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PROCEEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF THE
ZOOLOGICAL SOCIETY
OF LONDON.
1908, pp. 431-983.
(MAY—DECEMBER.)
ZALS SI
PRINTED FOR THE SOCIETY,
AND SOLD AT THEIR HOUSE IN HANOVER SQUARE.
LONDCN:
MESSRS. LONGMANS, GREEN, AND CO,
PATERNOSTER ROW.
1 tS) aL
OF THE
COUN Clin AN DO F isl Cis
OF THE
ZOOLOGICAL SOCIETY OF LONDON.
1908.
COUNCIL.
His Grace THe Duke or Beprorp, K.G., President.
Grorce A. BovuLencer, Esq.,
FE.R.S., Vice-President.
Prof. J. Rose Braprorp, M.D.,
D.Sc., F.R.S., Vice-President.
ALFRED H. Cocks, Esq., M.A.
THe Rr. Hon. THe Hart oF
Cromer, P.C., G.C.B.
CHARLES DRUMMOND,
Treasurer.
Str Epwarp Duranp, Br., C.B.
FREDERICK GILLETT, Esq., Vice-
President.
fF, Du Cane Gopman, Esoq.,
D.C.L., F.R.S., Vice-Presi-
dent.
Tae Marquis or Hamrnron,
Ms
Ksq.,
K. G. B. Mnape-Watpo, Esq.
Pror. Epwarp A. MIncHIn,
M.A., Vice-President.
P. CHatmers MitcHEnu, Esq.,
MAS DSc. De aheees
Secretary.
Tue Lorp Newron.
W. R. Ocitvie-Grant, Esq.
ALBERT Pam, Hsa.
Tur Hon. N. Cuartes Rora-
SCHILD, M.A.
Davin Seru-Suira, Esq.
OLDFIELD THomas, EsqQ.,
1D RAS.
A. Trevor-Batrys, Esq., M.A.
Henry Woopwarp, Hsq., LL.D.
F.R.S., Vice-President.
PRINCIPAL OFFICERS.
P, CHAtMers MurcHett, WE Nos IDSC,
Secretary.
LL.D, RSE
Frank E. Bepparp, M.A., F.R.S., Prosector.
R. I. Pocock, F.L.S.
Henry G.
JouN Barrow, Accountant.
W. H. Coun, Chief Clerk.
, Superintendent of the Gardens.
Purr, M.R.CS., Pathologist.
F. H. Watzurnovss, Librarian.
ARTHUR THOMSON, Assistant Superintendent of the Gardens.
LIST OF CONTENTS.
1908, pp. 431-983.
May 12, 1908.
Page
The Secretary. Report on the Additions to the Society’s
Menagerie during the month of April 1908 ............... 431
Mr. W. Woodland, F.Z.8. Exhibition of, and remarks
upon, preparations of a new gland in certain Teleostean
BTSs i era eA? Yeates bag de Si wag 2 Ya wanes ot 43]
Mr. T. A. Coward, F.Z.S. Exhibition of a specimen of
Estrelata neglecta Schleg., from Tarporley in Cheshire . 433
Mr. C. Davies Sherborn, F.Z.8. Exhibition of a specimen
of chert from the Middle Culm-measures of Christow
WowareWevomshiree Leys ssc Nate cles wtiaaels s actedtslannereer 433
Mr. R. Lydekker. Exhibition of, and remarks upon, a skin
Olea Witla @att trom Sze-ChUCM Si, .aceadc.. weet «arene 433
Mr. J. T. Cunningham, M.A., F.Z.8. Abstract of a paper
entitled ‘‘ The Heredity of Secondary Sexual Characters
* in Relation to Hormones, a Contribution to the Theory
DIMAS Dyas toes see se retin. Geta one ve oe ccleten.s waisisatgienieerste icy 434
1. The Marine Fauna of Zanzibar and British Hast Africa,
from Collections made by Cyril Crossland, M.A., in the
Years 1901 and 1902.—The Caleareous Sponges. By
OTE “TSIEN Gece OB BEAR RAB RR Ore OR conri.a <utUbdncorwnnes Manet ead 434
i)
bo
co
Or
lv
Notes on the Australian Fossorial Wasps of the Family
Sphegide, with Descriptions of new Species. By Row-
LAND E. Turner, F.Z.S. (Plate XXVI.) .....------+-
May 26, 1908.
. The Rudd Exploration of 8. Africa.—X. List of Mammals
collected by Mr. Grant near Tette, Zambesia. By OLp-
FIELD Tuomas, F.R.S., F.Z.S., and R. C. WroucHton,
TEVA ie inc: Nee ee eas SON EEE iia Ga BO eee Rag GoOCD 0
Zoological Results of the Third Tanganyika Expedition,
conducted by Dr. W. A. Cunnington, 1904-1905.—
Report on the Isopoda terrestria. By the Rev. THomas
R. R. Sreppine, M.A., F.R.S., F.LS., F.Z.S. (Plate
DOG Yl ke) ss eR BE REE pr Asee ere BBA auninnatrcAee oon cudcin0
. On the Anatomy of Antechinomys and some other Mar-
supials, with special reference to the Intestinal Tract
and Mesenteries of these and other Mammals. By
BRAN TEE EO EDDARDS ViicAte (HE Sos Ano 50 re eee eee
. The Armour of the Extinct Reptiles of the Genus Pareia-
saurus. By H. G. Sernny, F.R.S., F.Z.8., King’s
College, London
Cee eee ee eee eee te twee eer eee e emer eseseesesesseeee
. Additional Evidence as to the Dentition and Structure of
the Skull in the South African Fossil Reptile Genus
Diademodon. By H. G. Surrey, F.R.S., F.Z.S., King’s
College, London
Sisisiele/stelvisysls)s\e/a2s\els|sivlelols)sie/elelelslalsinicjslsistersleietalslateterd
. New Siphonaptera. By the Hon. N. Cuartes Rors-
SCHILD, M.A., F.Z.S. (Plates XX VITI.-XXXI_)
June 16, 1908.
The Secretary. Report on the Additions to the Society’s
Menagerie during the month of May 1908
Dr. A. Smith Woodward, F.R.8., F.Z.8. Exhibition of
photographs and fragments of skin and bone of a
Mammoth and a Rhinoceros from Starunia, Galicia
Page-
AD7
5D4
561
611
629
... 630
Vv
Dr. C. I. Forsyth Major, F.R.S., F.Z.S. Exhibition of, and
remarks upon, the lower jaw of a young Canadian
BEAMCR Mee eater todas. Lith Et. CO Teetnh Ae Dae eR
Dr. C. I. Forsyth Major, F.R.S., F.Z.8. Exhibition of draw-
ings of, and remarks upon, some remains of two species
of Castor from the East Runton Forest-bed ...............
Dr. C. I. Forsyth Major, F.R.S., F.Z.8. Exhibition of photo-
graphs of Phocene Bovine in the Florence Museum
1. The Duke of Bedford’s Zoological Exploration in Eastern
Asia.—X. List of Mammals from the Provinces of
Chih-li and Shan-si, N. China. By Oxprreitp THomas,
sbos Ses0 Eee Shy (bm laGe: SONOS ie eae i eadacee snceher soeeen
bo
. Ona Case of Impertect Development in Hehinus escu-
lentus. By James Rrrconiz, M.A., B.Sc., the Royal
Scottish Museum, and D. C. McInrosu, M.A., B.Sc.,
SEs ee Sys vac (Eslatties SMONO CUTE ica tna Aiea. oe cetera) oa
3. Observations on the Minute Structure of the Spicules of
Calcareous Sponges. By H. A. Mincutn, M.A., V.P.Z.S.,
Professor of Protozoology, University of London, and
D. J. Rew, M.B., C.M., F.Z.S. (Plates XXXIV.—
DE CSOT EL ot crantesSte cies Armee ne ee eee aa at em
4, Two New Genera (anda New Species) of Indian Lycenids.
By T. A. Coapman, M.D., F.Z.S. (Plate XX XVIII.) .
ox
. A Contribution to the Knowledge of the Batrachian
Rhinoderma darwint. By Frank E. Bepparp, M.A.,
Esa Ske al Sle ot Me Ree pie tM oe OR ae
6. Some Notes upon the Anatomy of Chiromys madagas-
cariensis, with references to other Lemurs. By Franx
He repppanos MBA. BRS WAS: Wo.) Ste a
7. Leucocytozoon musculi, sp. n., a Parasitic Protozoén from
the Blood of White Mice. By Anniz Porter, B.Sc.
Lond., Zoological Research Laboratory, University
Wollece. bondonn), “(Plate XX XIX.) eerie.
8. Deseriptions of African Micro-Lepidoptera. By E.
JVTTERY SCZ A TB fe eal eS] seal ley Sah ea Se) ea a
Page
630
. 635
646
661
676
678
694
v1
Page:
_ On Collections of the Cape Verde Islands Fauna made by
: Bie Crossland, A (ania. B.Sc. (Lond.), BLS.
(late of the Gatty Marine Laboratory, St. Andrews
University), from July to September US Vie= = line
Caleareous Sponges. By A. G. THACKER, fee
(Lond.), Research Scholar in Zoology at the Roya ia
College of Science. (Plate XLL.).........::::----:sre 5
November 3, 1908.
The Secretary. Report on the Additions to the Society’s
Menagerie during the months of June, July, August, 2
and September, 1908 ........-..--.---..50+++ De ee See 5 cs 783
Prof. E. A. Minchin, M.A., V.P.Z:S. Exhibition of some
drawings of trypanosomes and trypanoplasms of some
fo) v Pie
BASSI OSUSIOSS yo dod cogpeanoeuassonpoooosscnacouncenansa boss he 785
The Secretary. Exhibition of a photograph of a young
Mellay amy Da pit cscs ecuesees Jugeds ocaaste dae eek eee 786.
1. Description of a new Species of Toad from Sumatra. By
Grorrrey Mrape-Watpo, B.A. (Plate XLI.) ......... 786.
+. On Mammals from Inkerman, North Queensland, presented
to the National Museum by Sir William Ingram, Bt.,
and the Hon. John Forrest. By Otprrenp THomas,
F.R.S., F.Z.8., and Guy Dotiman, B.A. (Platé XLII:). 788:
3. The Sze-chuen and Bhutan Takins. By R. LypreKKer.
(Een D.C EE) aa See ee Ee eM aR EIA ae ic 795.
4, On an Indian Dolphin and Porpoise. By R. LypEKKER.
(Plates XLIV. & XLV.)
November 17, 1908.
The Secretary. Report on the Additions to the Society’s
Menagerie during the month of October 1908
Vil
Page
My. HK. E. Austen, F.Z.S. Exhibition of living specimens of a ;
Fly, Hermetia illucens L., caught in Manchester ......... 809
Notice of the Reports on the Ruwenzori Expedition Collec-
IsLOT Seah ek WM Eel Mohd. chin. aelsers's 3 oe SBME aie s gat 809
1. Contributions to the Morphology of the Group Neritacea
of Aspidobranch Gastropods.—Part I. The Weritide.
By Prof. Gitpert C. Bourne, D.Sc., F.Z.8. (Plates
SCL AV el DBO) one ce Ban en ae Rete Teton tan a a meee? ee 810
2. An unknown Lemur from the Lushai Hills, Assam. By
N. Annanvate, D.Sc., C.M.Z.S., Superintendent, Indian
Vn enumar ers AM en a ee eel creme cen ten arhietad a sc wes hacen 888
December 15, 1908.
The Secretary. Report on the Additions to the Society’s
Menagerie during the month of November 1908 ......... 890
Mr. Frederick Gillett, V.P.Z.S. Note on a Hunting Trip
Ome eed Uiitearae Sia Tes Sak neeee eee cs. 5.0 sue aisle ot ie claves &90
Mr. R. 1. Pocock, F.L.S., F.Z.S. Exhibition of photographs
of a Sumatran Tiger, and remarks upon this animal and
upon other Tigers living in the Society’s Gardens ...... 890
1. Some Notes on the Muscular and Visceral Anatomy of the
Batrachian Genus Hemisus, with Notes on the Lymph-
Hearts of this and other Genera. By Frank E.
TS roto), INT, Aad hI ele ios al ey AS), 1 cee ee) 894
2. Description of a new Species of Lacerta from Persia. B
G. A. Boutencer, F.R.S., V.P.Z.8. (Plate LXVII.)... 934
3. Remarks on some Wart-Hog Skulls in the British
Museum. By Dr. Eryar Lonnpere, C.M.ZS. &e. ...,.. 936
4, On Two Chinese Serow-Skulls. By R. LyppkKer......... 940
vil
Page
5, Warning Coloration in the Musteline Carnivora. By R.
I. Pocock, F.L.S., F.Z.8., Superintendent of the Zoolo- .
gical oclebys" Gardens: tartan aie ays en. «oe ace eee
944
6. On a new River-Crab of the Genus Gecarcinucus from
New Guinea. By W. T. Catan, D.Sc., F.Z.S., British
Museum (Natural History). (Plate LX VIII.) ......... 960
7. The Duke of Bedford’s Zoological Exploration in Eastern
Asia.—XI. On Mammals from the’ Provinces of Shan-si
and Shen-si, Northern China. By OxprreLp THomas,
FEARS GE Zi Si ie ieealen. 2. eh one ces 0 ae er 963
ALPHABETICAL LIST
OF TIE
COME WO Rs,
With References to the several Articles contributed by each.
(1908, pp. 431-983.)
ANNANDALE, Netson, D.Sc., C.M.Z.S8., Superintendent,
Indian Museum.
An unknown Lemur from the Lushai Hills, Assam ...
Austen, Ernest H., F.Z.8.
Exhibition of living specimens of a Fly, Hermetia
diaccosmbereruciat, 1m Wiamehester, ss62.2....0:444-0.++55se- 00
BepDDARD, Frank E., M.A., F.R.S., F.Z.8., Prosector to the
Society.
Y On the Anatomy of <dAntechinomys and some other
Marsupials, with special reference to the Intestinal Tract
and Mesenteries of these and other Mammals ............
A Contribution to the Knowledge of the Batrachian
DR UOUCTATUGN COMPU UIDO, sors ao ioe oss es 1 sa Sis s ager oleet emia
VY Some Notes upon the Anatomy of Chiromys madagas-
cariensis, with reference to other Lemurs ..................
Page
888
561
x
Page
BEDDARD, Frank E. (Continwed.)
Some Notes on the Muscular and Visceral Anatomy of
the Batrachian Genus Hemisus, with Notes on the
Lymph-Hearts of this and other Genera.............-.-.++-- 894
Boutencer, Grorce ALBert, F.R.S., V.P.Z.8.
Description of a new Species of Lacerta from Persia.
(leit e MAX AINE Weenie oe a: saved. cde Sa eae arate eee 934
Bourne, Prof. Gitpert C., D.Sc., F.Z.8.
Contributions to the Morphology of the Group
Neritacea of Aspidobranch Gastropods.—Part I. The
mvenitid oasis ((Plaibes eXeI Vil Tak Vel) see ete eee eee 810
Cauman, WiuuiamM T., D.Sc., F.Z.8., British Museum
(Natural History).
On a new River-Crab of the Genus Gecarcinucus from
NewaG umes: ) (Plate ix Vell) 2. nberseee ee eae eee 960;
CHapman, Tuomas A., M.D., F.Z.8.
Two New Genera (and a New Species) of Indian
iliyceemid s(t e XeXOX VALI |i2)a se oye oe 676:
Cowarb, Tuomas ALFRED, F.Z.S.
Exhibition of a specimen of Wstrelata neglecta Schleg.
mom elar non eyaual Oheshiceueee asa) ae i eee 433,
CunnincHAM, JoszrH T., M.A., F.Z.S.
Abstract of a paper entitled “The Heredity of
Secondary Sexual Characters in Relation to Hormones, a
Contribution to the ilteony oiellereditiy Jae cans eee A434
Dotiman, Guy, B.A., and Tuomas, Ouprrecp, F.R.S., F.Z.S.
On Mammals from Inkerman, North Queensland,
presented to the National Museum by Sir William
Ingram, Bt., and the Hon. John Forrest, (Plate XLIL.) 788
; Page
GILLert, FREDERICK, V.P.Z.S.
Note on a Hunting Trip to the Thian Shan ............ 890
JENKIN, C. F.
The Marine Fauna of Zanzibar and British East
Africa, from Collections made by Cyril Crossland, M.A.,
in the Years 1901 and 1902.—The Calcarecus Sponges... 434.
’ Lénnpere, Dr. Exar, C.M.ZS. ce.
Remarks on some Wart-Hog Skulls in the British
TTR STF Ean O° SOR ae SUSE ees eee Renee ona hk re Serch ce ee oe eee 956.
LyDEKKER, RICHARD.
Exhibition of, and remarks upon, a skin of a Wild Cat
EROUMBS ZS. CHUCIIN pesca cre Nace ass eMac leseicin'sisle cceteigsetaielein s niaioceles 433.
” The Sze-chuen and Bhutan Takins. (Plate XLIIT.)... 795
¥ On an Indian Dolphin and Porpoise. (Plates X LIV.
Ay DECILNT 3) 0g helco Ba aac TORR C nats SeRtC NEARS glen See Ronee ee 802.
¥ On UworChinese Serow-=Skulls Hes. es... 940
McInrosu, D. C., M.A., B.Sc., F.R.S.E., and Ruircuiz,
James, M.A., B.Sc., The Royal Scottish Museum.
On a Case of Imperfect Development of Hchinus
CSCUNCHULUS:) (Es ATEN ONC TI P2Fe ia Fee oie ne an 2 duis 646.
* Masor, Dr. C. I. Forsyru, F.R.S., F.Z.S.
* Exhibition of, and remarks upon, the lower jaw of a
poume, Canadian Beavercs........$.0c2-.. 2... Je sseeee evecneeeent 630°
Exhibition of drawings of, and remarks upon, some
remains of two species of Castor from the Kast Runton
TNGIASSES) OPER aa ane eer not 1S 8 On Men DMR PELE na ton Anant ty 630:
Exhibition of photographs of Phocene Bovine in
tnemEornemeer MAIS UIN jbtysxh crak sores ae ea ee is Seis 635.
xu
Page
Muapre-WAtbo, Grorrrey, B.A.
Description of a new Species of Toad from Sumatra.
(Blois XCLL) onarecomcczeacce ceo tac ncemmmgara gee ees: 786
Meyrick, Epwarp, B.A., F.R.S., F.Z.8.
Descriptions of African Micro-Lepidoptera............--- 716
Mixcuty, Prof. Epwarp A., M.A., V.P.Z.8.
Exhibition of drawings of trypanosomes and trypano-
plasms of some freshwater fishes ...........-+::01esssesereeee 785
Mincury, Prof. Epwarp A., M.A., V.P.Z.S., and Ren,
Duncan J., M.B., C.M., F.Z.8.
Observations on Minute Structure of the Spicules of
Caleareous Sponges. (Plates MX XIV =X XM VILL) ee 661
Mircuent, P. Caatmers, M.A., D.Sc., LL.D., F.RS.,
Secretary to the Society.
Report on the Additions to the Society’s Menagerie
durins the monthyot April 90821. ca saas-e cee eee 431
Report on the Additions to the Society’s Menagerie
dun ssthe months or May, W008 enn. enero eee 629
Report on the Additions to the Society’s Menagerie
during the months of June, July, August, and September
1908
Exhibition of a photograph of a young Malayan Tapir. 786
Report on the Additions to the Society’s Menagerie
during the month of October 1908 ...2.....2.2cecheseeseseeee 809
teport on the Additions to the Society’s Menagerie
during the month of November 1908
xill
Pocock, Rueinaw I., F.L.8., F.Z.8., Superintendent of the
Gardens.
‘Exhibition of photographs of a Sumatran Tiger, and
remarks upon this animal and upon other Tigers living
inh thersieerenysteerdemst 0.04. 0). Bie... Ne eee
‘ Warning Coloration in the Musteline Carnivora ......
Porter, AnnizE, B.Sc.Lond., Zoological Research Laboratory,
University College, London.
Leucocytozoon musculi, sp. n., a Parasitic Protozoén
from the Blood of White Mice. (Plate XX XIX.) ......
Rem, Duncan J.. M.B., C.M., F.Z.S., and Muncuin,
Prof. Epwarp A., M.A., V.P.Z.S.
Observations on the Minute Structure of the Spicules .
of Calcareous Sponges. (Plates XXXIV.-XXXVIL.)...
Rircuiz, James, M.A., B.Sc., The Royal Scottish Museum, |
and McInrosu, D. C., M.A., B.Sc., F.R.S.E.
On a Case of Imperfect Development in Hchinus escu-
Weiicismare Ge labor XOXCNGW TNs sos eae aes oes oSNswu ake ceSainsode <5 c=
Roruscuitp, The Hen. N. Cuaruzs, M.A., F.Z.S.
New Siphonaptera. (Plates XX VIII.-XXXI.) ......
SEELEY, Prof. Harry G., F.R.S., F.Z.8., King’s College,
London.
The Armour of the Extinct Reptiles of the Genus
eGo MiG Same ne se rnrNer tae att ae shove sont/au cy seennetntes he
Additional Evidence as to the Dentition and Structure
of the Skull in the South African Fcssil Reptile Genus
ID GUT DECT AEE SESE Ee A Ae Sele eck elena
Page-
890.
944
703.
661
646.
X1V
‘Suerporn, C. Davigs, F.Z.8.
Exhibition of a specimen of chert from the Middle
Gulm-measures of Christow Down, Devonshire .......-.-.-
Srepprne, The Rev. Toomas R. R., M.A, ERS ahaa Ss
¥.ZS.
Zoological Results of the Third Tanganyika Expedition,
conducted by Dr. W. A. Cunnington, 1904-1905.—
Report on the Isopoda terrestria. (Plate XXVII.)......
Tuacker, A. G., A.R.C.S.(Lond.), Research Scholar in
Zoology at the Royal College of Science.
On Collections of the Cape Verde Islands Fauna made
by Cyril Crossland, M.A. (Cantab.), B.Se.(Lond.), F.Z.8.
(late of the Gatty Marine Laboratory, St. Andrews Uni-
The Calcareous
versity), from July to September 1904.
Sponges. (Plate XL.)
i ii i i i iC ii i i iii i iri eric nna)
Tuomas, OLDFIELD, F.R.S., F.Z.8.
The Duke of Bedford’s Zoological Exploration in
Kastern Asia.
X. List of Mammals from the Provinces
of Chih-li and Shan-si, N. China. (Plate XXXII.)......
Vv The Duke of Bedford’s Zoological Exploration in
HKastern Asia.—XI. On Mammals from the Provinces
of Shan-si and Shen-si, Northern China
eee eee meres eves esevns
Tuomas, OnpFIeLD, F.R.S., F.Z.8., and Dotimay, IUY,
B.A.
\
On Mammals from Inkerman, North Queensland,
presented to the National Museum by Sir William
Ingram, Bt.,and the Hon. John Forrest. (Plate X LIT.)
rT ) f af t KB ; |
Pi0MAS, OLDETELD, F.R.S., F.Z.S., and Wrovueuton, Roper
Cy Lag
The Rudd Exploration of 8. Africa.—X. List of
Mammals collected by Mr. Grant near Tette, Zambesia .
Page
433
DD4
635
963
535
XV
’ Page
Turner, Rownann E., F.Z.8.
Notes on the Australian Fossorial Wasps of the
Family Sphegide, with Descriptions of new Species.
(CEs ibe NORGE) etre ce tle esac endo pneu evs a ata de ease gee ADT
Watpo, G. Mzuape-. See MeapE-WALDo, G.
WoopiLaANn, WiLuiAM, F.Z.S8.
Exhibition of, and remarks upon, preparations of a new
alamd wa, certam Leleostean Wishes) 2. 2.........:0..cceeeseuss 431
Woopwarp, ArtHuR SuitH, LL.D., F.R.S., F.Z.8.
Exhibition of photographs and fragments of skin and
bone of a Mammoth and a Rhinoceros from Starunia,
(Cin MCLED A ee CEN, SN Se tel ear 630
WrovucHton, Ropert C., F.Z.8., and THomas, OLpDFrreLp,
E.R.S., F.Z.8.
The Rudd Exploration of §$. Africa.—xX. List of
Mammals collected by Mr. Grant near Tette, Zambesia . 535
LIST OF PLATES.
1908, pp. 431-983.
Plate Page
OXI or Au strains pieaides 2a ot ooh, vgn sos solos 457
XXVII. A. Anchiphiloscia karonge, sp. n. ‘B. A, cunningtoni,
sp.n. C, Periseyphis convexus (Budde-Lund) .... 554
XXVIII. }
XXIX,
XXX. New Siphonap beta. soe temre ate ee ccsiy we tad as: 617
XXXL J
ONONG EC opredius Oedfondd. Oe shes see ome o sc nend ein: 635
DOXOMPU. Abnormal Bekins esculentus 00.0 d pec e hese ss 646
XXXIV. 1
ae rSpicules of Calcareous Sponges. 225.4455 .46 1--.- 661
XKXVILJ
XXXVIII. eee appendages of: 1. Bothria chennellit, x 45;
i a Notarthrimus buighani, 40 ove nce sc es ee 676
XXXIX. pao UEOROORRINUSCUUDN ma ey Gis a, pce ess 6 de ae es 703
Mle CapesVerde islands Caleared V5 055 cc te... es oe wots 757
XLI. Bufo valhalle, sp. nov. 1. Upper surface of head.
De Wmidersides Ol fOOtm saat css clsgs. s0's + es eee oe as 786
OWL SVG CropusiuMlaVOlUs MOTOR oo ve ued ve eosin 788
XLII. The Chinese Takin (Budorcas tibetanus), 1.3,2.2. 795
XLIV. 1. Tursiops dawsont. 2. Neophocena sp. .......... | 802
PROG iod SOLANGE TLEGUOSMee. Ger co ok oa is vz ome teaaenn eae (ear
POU Mee SCLCELG GOT UOMICO® a isin vss: sia = aa 4 Mie whale alesis, overeat =
XLVII. Ccelom and oviduco-ccelomic funnel of Septaria ....
XLVIII. : ve 2 scams
XLLX. Genital complex of Septaria bougammvilleti 9 ........ (810
L. Female ducts and osphradium of Septaria ........
Witaekidney of Septaria depiessd. wesw ew rede ds eee - |
Mee Gull-lamelleseteiof Septaria, svev. disci. eee ni cw J
Proc. Zoou. Soc.—1908. b
XVIll
Plate Page
LIT. Histology of genital ducts of Septaria......++....+: )
LIV. Anatomy of Nerita, Paranerita, and Neritina ...... |
LV. Alimentary tract and nervous system of Paranerita. . |
LVI. Anatomy and Histology of Neritide .............- |
ILI, JARRE GOLUES ~oo amo acovooooondnnDOOKeOOOOCS
LVIIiI.| P 5 ;
LIX. { aranerita vartegata and gagates..........e scenes
: 810
LX. Male ducts of Nerita and Paranerita ............++
ID, Wine GhiGis CE IRURAIGRIEG Oboonacsepbadononen todo:
LXII. Anatomy and Histology of Neritiede ...........5-. |
LXIII. Reproductive organs of ParanerttaQ ..........+-
MIs Hemalevorganslof Ean onenia c\.w. «lr acer el iereeete |
a
ee Reproductive organs of Nerita plicata Q .......... i)
LWA, LGMearses COUMPOGUSIGP “Sac udocvuodxcoodeodoonaboGK0$ 954
LXV, GIRTON (MRED. aodacoonsaboaonoasoondaane 960
a
LIST OF TEX'T-FIGURES.
1908, pp. 431-983.
79. Diagrain of the construction of the gas-oland rete mirabile .. ye
80, Cells of the new gland situated round and in contact with
2 WED RAE SBOE AANA RS Sie STORE en ere en 432
SMMC LUMMUTLNAUNUUNUD enc Nt wha oie et MLteak eeu Pile Ginn ane Et ok ae can, 436
82, Clatlaina darwin, spicules oo... eee cee ees es ne 436
SRR UIUD: COMULORUG «ik strutece steie os SSR hecinee Wiehe ass é xe kw Ss 437
84. Clathrina contorta var. spinosa, spicules .................. 438
2D Coho OUGH ALOT UL OUI ELAT Ws OR eer ota CR EaP DA ei =A a a 459
COLAO OLANGALSPICUICS 5. seer nai ais yall ata. «sect oie cs. 440
MeN UICOSGICHICL UNL ECQLULUTIS'. = nivd cia siacieate erases) setae oss, soo 44]
89. Leucosolenia wrregularis, skeleton of dermis ................ 44]
OWneieucocolenianncquidmis, SMICUIES| &2 64.5 ye. se ete tensa. . 4492
MIB OS COU, MOVIE, SIV. Ws SPICULES > (a-actss aicpe pes ap ceo eae + cw wae 443
PICCOLO TA y MMOS SPUCULCS ac Fre irene Sete ayrc aghivgs 6 oat nk me 445
Sen Gr ORULCS SCUUSUNUILCES S Deplcge ay elo cust Crate tues fycia t cae asy cin ke & Whe 446
94, Girantessa simplex, skeleton of body-wall .................. 447
OjGnannessm sumpler. SkelevOM OL GEYMIIS . cts ceases setae A447
96, Girantessa simplex, sastral skeleton 00... 0. ee cee ew om 448
AG ROMECSSO SUNPICD ISPICULES ccs retin: 6 fas «60 6 ces 2 woh ee 448
MI SemM GOTLLCSSO EO Me QDONIS. SPre Maa cuctaysia) sy s\¢.r cots e vk # tees ode en ee 449
99. Grantessa zanzibaris, skeleton of body-wall ................ 450
100. Grantessa zanzibaris, skeleton of dermis .................. 451
101. Grantessa zanzibaris, gastral skeleton..................+.0. 451
OZ GROMUCSSO, 2ONMOATIS, SPICULES . 2.2... ee ens acts cues son ess ADD
103. Heteropegma nodus gordi, spicules .............0..0+.++-.. 453
NGATAMPLEUEUIARLUSINCRS?S; SPICULES). eiccse nqnt uct eu cee. oe o% 454
VOLES LOG weOsinensts, SPICULES wc. 5 js cewitae ee ice sy sc5 455
XX
Page
105. Neuration of fore wing of Stigmus queenslandensis, Tachysphex
pilosulus, Zoyphium kohli, Gorytes tcarioides, Nitela kurande,
Pison (Parapison) aberrans .. 111+ reserve reeeces er eee 458
106. 1. Harpactophilus sulcatus (head). 2. H. steindachneri Kohl:
@ OV@ADS fee ees deen eee eee oe tlhe carck Stoic Neen ONG o 2-0" 46]
107. Cerceris: pygidial area of Q. 1. C. anexpectata. 2. C. labe-
culata. 8. C. predura. 4. C. multiguttatd ...-eerreeeee- 469
108. Antenna, Maxilla and palpus, Labial palpus, and Mandible of
Zoyphium rufontgrum, So vvrvecerccaceeeccrecrrereceees 495
109. Bembex, 3: second ventral segment of B. furcata, B. tubercul-
ventris, B. littoralis, B. palmatd .......+ esse eeee cece eee 503
110. Pison (Aulacophilus) difficile. .......44+ Rh een PRE Gio 520
111. Alimentary tract of Antechinomys laniger .....+0.sss000es 562
112. Liver of Antechinomys laniger, abdominal aspect .......... 563
113. “Umbilical cord” and adjacent viscera of Phascologale
PILLCAONCLLONSIS Lote nso) sys oie nuns Sete in Tole isbore ae wich nA em LE
114. Cxcum, colic spiral, &e. of Galago garnetti ............ Be Sy ic)
115. Alimentary tract of Hyrax capensis ........... Procoanel eae 582
116. Part of alimentary tract of Scrwrus cinereus ...... Sir ciap elena 585
i bart oialimentanytrach Or Avlacodes wer ann ieairee ieee 586
118. Part of alimentary tract of Hydrocherus capybara ...... Ph lo,
119. Part of alimentary canal of Dasyprocta punctata ....... a DOS
120. Diagrams of primitive Mammalian gut.............,...... 591
il, IDieyeren Or Miami Mth ~ Soopecondoocesacotadebensan- 592
122 Wepre OF lemme eG Ghooooossnooobssosencacduccs 593
123. Disoram of Mammalian intestine ................-+..s.0, 594
124. Suprarenal veins in various Mammals..... Se Oe 0S ches a a 601
125. Dorsal armour of Pareiasauwrus steenkampensis ..........0... 606
126. Dorsal seute of Pareiasaurus steenkampensis.......... ala 608
127-129. Anterior, posterior, and lateral views of caudal ee
OLeParelasaurus Steenisampensis) yo. ss) see ee 609
a See 2 Me skull of Diademodon entomophonus ...... 615
Q 3 ‘ of right lower jaw of Castor canadensis ........ 630
182. Castor plicidens Maj. East Runton Forest-bed.
of right lower molar series
ae Castor sp. East Runton Forest-bed. Upper view of left m, . 631
134, Castor fiber L. Peat deposit, England. Upper view of ri ‘nt
lower true molars ;
Upper view
135. Castor fiber L. Lower view of right upper se santien mee: an
ae Castor Jiber L. Lower view of right upper true molars . . y. 633
a : es Fae Side view of anterior portion of skull .... 634
He : Sher in Ambulacral Areas of Echinus esculentus 650
- 4 He pnormalities in Ambulacral Areas of Echinus esculentus . 651
0, 141. Abnormalities in Ambulacral Areas of Echinus esculentus, 652
nee = OE
Xxi
Page
148.. Alimentary tract of Rhinoderma darwini partly opened to
Shows Lold spore kimin ome my PTATS: 5). s: 5: 2h0seacerovenoh a nave dea eet 680
144. Alimentary tract of Rhinoderma darwini, to illustrate the
shortness of that of the male and the greater leneth of that
Ol Mepemalempn eka Gh oa so Se ea Mae os on ooo es 681
145. A portion of the musculature of the ventral surface of
SPU ULOM CRULOMLAMCUELID ogee AS GN torch ee etch ech io) hc oe Le, 685
146. Musculature of floor of mouth of Rkinoderma darwint........ 686
147. Dorsal musculature of Rhinoderma darwint ...........0405. 689
148. Thigh-muscles of Rhinoderma darwini exposed from the inside. 691
149, Thigh-muscles of Rhinoderma darwint exposed from the
OUUSIOC Meet a Asda Nite edad a ao Syme na kas oy ROD MES: 692
150. A portion of the intestinal tract of Chiromys................ 695
151. A portion of the intestinal tract of Lemur rufifrons.......... 697
152. The commencement of the aorta of Chiromys cut open to
show origin of intercostals. Azygos of Chiromys.......... 699
153. Postcaval vein of male and female Chiromys................ 701
154) Living Leucocytozodn muscult 0.50000 ec el eee acne 708
loos spicules) trom Leucosolenta panis yen)... ae 759
156. Spicules from Leucosolenta atlantica 1.0.0.6. cc ee ee 761
157-160. Spicules from four specimens of Leucosolenia canariensis,
showing variation in the size of the spicules in different
SIEGE TIS a MOEN. th cheers! ASIN. cE RNIIRINED, OPEL LLC RONN Paty 765
Holerspicules trom sSycon.camimatum: 1... cee es oes hs wae he. 769
162 opicules from Grantiaintermedia yee ee ee de. 2 771
iGs, Spicules from. Leweandra.verdensis. ....202510000.15 00000 ess 772
G4 spicules from: Leucandra rudujera ees. th ee, 774.
165. Spicules from Leucandra crosslandi ........ boo. 50. 778
166. Spicules from Leucandra gemimipara ..... 0. cc cee 780
NGA Mouse Malavane la pire a.£ yesh oR OW OOo Baloo 786
168. Imperfect skull and horns of adult male of the Mishmi
alsin nye eter iy, ee ee ee ke oe ol. oy wees 796
169. Skull and horns of old male of the Sze- oieen ALO ci memes 798
170. Skull and horns of old male of the Bhutan Takin .......... 799
ie skulllos very, youne Sze-chuen Takin ..5.5.......<.:0.e.0- 801
172. A dissection of Nerita plexa, showing the course and distri-
bution of the visceral nerves and the visceral, branchial, and
osphradial ganglia: semidiagrammatic .................. 855
178, An unknown Lemur from Assam ............. a MRE estes 888
Poe SO GL MeN is teehee sdk a bedded de od nab Aue ene BAN: 891
Em ep ULES OMAN OTS TI eae nies s,'ote «soe as's.'s) 2 6 Sane eae tte aed nara. 892
176. Ventral musculature of Hemisus guttatum ............44.. 899
177. Ventral musculature of Hemisus guttatum ............0... 901
178, Ventral musculature of neck-region in Hemisus guttatum .... 906
179. Hyoid of Hemisus guttatum. A. Ventral surface partly cleared
of muscle. B, Longitudinai section
XXil
Page
180. Hyoid of Hemisus guttatum and its musculature. Ventral
PRHIGOO Mo odopeneesocoooe noon Hooo dno hombeaccgocc.0 0.0 ¢ 909
181. Alimentary tract of Hemisus guttatum, opened to show the
varying characters of the lining membrane in different
TMERMONS gun mead se se nnnd goo or eos ero OmmmODNOA BORO: 914
182. Dorsal view of Hemisus guttatum, with the skin of the back
partly removed .....+..-eceee scent tees nett eee ee tees 917
183. Saccus iliacus of Hemisus guttatwm «1... 0. renee reese reees 919
184, Saccus iliacus of Hemisus guttatum ..- +... eee cece e ee eee 920
185. Saccus iliacus of Rana guppyt opened ....-- cece ee eee e ees 921
186. Lymph-heart (posterior) of Hemisus guttatum and associated
SAGER “Go og nee od me DOO REO GD OOM oc eccoOrc C6568 925
187. Hinder part of body of one side of Xenopus levis .......... 924
188. Hinder part of body of another individual of Xenopus levis .. 925
189. Iliac fat-body of Xenopus levis dissected to show three
IN piN IEA “ho aos ese op onbOesSoooOCacoDdeaDeEnoasl 5 927
190. Iliac fat-body of Xenopus levis, cut longitudinally into two
halves to show cavities of lymph-hearts.................. 928
191. Female skulls of Nemorhedus sumatrensis milne-edwardsi and
IN MGR GYOCK@ECS, TEOMN S26 -ClUeT) 2) 41ehisey ile eet ate 941
192. A female of the Sze-chuen race of the Serow, Nemorhadus
SUMGAUTENSIS) MUNC-COWANOSY a xra)iele) «shee ells) cles elie life et ela 942
193. Cape Zorilla (Ictonyx capensis) and Cape Weasel (Pecilogale
CUDUNUCH ON Hale ees oR a a ¥ 6 Gl ONSEN Oot 947
194. Teledu (Mydaus javanensis) and Ferret-Badger (Helictis
ORUETUAUS) 0 Teva sracaiaiere ts a scsetehe a < covey ata vtelitc as eset, See 949
195. White-backed Ratel (Mellivora ratel) .......00.... ce eeeee 950
196. Grison (Grison furav) and Patagonian Weasel (Lyncodon
A WPALAGONICUS)) ” Nialiince k assis eae tiahaur lace ecuenee eso ee 952
197. Bush-Dog (Speothos venaticus) and Tayra (Galera barbara) .. 954
93 Hinelish- Bad oers(Melestneles)) aes sates eee ee eee 956
LIST OF NEW GENERIC
TERMS
PROPOSED IN THE PRESENT VOLUME (pp. 431-983).
Page
Anchiphiloscia (Crustacea) ... 555
Bothria (Lepidopt.) .......0..0 677
Dragmatucha (Lepidopt.)...... 726
Ephysteris (Lepidopt.) ......... 724
Epiphractis (Lepidopt.) ...... 732
Eporycta (Lepidopt.) ......... 728
Hyoprora (Lepidopt.) ......... 758
Page
Hyostola (Lepidopt.)............ 730
Triothyrsa (Lepidopt.) ......... 736
Malacyntis (Lepidopt.) ...... 738
Notarthrinus (Lepidopt.)...... 677
Pitharcha (Lepidopt.) ......... 751
Pseudurgis (Lepidopt.) ...... 741
Thranitica (Lepidopt.) ......... 743
o
=
Beak
Acanthogenys
rufigularis, 783.
Acanthorbynchus
tenuirostris, 783.
Accipiter
melanoleucus, 809.
Acrocercops
dasmophora, 735.
Actzeon, 881.
/Bpyceros
melampus, SOI.
/ipypryimnus
rufescens, 591, 794.
Alcadia, 823.
Allactaga
mongolica, 979.
Amblysomus
corre, 5d2.
Ammopbhila |
ardens, 465, 466.
argentea, 466.
aurifera, 464, 535.
basalis, 464.
clavus, 463.
eyrensis, 460.
timpatiens, 466.
instabilis, 464, 466. |
suspiciosa, 466.
Amphibolurus
decresit, 784.
Ampulex, 457.
Amydria
leontopa, 752. |
opifica, 752. |
optania, 752.
percastis, 753.
trophias, 753.
Anchiphiloscia, gen. nov., |
5a5. |
cunningtoni, 557, 5d8,
560.
karonge, 556, 560.
-simont, 55d.
INDEX.
Antechinomys, 561-695,
697.
Antechinus, 561.
Anthropopithecus
troglodytes, 629.
Aphroceras
sericatum, 776.
Apodemus
agrarius, 642.
— coree, 642.
— pallidior, 973.
speciosus, 641, 973,
978.
— chevrieri, 642.
— draco, 642.
— peninsule, 641.
Arbacia
punctulata, 657.
Arctomys
marmotta, d85.
Arctonyx, 944.
collaris, 95d.
Argyroploce
orthacta, 718.
Avistotelia
acrena, 724.
peltosema, 724.
strota, 724.
Arvicanthis,
596, 604.
dorsalis, 545.
— calidior, 545.
pumilio, 583, 589.
Ascaltis
canariensis,
766.
compacta, 762, 763.
coriacea, 762, 766.
darwintti, 436.
Ascandra
contorta, 456.
falcata, 666, 670, 673,
676.
572, 581,
762, 763,
Ascandra
panis, 799.
Ascetia
blanca, 438.
coriacea, 762, 766.
primordialis, 436.
spinesa, +56.
Astata
migerrima, 491.
Ateles
melanochir, 577.
Aulacodus, 595, 604.
swindernianus, 586.
Aulacophilus
vespoides, 522.
¢
Bembex
atrifrons, 503.
crabrontformis, 502.
flavifrons, 505.
flavipes, 502.
Surcata, 502.
littoralis, 502, 503.
museca, 503, 504.
palmata, 503, 504.
pectinipes, 504, 505.
raptor, \02.
saussurei, 504, 505-
tridentifera, 504.
tuberculwentris,
503.
variabilis, 502.
Bettongia
ogilbyt, 601, 603.
Borkhausenia
galacteéa, 732.
Bos
indicus, 629.
Bothria, gen.
677.
chennelliz, 677, 678.
Box, 432. -
902,
noy.,
XXV1
Brachinia
musicopa,
serialis, 727.
sterictis, 727.
Bradypus, 571.
Breviceps, 679, 681, 682,
683, 684, 688-694, |
(706
894, 903, 904, 907,
909-913, 916, 917,
922° 933, 934.
verrucosus, 683, 687
916, 918.
Bucapra
daviesi, 800.
Buceulatrix
porthnis, 737.
Budoreas
sinensis, 799, 798.
taxicolor, 795, 796,
797, 800.
— mitchell, 79.
— tibetanus, 790,
796.
— whitet, 798.
tibetanus, 797, 798,
802.
Bufo
andersonii, 787.
calamita, 788, 929,
olivaceus, 787.
stomaticus, 787.
sumatranus, 787.
valhalle, 786,
788.
787,
Cacoecia
adustana, 723.
hedrastis, 722.
Callula, 911.
Canis
dingo, 789.
lupus tschiliensis, 967.
Capra
Meyaceros
Capreolus
bed fordi, 645.
capreolus, 645.
pygargus, 645,
tianshanicus, 645.
Capromys
thoracatus, 784.
783.
> 2
Capulus, 848. |
Castor, 54, |
sp., G31.
albicus, 632
halticus, 632. 634.
canadensis, 680, 699,
europaeus, 632,
fiher
65:
Gol, 683,
682,
INDEX.
Castor
galhe, 632.
plicidens,
632.
vistulanus, 632.
Casuarius
CUSANES,
Cebus, 573.
Cemoria, 874, 880.
Centetes, 592, 593, 597.
ecaudatus, 569, 570,
BYE.
Cephalophus
grimmi, dd1,
melanorheus, (85.
Ceratophrys, 898.
Ceratophyllus
andersoni, 628, 629.
anisus, 627.
argus, 627,
629.
hirundinis, 617.
imdages, 628, 629.
melis, 027.
scturorum, 627.
Cerceris
antipodes, 471.
australis, 473.
eneapecta, 469, 535. »
laheculata, 469, 470.
multiguttata, 469,
AML
predura, 472.
venusta, 473.
Cercocytonus
converus, 559.
Cercoleptes
caudivolvulus, 571.
Cercopithecus
denti, 783.
flavidus, 53
630, 631,
431.
55
pygerythrus. “537,
— rufoviridis, 537.
rufoviridis, 537.
Ceromitia
amphichroa, 755.
qlandularis, 756.
libropis, 750.
palyntis, 755.
spulodesma, Ta4.,
sporea, Td6.
strathnodes, T54.
tyrochtora, 756.
Cervus
duvaucelli,
pygargus
Cus, 645.
Cheetopsylla
globiceps, 628.
Chalinolobus
729,
gouldi, 729
2
784.
mantschuri-
Cheirogaleus, 696.
coquereli, 17
Chinehilla, 585.
lanigera, 699.
Chiromys
madagascariensis, 694—
702, 78d
Chodsigoa
hypstbia, 639.
Clicerephon
limbatus, 541.
Chry sothrix, aoe 598.
sciureus, O12, dT,
604.
Citellus
alaschanicus, 971.
mongolicus, 970, 971.
— wmbratus, 970.
obscurus, 971.
umbratus, 971.
Clathrina
blanca, 435, 488, 489,.
440.
clathrus, 666, 670,
671, 676.
contorta, 435, 662, 666,.
667, 671-676.
— spinosa, 437, 438
coriacea, 437, 665,
670.
darwinti, 435, 436,
437.
primordialis, 435, 436,.
440.
Climacteris
scandens, 783.
Coceuthraustes
robustus, 783.
Ccelogenys
paca, O87.
Coendu, 946.
Coesyra
balantias, 731.
Coleophora, 736.
Collyriocincla
harmonica, 783.
Columba
albiqularis,
Conchylis
africana, 722.
Conepatus, 946,
780..
eConothoa, 982.
Cosmopteryx
bactrophora, 733..
tabellaria, 733.
Crabro
bivittatus, 534.
cinctus, 531, 535..
conglohatus, 533.
doddit, 529.
hebetescens, 530.
INDEX. XXV11
‘Crabro | Dasyprocta, 583. | Elachista
mackayensis, 532. aguti, 588, 589. crocogastra, 733.
ordinarius, 532. | punctata. 588, 589. Enarmonia
palitans, 534. Dasypus, 570, 592, 593. batrachopa, 718.
perlucidus, 529, 535, vellerosus, 591, 604. Engystoma
proscpoides, 528. | Dasyurus carolinense, 911,
tricolor, 524. | hallucatus, 794. Ephysteris, gen. nov.,
(Rhopalum) agilis, macrourus, 568. (2.
528. | amaculatus, 568, 569. chersea, 725
(—) conater, 526. | maugei, 569, 600, 601, | Epicephala
(—) frenchii, 526. | 602, 700. pyrrhogastra, 734.
(—) tdoneus, 527. | wrsinus, 568. Epimactis
(—) inilitaris, 523. | -viverrinus, 569, 601. | metazona, 730.
(—) tenuiventris, 524. | Dawsoniella, 812, 813, Epiphractis, gen. noyv.,
(—) transiens, 525. | 817. 732.
(—) tricolor, 524. Deianira, 812. phenicis, 732.
‘Craseomys Dendrolagus, 563. Epomophorus
regulus, 643, 644. | bennetti, 567. erypturus, 38.
rufocanus, O44, | tnustus, 567, 568. Eporyeta, gen. nov., 728.
shanseius, 643, 978. | Deshayesia, 811, S12. tarbalea, 729.
‘Cricetulus | Diademodon | Equus
andersoni, 642, 974. | brachytiara, 611. grevyi, 629, 784.
bed fordie, 974. | brownt, 611, 612, 613, hemionws, 784.
dichrootis, 648. : 614, 616, 617. Erinaceus
griseus, 642, 974. entomophonus, 615, dealbatus, 965, 966.
longicaudatus, 642, | 617. hanensis, 966.
643. mastacus, 611, 616. hughi, 966, 967.
obscurus, 643. tetragonus, 611. miodon, 965, 966.
roborovuskit, 975. Didelphys, 592, 593, orientalis, 966.
triton, 975, 974. 604. tschifuensis, 966.
— incanus, 973. virginiana, 569, 575. | Erithizon, 946.
‘Crocidosema | Diphyllodes Ethmia
plebeiana, 720. | _ hunsteini, 809. ballistis, 732.
‘Crocidnra, 541. | Diplopelma | Eucosma
core@, 639. | berdimoret. 911. leucoptera, 719.
‘Crossarchus ornatum, 911. | Hutamias
fasciatus, 542. | Dipus, 963. | astaticus, 968.
Cryptolechia, 732. | sowerbyi. 979. | — intercessor, 969.
‘Ctenophthalmus | Dorocidaris | — ordinalis, 968, 970.
agyrtes, 624. palpillata, 659, 660. —— senescens, 963, 968.
rettigt, 624, 629. | Dragmatucha, gen. nov., | Zineatus, 968.
‘Ctenopsyllus 726. | ortentalis, 968, 969,
allophylus, 625, 629. proaula, 726. 970. .
Cyaniris Dromiciops senescens, 640, 968,
chennellit, 676, 677. australis, 626. 969, 970.
678. Dysopes k uthensis, 968, 970.
‘Cyanocephalus limbatus, 541. Everes, 677.
porcarius, 576. Exeirus, 457.
Cyclophorus, 881. Kccopsis
‘Cyclorhamphus acrocosma, 717 Felis
marmoratus, 910. Echidna, 574, 575, 596. aurata, 433.
Cyclostoma, 881. acanthion, 794. caligata, 553.
‘Cydia setosa, 783. catus, 967.
isogramma, 720. | Hchinobrissus | chrysothrix, 433.
psammacta, 721. orbicularis, 657. cottont, 434.
tumulata, 720. Hehinus dominicanorum, 433.
Cynictis esculentus, 646, 650, Tco, 431.
selousi, 553. 651, 652, 635, 657, longipilis, 893.
‘Cynomys | 660, 661. ocreata, 553,
ludovicianus, 584, | melo, 657. onca, 785.
589. ' Eira, 944. pardus, 431.
XXV111
Felis
serval, O41.
sondaica, 893.
temminckt, 435.
-— dominicanoruin,
43-4.
— milchelli, 430.
tigris, 893.
— mongolicd, 893.
nigra, 891.
sondaica,
virgata, 89
virgata, 893.
Fierasfer, 432.
Fissurella, 844, 848.
Funambulus
pennantii, 713.
Funisciurus
cepapt, 543, 544.
— sindi, 543, b44.
= 785.
2
Galago, 604, 702.
cr assicaudata, 578.
garnetti, 578, 702.
granti, O37.
moholi, 537, 538, 579
mossambicus, 537.
Galera, 944.
barbara, 952,
957.
Galictis, 700.
barbara, 9)2.
vittata, 952.
Gecareinucus
ingran,
963.
Jacquemontii,
962.
Gelada
ruecppelli, 576.
Gelechia
arotrias,
trisiqnis, F20.
