contact in the middle line. The whole of the inner side of
the wrist as far forward as the base of the second digit is covered
with very short hair as in Dasyprocta and Kerodon. (Text-
fig. 13, A.)
The hind foot is also five-toed, but is perissodactyle. The first
and fifth digits are short and slender, the first being a little
smaller than the fifth ; and they arise nearly opposite one
another far up the sides of the foot, only a little in front of the
metatarsal pads and a long way behind the main mass of the
plantar pad. The second, third, and fourth digits are thick and
strong, the second and fourth being shorter than the third which
hes between them. There is a shallow web between them at the
base. The claws and digital pads are similar to those of the fore
foot. The main portion of the plantar pad is an irregularly
shaped cushion-like mass with two especially strongly papillate
areas corresponding to the interdigital spaces and attesting the
origin of this pad from two plantar interdigital elements. The
remaining two interdigital elements, seen in Octodon, for example,
have travelled away from the main portion of the pad, and are
represented by small pads at the base of the first and fifth digits,
although the one at the base of the first digit (pollex) is not
always distinguishable. The metatarsal area is covered by a
large horny shield, distinctly double at its distal end, but only
indistinctly divided in two elsewhere. (Text-fig. 13, B.)
The feet of Dasyprocta are in many respects very different
from those of Celogenys, but seem to be derived from that type.
They are much thinner and longer. The fore foot is perisso-
dactyle ; the pollex is even more reduced than in Celogenys. The
fifth digit also is relatively smaller; it is, moreover, set high up
the foot, almost on a level with the pollex, some distance behind
the plantar pad and slightly in advance of the carpal pad. The
second, third, and fourth digits are well developed, the second is
slightly shorter than the fourth, and both are shorter than the
third which lies between them. There is no visible basal webbing.
392 MR. R. I. POCOCK ON THE EXTERNAL
The claws and pads on these three digits and on the fifth are
well developed, the pads being smooth. The plantar pad is very
different from that of Cologenys, consisting of two elements
opposite the interdigital spaces of the three main digits. The
third, or outer. element of this pad is small and circular, and has
accompanied the backward migration of the fifth digit. The
carpal pads, two in number, are narrow, indistinct Behind, and in
contact mesially, the inner pad being larger than the outer and
projecting further distally. They are separated by a longish
area of naked skin from the plantar pad. The inner side of the
wrist is covered with short hair, sometimes almost naked, as in
Celogenys. (Text-fig. 13, C.)
The hind foot is like that of C wlogenys, except that the first
and fifth digits have entirely disappeared, the three main digits
are longer, thinner, and unwebbed, the plantar pad is carmela
smaller, consisting of two larger but smal! pads at the base of
the inner and median digits, and of a very small circular pad
at the base of the fourth or outer digit. Assuming this pad
to be the homologue of the small pad at the base of the fifth digit
in Celogenys, it occupies a very different position. The meta-
tarsal pad is a long way behind the plantar pad, and shows at
most indistinct signs of a divisional line on its inner side. (Text-
fig. 13, D.)
"The feet of Chinchiila are derivable from the type seen in
Ceiogenys, with the basal webbing eliminated. In the fore foot
the digits are thinner, with relatively larger compressed pads
and shorter claws adapted to a life amongst rocks, but their
number, relative length, and disposition are similar. There is a
distinctly three-lobed plantar pad and a large two-lobed carpal
pad, which, however, is larger than the plantar pad and not
smaller as in Celogenys. The wrist, however, is uniformly
covered above and laterally with long hairs. The hind foot is
longer and thinner, and the hallux has entirely disappeared ; but
the remaining four digits are similarly placed, although, as in
the front foot, they have relatively broader pads and much
shorter claws. The plantar pad is two-lobed, its external element,
larger than im Calogenys, has travelled up the foot in company
with the fifth digit and lies laterally nearly midway between the
plantar and metatarsal pads, the latter being defined by a sulcus
which expands into a crotch distally. A specialisation is the
development of fringes of longer hairs on the inner sides of
the second and third digits, those on the second forming a stiff
comb. (Text-fig. 14, A, B.)
In general features the feet of Lagidiwm ave like those of
Chinchilla, except that the under sides of all the digits, apart from
the digital pads, are thickly covered with hairs iad Die digital
pads themselves are compressed and furnished, as in Cerodon,
with a rounded median keel. The three elements of the plantar
pad of the fore foot are very iarge and in contact. Behind these
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 393
are two very large carpal pads which are also in contact. In the
hind foot there is a very large bilobed plantar pad; the third or
Text-figure 13.
SSS
ESS
SS
ony
\\ * if Y
\ “ jy y,
AN YY
\y
a\
A. Right fore foot of Celogenys paca. Xi.
Ws gy dm ogy eH a
C. ., fore foot of Dasyprocta sp.? X
De 5 Buhal
Cele
39 Ey) 29
394 MR. R. I, POCOCK ON THE EXTERNAL
outer lobe, which is also very large, has separated itself from the
main mass of this pad and travelled up the foot in company with
the fifth digit, as in Chinchilla; and, as in the case of the front
foot, the hairs of the sides encroach laterally more over the sole
than in Chinchilla. (Vext-fig. 15, C, D.)
In Lagostomus the fore foot differs from that of Chinchilla
and Celogenys in the suppression of the pollex, but is otherwise
Text-figure 14.
A. Right hind foot of Chinchilla lanigera.
B. 29 fore 39 39 ”
C. ,, hind foot of Lagostomus trichodactylus.
Deore
33 29 22
tolerably similar in type, with adaptations for a life in the open
plains as opposed to a rocky or forest habitat. The digits are
shortish with well-developed pads and claws; the third and
fourth are subequal and longer than the second and fifth;
the second also is longer than the fifth, but not quite to
the same extent as in Chinchilla and Celogenys. They are thus
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 395
more symmetrically artiodactyl. The plantar pad is wider as
compared with its length than in Celogenys, and its three com-
ponent elements are less well defined than even in that genus,
and markedly less so than in Chinchilla. As in Chinchilla, the
carpal pad is larger than the plantar pad and closer to it than
in Celogenys, but it is not so distinctly divided into two as in
those genera, and, as in Chinchilla, the outer side of the wrist is
normally hairy.
The hind foot differs from that of Chinchilla and Celogenys in the
complete suppression of the fifth digit and of the corresponding
element of the plantar pad. The second digit, moreover, 1s very
markedly shorter than the fourth *, and the latter, too, although
long, is shorter in comparison with the long third digit than in
Chinchilla and Cologenys. The claws of these three digits are as
powerful relatively as in Celogenys. ‘The plantar pad, as in those
genera, 1s composed of two elements, but they are very indistinctly
defined, and the whole pad is three-lobed with emarginate
antero-lateral and posterior borders. The metatarsal pad is large
and horny, but undivided. A peculiarity of the foot is the
encroachment of the hairs of the outer side nearly up to the
middle line, overlapping, with those of the opposite side, the
area between the plantar and metatarsal pads; and a further
difference from Chinchilla is the development of a thick bunch
of stiff bristles on the inner side of the third digit. Ohinchilla
has long hair in this position, but the brush, or comb, is
specially developed upon the second digit. The feet of this genus
are interesting, because they connect those of the Chinchilline
with those of the typical Caviine section of the group of Rodents.
The hind foot is also modified on much the same lines as that of
Dasyprocta, although the fore foot is very different. (Text-
fig. 14, C, D.)
The tect of Cacia porcellus, the domesticated Guinea-pig, have
been figured and described by Mivart and Murie (P. Z.S. 1866,
pp- 383-417), and by Tullberg. In the artiodactyle fore foot the
pollex is suppressed and the four remaining digits are moderately
long and subsymmetrically arranged, the third and fourth being
subequal and longer than the second and fifth, which are also
subequal. The claws project well beyond the normally shaped
digital pads. The piantar pad is well defined and three-lobed,
the median lobe being the largest. Behind the plantar pad
there is a single carpal pad, separated from it by an area of
naked skin. The hind foot is perissodactyle, with only three
toes, whereof the median (the third) is the longest. The claws
are longer than in the fore foot. The plantar pad is moderately
well defined and two-lobed, the inner lobe being much smaller
than the outer. The metatarsal area has no large horny plate
representing the pads, which are merely indicated by a single
* Tt must be remembered that the numerical terms applied to the digits through-
out this paper are used in a homological sense, as if the limbs were normally
pentadactyle.
396 MR. R. I. POCOCK ON THE EXTERNAL
median semicircular pad near the centre of the area some
distance behind the plantar pad. In a wild-caught example
‘of Cavia aperea, the feet agree in all essentials with those* of
C’. porcellus; as also do those of Galea.
Text-figure 15.
Se
SS
TS
Sa
WS
SS
SS
SS
SN
Z
SS
SSS
SS
A. Right fore foot of Kerodon rupestris, from dried skin.
B. 2° hind 29 33 33 3 33
C. ,, fore foot of Lagidium peruanum ,, 3
D. bb) hind 33 re) 33 23 33
The feet of Aerodon differ from those of Cavia in some
respects. The number and disposition of the digits is the same,
but the claws are much shorter and blunter, especially on the
fore foot. The digital pads are larger, compressed, and some-
what pointed at their distal ends. The plantar pads are very
large, well-defined, cushion-like thickenings, separated by a short
strip of wrinkled integument from the carpal and metatarsal
pads. The carpal pads are very well developed and two in
number, a larger outer and a smaller inner, the two together
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 397
approximately equalling the plantar pad in size. On the inner
side of the fore foot there is an area covered with short hairs,
as in Dasyprocta and Calogenys. In the hind foot the greater
part of the metatarsal area is covered with a large horny shield,
like that of Dasyprocta. (Text-fig. 15, A, B.)
In Dolichotis patagonica the fore foot is symmetrically artio-
dactyle, the second and third digits being subequal and longer
than the fourth and fifth, which are likewise subequal. The
pollex is absent. The claws are moderately long but blunt, and
the digital pads are very well defined. The plantar pad is a
large, thick, irregularly hexagonal cushion set far forwards
beneath the digits. Its edges are emarginate opposite the inter-
digital spaces, and its posterior border is mesially notched. It
is exceedingly deep, and the gait of this genus is markedly
digitigrade. The metacarpal area behind the plantar pad is
remarkably long, and there is a single very small carpal pad
remote from the plantar pad. The hind foot, as in Cavia,
Dasyprocta,and Hydrocherus, is perissodactyle and furnished with
three digits, which resemble those of the fore foot in essential
particulars. The plantar pad is relatively as large and high as
in the fore foot, but differently shaped; its edges are more
evenly convex, and there is a single median precess in front
corresponding to the median or third digit. It is composed of
two indistinctly defined elements. This foot, like the fore foot,
is markedly digitigrade, the posterior portion of the plantar pad
projecting like a heel. The metatarsal area is very long, and
its posterior half is covered with a single horny shield, upon
which the animal squats, like Hydrocherus, Dasyprocta, Kerodon,
and others. In both the fore and the hind foot the naked lower
side of the digits and of the area behind the plantar pads is
overlapped by the hairs of the sides of those parts. (Text-fig. 16.)
Dolichotis salinicola has feet closely resembling those of pata-
gonica, except that, judging from the single example examined,
the metacarpal area of the fore foot is shorter and the carpal
pad a little larger and closer to the plantar pad.
If the habits of Dolichotis were unknown, it would not be
difficult to infer from the structure of its feet that the animal
is adapted for swift running over hard ground. The differences
the feet present from those of Cavia may be ascribed to adapta-
tion to that mode of life.
The feet of Hydrocherus differ in several important respects
from those of Cavia. This is particularly the case with the
fore foot, which, as in Cavia, has no pollex, but is perissodactyle,
the third digit being the largest and situated in the middle line,
with the second and fourth, which are subequal, flanking it
laterally ; and these three digits are united by narrow webbing
up to the digital pads. The latter, however, are scarcely recog-
nisable as such, being represented by a softish thickening of
integument blending without line of demarcation with the skin
of the digit behind and with the claw in front and forming a
Proc. Zoou. Soc.—1922, No. XX VII. Py f
398 MR. R. I. POCOCK ON THE EXYERNAL
kind of heel to the claw, which, although narrowed distally, is
broad and hoof-like, and horny below as well as above. The
plantar pad forms a cushion provided with two flat horny plates.
Hence it is devived from two elements, not from three as in
Cavia, and it is set far forwards beneath the three digits, which
Text-figure 16.
4;
ZN
A
A '®&
O2|—
°
A. Right hind foot of Dolichotis patagonica. X
B. Side view of extremity of the same.
C. Right fore foot of the same.
D. Side view of the same.
thus appear to be very short when viewed from the under side.
The fifth digit, on the other hand, arises considerably behind the
fourth, approximately on a level with the posterior line of the
plantar pad, and the element of the plantar pad corresponding
to this digit is a comparatively small horny pad entirely separated
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 399
from the main mass of the plantar pad, as in the hind foot of
Celogenys, Chinchilla, and Lagostomus. Owing to the backward
position of the fifth digit and the forward position of the plantar
pad, the Capybara walks on hard soil upon that padand upon the
second, third, and fourth digits, the fifth scarcely reaching the
ground. ‘There is, as in Cavia, a single conical or semicircular
Text-figure 17.
A. Right hind foot of Galea littoralis.
Be. 3% fore Pe 5 3
C. ., hind foot of Hydrocherus capybara.
DS es) Store & 3 Ae
(In some examples of Hydrocherus, the horny plates on the plantar pads are
much more clearly represented than in C and D.)
carpal pad, which is set well on the outer or ulnar side of the limb
above the base of the external border of the fifth digit. The
area between the carpal and plantar pads is covered with wrinkled
skin overlapped by the sparsely growing long hairs of the sides
of the foot. (Text-fig. 17, D.)
my fa
400 MR. R. I. POCOCK ON THE EXTERNAL
The hind foot differs from that of Cavia in the webbing of
the digits and the hoof-like claws, which are like those of the
fore foot, except that the webs are wider and, owing to the more
backward position of the plantar pad with its two horny"plates,
Text-figure 18.
A. Right fore foot of Ctenodactylus gundi.
ie ay Imvvel g, 5 ”
C. Inner or second digit of hind foot of same, showing combs.
D. Right fore foot of Clenomys mendocinus.
195) gy Tail ss Fs x
KF. Tip of second digit of hind foot of same, showing combs.
apparently longer. The metatarsal area resembles that of Kerodon
in having the posterior three-fourths of its length covered by a
thick horny plate. (Text-fig. 17, C.)
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 40]
‘In Clenomys the fore feet ave fossorial. The four main digits
are short and thick, and armed with long and strong claws. The
digital pads are not well defined from the thickened transversely-
grooved skin on the under side of the digits. Digits 2, 3, and 4
are not markedly unequal in length, but 5 is considerably shorter.
The pollex is abbreviated, but armed with a strong, sharp. curved
claw. The plantar pad is irregularly three-lobed and not well
defined, but the two carpal pads close behind it are well developed
and conical, especially the inner, which projects as an excrescence
immediately behind the base of the pollex. (Text-fig. 18, D.)
The hind foot is short and wide, and narrows behind to the
heel, which like the rest of the lower surface is naked. The four
main digits are subequal in length, but thinner than on the fore
foot, but otherwise similar, except that the claws are relatively
and actually smaller, although that of the 4th is elongated and
curved. Digits 2, 3, and 4 are set nearly in a straight line and
evenly spaced, but digit 5 is set further back and much more
widely separated from 4 than the latter is from 3. Digits 2
and 3 are thickened above for the accommodation of the two
superimposed combs of stiff bristles which overhang the claws.
Similar bristles, but thinner and less modified, are found on
digits 1 and 4, and in the case of the latter the function of
combing the fur is no doubt performed by the long curved claw,
the point of which reaches as far as the distal end of the
bristle-combs on the 2nd and 3rd digits. The plantar pad is
almost suppressd, being mainly represented by four small hemi-
spherical tubercles, two in front just behind digits 2, 3, and 4
and one at the base of digits 1 and 5 respectively. The area
between these tubercles is wrinkled and irregularly papillate ;
but the area behind them is smooth and mostly covered by a
large callous external metatarsal pad, which terminates anteriorly
in a tubercle, and posteriorly falls short of the heel, which, like
the inner side of the foot, is covered with thinner skin, the
internal metatarsal pad being represented by a tubercle bebind
the pollical tubercle of the plantar pad. (Text-fig. 18, E, F.)
The feet of Thrynomys are not like those of any other genera
of the group; both fore and hind are perissodactyle. In the
fore foot the pollex is minute. The second, third, and fourth
digits are short, thick, and armed with long strong claws, the
second and fourth being subequal and shorter than the third,
which lies between them. They are free from webbing. The
fifth digit is much smaller than the fourth, but set close behind
it a little in front of the level of the pollex. The plantar pad is
wide, three-lobed. convex in front, and concave behind. The
carpal pad, which is separated mesially, at all events, from the
plantar pad by a moderately long membranous space, is large
and indistinctly divided by a groove and notch in front. (Text-
fier 19, C.)
The hind foot has lost all trace of the hallux externally.
Otherwise the digits are similar to those of the fore foot in
A()2 MR. R. I. POCOCK ON THE EXTERNAL
number, shape, and disposition, although longer. The ,plantar
pad is well developed and supplied with three smooth, suboval,
‘Vext-figure 19.
rei
SAN
YY
a
A. Right hind foot of Dinomys branickii, copied from Peters.
BOLE
2 39 9 29 39
C. ., fore foot of Thrynomys swinderianus, trom dried skin. Xid-
DE gy Taal of Pe zt Ph
isolated, interdigital areas, the innermost, tolerably large, at the
base of the second digit, the median equalling it in size behind
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 403
the space between the third and fourth, and the outer smaller,
rounder, and set further back behind the base of the fifth. The
metatarsal area is mostly covered with a smooth horny plate, with
bilobed anterior border separated from the plantar pad by a
shortish naked area of skin. (Text-fig. 19, D.)
The fore foot of Otenodactylus gundi has only four tolerably
long and slender digits, which are subequal in length, sub-
equally spaced, and widely separable. The pollex is absent. The
skin of the lower side of the digits is smooth, the digital pads
are well developed, and marked in their posterior half by two
transverse grooves. The claws are short, sharp, and curved.
The plantar pad is very large, markedly three-lobed, and very
coarsely areolate. Immediately behind it there are two large
smooth carpal pads in contact in the middle line, or nearly so,
each longer than wide and the two together as wide as the
plantar pad. (Text-fig. 18, A.)
The hind foot has digits similar in number and shape to those
of the fore foot, except that they are relatively much longer with
reference to the plantar pad, which is in a general way like that
of the fore foot in shape and sculpturing. The claws are short,
sharp, and curved ; and the two inner digits, the 2nd and 3rd,
are provided with combs of bristles similar to those of Cienomys,
but there are three of them on each digit instead of two. The
tips of the remaining digits also have long bristles, but these are
much thinner than those of digits 2 and 3. There are two
elongated smooth metatarsal pads, separated by a definite area of
transversely wrinkled skin from the plantar pad. They are in
contact throughout their length, and the outer of the two
extends back to the heel. (Text-fig. 18, B, C.)
Genital Organs of the Male.
A peculiarity of the penis of the Hystricomorphs is the
presence at the tip of the gland, behind and below the orifice, of
a wide slit leading into a sac, with laminated or corrugated walls,
which can be evaginated and withdrawn again by the action of a
pair of tendons. In some genera, as recorded below, this sac
is provided with a pair of horny spikes arising from its floor, as
was recorded long ago in the case of Cavia, Calogenys, and Dasy-
procta. Tullberg’s conclusion as to the generic incidence of
these spikes does not tally at all with mine. He says they are
distinctive of the family Caviide, which for him included the
genera Dasyprocta, Coelogenys, Cavia, Dolichotis, and Hydrocherus.
He does not, however, appear to have seen the penis of Dolichotis.
Hence his generalization with respect to it must have been
merely inferential. The spikes were not present in the example
of Dolichotis examined by me; they were also absent in two
specimens of Hydrocherus, although, according to Tullberg, the
penis of this animal resembles that of Cavia in essentials. I
have also found these spikes well developed in genera which fall
404 MR. Re I. POCOCK ON THE EXTERNAL
Text-figure 20.
tay -
\ “Nuel 2
| cG
i
Bi
Me
re
\
aun
aa
. Glans penis of Acanthion longicauda, from beneath.
. The same with the orifice of the glandular pouch dilated.
The same with the glandular pouch evaginated.
The same as C, from the side.
Anal area of the same, with anus dilated to show glands.
. Lateral view of anal area of young Hystrix africe-australis.
. The same of Coendw nove-hispania, with penis protruded.
. Posterior view of the same with glans retracted.
Lower view of glans of the same.
. The same with the glandular pouch cut open, showing a pair of short
spikes at the bottom.
ARO ee Ob
a,anus ag, anal gland; gl, glandular pouch; gw, genito-urinary
orifice ; , penis.
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 405
outside the limits of the Dasyproctide and Caviide, namely, in
Uctodon and Ctenomys, where they are of large size, and in
Acanthion and Hrethizon, where they are short. I do not know
what the explanation of these differences may be. Probably,
however, these spikes are only fully developed in mature males ;
but, since my examples of Dolichotis and Hydrocherus were fully
mature, 1t also occurs to me as a possibility that they may be
broken off during copulation and reproduced subsequently. Also
I do not know what their function may be, unless it is to
penetrate the orifices of the Fallopian tubes, either to make a
passage for the semen or, if shed, to block the orifices after the
introduction of the semen. However that may be, one thing is
clear, namely, that the presence of these spikes in such widely
separated genera as Acanthion, Hrethizon, and Octodon does away
with their importance as evidence of kinship between such genera
as Dasyprocta, Cologenys, and Cavia.
One other point to note is that the testes never pass into a
scrotum in the Hystricomorpha.
The penis of Hystrix and Acanthion is retrospective, when at
rest, being bent backwards on itself so that the prepuce forms a
swelling a littie beneath the anus. When erected the penis pro-
trudes a long way from the prepuce and projects forwards. It is
furnished with an apical baculum. ‘The orifice, when dilated,
reveals two apertures, an upper and smalier one immediately
beneath the tip of the baculum and a lower one forming a trans-
verse slit which leads into a tolerably deep glandular pouch, with
puckered walls and a pair of small spikes at the bottom. This
pouch can be evaginated, and in this condition it projects well
beyond the normal tip of the penis. It can be withdrawn by the
action of a couple of tendons running backwards along the lower
half of the penis beneath the urethral canal. The epithelium of
this glandular pouch and of the penis itself is beset with minute
spicules, but the epithelium at the bottom of the pouch round
the tooth-like spikes is smooth. (Text-fig. 20, A—F.)
According to Parsons (P. Z. 8. 1894, pp. 251-296) the male
genitalia of Atherura resemble those of Hystria.
In the male of Coendu nove-hispanie the penis opens at the
lower extremity of the naked area of skin common to it and the
anus, as described below (p. 417). When at rest the penis is
retrospective and completely withdrawn within a short naked
prepuce. The structure of the penis is almost exactly the same as
in Acanthion longicauda. The orifices of the urethra and of the
glandular sac are normally concealed by the epithelium at the
tip of the glans penis, but when this is spread the orifice of the
urethra is exposed just beneath the tip of the baculum, with
that of the glandular pouch behind it. This pouch has longi-
tudinally corrugated walls and a pair of small spikes at the
bottom. (Text-fig. 20, G—K.)
In Myocastor the penis opens retrospectively a little distance
below and in front of the anus and is normally entirely
406 MR. R. I. POCOCK ON THE EXTERNAL
withdrawn, the prepuce being a mere low rim of naked {skin.
The apex of the penis is attenuated and pointed, with the genito-
uvinary orifice just behind the tip. The orifice of the glandular
pouch is a slit with puckered lips, and the walls of the pouch
are longitudinally corrugated, but there is no trace of spikes at
Text-figure 21.
Wt
Ma i), _ ale
mn, a
aap i
SSE Wis,
SS iS
. Anal and genital area of Capromys pilorides.
. Lateral view of glans penis, projecting backwards, of the same.
5. Lower view of tip of glans penis, with glandular pouch cut open.
. Lower view of tip of glans penis of the same, with pouch cut open.
. Anal orifice of the same dilated to show gland beneath.
G. The same from behind with glandular pouch protruded.
H. The same from the side.
A
B
C
D. Tip of glans penis of Iyocastor coypus, in same position as in B.
E
F
a, anus; ag, anal gland; gp, glandular pouch; gw, genito-urinary orifice.
the bottom of it. Round the region of the pouch the penis is
expanded, becoming gradually attenuated to the apex distally
and narrowed proximally, the expanded portion being covered
with minute recurved spicuies. (Text-fig. 21, D, E.)
In Capromys the prepuce is long and pendulous and some little
distance from the anus. The penis itself is finely spicular,
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 407
truncated at the apex, and furnished with a distinct lappet
beneath the genito-urinary orifice, between the labia of which it
projects as in Octodon and Ctenomys. The apex of the glans is
truncated, a little attenuated from the inferior aspect, but not at
all from the lateral view. The glandular pouch is mederately
deep, with laminate walls, two of the lamine extending from
Text-figure 22.
E
A. Tip of the glans penis of Octodon degus, with glandular pouch cut open
and the spikes turned to show the two prongs. In the normal position
one prong lies behind the other.
B. Side view of anal and genital region of the same.
C. Vertical section of anus of the same, showing median gland.
D. Side view of anal and genital region of Ctenomys mendocinus.
E. Anus and penis, with glans protruded, of the same.
F. Side view of glans penis of the same.
Lettering as in text-fig. 21.
the tip of the glans bemg thick and ridge-like. There is no
trace of the horny spikes seen in Octodon and Ctenomys. (Text-
fig. 21, A-C.)
The prepuce of Octodon projects backwards a little way below
the anus. The penis is cylindrical with a conical apex, which is
smooth, the rest being closely covered with spicules. Between
the labia of the orifice there is a small soft rounded process
underlying the genito-uvinary orifice. The pouch has corrugated
408 MR. R. I. POCOCK ON THE EXTERNAL
walls and is armed at the bottom with two long spikes, but each
of these instead of being simple is divided into a pair of subequal
branches rising from acommon base. There are thus four spikes,
a peculiarity not recorded elsewhere in the Hystricomorpha.,
(Text-fig. 22, A, B.)
The prepuce of, Ctenomys is closer to the anus than in Octodon
and the penis itself is thinner, and when viewed from behind or
below is seen to be tolerably evenly attenuated to the narrowed
apex, but from the side its distal end is a little expanded behind
the pointed tip. It is beset with spicules, and the pouch is
corrugated and armed with a single pair of long spikes, as in
Dasyprocta. (Vext-fig. 22, D—-F.)
In Pasyprocta the glans of the penis, figured by Tullberg, is
subeylindrical, with a bluntly rounded apex, and is beset, espe-
cially beneath, with minute recurved sharp papille; but on each
side of it there is a long lustrous horny blade-like lamina, attached
by one edge to the epithelium of the glans, the other edge being
free and finely serrate. This plate is capable of erection, and in
this state the free edge stands away from the glans like a ridge.
Presumably its function is to fix temporarily the penis in the
vagina during copulation. (Text-fig. 23, L.)
The penis of Calogenys, Aeeenbed by Owen and others,
tolerably closely resembles that of Dasyprocta, except that the
horny plate is represented by a short, hard, erectile plate, the free
edge of which is armed with about five comparatively large,
widely-spaced, thorn-like teeth. In both genera there is a pair
of long spikes in the glandular pouch. (Text-fig. 23, I, K.)
In Cavia porcellus and Galea littoralis the prepuce is at the
lower extremity of the naked subeaudal tract. The glans penis
is subeylindrical and apically rounded. On each side of it
towards the apex there is a long narrow flap of epithelium, the
edge of which is serrulate with soft papille, and above this there
are a few shorter smaller flaps. The longer larger flap, no doubt,
represents the horny lustrous plate on the penis of Dasyprocta,
but it is quite soft and pliable, not rigid and horny as in that
genus. The glandular pouch is deep and provided with a pair
of long spikes lying between two strongly developed ridges on
the upper or anterior wall of the pouch. (Text-fig. 23, F-H, M.)
The penis of Dolichotis projects backwards from the lowe
portion of the naked area of skin extending up to the root of the
tail. The prepuce is moderately long, but naked. The glans
penis is slightly narrowed apically, its surface is reticulated and
pitted, and the genito-urinary orifice, just beneath the tip of the
baculum, is large. The deep, proximally narrowed, glandular
pouch is lined with rows of elongated papille representing the
laminate ridges of other genera, but there is a pair of low
thickened ridges, corresponding to the large ridges of Cavia and
Galea, passing down the anterior or upper wall. No spikes are
present and no trace of the erectile papillate lateral ridge seen in
Cavia and Dasyprocta was detected. (Text-fig. 23, A-D.)
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 409
Text-figure 23.
DP
in
Tees
. Anal region of Dolichotis patagonica, with anus dilated.
. Section of anal gland of the same.
Side view of anal region of the same, with penis in natural position.
. Glans penis of the same from the side.
Tip of glans of the same, with glandular pouch cut open.
Anal region of Galea littoralis.
. The same of Cavia porcellus, with anus concealed in glandular crease.
. The same with glandular area dilated.
s
Side view of glans penis of Calogenys paca, with glandular pouch evaginated.
. Tip of glans penis of the same, with glandular pouch cut open.
Side view of glans penis of Dasyprocta sp.?
. Tip of glans penis of Galea littoralis, with glandular pouch cut open.
a, anus; ag, anal gland; gp, glandular pouch; gw, genito-urinary orifice;
o, orifice of anal gland ; p, penis.
410 MR. R. I. POCOCK ON THE EXTERNAL
In Hydrocherus the tip of the penis when contracted is retro-
spective and close below tie anus, in contact with the perineal
integument between the orifices of the pouches of the anal glands.
When distended the base of the penis is seen to arise just below
Text-figure 24.
fie _ =»
1
i)
i
v4 Tighe
my)
= —4N
= -2%
——S—s
\S
SS
< 12-0 GF,
A es
SG
CAT
SSS
EN
SS
cS
\
—
=
>
~
. Anal and genital region of Hydrocherus capybara, with penis retracted.
. The same showing the base of the penis extended and the orifices of the anal
oy
pouches partially opened.
The same with penis omitted and the anal pouches distended.
. Secreting area at the bottom of anal pouch.
. Transverse section through one of the pores of the glandular area.
. Lower view of glans penis.
. The same with glandular pouch cut open.
a, anus; ag, anal gland; gw, genito-urinary orifice ; gp, glandular
pouch; p, penis.
QHemtee
those orifices in a position similar to that of the vulva (p. 418).
The penis is large, smooth, and cylindrical, with a narrowed apex,
beneath which lies the large genito-urinary orifice in the form of
t=)
a longitudinal slit when undilated. Behind this is the wide,
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 4\]
transverse slit-like aperture of the glandular pouch, which is
rather shallow, narrowed at the bottom with weakly ridged or
Text-figure 25.
B
ee ey Z
ZZ
= SESS
-
. Posterior region of body of Chinchilla lanigera, denuded of hair.
. Lateral view of glans penis of the same, projecting forwards.
Tip of glans penis of the same, with glandular pouch cut open.
. The same with glandular pouch entire.
. Posterior region of body of Lagostomus trichodactylus.
Anal area of the same.
End of penis of the same frem below, with prepuce slit to show tip of
slender glans.
. Penis of the same, dissected to show long, slender glans lying in preputial
sheath.
QtHtyan>
x
a, anus; ag, anal gland; gp, glandular pouch ; gw, genito-urinary
orifice ; p, penis.
wrinkled lateral and inferior walls, but without the spikes
present in Caria, Dasyprocta, Celogenys, and others. (Text-
fig. 24, A, G, F.)
412 MR. R. J. POCOCK ON THE EXTERNAL
In Chinchilla the penis lies some distance in front of and
below the anus on the pubic area between the hind legs, and is
represented externally by a long hairy prepuce projecting down-
wards and slightly backwards. When extended from the prepuce
it is seen to be subcylindrical, with a slightly expanded, truncated
extremity and with the epithelium beset with minute spicules.
The genito-urinary orifice is terminal, and immediately beneath
it is the lar ger orifice of the glandular pouch, which has longi-
tudinally laminate walls but no spikes at the bottom. Thus,
apart from its elongated prepuce, projecting freely from the
pubic integument, the penis of Chinchilla structurally resembles
that of typical Hystricomorph Rodents. (Text- -fig. 25, A-D.)
The penis of Lagostomus, however, differs in one very important
point, and is unique so far as my observations go. As in
Chinchilla, it is represented externally by a long, pendulous, hairy
prepuce, which, however, is abdominal in position, being set
much further forwards than in Chinchilla. But the penis itself,
sheathed in the prepuce, is an exceedingly long and slender rod,
apically attenuated so as to be almost filiform at the tip and
much thinner than the lumen of the preputial sheath. The
pouch at the tip of the penis appears to have aborted. (Text-
fig. 25, E—H.)
_Tallberg described the glans of the penis in a young example
«diinn und stark zugespitz, fast lanzettenformig.”
The following tabulation, setting forth the principal variations
in the structure of the penis, is based entirely upon my own
observations. In many respects it would differ materially from
a table based upon the recorded observations of Tullberg :—
a. Glans of penis exceedingly long and slender, without glandular
pouch; elongated prepuce “situated far forwards on the
abdomen......... .... Lagostomus.
a’, Glans of penis comparatively short and stout, with glandular
pouch; prepuce long or short, but pelvic in position and
gp upitavely near the anus.
. Glandular pouch with a pair of spikes at the bottom.
c. Spikes very short ....0. 6.0.0.0... .ceeseceueeeeeeeeesse--- Acanthion, Erethizon.
c’. Spikes very long.
d. Spikes biramous ...... Be te Ad BS I TRE. SR RS NO chodon.
d’. Spikes simple, undivided.
e. Spiniform papilla on penis not arranged in definite
rows; a soft pointed lappet below urino-genital
orifice, as in Octodon ....... . Ctenomys.
e’. At least one long row of spiniform “papillae on an
erectile ridge on side of penis; no lappet below
orifice of penis.
Ff. The erectile ridge with its papille quite soft and
uncornifieds: 2882 ee eee eee Galea, Cavia.
f’. The erectile ridge and its papilla forming a hard
horny plate.
g. Horny plate long with serrulate free edge ......... Dasyprocta.
g'- Horny plate short, with about five larger spines
oniitsfreeredge 22. eis. ake seee cee a en Oaelonenyss
We Me spikes at bottom of pouch. —
h. Prepuce, when penis is contracted, withdrawn into sac,
mvolving, thevanws 1 0.)5/52). ee eee ae ee ET cea
CHARACTERS OF SOME HYSTRICOMORPH RODENTS, 413
h’, Prepuce not withdrawn into common integumental sac
with anus, but remote from that orifice.
j. A small pointed soft lappet in the middle line below the
urino-genital orifice and projecting between its labia,
as in Octodon and Clenomys .........0...c cee eee cnn eee eee ees Capromys.
j’. No lappet below the urino-genital orifice.
k. Walls of glandular pouch spicular, not distinctly
laminated, with merely two low rounded ridges
running down the anterior wall...................:....... -Dolichotis.
k’. Walls of glandular pouch with numerous compressed
ridge-like lamin.
1. Apex of glans broad and truncated ..................... Chinchilla.
1’. Apex of glans attenuated 0.0.0.0... eee eens Myocastor.
The Genital Organs of the Female.
The genital apparatus in the female Hystriw and Acanthion
appears externally as a large triangular prominence, marked on
its posterior surface by a longitudinal slit. Separation of the labia
of the slit reveals the orifice of the vulva above, and just beneath
the latter the orifice of the urethral canal. Some distance beneath
the urethra and near the apex of the integumental prominence is
a trilobed clitoris, with a shallow glandular depression above or
in front of its apex. The lateral lobes can be folded over so as
to meet each other in the middle line and form a partial tube for
the passage of the urine. In the unpaired female only one orifice
is superficially detectable above the clitoris. The condition is
very similar in Atherura, except that the clitoris has a simple
conical apex with the orifice of the urethra near its tip, and
therefore farther from the vulva than in Hystrix. (Text-fig. 26,
AS Dyas)
In Erethizon and Coendu the anal and genital area of the
female is bordered on each side by a ridge of integument running
from the root of the tail to the clitoris. The vulva is a tolerably
large orifice situated a little below the anus, and the urethra
is a smaller orifice beneath the vulva and above the tip of the
clitoris, which has a glandular pouch just below and in front
of it, and the clitoris is protected by a short projection of hairy
integument corresponding to the prepuce of the male. The
condition is thus very much the same as that of Hystria# and
Atherura. In Coendu the integument round the urino-genital
orifices is naked, whereas in Hrethizon there is a considerable
quantity of hair round the orifices. (Text-fig. 26, F, G.)
In Myocastor the vulva is an arched transverse orifice close
beneath the anus in the middle of the naked area above described.
A little way beneath it is the short, conical, slightly hairy clitoris,
which is perforated at the tip by the orifice of the urethra.
(Text-fig. 27, A, B.)
In Oapromys the vulva is a little way beneath the anus, and
below it the prepuce of the clitoris, which is long in the adult.
projects downwards and backwards. It has a few hairs at the
tip, which bears a large bilabiate orifice. (Text-fig. 27, C, D.)
In Octodon the vulva is a large transverse orifice close beneath
Proc. Zoou. Soc.—1922, No. XXVIII. 28
414 MR. R. I. POCOCK ON THE EXTERNAL
the anus, and the prepuce is a short conical projection, perforated
at the tip, beneath it. (Text-fig, 27, E.)
Text-figure 26.
A. Anal and genital area of Acanthion longicauda, 2 (unpaired), with the
orifices of the urethra and vagina, dotted in, covered with mtegument.
B. Section of anal gland of the same.
C. Orifice of the anal gland of the same, distended.
D. Genital area of Hystrix africe-australis, 2 ad.
E, Anal and genital area of Atherura africana, 2 ad.
¥. The same of Coendu prehensilis, 9 ad.
G. The same of Hrethizon dorsatum.
a, anus ; ag, anal gland; e, clitoris; , prepuce ; w, urethra; v, vulya.
The vulva of Celogenys and Dasyprocta is a large wide orifice a
little beneath the anus. From it a wide inferiorly narrowing
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 415
groove extends downwards to the apex of thé short, but pro-
jecting, hairy prepuce, just above the tip of which posteriorly
may be seen the urinary orifice. (Text-fig. 28, D.)
In a female Dolichotis there is a single small genito-urinary
orifice in the lower half of the naked subcaudal area, and there
is no external sign of clitoris or projecting preputial integument.
This is the normal condition presented by immature or unpaired
female Hystricomorphs: Possibly the vulva is closed by mem-
brane out of the breeding season. In the present instance
its orifice lay, no doubt, beneath the skin between the anus
above and the genito-urinary aperture mentioned above. (Text-
fig. 28, C.)
In Cavia porcellus and aperea the vulva is near the centre of
the naked glandular anal area and separated from the anus by
the depression of the glandular sac. The very short prepuce
of the clitoris is situated some distance below the vulva. Before
pairing takes place, or out of the breeding season, the vulva
is covered with thin membrane, which easily ruptures when
stretched. (Text-fig. 28, B.)
In the female Hydrocherus, as stated below (p. 419), the vulva
is situated a short way beneath the anus, near the lower border
of the ano-genital area. It is a large, transversely extended,
dilatable orifice, containing and concealing a large, soft, fleshy,
linguiform clitoris on its ventral or anterior wall, and the orifice
of the urethra is some little distance away from the tip of this
clitoris. (Text-fig. 28, A.)
In Ctenodactylus the vulva is situated a little below the anus,
and the prepuce is elongated and perforated at the top. (Text-
£ ‘ i 9
fig. 28, H.) (See also pp. 419-420.)
a. Orifices of genital and urinary organs visible externally as
separate apertures; the prepuce and clitoris not involved in a
sphincter embracing also the anus.
6. Urinary orifice visible beneath the vulva, not concealed
within a tubular prepuce.
e. Orifice close beneath the vulva, remote from the tip of the
(REIOUIGE: WiC aennne con sbroed aubtsa can adage ao enesaccdnot Hystriv, Hrethizon, Coendu.
ce’. Orifice not close beneath vulva, but nearer to the tip of
iSIUYES U2) LDN eam OR Uae beat ehoeibnnlan dln GeuontumelieeBacee . .ae au wren: Laer
b’. Urinary orifice at apex of closed prepuce. Octodon, Capromys, Dasyprocta,
; Celogenys, Cavia, Kerodon, Ctenodactylus.
a’, Ovifices of genital and urinary organs involved in a common
integumental fold, so that externally there is a single large
urino-genital aperture, and this is involved in a sphincter
CUMHNGUNG NSLS Goo nnc dos aneladvv os seebasboncasobesdo say csovosussesceee Hydrocherus.
According to Tullberg, Thrynomys by this table falls under b’.
The Anus.
The anus in Hystrix and Atherwra opens in the middle of a
naked area, surrounded laterally and above by hairs and spines,
and inferiorly reaching to the genital orifice. There is a pair of
large, solid, anal glands opening by a slit-like orifice just within
the anal aperture, but below the termination of the rectum. The
28*
416 MR. R. I, POCOCK ON THE EXTERNAL
orifice, which has puckered lips, leads into a short narrow duct
into which the secretion of the gland is poured. These glands
appear to be equally well developed in the male and the female.
(Lext-figs. 20, EH, F; 26, A-C, E.)
The anus in the female of Coendw and Hrethizon opens near
the centre of an area, common to it and the genito-urinary
Text-figure 27.
eS Rey INE a Le
—— Qe
—— SOG ee
Sy - g
B.
ean : a.
fe os : a :
at. aX nN ff bn 5 .
a Mir
v. iO
U7
Cc
A. Anal and genital area of Myocastor coypus, 2 ad.
B. The same from the side.
C. Anal and genital area of Capromys pilorides, 2 ad.
ID. The same from the side.
ii. Anal and genital area of Octodon degus, 9 ad.
a, anus; p, prepuce: uw, urethra; v, vulva.
orifices and defined on each side by a ridge of integument. The
area above the anus is naked in Coendu, but laterally hairy in
Erethizon; and in the latter there is in addition a half-circle of
hairs just above the orifice. I did not examine fresh specimens
for the anal glands; but failed to distinguish them, if present,
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. AIT
in spirit-preserved material. In a male of Coendu nove-hispanie,
however, in which the anus opens near the centre of a very
large naked area, without integumental ridges, and with the
penis at its lower extremity, there appeared to be a median
longitudinal mass of glandular tissue, without definite orifice,
showing through the skin of the longish perineal area between
the anus and the penis. Possibly this represents the coalesced and
partially aborted paired glands of Hystrix. (Text-figs. 20, G, H
20 Gran)
In Myocastor the anus, with a few hairs round the orifice,
forms a projection in the centre of a naked area a little distance
below the root of the tail. In the female this naked area is
continued round the genito-urinary region; but in the male the
long perineal area is hairy. The anal glands in both sexes form
a solid median mass, opening by four pairs of small papillate
orifices into a sac which can be extruded by evagination just
beneath the anus. (Text-figs. 21, F-H; 27, A, B.)
In Capromys the anus is on an eminence a little way below
the base of the tail in the upper half of a naked area of skin
which extends downwards to the prepuce of the penis or clitoris.
When it is dilated the single orifice of the anal gland may be
seen immediately beneath it within the sphincter. This orifice
leads into a small pocket filled with the secretion of the gland
which lies mainly behind the pocket. The gland seems to resemble
in all respects that of Octodon,and differs from that of Jyocastor
in being much smaller, apparently unprovided with papille and
also incapable of evagination. (Text-figs. 21, A; 27, C, D.)
In Octodon the anus also opens on a prominence a little way
below the base of the tail and about the same distance above the
prepuce in both sexes, the perineal region and the area round
the prepuce being naked. The anal gland is a median mass with
a small sac opening just beneath the anus by a median orifice,
but concealed from view unless the latter is distended. (Text-
figs. 22, B, C; 27, EB.)
In Ctenomys the anus is more dilated than in Octéodon, and its
walls inside are symmetrically wrinkled ; but in the single male
example examined I was unable to satisfy myself as to the
presence of a gland such as is seen in Octodon. According to
Tullberg, however, a gland similar in position and structure
to that of Octodon is found in Ctenome Ys, Lichimys, Nelomys,
Cannabateomys, and Abrocoma. (Text-fig. 22, K, D.)
Tn Lagostomus the anus lies just Beneath a hairy exerescence
some distance below the root of the tail. Just within the
sphincter may be seen the apertures of the anal glands opening
just below the orifice. From each side of the subeireular anus
a ridge of skin runs downwards, forwards, and obliquely inwards
to meet its fellow of the opposite side in a point, the two
forming the lateral margins of an acutely angled glandular area
covered with short hair. (Text-fig. 25, E. F.)
In Chinchilla the anus also forms a marked projection some
ANS) MR. R. I. POCOCK ON THE EXTERNAL
distance below the tail, but it is not overlapped by a projecting
flap of integument. The anal glands were not detected in the
Text-figure 28.
= es ie re ~, byt
LY) Neruupet
ts Lhiy = te)
BZ
A, Anal region of Hydrocherus capybara, 2, with the orifices of the anus,
anal glands, and genital organs pulled apart.
3. The same of Cavia porcellus, ° .
C. The same of Dolichotis patagonica, 2 (? immature and unpaired).
D. The same of Cawlogenys paca, 2 ad.
HW. The same of Ctenodactylus gundi, 2 ad.
a, anus; ag, anal gland; g, common genito-urinary orifice; p, prepuce
with urinary orifice: ¢, tail; v, vulva.
single spirit-preserved example examined ; but, according to Tull-
berg, the gland resembles that of Octodon. (Text-fig. 25, A.)
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 419
In Celogenys the anus formsa large prominence near the middle
of a naked area of skin which extends in front of the tail above
and of the penis or clitoris below. The anal glands are repre-
sented externally by a pair of small pouches opening one on each
side of the termination of the rectum within the sphincter of
the anus. (Text-fig. 28, D.)
Dasyprocta seems to resemble Celogenys in having paired
lateral anal glands.
The anus in both sexes of Dolichotis is in the upper half of an
area of naked skin which extends inferiorly beneath the genito-
urinary organs, and is separated from the tail above by a fringe
of hairs. When the anus is opened the orifices of the anal
glands appear as a pair of oblique slits set one on each side above
tne termination of the rectum. The orifice leads into a shallow
hair-lined pouch. (Text-figs. 23, A, B; 28, C.)
In Cavia porcellus the anus is situated near the summit of a
large naked area which extends downwards to the prepuce in
both sexes; but it is normally concealed from view by being
folded into a depression common to it and the anal glands. This
depression appears superficially as a median groove. When this
is dilated it resolves itself in the female into a pair of pits
separated by a low partition, and situated between the anus and
the vulva. In the male the pits are much larger and longer
and when distended to the fullest extent appear as a single
capacious pouch owing to the depression of the partition between
them. When partially closed the floor of the pouch rises to
form a low partition between the pouches. A female C. aperea
resembles C. porcellus in the features mentioned above. But a
male example of Galea littoralis differs very considerably from
the male of C. porcellus. The anus is at the summit of a large
naked or nearly naked area extending down to the penis; but
this area, overlying the testes, shows no trace of glandular
depression, the anus being exposed, as in Dasyproctw and other
genera. I failed to find the anal glands in a spirit-preserved
specimen. Aerodon apparently resembles Caria in the structure
of the anal region, but I have seen no fresh material. (Text-
figs. 23, F-H.; 28, B.)