Genetta
Jelina, 542.
lelabe, 542.
ludia, 542.
pardina, 890.
ruliginosa, 641,
O71.
senegalensis, 42.
tigrina, 42.
zambesiana, 542.
Genioglossus, 905.
Gerbillus
namaquensis, DAS.
Glyeiphila
Julvifrons, 784.
tobius, 452.
Gomphognathus,
615,616.
954, 95d,
960; 962,
960, 961,
542,
Giz;
{
if
INDEX.
Gomphognathus
mastacus, 611.
polyphagus, O14.
Gorytes
cygnorum, 590.
decoratus, 500,
duboulayi, 496.
frenchii, 501.
icarioides, 458, 499.
lucidulus, 498, 585.
ornatus, 500.
sanguinolentis,
499, 535.
thoracicus, 498.
Grantessa
intusarticulata, 450.
simplex, 435, 446, 447,
448, 452.
497,
canzibaris, 435, 449,
440, 451, 402
Grantia
ciliata, 62.
CES Ta, eno,
771. 182.
a OSM, 757.
Graucalis
mentalis, 783.
Grison, 953.
furax, 951, 952.
Guancha
blancha, 438.
xymnodactylus
nuliust, 784.
Heematopinus
spinulosus,
eb ilGe
Heemogregarina,
joe, TUL,
703,
hal fouri. 715.
jaculi, T14.
Haliotis, 816, 827, 874
tuberculata, 883.
Hapale, 595, 598.
albicollis, 697.
penicillata, 577, 697.
Hapalemur, 696.
Hapsifera
glebata, 751.
meliceris, 750.
ochroptila, 749.
pardalea, 749.
septica, 750.
Harpactophilus, 457.
arator, 460, 461, 535.
bicolor, 459.
kohlvi, 459, 461.
steindachneri, 459, 460,
461, 462, 463.
sulcatus. 460, 461.
tricolor, 462, 535.
Hatteria, 597.
Helicina
japonica, 822, 823.
Helictis
orientalis, 949.
personata, TOU.
Helix, 815.
Hemiaster
hatnensis, 657, 658.
Hemisus, 894-934.
guttatum, 894, 899,
901, $06, 908, 909,
Gay Oia, Bus), S20)
923.
Hermetia
illucens, 809.
Herpestes
gracilis, DO.
— rufescens, 543.
ornatus, DAS.
ruddi, 552.
Heteropeema
nodus gordii, 435, 458,.
666, 671, 676.
Hippopotamus
amphibius, 551.
Hipposiderus
caffer, 53%,
Hydra, 851.
Hydrocena, 812.
Hydrocheerus, 604.
capybara, 887.
Hydromys
chrysogaster, 789, 790.
— regine, (88, 789.
Fulvolavatus, 790.
Sulvo-venter, (90.
irma, 790.
leucogaster, 790.
lutrilla, 790.
Hylobates
aguis, 629.
Hylocheerus, 940.
5o2.
Hyoprora, gen. nov.,
753.
erymodes, 754.
Hyostola, subgen. nov.,
730.
Hypsiprymnus, 593
cuniculus, 56S.
gainardt, 568.
Hyrax, 573, 574, 596,.
605.
capensis 576, 579, 581,
582, 584, 604.
Hystrix, 946.
cristata, 574, 698.
Tetonyx, 948, 951, 933.
capensis, D7 rae 947
954.
Tetonyx
libyca, 947, 956.
Indris, 579, 596, 889.
Triothyrsa, gen. nov., 736.
melanogma, 730.
Tsoodon
macrourus, T94.
obesulus, 788.
dorosus, 788, 794.
Jaculus, 979.
714.
Karyolysus,
Lacerta
chlorogaster, 934-936.
praticola, 9305.
taurica, 939.
Lagomys
ogotona, 982.
Lagopus
scoticus, 715.
Lagorchestes
conspicillatus
ardti, 793.
— pallidior. 793.
Lagostomus, 574.
trichodactylus,
585, 604.
Lama
vicugna, 809.
Larra
australis, 474.
chrysonota, 475.
crassipes, 475.
Jemorata, 475
mansucta, 473
mendax. 473.
nigripes, 473, 474.
pacificalrix, 474.
rupfipes, 473.
scelesta, 474.
Lasiopodomys, 976,
Laspeyresia
delineana,
isacma, 721.
tricentra, 721.
Lecithocera.
cholopis, 726.
Lemur, 577, 696, 697.
albifrons, 578, 697.
brunneus, 697, 698.
flavifrons, 578.
macaco, 978.
mongoz, 78.
— nigrifrons, 57
rujifrons, 697.
selatert, 578.
Lepidosiren
paradoxra, 785.
leich-
576,
Gene
“
8.
INDEX.
| Leptodactylus
| lovati, 713.
ocellatus, 713,
|
|
|
|
|
|
|
Lepus
jilchneri, 980,
980.
monticularis, 93.
sechuenensis, 980.
stegmanni, 980.
swinhoci, 644,
kozlovi,
Leucaltis
subluteus,
714.
980.
979.
floridiana, 453, 454.
solida, 664.
Leucandra
alcicornis, 664.
Dy
435,
ananas,
445,
aspera,
676,
664, 665,
444,
776, 78l.
79.
cra AT
crosslandi,
778, 781, 782.
fistulosa, 445.
gemmipara,
757,
f
780, 781, 782
panis, 758.
rudifera, 737,75
774, 782.
sericata, TdT,
776.
typica, 157, 758. 776.
— massa, 777.
— tuba, 716, TTT.
verdens
772,
(WP
Meaiviee
as,
=
(
782.
Horidiana, 435, 4
wasinensis,
AAD,
tee
Leucoeytozoon
canis,
714.
705,
LOT
funambuli, 713.
Jelis, 713.
nuris, 714.
musculi,
ranarum,
Leuconia
rudifera,
sericatum,
aon
Frias
-— MASSA, § 4
1ypica,
TT
77
o:
— tuba, 776,
THER eGnOe: 637.
Leucosole
nia
atlantica,
461, 782
be i
canariensis, 75
762;
763,
766, 782
5
(>)
776.7
777
~
(
6.
ry
64,
703-716.
71s.
a7, 77
666,
2
XX1IX
Leucosolenia
canariensis tenuipilosa,.
764, 766.
complicata, 666.
contorta, 456.
coriacea, 762, 764, 766.
— ceylonensis, 764.
wrregularis, 485, 440,
441, 442.
laxa, 761.
lieher hithnit, 666.
nansent, TAT, T62, 763,.
764, 765, 766.
panis, T57, 758, 759
eZ
papillata, 763.
tenuipilosu, TO8, 762,
764,
(Ascaltis) canariensis,.
762.
Limneecia
ichnographa, 734.
Liris
hemorrhoidatis, 473.
magnifica, 473.
Lissochilus, 811.
Littorina, 881.
Lobe-ia
acolopa, 716.
Loris, S89.
gracilis, 5T9
Lozotznia
dorsiplagana,
Lupus
tschiliensis, 967.
Lutra
vulgaris, 702.
Lycopsylla, 618.
Lyncodon
patayonicus, 052, 953:.
O92
123.
Macarostola_
onychota, 7
Maclurea, 819,
Macropus, 592, 929.
agilis, 601, 788, 793..
bennettii, 573.
billardieri, 568.
brachyurus, 56S.
dorsalis, 601, 602.
giganteus, 568, 792,
hagenbecki, 575.
melanops, 568, 575.
parryi, 568, 795.
robustus eruhescens,7 92.
ualabatus, 602, 793.
-— apicalis, 792, 793.
— ingrami, 788, 792.
woodwardi, 575.
Macroscelides
Juscus, 541.
a0),
XXX
Malacyntis, gen. nov., |
738.
stibarodes, 738.
Malurus
cyaneus, 783.
Martes ;
flavigula borealis, 967.
— huatunensis, 987.
Megalophrys, 898, 908.
nasuta, 686, 689.
Melasina
abacodes, 749.
edifica, 744.
amica, 748.
cyclatma, TAG.
dissoluta, 747.
halieutis, 743.
zmmanis, 746, T47.
inimica, 747.
isospila, 746.
liochra, 748.
morbida, T44.
mylica, 748.
paraphrictis, 744.
sauropa, 7405.
stahularia, 745.
steélitis, 743.
systolea, 747.
Meles, 944.
anakuma, 627, 628.
hanensis, 967.
leptorhynchus, 967.
meles. 9d), YDG.
siningensis, 967.
Melipbaga
noveé-hollandie, 784.
Mellivora, 949.
capensis, 950, 951.
cottoni, C57.
indica, 951.
Melursus
ursinus, 957.
Mephitis, 946.
mephitica, 947, 954.
Meriones, 963.
auceps, 640, 971, 972.
erythrurus, SAO.
leucogaster, S45.
psammophilus.
64), Wel; 972:
640,
unguiculatus, 640, 641, |
One,
Mesocricetus
newton, 625.
Metoponorthus
prumosus, 554.
Microcebus, 592, 604,
696, 701.
smithit, 577, 700.
Microtus
brandti, 976.
INDEX.
Microtus
inez, O77.
mandarinus, 976.
nelanagaster, 977.
podomys, 976.
(Kothenomys)
964, 976.
| (—) melanogaster, 976,
977.
Miniopterus, 637.
| australis, 789.
chinensis, 638.
Japonie, 638.
schreibersi, 798.
— chinensis, 638.
| — fuscus, 639.
— japonie, 638.
Miscothyris, 497, 498.
Monopis
megalodelta, 737.
Mungos
auratus, 543.
cauut, 543.
melanurus lastti, 548.
ratlamuchi, 543.
Mus
sp., 791.
arborarius, O48.
assimilis, 620, 790, 791.
auricomis, 548.
— centralis, 548.
avarillus, 547.
chrysophilus, 546, 547,
548.
— acticola, 547.
— ineptus, 546, 547.
confucianus, 641, 972.
— luticolor, 972.
— sacer, 641, 972.
coucha, 545.
— culuensis, 546.
culnorum, 790.
decumanus, 567, 589,
704, 713.
delicatulus, 791, 792.
Jforresti, 791, 792.
greyt, 790, 791.
hermannsburgensis,7 91,
792.
lehocla, 548.
microdon, )45, 546.
| mordax, 621, 622.
| musculus, 703, 792.
namaquensis, 547, 548.
| — auricomis, 548.
— centralis, 548.
— lehocla, 548.
nove-hollandie, 792.
patrius, 791.
rattus, 589, 704, 713,
| 714, 790.
inez,
Mus
sacer, 972.
terre-regie,
791.
tunneyt, 790, 791.
wagneri mongolium,
641, 972. ‘
(Leggada) gansuensis,
641.
Mydaus, 948.
javanensis, J49.
Myospalax
cansus, 978.
fontanieri, 978.
psilurus, 978.
Myotalpa
cansus, 978.
Myotis
sp. 964. ‘
mystacinus, 964.
ricketti, 637.
(Leuconoe) capaccinit,
638.
(—) pequinius, 637.
Myoxus, 572, 590.
Myrmecophaga
jubata, 570, 592, 784.
Myzantha
Hlaviguia, 784.
Nandinia
binotata, 571.
Nardoa
canariensis, 762.
rubra, 762.
spongiosa, 662.
sulphurea, 7&2.
Nasilio
brachyrhynchus, 541.
Nasua
rufa, 601, GO2.
Natica, 811.
Naticopsis,
817.
Navicella, 816, 825.
porcellana, 882.
Nemophora, 754.
Nemorhcedus
argyrochetes, 940, 941,
944.
milne-edwardsi, 943.
sumatrensis, 940, 942,
943.
— inilne-edwardsi, 941.
942, 944,
Neophocena
sp., 806, 808.
phocenotdes, 806.
Nephautis
«ystopala, 729.
Neridomus, 812.
812, 813,
Nerita, 811-818, 820,
825, 828, 835, 837,
839, 840, 8438, 845,
849, 853, 861, 862,
863, 864, 872, 873,
874, 876, 877.
gagates, 847, 864, 885,
886.
lineata, 813, &46, 850,
865, 868, 870.
longispina, 847, 885,
387.
melanotraga, 847, 850,
865, 868, 870, 871,
875, 885, 886.
ornata, 821, 822, 829,
836, 852.
peloronta, 819, 820,
833, 846, 850, 860.
pica, $22. 823, 82%,
869, 870, 871.
plexa, 846, 850, 855,
870.
plicata, 832, 847, 850,
860, 868, 870, 871,
872, 887.
polita, 820, 882.
variegata, 832, 847,
885.
Neritella
(Neritina) pudlligera,
820.
Neritina, 811, 828, 835,
839, 877, 878.
aculeata, 815.
auriculata, 814, 815.
bicanaliculata, 815.
cariosa, 819, 849.
cornea, 815.
crepidularia, 815.
dilatata, 815.
fluviatilis, 810, 814—
818. 820, 823, 825,
827, 829, 851-841,
845-848, 850, 852-
857, 860, 861, 863,
865, 867, 868, 872-
876, 882, 883, 885.
gagates, 832, 834, 885,
886.
granosa, 815.
intermedia, 815.
latissima, 815.
longis 832.
morio, 815.
owent, 820.
oweniana, 816.
peloronta, 832.
picea, 815.
picta, 814.
plexa, 852.
INDEX.
| Neritina
polita, 815.
pulligera, 820.
recluziana, 815.
subsulcata, 815.
tahitensis, 815.
waluensis, 814.
virginea, 814.
— listeri, 814.
viridis, 814, 876.
Neritodonta, 812.
| Neritoma, 812.
Neritopsis
radula, 812.
Nesotragus
livingstonianus, 551.
Nitela
kurande, 458, 508.
reticulata, 508.
Notarthrinus, gen. nov.,
Mie
binghami, 677, 678.
Notogonia
abbreviata, 481.
agitata, 477.
basilissa, 476.
chrysonota, 475.
commiata, 480.
obliquetruncata, 479.
regina, 475, 539.
retiaria, 479.
serena, 478.
Nycteris
fuliginosa, 538.
Nyeticebus, 589,
888, 889.
604,
tardigradus, 579, 702,
889.
Nyeticejus
plamrostris, 538.
viridis, 539.
Nyctinomus
nor folcensis, 789.
planiceps, 789.
Nyctotherus, 710.
Nysson
(Acanthostethus) pane-
tatissimus, 505, 535.
(—) spiniger, 507.
| Ochotona, 9638.
alpina, 982.
bedfordi, 981.
cansa, 982, 983.
dauuwrica, 981, 982.
huangensis, 981.
ladacensis, 981.
ogotona, 981.
pallasi, 982.
Proc. Zoou. Soc.—1908, No. LXIIT.
| Ornithorhynchus,
XXX1
Ochotona
pusilla, 985.
roylei, 982.
sorella, 982.
Odites
citrantha, 729.
procellosa, 729.
Odobzenus
rosmarus, 890.
Cistrelata
neglecta, 433.
Ogotoma, 982.
Onehochilus, 812.
Oniscus
muscorum, DDD.
sylvestris, 5d5.
Opogona
chlorophanes, 737.
pheochalea, 737.
596,
Orycteropus, Nis, Biseh
592, 605.
| Oryctolagus
crassicaudatus, 553.
Otolienus
mossambicus, 537.
Otomys, 553, 604.
bisulcaius, 589.
irroratus, 539.
unisulcatus, 589.
Otostoma, 811, 877.
Ovis
aries, 629.
Oxybelus, 457.
Paltodora
psacasta, 723.
Paludina, 838, 863, 873,
_ 880 881.
vivipara, 882.
| Pamplusia
|
|
sardonia, 718.
Paradisea
raggiana, 809.
| Paradisornis
rudolphi, 809.
| Paranerita, 847, 849, 853,
862, 867, 870, 871,
873, 874. 876, 878.
cornea, 877.
gagates, 850, 852, 855,
860, 863, 865, 866,
868, 869.
longispina, 852,
£68, 869.
variegata,
868, 869.
Paraphiloscia
stenosona. 556.
63
865,
852, 865,
XXXil
Pareiasaurus
baini, 605, 606, 607,
609, 610.
steenkampensis,
607, 608, 609.
Pariodontis, 618.
Parotia
lawesi, 809.
Patella, 848.
Pelea
capreolus, 431.
Pelobates, 898, 909.
punctatus, 907.
Pelodytes, 907, 909, 910,
Peragale
lagotis, 601, 602.
Perameles
nasuta, 568, 783.
torosus, 794.
Periscyphis
albescens, 559.
converus, 559, 5H0.
trivialis, 559. 560.
Perodicticus, 579, 889.
Petalia
capensis, 538.
Petaurista
leucogenys, 627.
Petaurus
breviceps, 567.
Petrogale, 563.
assimilis, 798.
penicillata, 602, 793.
Phacochcerus
eliani, 936, 937, 940.
ethiopicus, 939, 940.
606,
africanus, 936, 937,
940.
delamerei, 940.
massaicus, 937, 9038,
940.
sundevalliz, 937, 938,
940.
Phaiomys, 976.
Phalangista
vulpina, 570.
Phascolarctos
cinereus, 567, 794.
Phascologale, 561, 564.
calura, 564.
Hlavipes, 783.
macdonellensis,
566, 567, 603.
penicillata, 564, 566,
567, 601, 603.
swainsoni, 564, 624.
Phascolomys
penicillata, 566.
Philanthus, 457.
Philoscia, 554.
annulicornis, 558.
565,
INDEX.
Philoscia
muscorwunn,
558.
suarezi, 556, 557.
Phonygama
purpureo-violacea, 809.
Phryniscus, 911.
Phthorimea
operculella, 726.
Pika, 982.
Pileolus, 812.
Pipa, 895, 900, 902,
Pison, 457.
auriventre, 512.
erythrocerum, O17.
fuscipenne, 510.
icarioides, 521.
ignavum, 511.
infumatum, 510.
insulare, 510.
iridipennis, 512.
marginatum, 513.
melanocephalun,
530.
nitidum, 510.
pallidipalpe, 512.
pelletier?, O14,
priscum, 510.
punctulatum, O12.
ruficorne, H14.
scabrum, 509.
simillimum, 513.
spinole, 509.
viroswm, O15.
westwoodi, 512.
(Aulacophilus) défficile,
520.
555, 556,
515,
(—) icarioides, 521,
535.
(Parapison) aberranus,
458, 519.
(—) caliginoswn, 518.
(—) noctulum, 516.
(— » pertinax, 517.
(—) tenebrosum, 518.
Pitharcha, gen. nov., 751.
chalinea, 751.
Pitymys, 976.
Pleurotomaria, 831, 845,
874, 880.
beyrichii, 884.
Plutella, 736.
Peecilogale, 953.
albinucha, 947, 948.
Polychrosis
harmonia, 716.
scorptodes, 717.
Polyhymno
tropea, 728.
Poreellio
pr winosus, 554
Potamon
(Parathelphusa)
dentatum, 963.
(Potamonautes) cunt
cularis, 963.
Potorous
platyops, 783.
Procavia
syriaca, 784.
Procolophon, 616.
Procometis
acharma, 731.
acutipennis, 730.
oxypora, 730.
terrend, 731.
trochala, 731.
Psenulus
interstitialis, 468, 535.
lutescens, 463.
tri-
| Pseudelaps
_ Pseudochirus
diadema, (84.
peregrinus, 572, 590,
698.
Pseudurgis,
TAL.
tectonica, TAL.
Pteropus
gouldi, 789.
scapulatus, 789.
Ptilorhis
intercedens, 809.
gen. Nov.,
| Ptilothyris
purpurea, 726.
Ptilotis
chrysops, 784.
chrysotis, 784.
leucotis, 784.
penicillatus, 784.
Pulex
globiceps, 628.
melis, 627.
Pupa, 812.
Putorius
desertorum, 625.
itatsi, 628.
sarmaticus, 956.
Pygiopsylla
afer, 618, 629.
ahale, 617, 621.
colossus, 617, 619, 620,
621.
echidne. 617, 618.
ferinus, 622, 629.
gravis, 620, 629.
hilli, 617.
laciniosus, 620, 629.
mordax, 621, 629.
nove-guinee, 617.
rainbowi, 619,
629.
620,
Pygiopsylla
robinsoni, 617, 619.
torvus, 617.
woodwardi, 617.
Pyrina
ovulum, 657.
Pyxicephalus
rufescens, 911.
Rana, 679, 680, 681, 684,
686, 687, 690, 692,
693, 694, 898, 904,
905, 907, 912, 918,
916, 922, 929.
breviceps, 911.
esculenta, 682, 686,
688, 691, 895, 896,
897, 915, 919-922,
930, 931.
guppy?, 688, 691, 894—
897, 900, 902, 915,
921, 926, 930, 931,
933, 954.
rufescens, 911.
temporaria, 929.
Raphiceros
sharpei colonicus, 551.
Rappia
sp., 910
Rhinoceros
sondaicus, 5716.
Rhinoderma, 894, 899,
903, 906, 907. 912,
913, 915, 933, 934.
darwini, 678-694, 905,
916.
Rhinolophus
hildebrandti, 538.
lobatus, 538.
ferrum-equinum
pon, 637.
Rhipidura
tricolor, 785.
ip-
Saccostomus
campestris, 900.
mashone, 590.
Salamandra
maculosa, 930.
Seardia
paracosma, 753.
Sciurotamias
davidianus, 639, 967.
Sciurus
cinereus, 584, 585.
macrurus, 584.
maximus, 5T4, 584.
tennanti, 584.
vulgaris, 585.
— orientis, 628.
INDEX.
Seoteinus |
greyt, 789.
minimus, 039, S41.
schlieflent, 539, 540. |
— albiventer, 540. |
— australis, 539, 540,
541.
— bedowin, 540.
Scotophilus
borbonicus, 538.
damarensis, 539.
dingant, 538.
nigrita, 538. 539.
— dingani, 588, Dabghe
— herero, 539.
viridis, 538, 589.
— damarensis, 539.
Scutellina
cinnamomea, 822, 823.
Semnopithecus, 577.
melalophus, 576, 604.
Septaria, 812, 815, 816,
818, 820, 821, 827,
829, 831-836, 842-
848, 853, 857, 860,
861, 862, 868-877.
borbonica, 825, 841,
875, 885.
hougainville?, 816, 825,
837, 838, 841, 875,
876, 885.
depressa, 825, 838, 863,
885.
parva, 822, 823, 838,
875.
suborbicularis, 822,
838.
(Navicella) porcelland,
| Sericophorus, ADT.
bicolor, 494.
Serpentarius
reptilivorus, +31.
Siredon
pisciformis, 930.
Sminthopsis, 603.
crassicaudata, 566.
larapinta, 566.
_ Sorex
sp., 622.
Soriculus |
hypsibius, 639.
(Chodsigoa) heresowsht,
639.
| Sotalia
ferqusont, 808. |
lentiginosa, 808.
Spalax
typhlus, 625.
Spatula
rhynchotus, 784.
|
XXx1il
Spectotheres
australis, 783.
Speothos
venaticus, 954, 955.
Sphex
clavus, 468.
diabolicus, 469.
gilberti, 468.
nigella, 467.
nitidiventris, 467, 468.
obscurella, 467.
prastinius, 467.
rufipennis, 469.
ustulatus, 407.
wallacei, 467.
(Isodontia) albohirtus,
466.
(—) nigellus, 407.
(—) obscurellus, AGT.
Sphodrotes, 457.
Spilogale, 946.
Spizaéfus
bellicosus, 785.
Spongia
(Grantia) compressa,
602.
(Leuconia) nived, 662.
Stagmatophora
semnostola, 734.
Stathmopoda
xanthoplitis, 733.
Steatomys
pratensis, 551.
Steganoptycha
obscura, 720.
Steno
lentiginosus, 808.
Stenops, 572, 579.
Stephanoeircus
dasyurt, 623, 624.
jarvisi, 628, 629.
mars, 629.
simsoni, 623, 624.
Stigmus
queenslandensts, 457,
458.
Struthio
molybdophanes, 783.
Struthisea
arcata, 742.
hormotris, 742.
omichlodes, 742.
Sulcus
lateralis, 930.
Suricata, 700.
Syeandra
ainpulla, 243.
ciliata, 435, 442.
quadrangulata, 767.
(Sycon) helleri, 769,
770.
XXX1V
Sycon
ampulluin, 435, 445.
caminatum, 757, 767,
769, 770, 782.
ciliatum, 435, 442, 443,
666, 676.
munitum, 435, 448.
quadrangulatum, 757,
766, 767.
raphanus, 443.
Syconella
quadrangulata, 766.
Sycortis
quadrangulata, 766,
767.
Syngnathus, 432.
Tachyglossus
aculeatus, {9A.
Tachysphex
debilis, 484, 486, 490.
JSortior, 484, 486, 487.
hypoleius, 484, 492.
imbellis, 484, 485.
mackayensis, 484, 487.
nigerrimus, 484, 491,
492.
pacificus, 484, 491.
pilostlus, 458, 484,
488.
pugnator, 484, 491,
rufo-niger, 485.
tenuis, 484, 489.
truncatifrons, 484.
walkeri, 484, 487.
Tachytes
approxtmatus, 483.
australis, 474.
Semoratus, 473.
fervida, 483.
formosissimus,
535.
migerrimus, 491.
rubellus, 482.
482,
Tamandua, 604.
tetradactyla, 570, 592.
Tamias
striata, 585, 584.
Tarroma
canariensis, 762.
rubrum, 762.
sulphureum, 762.
Tarsius, 570, 597, 889.
Tatera, 553.
lobengule, 544.
— bechuane, 544,
— mashone, 544.
INDEX.
Taxidea, 944.
emericana, 955.
Teras
capensana, "723.
meridionana, 723.
_ reciprocana, 723.
Thamnomys
cometes, 549, 550.
ruddi, 549, 550.
rutilans, 550.
surdaster, 550.
Theodoxus, 846.
Theropitbecus
gelada, 576, 604.
Thranitica, gen. nov.,
743.
hemicopa, 743.
Thylacinus, 601, 603.
Tigris
sondaica, 891.
Tierisoma
brasiliense, 785.
Tinea
homestia, 741.
enopis, 740.
othello, 741.
paraxena, 740,
tanystis, 741.
Tineola
chloristis, 740.
holopsamma, 739.
leucastis, 738.
marcescens, 739.
melanostoma, 739.
meretrix, 739.
ochropsamm«a, 739.
phocina, 740.
xanthastis, 738.
Titiscania, 816, 820.
Yomopterna
breviceps, 911.
Tortrix
africana, 722.
agroeca, "723.
capensana, 723.
cedrota, 722.
Zocoma, 723.
Trachydomia, 812, 813,
817.
Tragulus, 701.
Trichosurus, 595, 604,
605.
vulpecula, 568, 572,
573, 575, GOL, 602,
605, 794.
Trigonoporus, 87+.
Trochus, 856. 874.
Trypanosoma
raje, T\0.
Trypoxylon, 457.
connexum, 22.
pileatum, 523.
placidum, 523.
Tursiops
abusulam, 804, 805.
catalania, 804, 805.
dawsont, 806, 80°.
fergusoni, 804. 805.
gephyreus, 805, 806.
gilli, B04, 805, 806.
parvimanus, 804.
tursio, 802, 804, 805.
Urocricetus
kamensis, 648.
Ursus
arctos, 572.
malayanus, 957.
syriacus, 571, 574.
torquatus, 957.
¢
Velates, 812.
Vespertilio
capensis, 538,
Viverra
tangalunga, 890.
Vulpes
vulpes, 639, 967.
Xenophrys, 60.
Xenopus, 894, 9O7, 922,
SAG, Sli), Beil, Osis.
934.
levis, 924,
928, 929.
Xerus
capensis, 585.
925,
Yponomeuta
glaphyropis, 736.
Zizera, 676, 677.
Zoyphium, 457.
erythrosoma. 493.
JSrontale, 496.
kohtit. 457, 495.
rufonigrum, 494, 495,
535.
Printed by Tayztor and Francis, Red Lion Court, Fleet Street.
927,
OF LONDON.
1908.
Pages 431-782.
Part III]. conTAINING PAPERS READ IN
MAY anv JUNE.
OCTOBER 1908.
Sia PRINTED FOR THE SOCIETY,
= oe ar THEIR HOUSE IN HANOVER SQUARE.
- LONDON:
es, MESSRS. LONGMANS, GREEN, AND co.,
4 PATERNOSTER ROW.
ss [Price Twelve Shillings. | ,
Peay
List OF, CONTENDS. 3
1908, pp. 431-782.
May 12, 1908.
May 12, Page
The Secretary. Report on the Additions to the Society’s Menagerie during the month of
April 1908 .....-+ he, Clee ee eR tigi SARIN 2.2 481
My. W. Woodland, F.Z.S. Exhibition of preparations of, and remarks upon, a new gland
in certain Peleostean Hishes ..- +. .+- sje: cote e cieee sc eee - > 40+ ne = -l/t ieee . 431
Mr. T. A. Coward, F.Z.8. Exhibition of a specimen of Gstrelata neglecta Schleg. from
(Olicshhire. ccs cpio vels dng sale Gob seme cone ode sips Beis giatnin eg.” nae ee 433
Mr. C. Davies Sherborn, F.Z.S. Exhibition of a specimen of chert from the Middle
Culm-measures of Christow Down .. ..--- eee cece etre reece terete ee re nee .. 483
Mr. B. Lydekker, F.R.S., F.Z.8. Exhibition of the skin of a Wild Cat from Sze-chuen ... 433
Mr. J. 1. Cunningham, M.A., F.Z.8. Abstract of a paper entitled ‘‘The Heredity of
Secondary Sexual Characters in Belvtion to Hormones, a Contribution to the Theory of
Heredity? (i. 2 ese ede lucene coil ce ered cee hiss dla scjr ace 0M <leir age 434
1. The Marine Fauna of Zanzibar and British East Africa, from Collections made by
Cyril Crossland, M.A., in the Years 1901 and 1902. —The Calcareous Sponges. By
C. FB. Jenkin Re ana gr rat ort ace isco arabe hore intel a edcter avails a eaten
2. Notes on the Australian Fossorial Wasps of the Family Sphegide, with Descriptions of
new Species. By Rowzanp E. Turwmr, F.Z.S. (Plate XXVI.) .....-- 2.2.12 0000s 457
May 26, 1908.
1. The Rudd Exploration of 8. Africa.—X. List of Mammals collected by Mr, Grant near
Tette, Zambesia. By Oxprierp Tomas, F.R.S., F.Z.S., and BR. C. Wroveurton, F.Z.8. 538
bo
. Zoological Results of the Third Tanganyika Expedition, conducted by Dr. W. A.
Cunnington, 1904-1905.—Report on the Isopoda terrestria. By the Rey. Tuomas R.R.
Srespine, M-A., F.RS., F.L.S., F-Z.8. (Plate XXVII.)..--.- -- 1.0 jee 554
1
3. On the Anatomy of Antechinomys and some other Marsupials, with special reference to
the Intestinal Tract and Mesenteries of these and other Mammals. By Frank H.
Brpparp, M.A., F.R.S., F.Z.8. .. -
4. The Armour of the Extinct Reptiles of the Genus Parciasawrus. By H. G. SnEney, a
ELR.S., F.Z.8., King’s College, London: ... 0... .s2..5 aces 2 oe ene | ee al
. Additional Evidence as to the Dentition and Structure of the Skull in the South African |
Fossil Reptile Genus Diademodon. By H. G. Suntuy, F.RB.S., F.Z.S., a Colle
London ec iifea th any aon aire COs eae
i, New Siphonaptera. By the Hon. N. Cuartes Roruscutty, MA, E.Z8. “(Plates
XXVITI.-XXXTI.) ..... :
COVES H ee ee ne td ce vere
oe 0) 00 81h a 0 ole ve 018 10/0 666 ec8 6 oe) ee (pe, ae #8) Wein e/iei hard uisliere
Contents continued. on nage 3 of Wra
PROCEEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.
(May to December, 1908.)
May 12, 1908.
F. Du Cane Gopman, Esq., D.C.L., F.R.S., Vice-President,
in the Chair.
The Secretary read the following report on the additions made
to the Society’s Menagerie during the month of April 1908 :—
The number of registered additions to the Society’s Menagerie
during the month of April was 135. Of these 75 were acquired
by presentation and 15 purchased, 43 were received on deposit,
and 2 were born in the Gardens.
The number of departures during the same period, by death
and removals, was 145.
Among the additions special attention may be directed to :—
A hybrid between a male Lion (felis leo) and a female Jaguar-
Leopard (Felis onca x Felis pardus), bred in the United States;
deposited on April 14th.
A Vaal Rehbok (Pelea capreolus) from the Drakensberg
Mountains, presented by Frederick Burgoyne, Esq., F.Z.8., on
April 3rd.
Two Secretary Vultures (Serpentarius reptilivorus) from South
Africa, purchased on April 21st.
Two Australian Cassowaries (Casuarius wustralis), presented by
Sir William Ingram, Bt., on April 24th.
Mr. W. Woodland, F.Z.S., exhibited preparations of a new
gland he had found in certain teleostean fishes, and made the
Proc. Zoou. Soc.—1908, No. XXVIII. 28
432 oN A NEW GLAND IN TELEOSTEAN FISHES. [May 12,
following remarks ._“This new gland is diffuse in form and is
intermingled with the veins and arteries which subdivide to form
the numerous parallel capillaries of the rete mirabile (text-fig. 79)
found in connection with all teleost ‘red bodies.’ It is quite
distinct from the gas-gland, and consists of rows of large columnar
cells, situated in close connection with the venous blood-vessels,
possessing large nuclei and nucleoli and packed with numerous
large spherical granules derived from the red-corpuscle dis-
integration concerned in the generation of the oxygen found in the
*e, a
"2goa582 9,
oF *,
R.M.
G.E.
Diagram of the construction of the gas-gland rete mirabile (“red body Ha)
R.M., rete mirabile; G.E., gas-gland epithelium. The new gland now described is
situated round the veins at the proximal pole (pole remote from the glandular
epithelium) of the rete mirabile.
Text-fig. 80.
D.L. Cc.B.G.
C.B.G., cells > new gl 1 1
Gees os eee nee gue suet round and in contact with a vem; D.L
longitudinal section of a duct of the new gl sontaining s i ranules
Pee een eet new gland containing spherical granules
om the gland-cells; D.7., a duct 1m transverse section
ee cae These granules, thus abstracted by the gland-cells
aoe a blood, are carried away by special ducts appertaining to
i ou (eee 2 ; The discovery of this important gland
several genera—G'obi Syngn 7H) ¢ :
g robius, Syngnathus, Pierasfer, B
others—confirms Jzeger’s view as ‘ a ee
: S e’s view as to the mode of g: i
oe 2 yas 1 de of generation of the
b xygen. The rete mirak raAS-2
g mirabile of the gas-gland apparatus is
1908. ] ON A WILD CAT FROM SZE-CHUEN. 433
to be explained as a mechanism for bringing the toxin secreted by
the gas-gland cells (Jeger) into contact with the erythrocytes before
these reach the region of the gas epithelium, so that when the
blood does reach this region, the oxygen, liberated by the action
of the toxin on the erythrocytes, has become freely mixed with the
plasma and is therefore in a condition to be abstracted by the gas-
gland cells.”
Mr. T. A. Coward, F.Z.8., exhibited a specimen of a Petrel,
“strelata neglecta Schleg., the property of Mr. Arthur Newstead,
of Cheshire, which had been picked up dead, yet in a quite
fresh condition, at Tarporley in Cheshire, on April Ist, 1908.
This bird is a native of the Southern Pacific, and has almost
certainly never been recorded from the northern hemisphere,
and certainly never from Europe before.
Mr. C. Davies Sherborn, F.Z.S., exhibited a specimen of chert
from the Middle Culm-measures (Carboniferous) of Christow
Down, near Doddiscombe Leigh, Devonshire, showing numerous
large and well-preserved Radiolaria.
On behalf of Mr. R. Lydekker, the Secretary exhibited the
tanned skin (without the legs and part of the tail) of a Wild Cat
obtained by the Hon. Mason Mitchell, of the American Consular
Service, in Sze-chuen, and sent by that gentleman to Mr. Rowland
Ward. Compared with a light-coloured skin of Felis temmincki
from Sikhim (B.M. No, 91.10.7.10), the Sze-chuen specimen differs
by the much lighter colour of the upper parts, which are golden
tawny, with a comparatively narrow dorsal streak of light rufous
(in place of a broad one of mahogany rufous)—the tail being
golden rufous above, different in tint from both the middle
and sides. The under parts are white with a few brown spots,
instead of pale buff with similar spots. The specimen is of
interest as extending the range of the Bay Cat into Sze-chuen ;
and may be regarded as representing a local race, for which
Mr. Lydekker suggested the name /. temmincki mitchelli.
It was likewise pointed out that the Bay Cat presents striking
resemblances to the African Tiger-Cat (2. chrysothrix, or
fF. aurata), from which it is distinguished by the broad white
band between a pair of darker ones on each cheek, and a some-
what similar mark onthe forehead. Sikhim and Nepal specimens
exbibit both a bright rufous phase with pale and spotted under
parts, and a wholly dark reddish-brown phase. <A grey phase is
represented by a skin (B.M. No. 0.6.30.1) from Upper Burma,
presented by Mr. C. W. A. Bruce, and by the Cat from Foochow
_ figured in plate i. of the Society’s ‘Proceedings’ for 1898 by
Dr. P. L. Sclater as a new species under the name of F. domini-
canorum. This phase corresponds with the one of F. chrysothrix
28*
434 MR. C. F JENKIN ON [May 12,
to which Mr. Lydekker gave the racial name cottoni. Whether the
name F. temmincki dominicanorum can be used for the Foochow-
Burma Bay Cat, or whether, as in the case of cottoni, it refers
merely to a colour-phase, remains to be proved.
Mr. J. T. Cunningham, M.A., F.Z.S., read a paper entitled
“The Heredity of Secondary Sexual Characters in Relation
to Hormones, a Contribution to the Theory of Heredity.” The
paper contained an examination and criticism of the most
important recent investigations and theories on the subject by
evolutionists of various schools, namely, the theory which attributes
such characters to constitutional causes such as male katabolism,
Prof Karl Pearson’s biometrical investigation of sexual selection
in man, Castle’s Mendelian theory of the heredity of sex, and
Geoffrey Smith’s views on dimorphism of males and _ parasitic
castration in Crustacea. The author maintained that all these
contributions were more or less inconsistent with the known facts
concerning the connection between the development of secondary
sexual characters and the functional activity of the primary
gonads. He drew attention to the recent discovery and experi-
mental proof on the part of physiologists that the development of
the characters was due to the stimulus of a chemical substance or
hormone produced by the testis or ovary, and passed into the
blood, and suggested that conversely hormones from parts of
the soma might affect the gametes in the gonads. In this way
the hypertrophy of a part of the body due to external stimulation
might modifiy the corresponding determinants in the gametes so
as to produce some hereditary effect in succeeding generations.
Mr. Cunningham added that his theory was an interpretation in
terms of modern physiology of Darwin’s theory of pangenesis.
The following papers were read :—
1. The Marine Fauna of Zanzibar and British East Africa,
from Collections made by Cyril Crossland, M.A., in
the Years 1901 & 1902.—The Caleareous Sponges. By
C. F. JENKIN *, ‘
[Received April 1, 1908. ]
(Text-figures 81-104.)
The Collection, made by Mr. Cyril Crossland at Wasin and
Zanzibar in 1901-2, passed through several hands and was finally
entrusted to the writer in the autumn of 1907.
* Communicated by Professor ARTHUR Denpy, D.Sc., F.L.S., F.Z.S.
1908. | AFRICAN CALCAREOUS SPONGES. 435
The collection consists of 25 specimens belonging to 14 species,
5 of which are new, as shown in the following list :—
Grade. | Family. Genus. Species. sg eae
| \pecimens.
| | |
Homocela ...| Clathrinide ...... Clathrina ..., primordialis (H.). 2
| 3 darwinii (H.). 3
. contorta (DMin.). 3
| x blanca (Mik.). 1
| Leucosoleniide | Leucosolenia | irregularis, sp. n. 1
Heterocceela .... Sycettide ......| Sycon .........| ciliatum Fabr. 2
|e ampullum (H.). 1
Werstye | munitum, sp. 1. 3
| Grantiide ......! Leucandra ... ananas (H.). 1
| Heteropide ...... Grantessa .... simplex, sp.n. 2
s | zanzibaris, sp. 0. 1
Amphoriscidx ...| Heteropegma | nodus gordii (Pol.). 3
| Leucilla ...... _ floridiana (#.). 1
| 39 | wasinensis, sp. 1. 1
|
Total specimens ...... 25
The classification 1s that proposed by Poléjaeff (2) for the
Homocela, and by Dendy (8) for the Heteroceela, with slight
modifications by Minchin (4).
The identification of calcareous sponges is very difficult and
unsatisfactory in the present state of our knowledge. Haeckel
in his great work (1) laid down hard and fast definitions of the
different species, which if they accorded with the facts would
make identification very easy, but unfortunately actual specimens
very seldom fall within his definitions, This has led to a useless
multiplication of species, since each specimen which did not
exactly comply with Haeckel’s definition has been called by a
new name. Haeckel has also omitted to mention many striking
features of his species, such as the subgastral quadriradiates in
many of the Sycandra, the characteristic dermal spicules of some
species, and the hair-spicules in most of the species in which they
occur (e.g. in Sycandra ciliata). He has also made numerous
wrong identifications (see Minchin 5). Under these circumstances
no identification can be considered as certain and nothing very
satisfactory can be done till Haeckel’s work has been revised.
The specimens in the Crossland Collection are unfortunately
not in a good state of preservation. It therefore seemed better
to place the specimens among existing species, even if the iden-
tification was doubtful, rather than to make new species based on
single specimens in a poor state of preservation. This has been
done as far as possible, but there remained six specimens belong-
ing to five species which could not be classed in this way; to
these new names have been given.
436 MR. C. F. JENKIN ON [May 12,
Description of the Specimens.
CLATHRINA PRIMORDIALIS.
Ascetta primordialis H.
Two specimens of this sponge were dredged in 6 to 8 fathoms
at Wasin. ‘The dimensions of the spicules agree with those given
by Haeckel of specimens from Australia. Most of the spicules
have rays from 160-180 p long x 16-20 p thick.
CLATHRINA DARWINII. (Text-figs. 81, 82.)
Ascaltis darwintt H.
Three specimens of this sponge were dredged in 3 fathoms in
Chwaka Bay, Zanzibar. The largest specimen is shown, twice
Text-fig. 81.
Clathrina darwinii (H.). X 2.
Text-fig. 82.
Clathrina darwinii (H.), spicules. X 130.
ap., apical ray.
oe size, in text-fig. 81. They were bright lemon-yellow
colour when alive, and are buff-white in spirits. All three
1908. ] AFRICAN CALCAREOUS SPONGES. 437
specimens consist of solid lumps of the anastomosing tubes
typical of the genus Clathrina. They are firm to the touch and
similar in appearance to C. coriacea as it grows in the Channel
Islands.
The facial rays of the tri- and quadriradiates (text-fig. 82) vary
from 60-120 long x 12-16» thick, the commonest size being
110x15p. The apical rays of the quadriradiates are nearly
straight, about the same length as the facial rays and about 8 pu
thick.
The spicules are considerably thicker than in Clathrina con-
torta var. spinosa (Min.), and agree better with Haeckel’s Clathrina
darwin.
CLATHRINA CONTORTA var. SPINOSA. (Text-figs. 83, 84.)
Clathrina contorta var. spinosa Minchin (6).
Three specimens of this sponge were dredged in 3 fathoms
in Chwaka Bay, Zanzibar. They were pure white when alive and
are translucent white in spirits. They are very delicate in texture,
and readily fall to pieces. The largest specimen is a solid lump
shown twice natural size in text-fig. 83; the others appear to
be fragments only. They differ considerably in appearance from
the specimens of Clathrina darwinit which were collected at the
same time.
Text-fig. 83.
Clathrina contorta (Min.). 2.
The spicules (text-fig.{84) agree fairly well with Minchin’s (6)
and von Lendenfeld’s (7) descriptions of Clathrina contorta var.
spinosa. No oxea were found.
438 MR. C. F. JENKIN ON [May 12,
The rays of the triradiates vary from 100-130 » long x 10-12 pz
thick, the commonest size being 125 x 11 p. i .
The facial rays of the quadriradiates vary from 80-150 p long
x 10-12 p thick, the commonest size being the same as of the
triradiates 125 x 11 mu.
Text-fig. 84.
-
Be gl
Clathrina contorta var. spinosa, spicules. 180.
ap., apical ray.
The apical rays are 50-65 p long x 5-7 p thick; they are thicker
than those in Minchin’s or von Lendenfeld’s specimens.
CLATHRINA BLANCA. (Text-figs, 85-87.)
Guancha blancha Miklucho.
Ascetta blanca Hi.
One specimen of this sponge was dredged in 10 fathoms at
Wasin. Its colour in spirits is pure white. Its shape is shown
(natural size) in text-fig. 85. The sponge is formed of flat fan-
shaped heads on the ends of a branching stalk. The largest head
is shown (x5) in text-fig. 86. It consists of a flat mass of
anastomosing tubes with several oscules on the outer edge ;
sections show that the head is about three tubes thick, and that
the tubes are so arranged that the spaces in the meshwork never.
extend right through it. The stalk is somewhat flattened and
1908. | . AFRICAN CALCAREOUS SPONGES. 439
Text-fig. 85.
Clathrina blanca (Mik.). Nat. size.
Text-fig. 86.
Clathrina blanca (Mik.). 5.
440 MR. C. F. JENKIN ON [May 12,
is solid, The flat shape of the specimen is remarkable ; the
ordinary shape of the heads of Clathrina blanca 18 more or less
spherical. Ree
Phe skeleton consists of regular and sagittal triradiates (text-
fig. 87). The spicules agree fairly well with the descriptions
given by Haeckel (1) and von Lendenfeld (7).
Text-fig. 87.
Clathrina blanca, spicules. X 220.
The body-spicules are mostly regular, but some have the basal
rays slightly the longest. Paired rays 65-70 4x4-6p. Basal
ray 70-110 px 4-6°5 p.
The stalk-spicules are almost all sagittal, the smaller ones on
the outside and the larger ones inside, all arranged with the
basal ray downwards. Paired rays 60-80 ux 5-9 p. Basal ray
100-160 p» x 70-110 p.
LEUCOSOLENIA IRREGULARIS, sp. n. (Text-figs. 88-90.)
One small specimen of this new species was found among the
Clathrina primordialis dredged in 6-8 fathoms at Wasin. It
consists of two erect tubes with a short rooting tube (text-fig. 88).
The skeleton (text-fig. 89) is rather remarkable. It contains no
triradiates but is made up of two types of equiangular quadri-
radiates, lying without orientation, together with a few large
oxea which project in all directions.
1908. | AFRICAN CALCAREOUS SPONGES. 44)
Most species of Zewcosolenia contain triradiates as well as
quadriradiates, and the facial rays of both sorts of spicule are
usually alate and regularly placed with the basal ray downwards.
Text-fig. 88
Leucosolenia irregularis. X.
Text-fig. 89.
3
Leucosolenia irregularis, skeleton of dermis. X 150.
All the spicules in the new species are large, two or three
times the size of those in the British species of Leucosolenia.
The facial rays of the larger quadriradiates are usually of the
442 MR, C. F. JENKIN ON [May 12,
p riradiates are often of
ame length, but those of the smaller quadriradiates are 0
ee val oie The oscules of both tubes are too much damaged
to furnish any indication of their structure.
Text-fig. 90.
Leucosolenia irregularis, spicules. X 110.
(For explanation of the letters see text below.)
Spicules (text-fig. 90).
The oxea are of one sort :—
(a) Nearly straight oxea sharply pointed at both ends, 300-
800 » long x 16-28 p thick ; the usual thickness is 21-24 p.
The quadriradiates are of two sorts :—
(6) Large quadriradiates. Facial rays approximately regular,
150-220 p long x 20 » thick. Apical rays usually slightly bent
near the point, 210-260 p long x 16—20 p thick.
(c) Small quadriradiates. Facial rays usually regular, occasion-
ally of unequal lengths, 100-200 p long x 10-16 p thick. Apical
rays very slender and sharply pointed, always bent near the point,
120-150 p long x7 p thick.
SYCON cILiaTuMm Fab.
Sycandra ciliata H.
Two small specimens of this species were dredged in 7 fathoms
in the Zanzibar Channel. Their dimensions are 8x 2 mm. and
5x 1z7 mm.
1908. AFRICAN CALCAREOUS SPONGES. 443
SYCON AMPULLUM,
Sycandra ampulla H.
One small specimen of this species was dredged in 6-8 fathoms
at Wasin. It is 11 mm. longx5 mm. diameter. Its structure
is typical of the genus. The spicules are too small for Sycon
raphanus, and there are none of the thin subgastral tri- and
quadriradiates which are the most characteristic features of that
species. Comparing it with Sycon ampullwm the spicules are
rather thicker than the dimensions given by Haeckel and the
oxea are also rather longer, but on the whole it agrees fairly well.
SYCON MUNITUM, sp.n. (Text-fig. 91.)
Three specimens of this new species were dredged in 7 fathoms
in the Zanzibar Channel. In external appearance they resemble
small specimens of Sycon ciliatum. Their dimensions are
7x3mm.,5x3 mm., and 3x1 mm.
The peculiarity of the species is the presence of quadriradiates
in the articulated tubar skeleton. Only a fewspecies of Sycon are
known with quadriradiates in this position; from these the new
species is differentiated by having two sorts of gastral quadri-
radiates, viz., small quadriradiates with short apical rays and
larger ones with very large apical rays.
Text-fig. 91.
Sycon munitum, sp. n., spicules. X 110.
(For explanation of the letters see text p. 444.)
Skeleton.—The gastral skeleton is a dense felt of small tri- and
quadriradiates, fairly regularly arranged round the apopyles, with
the basal rays aborally directed and the small apical rays pro-
444 MR. C. F. JENKIN ON [May 12,
jecting into the gastral cavity. The apical rays being short, many
of them hardly reach further than just through the thick gastral
layer. Among these spicules, every here and there, lies one of
the large quadriradiates, with its large apical ray projecting far
into the gastral cavity.
The articulated tubar skeleton is built up of tri- and quadri-
radiates. The short apical rays of the quadriradiates project
into the flagellated chambers. The tops of the chambers are
crowned with tufts of small oxea.
The oscule has a thick fringe of thin straight oxea. The
flagellated chambers get shorter near the oscule and there is no
collar. There are remains of a diaphragm across the oscule.
Spicules (text-fig. 91).
The oxea are of one sort :—
(a) Oxea from the ends of the flagellated chambers, nearly
straight, pointed at both ends, 170-400 p long x 8 » thick. Some
of these are more refringent than others. The refringent spicules
are quite straight.
The triradiates are of three sorts :—
(6) Alate triradiates from the tubar skeleton. Basal rays
straight, 110-170 » long x 5-6 4 thick. Paired rays bent upwards,
60-100 » long x 5-7 p thick. Oral angle about 140°.
(c) Alate subgastral triradiates. Basal rays straight, 180-230 pu
long x6» thick. Paired rays bent downwards, 80-100 p long x
4-6 pw thick.
(d) Alate triradiates from the gastral layer. Basal rays straight,
80-210 » long x 6-8 thick. Paired rays bending upwards,
sometimes unequal in length, 70-130 pw long x 6-8 p» thick.
The quadriradiates are of three sorts :—
(ec) Alate quadriradiates from the tubar skeleton. The facial
rays are similar to (4) but larger. Basal rays 140-220 p long.
Paired rays 70-100» long. Apical rays slender, slightly bent
near the point, 50 uw long x 3 pw thick.
(f) Small alate quadrivadiates from the gastral layer, similar
to (d) with the addition of an apical ray 60 p» long x 6 p. thick.