In both sexes of Hydrocherus the anus and external genitalia
are packed closely together, as if contained by a common sphincter,
upon a nearly naked prominence, some distance beneath the tail.
Between the anus and the penis or vulva there is a short perineal
area, hairier in the male than in the female, and on each side of
this lies a long vertical slit which leads into a large pouch, lined
with hair, which projects from the orifice, and filled with secre-
tion. At the bottom of each of these pouches there is a strip of
naked skin with a row of four or five little pits, and beneath
this strip the dermis is thickened and glandular. (Text-figs. 24,
A, E; 28, A.)
In Ctenodactylus the anus opens just beneath the root of
the short, tapering, uniformly hairy tail, at the summit of an
420 MR. R. I. POCOCK ON THE EXTERNAL
elongated area of naked skin, from the lower half of which the
clitoris projects. The clitoris is a closed tube perforated by the
urethra. “When cut open in an unmated female the orifice of
the vagina may be seen to lie a little below the anus, and a little
lower down is the urinary orifice. (Text-fig. 28, E.)
The following table shows the variation in position of the
apertures of the anal glands, and other points connected with
them, in the genera in whieh I have detected them :—
a. Apertures or aperture of anal glands concealed within the
ne of the anus.
. A pair of moderately peel septiata) aperiaie above the anal
orifice ........ debe dhe TM alseen oer ee ERR oe ean letelc eae) DOLLCHOLIS:
’, Apertures not ‘above the anus.
ce. Aperture on each side of the anus .................... _ Dasyprocta, Cologenys.
c’. Aperture or apertures beneath the anus.
p
d. ‘Two apertures, one on each side of middle line .... Hystrix, Lagostomus.
d’. A single median aperture.
e. Anal gland small, incapable of evagination ......... Octodon, Capromys.
e’. Anal gland large, capable of evagination and provided with
four pairs of papille .......... .... Myocastor.
a’, Apertures of anal glands forming a pair of elongated slits outside
the sphincter of the anus and extending down the perineal area.
Cavia, Kerodon, Hydrocherus.
The interesting point to note in connection with this table
is the difference between Dolichotis and Cavia+ Hydrocherus
with respect to the glands.
According to Tullberg’s descriptions, Thrynomys falls by this
table with Cavia and Hydrocherus under a and Ctenodactylus
with Dasyprocta and Celogenys under c.
The Tail.
The tail supplies useful systematic characters. If, as is
probable, a long cylindrical tapering tail covered with scales and
short hairs is the pr imitive type, that type is prevalent in the
Octodontide, occurring in many of the mouse-like genera
Within the family, howerer, variations in the length of ane tail,
jn adaptation to habit, and in growth of the hairs are numerous.
In Octodon, for example, although the organ is longish, the hairs
in the distal portion are developed to form an elongated tuft.
In Ctenomys it is shorter, but thicker and somewhat compressed
with a very distinct crest of short hairs extending along the
posterior two-thirds of the upper edge.
In Thrynomys the tail is of the primitive type, but compara-
tively short, and the same applies to Capromys, but in this
seansorial genus the hairs on the underside are stiffened to aid
in alias. a variation foreshadowing that which is seen in a
much more marked degree in Coendu and Chetomys described
below. In one species (C. prehensilis) the tip of the organ
is said to be prehensile, but I am not aware whether the tip
is curled downwards over a branch or upwards as in the so-
called arboreal Poreupines. In Myocastor the tail is moderately
long, cylindrical, and rather stout, but shows no modifications
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 421
subservient to aquatic life. It is used mainly as a rudder, hardly
as a propeller, in swimming.
In the Hystricide the tail is always provided with a sounding-
organ formed of modified hairs. In the most primitive type
Trichys, in which the armature of the body-skin consists of a
coating of very coarse sharp bristles, nearly uniform in length, the
tailis long and cylindrical, and covered, except at the root and
tip, with scales and short hairs; and the sounding-organ or rattle
at the tip consists of a brush of long flattened hairs or spines
somewhat resembling dried blades of grass. In Atherura the
tail, although still scaly and hairy, is much shorter, and each of
the constituent parts of the terminal brush consists of a filiform
axis expanding along its course into a series of compressed but
hollow fusiform swellings, from two to seven or eight in number,
according to the length of the blade. These swellings vary in
size, but the one at the end is always much larger than the next
of the series.
In Thecurus, Acanthion, and Hystriv, in which the spine-
armature reaches its maximum of development, the tail is quite
short and thick and without scales and hairs; but twenty or
more of its terminal quills are expanded into hollow flattened
lamin or cylinders, for the most part open at the end. This is
the most highly specialised rattle in the group.
The so-called arboreal Porcupines of America (Erethizontide)
fall into two sharply defined groups by their tails. In the North
American genus Hrethizon this organ is quite short and armed
throughout with spines resembling those on the body. By
swinging it to right and left, the animal uses it as a weapon of
defence. Inthe tropical American genera Coendu and Chetomys
the tail is long, subeylindrical, tapering, and covered with spines
and hairs and distally with scales, although the extremity, which is
upceurled and prehensile, is naked above. The underside at the
base is thickly covered with close-set, stiff, sharp bristles, the
function of which, as Waterhouse rightly supposed, is to help
in the ascent of vertical or steeply sloping branches, and to give
support to the body when the animal is at rest. They are func-
tionally comparable to the caudal scales of the Anomaluride.
In the three genera assigned to the Chinchillide, namely
Chinchilla, Lagidium, and Lagostomus, the tail is also charac-
teristic. It is moderately long, covered with hairs, which are
comparatively short and soft all along the underside, but on
the upperside are coarse and very long—forming, as it were,
a brush.
In the Dasyproctide and Caviide the tail is reduced. In
Myoprocta it is at most a few inches long, but quite slender.
In Dasyprocta it is only about an inch or less. In Calogenys
it is about the same. In Dolichotis it is also quite short, but
constricted at the base and oval in outline from above or below.
In Hydrocherus it is at most a short conical excrescence as it is
in Cavia, Galea, and Kerodon, and is sometimes absent.
493 ‘MR. R. I. POCOCK ON THE EXTERNAL
Notes on the Families and Subfamilies.
Family Hysrricipa.
Since Lyon (Proc. U.S. Nat. Mus. xxxii. p. 576, 1907) has com-
paratively recently classified the Hystricide into the two sub-
families Atherurine (Atherwrus, Trichys) and Hystricine
(Thecurus, Acanthion, Hystrix), mainly by the structure of the
tail, I need not refer to these animals further beyond expressing
my complete concurrence with his opinion as to the systematic
value of the characters he uses as a basis for the separation of
the two subfamilies.
Family HREeraizontipé.
By their external characters the three genera (Chetomys,
Coendu, Hrethizon) composing this well-marked family fall into
two groups, the first represented by Hrethizon, the second by
Coendu and Chetomys. By cranial and dental characters, how-
ever, Mr. Thomas separated Chetomys from the others as repre-
senting a special subfamily Chetomyine, and assigned Ooendu
and Hrethizon to the Krethizontine. Adopting the separation of
Chetomys from Coendu, I think the latter should be similarly
separated from WHrethizon and the family divided into three
subfamilies, which may be diagnosed as follows * :—
a. Tail long, cylindrical, comparatively slender, and prehensile;
hind feet with a large movable lobe on the inner side and
minute hallux; fore feet also expanded on the inner side;
nostrils widely separated; ear specialised, with large antitragus.
a’. Body covered with flexible spine-like waved bristles; skull
with smal] orbit circumscribed by frontal and jugal ae
orbital processes; jugal arch very deep ; palate, tooth-rows
and symphysis of mandible Jong ........ See aeekeee eens RC iucenomnanivees
b'. Body armed with comparatively sistas fneaell spines ;
orbit large, no eae processes; jugal arch not deep ;
palate, tooth-rows, and mandibular sy Pee much
INOUE sad scoot SEE SSHOe BS a. \Coendine:
6. Tail short, thick, noe arlhansille iin foot “afl at me asmalll
inner lobe, but with well-dev eloped ha!lux; fore feet without
expansion on inner side; nostrils with narrow septum ; ear
SIMPLE: Ketone eae peat de | ces iect RON: co elen bomb acem eee. LeRen CP EeeEE Me REME nCUILEZ ONLI.
At present each of these subfamilies contains a single genus ;
but it seems to me to be probable that the first modern syste -
matist who has adequate material of Coendw, and the time to
devote to the study, will find characters justifying the separation
of that genus into two or more genera. There appear to me to
be no good reasons for thinking the Erethizontide especially
related to the Hystricide—a conclusion to which other authors
have come. The Erethizontide are, I think, probably very
* The external characters, apart from the ear, here made use of were long ago
pointed out by Waterhouse. It is, however, merely a guess on my part that the
nostrils and ears of Chetomys are like those of Coendu.
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 423
Specialised descendants of forms akin, or belonging, to the Octo-
dontide, whereas the Hystricide do not seem to be specially
related to.any of the South-American groups. In that case the
resemblances between the Poreupines of the Old and New Worlds
which have led to their affiliation must be due to convergence or
to the common inheritance of ancestral characters. In their
Spine- armature, for instance, it may be noted that in the two
genera which appear to me to be the most primitive of the two
families respectively—namely, Chwiomys and T’richys—the spines
are little more than very stiff bristles. It must be rememberec,
however, that there is one character, not previously recorded
apparently, in which the two families are alike and differ from
other Hystricomorphs—the prepuce in the female does not form
a closed tube, the orifice of the urethra being exposed beneath
the genital aperture.
Families Ocropontip#, PErRomyID&, and CrenoDACTYLID®.
I have seen too few examples of the Octodontide to offer any
opinion as to its subdivision into subfamilies. Petromys, too, I
have not seen, and I do not know whether it should be referred
to the Octodontide, where it was originally placed, or, in accord-
ance with Tullberg’s views, made the type of a special family.
Probably the latter is the better way of regarding it for the
present, in view of the differences of opinion that prevail concern-
ing its status. The structure of the ear alone seems to me to
justify its separation from Ctenodactylus, with which Thomas
associated it. The claims of Ctenodactylus, indeed. to a place in
the Hystricomorphs seem to me to be more than questionable.
Families CApRoMYID& and Myocasrorip&, nov.
The family-name Capromyide may be restricted to Capromys and
related genera, like Procapromys, Geocapromys, and, 1 presume,
Plagiodontia.
Myocastor, formerly associated with these genera, may, I think,
be regarded as representing a family by itself, Myocastoride.
This course, however, merely amounts to giving greater systematic
value to the characters used by Tullberg when he established the
subfamily Myopotamini.
Family TaryNnomyip&, nom. nov.
(=Aulacodide of Tullberg).
The genus Thrynomys, formerly classified with the Octodontide,
and later with the Capromyide, was separated as the represen-
tative of a special family Aulacodide by Tullberg, who employed
its old name Awlacodus. Agreeing with this decision, I adopt
Thrynomyide as the family-title.
AQ4 MR. R. I. POCOCK ON THE EXTERNAL
Judging from dried skins, the genus is distinguished by the
cleavage of the upper lip by the rhinarium, the valvular supra-
tragus—a character repeated in the Caviine, and the structure
of the feet. ‘The affinities of the family appear to me to be quite
doubtful.
Family Dinomyip&.
I can give no opinion about Dinomys beyond expressing my
pony Clare that it is not in any way nearly related, as Peters
thought, to the Dasyproctide, Chinchillide, or Caviide in the
sense in which he understood those terms. It must remain as
the sole representative of a well-marked family.
Families DAsyprocr1p# and Ca@iLoGENyIDA, nov.
So far as Iam aware, no suggestion has been made to break
up the family Dasyproctide as understood by earlier authors.
The tendency rather has been to merge it with the Caviide, as
was done by Winge, Tullberg, and Weber. Three genera are
now admitted — Das, yprocta, Myoprocta, and Calogenys. The first
two are closely related ; but it appears to me that the value of
the well-known distinctive features of Celogenys have been
greatly underrated. I propose to erect it to the rank of a family,
which may be distinguished from the Dasyproctid (s.s.) as
follows :—
a. Antemolar portion of palate broad and nearly flat; preeorbital fora-
men, zygomatic arch, and orbit without special modifications ;
nasals as long as frontals or nearly so; postorbital part of skull
comparatively long and narrow; occipital crest irregularly semi-
elliptical. No cheek-pouches. Feet long and slender; fore foot
nearly perissodactyle, with fifth digit tolerably widely separated
trom fourth; plantar pad bilobate, the outer lobe, if represented,
indicated merely by a small circular scale at the base of the fifth
digit and remote from the rest of the Bie Hind foot with only
three toes and a small plantar pad . Pshobooa es csabopnondea MOVASIvcenrNoxourine yan)
a’. Antemolar portion of palate strongly ieaneeeete eae a median
ridge deeply grooved and bicarinate almost to the incisors; the
whole zygomatic arch profoungly modified mainly by the out-
evowth from its maxillary and’ malar portion of a great bony
lamina forming a cheek-plate extending downwards to overlap and
conceal the greater part of the mandible; the maxillary portion of
the arch is deeply hollowed beneath and is continued forwards as
far as the premaxilary suture, the roof of the hollow forming the
floor of the preorbital foramen which is converted into a long deep
channel in front of the orbit, and the orbit itself is reduced in
dimensions by the upgrowth of its inferior edge and looks obliquely
outwards and upwards; nasals much shorter than frontals, post-
orbital portion of skull wide and short, occipital crest nearly sem1-
circular. Large cheek-pouches present. Feet short and robust,
fore foot nearly artiodactyle, with four main digits evenly spaced,
the plantar pad large with outer lobe well developed and confluent
with median ; hind foot with five digits ....................... C@LOGENYIDS.
So far as the feet are concerned, Calogenys is a much more
primitive type than Dasyprocta, and might be regarded as
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 425
ancestral to the latter genus; but with respect to the modifications
of the skull Calogenys is extraordinarily specialised and absolutely
isolated, nothing approaching an intermediate type between it
and Dasyprocta being known,
Family CHINcHILLIDA,
In all the classifications quoted above this family contains the
three genera Chinchilla, Lagidiwm, and Lagostomus. Of these
Chinchilla and Lagostomus are the only ones I have been able to
examine in a fresh state, Lagidiwm being known to me solely
from a dried skin, a skull, and the descriptions of other authors,
which possibly from want of material do not contain the points £
want. But, judging from the cranium and the teeth and the
external characters revealed by the dried skin, Lagidiwm is more
nearly related to Chinchilla than it is to Lagostomus—that is to
say, the family may be divided into two groups or subfamilies,
one containing Lagostomus alone, the ether Chinchilla and
Lagidium. In using the external genitalia of the male as
perhaps the most important difference between these subfamilies,
fT assume provisionally that Lagidiwm will be found to agree with
Chinchilla. The characters may be tabulated as follows :—
a. Penis normal in position and structure, glans stout and mcde-
rately long, with well-developed glandular pouch. Fore feet
with trilobed plantar, bilobed carpal pads, and compressed
digital pads; hind foot with fifth digit retained, with a large
pad on the sole of the foot at its base, digital pads com-
pressed ; principal digital brush on the second digit. Root
of ear raised externally so as to stand high above auditory
aperture. No coarse vibrisse on cheek. Molar teeth tri-
laminate. Postorbital area of skull sloping backwards so
that the weak occipital ridge is approximately on a level with
the middle of the orbit; naso-premaxillary region not down-
curved; ‘“ prepalatine foramina” long and ‘“‘Steno’s fora-
men” not remote from MCISOYS ..,...-.. see eee eee eeeeeecneeeseeeane Chinehilline.
a’. Penis very abnormally piaced, the prepuce being abdominal
in position; glans exceedingly long and slender, with pouch
apparently undeveloped. lore feet with indistinctly lobed
plantar and carpal pads and digital pads not compressed ;
hind foot without fifth digit and no trace of isolated pad on
the sole. An immense digital brush on the third digit.
Root of ear not raised so as to conceal auditory orifice. A
mat of coarse vibrissee on the cheek; normal genal vibriss
stout. Molar teeth bilaminate. Postorbital area of skull
not sloping backwards, the very strong occipital crest as high
as the summit of the orbit ; naso-premaxillary region arched
downwards; “ prepalatine foramina” comparatively short
and “ Steno’s foramen” remote from the incisors ............... Lagostomine
Families Cavitp® and HyprocH@rip&, nov.
Tn the current text-books the Caviide contain the three genera
Cavia, Dolichotis, and Hydrocherus; but the comparatively recent
severance of Cavia into some half-dozen genera—Cavia, Caviella,
Monticavia, Galea, and Kerodon—enhances the value of the
426 MR. R. I. POCOCK ON THE EXTERNAL
characters which distinguish Dolichotis from Cavia in the old
sense, and Hydrocherus from both. In the first place, Zydro-
cherus clearly differs from Dolichotis and Cavia much more than
these two differ from each other. As set forth in the table that
follows, Dolichotis also in certain well-defined particulars stands
aside from all the genera into which Cavia has been split up. I
propose to express these differences by severing Hydrocherus as
the representative of a distinct family * from the Caviidee, and to
divide the latter into two subfamilies Dolichotine and Caviine.
The distinguishing characters of these groups may be briefly
stated as follows :—
a, Last upper molar longer than the three others taken together; palate
long, its median line about twice its width, the palatine bones
not deeply excised by short mesopterygoid fossa. Feet webbed
up to ill-defined digital pads, which with the claw form a hoof-
like termination to the digits; digits of fore foot unevenly
spaced, the fifth set high above the fourth and carrying a
reduced and separate interdigital element of the plantar pad ;
the third digit markedly larger than the second or fourth and
submedian ; no additional ridge on ear ........................ HypRocH@rip a.
6. Last upper molar much shorter than the combined length of the
others ; palatine bones so deeply excised by the mesopterygoid
fossa that the median length of the palate is at most about equal
to the width between the last molars. Feet not webbed; dizital
pads well defined and distinct trom the compressed claws; digits
of fore foot tolerably evenly spaced; third and fourth digits
nearly evenly paired, subequal, the third not markedly larger
than second and fourth; plantar pad with external lobe con-
fluent with median lobe; a strong ridge beneath the supratragus
LENS: Gar Te eave ase 1A nes NL En ena oe Na Re an Pat a Sa OUACW TIED OIDs
a*. Skull with nasals narrowed and strongly convex in front, the
distal edge deeply excised; the interorbital region very wide
owing to an expansion, notched anteriorly and defined by a
notch behind, overhanging the orbits; upper edge of malar
geniculate, paroccipital processes long. Har long, with simple
supratragus. Nostrils expanded. A single pair of widely
separated anal glands above the anus. Digits short, legs long,
with area behind plantar pads overlapped by hairs; plantar
pads very large, projecting behind; carpal pad very small;
audistinetytan! Ress ee eee ea eee inner een en eam OT CHG iIICEs
é?. Skull with nasals nearly parallel-sided, not strongly convex in
front, and distal edge with shallow excision; no appreciable
bony growth roofing the orbit, upper edge of malar not geni-
culate ; paroccipital processes comparatively short. Ear short,
with valvular supratragus. Nostrils wot expanded. Anal
glands, where known, opening beneath the anus. Digits more
elengated, legs short, naked beneath, plantar pads not project-
rouge lorelonncsls Kevyn jakaGl Tee, TVG) WANT. sasnooanoon soonaacnosaan: Caviine.
Other differentiating characters might have been added to
those given above. But these must suffice.
* T submit that the characters upon which Hydrocherus may be separated as a
family from the Caviidee are of higher systematic value than those upon which
Castor has heen separated as a distinct family from the Sciuride (see Miller, Mamm.
of Western Europe, p. 947, 1912).
CHARACTERS OF SOME HYSTRICOMORPH RODENTS. 427
Mainly by characters supplied by the skulls and teeth, Thomas,
in the paper quoted above, showed how the genera of Caviin
might be distinguished. The three of which I know the external
characters may be differentiated as follows :-—
a. Ridge of ear beneath supratragus simple, not valvular: interramal
vibrisse consisting of two pairs of long seta set on the posterior
border of a large nearly naked area behind the chin ; anus exposed
at the summit of a naked or nearly naked area of skin which lies
over the testes and shows no trace of glandular depression ; penis
DLO;EctinaAROMAlOweendio: this area ee ee) 8 Conair ie
a‘. Above-mentioned ridge valvular; interramal vibrisse absent or at
all events typically indistinguishable ; anus concealed in the sub-
jacent glandular area.
5. Claws elongated; digital pads flat or lightly convex; a single
carpal pad Pitas Ponts : Cavia.
b'. Claws quite short ; digital pads compressed ; carpal pad double... Kerodon.
valor sas lama o sg
rin
Ni hk — F + ras nike poly
at A “ ee Mes ioe
ey ' Ni fan fe cA \
4h
th cenit sin
iat ag ey
ON THE ANATOMY OF THE DRILL. 429
22. On the Anatomy of the Drill (Mandrillus leucopheus).
By Cuarues F. Sonnrac, M.D., F.Z.8., Anatomist to
the Society.
[Received March 23, 1922: Read April 25, 1922.]
(Text-figures 9-24.)
ConrTents.
Page
My ology gieererry ceriteeen enter rete en at oN Snakes castashniie yn LOO
Alimentary Canali wea rolte tse sys oc so tnorsasaatosn aa 4D
Salivary Glands, Pancreas, Liver .............................. 488
Wwctless:Glandope ea mere esd. 1. ae eter issn ast ABO
Whotnoerermieell (OWENS convce 2 apocbesdecedesvenanesanasancoosecena LD
OeENS OF CHCTIOD sonscocosese coo peaconcou san cos sasvoaccoconace GLIl
OvFEHNAS OFF INGE VUE cs ccnnvse 7 bopn0so8s02n0n eon dehonbaoncnasen ZlZldh
NervyousiSystenmacearr ert ts cat nou ei eee aac soocusimtucn ss AAA
SummanygandaConclustonspeeeteee step eeee perce eee ene 4 O2
Bibliooxarp lity meena cesc: ses. ack teens ene aencr etetteas 458
The literature of the Primates contains very few references to
the structure of the Drill, so an account of its anatomy should
be useful. The present description is based on the examination
of an adult male which died in the Society’s Gardens. It had |
the following proportions :—
Length from tip of nose to root of tail ... 29-5 inches.
Teall “14. ees ne ee ta acc oo SOEs
ene tht, oftheadiy saass. ea ee 9 Be
ie. DOE AAR a gee AMM ene D ADR) oe
CASEIN Sesleri O 8 a
Horeanis 7 eee e OPER aie re anaes Ne SiO ges
AN ch sd Ammer MRSC ic io, Ac ee UN ne 2 ee Seal an
LOS a ge oaiiciuccadls Sateevecvgmeen ty Witics yoke Aa tnat sed Sua ts:
MyoLoey.
The platysma myoides is strongly developed in both neck and
face, and has a well-marked attachment to the lips. The muscles
of the lips and frontal region are also very strong and thick; and
it is to their great development that the frequent movements of
scalp and exposure of the teeth are due.
The temporal and masseter nvuscles are powerful, and the upper
part of the latter is concealed by the parotid gland (text-fig. 9).
But the full, rounded muscle is seen beyond the limits of the
gland. The former appears as a well-marked prominence above
the zygoma as in many of the Carnivora.
The arrangement of the vessels and nerves on the surface of
these muscles differs from that in many Primates and Carnivora,
Proc. Zoot, Soc.—1922, No. XXIX, 29
430 _ DR. C. F. SONNTAG ON THE
Sterno- and Cleido-mastoid:—Both muscles are powerful, but
there is no gap between them at any point. The former arises
from the anterior border of the manubrium sterni and margin of
the sternal end of the clavicle (text-fig. 10 A,8.M). The latter
originates from the inner fourth of the clavicle, and gradually
gains the deep surface of the sterno-mastoid. Both muscles are
inserted into the mastoid region and occipital crest. There is no
separate cleido-occipital. The spinal accessory nerve is deep to
both muscles. In some Primates there is no clavicular fascicle.
The omo-hyoid is flat, thin, undivided into bellies, and has no
central tendon. The hyoid and scapular attachments show
nothing peculiar, but there is no fascial connection to the
sternum.
The pretracheal muscles show nothing peculiar.
Text-figure 9.
,
< \\
\
CORY
CH
ry
yd
\ ;
0 \
@ \\
of
>
e
Os)
ti
ine
ct
The superficial anatomy of the side of the head, after the skin and platysma have
been removed. G.A.N: great auricular nerve; O.N: occipital nerve; S.M:
sterno-mastoid; P.G: parotid gland; O.A : occipital artery; E.C.A : ectocarotid
artery ; I.P.V : inferior labial vein; S.T.V: superficial temporal vein; E.J.V:
external jugular vein; M.M: masseter muscle.
The digastric belongs to Parson’s first type (4), but no fascia
unites it to the hyoid bone. And there is no essential difference
between it and that in Macacus rhesus*. The posterior bellies
are very strong, and the tendinous arch is long and powerful.
The stylo-mandibular ligament is partly ossified, and gives
origin to the stylo-hyoid muscle which is tunnelled by the inter-
mediate tendon of the digastic. The stylo-hyoid ligament, how-
ever, is not ossified.
The mylo-hyoid is strong and thick, but its attachments show
nothing peculiar.
The Ayoglossus separates the hypoglossal nerve from the
* C, F, Sonntag, P. Z.S, 1919, p. 437.
ANATOMY OF THE DRILL. 431
lingual artery and branches of glosso-pharyngeal nerve as in Man,
and the well-marked submaxillary gland lies on its surface. It is
united to its fellow to form one sheet. It is continuous with the
sterno-hyoid.
The pectoralis major (text-fig. 10 A) is strong and complex, and
consists of several parts. ‘The first is strong and thick, and
arises from the margin of the sternal end of the clavicle, the
entire length of the sternum, the inner ends of the costal carti-
lages, and the aponeurosis over the rectus abdominis in the infra-
sternal fossa. Its superficial fibres fuse with those of the deltoid.
Text-figure 10.
Muscles of the shoulder-girdle. A: superficial aspect; B: the humerus on section
across the pectoral crest ; C: structures under the pectoralis major and deltoid
C.M: cleido-mastoid; P.M: pectoralis major ; Su.M: subclavius; T: trapezius ;
T.A: thoracic axis artery; C.V: cephalic vein; Cl: clavicle; D.F: deep
fascia; R.L: first rib; T.M: teres major; L.H.B: long head of biceps; P.C:
pectoral crest.
The second part covers the inner part of the chest-wall and part
of the external oblique aponeurosis, and joins the deep aspect of
the first part which is inserted into the capsule of the shoulder-
joint and the prominent pectoral ridge on the humerus. A third
part arises from the second, third, fourth, and fifth costal carti-
lages and intercostal muscles in the intervening spaces, and is
inserted into the capsule of the shoulder-joint and deep part of
29*
432 DR. C. F. SONNTAG ON THE
the preceding part. It works over a strong cord running from
the first rib to the head of the humerus (text-fig. 10 C, C.C.L).
And this ligament may replace the costo-coracoid ligament in
Man. The first and third parts receive separate nerves from the
median nerve, but the latter probably only serves as a track from
the brachial plexus (text-fig. 23).
The pectoralis minor may be replaced by the third part of
pectoralis major.
A long, narrow muscle, composed of two bellies, takes origin
from the free edge of the latissimus dorsi, crosses the axillary
vessels and brachial nerves, and spreads out into a thin fan-
shaped aponeurosis. The latter covers the coraco-brachialis and
biceps, and is inserted into the deep surface of the insertion of
pectoralis major. It has a double nerve supply from the second
intercostal nerve and brachial plexus (text-fig. 11, G).
The subclavius (text-fig. 10C) is very strong. It arises from
the inner end of the first rib by a strong pointed tendon, and is
inserted into the inner five-sixths of the posterior surface of the
clavicle.
The delto-pectoral triangle (text-fig. 10 A, D.P.T) is long and
narrow. It contains the cephalic vein and thoracic axis artery,
but no lymphatic glands.
The skin over the pectoral region is supplied, as in Man, by
the supraclavicular nerves from the cervical plexus.
The trapezius arises from the inner part of the occipital crest,
the inion, ligamentum nuche, seventh cervical and upper six
dorsal spines, and supraspinal ligaments. And the thick, strong
fibres converge,asin Man. They are inserted as in Man, but the
fibres which are attached to the root of the spine of the scapula
end in a very strong triangular tendon, under which the spinal
accessory nerve passes to the posterior part of the muscle. The
sub-trapezial nerve plexus formed by the spinal accessory and
cervical nerves is well-developed.
Latissimus dorsi:—No fibres are derived from ribs and the
posterior angle of the scapula, and the upper fibres of origin are
under cover of the trapezius. They extend from the fourth to
the last dorsal vertebra, behind which they arise from the lumbar
aponeurosis, but no fibres extend back to the ilium. The fibres
converge, wind round the lower border of the teres major, and end
in a very strong, flat tendon which is inserted into the humerus
at the base of the pectoral crest (text-figs. 10, L.D and 11, H).
The tendon is intimately connected to the teres major and dorso-
epitrochlearis.
The levator anguli scapule arises from the dorsal surfaces of
the transverse processes of the first four cervical vertebra, and
the slip from the first one is the largest. The fibres unite to
form a thick muscle, which is inserted into the vertebral border
and deep surface of the scapula in its anterior part.
The rhomboids form a strong muscular sheet arising from the
vertebral spines from the seventh cervical to the fifth dorsal ;
ANATOMY OF THE DRILL. 433
and it is inserted into the vertebral border and margin of the
deep surface of the scapula. It is impossible to separate it into
major and minor rhomboids. The sheet does not extend to the
occiput as it does in many Primates.
The serratus magnus arises from the first ten ribs and inter-
spaces, but the origin forms a uniform muscular line instead of
pointed slips as in Man. Its insertion into the vertebral border
of the scapula is thick and muscular.
The omo-trachelian is a powerful muscle attached to the supra-
spinous fossa and root of the spine of the scapula.
The dorso-epitrochlearis (text-fig. 11, J) is a very strong muscle.
It is 42 inches long and 12 broad. Proximally it is intimately
connected to the latissimus dorsi and teres major. Its rounded
distal border is bound by a strong tendon to the internal condyle
of the humerus and the olecranon. It is supplied by a branch of
the musculo-spiral nerve (text-fig. 23, D.E.M).
Text-figure 11.
The structures in the axilla and arm. A and C: short and long heads of biceps ;
B: coraco-brachialis; D: fascia; E: pectoralis major; F.G: axelbogen; J.K:
dorso-epitrochlearis; L: internal condyle; M: olecranon; N.O.P: axillary
vessels and nerves; R.S: long subscapular artery and nerve. Other letters
in text.
The deltoid (text-fig. 10, D.M) is very powerful, and its super-
ficial fibres are fused with those of the pectoralis major. It has
an extensive origin from the whole of the ventral surface of the
clavicle except the sternal end, the acromion, and outer two-thirds
of the spine of the scapula. And the fibres arising from the
acromion are tendinous. All converge to the usual deltoid
insertion.
The teres major (text-fig. 11, I) has the same origin and insertion
as in Man, but it is intimately blended with the latissimus dorsi
and dorso-epitrochlearis. The teres minor has also an origin
similar to that in Man, but its tendon of insertion is pushed on te
the humerus distal to the greater tuberosity.
434 DR. C. F. SONNTAG ON THE
The subscapularis arises from the entire subscapular fossa, and
has a great admixture of strong tendinous fibres. The latter
converge and pass over the lesser tuberosity to be inserted into
its upper border. The tuberosity with these tendons presents a
full, rounded appearance.
The supraspinatus arises from the outer two-thirds of the supra-
spinous fossa and is inserted into the upper part of the great
tuberosity of the humerus. The infraspinatus arises from the
outer two-thirds of the infraspinous fossa. It is inserted by two
heads into the great tuberosity of the humerus and the bone
beyond it, so as to displace the insertion of the teres minor
distally (text-fig. 12 C).
The biceps arises by two heads as in Man, the short one joining
the long one in the upper third of the arm. The muscle con-
verges to a long, flat tendon which is inserted into the dorsal
Text-figure 12.
A+: manubrium sterni of Drill and Mandrill*. B: flexors of elbow-joint; C:
muscles attached to the great tuberosity of the humerus. B.A: brachialis
anticus; Bi: biceps; D.M: deltoid; I.S: infraspinatus; S.8: supraspinatus;
T.Mi: teres minor; I.L.L: internal lateral ligament.
aspect of the proximal part of the radius from the head to the
point where the bone begins to exhibit its marked forward con-
vexity (text-fig. 12B). It first narrows slightly and expands
later. There is no bicipital fascia as in Man.
The coraco-brachialis has a long, powerful tendon of origin and
an elongated fleshy belly inserted into the shaft of the humerus
close to the deltoid insertion.
The brachialis anticus arises as in Man. It has a very strong
tendon which winds round the internal condyle of the humerus,
and is inserted into the upper inch of the inner surface of the
shaft of the ulna, and the internal lateral ligament of the elbow
joint (text-fig. 12 B).
The triceps is a very powerful muscle. Its long head arises
* Vrolik states that the manubrium is absent in the Mandrill.
ANATOMY OF THE DRILL. 435
from the lateral two-thirds of the axillary border of the scapula,
and some fibres fuse with the proximal part of the dorso-epitro-
chlearis. The two humeral heads have extensive areas of origin.
The strong tendon of insertion passes to the dorsal aspect of the
extremity of the olecranon, but some fibres pass along the side
of the tendon to be attached to the proximal inch of the lateral
border of the ulna.
As the hands were not at my disposal, no attempt was made to
dissect out the muscles of the forearm.
The sartorius arises from a narrow area on the vertral border
of the ilium; and its origin bounds an arch through which the
Text-figure 13.
Muscles of the thigh. A.M: adductor magnus; A.lL: adductor longus; G: gracilis ;
S: sartorius; Q.E: quadriceps extensor attached to internal femoral condyle
(1.C.F.) and patella (P.); A.C.N., E.C.N., G.S.N: anterior crural, external
cutaneous and great sciatic nerves.
lateral cutaneous nerve of the thigh emerges. It is inserted into
the front of the shaft of the tibia distal to the imsertion of the
uadriceps extensor. ‘To reach that point it describes a curve
like the italic f. Jt helps to bound three triangles—a deep
Scarpa’s Triangle whose boundaries areas in Man, a triangle whose
other boundaries are the adductor longus and gracilis, and a tri-
angle whose other boundaries are the quadriceps tendon and
internal femoral condyle. In the latter triangle the tendon of
the adductor magnus is seen passing the internal condyle to reach
the tibia (text-fig. 13).
436 DR. C. F. SONNTAG ON THE
The gracilis (text-fig. 13) fuses with sartorius at its insertion,
and the combined muscles are inserted along one-third of the
length of the ventral border of the tibia.
The adductor longus (text-fig: 13) has a long attachment to
the femur, and its distal limit is about half an inch above the
internal condyle.
The adductor magnus (text-fig. 13) is a long, narrow muscle
whose tendon is attached to the upper end of the tibia.
The great sciatic nerve divides at the bottom of the triangle
formed on the inner aspect of the thigh by the sartorius, gracilis,
and adductor longus.
The anterior crural nerve divides into two main branches in
Scarpa’s Triangle.
The external cutaneous nerve breaks up into a number of
branches over the quadriceps, and communicates with the anterior
crural nerve.
The quadriceps extensor is inserted into both condyles of the
femur and upper border of the patella, and the ligamentum
patelle is attached to the tibia over a length of two inches. At
the lower limit of the latter is the long strip of attachment of
the sartorius and gracilis. The vastus internus component is
extremely powerful.
The glutei have coarse, thick fibres, but they are not voluminous
as in Man. The gluteus medius is much thicker than the
maximus.
The levator ani is a very stong, thick muscle, but the pyri-
jformis is very long, thin, and slender. It cannot be employed in
subdividing the structures passing from the pelvis to the thigh
as in Man.
The great sciatic nerve gives off a very thick cord to the ham-
strings after it winds beyond the femoral trochanter. These
muscles are inserted into the upper half of the dorso-lateral
border of the tibia.
The gluteus minimus and lateral rotators of the thigh form a
thick fan-shaped muscle, with many tendinous fibres originating
along the entire vertical length of the ilium.
THe ALIMENTARY CANAL.
Fleshy dips are connected to the gums by well-marked frenums,
but no labial tubercles are present.
The oral vestibule is smooth and has well-developed cheek-
pouches.
The hard palate is crossed by twelve pairs of incomplete ridges
radiating from the mid-line, but there is no prominent median
raphe, and no incisive pad; the ridges become more curved on
the posterior part of the palate, and those of the last two pairs
are sinuous.
The soft palate has « well-marked uvula.
ANATOMY OF THE DRILL. 437
The tonsils are two round bodies, the size of large peas,
situated in the fauces, and the mucous membrane over them is
fenestrated.
The esophagus does not project from behind the trachea in the
neck, and it is very capacious in the posterior part of the thorax.
Its intra-abdominal part is short. ‘The mucosa is smooth through-
out. Many branches of the vagi ramify over its surface just
before it passes through the diaphragm.
The stomach (text-fig. 14 A), which was empty in this specimen,
has no well-marked fundus, but the posterior part of the body
was directed posteriorly and to the left. The pyloric region is
not firmer than the rest of the stomach, is directed forwards
Text-figure 14.
The stomach (A,) and cecum (B.).
(craniad) and slightly to the right, and is marked off from the
duodenum by a constriction. Vrolik (3) describes a globular
stomach in the Mandrill.
When the stomach is opened along the greater curvature it is
seen how there are no rug, and no marked pyioric sphincter.
The duodenum makes a sharp bend and runs caudad. There is
no well-defined loop, and it passes insensibly into the beginning
of the jejunum. Valvule conniventes are absent, and there is
no bile papilla, but merely an orifice.
The entire small intestine is 82 inches long, and the large
intestine measures 60 inches without the cecum.
438 DR. C. F. SONNTAG ON THE
The cecum (text-fig. 14B), 3 inches long. is wide and
capacious, and is sacculated by longitudinal bands on its dorsal
and ventral surfaces. It has a well-marked mesentery, but no
artery occupies its free edge... There is no appendix vermi-
formis. :
The ileo-ewcal orifice is guarded by a circular valve.
Peyer's Patches ave absent entirely. I also found no trace of
them in Papio anubis.
There is nothing peculiar to note about the peritoneum. ‘The
mesenteric vessels run straight to the gut, and do not form tiers
of arterial arcades. Reaching the bowel they bifurcate, and the
halves unite with those of the neighbouring vessels. Several
small lymphatic glands are situated along the vessels at the
point where they reach the gut.
The entire mesentery and great omentum are devoid of fat.
THE SALIVARY GLANDS.
The parotid gland (text-figs. 9 & 17) consists of a large trian-
gular superficial part, with small closely-packed Iebules, and a
deeper part, with larger lobules. In the latter there are many
large vessels and nerves.
The sublingual glands ave the size of hazel nuts. They are
closely applied to the inner border of the mandible, and the
branches of the lingual nerves disappear under cover of them.
The submavillary glands are large, circular, and flat, and have
a large blood supply from the external carotid arteries. They
lie immediately posterior to the fused anterior bellies of the
digastric muscles. ‘hey are, however, relatively smaller in pro-
portion to the size of the animal than those in many other
Cercopithecide.
THE PANCREAS.
One of the most notable features in the anatomy of the Drill
is the small size of the pancreas. It is almost black in colour,
and its component lobules are just visible to the naked eye. It
has a thin tail and an expanded body, but there are no lateral or
accessory parts. The total length is three and a quarter inches,
and the thickness is two millimetres. The head is oneinch wide,
and the tail is a quarter of an inch across. The duct unites with
the common bile-duct.
Many arteries run into it from the splenic artery.
Tue Liver (text-fig. 15 A).
The liver is square in shape, as in Papio anubis (2), but it is
i 7 ] 5 } yy . A =| 1 1 1
thin throughout. The gall-bladder is pyriform, contained in a
well-marked fossa, and continued into a long, slender bile-duct
which unites with other hepatic ducts to form a long common
bile-duct. The latter unites with the pancreatic duct, but there
is no bile papilla in the duodenum. ‘The lobes of the liver differ
ANATOMY OF THE DRILL. 439
from those in the Baboons. ‘he left lateral lobe is smaller; the
left central lobe is not so wide, and its free margin is notched.
The right central lobe is intimately united to the left central
lobe, and its free edge is more rounded. The right lateral lobe,
caudate and Spigelian lobes are larger relatively to the others
than in the Baboons.
Text-figure 15.
The liver (A.) and trigone of the urinary bladder (B,
umbilical notch; left lateral (.L.), left central (
right central (R.C.), caudate (C.), and Spigelian (S.)
). G.B: gall-bladder; U:
L.C.), right lateral (R.L.),
]
obes.
The lateral fissures are deep, the fissure of the ductus venosus
is marked, and the umbilical fissure is bridged over. The vena
caval fissure is not bridged over, and the vein is superficial.
THe DuctriEss GLANDS.
The spleen (text-fig. 17 B) is flat and triangular, with rounded
angles. The hilum is not linear, as in many Mammalia, but
occupies an oval area into which tortuous vessels and sympathetic
nerves enter.
The suprarenal capsules are smali, thin, flat, and helmet-
shaped, and lie in the usual positions. Hach receives two small
arteries from the abdominal aorta and one’ from the renal artery.
The sympathetic plexuses, especially those to the left gland, are
very rich and closely related to other abdominal plexuses.
The oval lateral thyroid lobes are unconnected, and lie against
440) DR. C. lh. SONNTAG ON THE
the sides of the posterior part of the larynx and first four
tracheal rings. ‘They receive an artery from the common carotid,
but none from the subclavian as in Man.
The thymus was atrophied.
THe UroGenrIraL OrGANS (text-figs. 15 B & 16).
The kidneys are small, measuring only 21:1 X-5 inches. Each
has two papille, and neither las any fat in its pelvis. The left
one is fixed, but the right one, supported by its vessels and
ureter, is freely movable among the intestines.
The bladder is capacious. On the inner surface is a thickened
T-shaped band containing slit-like ureteric and circular urethral
orifices (text-fig. 15 B).
The testes have large epididymes, and the tunica vaginalis
is well-marked. When the parts are relaxed it is seen that
Text-figure 16.
PROSTATE
\ HF. *
et Ke
7
PU.
The generative organs. A: general view; B: bladder; P.U: prostatic urethra.
B: magnified view of the ducts. Other letters in text.
a considerable length of thick spermatic cord extends from
the epididymis to the external abdominal ring, which is
triangular.
The vasa-deferentia (V.D) become confluent, and unite with
the ducts of the vesicule seminales (D.V.S) to open on a single
large papilla in the dorsal wall of the prostatic urethra. No
definite vesicula prostatica was seen,
ANATOMY OF THE DRILL. 44]
The prostate (P.G) consists of a firm pyramidal part enclosed
in a capsule, and a triangular part which is softer and lobulated
(bse,
The vesicule seminales (V.S)are two immense masses of lobules
bound together by connective tissue and ductules. Many nerves
run into them. Their ducts are long, and fuse with the vasa
deferentia.
The corpora cavernosa penis have large expanded bases.
THE ORGANS OF CIRCULATION.
The heart (text-fig. 17 A) is 3°5 inches long, 2:5 inches wide,
and 2 inches thick. Its apex is less pointed than in some
Primates, and is composed entirely of the thin-walled right
ventricle. The thick-walled left ventricle falls short of the apex
by half an inch. The right auricular appendix is pointed, and
separated from the aorta by a distinct interval. It is slightly
Text-figure 17.
The heart and aorta (A.), and spleen (B.). .V. and R.V: left and
right ventricles. Other letters in text.
notched as in some Marsupials. There are only a few small
columne carnese arranged along the inner sides of its walls. The
right auricle has few columne. Between the orifice of the vena
cava inferior, which is not guarded by a valve, and the tricuspid
valve is a wide coronary opening. The right ventricle is capacious
and five papillary bundles give the chords tendinee to the
4492 DR. C. F. SONNTAG ON THE
tricuspid valve. Several small muscle bundles connect the
ventricular walls. To the left of the valvular muscles les the
wide, capacious antrum of the pulmonary artery which has no
columns carnee. The left auricular appendix has subdivided
edges, and no columnee carne. . The left ventricle is thick-walled,
and its small cavity is greatly subdivided close to its apex by
innumerable small muscular bundles.
The aortic arch gives off the innominate (I.A) and left sub-
clavian (L.8.A) arteries and upper three left intercostals, and the
former, after giving off the left common carotid artery (L.C.A),
divides into the right subclavian (R.S.A) and right common
carotid (R.C.A) arteries. This arrangement resembles that in
many Primates, Carnivora, and Marsupialia.
The common carotid arteries (text-fig. 18) divide at the level of
the upper border of the thyroid cartilage into external carotid
Text-figure 18.
The vessels of the head and neck. O.H: omo-hyoid muscle
Other letters in text.
(E.C.A), internal carotid (I.C.A), occipital (O.A) and superior
thyroid (S8.T.A) arteries, and a sympathetie nerve plexus accom-
panies the branches. Of these, only the external carotid requires
special mention.
The external carotid artery (H.C.A) gives off a trunk which
divides into lingual (L.A) and external maxillary (E.M.A)
arteries, and is continued as a trunk which divides into super-
ficial temporal (8.T.A) and internal maxillary (I.M.A) arteries.
The external maxillary is distributed to the lower lip, parotid,
and masseteric regions, xnd the superficial temporal replaces the
part which, in most mammals, extends to the inner canthus of
the eye. The lingual artery gives off a well-marked branch to
the frenal lamella (text-fig. 19, L.B).
The subclavian arteries have the usual course and relations
and it is noteworthy that the Annulus of Vieussens surrounds
ANATOMY OF THE DRILL. 443
them and their prominent vertebral branches. In most mammals
the Annulus, when present, surrounds only the subclavians.
Intercostal arteries:—The upper two spaces on each side are
supplied by the superior intercostal branch of the subclavian
artery, and the internal mammary artery. The third, fourth,
fitth, and sixth left spaces are supplied by the intercostal arteries
from the aortic arch and the internal mammary artery. The
lower six left spaces are supplied by intercostal branches of the
descending aorta and the musculo-phrenic arteries. The right
spaces from the third to the twelfth are suppled by intercostal
branches of the descending aorta, internal mammary and musculo-
phrenic arteries. The internal mammary artery gives off a large
branch along the anterior rib, and a thin one along the posterior
rib bounding each space supplied by it.
The aortic intercostal arteries are arranged as in those
Primates possessing thirteen pairs of ribs.
The abdominal aorta gives off branches in the following
order :—Inferior phrenics, celiac axis, splenics, superior mesen-
terics, renals, spermatics, inferior mesenteric. At intervals it
gives off four pairs of lumbar arteries. The first left lumbar artery
passes postero-laterally as an ilio-lumbar artery. The aorta ends
by dividing into two common iliac arteries, and does not give off
a middle sacral (caudal) artery. Lateral sacral branches of the
hypogastries form the caudal artery which passes through chevron
bones.
The Veins. .
The inferior labial veins (text-fig. 18, 1.P.V) unite to form
a long superficial trunk which opens into the superficial temporal
vein (S.T.V); and the latter drains the frontal, temporal and
orbital regions, and the upper lip. ‘The inferior labial vein com-
municates with the mylo-hyoid veins which drain the muscles
and the tongue. The mylo-hyoid trunk (M.H.V) divides into
two. One part enters the internal jugular vein (1.J.V), and the
other unites with a vessel formed by the internal maxillary
(I.M.V) and deep facial (D.F.V) veins. And the resulting
vessel opens into the internal jugular vein. ‘The arrangements
and relations of these veins are shown in text-figs.