(g) Large alate quadriradiates from the gastral layer. Basal
rays straight, over 200» long x10, thick. Paired rays nearly
straight, 180 longx 9-10, thick. Apical ray slightly bent
orally, 320-380 » long, oval in section 16 pw deep x 8 ps thick.
Leucanpra ANANAS H. (Text-fig. 92.)
Leucandra ananas H.
One specimen of this species was dredged in 3 fathoms in
Chwaka Bay, Zanzibar. It is flask-shaped, 20 mm. long x
13 mm. diameter, with an oscule 5 mm. diameter. It was dirty
white in colour when alive. The body-wall is about 3 mm. thick
near the middle and surrounds a gastral cavity about 7 mm.
diameter.
1908. ] AFRICAN CALCAREOUS SPONGES. 445
Canal-system.—There are large incurrent chambers under the
dermis from which the large incurrent canals run radially
inwards. The excurrent canals are also large and run radially
between the others; they communicate with the gastral cavity
through large ports.
Text-fig. 92.
Leucandra ananas, spicules. X 56.
(For explanation of the letters see text below.)
Skeleton.—The dermal skeleton consists of a thin layer of
delicate triradiates lying without orientation. Tufts of large
oxea project radially from slightly raised papille on the dermis.
The body skeleton consists of irregularly placed large triradiates.
The excurrent canals are lined with quadriradiates, the apical
rays projecting into the canals. The gastral skeleton consists
of a dense layer of quadriradiates with the apical rays projecting
into the gastral cavity.
The specimen agrees fairly well with Haeckel’s description of
Leucandra ananas, though the spicules are rather larger, and the
dermal skeleton differs from the body skeleton, which is not men-
tioned by Haeckel. Haeckel, however, makes a similar omission
in other cases, e.g. in his description of Leucandra fistulosa.
Spicules (text-fig. 92).
The oxea are of one sort :—
(a) Nearly straight sharply-pointed oxea, 700-3000 p» long x
28-46 yw thick.
446 ON AFRICAN CALCAREOUS SPONGES, [May 12.
The triradiates are of two sorts :—
(6) Slender dermal trivadiates, subregular. Rays 160-280 p
long x 5-10 w thick.
(c) Subregular triradiates from the body. Basal ray straight,
120-400 » long x 20-32 » thick. Paired rays almost straight,
140-550 pw long x 16-26 pw thick.
The quadriradiates are of two sorts :—
(d) Subregular quadriradiates lining the excurrent canals.—
Basal rays straight, 180-250 p long x 10-20 » thick. Paired rays
nearly straight, 200-280 w long x 6-17 » thick. Oral angle 120°.
Apical rays 170 x 6-8 p thick.
(ec) Alate quadriradiates from the gastral skeleton, similar
to (2) but with a larger oral angle, about 135°. Apical ray
50 p long x 6-8 pw thick.
GRANTESSA SIMPLEX, sp. n. (Text-figs. 93-97.)
One specimen of this new species was dredged in 6-8 fathoms
at Wasin, and one specimen was collected on the shore of Chwaka
Bay, Zanzibar.
They both consist of a confused mass of anastomosing tubes
(see text-fig. 93). The oscules are at the free ends. The tubes,
as preserved, are a good deal flattened and vary in size from
2-5 mm. in diameter. Both external and internal surfaces are
smooth.
Text-fig. 93.
Grantessa simplex, sp.n. Nat. size.
The structure of the body-wall is very regular and typical of
the genus Grantessa (see text-fig. 94).
The skeleton is formed entirely of triradiates. The dermal
skeleton (text-fig. 95) consists of alate triradiates lying tangen-
tially, without orientation. The subdermal triradiates (c, text-
fig. 97) are modified dermal spicules; the centripetal ray is one
of the paired rays, not the basal ray*; it is considerably longer
* Of, Poléjaeff (2).
Text-fig. 94,
simplex, skeleton of body-wall. X 100.
Grantessa
Text-fig. 95.
simplex, skeleton of dermis. X 160.
29
Proc. Zoot, Soc.—1908, No. XXIX.
Grantessa
_ Text-fig. 96.
ex, gastral skeleton. XX 150.
impl
mtessa s
Gra
Text-fig. 97.
spicules. > 100.
simplex,
ntessa
Gra
(For explanation of the letters
see text p. 449).
1908. ] ON AFRICAN CALCAREOUS SPONGES. 449
than the other paired ray. The basal ray is bent sharply near the
root; but for this bend the point would project beyond the dermal
layer. A few spicules of a similar form are found lying tan-
gentially among the dermal triradiates. The subgastral triradiates
are alate, with equal paired rays and an oral angle of 155°. The
gastral skeleton (text-fig. 96) is a thick layer of alate triradiates
similar to those in the dermis, but slightly smaller and rather
more regularly placed. The oscule has no special skeleton. The
body-wall ends abruptly, and there is hardly a trace of a collar.
Spicules (text-fig. 97).
The triradiates are of four sorts :—
a) Alate triradiates from the dermis. Basal rays straight,
210-250 » long x 10-12 » thick. Paired rays straight, except for
a slight curvature near the junction of the basal ray, which
rounds the oral angle smoothly, 200 » long x 8-10 » thick. Oral
angle 110°. iY
(0) Alate triradiates from the gastral layer, similar to (a) but
smaller. Basal ray 180, (occasionally much shorter). Paired
rays 140-150 long. Oral angle 115°-120°.
(c) Subdermal triradiates. Basal ray (lying in the dermis)
straight, except for an angular bend near the root, 130-190 «
long x 14 » thick. Centripetal paired ray straight, 240-420 w
long x 11-13 w thick. Dermal paired ray considerably bent near
the root, 160-190 » long x 10-124 thick. Oral angle 100°.
(d) Subgastral triradiates. Basal ray straight, 360-390 pw long
x 13-16 » thick. Paired rays, equal, bent downwards near the
root, 180-240 » long x 10-124 thick. Oral angle 155°.
GRANTESSA ZANZIBARIS, Sp. n. (Text-figs. 98-102.)
One specimen of this new species was dredged in 6-8 fathoms
Text-fig. 98.
Grantessa zanzibaris, sp.u. Nat. size.
at Zanzibar. It consists of a mass of branching tubes of various
29*
450 ON AFRICAN CALCAREOUS SPONGES. [May 12.
sizes; there is no anastomosis between the branches ee pe
fig. 98). The tubes, as preserved, are flattened so - at He
opposite sides touch ; they vary in size from 1-3 mm. wide. 1
external surface is smooth; the internal surface is lined wit
quadriradiates with the apical rays projecting inwards.
Text-fig. 99.
—S——————
BE
Grantessa zanzibaris, skeleton of body-wall. X 200.
The structure of the body-wall is shown in text-fig. 99. The
opposite rays of the subdermal and subgastral triradiates lie
beside each other, forming a typical non-articulated skeleton ;
in addition there are two or three intermediate rows of tri-
radiates, apparently the remains of an articulated skeleton. The
structure agrees closely with that of Grantessa intusarticulata
described and figured by Dendy (8).
452 MR. C. F, JENKIN ON [May 12,
Skeleton.—The dermal skeleton (text-fig. 100) consists of a thin
layer of alate triradiates lying tangentially, without orientation.
The subdermal triradiates are modified dermal spicules, as In
G. simplex. The subgastral triradiates are alate, with equal paired
rays and an oral angle of 165°. The gastral skeleton (text-
fig. 101) consists of a thin layer of sub-regular alate quadri-
radiates, lying without orientation.
Text-fig. 102.
Grantessa zanzibaris, spicules. X 120.
(For explanation of letters see text below.)
Spicules (text-fig. 102).
The triradiates are of four sorts :—
(a) Alate triradiates from the dermis. Basal rays straight,
65-190 w long x 8-10 thick. Paired rays nearly straight,
except for a slight curvature near the junction of the basal ray,
rounding the oral angle, 60-130, long x 8-9 thick. Oral
angle 110° to 125°. :
(6) Alate triradiates from the body similar to (a) but with
es curved paired rays considerably folded. Oral angle 125°
to .
(c) Subdermal triradiates. Basal ray, lying in the dermis,
straight, except for an angular bend near the root, 90-140 pw long
x 8-9 pw thick. Centripetal paired ray straight, 120-200» long
1908. | AFRICAN CALCAREOUS SPONGES. 453.
x 6-8 thick. Dermal paired ray considerably bent near the
root, 80-130 w long x 6-8 » thick. Oral angle 95° to 110°.
(d@) Subgastral triradiates. Basal ray straight, 100-260 yw long
x 7-9» thick. Paired rays equal, bent downwards near the root,
80-140 » long x 7-8 p thick. Oral angle 165°.
The quadriradiates are of one sort :—
(e) Quadriradiates from the gastral layer. Basal rays straight,,
130-150 » long x9 » thick. Paired rays straight, 90-130 » long
x 8-10 thick. Apical rays 80-130 » long x 4 p» thick.
HETEROPEGMA NODUS GoRDII Pol. (Text-fig. 103.)
Three specimens of this sponge were dredged in 6-10 fathoms
at Wasin. In form and size they agree closely with Poléjaeff’s
description and figures (2). The spicules (text-fig. 103) also agree
with his description with the following slight differences. The
minute tri- and quadriradiates in the Wasin specimens are about
4 thick, whereas Poléjaeff gives 2. There are none of the
intermediate forms mentioned by Poléjaeff between the remark-
able alate tri- and quadriradiates which line the gastral cavity
and the regular tri- and quadriradiates which are scattered
through the body.
Text-fig. 103.
Heteropegma nodus gordii, spicules. X 40.
LEUCILLA FLORIDIANA.
Leucaltis floridiana H.
One specimen of this sponge was dredged in 10 fathoms at
Wasin. It is irregular in shape, 10x75 mm., and has an
454 MR. C. F. JENKIN ON [May 12,.
oscule 3 mm. diameter flush with the surface; there is no collar:
or fringe.
The spicules agree very well with Haeckel’s description of
Leucaltis foridiana. The apical rays of the large quadriradiates.
in the dermis point inwards, the species must therefore be placed
in the genus Leucilla. The small quadriradiates line the excurrent.
canals, with their apical rays projecting into them.
LEUCILLA WASINENSIS, Sp. n. (Text-fig. 104.)
One specimen of this new species was dredged in 6-8 fathoms
at Wasin. It is ovoid in form, 16 mm. long x7 mm. diameter,
with a fringed oscule, 2 mm. in diameter. It is white, as pre-
served in spirit. The body-walls are about 2 mm. thick, leaving
a gastral cavity 3 mm. diameter in the centre. Externally it is
rough with the stumps of large projecting oxea. The oscule is
protected by a dense fringe of thin oxea, surrounded by a few
thick ones like those projecting from the dermis.
Text-fig. 104 A.
Ng
Leucilla wasinensis, spicules. X 40.
(For explanation of the letters see text p. 455.)
Canal-system.—There are large subdermal chambers from which
branch the incurrent canals. The excurrent canals are also
branched. The canal-system is similar to that most usual in the
genus Leucandra.
1908. | AFRICAN CALCAREOUS SPONGES. 455
Skeleton.—The dermal skeleton consists of a thin layer of alate
triradiates lying tangentially without orientation ; amongst them
are a few quadriradiates, with the apical rays directed inwards,
some of very large size. The gastral skeleton consists of a thick
layer of large alate quadriradiates, regularly placed with the
basal rays directed aborally and the apical rays projecting into
the gastral cavity. The skeleton of the central mass of the body-
wall between the dermal and gastral layers is a closely packed
mass of irregularly placed tri- and quadriradiates. The quadri-
radiates, some of which are very large, are mostly arranged round
the mouths of the excurrent canals, with the apical rays projecting
into the canals. The large projecting oxea are bunched into
little tufts. The inner ends usually pierce nearly through the
wall, and occasionally right through into the gastral cavity.
But for the presence of the large dermal quadriradiates this
sponge would be a typical Leucandra.
Text-fig. 104 B.
Lasav l U
Leucilla wasinensis, spicules. X 110.
(For explanation of the letters see text below.)
Spicules (text-fig. 104 A & B).
The oxea are of one sort :—
(a) Nearly straight oxea, sharply pointed at the inner end,
outer end snake-headed. The largest fragment is 2°8 mm. long
x 45 pw thick.
456 ON AFRICAN CALCAREOUS SPONGES. [May 12, |
The triradiates are of two sorts :—
(6) Alate trivadiates from the dermis. Basal rays straight,
150-240 p long x 10-13 w thick. Paired rays, curving slightly .
upwards, 160-320 p long x 9-13 p thick. Oral angle 110°.
(c) Subregular triradiates from the body. Rays very sharply
pointed, 600-950 pw long x 35-40 p thick. Individual spicules
often have their three rays of different lengths, but the angles,
are always approximately 120°.
The quadriradiates are of three sorts :—
(2) Subdermal quadriradiates. These spicules vary enormously
in size, and are of peculiar form as shown in the drawing. The
facial rays are folded inwards, 7. e. towards the side from which’
the apical ray projects. The maximum facial ray found is
700 » long x 26 thick. The maximum apical ray is 500 p long
x26 pw thick.
(e) Alate quadriradiates from the lining of the excurrent canals.
Basal rays straight, 280-480 ». long x 20-28 w thick. Paired rays
nearly straight, 340-420 long x 14-24 thick. Oral angle
130°.
(f) Alate quadriradiates from the gastral layer. Basal rays
straight, 300-560 « long x 10-12 thick. Paired rays, bending
upwards, slightly irregularly bent, 240-450 w long x 8-10 pw thick.
Apical ray, curved orally, 180-260» long x 8» thick. Oral
angle 110° to 120°. ae
List of Literature referred to in the paper.
(1) Haxcken.—Die Kalkschwimme. Berlin, 1872.
(2) PottsaErr.—Challenger Report, Monogr. vii., 1883.
(3) Denpy, A.—“ Studies on the Comparative Anatomy of
Sponges. V. Observations on the Structure and Classifi-
cation of the Calcarea Heterocela.” Q.J.M.S. vol. xxxv.
(1893) pp. 159-257, pls. x.—xiv.
(4) Mivcury, E. A.—A Treatise on Zoology. Edited by EH. Ray
Lankester. Part II. The Portfera and Coelenterata.
Chap. III. Sponges, by KE. A. Minchin, M.A., 1900.
(5) Mincuin, E. A.—‘‘The Characters and Synonymy of the
British Species of Sponges of the Genus Lewcosolenia.”
Proc. Zool. Soc. Lond. 1904, vol. ii. pp. 349-396.
(6) Mincuin, E. A.—‘“‘On the Sponge Leucosolenia cortorta
Bowerbank, Ascandra contorta H., and Ascetta spinosa
von Lendenfeld.” Proc. Zool. Soc. Lond. 1905, vol. ii.
pp. 3-20.
(7) LenpenreLp, R. voy.—“‘ Die Spongien der Adria: I. Die
Kalkschwimme.” Zeitsch. wiss. Zool., lili. Bd., 2 u. 3
Heft., 1891, : ye
PZ.S. 1908. Pl xem
West, Newman chr.
H. Knight ad nat.del.
SPHEGIDA.
AUSTRALIAN
ia
1908. ] ON AUSTRALIAN FOSSORIAL WASPS. 457
2. Notes on the Australian Fossorial Wasps of the Family
Sphegide, with Descriptions of new Species. By
Row Land EH. Turner, F.Z.S.
[Received April 6, 1908.]
(Plate XX VI.* and Text-figures 105-110.)
The Sphegide of Australia have attracted very little notice
since the time of F. Smith ; the cnly works in which any number
of species have been dealt with being Kohl’s Monograph of the
genus Sphex and various papers by Handlirsch on Bembex and
the allied groups. In consequence, it has been impossible to get
any reliable idea as to the characteristic points which might
attract notice as to the peculiarities of Australia in this group.
The present paper, although necessarily very incomplete from want
of sufficient material, is based upon the large collection formed
by the late Gilbert Turner in North Queensland together with
smaller additions from other sources. Considering the sandy
nature of much of the continent, usually so favourable to fossorial
wasps, Australia is rather poor in Sphegide, and several wide-
ranging genera represented in almost every other region seem to
be entirely absent. The most notable instances are the genera
Philanthus and Oxybelus, but Ampulex, so conspicuous in the
Indian and African regions, is also absent. Nor is it likely that
the absence of these genera in collections is due to insufficient
field-work, for North Queensland is the most likely part of the
continent for these wide-ranging genera to occur in, the line of
migration having been through New Guinea. Thus I am able to
record two species of Zrypoxylon closely allied to Indian and
Malayan forms, no species of the genus having yet been discovered
in the southern or western districts. On the contrary, genera
peculiar to Australia are not numerous and seem to be poor in
species, though probably more exhaustive collecting in Western
Australia would add to the number. Most characteristic of the
country are Sericophorus and the allied genera Zoyphiwm and
Sphodrotes, also the large Hueirus; but the worldwide genus
Pison is much richer in ‘species in Australia than elsewhere.
Harpactophiius is apparently a tropical genus, and will probably
be found to have its headquarters in New Guinea. The small
amount of material available from Central Australia seems to
point to the plentiful occurrence of a few wide-ranging species,
mostly identical with those of Western Australia.
STIGMUS QUEENSLANDENSIS, Sp. 0.
Q. Clypeus strongly convex, raised in the middle into a broad
carina, which is more narrowly continued on the front to the
anterior ocellus. Clypeus and vertex almost smooth, front finely
* For explanation of the Plate see p. 535.
458 MR. R. E. TURNER ON [May 12,
and closely punctured-rugose, slightly concave, with a depressed
row of large punctures, bordered within by a carina, along the
inner margin of the eyes. Eyes large, reaching the base of the
mandibles and slightly convergent towards the vertex, the
posterior ocelli rather farther from each other than from the eyes.
Antenne inserted nearer to the eyes than to each other, the
flagellum three times as long as the scape. Pronotum very short,
much narrower than the head, the anterior margin raised and with
prominent angles, a transverse row of large, deep punctures
behind it; the propleure obliquely striated, the mesopleure
rugose. Mesonotum and scutellum closely and very finely punc-
tured. The median segment as long as the thorax, vertically
Text-fig. 105.
Neuration of fore wing.
1. Stigmus queenslandensis. 2. Tachysphex pilosulus. 3. Zoyphiwm kohlii.
4. Gorytes icarioides. 5. Nitela kurande. 6. Pison (Parapison) aberrans.
truncated posteriorly, coarsely transversely striated; a triangular
space reaching from the base to the apex bordered by raised
carine, with two parallel longitudinal carine close to the middle
reaching from the base to the apex. Abdomen subsessile, not
petiolate, smooth and shining, about as long as the thorax and
median segment combined. The second cubital cell is very small,
triangular, and almost petiolate, the recurrent nervure is received
at about one-third from the apex of the first cubital cell, which is
longer than is usual in the genus.
Black; the mandibles pale yellow ; antenne, legs, and tegule
1908. | AUSTRALIAN FOSSORIAL WASPS. 459
pale ferruginous. Wings hyaline, slightly iridescent ; nervures
pale ferruginous.
Length 5 mm.
Hab. Mackay, Queensland (Z'urner); April.
Differs from typical Stigmus in the non-petiolate abdomen.
HARPACTOPHILUS STEINDACHNERI Kohl.
Q. The posterior ocelli are nearer to the posterior margin of
the head than to each other.
dg. The mandibles, antennz, and legs are ferruginous; the
front, cheeks, head beneath, and prosternum golden. The front is
much more closely and finely punctured than in the female. The
apical dorsal segment of the abdomen is strongly emarginate.
As in all the species of the genus, there is a semicircular depression
at the base of the second ventral segment.
Hab. Mackay, Queensland ; October to May. Also from Cairns
and Cooktown.
HARPACTOPHILUS BICOLOR Sm.
@. Very near H. steindachnert, from which it differs in the
greater breadth of the thorax, which is almost as wide as the head,
the more prominent angles of the pronotum, and the fuscous
colour of the wings.
Hab. Mysole.
HARPACTOPHILUS KOHLII, sp. n.
9. Mandibles bidentate at the apex, the teeth short and feeble,
the imner tooth a little the longest. Clypeus convex, with a
longitudinal carina from the base almost reaching the apex, at the
extreme apex there isa small, smooth, triangular truncation ; the
sides of the clypeus are punctured. Above the clypeus is a very
prominent, narrow, longitudinal carina, on each side of which are
short curved strie. The front between the eyes and the base of
the antenne is very closely punctured and covered with short,
thin, greyish pubescence. Antenne inserted very low down on
the sides of the clypeus, far apart, as far from each other as from
the eyes, the scape as long as the first five joints of the flagellum
and equal in length to a little more than two-fifths of the distance
between the eyes at the base of the antenne. Vertex very
coarsely rugose-striate, the cheeks very broad and coarsely striated.
The ocelli situated in an almost equilateral triangle, the posterior
ocelli almost in a straight line with the summit of the eyes,
half as far again from the eyes as from each other, and more
than twice as far from the posterior margin of the head as
from each other. Kyes surrounded by a narrow sulcus, which
is coarsely and closely punctured. The posterior margin of the
head broadly emarginate, and about one-third broader than
the mesonotum. Pronotum depressed below the level of the
mesonotum and invisible from above. Mesonotum punctured-
rugose, more than half as broad again as long; the scutellum
460 MR. R. E. TURNER ON [ May 12,
almost smooth. Mesopleure rather finely rugose-striate. Median
segment much broader than long, narrowed towards the apex and
vertically truncate posteriorly, shorter than the mesonotum; a
large triangular space occupying almost the whole of the dorsal
surface very coarsely reticulate and enclosed by carinee ; the sides
of the segment striated, the surface of the posterior truncation
coarsely rugose. Abdomen smooth and shining, shorter than the
thorax and median segment combined. Legs not spinose.
Black; the mandibles (except the extreme apex), the scape of the
antenne, the apical half of the femora, the tibie above, and the
tegule yellow; the apex of the mandibles, the flagellum, the
abdomen, the base of the femora, the tibize beneath, and the tarsi
ferruginous. Wings hyaline, nervures pale ferruginous.
The second cubital cell is more pointed on the radial nervure
than in H. steindachneri Kohl, from which it may also be dis-
tinguished by the small triangular truncation at the apex of the
clypeus, the larger head, which is much more produced posteriorly,
and the absence of a visible pronotum above. It is somewhat
intermediate between H. steindachneri and H. arator, but is
nearer to the former.
Length 7-8 mm.
Hab. Mackay, Queensland (Z'wrner).
HARPACTOPHILUS SULCATUS, Sp. n.
Q. Clypeus triangular, convex, with a longitudinal carina from
the base not quite reaching the apex, where there is a very small
oblique triangular truncation. Mandibles feebly bidentate at the
apex, the outer tooth a little thelongest. Antenne a little longer
than the head, the length of the scape equal to nearly half the
distance between the eyes at the insertion of the antenne ; the
first four joints of the flagellum much longer than broad, the
second joint twice as long as the first and more than half as long
again as the second. Head very large, coarsely longitudinally
striated, the strie curving round the summit of the eyes, a narrow
depressed line divided by deep punctures round the margin of the
eyes, the small space between the eyes and the base of the
mandibles smooth. The posterior ocelli nearly twice as far from
the eyes as from each other and about the same distance from the
posterior margin of the head as from the eyes. The posterior
margin of the head depressed and broadly emarginate. Pronotum
depressed below the mesonotum ; the mesonotum about one-third
narrower than the head, about half as broad again as long,
coarsely longitudinally striated in the middle, coarsely rugose on
the sides. Mesopleurz obliquely striated; scutellum almost
smooth. Median segment a little shorter than the mesonotum, |
the large enclosed triangular area on the dorsal surface very
coarsely rugose, the segment strongly narrowed to the apex and
vertically truncate posteriorly, the sides coarsely striated.
Abdomen shining and almost smooth. The carina on the front
of the head, between the grooves for the scape, is less strongly
developed than in the allied species.
1908. ] AUSTRALIAN FOSSORIAL WASPS. 461
Black ; the base of the scape yellow in front; the mandibles
(except the extreme apex), the antenne, abdomen, tibia, tarsi, and
the apex of the femora ferruginous. Wings fusco-hyaline,
nervures dark ferruginous.
The second cubital cell on the radial nervure is about one-third
of the length on the cubital nervure, the radial cell is longer than
in the allied species, and the nervures both on the fore and hind
wings are produced, reaching much nearer to the margin. The
recurrent nervure is received just before the apex of the first
cubital cell.
Length 8 mm.
Text-fig. 106.
1. Harpactophilus sulcatus (head). 2. H. steindachneri Kohl: ¢ organs.
Hab. Kuranda, near Cairns, Queensland (Zurner); January.
Most nearly allied to H. arator and H. kohlii, but the head is
much shorter and broader than in the former species ; the antennze
are longer than in other species of the genus and the proportions
of the basal joints of the flagellum very different.
HARPACTOPHILUS ARATOR, sp. n. (Plate XX VI. fig. 1.)
©. Mandibles bidentate at the apex, the teeth short and feeble.
Clypeus very slightly advanced, obliquely triangularly truncate
from the centre to the apex. Head very large, subquadrate,
emarginate posteriorly, half as broad again as the mesonotum,
exceedingly coarsely striated, the striz longitudinal, curving
round the summit of the eyes; a very strong longitudinal carina
starting just’ below the anterior ocellus and produced in the shape
of a ploughshare prominently overhanging the base of the clypeus.
Eyes not quite reaching the base of the mandibles; the posterior
ocelli nearly twice as far from the eyes as from each other
and more than half as far again from the posterior margin of the
head as from theeyes. Antennz inserted lower than the base of
the clypeus, far apart, as far from each other as from the
eyes. Pronotum depressed below the mesonotum, which is
coarsely longitudinally striated; the scutellum short, almost
smooth, with a feebly impressed median line. Median segment
462 MR. R. E. TURNER ON [May 12,
shorter than the mesonotum, narrowed and truncate posteriorly,
the dorsal surface exceedingly coarsely reticulate, the sides of
the segment and the mesopleure coarsely obliquely striated.
Mesosternum transversely striated. Abdomen hardly as long as
the thorax and median segment combined, smooth and shining,
only four segments visible from above, the fifth segment beneath
very deeply emarginate, the sides almost encircling the apical
segment. ‘Tibie and tarsi not spinose. Black; the mandibles,
antenne, tibie, tarsi, posterior femora, and abdomen ferruginous.
Wings hyaline, the basal half tinted with fulvous, nervures
ferruginous. The stigma is larger than in the common H. stew-
dachneri Kohl, and the recurrent nervure is received just before
the apex of the first cubital cell.
Length 8 mm.
Hab. Cairns, Queensland (Dodd).
Described from three specimens in the British Museum.
HARPACTOPHILUS TRICOLOR, sp. n. (Plate XX VI. fig. 2.)
Q. Mandibles slender, bidentate, the innertooth short. Clypeus
smooth and shining, triangular, convex in the middle at the base,
with a small triangular truncation anteriorly. Front very broad,
with a short prominent carina above the base of the clypeus.
Antenne shorter than the head, the scape stout and nearly half as
longas the flagellum and less than half aslongas the distance between
the eyes at the base of the clypeus, the antenne at the base as far
from each other as from the eyes. The posterior ocelli nearer to
each other than to the eyes, situated on the vertex a little behind
the summit of the eyes and fully half as far again from the
posterior margin of the head as from each other. Front finely
longitudinally striated, most strongly just below the ocelli; vertex
almost smooth, very finely striated behind the eyes. Eyes not
quite reaching the base of the mandibles. The posterior margin
of the head strongly emarginate. Pronotum transverse, very
short, narrower than the head by about one-third, the anterior
angles acuteand prominent. Mesonotum punctured, shorter than
the median segment and a little longer than broad ; scutellum
short and broad, almost smooth. Median segment rectangular,
longer than broad, vertically truncate posteriorly, as broad as the
mesonotum, rugose; with a triangular space enclosed by carine
and divided by a longitudinal carina from the base to the apex,
the surface of the truncation coarsely transversely striated.
Abdomen smooth and shining, as long as the thorax and median
segment combined, the six dorsal segments all well defined.
Mesopleurz almost smooth. Legs without spines, except the
apical spines of the tibie. :
Black; the mandibles, clypeus, front, antenne, cheeks, the
head beneath, the tegule, and the anterior and intermediate legs
yellow; the abdomen and the posterior legs light ferruginous.
Wings hyaline, iridescent ; nervures testaceous.
The neuration is similar to that of H. steindachneri Kohl, but
1908. ] AUSTRALIAN FOSSORIAL WASPS. 463
the second cubital cell is not pointed on the radial nervure, being
about one-third as long on the radial as on the cubital nervure.
Length 5-6 mm.
Hab. Mackay, Queensland (Zwrner); September—December.
Described from two specimens.
This isa much slenderer species than /. steindachneri and is
much less coarsely sculptured. The pronotum is less depressed
and has the anterior angles strongly prominent, in these characters
showing an approach to the genus Spilomena.
PSENULUS INTERSTITIALIS Cam. (Plate X XVI. fig. 4.)
Psenulus interstitialis Cam. Tijdsch. v. Ent. xlix. p. 222, 1906.
Psen. luiescens Turner, Ann. & Mag. Nat. Hist. (7) xix. p. 273,
1907.
Hab. Etna Bay, New Guinea ; Mackay and Cairns, Queensland.
I have not seen Cameron’s type, but his description is quite
sufficient.
AMMOPHILA CLAVUS Fab.
Sphex clavus Fab. Syst. Ent. p. 348. n. 12, 1775.
@. Mandibles quadridentate; clypeus shining, very sparsely
punctured, with a few coarse black hairs, a broadly triangular,
oblique truncation at the apex, the apical margin very feebly and
rather broadly emarginate in the middle. Second joint of the
flagellum half as long again as the first and third combined.
Eyes parallel on the inner margins; the posterior ocelli nearly
half as far again from the eyesas from each other. Head opaque,
finely punctured ; a fine, impressed and slightly curved, transverse
line behind the posterior ocelli ; the front concave, with a delicate
sulcus from the anterior ocellus, clothed with rather sparse greyish
pubescence. Pronotum and mesonotum coarsely transversely
striated, the posterior half of the mesonotum with oblique strize
converging towards the middle of the posterior margin. Scutellum
longitudinally striated ; the mesopleurz and metapleure vertically
striate-rugose. Median segment transversely striated, the striz
somewhat oblique, the middle of the segment coarsely reticulate
and rather strongly convex at the base; the extreme apex of the
segment with a little short, pale, fulvous pubescence. Abdomen
shining, the four apical segmenis pruinose ; the petiole two-jointed,
equal in length to the posterior tibia and basal joint of the tarsus
combined. Tarsal ungues simple, without a pad; the comb of
the anterior tarsi with four slender spines on the outer margin of
the basal joint, excluding those on the apical process.
Black; the mandibles in the middle fusco-ferruginous; the
scape and two basal joints of the flagellum, the tegulz, the legs
(except the coxee and spines of the tarsi), the petiole and the first
following segment of the abdomen bright ferruginous; the base
of the second joint of the petiole black ; the four apical segments
steel-blue. Wings pale flavo-hyaline, lighter at the apex, nervures
testaceous.
Proc. Zoo, Soc.—1908, No. XXX. 30
464 MR. R. E. TURNER ON [May 12,
¢. Mandibles bidentate ; clypeus and front clothed with fine,
short, silvery pubescence ; the clypeus as long as the breadth at the
middle, broadly emarginate on the apical margin. Second joint
of the flagellum only slightly longer than the first and third com-
bined. Eyes rather strongly convergent towards the clypeus.
The whole of the mesonotum transversely striated ; mesopleure
and metapleure rugose. Petiole very long, equal in length to the
posterior tibia and three basal joints of the tarsus combined.
Black ; the tegule, the second joint of the petiole beneath, and
the first following segment of the abdemen at the base above and
beneath ferruginous ; the abdomen beyond the petiole pruinose,
dark steel-blue. Wings hyaline, clouded on the outer margin ;
nervures black.
Length, 2 23 mm., ¢ 22 mm.
Hab. Mackay to Cape York, Queensland (Zwurner); November
to May.
Very near A. instabilis Sm. in the female sex, but differs inthe
form of the clypeus and in the proportionate length of the joints
of the flagellum. The petiole isalsolonger. Also near A. basalis
Sm. from India.
A common species throughout the coastal districts of North
Queensland. The male is very different from those of other Aus-
tralian species, and allied to Indian forms.
AMMOPHILA AURIFERA, sp.n. (Plate XX VI. fig. 3.)
@. Mandibles with a very long acute apical tooth, the imner
margin with three teeth, the one nearest the apex very short and
broadiy truncate, the second broad and rounded, the basal one
very small and acute. Clypeus very slightly convex, depressed at
the apex and subtruncate, very sparsely punctured, the punctures
very large. Antenne inserted about two and a half times as far
from the eyes as from each other, about one-third further from
the anterior ocellus than from the apex of the clypeus; the second
joint of the flagellum about five times as long as the first and
twice as long as the third. Posterior ocelli about one-quarter
further from the eyes than from each other. Head opaque,
almost smooth, with a longitudinal sulcus from the anterior ocellus
to between the antenne; the front round the base of the antennee
slightly concave. Pronotum coarsely transversely striated ;
mesonotum very obscurely transversely striated, the strize almost
obsolete on the disc, a deep and broad longitudinal suleus from the
anterior margin not reaching the posterior margin, a raised curved
carina above the tegule. Mesopleure rugose; mesosternum
transversely striated. Scutellum longitudinally striated; median.
segment transversely striated, the striz arched in the middle and
rather obscure, as long as the mesonotum and pronotum combined.
Abdomen coriaceous, the petiole two-jointed and as long as the
posterior tibia and first tarsal joint combined ; the apical segment
with long black hairs on the sides. Tarsal ungues simple, not
bidentate, and without a pad.
1908. } AUSTRALIAN FOSSORIAL WASPS. 465
Bright golden ferruginous ; the head (except the basal half of the
mandibles), the scape, and five basal joints of the flagellum black ;
the mesosternum, the four apical abdominal segments, a spot on
the middle of the petiole, and part of the ventral surface of the
petiole and the next segment black. Wings pale flavo-hyaline,
very faintly tinged with fuscous at the apex ; nervures ferruginous.
Length 28 mm., exp. 29mm. Breadth of mesonotum between
the tegule 3 mm.
Hab. Port Darwin (Turner) ; December.
Allied to A. ardens Sm., but is a much less robust species and
less strongly sculptured.
The male has the mandibles bidentate, the clypeus longer than
broad, the second joint of the flagellum less than twice as long as
the third, the eyes convergent towards the clypeus, not parallel on
the inner margins as in the female, the mesonotum and median
segment much more strongly striated. The antenne are almost
wholly black, as are also the disc of the mesonotum and the sides
of the median segment; the apical joints of the posterior tarsi
are fuscous.
Length 24 mm.
AMMOPHILA EYRENSIS, Sp. n.
@. Mandibles quadridentate, the outer tooth very long and
acute. Clypeus short and broad, the apical margin depressed,
very shallowly emarginate in the middle; sparsely punctured and
with a few long black hairs. Antenne inserted close together ;
the second joint of the flagellum more than twice as long as the
third, which is twice:as long as the first. Eyes parallel on the
inner margin, the posterior ocelli a little further from the eyes
than from each other. Head subopaque, almost smooth, a very
fine transverse sulcus behind the posterior ocelli and not extending
beyond them, a longitudinal frontal sulcus from the anterior
ocellus, the front between the base of the antenne and the
eyes clothed with short, thin, cinereous pubescence. Pronotum
coarsely, mesonotum more finely transversely striated ; a longi-
tudinal median sulcus on the anterior half of the mesonotum, in
which lies a short longitudinal carina. Scutellum longitudinally
striated ; the mesopleure and metapleure rugose-striate, the
mesopleure with a short, fine and shallow, vertical sulcus below
the anterior wings. Median segment transversely striated, the
strie slightly oblique, coarsely reticulate along the middle, broadly
at the base and narrowly at the apex. Abdomen shining and
almost smooth, the apical segments not pruinose; the petiole two-
jointed, the first joint seen from above very slightly longer than
the second, the twe combined a little longer than the posterior
tibia and the basal joint of the tarsus combined. Tarsal ungues
simple, without pads, the comb of the anterior tarsi rather long,
with four spines on the basal joint, excluding those on the apical
process.
Light ferruginous; the head black; the mandibles except at
the apex, the basal half of the antenne and the mouth-parts
30*
466 MR, R. E. TURNER ON [May 12,
ferruginous ; the posterior half of the mesonotum, a median
longitudinal line on the anterior half, and the spines and ungues
of the tarsi black; the three apical abdominal segments shining
steel-blue; a spot at the base of the second joint of the petiole
black. Wings hyaline, the base of the anterior pair pale flavo-
hyaline; nervures testaceous.
The second and third cubital cells are equal in length on the
radial nervure.
Length 18 mm.
Hab. Killalpanima, 8. Australia, 100 miles east of Lake Eyre
(H. J. Hillier).
Type in British Museum. Described from two specimens.
Near A. instabilis Sm., but is a smaller and more slender
insect; the second joint of the flagellum is longer in, proportion,
and the dorsal surface of the median segment is not sharply
raised towards the median line as in typical instabilis.
AMMOPHILA INSTABILIS Sm.
Ammophila instabilis Sm. Cat. Hym. B. M. iv. p. 214. n. 36,
1856, 2.
Ammophila impatiens Sm. Trans. Ent. Soc. London, p. 247.
n. 1, 1868, ¢ (as 2).
These are without much doubt the sexes of one species.
Hab. Champion Bay, W. Australia.
AMMOPHILA ARDENS Sm.
Ammophila ardens Sm. Trans. Ent. Soc. London, p. 247. n. 2,
1868.
Hab. Swan River (Du Boulay), Mackay, Queensland (7'urner).
AMMOPHILA SUSPICIOSA Sm,
Ammophila suspiciosa Sm. Cat. Hym. B. M. iv. p. 214. n. 38,
1856. |
_ This is the only Australian species known in which the petiole
is one-jointed. It is, as Smith points out, scarcely distinct from
the N.-African species 4. argentea Brulle.
Hab. Melbourne; Lake Eyre district; Perth ; Tasmania.
SpuEx (IsoDONTIA) ALBOHIRTUS, sp. 0.
2. Mandibles broad and tridentate. Clypeus rather sparsel
punctured, clothed with very short, close pubescence which hee
as silver in some lights, with sparse, long, brown hairs; slightl
convex at the base and subcarinate, very broadly and ‘halle
emarginate at the apex, with a small and narrow etinngenente
in the middle of the apical margin, the angles of the emargination
produced into minute spines. Second joint of the flagellum as
long as the third and half of the fourth. Eyes slightly con-
vergent towards the clypeus, separated on the vertex by A ace
equal to the length of the third and fourth joints of the fla sien
combined, and by nearly the same distance on the biyadeet the
1908. | AUSTRALIAN FOSSORIAL WASPS. 467
posterior ocelli a little nearer to each other than to the eyes.
Head shining, finely and rather sparsely punctured, the front
with very short silvery pubescence, the sparse long hairs on the
vertex brownish grey. Pronotum transverse and vertically
depressed ; the mesonotum sparsely, the mesopleure closely punc-
tured ; scutellum flat. Median segment punctured-rugose, with a
short longitudinal sulcus near the apex, without a sulcus from the
stigma. Petiole as long as the basal joint of the posterior tarsus ;
the pubescence on the petiole and the sides of the median segment
long and whitish, on the dorsal surface of the median segment
very pale brown and very sparse. Abdomen shining and almost
smooth, the first segment as long as the second, the apical segment
finely punctured and subopaque. The first recurrent nervure is
received near the apex of the second cubital cell, the second near
the base of the third cubital cell; the second cubital cell is very
large, subrectangular, more than half as long again on the cubital
nervure as high.
Black; the abdomen with obscure blue reflections. Wings
dark fusco-hyaline flushed with purple, nervures black. A patch
of short fulvous pubescence at the apex of the posterior tibie.
¢. Similar to the female, but the emargination on the middle
of the apical margin of the clypeus is very faintly indicated; the
abdomen is pruinose.
Length, 9 21 mm., ¢ 19 mm.
Hab. Mackay and Cairns, Queensland (Zurner); December to
March.
The wings are rather paler in Cairns specimens.
Allied to S. ustulatus Kohl, from Timor, also to S. morosus
Sm. and S. praslinius Guér. The last two may prove to refer
to one species. The emargination of the clypeus and the pale colour
of the pubescence will serve to distinguish the present species.
SpHEX (IsoponTIA) NIGELLUS Sm.
Sphex nigella Sm. Cat. Hym. B. M. iv. p. 255, 1856.
Hab. Mackay and Cairns, Queensland (Z'urner).
This species seems to occur throughout Southern Asia and also
in W. Australia.
SpPHEX (IsoDONTIA) OBSCURELLUS Sm.
Sphex obscurella Sm. Cat. Hym. B. M. iv. p. 251, 1856.
Hab. Tasmania.
Very near S. nigellus, from which it may be distinguished by
the much shorter petiole and by the shape of the third cubital
cell, which is as long as the second on the radial nervure; the
elvypeus is also rather longer.
SPHEX WALLACEI, nom. nov.
Sphex nitidiventris Sm. Proc. Linn. Soc., Zool. ii. p. 158, 1859
(nec Spinola).
Kohl states that nitidiventris Spin. also occurs in Java and
468 MR. R. E. TURNER ON [May 12,
Luzon, though a 8.-American species. The species in the British
Museum which is identified as nitidiventris Spin., rightly in my
opinion, though somewhat resembling Smith’s species, has the
petiole much longer and the clypeus different.
Hab. Ayu (Wallace); Mackay to Cape York (Turner).
A common species in North Queensland.
SPHEX GILBERTI, sp. n.
2. Clypeus convex, triangularly flattened from the middle to
the apex, the apical margin very feebly and narrowly emarginate
in the middle, the length equal to about three-quarters of the
breadth at the apex; sparsely and rather coarsely punctured, with
a feeble longitudinal carina from the base not reaching the middle,
very sparsely clad with long, stiff, black hairs, the sides in some
lights showing silvery reflections on very short, fine pubescence.
Front clothed with very short, fine pubescence, silvery below the
base of the antenne, golden above, with very sparse, long black
hairs above the base of the clypeus ; the vertex bare, opaque, and
almost smooth. Antenne twice as far from the eyes as from
each other; the first joint of the flagellum very short, the second
and first together about equal in length to the third and fourth.
The inner margins of the eyes almost parallel, converging slightly
on the vertex ; the posterior ocelli are nearer to each other than
to the eyes. Pronotum slightly depressed below the level of the
mesonotum, very steeply sloped anteriorly, opaque and smooth,
the posterior margin with very short silvery pubescence. Thorax
opaque, very finely and closely punctured ; the mesonotum slightly
depressed anteriorly in the middle and with an obscure longi-
tudinal suleus on each side above the tegule; the scutellum flat,
with a very faint longitudinal line in the middle; postscutellum
transversely depressed at the base; the mesopleure subopaque
and very shallowly punctured. Median segment opaque, sparsely
punctured, with sparse, stiff, blackish hairs; an obscure longitu-
dinal sulcus from the base to the apex, the dorsal surface bordered
by a very narrow shallow sulcus ; a deeper and broader sulcus, in
which are regular transverse striz, on the metapleure from the
stigma to the posterior angle. Abdomen shining, sparsely and
very finely punctured ; the epipygium coarsely punctured, with
sparse and very long black hairs, and convex; the petiole not
quite equal in length to the third joint of the posterior tarsi.
Tarsal ungues with two small teeth near the base ; the spines of
the anterior tarsus long. The length of the third cubital cell on
the radial nervure is about equal to the distance of the first
transverse cubital nervure from the stigma; the first recurrent
nervure is received at about one-fifth from the apex of the second
cubital cell, the second at two-fifths from the base of the third
cubital cell,
Black; the abdomen steel-blue. Wings flavo-hyaline, broadly
pale fusco-hyaline at the apex; nervures ferruginous.
Length 26-28 mm.
1908. | AUSTRALIAN FOSSORIAL WASPS. 469
Hab, Mackay, Queensland (Z'urner); February.
Very near S. diabolicus Sm., the Oriental form of rufipennis
Fab., but differs in the postscutellum, which is not longitudinally
divided or raised into a small tubercle on each side near the
middle ; the petiole also is rather shorter, and the abdomen is
shining instead of opaque.
CERCERIS INEXPECTATA, sp.n. (Plate XXVI. fig. 5.)
2. Mandibles blunt at the apex, with a large triangular tooth
on the inner margin nearer to the base than to the apex. Clypeus
sparsely punctured, broadly truncate at the apex, the central lobe
slightly convex, bluntly produced on the apical margin at the
junction of the lateral lobes, nearly twice as broad at the apex as
at the base. The antenne are inserted above the clypeus, at a
distance from its base equal to three-quarters of the greatest
length of the clypeus; the second joint of the flagellum is at
least twice as long as the first and half as long again as the third ;
at the base the antenne are separated from the eyes bya distance
equal to the length of the scape, and from each other by about
two-thirds of that distance. The eyes are slightly convergent
Text-fig. 107.
Cerceris: pygidial area of 9.
1. C. inexpectata. 2. C. labeculata. 3. C. predura. 4. C. multiguttata.
towards the vertex; the posterior ocelli are about one-third
further from the eyes than from each other, and as far from the
posterior margin of the head as from the eyes. Head very broad,
at least one-third broader than the mesonotum, the vertex coarsely
punctured-rugose, the front sparsely and shallowly punctured,
with an elevated carina between the antenne. Pronotum very
short, strongly rounded at the angles; mesonotum rather sparsely
punctured, with a very shallow sulcus from the anterior margin
not reaching the centre; mesopleure very coarsely punctured ;
seutellum short, very broadly truncate at the apex. Median
segment strongly but not very closely punctured at the sides, with
a suleus from the base to the apex; the triangular space at the
base smooth. Abdomen coarsely punctured, most closely on the
third and fourth segments ; the first segment rectangular, a little
broader than long, nearly half as broad as the apex of the second
segment ; the dorsal pygidial area rugose, truncate at the apex,
where it is half as broad as at the base.
47() MR. R. E.“TURNER ON [May 12,
Black ; the base of the mandibles, clypeus, front as high as the
base of the antenne, the frontal carina, the sides of the pronotum,
the postscutellum, a large spot on each side occupying the whole
length of the median segment, the sides and apex of the first
abdominal segment, the second segment except a triangular mark
at the base, the apex of the fifth segment very broadly, and the
two basal ventral segments yellow; the antenne, the outer orbits
of the eyes connected with a broad band on the posterior margin
of the head, a large spot on each side occupying the greater
part of the mesonotum, the scutellum, the apex of the fifth ventral
segment, and the legs dull ferruginous; the coxe above black.
Wings pale flavo-hyaline, nervures light testaceous, a faint clouc
in the radial cell.
Length 14 mm., exp. 24 mm.
Hab. Mackay, Queensland (Turner); November.
CERCERIS LABECULATA, Sp. 0.
¢. Mandibles simple, blunt at the apex. The central lobe of
the clypeus slightly convex and feebly porrected at the apex,
narrowly emarginate anteriorly, the angles of the emargination
produced into short teeth, narrowly truncate at the base, the
length almost as great as the greatest breadth. Antenne inserted
high up on the front, nearer to each other than to the eyes, from
which they are separated by a distance less than the length of the
scape, which is short ; the second and third joints of the flagellum
nearly equal in length. The antenne are separated from the
base of the clypeus by a distance equal to two-thirds of the length
of the clypeus. Inner margin of the eyes parallel; the posterior
ocelli nearly as far from each other as from the eyes and nearer
to the posterior margin of the head than to each other. Front
sparsely and shallowly punctured, the carina between the antennse
not very strongly raised; vertex, mesonotum, mesopleure, and
sides of the median segment coarsely punctured-rugose. Pro-
notum strongly rounded at the angles; the triangular space at
the base of the median segment finely obliquely striated. First
abdominal segment longer than broad, depressed at the base; the
abdomen deeply and closely punctured ; the pygidial area finely
punctured, narrowly truncate at the apex, where it is only half
S wide as at the base, rather broader in the middle than at the
ase.
Black; the mandibles at the base, the clypeus except the base
and the extreme apical margin, the apex of the interantennal
carina, the inner orbits of the eyes as high as the base of the
antenn very broadly, and the scape of the antennse pale yellow ;
the posterior margin of the pronotum interrupted in the middle,
the postscutellum, the apical margin of the first, second, and
fourth abdominal segments, and a short vertical streak on the
mesopleuree, orange-yellow ; the flagellum, tegule, scutellum, first
abdominal segment, the middle of the second, the apex of the
fifth, the pygidium, and the legs (except the coxee, trochanters,
1908.} AUSTRALIAN FOSSORIAL WASPS. 471
and base of the femora) ferruginous. Wings hyaline, stained
with fuscous along the costa, nervures black.
3. As in the female, but the clypeus is rather longer and not
porrect at the apex, and the apical half of the fifth abdominal
segment is yellow.
Length, 2 9mm., ¢ 8 mm.
Hab. Cairns, Queensland (Zurner) ; December and January.
Somewhat related to C. antipodes Sm., but the head is much
narrower and it isa much smaller species. The clypeus is also
very different.
CERCERIS MULTIGUTTATA, Sp. Nn.
©. Clypeus more than twice as broad as long on the central
lobe, short, shghtly porrect at the apex, the apical margin very
slightly and broadly emarginate, with four minute teeth. Antenne
inserted nearer to each other than to the eyes, separated from the
eyes by a distance equal to three-quarters of the length of the
scape and about the same distance from the base of the clypeus;
the second joint of the flagellum twice as long as the first and
half as long again as the third. The inner orbits of the eyes
very nearly parallel; the posterior ocelli a little further from the
eyes than from each other, and about the same distance from the
posterior margin of the head as from each other. Head, thorax,
and median segment coarsely and closely punctured, the frontal
carina strongly raised between the antenne. Pronotum broadly
rounded anteriorly ; scutellum very broadly truncate at the apex.
Median segment with a shallow sulcus from the base to the apex,
the triangular space at the base smooth and shining. Abdomen
coarsely punctured, the fourth and fifth more shallowly than the
basal segments; the first segment broader in the middle than
long, slightly narrowed to the base and apex and obliquely de-
pressed at the base, not as long as the second segment. Pygidial
area finely rugose, twice as long as broad, the sides nearly parallel,
broad]y rounded at the apex.
Black ; the mandibles at the base, clypeus, outer orbits of the
eyes, the inner orbits broadly to a little above the base of the
antenne, a narrow line from the anterior ocellus to the base of
the clypeus, an oblique spot on each side of the vertex, the pro-
notum narrowly interrupted in the middle, the tegule, a spot on
the mesopleure, a large round spot on each side of the scutellum,
the postscutellum, a small spot on each side on the triangular
space at the base of the median segment, a large spot on each side
at the apex extending on to the sides, the first abdominal segment
except at the base, the apical half of the second segment, the
three apical abdominal segments (narrowly black at the base), the
tibie, tarsi, and the apex of the femora all dull brownish yellow.
Wings hyaline, tinged with fulvous, darker on the costa; nervures
ferruginous.
Length 14 mm.
472 MR. R. E, TURNER ON [May 12,
a
The male is similar to the female, but the central lobe of the
clypeus is nearly as long as broad, truncate at the apex, and the
yellow spots on the vertex are smaller.
Length 9 mm,
Hab. Mackay, Queensland (Zurner); October.
CERCERIS PREDURA, Sp. 0.