The eaternal jugular vein (K.J.V) is formed on the surface of
the sterno-mastoid muscle by the union of superficial temporal
and inferior labial veins. It has the usual course through
the neck, and unites with the subclavian vein to form the in-
nominate vein. The internal jugular vein (1.J.V) drains the
thyroid gland, and opens into the innominate vein.
The left innominate vein crosses the anterior part of the thorax
obliquely and unites with the more vertical right innominate
vein to form the superior vena cava. The latter receives the
vena azygos major in the usual way and ends in the anterior
part of the right auricular appendix.
The intrathoracic part of the inferior vena cava is short.
444 DR. C. F. SONNTAG ON THE
Tat ORGANS OF RESPIRATION.
Laryne.—The epiglottis is quadrangular, with rounded angles,
and is attached to the tongue by a prominent glosso- epiglottic
fold. At its base, just above the false cords, a transverse slit
leads into a cavity formed by the excavated hyoid bone. From
the right side of the cavity a laryngeal sac protrudes. The false
cords are thick, the true ones are thin, and the ventricles are
deep. ‘The arytenoids are prominent. The upper border of the
thyroid cartilage has two small elevations for articulation with
the hyoid,
The trachea has twenty-four rings which are all incomplete
behind.
Lungs.—The left lung has two lobes, and the right one has
four, of which one is the azygos appendage.
Tue Nervous System.
The Glosso-pharyngeal Nerve (text fig. 19, 1x) has the usual
relations to other nerves at the base a the skull, and it com-
municates with the vagus, hypoglossal, and sympathetic. It
winds round the stylo- hyoid ligament, runs antero-mesially, and
breaks up into lingual nerves ‘which disappear under the hyo-
glossus. It sends branches to the pharyngeal and carotid
plexuses (P.P and C.A.P), the tonsil, and soft palate.
Text-figure 19.
The nerves in the anterior part of the neck. L.B.T: lingual branch of the
trigeminal; L.A: lingual artery. Other letters in text.
The Vagus Nerve (text-figs. 19-21) is separate from the
sympathetic in the neck. And its course in the neck and thorax
is similar to that already described in my papers on the Mar-
supialia. The ganglion nodosum (text-fig. 19, G, N) is well
marked on each mae and the following bemmanes are given off ;—
ANATOMY OF THE DRILL. 445
1. Communicating to the glosso-pharyngeal, accessory, hypo-
glossal, and sympathetic.
2. Pharyngeal nerve (a) which breaks up into the pharyngeal
plexus in which it meets branches of the glosso-pharyngeal and
sympathetic.
3. Superior laryngeal nerve (b) which gives off one external
branch and a brushwork of internal fibres, and the latter pierce
the thyro-hyoid membrane. No external branch connects it to
the recurrent nerve, and no depressor nerve arises from it.
Communications unite it to the ganglion nodosum and carotid
plexus.
No cardiac nerves are given off in the neck.
Text-figure 20.
Lower cervical and upper thoracic parts of the vagus and sympathetic. D.C:P. and
S.C.P: deep and superficial cardiac plexuses; S.A. and V.A: subclavian and
vertebral arteries. Other letters in text.
4. The right recurrent nerve (text-fig. 20, d) has the usual
origin, course, and relations. It communicates with the sym-
pathetic (8), but no branch connects it to the left recurrent
nerve (e). The latter has the usual course, but it does not com-
municate with the cardiac branches of the left vagus. Cae
5. Cardiac Nerves (f):—The left vagus gives off two large
branches to the superficial cardiac plexus, and these end in small
ganglia which also receive sympathetic filaments. The right
Proc. Zoot. Soc.—1922, No. XXX, 30
446 DR. C. F. SONNTAG ON THE
vagus gives off two thinner branches which break up into several
twigs to a deep cardiac plexus.
6. Pulmonary Nerves (text-figs. 20 & 21, g and g'):—-The left
vagus gives off two anterior pulmonary nerves before it passes to
the dorsal surface of the root of the left lung (g'). On the dorsal
surface of the pulmonary root it gives off three posterior
pulmonary nerves (g). The right vagus gives off one anterior
pulmonary nerve under cover of the vena azygos major, where it
curls over the root of the right lung to join the superior vena
cava. Behind the pulmonary root it divides into two branches.
One of these (/) is distributed to the posterior pulmonary plexus,
and then rejoins the other division.
7. Esophageal nerves (text-fig. 21, O.N):—The left vagus gives
off several cesophageal branches which form a plexus gule. The
right vagus gives an ascending cesophageal nerve (O.N'), and some
small branches which anastomose with those of the left nerve.
The left vagus runs through the ventral part of the cesophageal
opening in the diaphragm, and the right vagus runs through the
dorsal part. A complicated series of anastomoses between their
branches takes place in the posterior part of the thorax (text-
fig. 20).
The left vagus gives many branches to the cesophagus and
ventral surface of the stomach, and twigs can be traced into the
splenic and superior mesenteric plexuses. Communications can
also be traced to the left splanchnic nerves (text-fig. 22).
The right vagus supplies the dorsal surface of the stomach, and
branches run to the solar plexus (text-fig. 21).
No direct communications run between the vagus and phrenic
nerves.
The Spinal Accessory Nerve (text-fig. 19, x1) has the usual
relations to other nerves at its emergence from the foramen
lacerum posticum. It lies on the dorsal aspect of the sterno-
mastoid and cleido-mastoid, and it gains the deep surface of the
trapezius. It passes postero-laterally and runs over the dorsum
of the scapula between the ventral border and root of the spine.
It can be traced into the most posterior fibres of the trapezius.
It forms a rich plexus with the cervical nerves, and com-
munications run between it and the ninth, tenth, and twelfth
nerves.
The Hypoglossal Nerve (text-fig. 19, x11) communicates with the
other nerves at the base of the skull and describes a wide loop.
Reaching the tongue by passing on the surface of hyoglossus it
divides into two branches. One is more superficial, and gives off
numerous fine muscular twigs. ‘The other is deeper and thicker,
and can be traced almost to the tip of the tongue. Before it
divides, the hypoglossal gives off the descendens (D.H) nerve and
communicates with the lingual nerve. The former enters as
usual into the ansa hypoglossi.
In text-fig. 19 the three distinct nerve supplies to the tongue
are shown.
ANATOMY OF THE DRILL. 447
Tae Symparueric Nervous Sysrem.
Both superior cervical ganglia (text-fig. 19, 8.C.G) are present
and communicate with the glosso-pharyngeal, vagus, hypoglossal
« loa aT ryye ” ays a ~ ty, x 1 {Pp
and superior laryngeal nerves (5), and the cervical plexus (C.C.P).
Branches run to the pharyngeal (P.P) and carotid (C.A.P)
plexuses.
Text-figure 21.
RV. LV.
g. 3: 1
So.P GB.
B.
)
SMP SRP
Sp.R
uf
G.
Posterior thoracic and abdominal parts of the vagus nerves. G.B: gastric branches ;
S.P. and So.P: branches to solar plexus; Sp.P: to splenic plexus; S.M.P: to
superior mesenteric plexus ; S.R.P: to left suprarenal plexus.
The middle cervical ganglion (text-fig. 20, M.C.G) is only
present on the right side, and the Annulus of Vieussens (A.W) ais
interposed between the sympathetics at the root of the neck and
the inferior cervical ganglia (I.C.G). The Annulus on each side
encircles the subclavian (S.A) and vertebral (V.A) arteries.
No fusion occurs between the vagus and sympathetic in the
neck, and the communications between them take place at the
extremities of the cervical course. The left sympathetic gives off
cardiac nerves (C.B.S) at the root of the neck, and the right one
gives off a cardiac cord from its middle ganglion (U.B.8’), tracheal
nerves (T.N), and communicating to the right recurrent laryngeal
nerve (C.R.L.N).
The inferior cervical ganglia give off ascending branches which
accompany the vertebral arteries (V.A.N) and transverse twigs
which run to the brachial plexus (C.B.P). The main sympathetic
cords emerge from their posterior parts (‘T.C.S).
30*
448 DR. GC. F. SONNTAG ON THE
The thoracic cords have fewer ganglia than the numbers of
intercostal nerves, and the rami communicantes are of consider-
able length. In text-fig. 20 B are shown the slender white ramus
communicans (w.7.c) emerging from the vertebral foramen, and
the thicker grey ramus g. T, ¢) twining round the psoas muscle
to join the last dorsal nerve (L.D.N).
The left abdominal sympathetic has five ganglia (text-fig. 22,
1-v). It gives off four splanchnic nerves (S. N) which unite to
form a loop whence several plexuses radiate. And the loop
receives fibres from the left vagus (L.V), Farther back three
branches run together to form a long narrow cord, and branches
run thence to form the inferior mesenteric plexus (I.M.P).
Branches from the right sympathetic join the latter.
The right sy mpathetic (R.S8) divides into the right abdominal
cord which has fou ganglia (1--1v),and the splanchnic which gives
off the phrenic plexus (P.P) and ends in the right renal plexus
(&.R.P).
The main sympathetic cords diminish rapidly in size when
they enter the pelvis. And each has one ganglion between the
pelvic inlet and the point where the two run “together at their
posterior extremities. No communicating vami were seen
running to the sacral plexuses.
Cervical Branches of the Sympathetic (text-figs. 19 & 20).
The anterior ends of the sympathetics give communicating
branches to the glosso-pharyngeal, vagus, hypoglossal, and favors. |
laryngeal nerves (b). Grey rami cenit una eaves run to the
upper cervical nerves (C.C. P). Branches go to the pharyngeal
(P.P) and carotid (C.A.P) plexuses.
The right sympathetic gives communications to the right
recurrent nerve (d), and the tracheal plexus (T.P).
Both sympathetics give off cardiac nerves (C.B.S) in the neck,
but none arise in the thorax. These meet with branches of the
vagus nerves in the superficial and deep cardiac plexuses.
The Superficial Cardiac Plenus (text-fig. 20, 8.C.P) is composed
of :—
1. 'I'wo thoracic cardiac branches of the left vagus.
2. Three fine cervical cardiac branches of the left sympathetic.
The Deep Cardiac Plexus (D.C.P) contains :—
1, A cervical cardiac branch of the left sympathetic.
2. A large cervical cardiac branch of the right sympathetic.
3. Two thoracic cardiac branches of the right vagus.
The Thoracie Branches of the Sympathetic (text-figs. 20 & 22)
consist of :—
1, Pulmonary nerves entering the anterior and posterior
pulmonary plexuses (P.B.S). They do not arise as separate
nerves, but are given off from the cardiac nerves.
2. Splanchnie nerves (text- fig. 22) which end in the solar plexus
on the left side, and the right renal plexus (R.R.P) on the right.
ANATOMY OF THE DRILL. 449
There is only one splanchnic nerve on the right side, but four
form a loop on the left. ‘The connections which they establish
through the solar plexus, and by direct branches to the organs, are
numerous and complicated, but careful dissection enables one to
see the following offshoots :—
a. Superior mesenteric plexus (8.M.P).
b. Splenic plexus (8.P).
c. Left renal and suprarenal plexuses (L.R.P).
d. Right renal plexus (R.R.P).
e. Aortic plexus (A.P).
Jj. Inferior mesenteric plexus (I.M.P).
Text-figure 22.
The thoracic and abdominal sympathetic. U.S. and R.S: left and right
sympathetic cords; L.V: left vagus. Other letters in text.
The Inferior Mesenteric Plewus sends branches to the large
intestine, and a strong cord runs down into the pelvis. Gaining
the dorsal aspect of the vesicule seminales it forms a crescentic
expansion, whence branches run to the left sacral plexus and
pelvic viscera.
The Brachial Plexus (text-fig. 23) is formed by the fourth to
eighth cervical nerves, and branches of communication run to the
cervical plexus and first dorsal nerve (C.C.P and C.D.I). In
Macacus rhesus and Homo the fourth nerve only communicates,
450 DR. C. F. SONNTAG ON THE
and the plexus is formed by the fifth cervical to first dorsal
inclusive.
The fourth nerve communicates with the fifth and becomes the
suprascapular nerve, and the phrenic nerve arises from the
junction of the fourth and communicating branches. Filaments
may run from the communicating twigs from the cervical plexus
to suprascapular and phrenic nerves. In Macacus rhesus,
according to Brooks (1), the large fourth cervical gives a branch
which joins the small fifth cervical just as the latter gives off the
phrenic nerve. In Homo the phrenic nerve receives definite fibres
from the third, fourth, and fifth cervical nerves, and the supra-
scapular nerve arises from the fifth and sixth cervicals.
The fifth, sixth, and seventh nerves quickly divide into anterior
and posterior divisions, but the eighth does not divide. So there
Text-figure 23.
UPPER==- ~~
SUBSCAPULAR -—
MID SUBSCAPULAR. ~
B.A.B. ~ we
ur. BB. C.B. Se Z
"LONG. SUBSC!
MEDIAN.
ULNAR. Bg
JONG THORACIC.
The brachial plexus. The three small fibres running up from C.1v are known in
the text as C.C.P. The two small twigs attached to the median nerve are the
anterior thoracics. C: communicating nerves; C.B: coraco-brachialis; B.B:
biceps; B.A.B: brachialis anticus; R.N: radial nerve; P.I: posterior inter-
osseous nerve.
is no trace of trunks as seen in Homo, and there is no upper trunk
as in Macacus rhesus.
The anterior division of the fifth nerve gives off the upper and
middle subscapular nerves, and unites with the anterior division
of the sixth to form the musculo-cutaneous and middle subscapular
nerves. In Macacus rhesus two small subscapulars come from the
posterior division of the upper trunk (C.v and C.yz1). It is
possible that the musculo-cutaneous is the only representative of
the outer cord of the plexus in Homo.
The posterior branches of the fifth and sixth nerves can be
traced into the musculo-spiral and circumflex nerves, and the
musculo-spiral nerve is joined by the anterior division of the
seventh. In Macacus rhesus the posterior of the upper trunk
unites with a branch of the seventh cervical nerve to form a
ANATOMY OF THE DRILL. 451
trunk which divides into middle subscapular and circumflex. The
musculo-spiral nerve is formed by the posterior division of the
conjoined fifth and sixth, the posterior division of the seventh,
and the posterior division of the conjoined eighth cervical and first
dorsal nerves as in Homo.
The posterior division of the seventh and the whole of the
eighth cervical nerves fuse to form a trunk which divides into
median and ulnar nerves. The median has externally only a
single head of origin, so it dit.ers from those of Macacus rhesus
and Homo, in which two were seen.
Text-figure 24.
Sy.
The lumbo-sacral plexus. S.G. and I.G: superior and inferior gluteal nerves;
Q.F : to quadratus femoris ; G.S: great sciatic; Sy : sympathetic.
The long subscapular nerve arises from the back of the musculo-
spiral nerve as in some examples of Macacus rhesus (1). And
Brooks points out that, although its origin may vary considerably,
it is always related in some way to the musculo-spiral.
The nerves to the pectoral muscles arise from the median nerve,
but they may simply run through the latter for a considerable
distance. ;
In my opinion the musculo- cutaneous nerve represents the
outer cord in the human plexus, the posterior cord is represented
by the separated circumflex and musculo-spiral nerves, and the
inner cord is the band which divides into median and ulnar
nerves. It is, however, difficult to explain how the median and
452, DR. C. F. SONNTAG ON THE
external anterior thoracic nerves have no connection with the
outer cord.
The musculo-cutaneous nerve gives off branches to the coraco-
brachialis, biceps, and brachialis anticus, and the latter is of
considerable length.
The circumflex nerve gives off a large nerve to the teres minor
(T.M), but no ganglion is present on it as in Man. ‘The
remainder runs round the humerus, and divides into two parts
which end in the deltoid (D.B).
The musculo-spiral nerve gives off wpper and lower branches to
the triceps (U.T.B and L.T.B), and a long branch to the dorso-
epitrochlearis (D.E.M). It turns sharply round the humerus, and
divides into radial and posterior interosseous nerves at a point an
inch proximal to the elbow joint.
The Verve of Bell (long thoracic) arises from the sixth nerve,
and receives a fine communicating branch from the fifth.
The Lumbo-sacral Pleaus (text- -fig. 24) is a long series of
anastomoses between spinal nerves, and the branches are long and
comparatively unbrauched till they leave the abdomen and pelvis.
The arrangements differ from those in Man, for there are more
spinal nerves, and the muscles differ.
The plexus communicates above with the last dorsal nerve.
The first lumbar nerve communicates with the last dorsal and
second lumbar nerves, and becomes the ilio-hypogastric nerve.
The second lumbar nerve communicates with the first and third
and becomes the inguinal. The second, third, and fourth enter
into the lateral cutaneous nerve of the thigh. The third and
fourth form the obturator nerve. The third, fourth, and fifth
form the anterior crural. The fifth and sixth unite to form the
lumbo-sacral cord which passes to the sacral plexus. The sacral
and coccygeal nerves give mesial and lateral branches. The
mesial ones include muscular nerves to levator ani, coccygeus, and
pyriformis. The lateral ones include both sciatics, both gluteals,
and the nerve to quadratus femoris.
SuMMARY AND CoNncLUSIONS.
Lhe following facts are described for the first time in the
Primates except Man :—
1. The delto-pectoral triangle and contents.
2. The costo-coracoid ligament replaced by a costo-humeral.
3. The double nerve supply of the Axelbogen.
4. Confluence of both ejaculatory ducts.
5. Absence of the sinus pocularis.
6. The right ventricle forming the cardiac apex.
7. The Annulus of Vieussens encircling the subclavian and
vertebral arteries.
8. The absence of any semilunar ganglia in the solar plexus.
9. Analysis of the lumbar plexus.
10. Absence of the mid-sacral artery.
ANATOMY OF THE DRILL. 453
The Drill differs from the other Old World Monkeys in the
following respects :—
1. The clavicular origin of the pectoralis major is reduced to
the extreme sternal end, whereas the deltoid has a longer clavi-
cular origin than in other Primates.
2. The rhomboids do not reach the occiput.
3. The subclavius is more powerful.
4, The latissimus dorsi does not arise from any ribs.
5: The long head of the triceps has a very extensive origin
from the scapula.
6. The teres major, latissimus dorsi, and dorso-epitrochlearis
are fused.
7. The pancreas is extremely small.
8. The vesicule seminales consist of lobules.
9, The arrangement of the great veins of the face and neck.
10. The arrangement of the branches of the brachial and
lumbar nerves.
BIBLIOGRAPHY.
1. Brooxs, W. T.—Journal of Anatomy, 1883, p. 329.
2. Firowrr, W. H.— Medical Times and Gazette, 1872, p. 295.
3. Vrouix.—Art. ‘“ Quadrumana” in Todd’s Cyclopedia of
Anatomy and Physiology.
4, Parsons, F. G.—-Journal of Anatomy, 1898, p. 436.
‘ 8 tre salar “a eos re
: 4 a) “pal, jets ree ce wae 4
Lay, &
7 et Break
NA ba ah emg ae iy bUaarek. 6 ite antag cha Wit
Mili 0) ae a
rele? suaites raid tes aus caine
yg 0 Baa ites ro Ridin br or ot ee . a i
dig 444 ht by .
: ee Heal aces 4 di re Depiey Pi +i ar Sw
i
Wien cons igo ena ‘bya abil Ayr tifa fen NB 1 ae
ees he B 4 ‘erty tis ee yt bi |
. iio wena Gesethass Shaicite {Ui UReNy? waaegt i ait
S09 Aa TB be drivin A ta 98 a mili iit ae
Yew ee 'Fixtos ws bistelo Cry bio aees Mua
cette hyp habs “eure pei ts
Elis, r
Hye hit ward ‘a NIN: tig Loe
ly ake je? KC gaettie Gilles qr W. : dit ‘koe
* etians Pye Dae Aarti be ” ne Vb. sa ptt oe
aa Byatt ae ih aie oiecer’ fii ‘ we b vere
) ORE te ne “Oe sepa wae nny COINS
Sour BU aia PL 4 Penns eee +o huivi Ase
Ringe i Athal !
i
Ane
MESOPTERYGOID IN CERTAIN REPTILIAN SKULLS, A455
23. On the Persistence of the Mesopterygoid in certain
Reptilian Skuils. By R. Broom, F.R.S., C.M.Z.S.
[Received April 4, 1922: Read April 25, 1922.)
(Text-figures 5-7.)
In 1878, Kitchen Parker described the structure and develop-
ment of the skullin the Common Lizard, and figured a section
of a fairly advanced Zootoca showing the region where the ptery-
goid articulates with the basisphenoid. A fairly large intercalary
cartilage is shown to be present, but Parker apparently did not
consider it to be of any important morphological value.
In 1901, Howes and Swinnerton described the development
of the skull of Sphenodon punctatus (Trans. Zool. Soc. xvi. p. 1.
1901), and showed that there is also a cartilage similar to that
noticed by Parker in Zootoca, interposed between the pterygoid
and the pterygoid process of the basisphenoid. They compare it
to the meniscus found in the articulation of the jaws of most
mammals, and give to it the name meniscus pterygoideus.
For many years I have been inclined to think that this little
cartilage would yet turn out to be of considerable morphological
importance, but until recently I have never been able to get any
clear light on it from either skeletogenesis or paleontology.
Within the last few years, however, two papers have appeared
which seem to show with hardly any doubt that this little
meniscus pterygoideus of Howes and Swinnerton is really a
persistent piscine mesopterygoid.
Before discussing the nature of the element, it will be well to
consider in some detail its structure and relations, as, so far as I
am aware, it has never hitherto been at all fully described or
figured.
Among living animals it is only known to occur in Lizards and
Sphenodon, and as it can be seen even better in the primitive
Lacertilian Agama than in Sphenodon, I will give a few figures
of it as it occurs in a larval Agama hispida.
Text-fig. 5 represents a graphical reconstruction of the middle
part of the base of the skull of the embryo Agama with a skull
length of 7mm.* The membrane bones of the skull are all
ossified at this stage, but in the cartilage elements, such as the
quadrate, ossification has just started as an exostosis. As is seen
in the figure given, the pterygoid is well ossified, and extends far
forward, though it does not meet the prevomer as in Sphenodon.
The basicranial cartilages are still unossified, and from the basi-
sphenoid region there passes outwards a well-developed basiptery-
goid process to support the pterygoid ; but it will be seen that it
only supports the pterygoid indirectly, as a large independent
* T am indebted to Prof. J. P. Hill for having kindly sectioned the Agama skull
for me.
456 MR. R. BROOM ON THE PERSISTENCE OF THE
cartilage lies between the basipterygoid process and the pterygoid.
This is the meniscus pterygoideus of Howes and Swinnerton, and
the element whose affinities we wish to examine. It is rather
longer than the face of the basipterygoid process, and, as will be
seen in the cross-sections given, it is of considerable thickness.
It completely separates the pterygoid from the basisphenoid, so
that no part of the pterygoid is in contact with the basicranial
cartilage. Further, the ‘‘ meniscus” has no direct relations with
the epipterygoid, the pterygoid completely separating the one
from the other. ‘To a considerable extent it is clasped by the
Text-figure 5.
Sy ~ DS aN
C7 t \ : —
eee MANNS
Base of skull of embryo Agama hispida var. aculeata. Head length 7mm. Recon-
structed from sections. On the right side, parts of the vomer (parasphenoid),
basipterygoid process, and of the pterygoid have been removed ; and a full view
is given of the mesopterygoid.
References to lettering in figures :—B.Pt.P. Basipterygoid process of Basisphenoid ;
E.P. Ectopterygoid (in B & C of fig. 6 the element is not yet ossified where
sectioned); #.P¢. Epipterygoid; 22.Pt. Mesopterygoid; Mr. Maxilla ;
Pa. Palatine; Pma. Premaxilla; Pt. Pterygoid; P.Vo. Prevomer; @. Quad-
vate; Tv. Trabecula; Vo=PaS. Vomer=Parasphenoid.
pterygoid, and the movement at the joint is between the
““meniscus” and the basipterygoid process and not between
the pterygoid and the ‘‘ meniscus.” In fact, it is pretty manifest
from the examination of this embryo alone that the element is
really a pterygoid element.