Q. Clypeus depressed on the anterior margin, widely and
shallowly emarginate, the central lobe twice as broad at the apex
as long. Antenne inserted half as far again from the eyes as
from each other, the distance between them and the eyes equal to
a little less than the length of the scape, the distance from the
base of the clypeus nearly equal to the length of the scape and
the first joint of the flagellum combined ; the second joint of the
flagellum is twice as long as the first and not quite half as long
again as the third. The inner orbits of the eyes nearly parallel ;
the posterior ocelli about one-third further from the eyes than
from each other, but as far from each other as from the posterior
margin of the head. Clypeus and front shallowly punctured, the
rest of the insect, except the scutellum, postscutellum, base of
the median segment, and pygidium, very coarsely and deeply
punctured. Pronotum very broadly and feebly rounded ante-
riorly, less deeply punctured than the mesonotum; scutellum
transverse, shallowly punctured; postscutellum smooth. The
triangular space at the base of the median segment is divided by
a suleus and is deeply but rather sparsely punctured. First
abdominal segment about half as long again as broad, not as long
as the second segment; pygidial area granulate, elongate-oval,
and very narrowly truncate at the apex.
Black ; the mandibles (except at the apex), the clypeus, front
below the base of the antenne, scape, the carina between the
antenne, the pronotum broadly interrupted in the middle, the
tegule, scutellum, postscutellum, the base of the second abdo-
minal segment broadly, the third segment (with the base broadly
black in the middle), the fifth segment above, and the legs (except
the base of the coxe) yellow; the flagellum, the apex of the first
abdominal segment, the pygidium, and the posterior femora
ferruginous. Wings hyaline, the radial cell and the apex beyond
it clouded with fuscous ; nervures fusco-ferruginous.
3. Asin the female, but the clypeus is longer, being as long
as the breadth of the central lobe at the apex and without the
shallow emargination, the base a little nearer to the antenne
than in the female. The scutellum is black; the first abdominal
segment ferruginous; the apex of the fifth and the whole of the
sixth yellow; the femora and the apex of the posterior tibiz
ferruginous.
Length, 2 7mm., ¢ 6 mm.
Hab. Mackay, Queensland (Zwrner); April and May.
Allied to O. predator Sm. from Celebes, but the sculpture is
different, also the shape of the pygidial area.
1908. | AUSTRALIAN FOSSORIAL WASPS. 473
CERCERIS VENUSTA Sm.
Cerceris venusta Sm, Ann. & Mag. Nat. Hist, (4) xii. p. 413,
1873.
Hab. Mackay and Cairns, Queensland (Zwrner).
A species very variable in colour. In specimens from Cairns
the yellow markings are much reduced in size.
In the male the yellow marks on the median segment are
sometimes wholly absent. The posterior ocelli in this species
are very near together, about twice as far from the eyes as from
each other.
CERGERIS AUSTRALIS Sauss.
Cerceris australis Sauss. Mém. Soc. Phys. & Hist. Nat. Geneva,
xiv. p. 1, 1854.
Hab. Tasmania (Saussure); Melbourne to Mackay (Z%rner).
LIRIS HHMORRHOIDALIS Fab.
Liris magnifica Kohl, Verh. zool.-bot. Ges. Wien, xxxiii. p. 356,
1883.
Though the colour of the pile on the female is more brilliant
and the sculpture on the median segment rather stronger in
females from Australia, I cannot detect any appreciable difference
in the male.
Hab. Mackay to Cape York, Queensland (Zurner).
LARRA NIGRIPES Sauss.
Larrada nigripes Sauss. Reise d. Novara, Zool. 11. p. 1, Hym.
p. 74, n. 3, 1867.
Larra psilocera Kohl, Verh. zool.-bot. Ges. Wien, xxxili, p. 355,
1883.
Hab. Tasmania (Saussure) ; Adelaide, 8. Australia; Mackay,
Queensland ; Adelaide River, Northern Territory.
Saussure’s description seems to have been taken from a head-
less specimen, but I think there is little doubt that it is identical
with Kohl's species. It is allied to Z. mansueta Sm. from New
Guinea.
LARRA FEMORATA Sauss.
Tachytes femoratus Sauss. Mém. Soc. Phys. et Hist. Nat.
Geneva, xiv. p. 1, p. 20, n. 9, 1854.
Hab. Sydney, N.S.W.; Mackay and Cairns, Queensland.
North Queensland specimens differ a little from southern ones
in the shape of the pygidium, pronotum, and third cubital cell ;
the antenne also are a little stouter. But the differences are
very slight and do not seem sufficient to merit specific rank.
L. rujipes Sm. from Celebes and Z. mendax Sm. from Halmaheira
are slight geographical variations of the same species.
A774 MR. R. E, TURNER ON [May 12,
LarRA SCELESTA, Sp. 0.
@. Clypeus very finely and closely punctured, the anterior
margin shining and very slightly and broadly rounded. Mandibles
strongly notched on the outer margin, very blunt at the apex,
with a fringe of short stiff hairs on the outer margin. Antenne
inserted nearly as far from each other as from the eyes; the
second joint of the flagellum only a little more than half as long
again as the first, the third intermediate in length between the
first and second, Eyes separated on the vertex by a distance not
quite half as great again as the length of the second joint of the
flagellum. Labrum bilobed. Front smooth and shining, with a
longitudinal furrow on each side, the anterior ocellus lymg in a
broad and shallow transverse depression ; the posterior ocelli very
indistinct, with an irregular, shining, transverse depression behind
them; the vertex minutely punctured, Thorax very finely and
closely punctured, the pronotum longer and less abruptly depressed
than in LZ. nigripes Sauss. Median segment much longer than
the mesonotum, longer than in ZL. nigripes, finely rugose, with a
carina from the base to the apex, most strongly marked at the
base, vertically truncate posteriorly, the face of the truncation
finely rugulose, with a median suleus. Abdomen shining, very
shallowly and minutely punctured, the three basal segments with
a band of white pubescence on the sides; the pygidial area
broadly triangular, rather sparsely but deeply punctured. Anterior
tibiz without spines. The second cubital cell is as long as the
third on the radial nervure; the third is nearly three times as
long on the cubital as on the radial nervure, the third transverse
cubital is oblique and almost straight.
Black ; the mandibles and the spines of the tibie fusco-ferru-
ginous ; tegul testaceous. Wings fusco-hyaline, faintly iridescent;
nervures black.
Length 14 mm.
Hab, Mackay, Queensland (Zurner); Adelaide River, Northern
Territory (J.J. Watker). ..
Differs from L. nigripes in the shorter antennz with differently
proportioned joints, in the much closer approach of the eyes on
the vertex, the longer pronotum and median segment, and the
broader pygidium. The pronotum and abdomen differ markedly
from those of Z. australis, with which it agrees in the smooth
anterior tibie, which are heavily spined in ZL. nigripes.
LARRA AUSTRALIS Sauss.
_Tachytes australis Sauss. Mém. Soc. Phys. et Hist. Nat. Geneva,
xlv. p. 19, 1854.
If my identification of this species is correct it is a Larra.
flab. 8. Australia; W. Australia.
LARRA PACIFICATRIX, sp. n.
oe
2. Clypeus very broadly truncate at the apex, opaque and
almost smooth, with a row of large punctures before the apical
1908. ] AUSTRALIAN FOSSORIAL WASPS. 475
margin. Second joint of the flagellum more than half as long
again as the first, the second and third subequal and rather
shorter than the fourth. Head subopaque, almost smooth, the
. front shining, the margins of the eyes broadly bordered with
briliant golden pubescence ; the eyes separated on the vertex by
a distance equal to three-quarters of the length of the second
joint of the flagellum. Pronotum depressed below the level
of the mesonotum, much higher in the middle than at the sides,
almost vertical. Mesonotum and scutellum shining and almost
smooth, only microscopically but very closely punctured ; a large
patch of short golden pubescence on the mesopleure below the
base of the anterior wings. Median segment longer than broad,
opaque, rather indistinctly transversely striated, the strie more
obscure towards the apex, a median carina from the base reaching
beyond the middle, vertically truncate posteriorly, the face of the
truncation coarsely transversely striated, with a deep median
suleus. Abdomen smooth and shining, highly polished, the
pygidial area closely punctured; the second ventral segment
strongly convex, flattened on the sides at the base and with
a longitudinal carina from the base not reaching the middle,
the apex of the first segment narrowly obliquely striated. The
second cubital cell is pointed on the radial nervure, and about
equal to the third in length on the cubital nervure; the two
recurrent nervures are received before the middle of the second
cubital cell, the first much further from the base of the cell than
from the second. The tarsal ungues are long.
Black ; the clypeus, scape, first joint of the flagellum, mandibles,
and legs ferruginous; tegule testaceous. Wings fusco-hyaline,
with a slight purple gloss and tinted with yellow; nervures
black.
Length 17 mm., exp. 28 mm.
Hab. New Hebrides.
Type in Oxford University Museum ex coll. Saunders.
This species is very near Votcgonia.
NovroGONIA CHRYSONOTA Sm.
Larrada chrysonota Sm. Trans. Ent. Soc. London, p. 304,
1869.
Larrada crassipes Sm. Ann. & Mag, Nat. Hist. (4) xii. p. 294,
1873.
Larra chrysonota Kohl, Verh. zool.-bot. Ges. Wien, xxxiv
p. 242, 1884.
Hab. Champion Bay, W. Australia ; Adelaide, 8. Australia.
NOTOGONIA REGINA, sp.n. (Plate X XVI. fig. 7.)
2. Clypeus opaque, shining on the apical margin, which is
slightly depressed, almost straight, very narrowly and shallowly
emarginate in the middle; a carina from the base not reaching
the apex. Head and thorax opaque, the front strongly raised on
476 MR. R. E. TURNER ON | May 12,
the inner orbits of the eyes, with a longitudinal median suleus ;
the anterior ocellus lying in a broad depression which almost
reaches the eyes; on the vertex the eyes are separated by a
distance less than twice as great as the length of the first joint
of the flagellum. The second joint of the flagellum is more than
twice as ‘long as the first and nearly half as long again as the
third: the fourth and fifth joints about equal to the third.
Pronotum depressed very much below the mesonotum, very
steeply sloped anteriorly ; mesonotum with a rather broad median
depression from the anterior margin to the middle. Median
segment a little longer than the mesonotum, nearly half as long
again as broad, obscurely transversely striated, the striz almost
obsolete at the apex, vertically truncate posteriorly, the face of
the truncation more strongly transversely striated, the sides
of the segment indistinctly striated. Abdomen subopaque, the
apical margins of the segments broadly depressed. Pygidial area
pubescent, with an obscure median carina, the lateral carinz
strongly raised, separated at the apex by a distance not quite
equal to the length of the first joint of the flagellum. Anterior
tibizee smooth, intermediate and posterior tibie with a double row
of three or four widely separated and feeble spines ; tarsi spinose,
the ungues simple and very long.
Black; the antenne orange; the pile on the head and abdomen
silvery, the abdominal segments with broad bands of pile on the
apical margin ; a pale golden sheen on the pile on the front and
a small patch of golden pubescence at the apex of the posterior
tibie. Wings fusco-hyaline, the apex of the anterior wings and
a broad band across beyond the middle fuscous, the apex of the
posterior wings broadly pale fuscous. Nervures black; testaceous
at the apex of the radial and third cubital cells.
The recurrent nervures are both received before the middle of
the second cubital cell, the distance between them not exceeding
one-sixth of the length of the cell on the cubital nervure. The
third cubital cell is about four times as long on the cubital as on
the radial nervure.
Length 19 mm., exp. 27 mm.
Hab. Cairns, Queensland (Zurner); December to March.
Mackay, Queensland ; October. Cape York, Queensland; May.
The specimen from Cape York has the wings fuscous from the
base, with a large hyaline patch near the apex of the anterior
wings; the four apical joints of the flagellum are black and the
spines on the tibiz stronger.
The specimens from Mackay are smaller, measuring only 12—
14 mm., the strie on the median segment are almost entirely
absent, except on the face of the truncation, and there is a delicate
longitudinal carina from the base almost reaching the apex. The
six apical joints of the flagellum are black.
NOTOGONTA BASILISSA, Sp. N.
2. Mandibles notched rather deeply on the outer margin; the
1908. | AUSTRALIAN FOSSORIAL WASPS. ATT
clypeus opaque, very minutely punctured, clothed with rather
coarse silvery pubescence, the apical margin transverse, very
feebly emarginate in the middle and broadly smooth and shining.
The second joint of the flagellum is a little less than twice as long
as the first, a little longer than the third, and a little shorter than
the fourth. Head opaque, the front clothed with rather coarse
silvery pubescence; the eyes separated on the vertex by a distance
rather less than the third joint of the flagellum. Pronotum
steeply, but not nearly vertically sloped, higher in the middle
than at the sides; the mesonotum broadly but shallowly de-
pressed anteriorly, with sparse pubescence of a dull silver colour
on the sides and in thedepression. Median segment opaque, with
a short longitudinal carina from the base not reaching the middle,
very finely and closely punctured and very sparsely covered with
short grey pubescence, a few short transverse striz near the apical
angles, the face of the posterior truncation very shallowly and
irregularly transversely striated, with a deep and broad median
sulcus, the sides of the segment almost smooth, delicately obliquely
striated at the base; the segment much longer than broad. The
four basal abdominal segments with broad bands of rather dull
silvery pubescence on the apical margin ; the pygidial area rounded
at the apex, rather broad, and clothed with shining whitish
pubescence; the second and third ventral segments convex, the
second subcarinate at the base. The distance between the two
recurrent nervures is less by about one-third than that between
the first recurrent nervure and the base of the second cubital cell;
the third cubital cell is more than half as long again as the second
on the cubital nervure.
Black ; the five or six apical joints of the flagellum, the apex of
the joints of the tarsi, and the whole of the apical joint beneath
fulvous ; tegule testaceous. Wings subhyaline, nervures fusco-
ferruginous.
Length 13 mm.
Hab. Mackay, Queensland (Zwurner); March.
NOTOGONIA AGITATA, Sp. 0.
Q@. Clypeus opaque, clothed with silvery pubescence, the apical
margin broadly smooth and shining, transverse. Second joint of
the flagellum a little more than half as long again as the first, the
second and third subequal, shorter than the fourth. Head opaque,
the front clothed with short silvery pubescence; the eyes separated
on the vertex by a distance equal to three-quarters of the length
of the second joint of the flagellum. Mesonotum closely and
microscopically punctured, longitudinally and rather broadly
depressed in the middle anteriorly, with a short and fine longi-
tudinal sulcus in the depression, the disc with sparse and very
short cinereous pubescence, the sides with closer pubescence of a
silvery colour slightly tinged with pale golden. Median segment
opaque, very finely granulated, much longer than broad; the
surface of the posterior truncation strongly transversely striated,
478 MR. R. E. TURNER ON | May 12,
with a deep median sulcus; a very obscure longitudinal carina
from the base of the segment not reaching the middle. Abdomen
subopaque; the four basal segments with broad apical bands
of silvery pubescence ; the pygidial area narrowly truncate at the
apex, clothed with silvery pubescence ; the second ventral segment
smooth and convex, with a longitudinal carina on the basal third.
The third cubital cell is half as long again as the second on the
radial nervure and as long as the second on the cubital nervure ;
the two recurrent nervures are received by the second cubital
cell before the middle, the first more than twice as far from the
base of the cell as from the second.
Black; the scape beneath fusco-ferruginous; the tegule and
the apical margin of the fifth abdominal segment testaceous, the
tarsi beneath, the apical joint of the tarsi above, and the tarsal
ungues ferruginous. Wings hyaline, the outer margin bordered
with pale fuscous, iridescent ; nervures fusco-ferruginous.
Length 10-11 mm.
Hab. Mackay and Cairns, Queensland (Zurner) ; December to
May.
NOTOGONIA SERENA, Sp. 0.
2. Mandibles very feebly notched on the outer margin; the
clypeus very broadly rounded at the apex, clothed densely with
rather coarse silver-grey pubescence, almost flat. Scape clothed
with short, shining pubescence, the apex beneath smooth and
shining ; the first joint of the flagellum half as long as the second,
the fourth a little longer than the third and about equal in length
to the second. Hyes separated on the vertex by a distance equal
to the length of the third joint of the flagellum. Head opaque ;
the pubescence on the front silver-grey and rather coarse, that on
the broad depressed space round the anterior ocellus pale golden-
brown. Pronotum very sharply sloped, almost vertical, much
higher in the middle than at the sides; mesonotum very minutely
punctured-rugulose ; clothed with pubescence, which is rather
coarse and silver-grey flushed with pale golden on the sides,
darker and shorter on the middle ; longitudinally depressed from
the middle to the anterior margin. Median segment longer than
broad, with a delicate longitudinal carina from the base to the
middle, rather obscurely transversely striated, the striz most
distinct at the base and the sides, more opaque than the thorax,
vertically truncated posteriorly, the face of the truncation in-
distinctly transversely striated, with a low carina from the base
not quite reaching the apex, the apical margin of the segment at
the base of the truncation slightly raised. Abdomen opaque,
pruinose, the apical margins of the four basal segments with broad
bands of silvery pubescence, which is coarser on the sides than in
the middle; the pygidial area more than twice as long as the
breadth at the base, about half as wide at the broadly rounded
apex as at the base, clothed with rather coarse golden pubescence.
The two recurrent nervures are received very close together before
the middle of the second cubital cell, the first is shghtly curved
1908. ] AUSTRALIAN FOSSORIAL WASPS. 479
outwardly near its apex, the second very strongly curved out-
wardly near its base; the second cubital cell is half as long on
the radial nervure as the third.
Black; the apex of the fifth abdominal segment and of the
pygidium testaceous, the tegule testaceous brown. Wings hyaline,
the apex very faintly tinged with fuscous; nervures fusco-
ferruginous.
Length 15 mm.
Hab. Mackay, Queensland (Zurner); April.
NOTOGONIA OBLIQUETRUNCATA, Sp. n.
2. Mandibles rather deeply incised on the outer margin ; the
clypeus opaque at the base, with a carina from the base to the
middle, clothed with short, fine, silvery pubescence, very broadly
rounded or almost truncate at the apex, the apical margin broadly
smooth and shining. The second joint of the flagellum equal in
length to the third and more than twice as long as the first.
Hyes separated on the vertex by a distance equal to the length of
the second and half of the first jomts of the flagellum. Head
opaque, the front clothed with very short silvery pubescence.
Pronotum very steeply sloped, much higher in the middle than at
the sides; the mesonotum microscopically punctured, depressed
in the middle anteriorly, the sides and posterior margin clothed
with very pale golden pubescence. Median segment opaque, with
short transverse striz at the apical angles, vertically truncated
posteriorly, the face of the truncation transversely striated, with
a longitudinal sulcus from the base to the apex. Abdomen sub-
opaque, the apical margins of the four basal segments with bands
of silver-grey pubescence, broadly interrupted in the middle on
the fourth; the pygidial area almost pointed at the apex, long
and narrow, shining and sparsely punctured at the base, finely
punctured and clothed with short, pale, fulvous pubescence at the
apex. The first recurrent nervure is received almost as far from
the second as from the base of the second cubital cell; the second
is moderately rounded outwardly. The second cubital cellis much
longer than the third on the cubital nervure and equal to it in
length on the radial nervure, the third transverse cubital nervure
oblique and scarcely curved, the radial cell short and broad,
obliquely truncate at the apex.
Black; the mandibles, the scape beneath, and the anterior
femora and tibize beneath fusco-ferruginous, the apical margin of
the abdominal segments narrowly testaceous. Wings hyaline,
faintly tinted with fuscous at the apex, most strongly in the
radial cell; nervures black, the stigma fusco-ferruginous.
Length 10 mm.
Hab. Port Darwin (Turner); November.
NOoOTOGONIA RETIARIA, sp. n.
2. Mandibles shining, deeply notched on the outer margin.
Clypeus transverse, clothed with silvery pubescence, almost flat,
Proc. Zoou. Soc.—1908, No. XX XT. 31
480 MR. R. E, TURNER ON [May 12,
the centre very slightly convex, the anterior margin bare and
shining, feebly produced in the middle, and narrowly and shallowly
emareginate. Antenne inserted at the base of the clypeus, a little
nearer to each other than to the eyes; the second joint of the
flagellum fully twice as long as the first, slightly longer than the
third, about equal in length to the fourth, but a little shorter than
the fifth. Eyes separated on the vertex by a distance equal to
the length of the fifth joint of the flagellum. Head subopaque,
very minutely and closely punctured; the front with a longi-
tudinal depression on each side, the orbit of the eyes elevated,
the anterior ocellus lying in a broad depression, the posterior
ocelli indistinct, with a triangular depression behind them. Pro-
notum higher in the middle of the posterior margin than at the
sides, the middle on a level with the mesonotum, steeply but not
vertically sloped anteriorly. Mesonotum microscopically and
very closely punctured and opaque, the mesopleuree and scutellum
subopaque, the former more distinctly punctured. Median segment
scarcely longer than the mesonotum, a little longer than broad,
coarsely reticulate, with an ill-defined carina from the base to the
apex, sharply but not quite vertically truncate posteriorly, the face
of the truncation transversely striated, with a median carina.
Abdomen shining, microscopically punctured, the three basal
segments with a band of white pubescence on the apical margin,
interrupted on the basal segment ; the pygidial area elongate tri-
angular, narrowly truncate at the apex. The sides of the median
segment are finely obliquely striated. The legs are slender, the
anterior tibize unarmed, and the tarsal ungues long. The two re-
current nervures are received very close together, before the middle
of the second cubital cell. The second cubital cell is at least half
as long again on the radial nervure as the third, the second trans-
verse cubital nervure is shghtly curved inwards, the third is
oblique and straight.
Black; the mandibles fuscous ; the pubescence on the pygidial
area very pale fulvous. Wings light fusco-hyaline, nervures
black, the tegulee fuscous.
Length 9 mm.
Hab. Perth, W. Australia.
Type in British Museum.
NoroGoNIA COMMIXTA, sp. N.
2. Clypeus opaque and very finely punctured at the base, with
a delicate carina not reaching the apex, the apex broadly smooth
and shining, the margin very broadly rounded, with a very narrow
and shallow emargination in the middle. Antenne rather stout,
the second joint of the flagellum twice as long as the first and
about one-quarter longer than the third. Head opaque, the
front and the base of the clypeus clothed with short fine silver
pubescence, which is only visible in certain lights; the eyes
separated on the vertex by a distance equal to one-third more
than the length of the first joint of the flagellum. Pronotum very
1908. } AUSTRALIAN FOSSORIAL WASPS. 48]
steeply sloped, not very strongly raised in the middle; the meso-
notum and scutellum subopaque, very closely and microscopically
punctured, a very short, impressed, longitudinal line on each side
of the mesonotum twice as far from the middle as from the
tegule. Median segment a little longer than broad, rather
indistinctly transversely striated, the strie more distinct at the
base than at the apex, with an obscure carina from the base
to the apex; the posterior truncation finely transversely striated,
with a deep median suleus. Abdomen subopaque, the three basal
segments with bands of dull silvery pubescence on the apical
margin; the pygidial area rather broad at the base, narrowly
rounded at the apex, clothed with pale fulvous pubescence; the
apex of the first ventral segment finely obliquely striated, the
second and third sparsely clothed with very short, fine, grey
pubescence. The third cubital cell is nearly twice as long as
the second on the radial nervure and a little longer than the
second on the cubital nervure; both recurrent nervures are
received before the middle of the second cubital cell, the first
more than twice as far from the base of the cell as from the
second.
Black; the tarsal ungues fusco-ferruginous. Wings fusco-
hyaline, faintly flushed with opalescent blue; nervures black.
Length 13 mm.
Hab. Cairns, Queensland (Zurner); February to May.
The mesonotum is only very slightly depressed in the middle on
the anterior margin.
NoToGONIA ABBREVIATA, Sp. Nn.
@. Mandibles very shallowly notched on the outer margin ;
the clypeus opaque, very finely and closely punctured, the apical
margin transverse and strongly depressed in the middle, the base
bare, the middle with rather long, sparse, silvery pubescence e
changing to fulvous at the apex. Scape with a little short white
pubescence beneath ; the first joint of the flagellum half as long
as the second, the fourth slightly longer than the second or third,
which are nearly equal in length to each other. Head opaque,
the front clothed with silvery ‘pubescence ; the eyes separated on
the vertex by a distance equal to three-quarters of the length of
the second joint of the flagellum. Pronotum almost vertical,
much higher in the middle than at the sides; mesonotum very
broadly and shallowly depressed in the middle anteriorly, opaque
and very minutely punctured; the pubescence short, sparse and
cinereous, that on the mesopleure also cinereous, with a patch of
silver-white pubescence below the base of the anterior wings.
Median segment more opaque than the mesonotum, very shallowly
and obscurely transversely striated, the striz only visible in a
strong light, with a rather shallow suleus from the base to the
apex, vertically truncate posteriorly, with a deep longitudinal
suleus on the face of the truncation, the segment is as broad at the
base as long, narrowed posteriorly. Abdomen opaque, with broad
31*
482 MR. R, E. TURNER ON [May 12,
bands of silvery pubescence on the apical margins of the three basal
segments; the pygidial area rather broadly rounded at the apex,
more than twice as broad at the base as at the apex, clothed with
sparse, stiff, black pubescence. The second ventral segment has a
a longitudinal carina from the base to the middle. The two
recurrent nervures are received very near together, just beyond
one-third from the base of the second cubital cell; the third
cubital cell is longer by one-third than the second on the radial
nervure.
Black. Wings light fusco-hyaline, nervures black.
Length 16 mm.
Hab. Cairns, Queenslaud (Zurner); February.
The median segment is shorter and broader than in most
species of Votogonia, but otherwise it shows all the characteristic
points of the genus.
TACHYTES RUBELLUS, sp. n.
3. Clypeus closely punctured, depressed and shining at the
apex, the margin slightly recurved. Head closely and very
minutely punctured; the front covered with shining white
pubescence, a deep longitudinal sulcus on the vertex broadened
just behind the posterior ocelli into a small triangular depression.
Eyes convergent above, separated on the vertex by a distance about
equal to the length of the first and second joints of the flagellum
combined ; the third joint of the flagellum of almost the same
length as the second. Thorax very minutely and closely punc-
tured, the sides thinly clothed with white pubescence ; the meso-
notum depressed in the middle anteriorly; the median segment
about one-fourth shorter than the mesonotum, narrowed and
truncate posteriorly, a deep longitudinal sulcus on the surface of
the truncation. Abdomen shining, minutely punctured; the apical
margin of the segments with interrupted bands of white
pubescence. The mesopleure, the sides of the median segment,
and the femora beneath with silvery pubescence.
Black; the mandibles, the anterior tibie beneath, the anterior
tarsi, the three orfour apical joints of the intermediateand posterior
tarsi, the spines of the tibie, and a spot at the apex of the inter-
mediate and posterior tibie ferruginous; the tegule testaceous ;
the abdomen entirely ferruginous red, paler on the apical margin
of the segments.
Length 7 mm.
Hab, Port Darwin (Turner); December.
TACHYTES FORMOSISSIMUS, Sp. n. (Plate XXVI. fig. 6.)
Q. Head, thorax, and median segment densely covered with
shining golden pubescence; the clypeus transverse and finely
punctured, the anterior margin shining and without pubescence.
A longitudinal sulcus on the vertex behind the ocelli; the eyes
convergent above, separated on the vertex by a distance slightly
1908. ] AUSTRALIAN FOSSORIAL WASPS. 483
exceeding the length of the two basal joints of the flagellum.
Pronotum depressed below the mesonotum, which is depressed in
the middle of the anterior margin; scutellum short, broadly
truncate at the apex. Median segment about one-third shorter
than the mesonotum, narrowed posteriorly and truncate, a deep
median sulcus starting from just above the base of the truncation
and continued along its surface to the apex. Abdomen extremely
minutely punctured, without pile.
Head and thorax black; the apex of the clypeus, the mandibles,
and the scape of the antenne ferruginous brown; the scutellum
and median segment fusco-ferruginous ; abdomen and legs light
testaceous red, the apical margin of the abdominal segments
paler; pubescence golden. Wings hyaline; nervures black,
testaceous red at the base and on the costa.
Length 10 mm.
Hab. Mackay, Queensland (Turner); February.
This beautiful species is allied to fervida Sm., but is much
smaller and less stoutly built.
TACHYTES APPROXIMATUS, Sp. n.
Q. Clypeus closely punctured, transversely depressed and
truncate on the apical margin ; the clypeus and front clothed with
pale shining pubescence, with a faint golden sheen. Antenne
inserted nearer to each other than to the eyes; the second joint of
the flagellum about three times as long as the first and equal in
length to the first and third combined, the fourth, fifth, and sixth
equal in length to each other, each slightly shorter than the third.
Hyes separated on the vertex by a distance about equal to the
length of the third joint of the flagellum; the vertex very
minutely punctured, with a fine longitudinal suleus from the
posterior ocelli to the posterior margin of the head. Pronotum
very much depressed and very small; the thorax and median
segment very minutely punctured; a broad line of short, pale
golden pubescence on the sides of the mesonotum, very narrowly
continued on the posterior margin, with a quadrate, narrowly
divided patch on each side of the middle of the anterior margin ;
the pubescence on the postscutellum and the sides of the median
segment pale and shining, the segment only a little more than
half as long as the mesonotum, steeply but not quite vertically
sloped posteriorly, with a median sulcus on the posterior face.
Abdomen shining, finely punctured and pubescent, the pubescence
short, thin, and dark, the apical margins of the four basal dorsal
segments with a broad band of pale and somewhat obscure goiden
pubescence ; the pygidial area long and not very narrow, narrowly
rounded at the apex and clothed with brilliant golden pubescence.
The basal joint of the anterior tarsus with five spines on the outer
margin. The third cubital cell is a little longer than the second
on the radial nervure, and about as long as the second on the
cubital nervure, narrow and much curved.
Black; the mandibles fusco-ferruginous; the tegule, tibie,
ABA MR. R, E. TURNER ON [May 12
tarsi, the apex of the femora, and the apical margin’ of the fifth
abdominal segment very narrowly ferruginous. Wings hyaline,
nervures ferrugimous.
3. As in the female, but the pubescence on the head and
thorax is longer and duller, the anterior margin of the clypeus is
more rounded, the apical margin of the abdominal segments 1s
broadly dull ferruginous with the band of pubescence as in the
female on the four basal segments, the pubescence on the apical
segment is whitish, and the posterior femora are black at the base
only. The abdomen is more closely punctured than in the female.
The eyes are yellowish green in both sexes in life.
Length, 9 14mm., g 13 mm.
Hab. Mackay, Queensland (Zurner); February and March.
Key to the Australian Species of Tachysphex.
A. Abdomen wholly black.
a. Tibie and tarsi light ferruginous ..... ji dieehinaed bonen tn LERUMOCLUESs
6. Tibie and tarsi almost entirely black
a, Abdomen opaque, covered with very short pubescence.
a, Mesonotum not visibly punctured; abdomen szb-
CO) UN [ON cT eames dreeaeiee aoe, ORME eS ecormerareterduacse ce mgt lio MO AEXOURe
63, Mesonotum very distinctly punctured ; abdomen
ODAGWO cooose sesee T. mackayensis.
62, Abdomen more or less shining, pubescent on the
apical margin of the segments only.
a, Dorsal surface of the median segment longitudinally
FSET (oT Uae Rtas eae emer ABR Gere bor Mas Aeeecilen Lo URGIINAZI
63, Dorsal surface of the median segment not striated.
a+. Thorax and median segment strongly pubescent . 7. pilosulus.
64, Thorax and median segment not pubescent.
a. Median segment longer than the mesonotum... 1. tenwis.
6°, Median segment not longer than the meso-
notum.
a®, Mesonotum subopaque ........................... 2. debilis.
6°, Mesonotum shining.
a’, Pronotum sharply sloped.
a8, Second joint of the flagellum much
longer than the third. Length 8mm.. 7. pacificus.
68. Second joint of flagellum shorter than
the third. Length 13mm................ LZ. nigerrimus Sm.
67, Pronotum gradually sloped.
a°. Head shining, almost smooth. Wines
and antennz short ............:............. 2. pugnator.
68. Head opaque, finely punctured. Wings
and antenne of moderate length ......... T. hypoleius Sm.
B. Two basal segments of abdomen ferruginous red ............_ T. truncatifrons.
TACHYSPHEX TRUNCATIFRONS, Sp. Nl.
2. Mandibles very strongly incised on the outer margin.
Clypeus broadly truncate on the apical margin, slightly convex in
the middle, obliquely and broadly triangularly truncate from the
middle to the apex, the base opaque and indistinctly punctured,
the surface of the truncation shining, with a few scattered
punctures. Antenne inserted as far from each other as from the
eyes; the second, third, and fourth joints of the flagellum about
equal in length, each at least three times as long as the first joint
1908. ] AUSTRALIAN FOSSORIAL WASPS. 485
Hyes separated on the vertex by a distance equal to the length of
the first two joints of the flagellum ; the posterior ocelli situated
on a convexity, divided by a fine longitudinal sulcus, with a rather
large, shallow, subtriangular depression on the vertex behind
them. Head opaque, the vertex very finely and closely punctured,
the depression behind the ocelli smooth and shining, the front
very delicately rugulose, slightly raised in the middle above the
base of the antenne, where it is divided by a longitudinal sulcus ;
the front almost flat. Thorax subopaque, very closely and deli-
eately punctured ; the pronotum not very much below the level
of the mesonotum, and very gradually sloped anteriorly. Median
segment opaque, a little shorter than the mesonotum, very finely
rugulose, truncate posteriorly, with a deep median sulcus on the
face of the truncation. Abdomen shining and almost smooth, the
three basal segments depressed on the apical margin, narrowly at
the sides, broadly in the middle; the pygidial area elongate and
pointed, shining, with scattered punctures. The tibie are very
feebly spined ; the comb of the anterior tarsi is long, but thin.
The third transverse cubital nervure is rather feebly curved.
Black ; the apical joint of the tarsi fusco-ferruginous; the two
basal abdominal segments above and beneath bright ferruginous
red. Wings hyaline stained with fuscous, nervures fuscous.
Length 9 mm.
Hab. Queensland.
Type in Oxford University Museum ex coll. Saunders.
Allied to 7’. rufo-niger Bingh. from India.
TACHYSPHEX IMBELLIS, Sp. Nn.
2. Mandibles incised on the outer margin; clypeus shining,
very sparsely punctured, the punctures large, feebly convex, the
apical margin subtruncate. Head, thorax, and abdomen shining
and almost smooth; a very delicate longitudinal sulcus reaching
from the anterior ocellus nearly to the base of the clypeus; the
posterior ocelli subovate, situated on a very slight elevation, with
a small and fairly deep depression on the vertex behind them ;
the eyes separated on the vertex by a distance about equal to the
length of the first two joints of the flagellum. Antenne inserted
nearer to each other than to the eyes; the second, third, and fifth
joints of the flagellum nearly equal in length, each less than twice
as long as the first, the fourth joint very slightly longer than the
others. Pronotum depressed below the mesonotum, a little more
than half as wideas the head. Median segment at the base a little
more than half as broad again as long, at the apex only as broad
as long, opaque and finely shagreened, almost vertically truncate
posteriorly, the face of the truncation finely and obscurely trans-
versely striated with a deep rounded depression at the base. The
apical segment of the abdomen shining, finely and very sparsely
punctured, pointed at the apex. The comb of the anterior tarsi
is very poorly developed, the sete being very few and far apart.
The third cubital cell is not much extended on the cubital nervure,
486 MR. R. E. TURNER ON [May 12,
being more than half as long on the radial as on the cubital
nervure; the radial cell is short, not more than equal to the length
of the second cubital cell on the cubital nervure. ts
Black; the mandibles except at the apex, the tegule, tibie,
tarsi, and the apex of the femora light ferruginous, the scape of
the antenne beneath fuscous. Wings hyaline, nervures fuscous.
A little grey pubescence on the sides of the abdominal segments,
otherwise bare.
Length 6 mm., exp. 10 mm.
Hab. Mackay, Queensland (Turner); November.
TACHYSPHEX FORTIOR, sp. nh.
@. Clypeus shining, strongly but sparsely punctured at the
apex, moderately convex at the base, then almost vertically
depressed, and subtruncate on the apical margin. Antenne
inserted as far from each other as from the eyes ; the second joint
of the flagellum twice as long as the first, a little shorter than the
third or fourth and about equal to the fifth. Eyes separated on
the vertex by a distance scarcely greater than the length of the
third joint of the flagellum. Head opaque, very minutely
punctured ; the front very feebly convex, divided by a rather
strong longitudinal sulcus which reaches the anterior ocellus and
is continued behind it to the posterior margin of the head.
Posterior ocelli subovate, situated on a convexity, with a small
deep depression behind them. Pronotum depressed below the
mesonotum, the slope almost vertical; mesonotum subopaque,
nearly twice as broad as long. Median segment opaque, shorter
than the mesonotum, finely shagreened, vertically truncate
posteriorly, the face of the truncation rather finely transversely
striated, with a median sulcus. Abdomen subopaque ; the apical
margin of the segments feebly depressed, very broadly in the
middle, and thinly clothed with grey pubescence; the apical
segment smooth, with a few scattered punctures near the hase,
subcarinate longitudinally in the middle and pointed at the apex.
Tarsi strongly spinose, the comb of the anterior tarsi well
developed; the legs stout. The first recurrent nervure is received
almost as far from the second as from the base of the second
cubital cell; the third cubital cell is strongly curved on the outer
“margin and reaches on the cubital nervure nearly two-thirds of
the way from the apex of the second cubital cell to the outer
margin of the wing.
Black; the pubescence greyish white; the tegule, the anterior
tarsi, the two apical joints of the posterior and intermediate
tarsi, and the extreme base of the tibie ferruginous.
Length 10 mm.
Hab. S.W. Australia (Du Boulay).
Type in British Museum.
_ Allied to 7. debilis described in this paper, but is a much more
stoutly built insect and the legs are stronger and much more
splnose.
1908. | AUSTRALIAN FOSSORIAL WASPS. 487
TACHYSPHEX MACKAYENSIS, Sp. nh.
@. Clypeus finely punctured at the base, very coarsely and
closely punctured, the apical margin very narrowly depressed and
shining, very narrowly emarginate in the middle. Antenne
inserted a little further from the eyes than from each other; the
second joint of the flagellum twice as long as the first, but a little
shorter than the third or fourth. Eyes separated on the vertex
by a distance slightly greater than the length of the third joint
of the flagellum ; the posterior ocelli situated on a convexity with
a small depression behind them. Head opaque, very finely and
shallowly punctured ; the clypeus at the base and the front below
the base of the antenne closely, the front above the base of
the antenne very sparsely, clothed with moderately long white
pubescence. Thorax very closely punctured; the pronotum
strongly depressed below the level of the mesonotum, almost
vertical, a little higher in the middle than at the sides.
Median segment almost as long as the mesonotum, narrowed
towards the apex and vertically truncate posteriorly, opaque and
finely punctured-rugose, with a little scattered pubescence; the
face of the truncation finely transversely striated, with a deep
median sulcus not reaching the apex. Abdomen opaque, covered
with extremely short grey pubescence, which is rather longer and
whiter on the depressed apical margin of the segments; the
pygidial area shining, with large and ccaiiaaad punctures, almost
pointed and rather narrow; the ventral segments smooth and
shining. The posterior tibie are rather stro ongly spined, the comb
of the anterior tarsi is long but rather thin. The third cubital
cell is very long on the cubital nervure, reaching nearly two-thirds
of the way from the apex of the second cubital cell to the margin
of the wing; the third transverse cubital nervure is much curved ;
the second cubital cell is longer than the third on the radial
nervure, but scarcely as long on the cubital.
Black ; the tegulz and the comb of the anterior tarsi testaceous.
Length 12 mm., exp. 19 mm.
Hab. Mackay, Queensland (7wrner).
Allied to 7. fortior described above, but the eyes are further
apart on the vertex, and the sulcus on the vertex is absent, the
mesonotum is punctured, the median segment longer and more
strongly narrowed posteriorly, the pygidium narrower, and the
posterior tibize more strongly armed.
TACHYSPHEX WALKERI, Sp. 0.
@. Clypeus shining, sparsely punctured, very finely at the base,
more coarsely at the apex, obliquely depressed from the middle to
the apex, where it is broadly truncate. Antenne long and slender,
nearly as long as the head, thorax, and median segment combined,
inserted further from the eyes than from each other; the second
joint of the flagellum more than twice as long as the first and
about one-quarter shorter than the third. Eyes separated on the
vertex by a distance not quite equal to the length of the second
488 MR. R. E. TURNER ON [May 12,
joint of the flagellum ; the posterior ocelli subovate, situated on a
convexity, with a small triangular depression on the vertex
behind them. Head opaque, the vertex smooth, the front slightly
convex, finely shagreened, with a delicate, longitudinal, median
sulcus; the space round the base of the antenne clothed with short
silver pubescence. Thorax subopaque, the scutellum shining; the
pronotum much depressed below the mesonotum, gradually sloped
anteriorly. Median segment as long as the mesonotum, narrowed
posteriorly, opaque and rather irregularly longitudinally striated,
the sides finely obliquely striated, vertically truncate posteriorly,
the face of the truncation transversely striated, with a median
suleus. Abdomen microscopically punctured, the apical margin of
the three basal segments slightly depressed, with a band of short,
rather thin, silver pubescence ; the pygidial area polished, with a
few fine and scattered punctures, elongate triangular. The
posterior tibize very feebly spined, the comb of the anterior tarsi
only feebly developed. Third cubital cell narrow, the third trans-
verse cubital nervure rather strongly curved inward ; the second
cubital cell much shorter than the third on the radial nervure,
the radial nervure strongly continued, the appendiculate cel! being
clearly defined and reaching nearer to the outer margin than the
cubital nervure.
Black; the scape beneath fusco-ferruginous ; the apex of the
tegule and the comb of the anterior tarsi testaceous.
Length 7 mm.
Hab. Sand Islet, Long Reef, N.W. Australia (Walker); June.
Type in British Museum.
TACHYSPHEX PILOSULUS, Sp. 0.
2. Clypeus very broadly rounded anteriorly, clothed with dark
grey pubescence, changing to silver in some lights, the apical
margin bare and recurved, with a shining transverse depression
before it. Antennz inserted about half as far again from the
eyes as from each other; the second joint of the flagellum twice as
long as the first and about equal in length to the third and the
fourth singly. Eyes separated on the vertex by a distance
scarcely exceeding the length of the second joint of the flagellum ;
the posterior ocelli oblong and oblique, situated on a convexity,
with no depression or sulcus on the vertex behind them. Front
rather strongly convex, clothed with long and rather close silver-
grey pubescence, a very fine longitudinal sulcus below the anterior
ocellus. The whole head opaque, very minutely punctured. Pro-
notum very much depressed below the level of the mesonotum,
very steeply stoped anteriorly. Thorax and median segment
finely shagreened, opaque, with sparse and rather long cinereous
pubescence ; the mesonotum large, about half as lon g again as the
median segment, which is scarcely narrowed posteriorly, and ver-
tically truncate, the surface of the truncation indistinctly trans-
versely striate-rugulose, with an almost obsolete median sulcus.
Abdomen subopaque, the two apical segments and the whole
1908. ] AUSTRALIAN FOSSORIAL WASPS. 489
ventral surface shining, the basal segment with sparse and rather
long grey pubescence at the base, the three basal segments with a
broad band of shining silvery pubescence faintly flushed with
golden on the apical margin, the fourth segment with a similar
band on the sides only. Pygidial area elongate-triangular, very
narrowly truncate at the apex, microscopically punctured and less
polished than the fifth segment. The comb on the anterior tarsi
is very long. ‘The first recurrent nervure is received at the same
distance from the base of the second cubital cell as from the
second recurrent nervure, which is received just beyond the middle
of the cell, The second cubital cell is about half as long again as
the third on the radial nervure; the third is much produced on
the cubital nervure, reaching two-thirds of the way from the apex
of the second to the margin of the wing; the second and third
transverse cubital nervures are both strongly curved. The cubital
nervure of the posterior wings branches off immediately beyond
the apex of the anal cell.
Black; the spines of the tibiz and tarsi whitish; the tegule
and the comb of the anterior tarsi testaceous. Wings hyaline ;
nervures ferruginous at the base, fuscous at the apex. Anterior
tarsi fusco-ferruginous.
g. Similar, but slenderer, the head and thorax less strongly
pubescent, the eyes a little nearer together on the vertex, and the
apical abdominal segments more opaque. The eyes in life are
bright green.
Length, 2 11 mm., ¢ 10 mm.
Hab. 9, Cape York, Queensland; ¢, Cairns and Mackay,
Queensland (Zurner); November to April.
TACHYSPHEX TENUIS, Sp. n.
2. Clypeus shining, almost smooth at the base, punctured near
the apex, moderately convex, the apical margin transversely
depressed and shallowly emarginate in the middle. Antenne
inserted nearer to each other than to the eyes; the second joint of
the flagellum less than twice the length of the first and distinctly
shorter than the third or fourth. Eyes separated on the vertex
by a distance equal to the length of the first and third joints of
the flagellum combined; the posterior ocelli ovate and oblique,
placed on a convexity, with a small, deep, triangular depression on
the vertex behind them. Head opaque, very minutely punctured,
the front slightly convex, with an obscure longitudinal sulcus from
the anterior ocellus; the space round the base of the antenne
clothed with short white pubescence. Pronotum moderately de-
pressed below the mesonotum, not very steeply sloped. Thorax
slightly shining and almost smooth ; the median segment opaque,
a little longer than the mesonotum and longer than broad, ver-
tically truncate posteriorly, the face of the truncation finely
transversely striated, with a deep median sulcus. Abdomen
slender, slightly shining, and very minutely punctured ; the apical
margin of the three basal segments depressed, most broadly in the
490 MR. R. E. TURNER ON [May 12,
middle, with a little short grey pubescence on the sides. Pygidial
area shining and sparsely punctured, narrow and almost pointed
at the apex. The comb on the anterior tarsi is not very strongly
developed. The second cubital cell is very short on the radial
nervure, only half as long as the third; the first recurrent ner-
vure isreceived at the same distance from the base of the cell as from
the second recurrent nervure, which is received close to the middle
of the cell. The third cubital cell is not much prolonged on the
cubital nervure, reaching only about two-fifths of the way from
the apex of the second cell to the margin of the wing. The
second and third transverse cubital nervures are scarcely curved.
Black ; the apical half of the flagellum and the apical joint of
the tarsi fuscous; the tegule testaceous. Wings hyaline,
nervures ferruginous.
Length 6 mm.
Hab. Port Darwin (Turner) ; December.
TACHYSPHEX DEBILIS, Sp. N.
@. Mandibles strongly incised on the outer margin; clypeus
short and broad, strongly transversely depressed on the apical
margin and feebly emarginate in the middle, opaque at the base,
shining at the apex, and sparsely punctured. Front round the
base of the antenne clothed with very short silvery pubescence,
with a shallow longitudinal sulcus reaching very obscurely to the
anterior ocellus. Head opaque, the front slightly convex; the
posterior ocelli subovate, situated on an elevation, with a small
and rather deep rounded depression behind them. Antenne
inserted much nearer to each other than to the eyes; the second
joint of the flagellum a little shorter than the third, the third
and fourth equal in length, the fifth equal to the second. EHyes
strongly convergent, separated on the vertex by a distance equal
to the length of the first two joints of the flagellum combined.
Pronotum scarcely more than half as broad as the head, depressed
below the level of the mesonotum, which is opaque and micro-
scopically punctured; the scutellum shining, very minutely
punctured. Median segment very opaque, finely shagreened,
nearly twice as broad at the base as long, almost vertically
truncate posteriorly; the face of the truncation indistinctly
transversely striated, with a median sulcus. Abdomen shining,
minutely punctured, the apical margin of the segments with
broadly interrupted bands of obscure silver pubescence; the
apical segment pointed, shining, and very sparsely punctured.
The tibize are very feebly spined, the anterior tibie smooth; the
tarsi more feebly spined than is usual in the genus, the comb of
the anterior tarsi only moderately developed.
Black, the tegule fuscous. Wings hyaline, nervures fuscous.
The third cubital cell is narrow and strongly extended along
the cubital nervure towards the margin of the wing.
Length 7 mm.
Vs Sarees ey VRS 4
Hab. Cairns, Queensland (Turner); February.
1908. | AUSTRALIAN FOSSORIAL WASPS. 49]
TACHYSPHEX PACIFICUS, sp. n.
2. Clypeus very broadly truncate at the apex, strongly and
closely punctured at the apex, sparsely and more finely at the
base. Antenne inserted as far from each other as from the eyes;
the second joint of the flagellum three times as long as the first
and about one-third longer than the third, the fourth and fifth
almost equal in length to the third; the distance between the
eyes on the vertex equal to the length of the second joint of the
flagellum ; the posterior ocelli elongate-ovate, situated on a very
feeble elevation, a small depression on the vertex behind them.
Head subopaque, closely and finely punctured, with a delicate
longitudinal sulcus on the front. Thorax shining, very closely
and minutely punctured ; scutellum smooth and shining. Median
segment opaque, very finely shagreened, the apex shining, almost
vertically truncate posteriorly ; the face of the truncation finely
transversely striated, with a delicate median sulcus. Abdomen
smooth and shining, the apical margin of the segments broadly
and feebly depressed, the apical segment triangular, the sides
longer than the base, sparsely punctured. The comb on the
anterior tarsi 1s composed of very few sete. The third cubital
cell is strongly produced towards the margin of the wing, being
nearly four times as long on the cubital as on the radial nervure.
The sides of the median segment are obliquely striated.
Black; the mandibles and tegule fuscous. Wings hyaline,
faintly tinted with fuscous ; nervures black.
Length 8 mm.
Hab. Melbourne.
Type in British Museum ex coll. Smith.
Near 7’. debilis described above, but the proportion of the joints
of the antenne is very different, and the thorax and abdomen are
more polished, and the sculpture, especially on the sides of the
median segment, is different.
TACHYSPHEX NIGERRIMUS Sim.
Tachytes mgerrimus Sm. Cat. Hym. B. M. iv. p. 302. n. 26,
1856.
Astata nigerrima White; Butler, Zool. Voy. Ereb. & Terror, ii.
pl. 7. fig. 14, 1875.
Hab. New Zealand.
I have received a specimen said to be from Victoria, but the
locality may be a mistake.
Smith took the name from White’s MS. referring to the above
work, of which the portion containing the Hymenoptera was not
published till many years later.
TACHYSPHEX PUGNATOR, Sp. n.
2. Clypeus shining and sparsely punctured, convex, strongly
depressed, and bent inwards on the apical margin, giving the
appearance of a wide and shallow emargination. Antenne
492 MR. R. E. TURNER ON [May 12,
inserted almost as far from each other as from the eyes, short and
rather stout; the second joint of the flagellum not much more
than twice as long as the first, and about equal to the third in
length but not quite as thick, the fourth and fifth joints as long
as the third. Eyes separated on the vertex by a distance about
equal to the length of the third and fourth joints of the flagellum
combined. Head shining and almost smooth, a very delicate sulcus
reaching from the anterior ocellus almost to the base of the clypeus.
Posterior ocelli ovate, with a small deep depression behind them,
from which a rather deep sulcus runs to the posterior margin of
the head. Thorax narrower than the head, smooth and shining ;
the pronotum depressed below the mesonotum, gradually but not
very steeply sloped ; the mesonotum with a longitudinal suleus on
each side, and a very short longitudinal sulcus from the middle of
the anterior margin. Median segment opaque, shorter than the
mesonotum, abruptly but not quite vertically truncate posteriorly,
the face of the truncation finely transversely striated, with a deep
median sulcus not reaching the apex. Abdomen almost smooth,
the ventral surface highly polished; the segments depressed on
the apical margin, the three basal segments very strongly so and
much more broadly on the middle than on the sides, the second
segment obscurely transversely furrowed before the depression ;
the margins of the three basal segments with scant white
pubescence (probably thicker in fresh specimens); the pygidial
area shining and sparsely punctured, very long and narrow,
almost pointed at the apex. The comb on the anterior tarsi well
developed. The radial nervure reaches nearer to the outer margin
of the wing than the cubital; the first recurrent nervure is
received nearer to the second than to the base of the second
cubital cell.
Black; the flagellum fuscous, the apex of the scape beneath
fusco-ferruginous; tarsi fuscous at the base, the apical joints
fusco-ferruginous; the comb of the anterior tarsi ferruginous.
Wings hyaline, tinged with fuscous ; nervures black.
Length 11 mm.
Hab. Adelaide, 8. Australia.
Type in British Museum.
Somewhat allied to 7. nigerrimus from New Zealand, but the
thorax is narrower in proportion to the head, and the antenne
are very different.
TACHYSPHEX HYPOLEIUS Sm.
2. Clypeus slightly convex, depressed towards the anterior
margin, which is broadly subtruncate, very minutely punctured.
Antenne inserted further from each other than from the eyes,
the space between them shining and almost smooth, above the
shining space is a very short longitudinal carina with a very small
tubercular prominence on each side of it. The first joint of the
flagellum very short, not more than one-quarter of the length of
the second, the second and third about equal in length, the fourth
1908. | AUSTRALIAN FOSSORIAL WASPS. 493
a little shorter. Mandibles notched on the outer margin. Eyes
separated on the vertex by a distance equal to the length of the
second joint of the flagellum. Front slightly convex, subopaque,
finely and shallowly punctured ; the front below the base of the
antennee, the inner margin of the eyes as high as the ocelli and
the outer margin clothed with short silvery pubescence. Posterior
ocelli elongate- -ovate, situated on a slight convexity, with a shallow
depression on the vertex behind them. Thorax smooth and
shining, mesopleuree subopaque ; mesosternum shining, with a few
scattered punctures; the pronotum depressed below the meso-
notum, very gradually sloped. Median segment distinctly longer
than the mesonotum, opaque and shagreened, with a very shallow
longitudinal depression near the apex, vertically truncate pos-
teriorly ; the face of the truncation transversely striated, with a
deep median sulcus. Abdomen shining and almost smooth, the
apical margin of the segments depressed, with a band of short
silver pubescence broadly interrupted in the middle; the first
segment rounded towards the base; the pygidial area very
sparsely punctured, very slightly convex in the middle, elongate,
blunt at the apex. Intermediate coxe very widely separated ;
the comb of the anterior tarsi very strongly developed. The first
recurrent nervure is received rather nearer to the base of the
second cubital cell than to the second recurrent nervure, which is
received close to the middle of the cell; the third cubital cell
is produced, reaching on the cubital nervure much more than
halfway from the apex of the second cell to the margin of the
wing.
Entirely black, Wings hyaline, tinted with fuscous, nervures
black.
Length 14 mm., exp. 22 mm.
Hab. 8. Australia; W. Australia.
Type in British Museum.
The median segment, although broader than long, is longer
than is usual in the genus.
ZOYPHIUM ERYTHROSOMA, sp. li.
2. Mandibles very deeply notched on the outer margin.
Clypeus broad, slightly convex in the middle, the anterior margin
depressed transversely, with two minute teeth on each side,
almost smooth, with thin and very fine silvery pubescence on the
sides. Antenne inserted about half as far again from each other
as from the eyes, scarcely longer than the head, gradually
thickened to the apex, the length of the scape no greater than
the distance between the antenne; the first joint of the flagellum
globose, a little shorter than the second, the third half as long
again as the second, the fourth a little longer than the third, the
apical joint very large and massive, as long as the first and
second combined. stihvesk inner margin of the eyes almost straight,
the distance between the eyes on the vertex slightly exceeding
two-thirds of the length of the flagellum. The posterior ocelli
rather more than foie as far again ‘from each other as from the
494 MR. R. BE. TURNER ON [May 12, ©
eyes. Front depressed round the base of the antenne; head
a little broader than the mesonotum. Head and thorax opaque,
very closely and microscopically punctured. Pronotum depressed,
seen from above linear and transverse, sharply obliquely de-
pressed anteriorly. Mesonotum much longer than the median
segment, the punctures on the mesopleure a little more distinct
than on the mesonotum. Median segment rounded, with a
median longitudinal sulcus from the base to the apex, a short
earina lying in the sulcus at the base; along the base of the
segment are a number of short oblique strie. Abdomen a little
longer than the thorax and median segment combined, closely
and minutely punctured, the apical margin of the segments
broadly depressed and smooth. The pygidial area is triangular,
much more strongly punctured, with short, stiff hairs springing
from the punctures. The first recurrent nervure is received at
one-quarter before the apex of the first cubital cell, otherwise the
neuration is as in Kohl’s figure.
Obscure blackish blue; the clypeus, the base of the mandibles,
the scape of the antenne, the apex of the femora, the tibiz, and
the tarsi yellow ; the flagellum, the abdomen, and stains on the
tibie and tarsi light ferruginous; the tegule fuscous. Wings
hyaline, faintly iridescent, nervures ferruginous.
Length 8-9 mm.
¢. Clypeus blue-black ; hypopygium pointed, with a spine on
each side close to the apex. Otherwise as in the female.
Length 7 mm.
Hab. Townsville, Queensland (Dodd); Mackay, Queensland
(Lurner); February and March.
Described from two males and two females.
Near Sericophorus bicolor Sm., from W. Australia, but the
radial cell in that species is appendiculate and the head much
broader.
ZoYPHIUM RUFONIGRUM, sp.n. (Plate XXVI. fig. 8.)
3. Mandibles deeply notched on the outer margin, acute at
the apex, not bidentate. Clypeus very broadly rounded at the
apex, the apical margin depressed, without minute lateral teeth.
Antenne about one-third as far again from each other at the
base as from the eyes, the first joint of the flagellum shorter than
the second, globular, the second shorter than the third, the apical
joint massive and longer than the others, the antenne are a little
longer than head and are gradually thickened to the apex. The
inner orbits of the eyes are nearly parallel, very slightly con-
vergent towards the vertex. The eyes on the vertex separated
by a distance scarcely equal to two-thirds of the length of the
flagellum ; the posterior ocelli twice as far from each other as
from the eyes. Head and thorax minutely and closely punctured
the front clothed with short, silvery pubescence. Head broader
than the thorax; the pronotum short but not linear, strongly
depressed anteriorly. Mesopleure rather more strongly punctured
1908. ] AUSTRALIAN FOSSORIAL WASPS. 495,
than the mesonotum. The median segment rounded at the apex,
with a deep median sulcus ; the sides of the segment and the meso-
pleure clothed with silvery pubescence. Abdomen minutely
punctured, the apical margins of the segments rather broadly
depressed and smooth; the hypopygium subtruncate, with a
minute apical spine.
Text-fig. 108.
Zoyphium rufonigrum, 3.
1. Antenna. 2. Maxilla and palpus. 3. Labial palpus. 4. Mandible.
Black; the mandibles, clypeus, pro- and mesothorax, and
median segment ferruginous red; the tibiz, tarsi, scape of the
antenne, the first and part of the apical joint of the flagellum
testaceous yellow. Wings hyaline, iridescent, nervures fuscous.
Length 4 mm.
Hab. Port Darwin (Zurner); December.
Described from two specimens.
ZOYPHIUM KOHLII, sp. n.
©. Mandibles acute at the apex, deeply notched on the outer
margin at one-third from the base, with two small teeth narrowly
divided from each other on the inner margin also at one-third
from the base. Clypeus short and broad, opaque and smooth, the
anterior margin transversely depressed, with a tooth on each side
opposite the two small teeth on the mandibles. Antenne nearly
half as far again from each other at the base as from the eyes,
gradually thickened to the apex; the first jot of the flagellum
globular, shorter and stouter than the second, the second a little
longer than the third, the apical joint large and longer than any
two of the others combined. Eyes moderately convergent towards
the vertex, where they are separated by a distance equal to about
half the length of the flagellum ; the posterior ocelli nearly twice
as far from each other as from the eyes. Head and thorax
opaque, almost smooth, the front covered with very short golden
pubescence. Pronotum much narrower than the head, depressed
slightly below the level of the mesonotum and strongly depressed
anteriorly. Median segment shorter than the mesonotum,
Proc. Zoou. Soc.—1908, No. XXXII. ay
496 MR. R. E. TURNER ON [May 12,
rounded posteriorly, with a median carina lying in a shallow
depression reaching from the base almost to the apex, with very
short oblique strie at the base, shining at the apex, the inter-
mediate space opaque. Abdomen elongate-ovate, minutely and
rather sparsely punctured, the apical margin of the segments
narrowly depressed, the pygidial area triangular with sparse
pubescence.
Black; the mandibles, antenne, tegule, and legs pale ferru-
ginous brown. Wings hyaline, iridescent ; nervures fuscous.
Length 6 mm.
Hab. Mackay, Queensland (Turner); January.
ZOYPHIUM FRONTALE, sp. 1.
@. Clypeus very broad, truncate at the apex; the antenne
inserted at the base of the clypeus, twice as far from each other
as from the eyes, thickened to the apex and no longer than the
head. Eyes very slightly convergent towards the vertex; the
posterior ocelli more than twice as far from each other as from
the eyes. Front very broad, slightly concave at the sides, a little
elevated inthe middle. Head, thorax, and abdomen very minutely
punctured, the abdomen shining. Pronotum short, narrower
than the head, and depressed below the mesonotum. Mesonotum
large, as broad as long and as broad as the head. Median segment
rounded at the sides, abruptly truncate posteriorly, much shorter
than the mesonotum, delicately obliquely striated, with a median
carina from the base situated in a depressed suleus. Abdomen
subconical, the second segment a little broader than the first.
Black ; the clypeus, the front and the scape of the antennz
yellow ; the mandibles, flagellum, tegule, and legs ferruginous ;
the apex of the scutellum and the postscutellum fusco-ferruginous.
Wings hyaline, iridescent ; nervures fusco-ferruginous.
There are only two cubital cells, the second transverse cubital
nervure being obsolete; the second cubital cell nearly twice as
long on the cubital as on the radial nervure, the length on the
radial about equal to the length of the first transverse cubital
nervure.
Length 5 mm.
Hab. Mackay, Queensland (Turner); March.
GORYTES DUBOULAYT, Sp. n.
2. Clypeus short and broad, closely and minutely punctured.
Head very minutely punctured, the front almost smooth, concave,
with a longitudinal suleus from the anterior ocellus to the base of
the clypeus. Hyes very large, diverging towards the clypeus and
the vertex; the posterior ocelli twice as far from each other as
from the eyes. Antenne inserted further from each other than
from the eyes, thickened to the apex; the second joint of the
flagellum half as long again as the third. Pronotum linear,
transverse, nearly as broad as the head. Thorax very minutely
and shallowly punctured ; the mesonotum half as broad again as
1908. ] AUSTRALIAN FOSSORIAL WASPS. 497
long. Median segment with a smooth, shining triangular space
at the base ; from the apex of the triangle a deep sulcus ruus to the
apex of the segment, the remainder of the segment shining, with
microscopic punctures. Abdomen oval, as long as the thorax and
median segment combined; the first segment narrow at the base,
no longer than the second segment which is very broad ; all the
segments finely and very closely punctured ; the pygidium elon-
gate-triangular, with sparse, short pubescence.
Black; the mandibles at the base, clypeus, scape of the an-
tenn, pronotum, tegule, a spot on the mesopleure beneath the
anterior wings, a short transverse line at the apex of the scutellum,
the postscutellum, a large spot on each side of the second abdo-
minal segment, a transverse band very narrowly interrupted in
the middle at the apex of the fourth segment, the apex of the
anterior femora, and a line on all the tibiz above, pale yellow ;
the flagellum, the apex of the mandibles, the legs, the first
abdominal segment (except at the base), and the two apical
segments ferruginous.
The first recurrent nervure is received near the apex of the
first cubital cell, the second close to the apex of the second. The
first cubital cell is longer on the cubital nervure than the second
and third combined ; the second as long as the third, but much
shorter on the radial nervure.
Length 7 mm. .
Hab. Australia, N.W. Coast (Du Boulay).
Type in British Museum.
This and the two following species form a group of allied forms
approaching the subgenus Miscothyris Sm., from which they may
be distinguished by the shorter and more oval abdomen and the
very large and broad second abdominal segment.
GORYTES SANGUINOLENTUS, Sp. n. (Plate XX VI. fig. 10.)
2. Clypeus broad, slightly convex, about two and a half times
as broad on the apical margin as long, truncate at the apex, the
labrum slightly prominent; minutely punctured. Eyes large,
strongly divergent towards the clypeus and towards the vertex,
separated on the front by a distance rather less than one and
a half times the length of the scape. The posterior ocelli twice
as far from each other as from the eyes, separated from the eyes
by a distance about equal to the length of the third joint of the
flagellum. Antenne inserted on the front, above the base of
the clypeus, about equal in length to the head, nearer to the eyes
than to each other; the scape as long as the first three joints of
the flagellum combined ; the second joint of the flagellum longer
than the third, the apical joints thickened. Head opaque, very
minutely punctured, vertex flattened. Pronotuin linear and
transverse, nearly two-thirds of the breadth of the head; meso-
notum more than half as broad again as long, finely punctured ;
scutellum large and broad, truncate at the apex. Median
segment of about the same length as the scutellum, rounded
32*
498 MR. R. E. TURNER ON [May 12,
and steeply sloped posteriorly, the sides finely and not very
closely punctured ; a smooth, shining, triangular area at the base,
produced at the apex of the triangle into a narrow shining median
line on the posterior slope. Abdomen ovate, closely and finely
punctured, the second segment the broadest and half as long
again as the third; the first segment steeply sloped anteriorly,
not truncate, about half as long as the second; the apical
segment small and pointed, with a fine longitudinal carina.
Black ; the base of the mandibles, the clypeus, the scape of the
antenne, pronotum, a spot on the mesopleuree near the base of
the anterior wings, the base of the tegule, the tibie above, and an
obscure spot on each side near the apical angles of the second
abdominal segment, pale yellow ; the flagellum, the tibize beneath,
the tarsi, and the abdomen ferruginous. Wings hyaline, iridescent ;
nervures ferruginous at the base and on the costa, black at the apex.
The first recurrent nervure is received near the apex of the
first cubital cell, the second at the apex of the second cubital cell,
almost interstitial with the second transverse cubital nervure.
The second cubital cell is of about the same length as the third
on the cubital nervure, less than half as long on the radial as on
the cubital nervure. The cubital nervure on the hind wing
originates far beyond the apex of the submedian cell.
Length 7 mm., exp. 12 mm.
Hab. Mackay, Queensland (Zurner); March.
This is nearer to Miscothyris than to any other section of
Gorytes, but differs much from thoracicus Sm., the antenne being
shorter, the second joint of the flagellum not unusually elongate ;
the eyes much more strongly divergent; the first abdominal
segment narrower and not truncate, the second segment much
larger and the apical segment pointed, not broadly subtruncate as
in thoracicus. Smith describes thoracicus as a male, but the type,
the only specimen I have seen, is a female and has the antenne
twelve-jointed, not thirteen-jointed as in Smith’s figure.
GoRYTES LUCIDULUS, sp.n. (Plate XXVI. fig. 11.)
2. Clypeus very broad, truncate anteriorly, finely and closely
punctured. Antenne inserted above the base of the clypeus,
nearer to the eyes than to each other, about as long as the thorax ;
the scape as long as the first two joints of the flagellum combined,
the second joint of the flagellum much longer than the third.
HKyes diverging towards the clypeus and towards the vertex; the
posterior ocelli further from each other than from the anterior
ocellus and twice as far from each other as from the eyes. Head
shining, almost smooth, with a delicate sulcus from between the
antenne to the anterior ocellus. Pronotum almost vertically
depressed, the posterior margin nearly on a level with the meso-
notum. Mesonotum half as broad again as long, finely and
closely punctured; scutellum short and transverse. Median
segment very short, steeply sloped, with a deep median sulcus
posteriorly, shining with a smooth triangular space at the base,
1908. | AUSTRALIAN FOSSORIAL WASPS. 499
the sides and apex very finely punctured. Abdomen subovate,
searcely longer than the thorax and median segment combined,
the second segment the broadest, the apical segment lanceolate.
The first recurrent nervure is received by the first cubital cell a
little before the apex, the second is interstitial with the second
transverse cubital nervure. The second cubital cell is scarcely
more than half the length of the third, the first is longer than the
second and third combined. The second on the radial nervure is
a littie more than half as long as on the cubital.
Black; the clypeus, the scape of the antenne, a large spot on
the mesopleurz beneath the anterior wings, the scutellum, post-
scutellum, and a spot on each side near the apical angles of the
second abdominal segment yellow. The tarsi and anterior tibiz
fuscous. Wings hyaline, nervures fusco-ferruginous.
Length 8 mm., exp. 15 mm.
Hab. Mackay, Queensland (Turner); October.
The intermediate tibize have two spines at the apex ; the tarsal
ungues are long, curved, and simple. The form is very similar to
sanguinolentus, but the scutellum is shorter and the whole insect
less robust.
A colour variety from Cairns is without the yellow spot on the
second abdominal segment and has the apical half of the clypeus
dark brown.
GORYTES ICARIOIDES, sp. 1.
2. Clypeus broadly truncate at the apex, subtriangular, but
truncate at the base, almost smooth. Eyes very large, diverging
towards the vertex, approaching each other most closely at the
base of the clypeus, where they are separated by a distance about
equal to the length of the scape of the antenna. Antenne
inserted rather high up on the front, as near to the eyes as to
each other ; the second joint of the flagellum nearly half as long
again as the third and quite as long as the scape, the four or five
apical joints forming a strong club, the jomts broader than long.
The posterior oceili far apart, two and a half times as far from
each other as from the eyes, from which they are separated by a
distance equal to the length of the first joint of the flagellum.
Head opaque, the space round the ocelli rather coarsely punctured.
Thorax and median segment closely punctured; the pronotum
depressed below the level of the mesonotum, the posterior angles
widely separated from the tegule. Median segment as long as
the mesonotum, truncate at the apex, with a longitudinal carina
on the surface of the truncation, the triangular area at the base
of the segment rather obscurely defined and very finely rugose.
Abdomen petiolate, the first segment half as broad again at the
apex as at the base, and nearly twice as long as the breadth at
the apex; second and third segments large and broad; the whole
abdomen rather closely punctured and opaque ; the apical margin
of the first segment smooth and shining, with sparse punctures.
Pygidial area triangular, with the margins slightly raised, covered
500 MR. R. E. TURNER ON [May 12,
with short, stiff pubescence. Both recurrent nervures are
received by the second cubital cell beyond the middle, the second
near the apex. The radial cell is narrow and pointed ; the first
cubital cell incompletely divided by a branch from the first trans-
verse cubital nervure, the second three times as long on the
cubital as on the radial nervure, the third nearly as long as the
first on the cubital and more than half as long again on the cubital
as on the radial nervure. The cubital nervure of the posterior
wing originates before the apex of the submedian cell. The
posterior tarsi are nearly twice as long as the tibie.
Black ; the clypeus, the front below the base of the antenne,
the scape of the antennz, and the posterior angles of the pronotum
yellow ; the mandibles, the four basal joints of the flagellum, the
tegule and a curved line above them, a transverse spot on the
postscutellum, the apex of the first and third abdominal segments
rather broadly and of the fourth and fifth more narrowly, and
the legs (except the cox and the posterior femora) ferruginous.
Wings hyaline; nervures black, ferruginous at the base.
Length 10 mm., exp. 19 mm.
Hab. Mackay, Queensland (7urner).
Nearest to G. decoratus Handl. (ornatus Sm.), from West
Australia.
GoRYTES CYGNORUM, sp. 0.
3. Clypeus broadly subtruncate at the apex, clothed with
silvery pubescence. Head finely and rather sparsely punctured,
with a delicate sulcus from the anterior ocellus nearly reaching the
base of the clypeus. Eyes strongly divergent towards the vertex,
less strongly towards the clypeus, separated from each other on
the front at the nearest pomt by a distance about equal to the
length of the scape and the first joint of the flagellum combined ;
the posterior ocelli about half as far again from each other as
from the eyes. Antenne shorter than the thorax and median
segment combined, moderately and very gradually thickened to
the apex, much nearer to the eyes than to each other. Pronotum
transverse and linear, slightly raised, and as broad as the head.
Thorax rather sparsely punctured; the postscutellum coarsely
longitudinally striated. Median segment much shorter than the
mesonotum, almost vertically truncate posteriorly, rugose, with a
median sulcus on the truncation, the area at the base coarsely
longitudinally striated. Abdomen elongate-ovate, very finely
and closely punctured; the first segment of about the same
length as the second, broadened to the apex, where it is more than
half as broad as the second segment. Both recurrent nervures
received by the second cubital cell, the distance between them
nearly two-thirds of the length of the cell. The second cubital
cell is as long on the cubital nervure as the third, but is much
narrowed on the radial nervure; the cubital nervure is sharply
bent before the reception of the first recurrent nervure and has
from that point the appearance of being continuous with the
1908. | AUSTRALIAN FOSSORIAL WASPS. 501
recurrent rather than with the cubital nervure. The cubital
nervure of the posterior wing originates far beyond the apex
of the submedian cell.
Black; the clypeus pale yellow; the mandibles, the scape of the
antenne, the two basal joints of the flagellum, the pronotum,
tegule, scutellum, legs, the apical half of the first and second
abdominal segments, the apical margins of the remaining seg-
ments (most broadly on the fifth), the second ventral segment
(except the extreme base), and the apical margin of the remaining
segments (except the first) orange. Wings hyaline, iridescent,
slightly tinged with fuscous ; nervures fusco-ferruginous.
Length 7 mm.
Hab. Swan River (Du Boulay).
Type in British Museum.
GORYTES FRENCHII, sp. n.
3. Clypeus slightly emarginate in the middle of the apical
margin, about twice as broad at the apex as long, minutely
punctured and thinly clothed with silver-grey pubescence. Head
delicately and closely punctured, with a fine suleus from the
anterior ocellus nearly reaching the base of the clypeus. Eyes
strongly divergent towards the vertex and less strongly towards
the clypeus, separated at the base of the clypeus by a distance not
exceeding the length of the scape; the posterior ocelli rather
more than half as far again from each other as from the eyes.
Antenne rather shorter than the thorax and median segment
combined, slightly thickened towards the apex ; the second joint
of the flagellum twice as long as the first, and half as long again
as the third. Pronotum transverse and linear; thorax as broad
as the head, sparsely punctured ; scutellum transverse. Median
segment short and rounded, the sides coarsely obliquely striated,
the triangular dorsal area coarsely longitudinally striated.
Abdomen subopaque, very minutely and closely punctured, more
than half as long again as the thorax and median segment com-
bined, the first segment half as broad at the apex as the second
segment and slightly constricted, the third segment nearly as
long as the second; the hypopygium pointed. Both recurrent
nervures received by the second cubital cell, separated from each
other by nearly two-thirds of the length of the cell; the first
transverse cubital nervure sends out a short obscure branch into
_ the first cubital cell; the second cubital cell is longer than the
third on the cubital nervure and is incompletely separated from
it, the second transverse cubital nervure only reaching a little
more than halfway to the radial nervure. The cubital nervure of
the posterior wing originates beyond the apex of the submedian
cell.
Black; the extreme apex of the scape of the antenne, the
pronotum, a rather narrow band at the apex of the first, second,
fourth, fifth, and sixth dorsal abdominal segments, very narrowly
interrupted on the first and fourth, yellow; the tegule, tibie,
502 MR. R. E. TURNER ON [May 12,
tarsi, and the extreme apex of the femora ferruginous. Wings
hyaline, nervures black.
Length 10 mm.
Hab. Victoria (french).
Type in British Museum.
Belongs to the mystacews group.
BEMBEX VARIABILIS Sm.
Bembex variabilis Sm. Cat. Hym. B. M. iv. p. 325, n. 39,1 856, 2.
Bembex crabroniformis Sm. Ann. Mag. Nat. Hist. xu. p. 296,
Woes Gc
Var. Bembex raptor Sm. Cat. Hym. B. M. iv. p. 326. n. 40,
1856, 3.
T have not been able to dissect a male of 5. raptor, but cannot
find any perceptible difference in structure between it and
variabilis. The colour-differences are not reliable, though to
a certain extent, at all events, local. The variety raptor occurs
at Roeburne, W. Australia, and in various localities in Central
Australia; the abdominal fascize are continuous, not interrupted
as in the typical form, which occurs along the Eastern Coast and
at Port Darwin. The second joint of the flagellum in the male is
a little shorter in var. raptor than in the typical form.
BEMBEX FLAVIPES Sm.
3. Colour as in the female, the clypeus white, black at the
base. Clypeus very prominent from the base, then vertically and
broadly semicircularly truncate and subconcave. The penultimate
joint of the flagellum is strongly produced beneath at the apex,
the apical joint strongly hooked and pointed. Anterior tarsi
normal, intermediate femora not serrate. Second ventral seg-
ment carinate longitudinally, the carina produced into a large
compressed tubercle, truncate broadly at the apex; the sixth
segment with a black triangular plate, poimted at the apex.
Hab. Mackay, Queensland (Turner).
BEMBEX LITTORALIS, sp. .
3. Clypeus moderately convex, clothed with very short pubes-
cence, which only shows in strong lights, in which it is silver ;
shining and finely punctured. Mandibles tridentate, the two
inner teeth very small, the labrum normal. Antenne inserted a
little further from each other than from the eyes, the front
between them slightly raised; the last joint of the flagellum
slightly curved, the second joint fully half as long again as the
third. Cheeks very narrow; the head finely punctured, with
sparse, long, grey pubescence. Thorax and abdomen finely and
closely punctured; the apical dorsal segment of the abdomen
broadly rounded and more sparsely punctured; the second ventral
segment rather coarsely punctured, with a prominent, compressed,
curved tubercle, pointed at the apex; the sixth segment with a
1908. | AUSTRALIAN FOSSORIAL WASPS. 503
raised triangular plate, pointed at the apex, the seventh with
a longitudinal median carina; the eighth segment terminating
in a short spine. Anterior tarsi simple, the basal joint with six
slender spines on the outer margin; the intermediate tibiz not
serrate.
Black; the mandibles (except at the apex), the labrum, the
extreme base of the scape, the flagellum beneath, a spot on each
side on the front below the anterior ocellus, the posterior margin
of the pronotum very narrowly, the tegule, the tibie and tarsi
beneath, and the femora beneath at the apex, testaceous brown ;
a narrow transverse band, broadly interrupted in the middle, on
abdominal segments 2-5 pale olivaceous grey. Wings hyaline,
nervures black.
Length 13 mm.
Hab. Port Darwin (Zurner); December.
Allied to B. museca Handl., and also to B. atrifrons Sm., of
which it may possibly prove to be the male, but the species in the
group are very closely allied, and atrifrons will probably prove to
be distinct.
BEMBEX TUBERCULIVENTRIS, Sp. Nl.
3. Eyes slightly divergent towards the clypeus. The clypeus
strongly convex at the base, very prominent at about one-third
from the base, thence almost vertically depressed to the apex,
shining and very sparsely punctured. Antenne inserted as far
from each other as from the eyes, the front between them with a
low longitudinal carina; the seventh joint of the flagellum very
Text-fig. 109.
Bembex, § : second ventral segment.
1. B.furcata Erichs. 2. B. tuberculiventris. 3. B. littoralis.
4, B. palmata Sm.
feebly prominent at the base beneath. Mandibles tridentate, the
intermediate tooth very small and short. Head finely punctured
and clothed with long grey pubescence. Thorax and abdomen
very finely and closely punctured, the punctures on the ventral
segments larger and shallow ; second ventral segment longi-
tudinally carinate, the carina produced downwards into a large,
compressed prominence, curved anteriorly and truncate broadly at
the apex; the sixth segment with a flattened triangular plate,
prominent and bluntly pointed at the apex; the seventh segment
504 MR. R. E. TURNER ON [May 12,
tricarinate longitudinally, broadened towards the apex; the
eighth segment very small and pointed. Anterior tarsi normal,
the basal joint with eight long and rather slender spines on the
outer margin; the intermediate tibie not serrate. Third cubital
cell broader at the apex than at the base, longer than the second.
The apical dorsal segment of the abdomen is broadly rounded at
the apex.
Black ; the mandibles (except at the apex), the labrum, the
clypeus (except a broad transverse mark at the base), the outer
orbits of the eyes narrowing to nearly the summit, the inner
orbits more broadly, not reaching to the summit, a median line
on the front broadening between and round the base of the
antenne, with a spot on each side of it at its base, the posterior
margin and the sides of the pronotum, with a large black spot
close to the posterior angles and a smaller one before it, a spot on
the mesopleuree below the anterior wings with a large curved
mark below it, the ventral segments of the abdomen (except the
base of the fifth and sixth segments and a black spot on each side
of the first and of the second at the base), the prosternum, the legs
(except the tarsal ungues, a black line at the base of all the tibize
and on the anterior and posterior femora), pale yellow. A narrow
transverse band interrupted in the middle and on the sides on the
first abdominal segment, a broader band (curved and narrowly
interrupted in the middle) on segments 2-6, and the sides of the
apical segment narrowly pale, dull, greenish grey. Wings
hyaline; nervures black, ferruginous at the base. Antenne
yellow beneath.
Length 13 mm., exp. 22 mm.
Hab. Cooktown, Queensland (Zurner) ; October.
Allied to B. museca Handl., but has the clypeus more prominent
and more spines on the anterior tarsi.
BEMBEX PALMATA Sm.
Bembex palmata Sm. Cat. Hym. B. M. iv. p. 325. n. 38, 1856, 3;
Handlirsch, Sitz. Akad. Wiss. Wien, cii. p. 751, n. 28,1893, ¢ @.
Bembea tridentifera Sm. Ann. & Mag. Nat. Hist. (4) xii. p. 298,
ISB, OQ
Hab. Mackay and Cairns, Queensland (Zurner).
BEMBEX PECTINIPES Handl.
Bembex pectinipes Handl. Sitz. Akad. Wiss. Wien, cii. p. 875.
n. 108, 1893, ?.
Bembex palmata Sm. Ann. & Mag. Nat. Hist. (4) xii. p. 298,
1873, 3 (nec Smith, 1856).
This is not identical with B. sausswrei Handl., in my opinion.
As in true palmata, the labrum has no median longitudinal sulcus
at the base, but the anterior tarsi of the female have twelve
spines on the outer sides of the first joints and the mandibles are
tridentate. In the male the basal joints of the anterior tarsi are
1908. | AUSTRALIAN FOSSORIAL WASPS. 505
very broad, with eighteen long spines on the outer margins, the
second joints produced into a broad lobe on the outer margins,
but the third and fourth not so. The sixth joint of the
flagellum is produced beneath at the apex and the seventh at
the base, the latter very gradually narrowed towards the apex
with an emargination in the middle. The second ventral segment
is longitudinally carinate with a tubercle at the apex, less strongly
produced than in flavifrons Sm. The intermediate femora are
not toothed, and the epipygium is truncate at the apex.
Hab. Port Darwin (Walker).
BEMBEX FLAVIFRONS Sm.
Bembex flavifrons Sm. Cat. Hym. B. M. iv. p. 324. n. 36,
1856, ©.
Bembex saussuret Handl. Sitz. Akad. Wiss. Wien, cil. p. 873.
n. 107, 1893, 9.
This may be distinguished from pectinipes Handl. by the
longitudinal grooves on the sides of the clypeus, feebly continued
on the labrum, and by a short median longitudinal sulcus at the
base of the labrum.
3g. The clypeus is very deeply longitudinally grooved on the
sides, the grooves strongly continued on the labrum, which also
has a deep, median, longitudinal sulcus near the base. The
antenne have the sixth joint of the flagellum strongly produced
beneath at the apex and the seventh at the base. The anterior
tarsi have the first joints strongl? broadened and flattened, with
sixteen long spines on the outer margin, the following joints are
not strongly lobed. The intermediate femora are distinctly
serrated. The second ventral segment is very strongly longi-
tudinally carinated beneath, the carina produced at the apex into
a very prominent tubercle, more strongly developed than in
pectinipes. The markings on the disc of the thorax are almost
obsolete, the band on the first abdominal segment very broadly,
on the other segments very narrowly interrupted. Epipygium
narrowly truncate at the apex.
Hab. Mackay, Queensland (Zurner).
Nysson (ACANTHOSTETHUS) PUNCTATISSIMUS, Sp. Ne (Plate XX VI.
fig. 9.)
@. Clypeus broadly truncate at the apex, the apical margin
narrowly transversely depressed, the angles very feebly produced ;
sparsely and rather deeply punctured, without carine, short and
broad. Antenne inserted nearly half as far again from the eyes
as from each other; the scape as long as the first two joints of
the flagellum combined, the second joint half as long again as the
first and about the same length as the third, the fourth joint longer
and thicker, the apical joint nearly twice as long as the second.
Eyes separated at the base of the clypeus by a distance equal
to about one-third of the length of the antenne, strongly diver-
gent towards the vertex and broadly and shallowly emarginate on
506 MR. R. E. TURNER ON [May 12,
the inner margin near the summit. The posterior ocelli a little
further from the eyes than from each other, and half as far again
from each other as from the anterior ocellus. Head rather closely,
but not very deeply punctured, the punctures large. A short
longitudinal carina on the front reaching from between the base
of the antenne much less than halfway to the anterior ocellus,
forked above, the branches reaching nearly halfway to the eyes.
Thorax coarsely punctured; the pronotum short and steeply
sloped anteriorly, very broadly emarginate posteriorly ; pro-
sternum produced posteriorly into two broad projections pointed
at the extremity and overlying the anterior coxe ; mesosternum
punctured, mesopleure coarsely punctured-rugose, the inter-
mediate coxze broadly, the posterior coxee narrowly separated.
Scutellum subrectangular, laterally produced at the anterior
angles, half as broad again as long ; postscutellum very short and
divided at distant intervals by longitudinal carine. Median
segment a little shorter than the scutellum, strongly produced at
the posterior angles into stout spines, strongly punctured; the
enclosed space bordered by strongly raised carine, converging
towards the apex, marked with about six rather lower longi-
tudinal carine; vertically truncate posteriorly, the face of the
truncation with several longitudinal carine. First abdominal
segment rather longer than the second, rounded broadly anteriorly
and strongly punctured, the second segment less deeply punc-
tured, the punctures on the remaining segments finer and very
shallow. Pygidial area granulated, broadly subtriangular, rather
narrowly truncate at the apex. First ventral segment with a
strong median carina, broadly emarginate at the apex and much
shorter than the second segment, which is coarsely but sparsely
punctured, the remaining segments finely punctured. The
posterior tibie are serrate from near the base, emarginate on
the outer margin, and smooth near the apex.
Black; the clypeus covered with very short silvery pubescence ;
the front and orbits of the eyes, pronotum, mesopleure, and
scutellum with very short pale golden pubescence ; the mesonotum
with silvery pubescence ; the sides of the median segment outside
the enclosed space with very pale golden pubescence ; the vertex,
mesonotum, and median segment dark blackish brown; the apical
margins of the abdominal segments above, most broadly at the
sides, luteous yellow, the base of the first segment with a large
rounded spot of very short golden pubescence on each side; the
legs, mandibles, antenne, tegule, and the apex of the pygidium
ferruginous. Wings hyaline, slightly iridescent; nervures black.
There are only two cubital cells; the second is pointed on the
radial nervure ; the first recurrent nervure is received at about
one-sixth from the apex of the first cubital cell, the second close
to the apex, almost interstitial with the first transverse cubital
nervure. ‘The angles of the ventral segments are not spined.
Length 9 mm.
Hab. Mackay, Queensland (Turner); February.
1908. ] AUSTRALIAN FOSSORIAL WASPS. 507
Nysson (ACANTHOSTHETHUS) SPINIGER, sp. 0.
@. Clypeus depressed to the apex, and broadly but shallowly
emarginate anteriorly, clothed with very short silvery pubescence.
Antenne inserted a little further from each other than from the
eyes; the three apical joints of the flagellum much thickened, the
apical joint twice as long as the second, which is longer than
the third. Eyes separated from each other at the base of the
elypeus by a distance equal to about two-fifths of the length of
the antenne, strongly divergent towards the vertex, but not
emarginate; the posterior ocelli half as far again from the eyes
as from each other. Head strongly and very closely punctured ;
an indistinct, very short, longitudinal carina between the
antenne; the front and pronotum with very short and sparse
white pubescence. Thorax punctured-rugose; the pronotum
short, gradually sloped, the anterior angles prominent ; the meso-
pleurz strongly and closely, the mesosternum feebly and sparsely
punctured. Scutellum subrectangular, very little broader than
long. Median segment very short, in the middle only about
half as long as the scutellum; the posterior angles strongly
produced, ending in an acute spine directed outwards and back-
wards, the enclosed space with convergent lateral carinze and
marked with about six longitudinal carine ; vertically truncate
posteriorly, with several longitudinal carine on the surface of
the truncation; the sides of the segment finely and sparsely
punctured, the dorsal surface outside the enclosed area clothed
with silvery pubescence. Abdomen closely punctured ; the first
segment longer than the second, subtruncate at the base, and
more strongly punctured than the second segment; the pygidial
area granulate, subtriangular, and narrowly truncate at the apex.
The first ventral segment with a very strong median carina and
a long spine on each side at the apical angles; the second
segment much longer than the first and strongly punctured.
Posterior tibize almost unarmed, slightly emarginate near the
apex.
Black; the mandibles, the basal third of the antenne, the
tegule, the legs, the basal segment of the abdomen, the apical
margin of the remaining segments obscurely, and the sides and
ventral surface of the pygidium, dark ferruginous; a short,
obscure, transverse band on the sides of the first and second
abdominal segments yellow. Wings hyaline, nervures black.
The second cubital cell is almost petiolate, the first recurrent
nervure is received by the first cubital cell at about one-fifth from
the apex, the second close to the apex, almost interstitial with
the first transverse cubital nervure. The anal cell of the hind
wing is short.
3. As in the female, but without the spine at the angles of
the first ventral segment, and the apical segment is widely
emarginate at the apex.
Length 5 mm.
Hab. Mackay, Queensland (Z'urner) ; October.
508 MR. R. E. TURNER ON [May 12,
NITELA KURAND4, Sp. 0.
@. Clypeus strongly convex, with an elevated carina from the
base to the apex finely continued on the front, and reaching
halfway from the base of the clypeus to the anterior ocellus.
Byes more strongly convergent towards the vertex than in other
species of the genus ; the posterior ocelli very close to the eyes,
fully twice as far from each other as from the eyes. Head
opaque, the vertex almost smooth, the front very minutely and
closely punctured-striate. Pronotum | much narrower than the
head, the anterior and posterior margins raised, the short space
between indistinctly transversely striated, the tubercles at the
posterior angles reach back to the tegule. Mesonotum and
scutellum opaque. Median segment longer than the mesonotum,
narrowed towards the apex and vertically truncate posteriorly,
coarsely reticulate, with a faint median carina; the face ot the
truncation transversely striated. Abdomen smooth and shining,
the second segment slightly transversely depressed at the base, the
apical segment compressed and pointed.
Black ; the scape of the antenne, the basal half of the flagellum,
the mandibles, the tegulz, and the legs pale ferruginous. Wings
hyaline, iridescent ; nervures testaceous. The recurrent nervure
is almost interstitial with the transverse cubital nervure.
Length 5 mm.
Hab. Cairns, Queensland (Zurner) ; January.
NITELA RETICULATA, Sp. 0.
@. Clypeus strongly convex, with an elevated carina from the
base to the apex, shining and very minutely punctured. Front
rugose, vertex almost smooth with a few indistinct transverse
strie ; the eyes moderately convergent towards the vertex, the
posterior ocelli about half as far again from each other as from
the eyes; a delicate carina from the anterior ocellus not reaching
the base of the clypeus. Pronotum short and transverse, much
narrower than the head, the anterior and posterior margins
strongly raised, the short intervening space strongly rugose and
divided in the middle by a carina. Mesonotum coarsely rugose,
irregularly transversely striated anteriorly, with an indistinct
carina from the middle to the posterior margin. Median segment
longer than the mesonotum, narrowed towards the apex and
vertically truncate posteriorly, longitudinally striated, the space
between the strize very coarsely punctured, giving an appearance
of coarse reticulation. Abdomen smooth and shining, shorter
than the thorax and median segment combined ; the first segment
long, truncate at the base, the second segment strongly depressed
at the base, the apical segment pointed.
Black; the mandibles, the antennz, except the four apical
joints which are fuscous, the tegule and the legs ferruginous.
Wings hyaline, nervures pale ferruginous. The recurrent nervure
is interstitial with the transverse cubital nervure.
1908. ] AUSTRALIAN FOSSORIAL WASPS. 509
Length 5 mm.
Hab. Mackay, Queensland (Zurner) ; May.
The antenne are inserted very low down close to the sides of
the clypeus. The tubercles at the posterior angles of the pro-
notum do not reach the tegul, differing in this point from
typical Vitela.
PISON SCABRUM, sp. n.
2. Clypeus rather more than twice as broad as long; the
anterior margin broadly rounded, narrowly and bluntly produced
in the middle, closely punctured and clothed with long greyish
pubescence, changing to silver in strong lights. Antenne inserted
about as far from each other as from the eyes, as long as the
thorax and median segment combined, very slightly thickened to
the seventh joint of the flagellum; the second joint of the
flagellum three times as long as the first and one-third longer
than the third. Eyes deeply emarginate, the distance between
them on the vertex slightly exceeding the length of the two basal
joints of the flagellum combined, the distance between them at
the base of the mandibles nearly half as great again as on the
vertex. Posterior ocelli nearly half as far again from the eyes
as from each other, a little nearer to each other than to the
anterior ocellus. Head closely and rather finely punctured, thinly
clothed with cinereous pubescence ; a short, longitudinal, median
carina on the front above the base of the antenne. Pronotum
very short, depressed below the level of the mesonotum. Thorax
shining, closely punctured, with an obscure longitudinal carina
on the scutellum. Median segment not more than two-thirds
of the length of the mesonotum, punctured-rugose, without a
median carina, with a small shining spot at the apex, truncate
posteriorly, with a median sulcus not reaching the apex, and with
a few transverse striz at the apex. Abdomen very minutely and
closely punctured, the third segment as long as the second, the
apical margin of the segments depressed, with a band of silvery
pubescence, the apical segment elongate and very sharply
pointed.
Black; the tegule, the apical margin of the fifth abdominal
segment, and the spines of the posterior tibize fusco-ferruginous.
Wings hyaline, faintly tinged with fuscous, a little darker at the
apex ; nervures black.
The recurrent nervures are interstitial with the first and
second transverse cubital nervures; the second cubital cell reaches
halfway from the cubital to the radial nervure; the third cubital
cell is more than half as long on the radial as on the cubital
nervure.
Length 15 mm., exp. 25 mm.
Hab. Mackay, Queensland (Zune).
This is allied to P. spinole, but differs in the sculpture of the
median segment, in the stronger punctures on the head and
thorax, and the greater breadth between the eyes. The clypeus
510 MR. R. E. TURNER ON [May 12,
is also broader. Also allied to P. fuscipenne Sm. from W. Aus-
tralia and P. nitidwm Sm. from Mysole, but the distance between
the eyes is much greater than in either species, and the antenne
are much longer than in Suscipenne.
PISON INSULARE Sm. st. PRISCUM, n. st.
2. Clypeus nearly twice as broad as long, bluntly produced in
the middle of the anterior margin. Antenne as long as the
thorax and median segment combined, inserted a little further
from each other than from the eyes ; the second joint of the
flagellum a little longer than the third, and about two and a halt
times as long as the first. HKyes narrowly and deeply emarginate,
fully half as far again from each other at the base of the
mandibles as on the vertex, the distance between them on the
vertex exceeding the length of the second joint of the flagellum.
The posterior ocelli as far from the eyes as from each other.
Head opaque, the clypeus and the front round the base of the
antenne clothed with shining white pubescence. Thorax shining,
minutely punctured. Median segment shorter than the meso-
notum, almost vertically truncate posteriorly, very finely and
sparsely punctured, shining, with an almost obsolete median
sulcus; the face of the truncation with a deep depression at the
base, finely punctured at the base, with a few indistinct trans-
verse strize at the apex. Abdomen shining, minutely punctured,
the first segment oblique and slender at the base, much longer
than the second, the second segment not depressed at the base ;
the apical margin of the segments feebly depressed, with a little
greyish-white pubescence on the sides.
Entirely black. Wings hyaline, faintly tinted with fuscous,
especially at the apex, and slightly iridescent ; nervures black.
The first recurrent nervure is received close to the apex of the
first cubital cell, the second is interstitial with the second trans-
verse cubital nervure. The second cubital cell does not reach
halfway from the cubital to the radial nervure; the third is
about half as long on the radial as on the cubital nervure.
Length 12 mm., exp. 20 mm.
Hab. Mackay, Queensland (Zurner); November.
The eyes on the vertex of typical P. insulare, from the New
Hebrides, are nearer together than the length of the second joint
of the flagellum; the clypeus is not bluntly produced in the
middle of the apical margin ; the median segment is less abruptly
truncate posteriorly, and the median sulcus on it is well defined.
The wings are also more hyaline.
These “differences hardly seem of full specific importance, though
quite sufficient to constitute a good geographical race.
PISON INFUMATUM, sp. 0.
2. Clypeus twice as broad as long, bluntly produced on the
middle of the apical margin, thinly covered with short grey
pubescence, Head opaque, with a very obscure longitudinal sulcus
1908. | AUSTRALIAN FOSSORIAL WASPS. 511
on the front. Hyes deeply emarginate, separated on the vertex
by a distance equal to the length of the two basal joints of the
flagellum, more than half as far again from each other at the base
of the mandibles; the posterior ocelli very near the eyes, nearly
twice as far from each other, but much nearer to each other than
to the anterior ocellus. Antenne inserted a little nearer to each
other than to the eyes; the second joint of the flagellum more
than twice as long as the first and about one-third longer than
the third. Pronotum not much more than half as broad as the
head, steeply depressed anteriorly, the posterior margin straight.
Thorax subopaque. Median segment almost smooth, very minutely
punctured, with very short striz at the base, and a very obscure
median carina from the base to theapex. Abdomen very minutely
punctured, the apical margin of the segments broadly depressed,
the second segment not depressed at the base.
Entirely black ; the tegule testaceous brown; the pubescence
white on the inner margin of the eyes below the emargination,
on the sides of the postscutellum, on the sides of the median
segment near the apex, and very sparsely on the sides of the
abdominal segments. Wings hyaline at the base, the apical two-
thirds fusco-hyaline ; nervures black.
The recurrent nervures are almost interstitial with the first
and second transverse cubital nervures. The second cubital cell
extends more than halfway from the cubital to the radial
nervure ; the third is about three times as long on the cubital as
on the radial nervure.
Length 7 mm., exp. 10 mm.
Hab. Port Darwin (Turner); December.
PISON IGNAVUM, sp. n.
@. Clypeus broadly rounded anteriorly, about twice as broad
at the apex as long, clothed with shining white pubescence.
Antenne inserted about the same distance from each other as
from the eyes, about twice as long as the mesonotum, thickened
to the eighth joint of the flagellum; the second joint of the
flagellum a little shorter than the scape, fully twice as long as
the first joint of the flagellum and very slightly longer than tne
third. Eyes deeply emarginate, about one-third further from
each other at the base of the mandibles than on the vertex, the
distance between them on the vertex equal to about twice the
length of the second joint of the flagellum; the posterior ocelli
nearer to each other than to the anterior ocellus, a little further
from each other than from the eyes, with a depressed transverse
line on the vertex behind them. Head opaque, a very faint
longitudinal suleus below the anterior ocellus, the front round the
base of the antenne and the inner orbits of the eyes as high as
the emargination clothed with shining white pubescence. Thorax
minutely punctured ; the pronotum transverse, with white pubes-
cence ; the mesonotum more than half as broad again as long.
Median segment a little shorter than the mesonotum, broader at
Proc. Zoou. Soc.—1908, No. XX XITI. 33
512 MR. R. E. TURNER ON [May 12,
the base than long, the sides steeply sloped, truncate posteriorly,
obliquely striated, with a longitudinal median carima: the face
of the truncation rather coarsely transversely striated, with a
median sulcus, the sides of the segment raised, forming carine, a
little white pubescence at the apex. Abdomen ovate, closely and
minutely punctured, the second segment transversely depressed
at the base; the apical margin of all the segments feebly depressed,
most broadly in the middle, with interrupted bands of silvery
pubescence.
Black; the spines of the tibie testaceous ; tegule fuscous.
Wings hyaline, faintly clouded at the apex, nervures black.
First recurrent nervure received near the apex of the first
cubital cell, second at the middle of the second cubital cell.
The second cubital cell reaches more than halfway from the
cubital to the radial nervure; the third is extremely short on the
radial nervure.
3. Similar in all respects to the female.
Length, 9 9mm., ¢ 7 mm.; exp., 9 14 mm., ¢ 12 mm.
Hab. Mackay and Cairns, Queensland (7wrner); March. Also
from Melbourne.
Near P. westwoodi Shuck. from Tasmania, but differs in the
presence of a carina on the median segment and in the position
of the recurrent nervures. I have not seen the type of P. west-
woodi, nor any specimen quite answering to the description.
The present species is also near P. dridipennis Sm. from Hawaii,
but the antenne are longer and the eyes further apart on the
vertex, the clypeus is also different. P. pallidipalpe Sm., from
Ceram, is a larger species, and has the clypeus very different.
PISON IRIDIPENNE Sm.
Described by Smith from Honolulu.
Hab. Mackay and Cairns, Queensland (7'urner).
In the female the posterior ocelli almost touch the eyes, which
are very close together on the vertex.
Pison puNCTULATUM Kohl.
Pison punctulatum Kohl, Verh. zool.-bot. Ges. Wien, xxxiii.
p. 336, 1883, nee Cameron.
Hab. Peak Downs, Queensland (Xoh/); Mackay, Queensland
ib hers my
(Turner).
PIsON AURIVENTRE, sp. Nn.
Q@. Clypeus very short, at least three times as broad as long,
convex, clothed with golden pubescence, changing to silver at the
angles. Antenne nearer to the eyes than to each other; the
second joint of the flagellum scarcely longer than the third, about
twice as long as the first. Eyes shallowly and rather broadly
emarginate, the depth of the emargination about equal to the
length of the first joint of the flagellum ; about half as far again’
from each other at the base of the mandibles as on the vertex.
1908. ] AUSTRALIAN FOSSORIAL WASPS. iB:
Posterior ocelli as far from the anterior ocellus as from each
other, nearly half as far again from each other as from the eyes.
Head opaque, the front closely covered with golden pubescence.
Pronotum about two-thirds of the breadth of the head, the
posterior margin straight, the angles not rounded, covered with
pale shining pubescence. Thorax opaque. Median segment as
long as the mesonotum, with sparse golden pubescence, silvery on
the sides, obscurely obliquely striated, with a median carina; the
posterior truncation irregularly transversely striated, with a
median sulcus, the sides closely and finely punctured. Abdomen
ovate, very minutely and closely punctured, the sides and apical
margins of the segments covered with short golden pubescence,
the pubescence at the apical angles of the basal segment silvery.
The first segment is subtruncate at the base, the second is the
longest and broadest.
Black; the mandibles ferruginous; the spines of the tibie, the
tarsal ungues, and the apical margin of the abdominal segments
very narrowly fusco-ferruginous. Wings hyaline, iridescent,
clouded with fuscous at the apex ; the nervures black.