The reconstruction of the base of the skull in this Agama
embryo shows a very interesting condition of the median basal
bony element which the large majority of morphologists call the
parasphenoid, but which I regard as the true homologue of the
mammalian vomer. Besides nearly covering over the pituitary
MESOPTERYGOID IN CERTAIN REPTILIAN SKULLS. 457
opening, it forms a large support to the basisphenoid cartilage,
especially in the neighbourhood of the basipterygoid process.
In front this median vomer or ‘“ parasphenoid” extends forward
asa support to the median septum almost exactly as does the
mammalian vomer.
The fossil Tetrapod skull that has given us the clue to the real
affinity of the “meniscus” is that of “ Seymouria baylorensis”
Broili, recently described by Watson. ‘* Seymowria” is one of the
most interesting primitive fossil types ever discovered; and
Text-figure 6.
Cc
B
Transverse sectious through the mesopterygoid and related elements in the skull
of the embryo Agama shown in Fig. 5.
A. Through the posterior part of the element.
L. A section near the middle.
C. A section near the anterior end.
Broili, Williston, and Watson have vevealed to us nearly every
detail of its structure. There are, however, two points on which
one would like to remark. Williston, v. Huene, and Watson all
agree that Seymouria baylorensis Broili is the same animal as
Conodectes favosus Cope. Cope’s type skull is in the American
Museum, and his description, though by no means such as would
enable anyone to identify readily a similar species, is fuller than
that of many types, and I cannot agree with Watson that Cope’s
458 MR. R. BROOM ON THE PERSISTENCE OF THE
name should be treated as a nomen nudum. Conodectes has eight
years’ priovity over Seymouria, and if those who have examined
both types ave satisfied that they belong to the same species,
IT think we will have to use Cope’s name Concdectes favosus in
preference to that of Brioli.
The second point is a much more debatable one. Watson
deseribes “ Seymouria” as “the most primitive known reptile,”
and everyone else who has written about the animal has also
been of the opinion that it is a reptile; and until Watson’s paper
appeared, I also accepted this opinion, having never seen any of
the actual specimens. I do not in the present paper wish to enter
at length into the discussion, but may state that the study of
Watson’s paper leads me rather to the conclusion that Conodectes
is a highly evolved and specialized Embolomerous Amphibian,
with a very few reptile-like characters which have been acquired
by a parallel development. For the present, however, it matters
little whether Conodectes is an Amphibian-like Reptile or a Reptile-
like Amphibian. It certainly lets us know more of the primitive
Tetrapod structure than almost any other known type, and for-
tunately the palate is well known, and has been fully described
by Watson.
There is a large pterygoid which meets its neighbour in front,
and it is supported indirectly by a basipterygoid process. Between
the process and the pterygoid is a distinct bony element, con-
cerning which Watson says :—‘‘ The short basipterygoid processes
of Seymouria supporting the pterygoid through the intervention
of a special bone are unique, nothing similar occurring in any
other known adult reptile or amphibian. The shape and position
of the pterygoid render it certain that these special bones cannot
be the epipterygoids, which in Dimetrodon ave known to articu-
late with the basisphenoid. Swinnerton and Howes showed that
in the development of the skull of Sphenodon special articular
cartilages are developed between the basipterygoid processes
and the pterygoid, and it is not impossible that these are the
representatives of the articular bones in Seymourica.
“On the other hand, Gaupp has shown that in Lacerta the
basipterygoid processes contain independent centres of ossification,
and it is feasible and attractive to regard the Seymouria bones as
permanently separate autogenous basipterygoid processes.”
With regard to this second suggestion, it may be mentioned
that the basisphenoid in lizards and many other animals ossifies
for the most part from the parasphenoid, and, so tar as I am
aware, no one has confirmed Gaupp’s observation. Parker
examined an embryo Zootoca (his 6th stage) which one would
fancy ought to show the basipterygoid distinct if it is ossified
from a separate centre; but there is no evidence of this. Howes
and Swinnerton figure a stage in the development of the skull of
Sphenodon (stage 8) where the basisphenoid is evidently ossified
from a pair of centres each of which embraces the basipterygoid
MESOPTERYGOID IN CERTAIN REPTILIAN SKULLS. 459
If Gaupp’s observation be confirmed, the ossifications
must be regarded, I think, as the paired centres of the true
basisphenoid. In fact, if one looks at the basisphenoid of an
embryo lizard with its huge median fontanelle (c/, text-fig. 5),
it is difficult to see how it could be ossified except by lateral
ossifications if it is not completely ossified by invasion from the
parasphenoid.
process.
The other suggestion made by Watson, that the meniscus
pterygoideus found in Sphenodon by Howes and Swinnerton and
previously figured by Parker in the lizard is the homologue of
the little bones in ‘‘ Seymouria,’ seems to me manifestly the
correct one.
The bony element which forms the cranial base is regarded by
Text-figure 7.
Vil ig Px:
eee eth Px
Oo ee To
Y aa oS
Qi Ke Z yY ‘
w\ZA % Zp
Pa. Z Z ae Zi, Mx.
Z g 7 ! |RVolPVo}',
g 4 4 Mx. ‘ ey
AY }
AY 4
BY 4
AY ,
j 4
pr.("% PA pe \ Y
ELM
Vo. y
=PzS, 4
, Pt.
yy YH ‘ : a"
Wa Yj > VAI. <j
B
A. The palate of the Rhipidistian Fish, Husthenopteron foordi. After Bryant.
B. The palate of the primitive Tetvapod Conodectes favosus Cope=Seymouria
baylorensis Broili. After Watson. In the opinion of Watson, Williston, and
‘others this is a very primitive reptile: the writer favours the view that it is a
highly developed Embolomerous Amphibian.
Watson as made up of parasphenoid in front and basisphenoid
behind. Though I have not seen the specimens, I incline to
think it probable that the whole will prove to be parasphenoid
or vomer.
But, in any case, there seems to me no doubt that the
little intercalated element is the homologue of the little one
we have in the lizard and Sphenodon.
When we go further down in the scale we have hitherto had
difficulties, as no good palate was known of any of the Rhipidistian
460 MESOPTERYGOID IN CERTAIN REPTILIAN SKULLS.
Crossopterygians. Now, however, this difficulty has been removed
by the description of the palate of Husthenopteron foordi by
Bryant*, and we have the clue to the little element in both
Conodectes and the lizards (text-fig. 7). As will be seen from the
figure, there is a large bone which has on its outer side a palatine,
and an ectopterygoid, and which is manifestly the pterygoid of the
higher forms. Internally it articulates with a long, slender bone
whieh is the mesopterygoid of the bony fishes. This mesoptery-
goid is supported by the parasphenoid or vomer. The relations
of all the bones is very similar to what we have in the higher
forms, and the only striking difference is that in the Tetrapods
the mesopterygoid has become greatly reduced and usually lost.
But when it is retained it is found, as in the Rhipidistian fish,
supporting the pyterygoid and itself articulating with the basi-
pterygoid process of the basisphenoid or the parasphenoid.
There is one other group of fossil reptiles in which it is
possible there is also a distinct mesopterygoid. SollastT, in
describing recently the skull of Jehthyosaurus from serial sections,
showed that there are two epipterygoid-like elements on each
side. The posterior pair are, I think, manifestly the true epi-
pterygoids. Not improbably the anterior pair will prove to be
mesopterygoids, which in the specimen examined by Sollas had
been slightly displaced outwards by crushing.
* “On the structure of Husthenopteron,’ Bull. Buffalo Soc. Nat. Sei. xii.
pp. 1-59 (1919).
+The skull of Ichthyosaurus, studied in serial sections,” Phil. Trans. vol. 208 B
(1916).
A NEW SPECIES OF GSOPHAGOSTOMUM. 461
24. A New Species of Qisophagostomum (sophago-
stomum xeri, sp. n.) from a Rodent (Xerus setosus).
By R. J. Orriepp, M.A., Research Assistant, Helmin-
thological Dept., London School of Tropical Medicine ;
Hon. Parasitologist to the Society.
[Received March 9, 1922: Read April 25, 1929. ]
(Text-figures 1-6.)
In January 1922, £ collected a number of Nematode parasites
from the ceca of two Bristly Ground Squirrels (Xerws setosus)
which had been received from South Africa, a month previously,
by the Zoological Society, Regent’s Park, London. These parasites
comprised 11 males and 17 females. They were irregularly
distributed in the cecal contents.
In the specimens, which were all killed by immersion in hot
70 pex cent. alcohol, the anterior end of the body is bent ventral-
wards in the form of a hook.
The cuticle 1s transversely striated, the distance between the
most anterior stri being slightly over 7 .; this distance increases.
posteriorly until a maximum of 27 is reached at about one-
seventh of the total length of the worm; after this the distance
gradually diminishes, so that in the middle of the body it is 12 p
and at the hinder end 7 p.
A mouth collar (text-figs. 1 & 2) is present; this is broad
and shallow, its depth being less than half its breadth. Immedi-
ately behind it there is a well-marked vesicular swelling about
as long as it is broad, but sometimes slightly longer than broad
(text-fig. 1). Transversely it shows fine cuticular striations. It
is terminated behind at a distance of about 220 from the
anterior end by a cervical constriction of the cuticle ; this con-
striction almost encircles the worm except for a short space on
the dorsal surface.
Two cervical papille are present; these are small, and are
situated behind the middle of the csophagus at about the
anterior margin of the posterior third of the cesophagus.
There are no lateral ale.
The females are slightly longer and thicker than the males,
measuring on an average 15mm. in length, the smallest being
13mm. and the largest 16°5 mm. long. In the males the average
length is 13mm.,, the smallest being 12mm. and the largest
14mm.
The females have their maximum breadth at the junction of
the posterior and middle thirds of the body, and this breadth in a
female 16mm. long is 510. From this point the bedy tapers
Proc. Zoou. Soc.—1922, No. XX XI, Bilt
462 MR. R. J. ORTLEPP ON A
towards the anterior end until it reaches the posterior margin of
the vesicular swelling, where it is then 190, broad. Posteriorly
the narrowing is more gradual until the level of the vaginal
opening, where the breadth is 360 4; from here the narrowing is
very marked, and ends in a pointed tail.
In a male 12°5 mm. long the breadth of the body at the base of
the vesicular swelling is 170. From here the breadth increases
until a maximum of 380, is reached about half-way down the
length of the worm; posteriorly the body tapers until the
anterior margin of the bursa is reached, where the breadth is then
227 pe.
Text-figure lL.
Lateral view of anterior end of body.
©.P. = position of cervical papilla; E.P. = excretory pore; Int. = intestine; M.C. =
mouth collar; N. = nerve ring; Oes. = esophagus; V.S. = vesicular swelling.
The mouth collar surrounds the mouth capsule (text-fig. 2);
this latter is a chitinous structure whose breadth is about three
times its depth; it is somewhat cylindrical in shape, having its
anterior opening slightly smaller than its posterior ; anteriorly it
bears a crown of ten large leaf elements, which pass through the
mouth and whose extremities are slightly recurved over the
anterior part of the mouth collar. An internal leaf-crown is
present, and it consists of about twice as many small elements as
the external leaf-crown. The mouth aperture is surrounded by
the usual six papille, two short and broad laterals, and four
longer submedians which pierce the mouth collar.
NEW SPECIES OF GSOPHAGOSTOMUM. 463
A well-developed wsophageal funnel is present; this is goblet-
shaped and is lined with chitin. The anterior margin of the
funnel has a thickened chitinous rim having an undulating
outline. Because of these undulations the depth of the
Text-figure 2,
i
un
Pen IRE
|
Optical section of head, lateral view.
E L.C. = external leaf-crown; Ex.P. = excretory pore; Fl. = esophageal flange ;
1.L.C = internal leaf-crown; M.Cap. = mouth capsule; Oes.F. = cesophageal
funnel; T. = dorsal tooth.
Text-figure 3.
Frontal view of head, diagrammatic.
C.R.O. = chitinous rim of cesophageal funnel; D.T. = dorsal tooth; E.L.C. =
external leaf-crown; M.C. = mouth collar; M.Cap. = mouth capsule.
mouth capsule, which rests on the rim, is not everywhere the
same. From theanterior border of this rim arise three recurved
tooth-like processes, each about 7 yw long, situated one on the
summit of each of the three undulations; these processes are
situated one dorsal and two latero-ventral, the three, however, being
31*
464 MR. R. J. ORTLEPP ON A
equidistant (text-fig. 3). The chitinous lining of the funnel is
provided with three flange-like chitinous plates, projecting into
the lumen in a’ longitudinal direction ; each flange is situated
about half-way between each pair of tooth- like processes.
The funnel passes into the cesophageal lumen, which, from its
commencement, is triradiate in.transverse section, the sides of
each ray being str ost chitinized. The wsophagus (text-fig. 1)
is a little less than ay of the total length of the worm ; it is slightly
swollen anteriorly round the funnel ; posteriorly it becomes
eradually thickened, and assumes the form of a club, its greatest
diameter being 110 p.
The intestine which follows has a dark colour; it passes
straight backwards to open to the exterior by a short rectum
lined intern: ly with chitin.
Text-figure 4.
Dorso-lateral view of male genital bursa.
D.R. = dorsal ray; E.D.R. = externo-dorsal ray; E.L.R. = externo-lateral ray ;
M.L.R. = medio-lateral ray ; P.L.R. = postero-lateral ray; P.P. = prebursal
papilla; V.R. = ventral rays.
The nerve ring is situated just behind the level of the cervical
groove, its dorsal part being slightly more anterior than its
ventral.
In the cervical groove, on the ventral surface, is situated the
opening of the evcretory duct; this duct passes obliquely back-
wards and inwards, and in so doing passes over the ventral
surface of the nerve ring.
The male genital bursa is ample, forming about =; part of the
length of the worm and being nearly twice as broad as it is deep
(text-fig. 4). It consists of two lateral lobes united by a small
median dorsal lobe. In each lateral lobe are situated six rays. The
two ventral rays are of the same size, run parallel, and are closely
NEW SPECIES OF GSOPHAGOSTOMUM. 465
apposed to each other to the rim of the bursa. The postero-lateral
and medio-lateral rays are also equal in size, and pass to the edge
of the bursa closely parallel to each other. The externo- iateral
ray diverges from the other lateral rays, so that eventually its tip
is about one-third of the distance between the medio-lateral and
latero-ventral rays. From the main trunk of the dorsal ray, about
half-way between its root and its bifurcation, the externo-dorsal
ray takes its origin; it also passes into the lateral bursal lobe,
but terminates a short distance from the margin of the bursa.
The dorsal ray supports the median lobe of the bursa. Its base
Text-figure 5.
a = spicules in situ. B. = bursa; Fl. = spicular flange; G. = gubernaculum.
6 = side view of gubernaculum.
is stout and about 56 u in diameter, but after the origin of the
externo-dorsal ray it is only about half this breadth. Tt is about
230 long and bifureates at its middle, and each branch so
formed bears on its outer margin a short, somewhat elbowed
branch which is slightly thicker than the rest of the main
branch.
The spicules (text-fig. 5a) are long and tubular, and are
protruded some considerable distance in almost all the males
obtained. They are similar and of equal length, measuring
1:29 mm. long, more or less straight, and both end in a sharp point,
466 MR. R. J. ORTLEPP ON A
The terminal third of each spicule is wound round its fellow,
and its tip is slightly bent. Each spicule is provided with a thin
chitinous and finely transversely striated flange; this flange starts
near the base of the spicule and accompanies it almost toits tip. It
becomes broader posteriorly, having its maximum breadth—about
15 p—atits termination, which is about 50 « from the tip of the
spicule. A gubernaculum (text-fig 56) is present; this in side
view is boomerang-shaped, with the posterior end terminating in
a sharp point. Along its outer curve it is 170, long, and its
maximum depth (at the bend) is 17 p.
Text-figure 6,
a = lateral view of posterior extremity of female. An. = anus; Int. = intes-
tine; Ovj. = ovijector; Ut. = uterus; Vul. = vulva. 6 = egg, just after being
deposited.
Two small prebursal papille ave present, one on either side
somewhat ventral, about 170 p in front of the anterior margin of
the lateral bursal lobe.
The posterior end of the female (text-fig. 6 a) tapers abruptly,
so that the tail is relatively short, measuring 114-142 y or about
rr; of the total length. Near its tip there is situated on either
side a small caudal papilla.
The vaginal aperture (text-fig. 6a) is circular, and is situated
a short distance—245 u.—in front of the anus. ‘The cuticle
bordering its posterior margin is slightly protruded. The vagina
is roughly sigmoid in shape and is about 240 pelong. Its lumen
NEW SPECIES OF GSOPHAGOSTOMUM. 467
is lined with chitin, The ovijectors lie in an antero-posterior
direction, being however slightly obliquely placed from side to
side, Inclusive of its sphincters, it measured,in a 2 15mm. long,
535 uw. The right uterus terminates in the anterior ovijector ;
the terminal portion of the posterior ovijector is bent slightly
forward, and the left uterus, which meets it, passes down the body
parallel to that of the right.
The eggs (text-fig. 66) are oval, thin-shelled, and of medium
size, measuring on an average 58x39 4. They commence their
development iz utero and, when laid, the embryo has just formed.
Since writing the above description, I have obtained from
nodules in the ceecum of a third Ground Squirrel, representatives
of a larval stage of O. xeri, sp.n. These larvee were all females
and almost full-grown, being only slightly smaller than the adult
forms. From this it would appear that the larval stages, as in
several other Cisophagostomes, are also passed inside the tissues
of the digestive tract.
From the above description it is apparent that this parasite
belongs to the genus Gsophagostomum ; further, the characters
of the mouth capsule and the cesophageal funnel with its
armature show that it is closely related to @. apiostomum. It,
however, differs from @. apiostomum in two important respects :
namely (1) the spicules of the male are much longer, measuring
nearly 1300 », whereas in @. apiostomum they are less than a
millimetre; (2) the tail of the female is much more abruptly
pointed than in @. apiostomum ; in the latter the distance from
the anus to the tip of the tail is greater than that from the anus
to the vulva, whereas in the new form just the reverse is the case.
These two characters are constant in all the parasites collected, and
may thus be regarded as of specific significance, especially also as
these parasites seem to be normal in a host belonging to a group
of mammals from which no CGisophagostomes had previously been
recorded. For this parasite I propose the name
sophagostomum xeri, sp. n.,
with the following specific description :—Wsophagostomum. Males
about 13mm. and females about 15mm. long. The maximum
breadth of the male is about midway, and is =), of the total
length ; in the female the maximum breadth of about 5 the total
length is reached at the junction of the posterior and middle
thirds of the body. Cuticle transversely striated. Mouth collar
about twice as broad as it is deep. Cervical swelling about as
long as it is broad. Lateral ale absent. Cervical papille
situated at the level of the anterior margin of the posterior third of
the esophagus. External Jeaf-crown of 10 elements; internal leaf-
crown small, and of about twice as many elements. Cisophageal
funnel well-developed ; its anterior margin thickened, and carrying
three recurved tooth-like processes—two subyventral and one dorsal,
468 MR. R. J. ORTLEPP ON A
Three longitudinal chitinous ridges project into the lumen of
funnel. The tail of the female is abruptly pointed, so that the
distance from the anus to the tip of the tail is shorter than that
from the anus to the vulva. Female genitalia typical for the genus.
Male bursa of the usual Cisophagostome type. Spicules long,
tubular, and simple, over 14mm. long, and each is provided with
a lateral flange. Larval stages occur in nodules in the wall of the
execum., ‘
Habitat. Cecum of Xerus setosus (South Africa).
On looking through the literature of Gisophagostomes, I find
that Parona and Stossich in 1901 described as Wsophagostomum
tuberculatum a parasite from Dasypus villosus (Argentine). I
have compared their description and figures with some parasites
I obtained from the same host, and I am satisfied that the
parasites collected by me are the same; further, I am able to
state that this parasite is not an @sophagostomwum, but probably
the representative of a new Trichostrongyle genus.
Literature consulted.
Crepuin. 1849. Nachtrage zu Gurlt’s Verzeichniss der Thiere
in welchen Endozoen gefunden worden sind, p. 54.
Geportst, L. 1916. Notes sur la Faune Parasitaire du Congo
Belge. Revue Zool. Afr. vol. v. fase. 1, Dec. 1916,
pp- 60-62.
Haun, M. ©. 1916. Nematodes parasitic of Mammals of the
orders Rodentia, Lagomorpha, and Hyracoidea. Proc.
U.S. Nat. Mus. vol. 50, No. 2131.
Lanr, C. 1917. Bunostomum kashinathi and the Ancylo-
stomide. Ind. Jour. Med. Res. vol. iv. pp. 414-439.
Leirer, R. T. 1911. The occurrence of Csophagostomum
apiostomum as an internal parasite of Man in Nigeria.
Reprint from Journal of Trop. Med. & Hygiene.
Lixstow, O. von. 1878. Compendium der Helminthologie.
1889, Compendium der Helminthologie Nachtrag.
——. 1906. Zwei wenig bekannte Ankylostomen und @sophago-
stomum dentatum. Centralbl. f. Bakt. Parasit. u. Infek.
vol. xli. Heft 1, p. 90.
Parona, C., & Srosstcu, M. 1901. Csophagostomum “ tuber-
culatum” n. sp. parasita dei ‘* Dasypus.” Bull. dei Musei
d. zool. e anat. comp. d. Univ. d. Genova, No. 110.
Rainier, A. 1895. Traité de Zoologie Médicale. 2. ed. Paris;
p. 418 & pp. 451-452.
Rarur A., & Henry, A. 1906. Sur les Hsophagostomes des
Primates. C. R. Soc. Biol. Paris, vol. lviii. (1) pp. 448-
450.
——. 1909. Sur la classification des Strongylide. IT. Ankylo-
stomine. C. R. Soe. Biol. Paris, vol. Ixvi. pp. 85-88 &
pp. 168-171.
NEW SPECIES OF (SOPHAGOSTOMUM. 469
Raiuuier A., & Henry, A. 1909. Une seconde espéce d’Ciso-
phagostome parasite de 1Homme. Bull. Soc. Path. Exot.
vol, 11, pp. 643-649.
——. 1911. Helminthe du Pore vecueilli par M. Bauche en
Annam. Bull. Soc. Path. Exot. vol. iv. pp. 693-699.
—-—. 1912. les Cisophagostomes parasites de l’Homme.
Arch. d. Parasit. vol. xiv. pp. 562-583.
——. 1913. Sur les Gisophagostomes des Ruminants. Bull.
Soc. Path. Exot. vol. vi. pp. 506-509.
Ransom, B. H. 1911. The Nematodes parasitic in the alimen-
tary tract of Cattle, Sheep, and other Ruminants. U.S.
Dept. Agr., Bureau of Anim. Industry. Bull. 127,
pp. 40-47.
Srines, C. H., & Hassaun, A. 1920. Index Catalogue of Medical
and Veterinary Zoology. Subjects: Roundworms. U.S.
Pub. Health Service. Hyg. Lab. Bull. 114.
Va i hnuhiedlaaiisksl sbadien
inom Mine oak aie inte
ah Pak AERA ADT En UENO, Oe a sto:
on cath vlog Tesla
Lacks: alee hla
“ceo BP dy ke gr itcaontgay i
OEE RR eae (qaral,
Wiebke: tari th
Hie Ae le ae
, TA MMe) a
ae . rdiheurtes a eda CE
ia Saar + th a, wha Baa cubes
Paving Wy i) be ere Te nase meee
Lemar df rahe nee
ik Me mila gp isin
i), eae wae smart gen
Pith in, Wika Sth ea tov biig ath ya) gnc
Ph ye TbiCY CHL a eS ey een ‘NSD Si
i Bey Ratt + pirsilivd.