The first recurrent nervure is received by the first cubital cell
very near the apex, the second is interstitial with the second
transverse cubital nervure. The third cubital cell is three times
as long on the cubital as on the radial nervure.
Length 8 mm.
Hab. Victoria (French).
Aled to P. marginatum Sm., from which it may be dis-
tinguished by the more shallow emargination of the eyes, the
front is also much narrower. As in marginatum the two or
three apical joints of the antenne are slenderer than the pre-
ceding ones. The allied West-Australian species ¢ibiale is a much
stouter built insect with a much shorter median segment. The
median segment is also longer than in P. simillimum Sm., in
which species the eyes are deeply emarginate as in P. marginatum.
PISON VIROSUM, sp. n.
Q@. Clypeus large, not more than twice as broad as long at the
broadest point, narrowed towards the apex, where it is sub-
truncate, feebly convex, clothed with dull golden pubescence.
Head and thorax opaque; the front broad, with a shallow, longi-
tudinal, median sulcus above the base of the clypeus, not nearly
reaching the anterior ocellus. Eyes separated at the base of the
clypeus by a distance half as great again as that separating them
on the vertex; the emargination shallow and rather broad, not
equal in depth to the length of the first joint of the flagellum.
Posterior ocelli a little further from each other than from the
eyes, with a delicately impressed transverse line above and
touching them. Antenne as long as the thorax and median
segment combined, inserted near together, twice as far from the
eyes as from each other, very slightly thickened to the apex;
the second joint of the flagellum as long as the scape, twice as
514 MR. R. E. TURNER ON [May 12,
long as the first joint and nearly half as long again as the third.
Pronotum straight, strongly depressed anteriorly, about two-
thirds of the width of the head. Median segment not quite as
long as the mesonotum, narrowed and almost vertically truncate
posteriorly, closely obliquely striated, with a median cara from
the base to the apex; the face of the truncation transversely
striated, with a deep median sulcus. Abdomen very minutely
punctured, the apical margin of the segments rather broadly
depressed, the second segment a little longer than the first or
third, the first subtruncate at the base.
Black; the front covered with fine golden pubescence, the
posterior margin of the pronotum with fine silvery pubescence ;
the antenne (except the apical joint), the mandibles, tegule, tarsi,
tibie, the apex of the femora, and the abdomen (except the base
of the first segment) ferruginous. Wings hyaline, faintly iridescent ;
nervures fuscous.
The first recurrent nervure is received close to the apex of the
first cubital cell, the second at the middle of the second cubital
cell. The third cubital cell is twice as long on the cubital as on
the radial nervure, and the second cubital cell reaches a little
more than halfway from the cubital to the radial nervure.
3. The clypeus produced into a pomt on the middle of the
apical margin, the pubescence on the front paler than in the
female, and the apical abdominal segment narrowly emarginate at
the apex. Otherwise as in the female.
Length, 2 9 mm., ¢ 7 mm.
Hab. Mackay, Queensland (Turner); September to February.
This seems to be near P. pelletiert Le Guillou, but is a smaller
species and the sculpture of the median segment is very different.
I have not seen P. pelletieri.
PISON RUFICORNE Sm.
2. Clypeus large, not more than half as broad again as long,
truncate at the apex, clothed with short silvery pubescence, very
slightly convex. Head and thorax opaque, the front above the
base of the antenne and the inner margin of the eyes below
the emargination clothed with silvery pubescence. Eyes more
strongly convergent towards the clypeus than towards the vertex,
separated on the vertex by a distance about one-third greater
than the distance separating them at the base of the elypeus, the
emargination narrow and exceeding in depth the length of the
first joint of the flagellum by about one-half. Antenne inserted
a little nearer to the eyes than to each other; the second joint of
the flagellum as long as the scape, twice as long as the first joint
but only slightly longer than the third, the apical joints
moderately thickened. Pronotum very short, the posterior
margin straight, depressed below the level of the mesonotum,
about one-quarter narrower than the head,.and thinly clothed
with short white pubescence. Median segment rather shorter
than the mesonotum, narrowed and vertically truncate posteriorly.
neti ain att ale
1908. | AUSTRALIAN FOSSORIAL WASPS. 515
rather coarsely obliquely striated, with a rather ill-defined median
carina from the base to the apex; the surface of the posterior
truncation transversely striated, with a deep median sulcus.
Abdomen not quite as long as the thorax and median segment
combined, almost smooth, the apical margin of the segments de-
pressed, with a little fine pubescence on the sides; the first
segment subtruncate at the base, the second a little longer than
the first or third, transversely depressed at the base.
Black ; the mandibles, palpi, antenne (except at the extreme
apex), tegule, tibize, tarsi, the apex of the femora, and the abdo-
men (except the basal half of the first segment and the extreme
base of the second) ferruginous. Wings hyaline, nervures black.
The first recurrent nervure is received by the first cubital cell
about one-fifth before the apex, the second at the middle of the
second cubital cell. The second cubital cell does not reach quite
halfway to the radial nervure from the cubital nervure ; the third
cubital cell is a little less than twice as long on the cubital as
on the radial nervure.
Length 8 mm.
Hab. Victoria (french).
In another specimen the second, third, and fourth abdominal
segments are strongly stained with black. The type was from
the Macintyre River, Queensland, and has much more black on the
abdomen. ‘The specimen in the British Museum collection is from
that locality and is probably the type, though not marked. I have
taken the description from a Victorian specimen. Jn Queensland
specimens the eyes are nearly, if not quite, as far apart on the
clypeus as on the vertex. In specimens from Mackay the median
segment is much more finely striated and the abdomen is brighter
in colour and entirely ferruginous. These differences, although
apparently constant, do not seem to me suflicient to deserve even
subspecific rank; it is probable that connecting forms will be
found to occur in intermediate localities.
PisON MELANOCEPHALUM, sp.n. (Plate X XVI. fig. 12.)
Q. Clypeus subtriangular, truncate at the base, convex,
obliquely triangularly depressed from the centre to the apex.
Head opaque, almost smooth, with a shallow longitudinal sulcus
reaching from the anterior ocellus to near the base of the clypeus.
Eyes deeply and narrowly emarginate, convergent towards the
base of the antenne, where they are separated by a distance about
equal to the length of the scape of the antenne, separated on the
vertex by a distance nearly twice as great; the posterior ocelli
nearly twice as far from each other as from the eyes. Antenne
inserted at the base of the clypeus, very close to the eves; the
scape short, about equal in length to the two basal joints of the
flagellum ; the second and third joints of the flagellum about equal
to each other in length, each more than half as long again as the
first joint, the apical joints slightly thickened. The eyes reach to
the posterior margin of the head. Pronotum about two-thirds of
516 MR. R. E. TURNER ON [May 12,
the width of the head, strongly depressed anteriorly. Thorax and
median segment impunctate, subopaque; the median segment
slender, half as long again as broad at the base, narrowed pos-
teriorly and oblique, with a median sulcus from the base not
reaching the apex. First abdominal segment slender at the base,
with a longitudinal sulcus not reaching the apex, nearly as long
as the second segment; the second and third segments strongly
transversely depressed at the base. The tarsi are without spines.
Light ferruginous brown ; the head black ; the antennz (except
the two apical joints) castaneous. Wings hyaline, nervures pale
ferruginous.
The second cubital cell is very small, only extending along the
transverse cubital nervure for less than one-third of its length,
the second recurrent nervure is received close to the base of the
cell; the first cubital cell more than three times as long as the
third on the cubital nervure, receiving the first recurrent nervure
beyond two-thirds from the base.
Length 5 mm.
Hab. Cairns, Queensland (Zurner); February.
This is closely allied to some of the species of the Parapison
section in which the second cubital cell is wholly absent. The
strong convergence of the eyes towards the base of the clypeus is
remarkable.
Prison (PARAPISON) NOCTULUM, Sp. Nn.
2. Clypeus narrowly convex in the middle, nearly twice as
bread as long, the apical margin very. broadly rounded, clothed
with very short silvery pubescence. Antennz as long as the head,
thickened towards the apex, inserted a little further from each
ether than from the eyes; the second joint of the flagellum twice
as long as the first and half as long again as the third. Eyes
rather deeply emarginate, half as far again from each other at the
base of the mandibles as on the vertex, separated on the vertex
by a distance about equal to the length of the two basal joints of
the flagellum. Head opaque, with a delicate longitudinal sulcus
below the anterior ocellus. Posterior ocelli half as far again from
each other as from the eyes, and as far from each other as from
the anterior ocellus. Thorax opaque, very minutely punctured ;
pronotum narrowed anteriorly and oblique, longer than in most
of the species of the genus, the posterior margin as broad as the
head and on a level with the mesonotum. Median segment
shorter than the mesonotum, narrowed and truncate posteriorly,
very delicately transversely striated, with a small smooth tri-
angular mark at the apex, from the apex of the triangle a carina
reaches almost to the apex of the segment; the surface of the
truncation is almost smooth, with a broad median suleus. Abdo-
men almost smooth, the apical margin of the segments depressed,
with sparse grey pubescence on the sides; the second segment
feebly transversely depressed at the base, equal in length to the
third. The mesopleure are opaque, very minutely punctured,
] 908. | AUSTRALIAN FOSSORIAL WASPS. 517
with the longitudinal sulcus below the base of the wings much
more feebly marked than in P. erythrocerwm Kohl.
Black ; the tegule fuscous. Wings hyaline, faintly clouded at
the apex ; nervures fuscous.
Length 7 mm.
Hab. Mackay, Queensland (Zwirner) ; February.
The pronotum is shaped as in erythrocerwm Kohl, but more
strongly narrowed anteriorly, and much less depressed than in
other species,
PIsoN (PARAPISON) PERTINAX, sp. n.
Q. Head and thorax opaque, not visibly punctured, a short and
obscure longitudinal sulcus below the anterior ocellus. Clypeus
short, fully twice as broad as long, convex, very broadly rounded
anteriorly, and covered with short silvery pubescence. Antenne
a little longer than the head, inserted at the base of the clypeus,
nearly half as far again from each other as from the eyes; the
scape short, scarcely longer than the second joint of the flagellum,
which is about half as long again as the first or third; the first
joint is stout and equal in length to the third. Eyes narrowly
emarginate, the depth of the emargination about equal to the
length of the first joint of the flagellum; the distance between the
eyes on the vertex equal to about three-quarters of the distance
between them at the base of the clypeus. The posterior ocelli a
little nearer to the eyes than to each other, situated very near to
the posterior margin of the head, which is slightly transversely
raised behind them. Pronotum short and strongly depressed
anteriorly, the posterior margin straight; scutellum broadly
truncate at the apex. Median segment rather slender, much
longer than broad, a little longer than the mesonotum, delicately
obliquely striated, a carina from the base to the apex lying in a
depressed sulcus, the lateral margins of the segment marked by a
carina; narrowed posteriorly and abruptly truncated, the surface
of the truncation transversely striated, with a deep median sulcus.
Abdomen as long as the thorax and median segment combined,
shining; the apical margins of the segments depressed, most broadly
in the middle, the second segment constricted at the base, longer
and broader than the first or third.
Black; the mandibles, antenne (except the two apical joints),
tibie, tarsi, and abdomen (except the middle of the third segment)
ferruginous. Wings hyaline, iridescent; nervures black.
The first recurrent nervure is received by the first cubital cell
beyond two-thirds from the base, the second at the extreme base
of the second cubital cell, which is almost pointed on the radial
nervure, being less than a quarter of the length of the cell on the
cubital nervure.
Length 7 mm.
Hab. Mackay, Queensland (Zurner); January to May.
In some specimens the abdomen is wholly ferruginous.
518 MR. R. E. TURNER ON [May 12,
Pigon (PARAPISON) TENEBROSUM, Sp. 0.
@. Mandibles acute at the apex ; clypeus short, more than
twice as broad as long, narrowly convex or subcarinate in the
middle, with a small blunt tubercle just before the apex, sub-
truncate at the apex, clothed with short silvery pubescence. Head,
thorax, and abdomen opaque; the eyes narrowly and shallowly
emarginate, separated on the clypeus by a distance about one-
quarter greater than that separating them on the vertex. Antenne
scarcely longer than the head, inserted a little further from. each
other than from the eyes, the scape as long as the first two joints
of the flagellum combined; the first joint of the flagellum two-
thirds of the length of the second, the second and third almost.
equal. Posterior ocelli twice as far from each other as from the
eyes. Pronotum almost straight, transverse, about three-quarters
of the breadth of the head; mesonotum with a broad, shallow,
longitudinal, median depression not reaching the posterior margin.
Median segment a little shorter than the mesonotum, narrowed
from the base, obliquely truncate posteriorly, with a longitudinal
carina from the base lying in a shallow depression and not
reaching the apex, obscurely obliquely striated ; the face of the
truncation minutely and very closely punctured, with a deep
median sulcus. The sides and apical margin of the abdominal
segments with sparse grey pubescence. The first segment sub-
truncate at the base, nearly as long and broad as the second.
Black; the mandibles dark ferruginous; the spines of the tibie
testaceous; the tegule, the anterior tibie, and the tarsi fuscous.
Wings hyaline, nervures fusco-ferruginous.
The second cubital cell is small, triangular, pointed on the
radial nervure, less than one-third of the length of the first on
the cubital nervure, receiving the second recurrent nervure near
the base. The first cubital cell receives the first recurrent nervure
beyond two-thirds from the base.
Length 5 mm., exp. 8 mm.
Hab. Mackay, Queensland (Zurner); January.
Pison (PARAPISON) CALIGINOSUM, Sp. .
@. Mandibles short and very broad, shallowly and widely
emarginate at the apex. Clypeus half as broad again as long,
very feebly convex, strongly rounded at the apex. Head and
thorax opaque, very minutely and closely punctured ; the antennze
inserted almost as near to each other as to the eyes, the scape about.
equal in length to the first two joints of the flagellum, the second
joint of the flagellum more than half as long again as the first and
a little longer than the third. The emargination of the eyes
exceeding in depth the length of the first joint of the flagellum ;
the eyes at the base of the clypeus nearly half as far again from
each other as on the vertex. Posterior ocelli nearer to each other
than to the anterior ocellus, but nearer to the eyes than to each
other. Pronotum not more than two-thirds of the breadth of the
1908. ] AUSTRALIAN FOSSORIAL WASPS. 519
head, very short, the posterior margin straight. Median segment
as long as the mesonotum, narrowed str ongly and ab: uptly
truncate posteriorly, with a median carina lying in a narrow
depression not quite reaching the base of the truncation, delicately
obliquely striated, punctured between the striz ; the surface of the
truncation with a deep depression at the base, finely punctured at:
the sides, finely transversely striated near the apex. Abdomen
shining, very minutely punctured, the first segment slender at the
base, as long as the second, and at the apex about two-thirds of the
breadth of the second; the second segment transversely depressed
at the base and longer than the third.
Entirely black, with a little sparse silvery pubescence on the
orbits of the eyes and the sides of the median segment and
abdomen ; a little fulvous pubescence on the sides of the two 0 apical
segments ; the silvery pubescence very short and fine, the fulvous
coarser. Wings hyaline, iridescent, faintly clouded at the extreme
apex ; nervures black.
The second cubital cell is very short on the radial nervure, but.
half as long as the first on the cubital nervure, receiving the
second recurrent nervure before the middle. The first cubital cell
receives the first recurrent nervure at about one-sixth from the
apex.
Length 7 mm., exp. 11 mm.
Hab. Kuranda near Cairns, Queensland (Zwrner) ; February.
Pison (PARAPISON) ABERRANS, Sp. 0.
6. Antenne no longer than the head, moderately thickened to
the apex, about the same distance from each other at the base
as from the eyes; the second joint of the flagellum scarcely longer
than the first, the scape longer than the first three joints of the
flagellum combined. Clypeus nearly three times as broad at the
apex as long, the apical margin almost straight, clothed with
shining silvery pubescence. Head and thorax opaque; an obscure,
impressed, longitudinal line on the front below the anterior
ocellus; the front round the base of the antenne and the inner
orbits of the eyes below the emargination clothed with silvery
pubescence. Eyes narrowly and rather shallowly emarginate, the
emargination hardly equalling in depth the length ‘of the first
joint of the flagellum; the distance between the eyes at the base
of the clypeus exceeding by more than one-third the distance
between them on the vertex; the posterior ocelli nearly twice
as far from each other as from the eyes. Pronotum strongly
depressed anteriorly, the posterior margin very broadly arched.
Scutellum smooth and shining, broadly truncate at the apex.
Median segment longer than broad, finely obliquely striated,
longitudinally depressed in the middle, with a carina in the
depression, truncate posteriorly; the surface of the truncation
transversely striated, with a median sulcus. Abdomen shining,
microscopically punctured, the segments feebly but broadly
depressed on the apical margin; the second segment transversely
520 MR, R. E. TURNER ON [May 12,
depressed at the base, broader but hardly longer than the first and
third.
Black; the mandibles and the tarsi ferruginous brown. Wings
hyaline, iridescent ; nervures black.
The first recurrent nervure is received by the first cubital cella
little before two-thirds from the base, the second at the extreme
base of the second cubital cell. The second cubital cell is extremely
small, almost pointed on the radial nervure, and not more than
one-sixth of the length of the first cubital cell on the cubital
nervure. On one side of the type specimen the second transverse
cubital nervure is absent, leaving the cell open.
Length 4 mm.
Hab. Mackay, Queensland (7urner); January.
Pison (AULACOPHILUS) DIFFICILE, sp. n.
@. Clypeus slightly produced and pointed in the middle of the
apical margin, nearly twice as broad at the apex as long, thinly
clothed with pale fulvous pubescence. Antenne inserted just
above the base of the clypeus, a little further from each other than
from the eyes, as long as the thorax and median segment
combined; the apical jomts not at ail thickened ; the second joint
of the flagellum longer than the third joint or than the scape.
Eyes rather deeply emarginate, convergent towards the vertex,
where they are separated by a distance equal to the length
of the second joint of the flagellum; the posterior ocelli nearer to
Text-fig. 110.
Pison (Aulacophilus) difficile.
the eyes than to each other. Head opaque, about equal in breadth
to the mesonotum, the front thinly clothed with short, pale,
golden pubescence. Pronotum short, strongly depressed and
clothed with pale golden pubescence. Thorax and median
segment very delicately punctured. The median segment longer
than the mesonotum, strongly narrowed and depressed to the
apex, with a deep sulcus from the apex nearly reaching the middle,
the sides with thin greyish pubescence. Abdomen petiolate,
pubescent ; the first segment twice as broad at the apex as at the
base, half as long again as the second segment, the third segment
1908. | AUSTRALIAN FOSSORIAL WASPS. 521
a little broader than the second; the apical margin of segments
1-4 with a band of pale golden pubescence interrupted in the
middle.
Black; the mandibles at the base, the antenne (except the two
apical joints), the tegule, the base and apex of the first abdominal
segment broadly, the apical margin of the remaining segments, the
tarsi, tibie, and the apex of the femora rufo-testaceous. Wings
hyaline, nervures dull rufo-testaceous.
The first cubital cell very long, receiving the first recurrent
nervure very near the apex; the second cubital cell very short
on the radial nervure, receiving the second recurrent nervure near
the base, less than three times as long on. the cubital as on the
radial nervure.
Length 11 mm., exp. 19 mm.
Hab. Mackay, Queensland (Zwrner).
This species resembles the following one P. icarioides in shape,
but the antenne are much longer and are not thickened at the
apex, and the shape of the second cubital cell is very different. It
does not approach so nearly to 4 ulacophilus, forming a connecting
link with true Pisoz.
Prison (AULACOPHILUS) ICARIOIDES, sp.n. (Plate XX VI. fig. 13.)
2. Clypeus broadly rounded at the apex, more than twice as
broad as long, clothed with rather long, shining, very pale
golden pubescence. Front round the base of the antenne, and
the inner orbits of the eyes as high as the emargination, clothed
with short silvery pubescence; a short, obscure, median carina
above and between the base of the antenne; the vertex opaque.
HKyes rather shallowly and broadly emarginate, the distance between
them at the base of the mandibles about one-third greater than on
the vertex ; the posterior ocelli almost as far from each other as
from the eyes. Antenne inserted a little further from each other
than from the eyes, scarcely longer than the head, much thickened
to the apex ; the second joint of the flagellum less than twice as long
as the first and one-third longer than the third. Thorax, median
segment, and abdomen opaque, very minutely and closely punc-
tured; the pronotum nearly as broad as the head, very gradually
depressed anteriorly; scutellum short and transverse. The median
segment rather slender, longer than the mesonotum, much
narrowed and rounded to the apex, not truncate, with a median
sulcus from the base tothe apex. First abdominal segment slender,
nearly twice as long as it is broad at the apex, about three times as
broad at the apex as at the base, with a deep sulcus from the base
to beyond the middle, the apical margin strongly depressed ;
second segment very large, nearly three times as broad at the
apex as the first, strongly convex; the apical margin of all the
segments depressed.
Black; the mandibles (except at the apex), the antenne at the
base, the apex of the clypeus, the tegule, the first segment of the
abdomen, the apical margins of the other segments, the tibia, tarsi,
and the apex of the femora dull ferruginous. The pubescence on
522 MR. R. E. TURNER ON [May 12;
the posterior margin of the pronotum, the postscutellum, the apex
of the median segment, the apical margin of all the abdominal
segments, and the whole of the third golden yellow, short and
dense. Wings very pale flavo-hyaline, slightly clouded on the
costa and at the apex; nervures ferruginous.
The first recurrent nervure is received by the first cubital cell
just beyond three-quarters from the base, the second by the second
cubital cell close to the base. The second cubital cell is extremely
short on the radial nervure, both transverse cubital nervures being
strongly curved and convergent. As in Parapison, the true
second cubital cell is obsolete.
Length 11 mm., exp. 18 mm.
Hab. Mackay, Queensland (Z'urner).
This species differs from <Azlacophilus vespoides Sm. in the
shape of the first abdominal segment, which is much shorter and
broader, and also in the shape of the second cubital cell and the
position of the recurrent nervures. I do not consider Aulaco-
philus of more than subgeneric value. The occurrence of species
in Australia is another instance of therelationship of the Hymeno-
pterous fauna of 8. America and Australia. The present species
shows most relationship to Pisom in the auratus group.
TRYPOXYLON CONNEXUM, sp. 0.
@. Clypeus broadly rounded anteriorly, broad and_ short,
densely clothed with short silvery pubescence. Antenne inserted
very near together, nearer to each other than to the eyes; the
second joint of the flagellum nearly half as long again as the
third, the apical joint conical, scarcely longer than the penultimate
joint. Eyes separated on the vertex by a distance about one-
third greater than that separating them at the base of the
clypeus; the posterior ocelli nearly twice as far from each other
as from the eyes, and further from the anterior ocellus than from
each other. The inner orbits of the eyes are clothed with silvery
pubescence as high as the deep and narrow emargination. The
anterior ocellus is surmounted by an elevated semicircular carina,
which is narrowly separated from the inner margin of the eyes and
reaches as low as the emargination, whence it is continued as a
V-shaped carina, much elevated at the apex and almost reaching
the base of the antennz ; from the apex it is continued as a very
high carina between the antenne, terminating abruptly at the
base of the clypeus; the space enclosed by the carinz is concave
and opaque; the vertex subopaque, with an obscure longitudinal
carina between the posterior ocelli. Pronotum on a level with
the mesonotum, short and broadly, but slightly, emarginate
anteriorly, the anterior margin raised and thickened. Mesonotum
opaque and finely, but not very closely punctured; the mesopleurz
with sparse silvery pubescence. Median segment slender, obliquely
truncate posteriorly, transversely striated, the posterior surface
with a deep median sulcus; the triangular space at the base
slightly convex. transversely striated, and divided by a longi-
1908. ] AUSTRALIAN FOSSORIAL WASPS. 523
tudinal sulcus. Abdomen shining, very minutely punctured and
feebly pubescent; the first segment very slender, slightly
thickened at the apex, longer than the second and third segments
combined.
Black; the posterior margin of the pronotum pale testaceous ;
tegule and the spines of the tibie testaceous. Wings hyaline,
nervures fusco-ferruginous.
¢. As in the female, but the apical joint of the antenne is
longer and slightly curved near the apex, fully twice as long as
the second joint of the flagellum.
Length, 2 10-12 mm., ¢ 9 mm.
fab. Mackay, Queensland (Zwrner); February and March.
Very near 7’. pileatum Sm. from India, but the frontal carinz
are much more strongly developed and the first abdominal segment
longer; the sculpture of the median segment is also coarser.
TRYPOXYLON PLACIDUM Sm.
Trypoxylon placidum Sm. Proce. Linn. Soc., Zool. vii. p. 35. n. 1,
63.
Hab. Mysole (Smith); Cairns, Queensland (Zwrner), April.
CrABrRo (RHOPALUM) MILITARIS, Sp. n.
3. Clypeus small, subtriangular, without a carina, without
spines on the anterior margin. Mandibles bidentate at the apex,
the inner tooth the longest. Head and thorax shining, minutely
punctured, the front smooth, the groove for the scape deep and
very narrowly separated from the eyes. Eyes almost touching
the base of the antennz, where they are separated from each other
by a distance equal to half the length of the scape. The second
joint of the flagellum half as long again as the first. Pronotum
not more than half as wide as the head, narrowed anteriorly, the
angles obtuse. Mesonotum broadly and very shallowly depressed
on the middle of the anterior half, with a very obscure longi-
tudinal carina in the depression; the scutellum divided by an
obscure longitudinal carina. Mesopleure shining, a little more
strongly punctured than the mesonotum. Median segment steeply
sloped posteriorly, smooth and shining; the longitudinal sulcus
dividing the triangular space at the base very shallow and obscure,
well-defined and deep on the posterior slope. Abdomen petiolate,
longer than the head and thorax combined, and slender; the first
segment very narrow and flat, swollen at the apex, longer than the
second segment, which is long and gradually broadened to the
apex, the fourth segment the broadest. Epipygium shining, very
narrowly truncate at the apex.
Black; the scape of the antenne, the anterior and intermediate
tibiee and tarsi, and the apex of the femora yellow; the apex of
the first abdominal segment, the second and third (except on the
apical margin), the posterior tibie and tarsi, and the apex of the
femora ferruginous. Wings hyaline iridescent, nervures black.
524 MR. R. E. TURNER ON | May 12,
The recurrent nervure is received by the cubital cell at two-
thirds from the base. The posterior tibize are much swollen
towards the apex.
Length 12 mm., exp. 17 mm.
Hab. Victoria (French); Tasmania.
The specimen from Tasmania has the wings fusco-hyaline.
CraBro (RHOPALUM) TRICOLOR Sm.
Orabro tricolor Sm. Cat. Hym. B. M. iv. p. 394. n. 14, 1855, 3.
2. Mandibles bidentate at the apex, the outer tooth very small.
Clypeus convex in the middle and subcarinate, produced in the
middle of the apical margin and emarginate, the angles of the
emargination forming strong blunt teeth, a small tooth on each
side; the whole clypeus clothed with shining silvery pubescence.
Antenne inserted close to the eyes, the distance between the eyes
at their base about equal to the length of the second joint of the
flagellum; the first joint of the flagellum about half as long as
the third, and a little more than half as long as the second. The
posterior ocelli nearer to each other than to the eyes and nearer
to the anterior ocellus than to each other, with a feeble longi-
tudinal carina between them. Head and thorax opaque, very
closely and minutely punctured, the front smooth and shining,
the groove for the scape occupying the entire breadth, the
posterior margin of the head broadly emarginate. Pronotum
short, strongly narrowed anteriorly ; the mesopleurz less opaque
than the mesonotum. Median segment steeply sloped posteriorly
and rounded, the space at the base opaque, very finely rugose,
with a sulcus from the base to the apex and a few very short striz
from the base. Abdomen petiolate, opaque, very minutely punc-
tured; the first segment long and slender, swollen at the apex,
a little longer than the second, the second and third gradually
widened, the fourth the broadest; the fifth dorsal segment clothed
with short, close, fulvous pubescence. The epipygium is tri-
angular, opaque, the sides slightly raised. The posterior tibiz
strongly swollen to the apex. The recurrent nervure is received
by the cubital cell just before three-quarters from the base.
Black; the scape of the antenne, the apex of the tegule, the
tarsi, the anterior and intermediate tibiz, and the apex of all the
femora and of the posterior coxe yellow; the base and apex of the
second abdominal segment and the base of the third ferruginous ;
the tarsal ungues black. Wings hyaline, faintly tinged with
fuscous ; nervures fuscous.
3. As in the female.
Length, 9 13 mm., ¢ 11 mm.
Hab. Woodford, Blue Mts, N. 8. Wales (G. A. Waterhouse) ;
Victoria (french) ; Tasmania (Smith).
CraBro (RHOPALUM) TENUIVENTRIS, sp. n.
9. Mandibles feebly bidentate at the apex. Clypeus clothed
1908. ] AUSTRALIAN FOSSORIAL WASPS. 525
with shining white pubescence, truncate on the apical margin,
convex at the base, with an oblique semicircular truncation at the
apex, the truncation smooth and shining, without pubescence.
Antenne inserted close together, touching the eyes; the second
joint of the flagellum longer than the first. Eyes just above the
base of the antenne separated by a distance about equal to the
length of the second joint of the flagellum; the posterior ocelli a
little further from the eyes than from each other and about the
same distance from the posterior margin of the head as from each
other. Head smooth and shining, the posterior margin broadly
emarginate. Thorax smooth and shining, the pronotum narrowed
anteriorly ; the median segment rounded posteriorly, shining and
very minutely punctured, with a median sulcus from the base.
Abdomen elongate, shining and minutely punctured; the first
segment very long and slender, of almost even thickness through-
out, the second and third gradually broadened, the fourth the
broadest. The recurrent nervure is received at the middle of the
cubital cell, the transverse cubital at about one-third from the
base of the radial cell.
Black; the mandibles and a mark on each side of the third
abdominal segment rufo-testaceous; the scape of the antenne, the
apex of the tegule, the tubercles at the posterior angles of the
prothorax, and the anterior and intermediate tibiz and tarsi pale
yellow; the four apical ventral segments testaceous. Wings
hyaline, nervures black.
Length 4 mm.
Hab. Mackay, Queensland (Turner); May.
CRABRO (RHOPALUM) TRANSIENS, Sp. 0.
3g. Clypeus broadly rounded at the apex, without a carina,
very feebly convex, and clothed with short silvery pubescence.
Hyes separated from each other at the base of the antenne by a
distance nearly half as great again as the length of the scape ; the
first joint of the flagellum longer than the second, which is equal
to the third. Posterior ocelli a little further from each other
than from the eyes and as far from the eyes as from the posterior
margin of the head. Headand thorax subopaque, microscopically
punctured ; the pronotum less than half as broad as the head and
narrowed anteriorly, mesopleure minutely punctured, the groove
rather shallow. Median segment smooth, rounded and steep
posteriorly, with a transverse row of coarse punctures at the base
and a short longitudinal sulcus at the extreme apex. Abdomen
shining, microscopically punctured, petiolate; the first segment.
very narrow, swollen at the apex, a little longer than the second
segment, the third and fourth segments the broadest. The re-
current nervure is received by the cubital cell at about three-fifths
from the base.
Black ; the scape of the antenne and the posterior angles of the
pronotum yellow; the first joint of the flagellum, the following
four joints beneath, the mandibles, tegule, first joint of the
526 MR. R. E. TURNER ON [May 12,
abdomen, the base of the second joint, the tibie, tarsi, trochan-
ters, and femora (except at the base) ferruginous. Wings hyaline,
brilliantly iridescent ; nervures fusco-ferruginous.
Length 5 mm.
Hab. Victoria (french).
CrABRO (RHOPALUM) FRENCHII, Sp. n.
2. Mandibles bidentate at the apex, the teeth short and feeble.
Clypeus short, slightly convex, without a carina, and clothed with
short silvery pubescence. Eyes separated from each other at the
base of the antennz by a distance equal to about three-quarters
of the length of the scape; the first joint of the flagellum longer
than the second. Posterior ocelli as far from the eyes as from
each other and a little nearer to the posterior margin of the head.
Head and thorax shining, microscopically punctured ; the pro-
notum short and transverse, about two-thirds of the width of the
head, theangles not prominent. Median segment shining, with
a median sulcus from the base to the apex. Abdomen petiolate,
as long as the head, thorax, and median segment combined, shining;
the three basal segments smooth, the fourth and fifth very closely
and minutely punctured; the first segment narrow, very slightly
swollen at the apex, more than twice as long as broad, about one-
third shorter than the second segment; the second gradually
widened to the apex ; the third and fourth segments the broadest.
Pygidium elongate, triangular. The posterior tibie are swollen
towards the apex. ‘The recurrent nervure is received close to the
middle of the cubital cell.
Black; the mandibles (except at the apex), the scape of the
antenne, the posterior angles of the pronotum, the trochanters,
and the anterior and intermediate tibie and tarsi pale yellow ;
the tegule, pygidium, and the posterior tibiz and tarsi fusco-
ferruginous. Wings hyaline, iridescent ; nervures fusco-ferru-
gimous.
Length 5 mm.
Hab. Victoria (French) ; November.
CrABRO (RHOPALUM) CONAYOR, Sp. n.
g. Mandibles bidentate at the apex, the teeth of about equal
length. Clypeus clothed with silvery pubescence, broadly rounded
anteriorly, with a delicate median carina. Hyes separated from
each other at the base of the antennez by about half the length of
the scape, diverging very broadly towards the vertex. Second
joint of the flagellum half as long again as the first and a little
longer than the third. Posterior ocelli a little further from each
other than from the eyes and more than: half as far again from
the posterior margin of the head as from each other. Head large,
subquadrate, broadly emarginate posteriorly, finely rugulose
behind the ocelli, rugose in front of the ocelli, with very thin
erect pubescence. Pronotum short, broadly and shallowly emar-
ginate anteriorly, the anterior angles slightly produced, narrower
than the head. Mesonotum shallowly punctured; mesopleure
1908. | AUSTRALIAN FOSSORIAL WASPS. 527
opaque, almost smooth, the groove narrow and punctured. The
enclosed space at the base of the median segment coarsely longi-
tudinally rugose, the sides and apex of the segment obliquely
striate-rugose. Abdomen opaque, petiolate; the first segment
very slender, twice as long as the second, a little swollen at the
apex, the second segment nearly as broad as the third.
Black; the scape of the antennez, a spot on each side of the
apex of the first abdominal segment, a broad band across each of
the remaining segments and the whole of the seventh, the tibiz,
tarsi, and the apex of the femora ochraceous yellow. Wings
hyaline, iridescent ; nervures fusco-ferruginous.
The recurrent nervure is received by the cubital cell beyond
two-thirds trom the base.
Length 7 mm.
Hab. Cooktown, Queensland (Zurner) ; November.
CraBro (RHOPALUM) IDONEUS, Sp. n.
9. Head and thorax opaque, very delicately punctured-rugu-
lose; mandibles bidentate at the apex, the teeth short, the inner
tooth the longest ; maxillary palpi five-jointed, labial palpi three-
jointed. Clypeus transverse, slightly produced in the middle of
the anterior margin. Front concave, the antenne inserted nearer
to the eyes than to each other. Pronotum transverse, rounded
at the angles; mesonotum with a depression from the middle of
the anterior margin to the centre ; the scutellum half as long as
broad. Median segment very short, rounded, and truncate
posteriorly ; the space at the base longitudinally striated at the
base, almost smooth and opaque at the apex, with a median carina
extending on to the surface of the truncation to the apex. Abdo-
men petiolate, very minutely punctured; the petiole about half
as broad as long, of even length throughout, nearly as long as
the second segment, which is slightly narrower than the third.
Epipygium deeply punctured, lanceolate.
Black; the clypeus, cheeks, the truncation of the median seg-
ment, and the fourth and fifth abdominal segments clothed with
pale golden pubescence; the mandibles fusco-ferruginous; the
flagellum (except the basal joint), the tegul, the first abdominal
segment, the pygidium, the apical margins of the other abdominal
segments, the posterior tibiz and tarsi, and the posterior femora
above ferruginous ; the scape of the antenne, the basal joint of
the flagellum, the pronotum very narrowly interrupted in the
middle and on the sides, the scutellum (except the apical margin),
a transverse line on the postscutellum, the anterior and inter-
mediate legs, and the posterior tibize beneath yellow. Wings
hyaline, iridescent, clouded in the radial cell; nervures ferruginous.
The recurrent nervure is received by the cubital cell at three-
fifths from the base. The radial cell is very broadly truneate at
the apex, the radial nervure indistinctly produced beyond the cell.
The eyes are separated at the Base of the antenne by a distance
equal to about three-quarters of the length of the scape; the
Proc. Zoou. Soc.—1908, No. XX XTV. 34
528 MR. R. E. TURNER ON [May 12,
first two joints of the flagellum are about equal in length. The
posterior ocelli are a little further from the eyes than from each
other and alittle further from the posterior margin of the head
than from the eyes.
3. Asin the female, but more slender; the head and thorax
shining, almost smooth; the second abdominal segment narrow,
about. one-third longer than the first, very little more than halt
as wide at the apex as the third segment; the fourth segment
the widest ; the posterior tibie more dilated than in the female ;
the scutellum wholly black, the second abdominal segment
ferruginous with a large black spot in the middle. The eyes are
only separated at the base of the antenne by about half of the
length of the scape and diverge towards the vertex less strongly
than in the female.
Length, ¢ 6 mm., @ 7 mm.
Hab. Mackay, Queensland (Turner) ; May.
Crapro (RHOPALUM) AGILIS Sin.
Crabro (Rhopalum) agilis Sm. Proc. Linn. Soc., Zool. 11. p. 18,
1858, 9. .
Hab. Celebes (Wallace); Mackay, Queensland (Turner).
CRABRO PROSOPOIDES, Sp. 0.
2. Clypeus slightly produced, with a median carina from the
base not reaching the apex, clothed with silvery pubescence. Eyes
separated at the base of the antenne by a distance scarcely
exceeding one-quarter of the length of the scape; the first jomt
of the flagellum fully as long as the second. Mandibles bidentate
at the apex, the teeth short; maxillary palpi six-jointed, labial
palpi four-jointed. The posterior ocelli about one-third further
from each other than from the eyes and about as far from the
posterior margin of the head as from each other. Head very
closely and finely punctured, the front smooth and concave.
Pronotum transverse and linear, the angles not prominent ; meso-
notum finely and closely punctured, the groove on the meso-
pleure well marked. Median segment steeply, but not vertically,
sloped posteriorly, the triangular space at the base smooth and
shining, with a shallow median sulcus, and a transverse row of
deep punctures at the extreme base; the pesterior slope with a
large, deep depression at the base, very delicately and closely
transversely striated. Abdomen very finely and closely punctured
the first segment one-third longer than the second, very narrow
at the base, the apex half as wide as that of the second segment
the second segment narrower than the third, the second and third
segments moderately constricted at the base, the apical segment
triangular. The recurrent nervure is received at the middle of
the cubital cell.
Black; the scape of the antenne, pronotum, tegule, scutellum,
postscutellum, tibize, tarsi, and the apex of the femora yellow ;
the flagellum light ferruginous. Wings hyaline, nervures fuscous,
1908. | AUSTRALIAN FOSSORIAL WASPS, 529
' 6. Asin the 2. The apical segment of the abdomen is
rounded.
Length, 2 8mm., 3 6 mm.
Hab. Mackay, Queensland (Zurier); March to May. Towns-
ville, Queensland (Dodd).
CRABRO PERLUCIDUS, sp.n. (Plate XX VI. fig. 15.)
@. Mandibles bidentate at the apex, the inner tooth a little
the longest. Clypeus strongly convex at the base, clothed with
silvery pubescence, with an obliquely depressed smooth truncation
to the middle of the apical margin. Antenne inserted nearly
twice as far from each other as from the eyes, the first joint of
the flagellum longer than the second; the eyes separated from
each other at the base of the antenne by a distance exceeding
half the length of the scape. Posterior ocelli a little nearer to
each other than to the eyes, half as far again from the posterior
margin of the head as from each other. Head and thorax smooth
and shining; the pronotum slightly depressed, rounded at the
anterior angles; scutellum very finely punctured. Median seg-
ment very short, steeply sloped posteriorly, the enclosed space at
the base irregularly obliquely striated, with a depressed, transverse,
coarsely punctured sulcus at the base; a median sulcus from the
base to the apex. Abdomen shining, very minutely punctured,
nearly as long as the head and thorax united; the first segment
as long as the second, twice as broad at the apex as at the base
and half as broad as the apex of the second segment. Pygidium
elongate triangular.
Black; the scape of the antenne and the anterior and inter-
mediate legs yellow, the intermediate legs stained with ferruginous;
the flagellum, posterior legs, and abdomen bright ferruginous ;
the tegule testaceous. Wings hyaline, nervures black.
The cubital cell receives the recurrent nervure at two-thirds
from the base; the radial cell receives the transverse cubital
nervure before the middle.
Length 7 mm., exp. 10 mm.
Hab. Mackay, Queensland (Turner); May.
CRABRO DODDII, sp. n.
@. Clypeus a little produced and truncate at the apex, with a
median carina, clothed with silvery pubescence. Antenne inserted
twice as far from each other as from the eyes ; the second joint of
the flagellum small, shorter than the first. Eyes separated from
each other at the base of the antenne by a distance equal to two-
thirds of the length of the scape, strongly divergent towards the
vertex. Posterior ocellia iittle further from each other than from
the eyes, but nearer to each other than to the posterior margin of
the head. Head finely punctured-rugose; the front opaque, with
sparse silvery pubescence. ‘Thorax rugose, the pronotum trans+
verse, almost as broad as the head, the anterior angles prominent.
Mesopleurz without a groove for the femora. The enclosed space
34*
530 MR. R, E, TURNER ON [May 12,
at the base of the median segment coarsely rugose with a deep
median sulcus. Abdomen short, not petiolate ; the first segment
subtruncate at the base, depressed on the apical margin, the
second segment the longest, All the segments opaque, very
closely and finely punctured, The recurrent nervure 1s received
near the apex of the cubital cell.
Black; the mandibles (except at the apex), the scape of the
antenne, the anterior and posterior angles of the pronotum, the
postscutellum, the tibize, and the apex of the femora yellow ; the
flagellum, the tegule, the base of the femora, the tarsi, and the
abdomen light ferruginous, the first abdominal segment with an
obscure yellow band at theapex. Wings hyaline, nervures fusco-
ferruginous.
¢. As in the female, but the pronotum is entirely yellow and
the postscutellum black.
Length, 9 8mm., 3 6 mm.
Hab. Townsville, Queensland (Dodd); February.
CRABRO HEBETESCENS, Sp. 0.
@. Mandibles tridentate at the apex, the inner tooth the
shortest, a strong acute tooth just before the middle of the inner
margin. Maxillary palpi six-jointed, labial palpi four-jointed.
Clypeus clothed with shining white pubescence, with a carina.
from the base to the apex, truncate at the apex, with two short
teeth on each side. Head large, finely and closely punctured,
the cheeks clothed with silvery pubescence. Antenne inserted
very close to the eyes, the second joint of the flagellum about half
as long again as the first or third. Eyes at the base of the
antenne separated by a distance about equal to the length of the
two basal joints of the flagellum; front very narrow, the groove
for the scape reaching the eyes; the facets of the eyes larger near
the base of the antennz than elsewhere. Posterior ocelli very far
apart, as far from the eyes as from each other, but nearer to each
other than to the posterior margin of the head, which is straight.
Pronotum more than two-thirds of the breadth of the head, very
short and transverse, the anterior margin raised. Thorax punc-
tured, the mesopleure strongly grooved for the anterior femora.
Median segment short, subtruncate posteriorly, the enclosed space:
at the base very broadly rounded and rugose; the face of the
truncation finely transversely striated, with a delicate longitudinal
carina. Abdomen ovate, shining and finely punctured, the apical
segment lanceolate.
Black ; the mandibles at the base, the scape of the antenne,
the anterior margin of the pronotum, interrupted narrowly in the
middle, the tubercles at the posterior angles of the pronotum, a
small spot on each side at the base of the scutellum, a spot on
each side of abdominal segments 1-5, the tibie above, and the
basal joint of the tarsi creamy white. Wings hyaline, nervures.
black, the tegule fuscous. The recurrent nervure is received by
the cubital cell at about three-quarters from the base, the cubital
5
1908. | AUSTRALIAN FOSSORIAL WASPS. Fyoil
nervure is sharply bent at the point of junction, the apical portion
appearing almost more like a part of the transverse cubital than
of the cubital nervure.
Length 9 mm.
Hab. Mackay, Queensland (Zurner) ; January.
CRABRO CINCTUS, sp.n. (Plate XX VI. fig. 14.)
@. Head very large, almost square, slightly rounded at the
posterior angles, extremely finely and closely punctured. Man-
dibles broad at the apex and tridentate, the inner tooth very short,
the middle tooth the longest. Clypeus slightly porrected, almost
vertically truncate at the extreme apex, the surface of the trun-
cation semicircular and slightly concave. Cheeks depressed along
the outer orbits of the eyes, the margins of the face and the
depressions on the cheeks clothed with short silvery pubescence.
A longitudinal suleus below the anterior ocellus and a longitudinal
carina above it, a shallow depression on each side of the carina
behind the posterior ocelli, the front clothed with short golden
pubescence, the inner orbits of the eyes slightly depressed.
Thorax narrower than the head; the anterior and posterior
margins of the pronotum raised and with the angles slightly
prominent, a deep transverse sulcus interrupted in the middle
separating the raised margins. Mesonotum and scutellum very
finely and closely punctured, the disc of the mesonotum with a
broad and shallow depression. Median segment short, truncate
posteriorly, with a deep, longitudinal, median sulcus from the base
to the apex; the enclosed space at the base clesely punctured
with a few very short striz at the base, broadly rounded at the
apex. Abdomen subpetiolate, coriaceous, pygidium narrow and
lanceolate.
Black ; the mandibles at the base testaceous yellow; the clypeus,
the scape of the antenne, the two basal joints of the flagellum,
and a spot on each side at the anterior angles of the mesonotum
yellow; the pronotum, a large mark on the scutellum, the post-
scutellum, a spot on the mesopleure, a short longitudinal line on
the middle of the basal abdominal segment, a broad transverse
band at the base of the second segment, and the three apical
segments above orange; the tegule and the apical ventral seg-
ment of the abdomen ferruginous brown. Legs yellow, stamed
with ferruginous brown. Wings hyaline, tinged with fuscous ;
nervures dark fuscous.
Length 12 mm.
Hab. Mackay, Queensland (Zwrner) ; April.
The posterior ocelli are two and a half times as far from the
posterior margin of the head as from each other; further from
each other than from the eyes. The antennz are inserted close
to the eyes, the second joint of the flagellum nearly twice as long
as the first and as long as the third; the distance between the
eyes at the base of the antenne equal to half the length of the
scape. The mesopleure are rugose, the depression for the femora
532 MR. R. E. TURNER ON [May 12,
not developed. The recurrent nervure is received by the cubital
cell beyond three-quarters from the base ; the radial nervure is
not continued beyond the end of the radial cell.
CRABRO MACKAYENSIS, Sp. n.
2. Mandibles tridentate at the apex, the inner tooth much the
shortest, the central tooth longer than the outer one. Clypeus
slightly advanced, truncate at the apex, with a longitudinal carina
from the base, clothed with short silvery pubescence. The pubes-
cence on the cheeks pale golden. Head finely and very closely
punctured ; eyes very large, the front between them narrow and
concave; a very shallowly depressed, longitudinal, and almost.
smooth mark on the inner margin of the eye near the summit ;
a short longitudinal suleus below the anterior ocellus and another
on the vertex almost reaching the anterior ocellus ; the posterior
ocelli as far from each other as from the eyes. Pronotum very
short, transverse, raised and thickened, not prominent at the
angles; mesonotum finely rugose. Median segment short, the
space at the base enclosed by carinz, longitudinally striated, with
an obscure median carina, broadly rounded at the apex; the
posterior truncation very obscurely transversely striated, with a
median suleus. Abdomen subovate, very minutely and closely
punctured; the first segment narrow at the base, the apex about
two-thirds of the breadth of the second segment, only a little
longer than the second segment; the apex of the fifth segment
clothed with pale golden pubescence ; the apical segment lanceo-
late, smooth and recurved at the sides, with a tuft of long golden
pubescence on each side.
Black; the mandibles (except at the apex), the scape of the
antenne, the pronotum (very narrowly interrupted in the middle
and more broadly on the sides), a spot on each side near the base
of the scutellum, a transverse line on the postscutellum, a spot
near the middle of the enclosed space on the median segment, a
broad transverse band near the apex of the first abdominal seg-
ment, a small spot on the sides of the second and fourth segments,
and a transverse band narrowly interrupted in the middle on the
fifth segment, yellow; the four basal joints of the flagellum, the
tegule, the apical margins of the abdominal segments, the tibie,
tarsi, and extreme apex of the femora and coxe ferruginous brown.
Wings pale flavo-hyaline, nervures pale ferruginous.
Length 10 mm., exp. 17 mm.
Hab. Mackay, Queensland (Z’urner); April.
The second joint of the flagellum is about one-third longer than
either the first or third, and nearly equal in length to two-thirds
of the distance between the eyes at the base of the antennz.
CRABRO ORDINARIUS, sp. n.
Q. Mandibles bidentate at the apex, the teeth long and of
about equal length. Clypeus densely clothed with silvery pubes-
cence, with a carina from the base to beyond the centre, with a
1908. | AUSTRALIAN FOSSORIAL WASPS. 533
smooth and shining, oblique, triangular truncation at the apex.
Head and thorax very closely and finely punctured, the front and
cheeks clothed with pale golden pubescence; the posterior ocelli
far apart, further from each other than from the eyes. The
anterior margin of the pronotum raised, with a transverse groove
behind it on the sides. A faint and broad depression from the
anterior margin of the mesonotum to the middle; the scutellum
faintly depressed in the middle. Median segment short, truncate
posteriorly, the enclosed space at the base broadly rounded at the
apex, obliquely striate rugose, with a very broad median sulcus in
which lie several transverse strie; the face of the truncation
pubescent, very finely rugose, with a strong median sulcus. Abdo-
men very closely and minutely punctured, not petiolate, the apical
segment lanceolate.
Black; a spot at the base of the mandibles, the scape of the
antenne, the first jomt of the flagellum, the anterior margin of
the pronotum narrowly interrupted, the posterior angles of the
pronotum, a spot on each side at the basal angles of the scutellum,
a transverse line on the postscutellum,a trilobed mark at the
base of the first abdominal segment, a transverse band at the base of
the third and fourth segments, a line on the anterior tibie, and a
spot near the apex of the posterior tibiz, yellow; the tegule and
the legs (except the coxze) ferruginous. Wings hyaline, nervures
ferruginous.
¢. Asin the female, but without the yellow mark on the first
abdominal segment, and there is a dull yellow transverse band on
each side of the fifth and sixth segments.
Length, 9 9-11 mm., ¢ 9 mm.; exp., ¢ 12 mm., 9 1/ mm.
Hab. Mackay, Queensland (Zurner); February—May.
The second joint of the flagellum is longer than the third in
both sexes and more than twice as long as the first. The eyesare
separated at the base of the antenne by a distance rather ex-
ceeding one-third of the length of the scape in the female and by
a little less in the male. The groove on the mesopleure is not
well-developed.
CRABRO CONGLOBATUS, Sp. Nl.
@. Mandibles broad and bidentate at the apex, the teeth of
about equai length. Clypeus small, advanced in the middle,
strongly emarginate at the sides, slightly convex, without a carina,
and clothed with pale shining pubescence. Eyes separated at the
base of the antenne by a space equal to one-third of the length of
the scape; the second joint of the flagellum about three times as
long as the first and half as long again as the third. Posterior
ocelli a little nearer to each other than to the eyes, and more than
half as far again from the posterior margin of the head as from
each other. Head and thorax very closely and finely punctured ;
the cheeks clothed with very pale golden pubescence. Pronotum
narrower than the head, transverse, the angles not prominent ;
mesopleure smooth and shining, with a well-marked groove.
534 ON AUSTRALIAN FOSSORIAL WASPS. [May 12,
Mesonotum half as broad again as long, with a broad shallow
depression from the anterior margin to the centre, in the de-
pression is a very delicate longitudinal carma. Median segment
very short, the posterior slope vertical, the space at the base
coarsely obliquely striated, with a median sulcus; the posterior
surface transversely rugose, with a deep median suleus. Abdomen
subovate, shining, very minutely punctured ; the first segment
about one-third longer than the second, very narrow at the base,
rather more than half as wide at the apex as the apex of the
second segment, the apical segment lanceolate. The radial
nervure is received not far from the apex of the cubital cell.
Black; the scape of the antenne, pronotum very narrowly
interrupted in the middle, the scutellum (except at the base), a
transverse line on the postscutellum and a transverse band on
each side of abdominal segments 2-4, least broadly separated in
the middle on the second segment, yellow; the tegule, legs,
mandibles, and the apical margins of ventral segments 2-5 ferru-
ginous. Wings fusco-hyaline, nervures black. A small spot on
the mesopleurze below the anterior wings yellow.
3g. Asin the 9, but the distance between the eyes at the
base of the antenne is equal to half the length of the scape, the
median segment is more coarsely striated and the apical abdo-
minal segment is rounded ; the yellow spot on the mesopleure is
absent.
Length, ¢ 9mm., exp. 14mm.; 9 10mm., exp. 17 mm.
Hab. Mackay, Queensland (Zwirner); April and May.
Nearly allied to C. palitans Bingh. from India.
CRABRO BIVITTATUS, Sp. Nn.
@. Mandibles bidentate at the apex. Clypeus small, clothed
with silvery pubescence, with a median carina, slightly porrect at
the apex, with a minute tooth on the apical margin on each side
of the carina. Head subquadrate, slightly emarginate posteriorly,
very finely and closely punctured, the front smooth in the middle,
clothed with silvery pubescence on the sides. The posterior ocelli
further from each other than from the eyes and at least
one-half further from the posterior margin of the head than
from each other. Pronotum narrowed anteriorly, the anterior
margin straight and a little raised with a slight groove
behind it; the mesonotum slightly depressed in the middle
anteriorly, minutely punctured. Median segment steeply sloped
posteriorly ; the space at the base divided by a very broad median
sulcus in which are a few transverse strie, with short longitudinal
strie at the base, the apex finely punctured ; the posterior slope
of the segment transversely and very finely striated, with a
median sulcus. Abdomen very closely and minutely punctured.
The recurrent nervure is received a little before the apex of the
cubital cell.
Black ; the scape of the antenne yellow ; the anterior margin
of the pronotum narrowly interrupted, a broad transverse band at
535
1908. ] ON MAMMALS FROM ZAMBESIA.
the base of the second and fourth abdominal segments and a
narrow band on each side of the fifth segment, orange; the tibiz,
tarsi, and the apex of the femora dark ferruginous, the tegul
rufo-testaceous. Wings hyaline, tinted with fuscous; nervures
black.
Length 9 mm.
Hab. Victoria (French).
Type in British Museum.
The eyes are separated at the base of the antenne by a distance
equal to about one-third of the length of the scape, the second
joint of the flagellum is nearly three times as long as the first and
fully half as long again as the third. The first abdominal
segment is narrowed to the base.
EXPLANATION OF PLATE XXVI.
Fig. Fig.
1. Harpactophilus arator, sp. nu.. | 9. Nysson (Acanthostethus) punctatis-
p-461. | simus, Sp. N., p. 505.
2, Harpactophilus tricolor, sp. n., | 10. Gorytes sanguinolentus, sp.n.,p. 497.
p. 462. | 11. Gorytes lucidulus, sp. n., p. 498.
. Ammophila aurifera, sp. 0., p. 464.
. Psenulus interstitialis Cam., p. 463.
Cerceris inexpectata, sp. n., p. 469.
Notogonia regina, sp. n., p. 475.
. Tachytes formosissimus, sp. n.,p.482. |
. Pisonmelanocephalum,sp.n., p. 515.
3. Pison
(Aulacophilus)
sp. n., p. 521.
icarioides,
. Crabro cinctus, sp. ., p. 531.
. Crabro perlucidus, sp. n., p. 529.
. Zoyphiun rufonigrum, sp. n.,p. 494. |
May 26, 1908.
Prof. E. A. Mincutn, M.A., Vice-President,
in the Chair.
The following papers were read :—
1. The Rudd Exploration of 8. Africa. —X. List of Mammals
collected by Mr. Grant near Tette, Zambesia. By
Orprrmnpy Proms, HRS. “RUZIS.” and’ Ra".
WrovgutTon, F.Z.S8.
[Received April 14, 1908.]
We now come to the final collection of the Rudd Exploration,
for after making it Mr. Grant had a severe attack of fever, and
by arrangement with Mr. Rudd he has now come home, so that
this magnificent exploration, which has been going on for the last
five years, thus comes to an end.
Further details of the papers written on the different collections
are appended to the present account, but we may here say that
536 MESSRS. THOMAS AND WROUGHTON ON [May 26,
the results of Mr. Rudd’s splendid generosity have far surpassed,
in their great and permanent value, our most sanguine expecta-
tions—a fact for which the fullest credit must also be given to the
collector, Mr. C. H. B. Grant, who has risen in the ablest manner
to the great opportunity afforded him by Mr, Rudd.
The total results: form the largest collection of Mammals ever
received by the National Museum from any one source, the
nearest approaches to it being the products of the Simons and
Robert expeditions to 8. America, and the Duke of Bedford’s
Exploration of Eastern Asia, the last-named being still in
progress.
In all 1541 mammals, exclusive of duplicates, have been
registered as presented to the National Museum by Mr. Rudd,
while duplicates have been presented to the Royal Scottish
Museum, Edinburgh, and the South African Museum, Cape
Town.
A considerable and quite unexpected number of new species
and subspecies have been discovered, and, what is quite as
important, most of the old species, insufficiently or imexactly
described on specimens now deteriorated, have heen definitely
identified by topotypes, and are represented by good modern
material, which may be made the basis of further progress.
In this connection the Tette series, of which we give an account
in the present paper, is of especial importance ; for every worker
on South African zoology has been hampered by the difficulty -
of making out with exactitude the species obtained during
Dr. Peters’s famous expedition to Zambesia, and described by him
in his ‘Reise nach Mossambique, of which the ‘ Siiugethiere ’
was published in 1852.
In order, therefore, to get a series of the species described by
Peters, Mr. Grant went to Tette, Peters’s chief collecting-place,
and formed the series enumerated below.
While we were working out this series, the definite determination
of Peters’s species has enabled us to sort out a number of the groups,
with the result that many forms hitherto assigned, in our papers
and elsewhere, to Peters’s species, now prove to need description.
Mr. Grant’s notes on the Tette district are as follows :—
“Tt was the driest time of the year when I reached Tetite, and,
except in the main rivers, there was practically no water any-
where, and as, on the Zambesi near Tette, there were too many
natives present for it to be possible to collect, I moved south-
wards and pitched my camp at the junction of the Luenya and
Mazoe Rivers, which is some 20 miles due south of Tette.
“The country there is exactly similar to that along the
Zambesi, being hilly, and in places somewhat mountainous ;
the soil is sandy and very stony, especially on the hill-sides, but
snee are no krantzes that would harbour dassies or red hares.
All the vegetation, except along the rivers, was dried and dead
and the trees leafless, the course of the rivers being plainly shown
from a distance by the verdure of the trees on their banks.
1908. | MAMMALS FROM ZAMBESIA. 537
“ Everywhere the veldt is well bushed, amounting to thickets
in most parts, with a fair amount of larger timber, mainly mopani
and ‘ cream-of-tartar,’ the latter being very plentiful and growing
to an enormous size.
‘“‘ Except in favourable situations, grass does not seem to grow
freely, and the cereal crops of the natives are not nearly so good
as In many districts to the southward, although tomatoes, onions,
&e. are grown freely on the banks and in the beds of the rivers.
“The natives are mixed local tribes variously known as
Nyungwis, Tongas, Barués, &. When not too lazy they give
much of their time to the capture of small buck, cats, squirrels,
rats, &c., the majority of which they utilize for food.
“The climate cannot be considered healthy, even in the dry
season, and the temperature is generally high during the day and
makes good collecting difficult.
“In the five weeks I spent in the Mazoe camp the average
temperature was 95°, 104° being the highest recorded; no rain
fell.”
1. CERCOPITHECUS PYGERYTHRUS RUFOVIRIDIS Is. Geoff.
2. 2003.
This specimen, a young female, is undoubtedly the same form
as those from Gorongoza mentioned in our last paper, and both
are almost certainly Cercopithecus flavidus of Peters. We follow
Mr. Pocock * in holding that that species isa synonym of C. rufo-
viridis, and accordingly adopt this name for the specimens from
Gorongoza and Tette. Mr. Pocock has pointed out (/.¢.) that the
Specimens obtained by Mr. Grant in the Knysna, Zululand, and
E. Transvaal are all typical C. pygerythrus Cuv., and to these we
may add the specimens received from Inhambane and Beira
since the date of Mr. Pocock’s paper. All former identifications
in the present series of papers must be modified accordingly.
““ Native name, ‘ Pusi.’
“Only two troops of this monkey were seen and they were
exceedingly wild.
“Generally frequenting the trees along the river-banks and
observed drinking in the middle of the afternoon.”—C. H. B.G.
2. GALAGO MossAMBICUS Pet.
3S. 2042. ©. 2028, 2029, 2030, 2031, 2043.
Topotypes of species.
These specimens, which represent “ Ofolicnus mossambicus,”
confirm what we have stated in describing G. granti in a former
paper 7, namely, that this latter species is readily recognizable
on account of its long muzzle.
From G. moholi Sm., to which it is no doubt closely allied,
G. mossambicus is > es by its smaller size (greatest length
* P. Z.S. 1907, p. 737.
+P Z. 8. 1907, p. 286.
538 MESSRS. THOMAS AND WROUGHTON ON [May 26,
of skull 38 mm., of upper tooth-row from front of canine to back
of last molar 13°7 mm., against 41 and 15 mm. mm moholt) and
proportionally much longer tail.
3. EPoMOPHORUS CRYPTURUS Pet.
Or So
A young specimen. Topotype of species.
‘Native name, ‘ Demanyundo.’
“Said to be common, but only the one specimen was seen, and
that was put up and shot in the daytime in a thicket on the bank
of the Mazoe.”—C. H. B.G.
4, RHINOLOPHUS LOBATUS Pet.
go. 2018. ©. 2013, 2014, 2019, 2021, 2025.
These are practically topotypes of Peters’s species, the technical
type-locality being Sena.
“Native name, ‘ Nyagelingwelingwe’; it is the same for all
insectivorous bats.”—C. H. B. G.
5. RHINOLOPHUS HILDEBRANDTI Pet.
®. 2009.
6. H1rprosIDERUS CAFFER Sund.
J. 2024. @. 2015, 2016, 2020, 2022, 2023.
“Both species of Rhinolophus (vide supra), these, and Petalia
(vide infra) were all taken out of one tree.”—C. H. B. G.
7. PETALIA* CAPENSIS Sm.
Ge 20M 2OI2 QOL. nO ZOO:
These specimens represent the Vycteris fuliginosa of Peters, the
type-locality of which was Boror.
8. VESPERTILIO CAPENSIS Sin.
fg. 2004.
An exceptionally large individual, but not, we think, separable
from V. capensis.
9. ScOTOPHILUS NIGRITA DINGANI Sm.
fo. 1994.
The various forms included by Dobson under Scotophilus
borbonicus = nigrita, in his ‘Catalogue of the Chiroptera,’ may
apparently be separated into two groups by their size—a larger,
represented by S. nigrita Schreb., and a smaller, the oldest name
for which is S. viridis Peters.
The present specimen is a topotype of Peters’s WVycticejus
planirostris, but we ave unable to separate it from S. dingand.
*« = Nycteris auctorum.
1908. ] MAMMALS FROM ZAMBESIA. 539
The forms of 8. African Scotophilus (including those noticed
below) may be arranged in a key as follows :—
A. Size larger (forearm 52-57 mm.; skull length
20-21 ; “maxillary tooth-row 7-7" 5) Mata: .. nigrita-group.
a. A strong suffusion of yellow in the colouring
making upper surface olive-green, belly bright
SUD chishacbos 55 son aee niece eee Hee odes eee eee S. nigrita dingant Sm.
b. No tinge of yellow in the color ations. S. nigrita herero Thos.
B. Size smaller (forearm 45-51 mm. > skull leneth
17-18 ; maxillary tooth-row 6°3-6° ) Slascae ns viridis-zroup.
a. A strong suffusion of yellow in the colouring,
making upper surface olive-green, belly bright
yellow ......... soreconeosioas Se GURAGIIS Tai,
b. No tinge of yellow i in the coloration ............ S.viridis damarensis Thos.
“Two species of this genus were secured, neither being
common.
“They appear early in the evening and their flight is strong ;
they hawk the country in wide and regular circuits.” —-C. H. B.G.
10. ScoroPHILUS VIRIDIS DAMARENSIsS Thos.
g. 1955, 1956, 2027, 2032.
Reference was made in the paper on the Inhambane Collection *
to the presence in the series of Scotophilus of certain specimens
smaller in size than the rest; as these smaller specimens possess
all the essential characters attributed by Peters to his Nycticejus
viridis, we accept them as representing that species, the type
locality of which is the island of Mozambique.
The present specimens are indistinguishable from S. damarensis
Thos., and as they also closely resemble the Inhambane specimens
except in coloration we rank them as a western race of S. viridis.
11. SCOTEINUS SCHLIEFFENI AUSTRALIS, subsp. n.
6. 1967, 1975, 1993, 2000, 2005, 2007. 2. 1995, 2001.
On laying out all the specimens of this species in connection
with the identification of the present series, it became evident
that there are several well-marked geographical races separable
on colour characters.
Typical S. schlieffent was based by Peters on a specimen from
Cairo. He described it as “supra rufescens, subtus ex albo
rufescens.” A second species, S. minimus, based on a 2 from
Tanganyika was described by Noack as “ oben olivengelbbraun
unten weissgelb. Seiten hell umbra.... ” Unfortunately we
have no undoubted specimen of either of these for comparison,
but we consider that we are justified in describing three forms
as certainly distinct from either typical S. schlieffeni or WS.
nunimus (which latter is at most a local race of the former).
These are: (1) a pale desert form from the Aden Hinterland ;
(2) a white-bellied desert form from Upper Egypt; and (3) the
present series from 8. Africa.
* P. Z, S. 1907, p. 285.
540 MESSRS. THOMAS AND WROUGHTON ON [May 26,
The following are descriptions of these three forms :—
ScCoTEINUS SCHLIEFFENI BEDOUIN, subsp. n.
Rather smaller in size than typical S. schlieffent.
Colour above nearest to “ wood-brown,” but a much paler shade
than that given by Ridgway ; below still paler, 7. ¢., the colour
containing more white.
Dimensions of type :—
Head and body 41:8 mm. ; tail 28°8 ; forearm 30; ear 9°).
Skull—ereatest length 12-2 ; interorbital breadth 3:3; brain-
case breadth 6:3; breadth across upper jaw at level of m®° 5:6;
post-canine tooth-row 3°5.
Hab. Lahej, near Aden.
Type. Adult. B.M. no. 95.6.1.53. Collected on_ the 12th
March, 1895, and presented to the Museum by Col. J. W.
Yerbury.
A second specimen taken at the same time only differs in being
slightly smaller. The difference in coloration between these
specimens and a series of four taken by Mr. W.’ Dodson, 18th Sept.,
1899, at Sheik Othman, only 10 or 12 miles distant—nearer the
coast—is most marked. These latter do not differ materially
from specimens from the south coast of the Red Sea, which we
provisionally refer to typical schlieffent.
SCOLPEINUS SCHLIEFFENI ALBIVENTER.
Size as in typical S. schlieffent.
Colour above “ ecru-drab,” below pure white.
Dimensions of type :—
Head and body 50 mm.; tail 30; forearm 32; ear 9.
Skull—greatest length 12°6; interorbital breadth 4; brain-
case breadth 7:3; breadth across upper jaw at level of m* 5°9:
post-canine tooth-row 3°8.
Hab. Naikhala, Upper Egypt.
Type. Adult male. B.M. no, 4.11.3.4. Original number 73.
Collected 13 Feb. 1904, and presented to the Museum by the
Hon. N. C. Rothschild.
SCOTEINUS SCHLIEFFENI AUSTRALIS.
Size about as in typical S. schlieffeni.
Colour above near ‘‘ mummy brown,” below the same colour but
aler.
‘i Dimensions of type :—
Head and body 50 mm.; tail 28; forearm 31; ear 12.
Skull—ereatest length 13; interorbital breadth 4; brain-
case breadth 7; breadth across upper jaw at level of m°* 6;
post-canine tooth-row 3°8.
Hab. South Africa (type from Inhambane).
Type. Adult male. B.M. no. 6.11.8.19. Original number
1595. Collected 5 Aug. 1906, by Mr. C. H. B. Grant (Rudd
Exploration).
1908. | MAMMALS FROM ZAMBESIA. 541
Mr. Grant took two specimens at Inhambane and the present
series of eight individuals at Tette. There is but little variation
throughout the series, the greatest difference being in size, the,
forearms ranging from 28 to 31 mm. There is absolutely no sign
of the green or olivaceous tinge implied by Noack’s description
of the colour of S. minimus, viz. ‘“ olivengelbbraun,” and the
underside of S. schlieffent australis could by no possibility be
characterised as ‘‘ weissgelb.”
It is worthy of record that in one specimen (2005) of the Tette
series there is present a well-developed second incisor on the left
side of the upper jaw, between the normal incisor and the
canine.
12. CH@REPHON LIMBATUS Pet.
So LB Qe Uae aS)
The type locality of Peters’s Dysopes limbatus was the island of
Mozambique, but he also records it from Sena.
13. NaAsILIO BRACHYRHYNCHUS Sm.
Q. 1974.
“ Native name, ‘ Nyumdundo.’
‘“ According to native report, common, although I was unable
to obtain more than the one specimen.
‘“‘ Inhabiting the more stony parts of the veldt.”—C..H. B. G.
It has been already suggested by Thomas* that Peters’s
Macroscelides fuscus, from Boror, was based on an abnormal
melanistic example of this species.
14. CrociDuRA sp.
@. 1960.
‘“‘ Native name, ‘ Sutsutsu.’
“¢ Apparently very scarce.
“ Frequenting the vegetation and reeds on the river-banks.”—
CV Ho BG.
15. FELIS sERVAL Erxl.
2. 2002.
“ Native name, ‘ Njanjanji.’
“Said to be plentiful, and certainly the spoor was frequently
seen.
‘““ Nocturnal only, often visiting the kraals at night.”—
CE. B..G.
16. GENETTA RUBIGINOSA Puch.
dg. 1968, 1982, 2035.
On collating all the 8. African Genets in the Museum
* Ann. Mag. N. H. (6) xiii. p. 70, 1894.
D42 MESSRS. THOMAS AND WROUGHTON ON [May 26,
Collection we find that they may be arranged in three well-
marked groups, as follows :—
A. Fore feet black.
a. Hairs of dorsal crest and tail long (at least 50 mm. near
base of tail); dorsal spots relatively small with a
distinct tendency to coalesce into longitudinal stripes ;
TMU EDE®) GSMAVINE on ogg ancroocasaasdor nes pooseppaeoan ~ > senos9eccdesnoe
}. Hairs of dorsal crest and tail short (not more than
35 mm. near base of tail); dorsal spots large, always
Gistinchemtanletipn lac kaeereeteeeeereer eee te ts Ceeee terete enn
B. Fore feet pale; hairs of dorsal crest and tail short (not
more than 35 mm. near base of tail); dorsal spots of
medium size, not coalescing; tail-tip black .................. rubiginosa-group.
felina-group.—In his ‘ Mammals of South Africa’ (p. 52, 1900),
Mr. Sclater records a species under the name of Genetta sene-
galensis, from Lake Ngami. The animal he described is probably
the same as Genetta ludia Thos. & Schwann* and is certainly a
member of our jelina-group. The distribution of this group
is thus the central plateau from Namaqualand to the Transvaal,
north of 30°S. lat.; within this area, it is represented in the
south by typical G. felina, and in the north by G. ludia.
tigrina-group.—Occupies the extreme south of Africa below
30° lat., scarcely varying at all, so far as we know, throughout its
range.
rubiginosa-group.—G. letabe Thos. & Schw., belongs to this
group, and it now seems doubtful whether it can be distinguished
specifically from typical G. rubiginosa.
In his unfinished monograph of the Genets + Prof. Matschie,
when establishing G. zambesiana, gives the habitat of G. rubi-
ginosa as “Caconda u. Kiiste von Deutsch Siid-West Afrika,”
but Pucheran distinctly states in the original description that it
was from the Cape of Good Hope. We have compared repre-
sentatives of this group from Natal, Inhambane, N. and E.
Transvaal, Beira, Gorongoza, Tette, and Angoniland, and can
find no essential variation, so that if Natal be taken as the type-
locality, both letabw and zambesiana may have to be considered as
synonyms of Pucheran’s species. The distribution of the group
is therefore all South Africa north of 30° lat. and east of
28° long., extending at least to Angoniland, in 16° S. lat.
“ Native names, ‘ Mpiswi’ and ‘ Mwilt.’
‘“¢Common, especially near kraals, where they cause considerable
annoyance by stealing fowls.
“ Strictly nocturnal, never observed in the daytime.”—
Can BG,
Felina-group.
tigrina-group.
17. CROSSARCHUS FASCIATUS Schreb.
‘“‘ Native name, ‘ Ndembo.’
‘“‘ Not common; found in small troops.
e ees the thickest parts of the bush as at Gorongoza.”—
Ee EB AG:
* P.Z.S. 1906, p. 579.
+ Verhandl. Internat. Zool. Congr. Berl. 1901, p. 1138, &c.
1908. ] MAMMALS FROM ZAMBESIA. 543
18. MtncGos AURATUS, sp. n.
Os LOKG, 9 96:
A brilliantly fulvous Mungoose about the size of WV. ratlamuchi,
but differing in having the hairs of the back annulated.
Size about as in WW. ratlamuchi. General colour above
“ochraceous buff,” darker on the back and tail; below “ ochra-
ceous buff.” Individual hairs of the rump and back, as far
forward as the shoulders, basally “‘ drab-grey,” then “ ochraceous,”
paling to “ buff” at the tip, with a subterminal black ring ; those
of the crown and face ringed buff, black, buff and tawny; those
of the nape, sides of the throat, shoulders, flanks, limbs, and belly
‘ochraceous buff” almost to their bases, which are “ mouse-grey.”
Tail coloured like the back for two-thirds its length, then dark
“tawny,” with a black tip 60-70 mm. long.
Skull as in WZ, ratlamuchi.
Dimensions of type :—
Head and body 324 mm.; tail 290; hind foot 62; ear 26.
Skull—condylo-basal length 62; basilar length 57 ; zygomatic
breadth 33; palate breadth across p* 21; length c—m' 21°5.
Hab. Tette, Portuguese Zambesia.
Type. Adult female. B.M. no. 8.4.3.46. Original number
1976. Collected August 26th, 1907.
A second specimen, a younger female, is quite like the type, and
Mr. Grant assures us he saw several more.
This beautiful new Mungoose is an unexpected discovery, as
Tette is the type-locality of Peters’s Herpestes ornatus, which
Mr. Grant supposed he had secured. But ornatus, as shown by
Peters’s figure and descriptions, is allied to and probably identical
with the much darker coloured J/. cawui Smith *, of which the
Museum possesses specimens from both north and south of
Tette.
‘* Native name, ‘ Runkoe.’
“Several of this species were observed, but were difficult to
trap.
“ Found everywhere, especially near kraals.
“ Certainly diurnal, perhaps nocturnal also.” —C. H. B. G.
19. FUNISCIURUS CEPAPI SINDI, subsp. n.
6. 1961, 2006, 2026. 9. 1941, 1969, 1985.
On laying out the avaiable specimens of J”. cepapi it becomes
evident that there are two well-marked geographical races, a
northern and a southern, separable on their coloration. The
type-locality is given by Smith as “the banks of the Marikwa R..,”
i. é., the upper basin of the Limpopo River, in the southern part of
* of Wroughton, Ann. Mag. N. H. (7) xx. p. 120, 1907. It may be noted here
that the subspecies from Zanzibar described by Wroughton in this paper as Wwngos
melanurus lasti is antedated by Herpestes ornatus rufescens Lorenz (Abh. Senck.
nat. Ges. xxi. Heft iii. p. 462, 1898), a name of which no indication is given in the
title to the article, and which has therefore been missed by all recorders and
bibliographers since.
Proc. Zoou. Soc.—1908, No. XXX V, 35
544 MESSRS. THOMAS AND WROUGHTON ON [| May 26,
the combined range, and we therefore separate the present series
as a northern subspecies under the name of /. cepapt sindi.
Size as in typical /’. cepapi. Fur rather shorter (5-7 mm. on the
back). Colour-pattern above asin true /’. cepapi; below pure white
all over instead of the white being limited to (at most) the chin,
throat, and chest as is the case in /’. cepapi, which has the belly
washed with clay-colour. Back of thighs and midrib of tail
beneath bright ochraceous. Individual hairs of tail ochraceous
with two black rings, so that when the hairs are spread out at
right angles to the midrib there are two black longitudinal bands
running the whole length of the tail (as seen from below) parallel
to the midrib; in typical 7. cepapi the ground-colour of the tail-
hairs is dull “clay-colour” with three black rings, and con-
sequently the resultant black longitudinal stripes are three in
number. ‘Tail equal in length to head and body, proportionally
somewhat shorter than in typical J’. cepapi.
Skull slightly smaller, brain-case broader and fuller.
Dimensions of type :—
Head and body 170 mm. ; tail 168; hind foot 39; ear 20.
Skull—greatest length 43; basilar length 33; interorbital
breadth 12; length of upper molar tooth-row (exclusive of
os
; Mab Lower Basin of Zambesi (type from Tette).
Type. Adult female. B.M. no. 8.4.3.51. Original number
1941. Collected 18th August, 1907.
The specimens from Gorongoza mentioned in our last paper on
the Rudd Exploration (P. Z. 8. 1908, p. 169) must be included in
this subspecies, though they show the distinctive characters less
markedly than the Tette series.
‘“‘ Native name, ‘ Sindi.’
‘‘Common, generally observed in pairs.
‘‘ Living on the berries, &c., of the trees and shrubs, for which
they may often be seen hunting on the ground.
“When alarmed they quickly make for some large tree and
disappear into a hole or cavity.
“Diurnal, active in the early morning and late afternoon,
resting during the heat of the day.”—-C. H. B. G.
20. TATERA LOBENGULH de Wint.
3. 1947, 1951, 1952, 1953. 2. 1942, 1943, 1944, 1945, 1946,
1954.
Externally these specimens cannot be separated from those from
Beira and the Limpopo Valley, 7. e. from 7. lobengule bechuane,
but while having the same narrow skull as that subspecies they
pproximate to 7’. lobengule mashone in having rather smaller
bulle than the Limpopo form.
“Native name, ‘ Mpynya.’
“Common and found everywhere, especially in clearings and
native lands.”—-C. H. B. G.
1908. ] MAMMALS FROM ZAMBESIA. 5A5
Peters’s ‘‘ Meriones leucogaster” (type-locality Mesuril) is a
member of the short-tailed group, of which most of the species
are found north of the Zambesi.
21. ARVICANTHIS DORSALIS CALIDIOR, subsp. n.
O. 1962,
Comparison of the series from the Zambesi Basin with those
from the Transvaal and Zululand, which, as the type shows,
represent the true Arvicanthis dorsalis of Smith, establishes the
fact that individuals of the former are easily separable by their
darker, warmer colouring, and we propose to separate them as a
geographical race under the name of Arvicanthis dorsalis calidior,
Size and fur as in typical A. dorsalis.
Colour-pattern richer and darker than in the southern form.
General colour “‘ chestnut” above, individual hairs dark slate with
“‘ vinaceous cinnamon ” band and black tip; in true 4. dorsalis the
general aspect is near “clay-colour” and the pale rings of the
individual hairs are the palest buff.
Dimensions of the type :—
Head and body 1385 mm.; tail 146; hind foot 27; ear 17.
Skull—greatest length 34; basilar length 27; zygomatic
breadth 16; diastema 8°5; upper molar series 6.
Hab. Zambesi Basin (type from Tambarara, Gorongoza
Mountains).
Type. Old male. B.M. no. 8.1.1.72. Original number 1817.
Collected 13th March, 1907.
The present specimen from Tette, though immature, is identi-
fiable as belonging to this subspecies, in which also should be
included the specimens dealt with in our paper on the collection
from Beira (P. Z.S. 1907, p. 779), a series in the Museum
Collection from Mashonaland, collected by Mr. J. ff. Darling,
others presented by Mr. C. F. M. Swynnerton from Chirinda, &e.
‘« Native name, ‘ Mhoni.’
““ Rare in this district, the specimen sent being the only one
taken or observed.”—C. H. B. G.
22. Mus micropon Peters.
3. 1964, 1965, 1966, 1972, 1973, 1983, 1984, 1987, 1988,
2008, 2. 1979, 1981, 1990, 1991, 1992.
Topotypes of species.
The characters of these specimens prove that the group of
South African multimammate mice is divisible into two species.
The present one, with a tail equal in length to the head and body
combined, extends, so far as we can judge from the specimens
available, from Natal and Zululand northwards along the coast
and throughout the Northern Transvaal and Rhodesia to the
Zambesi. The second species, Mus coucha, recognisable by its
proportionally much shorter tail, is represented in the Museum
35*
546 MESSRS. THOMAS AND WROUGHTON ON [May 26,
Collection from the South-West Transvaal, Bechuanaland, Orange
River Colony, Basutoland, and as far south as Deelfontein and
King William’s Town, in Cape Colony (i. ¢., about 33° S. lat.).
Mr. Sclater in his ‘Mammals of South Africa’ records it from
the Cape and Namaqualand, but Mr. Grant failed to obtain it
in either of these localities.
We are doubtful if Mus coucha zuluensis Thos. & Schw.* can
be retained as a subspecies distinct from true J/. microdon, with
which its describers had not an opportunity of comparing it.
‘“¢ Native name, ‘ Ntisha.’
“¢ Abundant everywhere ; habits similar to those of JZ, coucha
zuluensis.”—C. H. B. G.
93, Mus CHRYSOPHILUS INEPTUS, subsp. n.
g. 1949, 1950, 1963, 1971. 2. 1978, 1989.
Like true chrysophilus, but with lower skull and narrower
brain-case.
Size as in the typical form, but hind foot on the average shorter
and tail proportionally longer. Colour also as in chrysophilus but
paler, the slaty bases of the hairs markedly paler both above and
below.
Skull about the same length as in the type form, but markedly
narrower and flatter; the brain-case much smaller; the whole
skull lower, height from alveolus of m* to crown 9 mm. against
10 in true IW. chrysophilus. .
Dimensions of type :—
Head and body 148 mm. ; tail 182; hind foot of ear ie
Skull—greatest length 37; basilar length 29; zygomatic
breadth 17; brain-case breadth 13; nasals length 16; diastema
9-5; molars 5:7.
Hab. Tette, Portuguese East Africa.
Type. Old male. B.M. no. 8.4.3.73. Original number 1949.
Collected 22nd August, 1907.
Tn the series of 7 specimens obtained by Mr. Grant several have
broken tails, but in those which are complete, the head and body
varying between 145 and 150 mm., the tail reaches 180, whereas
in typical I. chrysophilus specimens of the same size have a tail-
length of about 170. The greatest breadth and brain-case breadth
in the type skull of the species are recorded by Mr. de Winton as
18 and 15 mm. respectively; a comparison with the similar
measurements given for this local race shows how markedly
narrow its skull is; moreover, in true J/. chrysophilus the
greatest breadth is at the posterior end of the zygomatic arch
while in W/. c. ineptus it is across its anterior end. The type is
distinctly older than the individual described by Mr. de Winton
‘‘ Native name, ‘ Kwisikwisi.’
eee de 2 aA
fs ee) y conn ane inhabiting both the bush and the native
% P Z.S. 1905, 1. p. 268;
1908. | MAMMALS FROM ZAMBESIA. 547
We take this opportunity of describing a second local race of
M, chrysophilus :-—
MUS CHRYSOPHILUS ACTICOLA, subsp. n.
A large coast form of JW. chrysophilus, the hind foot always
markedly longer than in that animal.
Size somewhat larger than in true J/. chrysophilus.
General colour as in the type form, but the slaty bases of the
hairs of the under surface markedly shorter and very much paler
than in the true chrysophilus, in which the slaty bases of the hairs
of the lower surface of the body do not differ materially in shade
from those of the back.
Skull larger than im the typical subspecies, but the bulle
slightly smaller. Height at m°’ 10 mm.
Dimensions of the type :—
Head and body 155 mm.; tail 202; hind foot 34; ear 23.
Skull— greatest length (c.) 39; basilar length 30:5; zygo-
matic breadth 19; brain-case breadth 15; nasals length (c.) 15;
diastema 10; molars 6:2.
Hab. Coast between Limpopo and Zambesi Rivers (type from
Beira).
Type. Adult male. B.M. no. 7.6.2.59. Original number 1752.
Collected 25th December, 1906, by C. H. B. Grant (Rudd Ex-
ploration).
The type of J. c. acticola is of about the same age as that of
true VW. chrysophilus. Ina long series of adult specimens from
Inhambane and Beira the hind foot is recorded as low as 30 mm.
in only three specimens, whereas in a large number of typical
M. chrysophilus it never exceeds 29. The tail would seem to be
proportionally quite as long as, or even longer than, in J/. c.
ineptus.
24, Mus AVARILLUS, sp. n.
2. 1980.
A mouse outwardly resembling J/. namaquensis, but with the
teeth of J. chrysophilus, of which it is probably a dwarf relative.
Size rather smaller than in I. namaquensis. Fur soft and
silky, but rather short (8-9 mm. on the back, 5-6 on the belly).
Colour as in I. namaquensis, but the shorter coat allowing the
basal slaty portion of the hairs to show through gives the upper
side a duller general colour and makes the belly greyish white
instead of the apparently pure white of namaquensis.
Skull a copy in miniature of W. chrysophilus. Teeth as in that
species.
Dimensions :—
Head and body 105 mm. ; tail 143; hind foot 26; ear 21.
Skull—greatest length 31; basilar length 23; greatest breadth
14; brain-case breadth 12°5; interorbital breadth 4°9; nasals
11°6; diastema 7°6; upper molar series 6; bullee 5:3.
548 MESSRS. THOMAS AND WROUGHTON ON [May 26,
Hab. Tette, Portuguese Zambesia. oh
Type. Adult female. B.M. no. 8.4.3.79. Original number
1980. Collected 27th August, 1907.
Outwardly this species has a most striking resemblance to
WM. namaquensis, but its skull and teeth characters separate it.
readily from any member of that group. The shape of the skull
and large size of the teeth point conclusively to its close affinity
to M. chrysophilus, of which it is no doubt a dwarf form.
[Mus arsorarius Peters.
As this species was based on specimens from Tette we have
given special attention to its identification. Peters in his
description * mentions two specimens, but as he quotes the
dimensions of the female in the diagnosis and figures its skull
(that of the male being still in the stuffed specimen 7) we adopt it
as the type. He gives an excellent figure of the skull (J. ¢.
pl. xxxv. fig. 7), and this agrees in all essential characters with
that of a specimen from Salisbury, Rhodesia, belonging to a
widely-spread species, of which we have been able to recognise at
least four local races, as follows :—
1. Mus wamaquensis Sm. Hab. Namaqualand. (= Mus
auricomis de Wint.,” Thos. & Schw. P. Z.8. 1904, p. 179.)
2. Mus NAMAQUENSIS CENTRALIS Schw. Hab. Deelfontein, C.C.
(= Mus auriconis centralis Schw. P. Z. 8. 1906, p. 107.)
3. Mus NAMAQuENsIS LEHocLA Sm. Hab. Kuruman. (= Jus
lehocla Sm. Type locality “ Latakoo.”)
4, Mus NAMAQUENSIS AURICOMIS de Wint. Hab. Mashonaland.
(= Mus auricomis de Wint. P. Z. 8. 1896, p. 802.)
We have unfortunately no material to enable us to judge
whether arborarius is identical with the Mashonaland race or
whether, as seems to us more probable, it forms a fifth geo-
graphical subspecies.
This identification altogether removes arborarius from the
Thamnomys group, to which it has been usually referred, perhaps
on the evidence of the stufted male, which may possibly prove to
be an example of the next species.
The skull of Mus namaquensis has, as Peters’s figure of
“ arborarius” shows, evenly divergent supraorbital ridges, cut:
back anterior zygomatic plate, small bullz, and comparatively broad
parapterygoid fossee, all these characters being in contrast with
those shown by the Tette Thamnomys next to be described.
The definite determination of the old types of Smith’s Gerbillus
namaquensis and Mus lehocla is one of the many advantages.
gained from the study of the Rudd Collection. ]
* Reis. Mossamb. p. 152, 1852.
+ At least when Thomas examined it in Berlin in 1887.
Wo}
1908. ] MAMMALS FROM ZAMBESIA. 54
25. THAMNOMYS RUDDI, sp. n.
3.1970. @. 2033, 2036.
A Thamnomys, belonging to the group in which the charac-
teristic third inner cusps of the upper molars are reduced to a
ridge, and with the mammary formula 1—2=6.
Size about as in Mus namaquensis auricomis. Fur soft and
fairly long (12 mm. on the back). General colour above near
“ clay-colour ”; below pure white. Individual hairs of the back
basally slate-colour for two-thirds their length, then buff; a small
proportion of black hairs scattered through the coat; belly-hairs
white to their bases. Hands and feet white.
Skull with a marked interval (2 mm. or more) between the
henselion and the commencement of the palatal foramina.
Dimensions of the type :—
Head and body 124 mm.; tail 160; hind foot 22; ear 20.
Skull—greatest length 32°5; basilar length 26; greatest
breadth 15°5; brain-case breadth 13:2; interorbital breadth 5;
nasals length 12; palatal foramina 7; diastema 8°5; upper molar
series 5; bullee 7.
Hab. Tette, Portuguese Hast Africa.
Type. Old female. B.M. no, 8.4.3.81. Original number 2033.
Collected 14th Sept., 1907. Three specimens examined.
“ Native name, ‘ Nsunto.’
“This species, although undoubtedly common, is difficult to
secure owing to its arboreal habits.
“Tt inhabits the hollows of decayed and dead trees, in which it
makes warm nests of leaves, &c., of no particular shape, merely
filling up the cavities with débris.
“ Strictly nocturnal; one specimen was shot at night whilst
climbing among the branches of a small tree when I was sitting
up for Galago.”—C. H. B. G.
In working out this Thamnomys we have found that the two
following forms also require description :—
THAMNOMYS COMETES, Sp. n.
A Thamnomys belonging to the same group and about the
same size as the last, but with a markedly longer tail.
Size as in 7’. ruddi, but tail one-third longer. Fur soft, shorter
than in 7’. ruddi (9-10 mm. on the back). Colour almost exactly
as in 7’. ruddi. Hands and feet white.
Skull with bulla and teeth smaller, and palatal foramina pro-
duced more forward than in 7’. ruddt.
Dimensions of the type :—
Head and body 124 mm.; tail 195; hind foot 24; ear 20.
Skull—greatest length 33:2; basilar length 26; greatest
breadth 15:8; brain-case breadth 13:6; interorbital breadth 5 ;
nasals length 12°5; palatal foramina 8; diastema 8-5; upper
molar series 4°5; bulle 6.
Hab. Inhambane, Portuguese East Africa.
950 MESSRS. THOMAS AND WROUGHTON ON [ May 26,
Type. Old female. B.M. no. 6.11.8.115. Original number
1644. Collected by Mr. C. H. B. Grant, 18th August, 1906 (Rudd
Exploration).
Three specimens examined. The proportionally very long tail,
the smaller teeth and bullae, and the extension forward of the
palatal foramina almost to the henselion serve to distinguish
cometes at once from ruddi, which in colour it so _ closely
resembles.
THAMNOMYS SURDASTER, Sp. 0.
A smaller Thamnomys belonging to the same group as the two
described above, but with smaller skull, teeth, and bulle, and
colour-pattern as in the rutilans-group.
Size somewhat smaller than in 7. ruddi. Fur soft, short
(7-8 mm. on the back). General ground-colour above “ clay-
colour,” with strong tawny suffusion on rump and lower back,
often extending forward even to the crown; below pure white.
Hands and feet buff, fingers and toes white.
Skull small, teeth and bulle very small.
Dimensions :—
Head and body (c.) 110 mm.; tail 160; hind foot 22; ear 18.
Skull—ereatest length 29; basilar length 22; greatest
breadth 14; brain-case breadth 12; interorbital breadth 4:5;
nasals length 10°6; palatal foramina 5:8; diastema 7:2; upper
molar series 4; bullee 5.
Hab. Nyasaland and North-East Rhodesia. (Type from
Zomba. )
Type. Adult. B.M. no. 93.5.2.27. Collected by Mr. A. Whyte
in October 1892 and presented by Sir H. H. Johnston.
Three specimens from Zomba examined. The Museum has an
example from Angoniland and one from the East Loangwa
District (collected by Mr. 8. A. Neave), which also appear to
belong to this species.
26. SACCOSTOMUS CAMPESTRIS Peters.
3. 1948, 1977.
Topotypes of species *.
These specimens confirm our opinion that in South Africa
there are two forms in this genus, the smaller ones represented
by S. campestris, and a larger, for which the oldest name is
S. mashone de Wint.
‘““ Native name, ‘ Psuku.’
‘Apparently rather scarce and usually taken in native culti-
vation along the banks of the rivers.
‘The cheek-pouches of the specimens sent contained sweet
potato.”—C. H. B. G.
* Reis. Mossamb. 1852, p. 167, pl. xxxiv. fig. 3 & pl. xxxv. fig. 12.
1908. ] MAMMALS FROM ZAMBESIA. 551
27. STEATOMYS PRATENSIS Peters.
6. 2038, 2040, 2041. 9. 2039.
Topotypes of species.
‘‘ Native name, ‘ Nsana.’
‘“‘ Not found nearer Tette than the southern side of the Luenya
and Mazoe Rivers, and even there not plentifully.
“Lying dormant in small burrows throughout the winter. The
specimens sent were dug out and were excessively fat and lazy.”—
CHEE Ie es
98. HippoporaAMUS AMPHIBIUS L.
3. Luenya River.
‘“‘ Native name, ‘ Umvu.’”—C. H. B. G.
29. CEPHALOPHUS GRIMMI L,
3. 1997.
‘‘ Native name, ‘ Mhemwi.’
“ Fairly common, but as it generally inhabits the thickets it is
not easy to shoot.
“ Feeding in the early morning and late afternoon and probably
throughout the night.
“Observed going to water just before sundown.”—C. H. B. G.
30. RAPHICEROS SHARPEI CoLONICUS Thos. & Schw.
@. 2034 (juv.).
‘“‘ Native names, ‘ Kesenyi’ and ‘ Gagoro.’
“Several of this species were seen, but always in such thick
country that it was impossible to get a shot.
“Tnhabiting broken and hilly country, thickly bushed.”—
Cor. BAG:
31, NESOTRAGUS LIVINGSTONIANUS Kirk.
, o- 2037 (juv.).
“« Native name, ‘ Mrumsa.’
“ Decidedly scarce, owing principally to the natives driving and
catching them in nets and also to there not being a great deal of
country suitable to their habits.
“Only found in the thickest bush.
“The Livingstone Buck is found near Beira, but is not found
in Gorongoza, where it is quite unknown to the natives.”—
CHE. B.G.
32. ANpyYcEROS MELAMPUS Licht.
3. 1940.
‘“‘ Native name, ‘ Impala.’
‘Only seen in small herds, seldom exceeding six in number,
they having been much shot out, and no old rams were seen.
“ Out feeding on the short grass in the open glades in the early
5d2 MESSRS. THOMAS AND WROUGHTON ON [May 26,
morning, where they can sometimes be approached within shot,
retiring soon after sunset to the thickest and more inaccessible
parts of the bush.
“ The alarm-call is a loud snort.”—C. H. B. G.
The following is a list of the papers which have been published
on the mammals presented by Mr. Rudd, and we have supple-
mented it by mentioning such other papers as have been
published both here (also largely based on Mr. Rudd’s specimens)
and in 8. Africa during the same period, thus making this a
bibliography of 8. African Mammalogy subsequent to the
publication of Sclater’s ‘ Mammals of 8. Africa.’
The Rudd papers are arranged chronologically under the
headings of the localities dealt with :—
J. British Namaqualand.
O. Thomas & H. Schwann, P. Z. 8. 1904, i. pp. 171-
183, pl. vi. (28 species.)
II. $.#. Transvaal—Wakkerstroom.
lid. P.Z.S. 1905, i. pp. 129-138. (26 spp.)
I. Zululand.
id. P. Z.8. 1905, i. pp. 254-276, pl. xvi. (49 spp.)
IV. Knysna.
lid. P.Z.8. 1906, 1. pp. 159-168. (31 spp.)
V. N.E. Transvaal—Klein Letaba and Woodbush.
lid. P. Z. 8. 1906, pp. 575-591. (51 spp.)
VI. E. Transvaal—Legogot.
Nid. P. Z. 8. 1906, pp. 779-782. (25 spp.)
VII. Inhambane—Coguno.
O. Thomas & R. C. Wroughton, P. Z. 8. 1907,
pp. 285-299. (39 spp.)
VIII. Beira.
lid. P.Z.8. 1907, pp. 774-782. (29 spp.)
IX. Gorongoza Mts.
id. P.Z.8. 1907, pp. 164-173. (32 spp.)
X. Tette, Zambesia (as above).
Tid. P.Z.8. 1908, pp. 535-552. (32 spp.)
Other papers based wholly or in large part on Rudd
material :—
Andersen, K. Five new Rhinolophi from Africa. Ann, Mag.
INSEE) xiv pape el OAs
— On Hipposiderus caffer Sund. and its closest allies. Ann.
Mag. N. H. (7) xvii. p. 269, 1906.
Thomas, O. A new Mungoose (Herpestes ruddi) from Namaqua-
land. Ann. Mag. N. H. (7) xii. p. 465, 1903.
—— A new Golden Mole (Amblysomus corre) from Knysna.
P.Z. 8. 1905, ii. p. 57. }
1908. | MAMMALS FROM ZAMBESIA. 5D3
Wroughton, R. C. On the various forms of Arvicanthis pumilio
Sparrm. Ann. Mag. N. H. (7) xvi. p. 629,
1905.
—— Notes onthe Genus Zatera, with descriptions of new species.
Ann. Mag. N. H. (7) xvii. p. 474, 1906.
— Notes on the Genus Otomys. Ann. Mag. N. H. (7) xviii.
p. 264, 1906.
-—— On three new Mammals from 8. Africa. Ann. Mag. N. H.
(7) xx. p.-31, 1907.
— On the African Mungooses usually referred to the
FHerpestes gracilis group. Ann. Mag. N. H. (7)
x DOs MOT.
Other papers bearing on 8. African mammalogy that have been
published during the last few years are :—
Broom, R. On some new species of Chrysochloris. Ann. Mag.
IN. EE. (7) sax. p.. 262, 1907.
— A Contribution to the Knowledge of the Cape Golden
Moles. Trans. 8. Afr. Phil. Soc. xvi. p. 283,
1907.
— Further Observations on the Chrysochloride. Ann. Trans-
vaal Mus. i. p. 14, 1908.
De Winton, W. E. On Cynictis selousi de Wint. P.Z.8. 1901,
[Seen Olle la
Gough, L.H. Ona new Species of Rhinolophus from Pondoland.
Ann, Transyaal Mus. i. p. 71, 1908.
Jameson, H. L. On a new Hare from the Transvaal. Ann.
Mag. N. H. (7) xx. p. 404, 1907.
Matschie, P. Ueber die Abinderungen der Ginsterkatzen
(Genetta). Verh. V. Internat. Zool. Congress,
Berlin, p. 1128, 1902.
Schwann, H. List of Mammals obtained? by Messrs. R. B.
Woosnam and R. E. Dent in Bechuanaland.
PZ. S, 1906, 1.9, 101. (25 spp.)
—— On Felis ocreata, better known as Felis caligata, and its
subspecies. Ann. Mag. N. H. (7) xii. p. 421,
1904.
Thomas, O. On two new Hares allied to Oryctolagus crassi-
caudatus. Ann. Mag. N. H. (7) x. p. 244,
1902.
— Onsome new forms of Otomys. Ann. Mag. N. H. (7) x.
p- 311, 1902.
—— The common Hare of Central Cape Colony. Ann. Mag.
N. H. (7) xii. p. 343, 1903.
—— On a remarkable new Hare (Lepus monticularis) from
Cape Colony. Ann. Mag. N. H. (7) xi. p. 78,
1903.
ddA THE REY. T. R. R. STEBBING ON [May 26,
2. Zoological Results of the Third Tanganyika Expedition,
conducted by Dr. W. A. Cunnington, 1904-1905.—
Report on the Isopoda terrestria. By the Rev. THomas
R. R. Srepsine, M.A., F.R.S., F.L.S., F.Z.8.*
[Received April 23, 1908. }
(Plate XX VIL. +
The small collection of African Land-Isopoda which Dr. Cun-
nington has asked me to examine includes only four species. ‘Two
of these are already known. ‘The other two appear to require the
institution of a new genus. Whether the species for which it is
instituted are themselves new may be more open to question.
It is with much reluctance that I accept the responsibility of
offering a decision. In the balance of uncertainties one has at
last to make up one’s mind. At present there are a bewildering
number of species in the genus Philoscia, many of them very
incompletely described, and few, if any of them, completely illus-
trated. A remedy for this state of affairs will not be easily
found. The creatures themselves put difficulties in the way of
the student. Their readiness to wander about the world under-
mines any systematic structure built on geographical distribution.
Their variability seems to separate forms which are specifically
identical. On the other hand, general resemblance seems to
unite forms which, on closer examination, are found to be
distinct. Among the appendages the antenne and uropods afford
especially useful characters, and these appendages are particularly
lable to be detached or broken. The structure of the pleopods,
especially those of the male, is more and more acquiring
systematic importance, but male specimens are not always avail-
able. Fortunately there are naturalists to whom difficulties are
not discouragements so much as incentives to action. This paper
will serve at least some useful purpose if it should induce any
such investigator to deal effectively with the genus Philoscia and
its immediate allies.
Fam. ONISCID 2.
Gen. Metorponortuus Budde-Lund.
1885. Metoponorthus Budde-Lund, Isopoda terrestria, p. 161.
METOPONORTHUS PRUINOSUS (Brandt),
1833. Porcellio pruinosus Brandt, Conspectus Crust. Oniscod.
pp. 19, 26 (Budde-Lund).
a
* [The complete account of the new genus diagnosed in this communication
appears here, but since the name and the preliminary diagnosis were published in
the ‘Abstract,’ the genus is distinguished hy being underlined.—Epr1rTor. |
; + For explanation of the Plate, see p. 560.
deayAersyeallecs OG) ea elegy OG All,
a “Stebbing. J. TRennie Reid Eth Edin?
A. ANCHIPHILOSCIA KARONGAE,n sp B.A CUNNINGTONLnsp.
; Sean iS cyPats CONVEXUS (BuddeLund).