Pel taiy FDA? TS eee aay a OV ee Ric! ee PR ,
{ “hl ae ‘ine
| ie. ey ¢ ha wee WE ea QWotAen whist, $y ha 1 wy
PERS (iy. Oy ea ied ure fa Tees Rane el le
feelin. Mannie Vins Me. Fels A Gay Ae Pas
) BOARS lias Sega Wea diese Mau. turcitidy al, Minearade ie
; alban = DL orl iat bi a solo pie, Aedes Eh vii cettat id i
ae oth AL i at Mi) 5. rh yy
at a T ae aten TEh wan) Lice aabeh Ate iit 2
, 5 Vad oss Aiea Mwai. gl) Ve Hic 4) nity
oy ; ba) Ee AS, ah? Garret) ay is vary henge
oh Neen dhs be
Pi ta ie me. Ava be iit nie rig and vit } Ath Ai. ae
fl . eet J LvoHn ee tes icon, ep
Ps ce Oras: ipa eeenctrt ntaal Halie ae vanes
: aE Pe hs ae TIO er BS ah an! Wa ey anes © ee i) Se ;
; tar Ne ee rey Tre ‘nv eahea Wie see a) SR Te Vike pag
a Siekrabin, 7 Pak Pein a, aa
a aa
i AL Sitohieee tae irate igs oe
FT eee i ed ce Lee eat hae Path: hea F
iC RAae az te ase cats ae Tih ry: Mi eiionae a Beh a
Bede eet A ee ea tae it fe vag af, Mt ouide ie o
Pe ee sti: ‘Adi i ee
Pate) 11, od el
mete i, Mtap sified ei
sbi: a Mee i) io a ae
hae
ON THE BAT-PARASITE CYCLOPODIA GREEFFI. 471
25. Note on the Dipterous Bat-Parasite Cyclopodia greeffi
Karsch, and on a new Species of Hymenopterous
(Chaleid) Parasite bred from it. By F. W. Uric
(Government Entomologist, Trinidad, B.W.I.) ; Huau
Scott, M.A., Se.D., and J. Wartrrston, D.Se., F.Z.S.
[Received January 26, 1922: Read March 7, 1922).
(Text-tigure 1.)
I. Inrropucrory REMARKS: by HucH Scorr.
The Nycteribiide are a family of wingless, pupiparous Diptera
of bizarre form, parasitic exclusively on bats. A tolerably com-
plete summary of what is knewn as to their biology will be found
in a paper by the writer in ‘ Parasitology ’ (Cambridge), vol. ix.,
no. 4, p. 593, 1917. The only species whose life-cycle has been
at all completely studied is the one which is the subject of this
present note, Cyclopodia greeffi Karsch (= C. rubiginosa Bigot :
see Scott, op. cit. p. 596, footnote). Its life-history was described
in a valuable paper by Rodhain and Bequaert, “ Observations
sur la biologie de Cyclopodia greeffi,” Bull. Soc. Zool. France,
vol. xl., nos. 8-10, pp. 248-262, 1915. An abstract of this work
formed a large part of the writer’s summary in ‘ Parasitology,’
referred to above.
Cyclopodia greeffi was originally described (1884) from the
island of San Thome and the adjacent islet of Rolas, its host
being the flying-fox Hidolon helvwm Kerr (= Cynonycteris
straminea Geoftroy)*. It was further described (1891), under
the name C. rubiginosa, by Bigot from Assinie on the Guinea
Coast (host not stated). The Imperial Bureau of Entomology
submitted to the writer 9 5 and 4 9, now recorded for the first
time, from Ossidinge, Cross River, Cameroons (A. W. Pomeroy
coll., 1. xi. 1915, from a large bat of undetermined species) ;
and Monsieur EH. Roubaud sent for determination in 1916 a
single ¢ from Agouagon, Dahomey (host not exactly named),
Rodhain and Bequaert made their observations at Leopoldville,
Belgian Congo, the host being Hidolon helvum.
In 1920 Mr. F. W. Urich, being in Europe on leave from
Trinidad, went ona mission to San Thomé. There he rediscovered
Cyclopodia greeffi on its original host-species of bat, not only
finding adult examples on the bats, but breeding specimens from
the puparia.
Mr. Urich also bred two examples of a Chalcid hyperparasite
from two of the Nycteribiid puparia. These are described
below by Dr. Waterston as a new species of Hupelmus. ‘They
* Karsch, Sitzb. Ges. Beférd. ges. Naturwiss. Marburg, 1884, p. 77. See also
Speiser, Arch. Naturg. 67. 1. 1901, pp. 54, 65.
472 MR. F. W. URIGH AND DR. H. SCOTT ON THE
are of quite special interest, as they are (so far as the writer
knows) the first insect hyperparasites of any kind recorded from
Nyeteribiidee *, The Hupelmus belongs to a group some members
of which are parasitic on scale-insects (Coccide), and it is
remarkable that the puparia of the Cyclopodia themselves bear
a very strong superficial resemblance to certain scale-insects
which are parasitised by other forms of Hupelmus.
As regards the bionomics of the Cyclopodia, Mv. Uvich’s obser-
vations confirm certain of those of Rodhain and Bequaert. He
mentions the agility of the movements of the parasites, when
disturbed, on the bedies of their hosts—a fact which has been
noted also concerning other species of sa es These
insects are, as stated above, “‘ pupiparous,” 7. ¢. the females give
birth to full-fed larvee, ae integument ae which immediately
hardens and darkens to form the puparium. It appears that
some species of Nycteribiide attach their larve to the bodies of
their hosts (see Scott, ‘ Parasitology,’ ¢ ¢. p. 598), but in
Cyclopodia greeffi this is not the case. The females of this
species attach thei Jarve to parts of the trees in which the bats
have their sleeping -places. Rodhain and Bequaert made their
observations on bats in captivity, and the majority of the
Nycteribiid larvee were fixed to the undersides of wooden perches
in the cages: these authors think that in a wild state the larve
are probably attached to the smooth trunks or branches of
Dracena-trees, which are the favourite sleeping-places of the
bats near Leopoldville. In San Thomé Mr. Urich found the
puparia fixed to the upper and lower surfaces of the smooth
leaves of certain dicotyledonous trees (name not stated) in which
the bats slept.
Rodhain and Bequaert describe how the female Cyclopodice
with the undersides of their bodies press their larvee down on to
the substratum, to which the larva firmly adheres. It imme-
diately assumes the shape of, and hardens and darkens to form,
the puparium. This is a half-ellipsoidal body, with elliptic
contour, rather narrower behind, convex dorsally and quite flat
ventrally, where it is “glued” to tne substratum. Its dorsal
and ventral surfaces are separated by an angular margin. The
dorsal surface becomes uniformly black, and the resemblance of
the puparium to a black scale-insect (such as Saissetia ole) is at
first sight very great, though with closer examination traces of
segmentation and the position of the two pairs of larval spiracles
are discernible. The puparium is figured by Rodhain and
Bequaert, op. cit. p. 258.
The flat ventral surface of the puparium remains colourless
and transparent, and if pupee are detached from their substratum
various phases in the development of the enclosed nymph can be
* As regards other organisms hyperparasitie on Nycteribiide, the latter some-
tines bear on their bodies Laboulbeniaceous fungi: see Speiser, "Arch. Naturg. 67.
mae p. 29; Seott, A Naturg. 79. A. 1913, Heft 8, p. 96 (1914), and Ann.
Wan 3 Nat. Hist. (8) xiv. p. 234 (1914).
BAT-PARASITE CYCLOPODIA GREEFFI., 473
observed through this ventral transparent covering. In the
material sent by Mr. Urich several stages can be seen, including
a fully-formed adult nearly ready to emerge from the puparium.
Sufficient has now been written to introduce Mr. Urich’s own
report of his observations.
TI. Opservations in San THomvf in 1920: by F. W. Uricu.
The host of the Cyclopodia was the flying-fox Widolon helvum
Kerr (= Cynonycteris straminea Geofir.), which was common
and numerous in the north-western part of the island of San
Thomé. These flying-foxes seem to have established themselves
in the woods situated near Morro Moguingui (about 3 miles from
the sea)—at least, that is where they passed the day. Every
evening, as soon as the last rays of the sun disappeared, they
sallied out in thousands. They all flew towards the interior,
those going far kept very high and the others whose objective
was near by just kept above the tree-tops; in the morning just
before sunrise, between 5 and 6, they could be seen returning to
their sleeping quarters. Their food consisted principally of the
ripe fruit of the papaya, Carica papaya, which grew wild all
over the island. At the island of Principé, where the same
species of flying-fox occurs, I was told that they attacked cacao
pods, papaya not being so common there. The trees selected for
sleeping were very tall and slender-stemmed, and the flying-foxes
were settled so close together that two shots from a fowling-piece
brought down 20 specimens. This roosting-place was known to
be used for years, and the small branches of the trees showed
very old scars and scratches from the flying-foxes’ claws; the
ground under the trees was covered with young papaya seedlings,
and a peculiar acrid odour from accumulated excrement prevailed
under the trees*. The dead specimens were put into cloth bags
as quickly as possible to prevent the escape of «ny Nycteribiid ;
the parasites moved about quickly on the bodies of their hosts,
but I did not observe any actually darting about. ‘The leaves of
the trees used for roosting were dotted with numerous puparia.
A hundred leaves collected at random showed 61 puparia on the
top surface and 96 underneath. The leaves were smocth on both
surfaces. The skin of the larva must dry and harden very
quickly ; dead specimens were placed in cloth bags at about
2 p.m., and four hours after two fully hardened and coloured
puparia were found sticking to the cloth. [Rodhain and Bequaert,
op. cit. p. 257, state that the hardening and darkening process is
complete in 20-30 minutes after the birth of the larva. |
* [Habits not unlike those here described have been noted among various kinds of
large fruit-eating bats by other observers. H. g., in the Seychelles the local flying-
foxes (Pteropus sp.) have regular sleeping-places in particular clumps of trees, and
“Camp Chanve-Souris ” has become a place-name in at least one island (Silhouette).
In Mahé (1908) the bats might frequeutly be seen in numbers passing northwards
along the west coast; of the island at sunset, on their way (it was said) from one of
their “camps” to their feeding-grounds, which were said to be places where bread-
fruit and other fruit trees were plentiful—H. Scorr. |
47 4. DR. J. WATERSLTON ON A NEW
No larvee or puparia were found on the flying-foxes. Newly
emerged adult specimens, kept in cloth bags with leaves, died
after 48 hours and never assumed the full coloration. When
placed on the skin of a rabbit they made feeble attempts as if to
feed, but desisted after a short time. [Rodhain and Bequaert.
found that specimens newly-emerged from puparia cannot survive
more than 48 hours unless they find a host ; and that examples
removed from their host-bats could not be induced to feed and
very soon died. |
Two examples of a hymenopterous hyperparasite issued, one
from each of two puparia. In emerging they made a rather
jagged-edged hole in the dorsal surface of the puparium. They
are described below by Dr. Waterston.
III. A new Lupelmus (£. wrichi, sp. n.) FROM SAN THOME :
by James WATERSTON.
(Offered for publication by permission of the Trustees of the British Museum.)
The species now described might well, owing to its antennal
and thoracic peculiarities, be made the type of a new genus. I
do not propose to take this step however, partly because it is not
yet clear how far the coalescence of funicular joints can be relied
upon for systematic purposes, but chiefly because only one sex is
at present available for study.
It should be noted in connection with the colour characters
given below, that the specimens studied had been in alcohol for
some time before mounting. It is likely that they now appear
to be somewhat duller than in life.
EUPELMUS URICHI, sp. n.
g. A blackish-brown to nearly black species with the follow-
ing parts more or less pale ferruginous: scape (except at tip),
mandibles (dental apices dark), all trochanters, knees of fore and
mid legs and tibial apices of the same, and tarsi except for the
5th jomt in fere and 4th and 5th of mid and hind legs whieh
are more or less infuscated. Metallic lustre, only distinctly
showing on propodeon and dorsum of abdomen, reflections on
head and thorax faintly eneous, more distinetly purplish green
on propodeon and base of abdomen.
The wings, especially the fore pair on distal half, are distinctly
infumated, with the veins still darker.
Head (15: 13): mouth opening half as broad as head ; the eyes
ave bare and occupy two-thirds of the depth. At their nearest
separated by five-twelfths of the breadth. Toruli just above the
base line of the eyes. The shallow straight-edged clypeal emargi-
nation is in length subequal to the distance between the toruli.
Except in the scapal grooves, the entire frons and face have a
moderately fine distinctly raised sculpture, the surface being dull,
with very numerous short, stiff bristles, of which two (1:1) above
the corners of the clypeal emargination are longer.
CHALCID PARASITE OF CYCLOPODIA GREEFFI. 475
The toruli at their nearest (superiorly) are separated by about
the long diameter of either and basally by about two diameters.
They stand from the orbits four-fifths of a diameter.
Text-figure 1.
Eupelnus urichi, sp. n. g:—a, antenna ; 6, propodeon (p, upper edge of peritreme) ;
e, spiracle at its greatest length and breadth seen from the direction indicated
by the arrow.
476 DR. J. WATERSTON ON A NEW
Antenna: length 1-25 mm. (see text-fig. 1), consisting of scape,
pedicel ving joint, five normal funicular joints and a large
sausage-like club (consisting of two funicular and three normal
club segments fused).
Mouth parts.—Labrum (3: 2) with convex edge and five stout
fringing bristles of which the longest equals the breadth of the
sclerite. Mandible (4:3) bidentate, the upper tooth smaller
and rather abruptly and obliquely truncated superiorly. Stipes
smooth, about a dozen bristles. Max. palp. (measured along
outside edge) 11, 12, 15, 36. In same ratio the breadth of
joints 1-3 is about 10 and of 4, 15. .Terminal bristles of 4th
about half the length of that joint which bears also 10 stout
bristles on apical half and about 20 finer more generally distri-
buted. Labial palpus 15, 9, 15 with a breadth at the joints of
12-14. Apical bristles distinctly longer than third joint which
bears also about a dozen other bristles mainly stout. There are
no fewer than 10 (5: 5) setigerous cells on the lingua.
Thorax.—Pronotum, in two separate rhomboidal sclerites each
with the inner anterior angle rounded off. The outer anterior
angle is weakly chitinized. The postero-lateral angle is folded
down enclosing the spiracle. Hach sclerite bears (mainly on the
posterior half) about 50 stiff bristles of which 6-8 stand along
the hind margin between the mid line and the spiracle. Pro-
sternum, bristles (8-9 : 8-9). Prothoracic surface generally
nearly smooth, duller and raised reticulate at the sides in front of
the spiracle. Mesonotum very broad, only a little longer (7 : 6)
than wide, divided almost equally by the transverse suture, the
scutellum and axille being relatively large. Parapsidal furrows
deep, mid lobe (measured with the prothorax removed) just
longer than the scutellum. The intercept of the parapsidal
furrows on the transverse suture (4 times the narrow base of
the scutellum) occupies rather over one-third of the breadth.
The pattern on the mid lobe is moderately coarse and raised ;
of the parapsides and axillee a little lower, while the scutellum is
nearly smooth with over 40 bristles equally distributed about
a bare mid line. Before the suture the surface is densely bristly
and there are about a dozen on each axilla. These notal bristles
are a little longer posteriorly with six (5:3) above the hind
edge of the scutellum distinctly stronger than the others.
Metathorax.—All over finely reticulate, pattern hardly raised,
the three regions well defined. Mid area posteriorly crenulate,
lateral areas with one bristle each.
Propodeon smooth with strong reflections, contoured as in text-
fig. 1, shortly rugose round the insertion of the petiole. With a
delicate percurrent mid line but no keel. Spiracle facing
anteriorly and outwards, only the edge of the peritreme visible
from above. There are about 15 bristles behind and beyond
the spiracle.
Mesosternopleure nearly all smooth, the only indications of a
raised pattern being on the prepectus (posterior half) and in
CHALCID PARASITE OF CYCLOPODIA GREEF I. AT7
front of the femoral furrow where there are 12-15 bristles.
About the same number stand in a triangular area on the
mesosternum posteriorly and there are one or two more at the
sides.
Forewings : length 1-3 mm., just over twice as long as broad,
submarginal, radius, postmarginal, 21: 13:5: 10, submarginal
13 bristles. Radius, 4 on stalk and 10 on club.
Legs robust. Fore-legs: femur (45 : 15), tibia (32:7) with
three peg-like apical spines or teeth, tarsus 52, 32, 24, 20, 28.
Mid legs: femur (nearly 5:1), slightly shorter (14:15) than
the tibia (15 : 2), spur as long as 1st tarsal joint, tarsus 51, 36,
27, 21,28. Hind-legs: coxa long, ? the femur (7% 2). Tibia
(9: 1) apical comb 12-14 spines; longer of the two spurs not
2 the Ist tarsal jomt. Tarsus 68, 38, 27, 21, 28. All legs
densely set with bristles.
Abdomen smooth. ist (3rd) tergite shortly incised on posterior
edge, more heavily chitinized antero-medianly (basally) with
traces of sculpturing on the thickened area. Other tergites
simple, transverse and entire. Spiracle minute, circular. Stylet
very short, 5 bristles. The petiole ventrally shows two chitinous
ridges. The sternites show a median weakly chitinized line.
Entire ventral surface very bristly.
Length about 2°3 mm.
Expanse about 3:2 mm.
Type ¢ (dissected on slide) in British Museum, one of two
3 reared ex puparia of Cyclopodia greeffi Karsch, a Nycteribiid
parasite of the flying-fox Hidolon helcwm Keryv (Oynonycteris
straminea KH. Geoftr.). W. Africa, San Thomé, Oct. 1920. F. W.
Urich coll.
ROC ACO SOC. 1922) No. XOX XU:
bo
Os
: (pe) .
ane %
yh i, sgh RA
: t
we a AGAR) Aa te:
; ’ 7m a
w
a a\ t ; py: { " a
ea, Oe Ne wa On
ay Le oe )
Poreee Marsiharaeyl oe ww
*
wil in 7)
ON METAMORPHOSED MEXICAN SALAMANDERS. 479
EXHIBITIONS AND NOTICES.
March 21st, 1922.
Dr. A. Smirm Woopwarp, F.R.S., Vice-President,
in the Chair.
The Secrerary read the following Report on the Additions to
the Society's Menagerie during the month of February, 1922 :—
The registered additions to the Society’s Menagerie during the
month of February were 126 in number. Of these 39 were
acquired by presentation, 81 were deposited, 5 were purchased,
and 1 born in the Menagerie.
The following may be specially mentioned :—
2 Elephants (Llephas maximus), from India, presented by
H.H. The Gaekwar of Baroda on February 18th.
1 Lioness (felis leo), bred in India, presented by H.H. The
Maharajah of Mayurbhanj on February 18th.
1 Allamand’s Grison (Grison allamandi), from Pernambuco,
presented by Lt.-Commander Rutherford Collins, R.N.R.
April 4th, 1922.
Prof. EK. W. MacBrinz, F.R.S., Vice-President,
in the Chair.
Mr. C. Tatr Recan, F.R.S., F.Z.S., exhibited, and made
remarks upon, a series of lantern-slides illustrating specimens of
various blind fresh-water. Fishes from eaves.
Mr. R. H. Burne, M.A., F.Z.S., exhibited, and made remarks
upon, a series of specimens demonstrating the recessus orbitalis
in Flat Fishes.
Dr. L. Hogpen, M.A., F.Z.8., exhibited some small examples
of metamorphosed Mexican Salamanders (Amblystoma tigrinum),
and remarked as follows :—
“ Significant advances have been made during the past seven
or eight years in the study of the part played by ductless glands
in Amphibian metamorphosis. That the thyroid seeretes an
iodine compound which stimulates the assumption of adult
characters, and that the removal of this gland results in a
480 THE SECRETARY ON ADDITIONS 'TO THE MENAGERIE.
permanent retention of the larval organisation may be regarded
as definitely established by the work of Gudernatsch, Bennet
Allen, Hoskins, Swingle, Morse, Uhlenhuth, and others. There
are ample ei eanione sels the thyroid gland is not alone in
influencing the developmental processes of “Batt ‘achians. Several
recent workers, nota bly Allen, have demonstrated. the failure of
Anuran tadpoles to undergo metamorphosis after the ablation
of the pituitary (anterior lobe) rudiment. But our knowledge of
the function of this organ is very slight.
“A short while ago the author, in collaboration with Mr.
Julian Huxley, independently confirmed Laufberger and Jensen’s
claim to have induced sexually mature larvee of the neotenous
Mexican Salamander to assume the adult form by treatment with —
a diet of mammalian thyroid. It was not possible after many
attempts to obtain this result with pituitary (anterior lobe) gland
substance administered per os. Recently it has been found
possible to induce year-old Axolotls in Prof. MacBride’s laboratory
at the Imperial College of Science to transform after tri-weekly
intr avenous injections of anterior lobe extracts. The injections
of -3 cc. Ringer extract of equivalent to 2 grains fresh gland (ox)
were made via the anterior abdominal vein. The animals were
fed on raw beef tri-weekly, and kept in deep water till the
completion of metamorphosis at the conclusion of one month
from the beginning of the treatment. The time required by this
method is thus the same as that in the case of thyroid dieting—
conspicuously less than with the shallow-water treatment of
Mlle. de Chauvin and KE. G. Boulenger.”
April 25th, 1922,
Sir Srpney F. Harmer, K.B.E., I'.R.S., Vice-President,
in the Chair.
The Secretary read the following Report on the Additions to
the Society’s Menagerie during the month of March, 1922 :—
The registered additions to the Society’s Menagerie during the .
month of March were 157 in number. Of these 61 were
acquired by presentation, 20 were deposited, 70 were purchased,
2 were received in exchange, and 4 were born in the Menagerie.
The following may be jasoeilly mentioned :—
3 Alpine Marmots (M/armota marmota), from Argentiéres,
Haute Savoie, presented by Major J. R. Peploe on March 27th.
3 Canadian Tree-Porcupines (Hrithizon dorsatus), from
N. peneriee, received in exchange on March 22nd.
2 Reindeer (Rangitfer tar andus), from Norway, purchased on
Maren 21st.
THE SECRETARY ON ADDITIONS ''O THE MENAGERIE. 48]
1 Black-gloved Wallaby (Macropus irma), from W. Australia,
presented by Capt. F. J. Ogilvie on March 31st.
1 Nepal Scops Owl (Scops bakkamena lettia) and 1 Indian
Kestrel (Cerchneis tinnunculus saturatus), from Darjeeling, new
to the Collection, presented by Mr. H. Stevens on March 11th.
1 Bonelli’s Eagle (Hntomactus fasciatus) and 1 Great Eagle-
Owl (Bubo bubo), presented by the Hon. Mrs. Willoughby Verner
on March 31st.
The Secretary exhibited, and made remarks upon, some
dressed skins of a Tyree-Hyrax (Procavia valida?) from Tan-
ganyika Territory.
Prof. P. 'T. Frynn exhibited, and made remarks upon, a cast
of the skull of a Squalodont Whale from the Tertiary (Miocene)
Strata, Table Cape, Tasmania.
The SecRETARY communicated on behalf of Mr. Arruur
LoveripG£, C.M.Z.8., an account of Mr. Loveridge’s experiences
while watching “ Lions at their Kill.”
Proc, Zoou. Soc.—1922, No. XX XIII. 33
aanricnhic hia
teat be Holt, fait de bk
ip r nate i otis
Eich dase piri
as RL 1 A
PE oo hin
on
a
ht ca ae”
Stead
aos fy mig of Suan
Ai . ne oa
pt
ZOOLOGICAL SOCIETY OF LONDON.
Tus Society was founded in 1826 by Sir Sramrorp Rarruss,
Mr. J. Sasine, 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.
Patron.
HIS MAJESTY THE KING.
COUNCIL.
HIS GRACE THE DUKE OF BEDFORD, K.G., F.RS., President.
Tae Hon. Cecrt Barine, M.A.
Lr.-Cou. 8. Monckton Copeman,
M.D., F.B.S.
Caartes Drummonp, Ksa.,
Treasurer.
Aurrep Hzra, Ese, O.B.E.,
Vice-President.
Hueu 8. Guapstons, Ksa., M.A.,
F.R.S.E.
THe Rr. Hon. rae Viscount |
Grey, K.G., P.C.
Sir Sipnuy F. Harmer, K.B.E.,
MAG Secale) HAR Sia: 0 hee
President.
Mason tHe Lorp Anasrarr
Rosert Iwnes-Kzur, D.S.O.
Pror.ERvust W.MacBring,D.Sc.,
LL.D., F.R.S., Vice-President.
Cor. Sir A. Henry McManon,
G.C.M.G.,
President.
HK. G. B. Muapz-Watpo, Ese.
P. Cuatmers Mircuenr, Esa.,
C.B.E., M.A., D.Sc., LbL.D.,
F.R.S., Secretary.
Tae Karn or Onstow, O.B.E.
Masor AtBerr Pam, O.B.E.
Tur Lorp QuEnnBoRoUGH.
K.C.LE., Vice-
Tur Marqusss or Sxi¢o, F.S.A.,
Vice-President.
Pror. G. Exziior Suira, M.A.,
M.D., F.RBS.
Ricuarp 8. Taytor, Esq.
AntHony H. WrneFrexp, Esa.
A. Smita Weopwarp, KEse.,
LL.D., F.R.S., Vice-President.
2
The Society consists of Fellows, and Honorary, Foreign, and
Corresponding Members, elected according to the By-Laws. It
earries out the objects of its foundation by means of its collection
of living animals, by its Library, and by its Scientific Publications.
The Office of the Society, Regent’s Park, N.W.%, where all com-
munications should be sent, addressed to “The Secretary,” is open
from Ten till Five, except on Saturdays, when it closes at OnE P.M.
The Library, under the superintendence of Mr. F. Martin Duncan,
F.Z.S., F.R.M.S. is open daily (except Sunday) from Ten a.m. till
Five p.m.; on Saturdays, Ten a.m. till One p.m.
The Library is closed from Good Friday to Easter Monday, and
upon all other Bank Holidays. It is also closed annually for
cleaning purposes during the whole month of September.
The Meetings of the Society for General Business are held in
the Meeting Room at the Society's Office on the third Wednesday
of the month at 4.30 p.m. except in September and October.
The Meetings for Scientific Business are held in the Meeting
Room at the Society’s Office fortnightly on Tuesdays, except in
July, August, September, and December and January, at half-past
Five o’clock p.m.
The Anniversary Meeting is held on the 29th of April, or the
nearest convenient day, at Four p.m.
The Society’s Gardens are open daily from Nine o’clock until
Sunset. Mr. R. J. Pocock, F.R.S., F.L.S., is the resident Super-
intendent and Curator of Mammals, Mr. D. Seth-Smith is Curator
of Birds and Inspector of Works, Mr. E. G. Boulenger is Curator of
Reptiles, Miss L. K.Cheesman, F.E.S., is Curator of Insects. Appli-
cations for anatomical material or facilities for work in the
Prosectorium should be addressed to the Secretary.
TERMS FOR THE ADMISSION OF FELLOWS.
Frttows pay an Admission Fee of £5, and an Annual Contri-
bution of £3, due on the Ist of January, and payable in advance,
or a Composition of £45 in lieu thereof; the whole payment,
including the Admission Fee, being £00.
No person can become a Frrnow until the Admission Fee and
first Annual Subscription have been paid, or the annual payments
have been compounded for.
Ferttows elected in November and December are not liable for
the Subscription for the vear in which they are elected.
PRIVILEGES OF FELLOWS.
Fetitows have Personal Admission to the Gardens. upon signing
their names in the book at the entrance gate, and may introduce
Two Companions daily.
The Wire or Houssanp of a Futtow can exercise these privileges
in the absence of the Fellow.
Until further notice, Frtnows will receive 40 undated Green
Cards, available on any Sunday or week-day up to the end of
February of the year following the year of issue, and 20 White
Cards available on any week-day up to the same date. Twenty
of the Green Cards may be exchanged for a book containing two
Orders for each Sunday in the year. Twenty White Cards 'may
be exchanged for a book of dated Week-day Orders, each Order
available for any day during the week except Sunday. Special
children’s tickets are no longer issued, but the Green and White
Cards are perforated, and each half is valid for a Child under twelve
years of age. It is particularly requested that Fellows will sign
every ticket before it goes out of their possession. Unsigned tickets
are not valid.
Frttows are not allowed to pass in friends on their written
order or on presentation of their visiting cards.
Futiows have the privilege of receiving the Society’s ordinary
Publications issued during the year upon payment of the additional
Subscription of One Guinea. ‘This Subscription is due upon the
lst of January, and must be paid before the day of the Anniversary
Meeting, after which the privilege lapses. FEttows are likewise
entitled to purchase these Publications at 25 per cent. less than
the price charged to the public. A further reduction of 25 per
cent. 1s also made upon all purchases of Publications issued prior
to 1881, if above the value of Five Pounds.
Ferttows also have the privilege of subscribing to the Annual
Volume of ‘The Zoological Record,’ which gives a list of the Works
and Publications relating to Zoology in each year, for the sum of
One Pound Ten Shillings. Separate divisions of volumes 39 onwards
can also be supplied. Full particulars of these publications can
be had on application to the Secretary.
Frettows may obtain a TRANSFERABLE Ivory Ticker admitting
two persons, available throughout the whole period of Fellowship,
on payment of Ten Pounds in one sum. A second similar ticket
may be obtained on payment of a further sum of Twenty Pounds.
“2
4
Any Fetxrow who intends to be absent from the United Kingdom
during the space of at least one year, may, upon giving to the
Secretary notice in writing, have his or her name placed upon the
“dormant list,” and will then be called upon to pay an annual
subscription of £1 only during such absence, or in the event of
returning to the United Kingdom before June 30th in any year to
pay the balance of the ordinary subscription. After three years a
Dormant Fellow must make a further application to be retained on
that list.
Any Frtiow, having paid all fees due to the Society, is at liberty
to withdraw his or her name upon giving notice in writing to the
Secretary.
Ladies or Gentlemen wishing to become Fellows of the Society
are requested to communicate with “The Secretary.”
P. CHALMERS MITCHELL, -
Secretary.
Regent’s Park, London, N.W. 8.
June, 1922.
MEETINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON
FOR
SCIENTIFIC BUSINESS.
1922.
LUESDAV. sUINE eee eee “13:
#6 OCTOBER ......... 24.
Bs NOVEMBER ...... 7 and 21.
The Chair will be taken at half-past Five o'clock precisely.
ZOORUGICAL, SOCIETY OF “LONDON,
LIST OF PUBLICATIONS.
THe scientific publications of the Zoological Society of London
are of two kinds—“ Proceedings,” published in an octavo
form, and “ Transactions,” in quarto.
According to the present arrangements, the ‘‘ Proceedings”
contain not only notices of all business transacted at the scien-
tific meetings, but also all the papers read at such meetings
and recommended to be published in the ‘“ Proceedings” by
the Committee of Publication. A large number of coloured
plates and engravings are issued in the volumes of the
“ Proceedings,” to illustrate the new or otherwise remark-
able species of animals described there. Amongst such
illustrations, figures of the new or rare species acquired in a
living state for the Society's Gardens are often given.
The “‘ Proceedings” for each year are issued in four parts,
paged consecutively, during the months of March, June,
September, and December. From January 1901 they have
been issued as two half-yearly volumes, indexed separately.
An ‘“ Abstract of the Proceedings ’”’ is published by the
Society on the Tuesday following the date of the Scientific
Meeting to whichit refers. It is issued along with the “ Pro-
ceedings,” free of extra charge, to all Fellows who subscribe to
the Publications, but it may be obtained on the day of publi-
cation at the price of Sixpence, or, if desired, sent post free
for the sum of Six Shillings per annum, payable in advance.
The “ Transactions” contain such of the communications
made to the Scientific Meetings of the Society as, on account of
the nature of the plates required to illustrate them, are better
adapted for publication in the quarto form. They are issued
at irregular intervals. —
Fellows and Corresponding Members, upon payment of
a Subscription of One Guinea before the day of the Anni-
versary Meeting, are entitled to receive the Society’s
Publications for the year. They are likewise entitled to
purchase the Publications of the Society at 25 per cent. less
than the price charged to the Public. A further reduction
of 25 per cent. is made upon purchases of Publications
issued prior to 1881, if they exceed the value of Five
Pounds.
Fellows also have the privilege of subscribing to the
Zoological Record for a sum of Two Pounds Ten Shillings
(which includes cost of delivery), payable on the 1st of July
in each year; but this privilege is forfeited unless the
subscription be paid efore the 1st of December following.
The following is a complete list of the publications of the
Society already issued.
TRANSACTIONS OF THE ZOOLOGICAL SOCIETY OF LONDON.
4to. 20 vols. and Index. ’ Price to Price to the
Fellows. Public.
Vols. I.-IV. (out of print).
Vol. V., containing 67 Plates.) (1862-66) 7.0.5 4 BP 61D 0
Be Wale s 92 the dete REO -OU)) te eamIN 0) 40) eneeeine en mer ee
3 ALLS, +) 73 ir feo OOO— (2) ecu. 4 Jp hued Le a0
5) WALLS Fp eare2 }, (0872274) pee Oo 88 idan tema ids 70
Ae MeL Nes i 99 3p ee ALOFT), TREE TRG 2 eee Gree aa
5 Gy " 95 0 Bev LETTE) Wg otape Ole 0) cekvrar irs dalle yaaa
Index sVidllss ILoX. faci teins fds fap ame (1883-79) 04) 0%. 6h. 010 0
Vol. XI., containing 97 Plates .. (1880-85) .... 912 0.... 12 16 O
nak. LAs Soleo 65 eto! (Uusihe ON) CL AS) ately Oi. TUA> 10
Be” Nee 5 62 eee CSOT JO) ines. COLetSune, 811 0
| WL * 47 se (ustewcyy Bo so namayey OS Oe
5 UE... ‘5 52 » © «- (1898-1901) 5. 6 16 6. 714 0
pee KEV IL a k3S 5) A A901—1908) 29985 Heir Om 7 4 0
Fj; ee, hal Sve yh Ag 4) GISOSAISIUD) nes Galle (mikey 0)
ILLES, wa Wad Pirie CISC ARID RY, sie iy Ons 5 8 0
5 MOLL XE, 5 24 pil Mag (LGOO=19L0) ea. LO Ore 13 12 0
5h ote) NO BOL DS ye GLOU2 SONS) eee ON Sy: 5397) 40
» XXIJ.—Part 1. (7 Plates & 12 Text-figs.)
(Sine AOC) Sear ae Da AO ay aus One am
now able to offer for sale, at the reduced price of £30, sets of Vols. V._XVI. inclusive, and
In consequence of a re-arrangement of the stock of the ‘Transactions,’ the Society is
eeparate papers, of which a list can be supplied, at about one-fourth their published price. |
PROCEEDINGS OF THE COMMITTEE OF SCIENCE AND
CORRESPONDENCE OF THE ZOOLOGICAL SOCIETY OF
LONDON. 8vo. 2 vols. (Letterpress only).
Price to Price to the
Fellows. Public.
Part I. 1830-31. 1 vol. 8vo., out of print.
“4 UU Relves 59 Mar ci aa. y's ER As; (Gis peng NOSs
PROCEEDINGS OF THE ZOOLOGICAL SOCIETY OF LONDON.
First SERIES.
Parts 1-XV. (1883-1847). 8vo. 15 vols. (Letterpress only.) Price to
Fellows : 4s. 6d. each part ; to the Public, 6s.
Index 1830-1847. Price to Fellows: 4s. 6d.; to the Public, 6s.
Parts I., VIL.-IX., XI., XIV., XV., out of print.
SECOND SERIES.
Parts XVI.-XXVIIT. (1848-1860). 8vo. 13 vols. (Letterpress only.)
Price to Fellows: 4s. 6d. each part ; to the Public, 6s.
Index 1848-1860. Price to Feliows: 4s. 6d. ; to the Public, 6s.
The Parts of this series containing Coloured Plates are out of print.
PROCEEDINGS OF THE SCIENTIFIC MEETINGS OF THE
ZOOLOGICAL SOCIETY OF LONDON. 8vo. 40 vols. and 4 Indices.
Letterpress only. With Plates uncoloured. With Plates coloured.
Price to Price to the Price to Price to the Price to Price to the
Fellows. Public. Fellows. Public. Fellows. Public.
SGI par sMmOds fF 3. OSs eas eene Qs. PPM eR EEA) cea Seat At NT 33s. 9d. .... 45s.T
1862 4s. 6d. .... 6s 9s, Deer s PLS). des. 9d. .... 45s
18638 GMOS as s.ts, OSL Me koala Qs. PSN are Ares. 38s. 9d. 45s.+
1864 ASOG. ..\:)1 OS. aha oe 9s. spre hoe dos. Od. .... 45s.
1865 ASHNGOSy ... ae OStipo meet 9s. ye LAS iene ae 30s. Od. .... 45s
1866 Als. NGG apeeste "OCG 6 0.0-0.0% 9s. be See ey Sa, YW, son, 4En8
SOMES conn. eee 9s. 12s.*, dos. 9d. .... 45s
NSOSM ek: & < od cape eee 9s Pgs Bee ee Bie, Wh sso. Ze
SCO MRE aS siesta 9s.» OR ROSS Fa S oes. 9d. 2.5.) 45s.
SORT: Ss 5.3. buch taarmnenne 9s. BUN LEY Med ag 33s. 9d. .... 46s.
Index5 1861-1870: yaie, Sarees AS PAG Gla ee 6s.
Sots ee RR es ee orca as 9s IDS eon OEE mee 29
LG 2) aE RMR me ST a daar A, 9s OE ea Slee 30s. 9d. .... 458.
TEST ae ae a ibe hale ee AER VS 9s 12s ete 30s. Od. .... 458
HS //<: Wie Le RRL R AR DYE LAL UNS 9s TA ee pie 36s 48s.+
I fey Saipan (oe > se, uate as LIME Boda 2 9s 1 he eee 36s 48s
SAG eis Res ele ks See Aes oe 9s Ie eens 36s 48s.
1 (7 GRR SE eR STANT het 9s Ds Mace 36s 48s
MSS kisi each ang aoe ae ee ee 9s VOD ees peers 36s 48s
LUCIA ee Mangnerer: sorteneren 08 1 he treat ih Ot 2 ahd 9s. je .§ 6 G8 Wa eu 36s. Ei aake AS st
TLEXS 0) ere RPMS on ths acer me SalI NE 9s. Rew MOS! eth a 36s. Mies, AGae
index, 1871=1880) ap os ae teases 4s, Gd. .... 6s.
1881-1890. Ten volumes .... Hach ................ 36s. Soom, ake,
Index, 1S81=1890 peewee eee As. Gdten!. 3 6s.
1891-1900. Ten volumes .... Bach ................ 36s. Mee Ass:
index, 18911900 eo eae - As Ou). G8.
* No perfect copies in stoek. t Out of print.
PROCEEDINGS or tar GENERAL MEETINGS ror SCIENTIFIC
BUSINESS or tox ZOOLOGICAL SOCIETY OF LONDON.
S8vo. 32 vols. and Index.
Priceto . Price to the
Fellows. Public.
1901-1910. Twenty volumes: Fives 1.0 irl Hachel ess eae 24s,
Index, 1901-1910 105... A PALAERESG te oe ce a a AS MOC seh 6s.
TOI, VOL. OU. : s.2 pice sage 40 3 Mipetelecaeta sie Mi <p oem RAS sah eee Re 24s.
* ah Milica nee falco ns aienie Go epere ne keneiia tones) o\'+) eel site Rama teMercs ve) <\i ame Mets Ss; Br tone
1912, ») Ds Se ie ee RE AT BA he hae 2 ee ee 133. eer 24s. *
¥ pe AL, ‘apo a tales one ee em Pre is eC he - ae 88. sere Le
1913; A TO, SLE a gaaeh ei oe ene VR ee eae 18st fee 24s.
5 rap Ul nese ts Sore hy ade oe cho oe Mee an oye LSS .0).. eee 24s
LOVE ie ees Reem Mr role WAAR oi atig MARS onl bok 18S: 02. 24s,
Pd Me RENE cnc 6 Gs Giri ruc CIC ce Aaah 09 cae Boo Hc ct LB Ss oc eee 24s,
TONG; oy Decides sini > ee Bee OMA ROO Ae. eee 18s... 54, ese
a Per | NR a RABE os eeciamn SG yo aadokes tG ues LSS: sige eae
TOUS E TBS Hee 0. BEES AE Oe teat ket ar es ae eam LS siicd tage 24s
si vopnn LEGS 2) SC Eon telae EPMA eC PO ats hoe a opejL Sse ape 24s
Ae ie Aarne ot bado so ake todas 4 et no trae 2 8a! eee 24s,
Pe AUR GARE 5, SERED Sag eae de ESTA oe Sa oly ee 933 hae 12s.
TES eed nner a at oan octkn Genes oo bp. dc.5, 0.4 OS: pox eee 12s.
Sli 0 DAN ona Nooo cise can ata 0 nicinbke Cuboaie ise os a Os & WA 12s.
ICC AA eel ROLE MSS C8 UN Gk a: hey Cueibla Gora alc asa to TOst4 ce OS
vias 0 Peco. ceo une: coe tg ay esc adits « 15s tees 20s
i112 (0) et gaa Ober Marae earch elo cs Mirae cit a9 Raaalerss a n'a 5" Whe) LOS. cents ee 20s.
ey UL Vivek GE ie ieee Pbe.1 oa yr cies RE Bette er ghysree ISS. seen 24s
OO SL, vc chee) ares +. cue) pe Megane reo ee nckeT = so Cee owen | Rey jamreiatrsiss 24s.
Be PEL Foca 4 Shes yc Be wae Cx. Sa sity Tare eee 18s: \eoaeee 2As,
ICS oy hi el ea a 3s 3 "> o alia dole cea gle Weak Snipa tes Osta ae 12s.
LISTS OF THE ANIMALS IN THE SOCIETY’S GARDENS.
List of the Vertebrated Animals now or lately Living in the Gardens
of the Zoological Society of London. (Kighth Edition.) 8vo.
1883. Cloth, 4s. 6d.
List of the Vertebrated Animals now or lately Living in the Gardens
of the Zoological Society of London. (Ninth Edition.) 8vo.
1896. Cloth, 6s.
CATALOGUE OF THE LIBRARY of the Zoological Society of
London (Fifth Edition.) 8vo. 1902. Cloth, 6s.
THE OFFICIAL ILLUSTRATED GARDEN GUIDE—20th Kdition
(Revised)—with (1) a Railway and Street Map; (2) a Plan of
the Grounds; (3) a short description of some of the principal
animals in the Collection, with 52 Photographic Illustrations
and Index. Price 1s. in Stiff Paper Cover, postage 2d.
THE HOUSE-FLY CAMPAIGN. 3rd Edn. Illustrated. Svo.
1916. 3d.
PRACTICAL ADVICE ON THE FLY QUESTION. 8vo. 1915.
2d.
ZOOLOGICAL RECORD.—Vol. 56, containing literature relating
chiefly to the year 1919, was published in November 1921.
Vol. 57, for the year 1920, is being prepared as usual.
P. CHALMERS MITCHELL,
Secretary.
Regent’s Park, London, N.W. 8.
June, 1922.
These publications may be obtained at the Soctzry’s OFFICE
or through any bookseller.
PAPERS.
Page
1. The Fauna of Hast Africa and its Future. By C. W. Wostuy, C.M.G., C.M.Z.S. .... 1
2. On the Temporal Arelres of the Reptilix. By R. Broom, D.Sc, F.R.S., C.M.ZS.
(Rext=figrames labora warehn ahbatareletetcrate sf sa ctna me celts aa ena alens 5 Hho Udine onitiba cuemriG d winisg a eet
3. Animal Communities in the Southern North Sea... By H. Birevap, Dr.Phil. (Text-
LERMAN Qieioeciew ei Pe Meee c tus aia pagerenalme al eeacaiaticc Pateiane Creuse widens aaa MCUs rors 27
4. Evolution within the Genus Dendronephthya (Spongodes) (Aleyonaria), with descrip-
tions of a number of Species. By W. Rat Sterrives, M.A., D.Se., F.L.S.
(Plates I-III. ; Text-figures 1-30.) ; 33
5. The Life-history of the Water-Beetle Pelibiws tardus Werbst. By Frayk Barrour-
BNO TO IN. NIRS IL. VAS (lead Wy ooecssugodeodbadbenccmmuc Pociers sues
6. On the Vagus and Sympathetic Nerves of the Edentata. By Crarizs F. Sonyrc,
MED ES ZAKS. = (extieiaumpeas =o.) “Scoccocenaccsc edocs Shas a Nal EMME het dS ona a 99
7. Contributions to the Morphology, Classifiextion, and Zoogeography of Indian
Oligocheta. By J. Srurirensoy, M.B., D.Sc., F.Z.8. (Plate I.; Text-figures 1-19.) 109
8. On the Vagus and Sympathetic Nerves of Hyrax capensis. By Cuaruus F, Sonntaa,
oe E.ZS. (CRON Nhe: Gach) i goiqalde-vocoreey Gane onbbtC So Ono Sess too 6.0 Hews 149
9, The Cichlid Fishes of Lake Victoria. By C. Tare Rueay, F.R.S., F.Z.8. (Plates 1-1V.;
~ Text-figures es —14.) Aico omar nae lavablc: aitaiarioredikepelaweielmiat Greek: ocay a: erick they aceatare eh aerate Bode. 1 dltaiey
10. Direct Development in a Dromiid Crab. By Srepmmn K. Monraomery, B.A., B.Sc.
(Rextieuires Mees) iisbel e's se Sc trele sa «ele Oianclsesanehens spsteiay letrots eietaiolenets Htnpogowcage «Ok
11. Notes on the Life-history of Cucuwlus eanorus, with exhibition of Eggs. By A. H.
LO
LIST OF PLATES. Sa
1922, Parr I. (pp. 1-203).
.
Page
W. Rae Suerrirrs: Pl. I. Dendronephthya clavaia Kikenthal .... 33
Pl. IL. Dendronephthya gigantea Vervill ...... 33
Pl. III. Dendronephthya annectens, sp.n. ..+++. 33 J
F. Batrour-Browye: Pls, I-III. Life-history of the Water-Beetle
Pelobius tardus Herbst. ..22.0-eeessseeses Fo 6
J. STEPHENSON : Pl. I. Morphology of Indian. Olsonen «app si LOSS
C. Tats Ruean: Pl. IT. MHaplochromis ultigents .....eceeeee Ga Alsi
Pl. If. Haplochromis serranoides ...0..+0..0% 1S 8
Pl. III. (1) Haplochromis dentex, (2) H.macro-
GNUERUS oo cca ve cene ce cacesnes 157
Pl. TV. (1) Haplochromis gr caneniite (2) Ei
ongirostvis ...see Sacicicesctes ee 157
NOTICE.
The ‘ Proceedings’ for the year are issued in for parts, prged consecutively,
so that the complete reference is now P. Z. S. 1921, p.... ‘The Distribution
is usually as follows :—
Part J. issued in March.
sneasit) bit e June.
pel Gl Gettin September.
spoon Vis ms December,
‘Proceedings,’ 1921, Part LV. (pp. 647-887), was published on January 27th,
922. ;
@
The Abstracts of the ‘ Proceedings,’ Nos. 223-225, are ;
contained in this Part. ‘
PhO
The dates of Publication of ‘ Proceedings’ 1830-1858 will be found in the
« Proceedings’ for 1898, page 486. ;
The dates of Publication of ‘ Transactions’ 18838-1869 will be found in the
‘Proceedings’ for 1913, page S14.
41 +. 2 nd a
®&, .
2
j ey 3 tee
Neg
Live Music Archive
Librivox Free Audio
Metropolitan Museum
Cleveland Museum of Art
Internet Arcade
Console Living Room
Books to Borrow
Open Library
TV News
Understanding 9/11