1908. | TERRESTRIAL ISOPODS FROM TANGANYIKA. DDD
1879. Metoponorthus pruinosus Budde-Lund, Prospectus Crust.
Isop. terrest. 4.
1885. Metoponorthus pruinosus Budde- Lund, Isopoda terrestria,
Pp. Loo:
Of this well-known and widely distributed species there is a
single specimen in the collection, found under stones on beach,
Island Camp, Birket el Qurun.
ANCHIPHILOSCIA Stebbing.
Abstr. P. Z.S. 1908, p. 28 (May 26).
Mandibles with setulose lobe adjoining the inner cutting-plate,
and with two to three plumose sete between this lobe and the
unjointed plumose lash which represents the molar. First maxille
with eight spines at the apex, and the strongly sinuous distal part
of the outer margin fringed with fine setules, which are much
longer proximally and a little longer distally than those in the
middle of the series. Second maxille with a small, clearly defined,
finely setulose inner plate, not longer than broad; the outer part
of the maxilla much broader, pellucid, almost unarmed, without
apical cleft. Maxillipeds with terminal joint of palp distinguish-
able from the second. Telsonic segment triangular.
The genus Philoscia was established by Latreille in 1804.
Both in French and Latin the name is on that occasion printed
Philoscie, but it has always been presumed that the Latin form
was due toa printer’s error. Oniscus sylvestris Fabricius, which
is the same as the earlier O. muscoruwm Scopoli, is the only species
mentioned im connection with the genus at its institution. In
any re-arrangement, therefore, of the very numerous forms which
are at present united under Latreille’s generic name, it is clear
that Scopoli’s species must be the standard for those to be
retained under Philoscia.
The new genus characterised above agrees in general with the
definition of Philoscia given by Budde-Lund in 1885 (Isopoda
terrestria, p. 207), A. Dollfus in 1897 (Feuille des Jeunes Natu-
ralistes, No. 317, p. 1), and Sars in 1898 (Crustacea of Norway,
vol. 1. p. 172). Thus, it has the body cval or elongate, scarcely
contractile. The lateral lobes of the head are sharply deflexed.
The first segment of the pereeon has the hind margin regularly
curved. The pleon is more or less abruptly narrower than the
pereon. The second antenne are slender, with three-jointed
flagellum. The rami of the uropods extend beyond the telsonic
segment, and the inner ramus is articulated to the peduncle not
far in front of the outer.
On the other hand, in the mandibles, where P. muscoruwm has
only one penicil or plumose seta, the new genus has three penicils
on the left and two on the right mandible. In this respect
it agrees with the minutely described and carefully illustrated
Anaphiloscia simoni Racovitza (Arch. zool. expérimentale, vol. vii.
556 THE REY. T. R. R. STEBBING ON [May 26,
p. 185, 1907), but it has the setulose lobe with which in
Racovitza’s genus the mandible is not equipped. The first maxille
have eight spines on the outer plate, whereas in Paraphaloscia
stenosoma Stebbing (Willey’s Zoological Results, part v. p. 648,
1900) they have only three apical spines on that plate. The
outer margin of that plate is more strongly simuous and less
uniformly fringed than in P. muscorwm. The second maxille
have the inner setulose lobe distinctly defined and the much
broader outer lobe undivided; while in P. muscorwm the outer
lobe is longitudinally cleft and the setulose lobe has no distinct
lower margin, nor is this margin mentioned by Racovitza, in
whose species the second maxilla otherwise resembles that of
Anchiphiloscia. Tn the maxillipeds the articulation between the
second and third joints of the palp is in this genus discernible
under high magnification. Racovitza declares this palp to be
clearly only two-jointed in A. simoni. In the latter species the
telson is almost semicircular, in the species referred to the present
genus it is triangular. The two species in question are in many
respects in close agreement, but one of them has the pleon
abruptly narrower than the person, with the lateral apices of the
pleon segments inconspicuous from above, while in the other
the narrowing of the pleon is far less abrupt, and the apices of
the third, fourth, and fifth segments as seen from above are well
separated and conspicuously displayed.
The generic name is compounded of dyye, near, and Philoscia.
ANCHIPHILOSCIA KARONGA, sp.n. (Plate XXVIL., A.)
This species shows a close resemblance to Philoscia swarext
Dollfus, from Diego Suarez in North Madagascar (Mém. Soe.
Zool. de France, vol. vill. p. 185, fig. 7 in text, 1895), of which,
however, the mouth-organs are not described. Independently of
these, other features seem to make the identification inadmissible.
These affect the second antennz, the uropods, and the telson.
The body is rather narrowly oval, the head not broad, the last
pereon segment strongly arched over the pleon and then turning
obliquely outwards on either side, not forming acute apices. The
pleon is brusquely narrower than the perzeon, and the adpressed
lateral angles of the third, fourth, and fifth segments are not
discernible from above. The telsonic segment is broader than
long, with the sides straight, but the point not blunted as in
P. swarezt.
The sides of the head curve in towards the front, with the eyes
situated at the angles so formed. The small first antenne have
the second joint intermediate in thickness between the stout first
and tapering third, the latter being scarcely longer than the
-second, and a little shorter than the first. The second antennz
are longer than half the body, the second joint a little longer
than the third but much shorter than the fourth, the fifth nearly
as long as the third and fourth combined and as long as the
flagellum; in this the first joint is longer than the second, but
1908. | TERRESTRIAL ISOPODS FROM TANGANYIKA. 557
searcely longer than the third (without reckoning the brush-like
setiform apical process), therefore not as in P. swarezi nearly
equalling in length the two together. In the example figured,
however, the third joint is a little shorter than the second.
The upper lip has its distal margin evenly rounded. The
cutting-edge of the mandible is divided into four horny-looking
blunt teeth. The secondary plate on the left mandible has the
same horny appearance, but that on the right is pellucid. The
other features have been already discussed. The first maxillee
have the two plumose sete seated on the inner part of the apex ;
of the spines on the outer plate the outermost is, as usual, the
strongest; of the three following the intermediate is short; to
these simple spines succeed four, each with a subapical tooth.
The second maxille have been already discussed. The maxillipeds
have the plate of the large and long second joint extending to the
apex of the palp’s second joint, notched at the inner distal angle,
where it has one minute spine-tooth, a little behind which a
regular spine rises from a little surface-lobe, and at the outer
distal angle the apical margin has two more spine-teeth, also of
very small size. he second joint of the palp is furnished with
one or two setiform spines, and the slender, faintly separated
third joint has setules on the outer margin and apex.
The first gnathopods have the third, fourth, and fifth joints
much broader than the sixth; the fourth and fifth subequal in
length, strongly spmed on the hind margin, the fourth with a
row of tiny denticles on the proximal part of its front margin,
the fifth with oblique rows of spinules on the inner surface, the
distal series projecting beyond the obliquely truncate distal
margin; the sixth jomt has the hind margin proximally furred
with spinules.
The fifth pereopods are, as usual in the allied forms, the
longest of the limbs. They have the fourth joint shorter than
the third or fifth, and the fifth shorter than the sixth. In the
young, taken from the mother’s pouch, with eyes and other parts
well developed, no trace of these limbs could be discerned.
Uropods with inner ramus attached on a level with the outer,
compared with which it is not exceptionally narrow as in
P. swarexi, and it is more than half as long. Each ramus is
tipped with a tuft of sete. Telson in young rounded.
Dimensions about 6 mm. in length by 2°75 mm. in breadth.
Locality. On damp decaying wood close to swamp, Kambwe,
near Karonga.
For the colouring comment will conveniently be made in
comparison with the next species.
ANCHIPHILOSCIA CUNNINGTONI, sp. n. (Plate XX VIL, B.)
This species is in striking agreement with the preceding in
regard to the antenne and all the minute details of the mouth-
organs, but the head is wider and the hinder segments of the
pereon have their postero-lateral angles more acute, the hind
558 THE REY. T. R. R. STEBBING ON [May 26,
margin of the seventh segment being uniformly arcuate and
strongly produced over the pleon; the third, fourth, and fifth
leon segments are not adpressed, but have their postero-lateral
angles well displayed; the convergent sides of the telson are
slightly incurved. The first gnathopods have the sixth joint a
little less widened distally than in the other species, but the
difference is slight. The uropods have the inner ramus attached
slightly in advance of the outer, which is fully double its length,
therefore relatively much longer than in P. annulicorms Budde-
Lund, as shown in the figure by Dollfus (Mém. Soe. Zool. de
France, vol. viii. p. 184, 1895), from Diego Suarez.
In the general colouring of brown, variegated with lighter
markings, on the back, this species agrees with Philoscia mus-
corum and several allied species. It has a thin light stripe
extending from the back of the head nearly to the tip of the
telson. In the pereon this is flanked on either side by a broad
piece of variegated pattern, while the side-plates are dark, each
with one conspicuous light patch, but not, as in P. annulicornis,
having the postero-lateral angles pellucid. In the pleon the first
two segments and the telsonic segment have each three lght
patches—one belonging to the medio-dorsal stripe, the others
separated from it by dark patches. The second antenne have the
long fifth joint proximally dark and distally light. The uropods
have the peduncles generally light and also the proximal part of
the outer ramus; but this is variable, as one specimen shows one
of these rami light to the apex, though the other is dark except
close to the base.
In A. karonge the colour is similar, but rather more dusky, the
medio-dorsal stripe being also less continuous, and in one specimen
the hinder segments of the perzeon have their apices clear instead
of dark.
Dollfus (loc. cié.) remarks of P. annulicornis: “This species is
variable; the specimens of the type (coming from Mount Ambre)
are all of great size (12 mm. by 6:5mm.). Generally, above all in
the plain, the length does not exceed 7 to 8 mm.; sometimes the
joints of the flagellum of the second antenne are subequal; but
above all there are varieties ew colore: the clear spaces at the
angles of the pereeon may be lost, or instead of them there may
be marblings or patches [such as those] situated in the median
region; the antenne are often either entirely light or entirely
dark.” The original says, ‘les taches claires des angles du
pereion peuvent s’effacer, ou étre remplacées par des marbrures
ou taches situées dans ja région médiane,” where a word or two
seems to be missing from the second clause.
Specimens of A. cunningtont measured 7 mm. by 3°5 mm., or
6 mm. by 3 mm., the antennz and uropods in each case not
included.
Locality. Under stones, near Niamkolo Bay, Lake Tanganyika.
1908. ] TERRESTRIAL ISOPODS FROM TANGANYIKA, 5d9
Fam. ARMADILLIDIIDG.
Gen. PreriscyPutis Gerstaecker,
1873. Periscyphis Gerstaecker, Die Gliederthier-Fauna des
Sansibar-Gebietes, van der Decken Exp., p. 526 (Budde-
Lund).
1885. Cercocytonus Budde-Lund, Isopoda terrestria, p. 42.
1885. Peryscyphis Budde-Lund, Isopoda terrestria, p. 293.
1898. Periscyphis Budde-Lund, Die Land-Isopoden Ost-Afrikas,
Vic Oi
1908. Periscyphis Budde-Lund, Swedish Zool. Exp. in Egypt,
No. 264, p. 10.
Tn his latest work Budde-Lund gives a new description of this
genus, having recognised that several of the species from German
East Africa which he allotted to it in 1898 require a different
systematic position. He gives some valuable illustrations of
details in his own two species, P. convexus and P. albescens, but
the typical species, P. trivialis Gerstaecker, remains unfigured,
It is obvious that under the new definition those species which
have the apex of the telsonic segment quadrate are excluded.
PeRISCYPHIS convExuS Budde-Lund. (Plate XX VIT., C.)
1885. Cercocytonus convexus Budde-Lund, Isopoda terrestria,
p. 44.
1885. Peryscyphis convexus Budde-Lund, Isopoda terrestria,
e203.
1908. Periscyphis conveaus Budde-Lund, Swedish Zool. Exp. in
Egypt, No. 264, pp. 2, 9, pl. 1. figs. 20-25.
Head with frontal line evanescent in the middle. The side-
plates or raised lateral borders of the first perzeon segment
broadest where flanking the eyes, narrowing downwards without
quite reaching the hind margin of the segment ; the second and
third segments narrowly rounded at the lower corners, the
following segments more squared, the seventh having the hind
margin very concave in the middle but rather flattened at either
side. The pleon short, with the segments in position successively
shorter in the middle till the telsonic segment, which has a
linguiform termination.
The eyes in our specimens do not appear to have more than
20 ocelli. Budde-Lund gives the genus about 25 to 30.
In the second antenne the joints of the peduncle in order of
length are the Ist, 3rd, 2nd, 4th, 5th, the last decidedly the
largest, and longer than the flagellum, of which the first joint is
longer than the second.
The mandibles have a trifid cutting-edge.
The lobes of the lower lip appear to be almost circular.
The first maxille agree closely with Budde-Lund’s figures, the
inner plate having 2 setze or penicils, of which the inner is longer
Proc. Zoou. Soc,— 1908, No, XXXVI. 36
560 ON TERRESTRIAL ISOPODS FROM TANGANYIKA. | May 26,
and stronger than the outer. Of the 9 spines on the outer plate,
the two outermost are the stoutest, and then the next but one.
I do not find any of them bifid at the tip. The subapical
incurvation of the outer margin is finely setulose, as shown by
Budde-Lund.
The second maxille have the inner plate very small, as broad
as long, and strongly furred with very little setules; the outer
plate membranaceous, broadly rounded, almost unarmed.
The maxillipeds seem to agree well with Budde-Lund’s figure.
The epipod, which he does not show, is large, reaching to the
distal end of the principal joint, or nearly so.
That which appears to be the most characteristic feature of
the species (unless it be shared with P. ¢rivialis) is found in the
uropods. Here the peduncle is large and quadrate, obliquely
rounded off at the distal outer angle, but having its minute outer
ramus inserted in and not protruding beyond a little notch near
the distal inner angle. The inner ramus depending from an
upper arm of the peduncle does not reach its extremity or the
apex of the telson.
The length about 7 mm.
Locality. Under stones on beach, Birket el Qurun.
EXPLANATION OF PLATE XXVII.
A. Anchiphiloscia karonge, n. g. et sp.
n.s. Lines indicating natural size of specimen eae in dorsal view.
a.t. Second antenna, with further enlargement of setiform terminal portion
and its armature.
me.1, ma.2, map. First and second maxille and maxilliped, with further enlarge-
ment of terminal portions.
gu. 1. First gnathopod.
urp., T. Uropod, and telsonic segment.
B. Anchiphiloscia cunningtoni, n. g. et sp.
ws. Lines indicating natural size of specimen fie ured in dorsal view.
m.m., me. 1, map. Mandibles, first maxilla, and maxilliped.
gn. 1. First gnathopod.
urp. Uropod.
C. Periscyphis converus (Budde-Lund).
Specimen in the centre figured in three-quarter dorsal view, without appendages.
a.i. Second antenna.
max.2. Second maxilla.
urp. Uropod.
T. Telsonic segment, with left uropod and part of fifth pleon segment.
The mouth-organs throughout are figured on a higher scale than the
other appendages.
1908. ] ON THE ANATOMY OF ANTECHINOMYS ETC. 561
3. On the Anatomy of Antechinomys and some other Mar-
supials, with special reference to the Intestinal Tract and
Mesenteries of these and other: Mammals. By Frank
E. Bepparp, M.A., F.R.S., F.Z.S8.
[Received April 28, 1908. ]
(Text-figures 111-124.)
ContTENTS.
I. On some Points in the Anatomy of Antechinomys, p. 561.
II. On some Points in the Anatomy cf Phascologale, p. 564.
III. On the Intestinal Tract in some Marsupials, compared with that of
other Mammals, p. 567.
IV. On the Existence of a Siprarenal Portal System in Marsupials,
p. 600.
YV. Resumé of new facts, p. 608.
I. On some Points in the Anatomy of Antechinomys laniger.
An example of this rare Marsupial was kindly placed in my
hands some time since by Mr. Beck of this Society, with the
request that I would examine its anatomy. The specimen
consisted of the carcase after the skin had been removed, and
was therefore incomplete as regards many of the muscles. But
the viscera were intact, and J am able theretore to contribute
something towards a fuller knowledge of this small Polyprotodont
Marsupial. The only memoir known to me which deals with its
anatomy is one by the late Mr. Alston in the ‘ Proceedings’ *
of this Society, written nearly thirty years ago. In this paper
a number of facts is given relating to the internal organs of the
body and there is also an account of its osteology. The viscera.
are somewhat summarily described by My. Alston; and there is
thus some scope for a fuller account of certain organs in the
hight of later work upon the anatomy of mammals.
The stomach of this Marsupial has been figured by Alston,
who has compared it with that of Antechinust. He has, more-
over, divided other related genera by the more globular or more
transversely elongate form of that organ. I submit herewith
(text-fig. 111) a drawing of the stomach of the specimen of
Antechinomys which I have myself dissected and which I hope
is accurate. It will be seen that it agrees more nearly with
Antechinus as represented by Mr. Alston than with Antechinomys
as figured by that naturalist. In particular [ would call attention
to the considerable calibre of the duodenum where it leaves the
stomach, and to the fact that the cardiac half of that organ is
considerably prolonged beyond and to the left of the entrance of
the esophagus. I am led, therefore, to doubt the value of the
* P.Z.S. 1880, p. 454. :
+ Now referred to Phascologale (Thomas, Cat. Marsupials Brit. Mus. 1885).
36*
562 MR. F, E, BEDDARD ON THE ANATOMY OF [ May 26,
characters of that organ in discriminating between these allied
genera.
Text-fig. 111.
Alimentary tract of Antechinomys laniger.
O. Omentum. O'. Splenic omentum. S¢. Stomach.
The intestine presents a number of features of interest. 1% is
in the first place very short, as Mr. Alston has pointed out.
Furthermore, there is no external differentiation into sections.
The tube has the same bore throughout, and the absence of any
trace of a cecum renders it impossible to fix the delimitation
between ileum and colon. The alimentary tract is not only
short but, comparatively speaking, rather wide. There is no
duodenal loop to be distinguished from the rest of the tube.
The interesting fact about the intestine is that the whole tube
is suspended from the middle line of the dorsal parietes by a
single continuous mesentery. This is absolutely uncomplicated
by any secondary attachments of any sections of the gut to each
other, or to the parietes. In other mammals there is at least (in
the majority of cases at any rate) one folding of the gut upon
itself. That is, in the region of the stomach the transverse colon
lies above the duodenum just after it issues from the stomach,
and there is in this region a more or less direct mesenteric
secondary connection between the duodenum and the colon or
mesocolon. ‘There is nothing of the kind in Antechinomys.
Another point of importance to be noted in relation to the
intestine is that the loops into which it is thrown are not fixed.
The whole intestine can be straightened, or rather of course
thrown into a continuous curve, and thus differs markedly from
many forms where the foldings of the intestine are permanent
foldings and cannot be straightened out without tearing the
supporting mesentery. Indeed, it is not too much to say that
the alimentary tract of this mammal is as simple as that of
any Lizard.
1908. ] ANTECHINOMYS AND OTHER MARSUPIALS. 563
Mr. Alston’s figure of the liver of Antechinomys does not agree
entirely with the appearances which I have observed, and the
description which he gives is very brief. I find (text-fig. 112) no
Spigelian lobe, but the caudate lobe is deeply bifid and largely covers
the right kidney. The gall-bladder, which Alston has correctly
stated to be present, appears to me to occupy an unusual position.
Instead of lying in a cleft in the middle of the right central
lobe, as is at any rate often the case among mammals, it lies, as in
some other Marsupials*, to the median side of the right central
lobe almost between it and the left central. I did not find the
right lateral lobe quite so large as Alston has figured it. It is
closely fitted to the caudate. The liver “formula” of this
Marsupial seems to me to be fairly accurately expressible as
follows :—viz., LL > LC = RC > RL = Ca.
Text-fig. 112.
Liver of Antechinomys laniger, abdominal aspect.
Ca. Caudate lobe. g.b. Gall-bladder. 2Z.C. Left central lobe. Z.L. Left lateral
lobe. #&.C. Right central lobe. R.Z. Right lateral lobe.
The spleen is of large size and shows no indication of a
triradiate form ; it is wider at the duodenal end and narrower
at the opposite extremity. The wide extremity of the spleen is
marked by two parallel longitudinal furrows which divide up
this extremity into several finger-shaped lobes not detached from
each other.
The pancreas is very diffuse and scattered, consisting of
numerous small lobules; it forms a more or less continuous
mass which lies partly in the mesoduodenum and partly in the
splenic omentum. It reaches also to the other side of the
stomach and a piece of it lies to the left of the cystic duct between
that duct and the duodenal end of the stomach.
The omentum (text-fig. 111, O) is very short and ends in
pulled out ragged edges as is shown in the drawing. It is
* B.g. Dendrolagus and Petrogale, Beddard, P. Z.S. 1895, p. 131.
564 MR. F. E, BEDDARD ON THE ANATOMY OF [May 26,
nowhere attached to the alimentary canal. The splenic omentum
(text-fig. 111, O') extends a little beyond the left kidney in its
insertion on to the median dorsal line.
The left Lidney is well below the right, and in fact its anterior
end is only just a trifle above the level of the posterior end of
the right kidney. The kidneys are rather large, about 10 mm.
long.
The organs of reproduction are not widely different from those
of other Marsupials. The example of Antechinomys dissected by
myself was a female and apparently fully formed im the organs
of sex. Iam able to fill upa lacuna in our knowledge of those
organs in the Marsupialia. The uterus of each side was swollen
into an almost spherical dilatation near to the origin of the
Fallopian tube into which it abruptly contracted. At the other
extremity this dilatation lessened more gradually, though at the
same time with some abruptness, into the rest of the uterus.
At the junction of the two uteri in the middle line the common
chamber thus formed is of greater calibre than is either of
the uteri just before the junction. There seemed to be a
yudiment of a median cul-de-sac. The two vagine are about
the same calibre as the thinner part of each uterus and are
not specially dilated anywhere. The funnel forms the usual
fimbriated expansion which is attached to the ovary for a very
short space. It and the Fallopian tube are supported by the
anterior ligament of the uteri which is attached to the parietes
in front of the ovary, and thus forms a pocket into which the
ovary can be pushed.
II. On some Points in the Anatomy of Phascologale
macdonellensis.
I also owe to the kindness of Mr. Beck a complete example
of this recently described species of Phascologale, which pre-
sents several features of interest in its anatomy. The genus
itself is not well known anatomically. So far as I have been
able to ascertain, there are only two recent memoirs extant
which deal with the visceral structures of species of Phascologale.
The first of these is in a paper upon Antechinomys laniger by
the late Mr. Alston to which reference has already been made,
and in which the stomach of Antechinus (= Phascologale)
swainsont and Phascologale penicillata ave described, and in the
case of the former figured, as well as some other viscera. In
the second paper—Dr. D. J. Cunningham’s Report upon the
Marsupials collected during the voyage of H.M.S. ‘ Challenger,’ *
some notes are given upon the thoracic viscera (not the abdominal)
of Phascologale calura and concerning the azygos vein.
The example which I dissected was a fully adult female with
no signs of immaturity The teats (3 on each side) were large,
* Vol. v. of the Reports.
1908. | ANTECHINOMYS AND OTHER MARSUPIALS. 565
indeed so large as to suggest that the animal had recently borne
young. Neither were there any signs of immaturity about the
organs of reproduction. Nevertheless, there was a large per-
sistent urachus or umbilical cord. There is no question here of
mere traces or of a minute rudiment of this structure. It was
large and conspicuous, as is plainly shown in the accompanying
figure (text-fig. 113). Whether this umbilical cord does or does
not contain any functional blood-vessels I cannot say. All
vestiges of blood-vessels were invisible on dissection. More-
over, although I have used the term “cord” to express this
structure, it is by no means an exactly descriptive word. When
the animal was dissected under water the membrane forming
the umbilical cord floated out loosely, being merely gathered
together at the point where it perforates the rectus abdominis
muscle to be attached to the skin.
Text-fig. 115.
“Umbilical cord” and adjacent viscera of Phascologale macdonellensis.
int. Intestine. O. Omentum. R. Rectus muscle perforated by umbilical
cord. S¢. Stomach.
The attachment of this membrane to the intestine fixes that
tube, as is natural, in a definite position; if it were not for this
persistent umbilical cord the intestine could be, as I imagine,
laid out in a continuous curve as in Antechinomys. The point
where the umbilical membrane is attached to the gut represents,
566 MR. F, E. BEDDARD ON THE ANATOMY OF [May 26,
I presume, the position of Meckel’s diverticulum. In this case
the greater part of the intestine will be referable to the small
intestine and the colon will be short. The umbilical membrane,
however, is not limited to that portion of it which is attached
to the intestine. In spreads out and is continuous with the
omenta both great and small ; there is absolute continuity between
the umbilical membrane and the stomach and spleen. It would
appear therefore, that Phascologale macdonellensis, like the majority
of Marsupials, has an umbilical placenta. In relation to this I may
say that there was no connection whatever between this umbilical
cord and the bladder. The umbilicus itself lies rather in front
of the last rib, and is situated 10 mm. from the end of the
sternum ; it is, on the other hand, 28 mm. from the anus.
One cannot but think that the case described here is ex-
ceptional, and that the genus or species generally is not
characterised by this persistent umbilical cord. It will be,
however, interesting to enquire how far the omentum is really
connected with the membranes of the feetus in development.
The intestine, as already mentioned, is held in position by the
ligament derived from the persistent umbilical vesicle. It has
a simple course like that of many Lizards, and forms only one
loop upon itself as is shown in the drawing (text-fig.113). It is
not without interest to notice that this single loop is rather like
the single loop thrown over the duodenal region during the
development of the human alimentary canal.
The liver of this Marsupial differs in a few points from that
of Antechinonys. There is in the first place a considerable
Spigelian lobe; the caudate is not so extensively prolonged over
the right kidney as it is in Antechinomys. Otherwise the liver
does not differ much. An examination of an example of the
larger species Phascolomys penicillata (for which I am also
indebted to Mr. Beck) shows the same characters. The liver of
Sminthopsis (of the species S. crassicaudata and S. larapinta)
agrees more with that of Phascologale than of Antechinomys.
In the former at any rate the Spigelian lobe was obvious.
The spleen of Phascologale is unlike that of Antechinomys in
that the Marsupial triradiate form was obvious, particularly in
P. penicillata. In the smaller species described here the bifurcate
end of the spleen, that turned towards the left side of the body,
was rather in the form of a triangular expansion of the end
of the spleen, the base of the triangle lying towards the right.
In Sminthopsis the spleen is more like that of Antechinomys, the
right expansion being little marked.
As in Antechinomys the right kidney is much above the left,
and the same asymmetry is seen in Sminthopsis. T examined
the female generative organs of P. penicillata as well as of the
species which forms the subject of the present notes. The same
very strong dilatation of the distal end of each uterus that I
have described in Antechinomys was to be seen in both of these
species. It was very marked indeed in both. In the larger
1908. | ANTECHINOMYS AND OTHER MARSUPIALS. 567
Phascologale penicillata there was also a considerable dilatation
at about the middle of each vagina. The two vagine and the
bladder open into the common external canal at about the same
level. I did not notice any trace of a median cul-de-sac of the
uterl in the larger species, whose genitalia appeared to be
rather more mature and were in any case larger than those of
P. macdonellensis.
III. On the Intestinal Tract in some Marsupials, compared
with that of other Mammals.
Some of the general features of the intestinal tract in many
Marsupials are already well known. It is well known through
the investigations of many (e.g., Owen*, Forbes 7, Beddard £,
Parsons §, Lonnberg ||, Klaatsch J, Mitchell a &e.) that the
large intestine of the Diprotodont Marsupials is as a rule ty very
long relatively, even—it may be—much longer than file small
intestine. In those Polyprotodontia in which a cecum 1s
present to mark the junction of the two regions of the gut, it
is plain that the large intestine is relatively shorter, but not
so short as in the Carnivora. These facts are shown graphically
for six species of Marsupials by Dr. Mitchell. Though doubtless
it is not asserted that the figures referred to exhibit with
absolute accuracy the relative lengths of the large and small
gut, it is clear that they make a very fair approximation to
accuracy.
Furthermore, it is at least highly probable that more accurate
figures could not have been compiled. For it is well known
that the relative lengths of the two sections of the gut are apt
to vary. For example it has been asserted by Brants (quoted by
Tullberg ft) that in 30 examples of Jus decwmanus the relative
lengths of the small and large intestine fluctuated between a
small intestine ten times the length of the large, and a large
intestine which was only one third of the length of the small
intestine. Tullberg himself, in a series of very careful measure-
ments of eight examples of the common rat, made upon specimens
prepared in exactly the same fashion, found not so ereat but
yet a considerable fluctuation. The extremes in two rats of
equal size were in one example a small intestine of 808 mm. and
a large intestine 201 mm., in the other the small intestine
835 mm. and the large intestine 186 mm. It is therefore
* Dendrolagus inustus, P. Z.S. 1852, p. 103.
v Phascolarctos cinereus, P. Z.S. 1881, p. 180.
t Dendrolagus bennetti, P.Z.S. 1895, p. 131.
§ P.Z.S. 1896, p. 683.
|| Several species, P. Z. S. 1902, i. p. 12.
§| Morph. JB. 1892.
** Several species, Tr. Z. S. 1905, vol. xvii. p. 437.
++ To which at any rate Petawrus breviceps, an insect-eating Diprotodont, is
an exception. See Lénnberg, loc. cit. p. 14.
tf “ Ueber das System der Nagethiere,” Nova Acta Reg. Soc. Upsala, ser. 11. 1899.
568 MR. F. E, BEDDARD ON THE ANATOMY OF [May 26,
impossible, as it would appear, to suggest any improvement Im
the figures of Dr. Mitchell from this side of the question.
Small Int. Large Int. Cecum.
Dendrolagus wiustus 2 ......... Ay: 73 4
Trichosurus vulpecula S ...... 702 81 153
Macropus melanops SB ......++ 156 162 18
Me GUpaMieUs: OO ease 172 54 ik
Pe brachyurus 2 ...... 52 21 24
= BullamOncrea) Meese. /ta- 146 3 24
Hypsiprymnus cuniculus Q.... 24 10 Ie
- gaimardi 3 ... 40 28 2
Apart from the relative lengths of the small and large intestines
and the form of the cecum, or its absence, upon which I have
no new facts of my own to record in the present paper, but
concerning which I take the opportunity of utilising a series of
measurements recorded by my predecessor Mr. W. A. Forbes (see
list above), the intestinal tract of mammals exhibits a series of
modifications in different groups and in members of those groups,
which may be considered under the following headings, viz. :—
(1) The relationship of the various coils of the intestine to
each other. (2) The fixity or freedom of the loops of the small
intestine. (3) The permanent loops of the colon.
Under all of these headings I have a few new facts to record
with respect to the Marsupials and to certain other mammals.
(1) Lhe relations of the coils of the intestine to each other.—lt
is clear from the descriptions given by Owen*, that as a rule
at any ratet the intestinal tract in the Marsupials is a freely
movable tract throughout, having no mesenteric connections
between the colon and other regions. As Sir Richard Owen was
particular to describe such folds and connections when they
occur in other mammals (as for example Rodents), the absence of
any such statements in his papers upon Marsupials leads to the
inference that such do not exist in those mammals; a statement
which I am able to confirm from my own dissections. Other
observers have noted a similar series of facts. Prof. Grant in
describing the anatomy of Perameles nasutat has noted the
dimensions of the several tracts of the intestine but has made
no comment upon any folds of the gut, which would certainly
have been mentioned (one assumes) had they been present, since
the same observer almost at the same time § carefully described
such folds in the Paca. Vvolik, in describing the anatomy |} of
Dasyurus ursiius, does not appear to have said anything about
* On Dasyurus macrourus (=D. maculatus), P.Z.S. 1835, p.7; on Macropus
parryi, ibid. 1834, p. 152; on Dendrolagus inustus, ibid. 1852, p. 103.
y This does not, however, seem to apply to the Wombat (see Owen, P. Z.S. 1836,
p. 49) which requires re-examination from this point of view.
{ Mem. Wernerian Soc. vi. 1832, p. 184. .
§ Loc. cit. & t. cit. p. 183.
|| Tijdschr. wis- en natuurk. Wetensch. 1851, p. 153; briefly abstracted in Ann.
Mag. Nat. Hist. ix. 1852, p. 245.
1908. | ANTECHINOMYS AND OTHER MARSUPIALS. 569
the presence of any folds in the gut, but gives measurements of
the lengths of the several parts of the gut.
The late Sir William Flower * by implication makes the same
statement; for he remarks of the alimentary tract of the Opossum
(Didelphys virginiana) that “the colon forms a single arch and
then passes directly down to the pelvis; but being very loosely
attached by mesentery it is very movable.”
None of these writers, however +, with the exception of
Sir Richard Owen, refers to the fact that among the Marsupialia
there are two grades in the complication of the alimentary tract
to be met with. That author says of Dasywrus niaculatus—< The
mesentery was one continuous duplicature of the peritoneum
extending from the pylorus to the end of the colon as in the
Reptilia.”
It appears to me that by this phraseology is indicated an
intestine like that of <Aztechinomys described in the present
paper £, comparable to an early stage in the development of the
alimentary tract of man before the hinder part of the gut has
been folded over the anterior part. In his Treatise on Com-
parative Anatomy the “reptilian” condition of the gut in
certain Marsupials is more plainly stated.
These two stages in the development of the intestine are
figured in Flower’s Lectures on the organs of digestion in the
Mammalia already referred to and in any textbook of human
anatomy. It is obvious that the conditions obtaining in Antechi-
nomys represent the first stage persistently retained. There
are not many positive facts which lead to the supposition that
any other Marsupial shows the same. The only figures of the
entire alimentary tract of a series of Marsupials known to me
are those of Mitchell already referred to, and of Klaatsch§. But
there is no indication here of differentiation between the lower
and higher types in the direction referred to. Klaatsch, indeed,
figures Dasyurus viverrinus as not possessing the “reptilian ”
form of gut, since it possesses the cavo-duodenal ligament. I can
quite confirm him from an examination of D. maugei.
That this simple form of gut is not due merely to its shortness
and to the mechanical difficulty implied by a folding over, is
proved by the occurrence of the same type in animals belonging
to other orders of mammals. In Centetes ecaudatus, for example,
among the Insectivora, there is precisely the same type. The
continuous mesentery of the gut is nowhere folded over, and the
whole intestine is arranged exactly on the plan of that of
Antechinomys, though it is much longer. Dr. Mitchell’s figure
of Centetes might at first sight appear to bear out my statement.
But if this figure be compared, e.g., with that of Phalangista
* In his Lectures on the Alimentary System of Mammals reported in Med.
Times and Gazette, 1872. .
+ Nor is the fact adverted to by Cunningham in his description of the gut ef the
‘Thylacine, &c., Rep. Zool. Challenger, vol. v.
{t Supra, p. 562. Ak
§ Morph. Jahrb. xviii. 1899, p. 622, fig. 2, p. 664, fig. 10, pl. xxii. fig. ie
570 MR. F. E. BEDDARD ON THE ANATOMY OF | May 26,
vulpina, no difference in this particular between the two forms
is there discernible. Now in Phalangista there is the typical
folding over of the hinder part of the alimentary tube upon the
anterior. The two figures, therefore, which have been referred
to, only hide the essential differences between the two types,
informing us merely of the existence in one and the absence in
the other of a cecum—a fact already well known. It is interesting
to note that Centetes has not always this simple arrangement of
the gut. I found it in one example but not in another, where
there was one folding over, but no further specialisation. This
is also interesting not merely from the point of view of fixity of
characters, but because in the Hedgehog the gut is folded over
upon itself as in most mammals, and is not a simple coil on a
continuous mesentery *.
Tam not quite clear from his description and interpretation
of Zoerner’s 7 results, whether Klaatsch places the Hdentate
Myrmecophaga in the same category as Antechinomys. But I
imagine not, since Z'arsius is described as showing “die einfachsten
Mesenterialverhiiltnisse ” among the Mammalia. In any case,
T can assert that some of the American Edentata also show a
very simple gut, comparable to that of Tarsius and Antechinomys.
In Myrmecophaga jubata and Tamandua tetradactyla the gut
can be laid out without removal from the body in a continuous
loop, precisely like the figure given of the same by Mitchellt. In
this feature the genera mentioned are to be contrasted with
Dasypus and other Armadillos. There is, however, no indication
of the differences in the figures of Mitchell excepting, indeed,
that the mesocolon is drawn on one side of the colon in
one form, and on the other in the other types. This cannot,
however, imply any such difference as is here recorded, since, as
already referred to, the different conditions obtaining in Centetes
and Phalangista are indicated by a diagram which would
imply complete similarity, while Phalangista and Macropus are
* It is possible, however, that Dr. Mitchell (Tr. Z. S. 1905, vol. xvii. p. 532) may
have forestalled me and have met with the same variation in this rather important
feature which I have noted above. In his “Summary of Systematic Inferences” he
writes that “in most .... Insectivora.... the intestinal tract .... (is) suspended by
a continuous mesentery.” ‘This would seem to imply the condition which I have
described above. On the other hand, he does not in the same section say the same.
of the American Anteaters, which in this respect resemble Centetes, so far as one of
the specimens of the latter which I dissected is concerned. And again in the
special section dealing with mammals individually, Dr. Mitchell makes no mention
of a continuous mesentery, but remarks that Hrinaceus is practically identical with
Centetes. It may well be that the recollection of more than one example led to
these two not precisely contradictory, but not altogether concordant statements.
I may recall the fact that some years ago I described (Novit. Zool. vol. viii. 1901,
p- 91) acontinuous mesentery in two other examples of Centetes ecaudatus, and did
not meet with the important variation recorded here.
y “‘Bau und Entwickelung des Peritoneum &c. von einiger Edentaten,” Inaug.-
Diss. Halle, 1881. Not seen by me.
= It is important to settle this matter definitely, for Flower is not conclusive in
his statements. On an early page of the “Lectures,” already quoted, he remarks
that the reptilian character of the gut is “only found among Mammals in some
Insectivora and insectivorous Marsupials.” Later, however, he refers to it as also
existing in Whales and in Myrmecophaga.
1908. | ANTECHINOMYS AND OTHER MARSUPIALS, 571
represented as different. I am disposed to believe from its
shortness that the gut of Bradypus will be found also to conform
to this plan*. The simple colon of all these forms of moderate
length, neither excessively long nor excessively reduced, is, as I
think, in agreement with Dr. Mitchell, a mark of low position
in the series, especially since no modification traceable to different
feeding-habits is recognisable.
On a superficial inspection the intestinal tract of many
Carnivora appears to be constituted on the same simple plan
as that of Antechinomys, Myrmecophaga, Centetes, &c. That is
to say, the gut can be laid out in one continuous coil without
removing it from the body or cutting any mesentery. There are,
however, variations in the degree of freedom of the gut. In
Cercoleptes caudivolvulus for example, the gut can be readily laid
out either to the left or to the right, and then forms a continuous
coil apparently with a continuous mesentery throughout. It
would seem in fact to have retained the primitive arrangement
altogether. In /ctonyx capensis, on the other hand, this spreading
out into a continuous fold is only possible on the left side; the
gut cannot be thus spread out on the right side. So too with
Nandinia binotata. In Ursus syriacus the gut can be readily
laid out on the left side; but I have unfortunately no note as
to whether it can be also spread out to the right so as to present
the appearance of a continuous mesentery like that of the simplest
mammals. In Genetia rubiginosa, however, the gut can, as in
Cercoleptes, be laid out on either side of the middle line without
tearing or unduly stretching any mesenteries. The disposition
of the gut, however, in these animals, though superficially as
has been said that of the simplest forms in the order, is in reality
different. It will be seen that where the end of the duodenal
loop comes near to the middle dorsal line, it is actually fastened
to the mesocolon by a short mesentery, as is shown in the
accompanying figure (text-fig. 120 B on p. 591). The comparative
freedom which the whole intestinal tract enjoys is clearly due to the
partial disappearance of this particular mesentery, the ligamentum
cavo-duodenale. If more extensive, the arrangement of the
intestine in a continuous coiled line from stomach to rectum
would be impossible. If, on the other hand, the mesentery
disappeared altogether, there would be actually a continuous
mesentery from end to end of the gut. I am inclined, therefore,
to believe that the intestine has in these forms undergone a
simplification approximating to the primitive state of the
gut with its mesentery. And indeed it may well be that even
the apparently simple forms lke Amtechinomys are in reality
the terminal stage in such a reduction, and not evidence of the
persistence of a primitive state of affairs. I believe, however,
that the coincidence of this apparently primitive state in such
widely removed types as Centetes and Tamandua is evidence in
* As to Cholcepus, however, Klaatsch observes that “das Lig. hepatocavoduodenale
ist schwach entwickelt.”
572 MR, F. E. BEDDARD ON THE ANATOMY OF [May 26,
the contrary direction. The conditions obtaining in the Carnivora,
and as it would appear in the whole of that group, cannot be
looked upon as a reduction due to small size. For it will have
been noted that the large Bear is quite on a level with the small
Cercoleptes or Ictonyx. It is not only the Carnivora which show
this simplification of the gut. For I have already remarked that
the same state of the intestine and its mesentery is to be seen
alternatively in Centetes. Even among the Primates it exists;
for in Chrysothrix sciureus, as I point out later (p. 577), the
intestine can equally well be laid out along a comparatively
straight mesentery to either right or left side. It must be noted,
however, that in this animal and in the Carnivora the colon is
very short. It is, in the specimen which I dissected, only 63
inches in length, a measurement which agrees exactly with that.
of Martin *.
Although the above facts concerning the Carnivora are I
believe correct, and indeed quite bear out Klaatsch’s figure? of
the intestinal tract of the Cat, where the mesentery attached
to the duodenum and to the middle line is figured and termed
“ligamentum cavoduodenale,” and his statements concerning other
genera. Max Weber has, however, described and figured a
different state of affairs in the Beart. The species examined
was Ursus arctos, and the gut is figured as turned over to right
and left without a trace of this ligament, and described in the
following words :—“ Der ganze iibrige Darm an einer einiachen
Mesenterialplatte (Mesenterlum commune) die mit einfacher
radix mesenterii an der Wirbelsaiile wurzelt aufgehaingt ist.”
There may of course be this difference between the two species
of Ursus, or the case may be analogous to that which I have
described above in Centetes ecaudatus. In any case it is clear
that the majority of the Carnivora (whether Arctoid or Atluroid)
do not bear out the statement of Max Weber with reference
to Ursus arctos.
In more differentiated forms a further complexity is introduced
in the existence of a special ligament joining the commencing
duodenum with the proximal end of the colon. For this Klaatsch
adopts Krause’s term ligamentum colicoduodenale. It is figured
by Klaatsch in several forms, in JM/youxus, Stenops, and human
embryo. Nor has Tullberg neglected this connection between
the small and large intestine in his figures of certain Rodents.
This structure is so persistent in the Rodents that it even occurs
in the case of the small Arvicanthis, where the colic coils are
reduced to a rainimum ; as indeed they are according to Klaatsch’s
figures in Myoxus. It is very important to note that even the
Marsupials with their little specialised gut show traces of the
same; in Vrichosurus vulpecula and Pseudochirus peregrinus
* PZ. S. 1833, p. 89.
+ “Zur Morphologie der Mesenterialbildungen, &c. ii Theil,” Morph. Jahrb. xvii.
1892, p. 646, fig. 4.
{ Die Saugethiere, p. 212.
1908. | ANTECHINOMYS AND OTHER MARSUPIALS. 573
I found this membrane, and I may take this opportunity of
remarking that the omentum also is attached, though for a very
short space, to the colon as in the genus Macropus. Neither
of the facts is represented in Klaatsch’s figure of the gut
of Trichosurus vulpecula*. There remains, however, a sub-
stantial difference between Trichoswrus and Pseudochirus on
the one hand and Macropus on the other, in that in the latter
the connection between omentum and colon is extensive, as is
duly figured by Klaatsch for Macropus bennettii. I cannot
agree with Dr. Mitchell in finding no connection of this kind
between the duodenum and colon in Hyrax. In examples of
this ‘“Subungulate” which I dissected some time since, the
duodenum near to its exit from the stomach was adherent to
the colon in the neighbourhood of (distal to) the paired ceca by
a ligament of some length. Moreover, there was also a fixation
of the omentum upon the same region of the colon. I take
this opportunity of remarking that Hyrax shows a particular
point of likeness to the Perissodactyle Ungulates as I venture
to think. In the latter group without exception, as has been
shown by many anatomists, the cecum is immediately followed
by a single simple loop of the colon. The same occurs in Hyraz,
though it is in that animal not quite so well-marked. The
colon immediately after leaving the single cecum, which I regard
as the equivalent of the Perissodactyle cecum, is folded back
along the cecum and connected with it by a ligament; the distal
limb of this loop is not quite so well established as in the
Perissodactyle, however. I discuss these facts more in detail on
a subsequent page.7.
The attachment of the great omentum to the colon is well-
known as an anatomical fact. Klaatsch has figured this in
several forms. Later in this communication I direct attention
to the same attachments in various Rodents and Lemurs.
J have found no such attachment in Carnivora, nor in
American Edentates (in which I am in accord with Klaatsch).
In Orycteropus, however,—and this emphasises the distinctness of
that type from its alleged relatives in the New World—there is
a fixation of the omentum along two separated lines. The most
anterior is on to the cecum and the very commencement of the
colen. Then follows a considerable area of colon unattached to
the omentum which is again attached to it further along. This
state of affairs reminds us of the transitory condition in the
human foetus figured by Klaatsch (¢: loc. cit. p. 694, fig. 16),
which is, however, permanent in Cebus as he has pointed out.
For other facts relating to the attachment of the omentum and
of other ligaments reference may be made to subsequent pages,
where they are described in several animals.
* Loc. cit. pl. xxii. fig. 7. 7 See below, p. 579.
{ Except in one or two cases where it appeared to me to be distinctly pathological.
The non-attachment of the omentum in Carnivora is not, of course, put forward as
anew fact. I confirm it by fresh instances.
574 MR. F. E. BEDDARD ON THE ANATOMY OF | May 26,
It is possible that the secondary attachment of the omentum
to the colon bears some relation to the formation of the permanent
loops of the colon. For this attachment at least offers a fixed
and more or less immovable area or length, which would permit
of unequal growth in this as compared with neighbouring tracts
of intestine. Further contractions of the omentum during its
growth would obviously tend to emphasise such loops and would
act in the direction of rendering them more permanent. In any
case it is important to notice that where there are no fixed anse
coli, there is at least frequently no secondary attachment of the
omentum to the colon. This is the case for example with
Carnivora, and with Carnivorous Marsupials, with Armadillos,
and Insectivora*. This rule, however, is not universal in its
application; for among the Apes, where there are no definite
anse coli, there is an attachment of the omentum to the colon.
On the other hand, where there are well-defined anse the
omentum is asa rule found to be inserted upon one of the ansz.
The more exact relations in a number of Mammals are as
follows:—Among the Lemurs where there is one colic fold,
whether simple or forming a spiral, the omentum is invariably
attached to the distal limb of the loop or spiral. Among
Rodents I found that in Lagostomus the omentum was attached
along the colon from the beginning of the distal limb of
the ansa paracecalis to the end of the proximal lmb of
the ansa coli dextra. In Hystrix cristata the omentum is
attached along one half of the distal limb of the ansa coli dextra,
the ansa coli sinistra having no such connection. In Sciurus
maximus, however, where the same two anse are present, there
is no such insertion of the omentum. These facts do not appear
to have been dealt with by Tullberg in the Rodents.
The lesser omentum of human anatomists is called by Klaatsch
ligamentum hepatogastro-duodenale. It is the ventral membrane
of the gut. This is universally represented among Mammals.
The posterior continuation of this, however, the ligamentum
hepato-cavoduodenale, is not universal. The cavo-duodenal part
of this has already been referred to in various mammals. The
ventral portion of the membrane, the hepatoduodenale, now
requires some consideration. This is described by Klaatsch in
Echidna, and sought for without finding it in certain Marsupials.
On the other hand, it is stated to be present in various Carnivora,
Rodents, and Lemurs. I found this as a distinct membrane
very plainly to be seen in Ursus syriacus. It is to be noted
that in this Carnivore the caudate lobe of the liver is prolonged
down to the level of the duodenal loop. And thus there was a
possibility, so to speak, of finding the membrane, which indeed
was slight though unmistakable and connected the extremity
* T do not, of course, put forward this non-attachment of the omentum as an
entirely new fact.
+ Klaatsch, Joc. cit. p. 667, fig. 11, would regard the omentum as occupying the
interspace of the loop also. J am not convinced of this.
1908. ] ANTECHINOMYS AND OTHER MARSUPIALS. 575
of the liver-lobe with the hgamentum cavoduodenale. The two
mesenteries were quite continuous. In Hechidna also and other
types in which this membrane exists, the liver descended to the
level of the recurrent duodenal loop, where it was attached by
the hgamentum cavoduodenale. In Hyrax, however, the extreme
end of the liver is distant from the end of the duodenal loop by
a space of fully three inches. There is thus an impossibility of
finding a state of affairs like that described above in Ursus. In
Hyrax the duodenum in the ascending limb courses over the
right kidney, to which it is firmly attached by membrane; the
same kidney is equally attached to the liver by an hepato-renal
ligament. Thus we have a series of membranes connecting
the liver with the ligamentum cavoduodenale. Klaatsch has
regarded this as the equivalent of the entire ligamentum hepato-
cavoduodenale, the arrangement of which with respect to its
several elements will evidently depend upon the relative positions
of the several viscera concerned.
(2) The Permanent Loops of the Colon.—As a rule, with but
few exceptions, the Marsupials possess none of those permanent
loops of the colon which have been termed by Tullberg “ ans
coli,” and by Klaatsch “ flexure coli.” This is evident from the
figures given by Klaatsch and Mitchell and from the descriptions
of others, to some of which I have referred above. Nevertheless,
the group is not absolutely to be characterised thus. I have
found in one example of Didelphys virginiana a distinct perma-
nent loop which was rather wide and lay at a considerable distance
behind the cecum. The specimen in which this occurred was a
male, and in a female of the same species there was no such loop.
I do not connect the variation with sex, but note its presence
as indicative of the commencing formation of these special loops
in the Marsupialia. I also observed something of the same kind
in an individual of Trichosurus vulpecula. Furthermore, in
several species of Macropus* (i.e. M. woodwardi, M. melanops,
and M. hagenbeckt) the colon shortly after it issues from the
cecum and just below the stomach is slightly flexed, and perma-
nently so, into a wavy outline of one or two undulations. This
again is perhaps to be looked upon as a commencement of the
anse coli of more highly differentiated forms.
The permanent loops of the colon have been described in a
great many mammals and by many zoologists. I have, however,
in the course of the past year accumulated a considerable number
of facts in this department of anatomy which partly confirm the
results obtained by others, are partly new (so far as I am aware),
and -in some cases enable me to distinguish between already
published descriptions that do not happen to be in entire
harmony. As to the latter section it must be borne in mind
* Quite possibly in others; but I have no notes on the point except of the species
mentioned above.
Proc. Zoou. Soc.—1908, No. XX XVII. 37
576 MR. F. E, BEDDARD ON THE ANATOMY OF | May 26,
that there is apt to be some variation in these loops, as I point out
more particularly in the case of the Vizcacha (Lagostomus tricho-
dactylus) and the Cape Hyrax (Hyrax capensis). Of the latter
species I have examined an unusually large number of examples,
a fact which naturally gives me some confidence in detailing
the characteristics of the alimentary canal and mesenteries of
this interesting form. My notes reter to the Anthropoidea, the
Lemuroidea, the Hyracoidea, and the Rodentia, which I consider
in the order named.
ANTHROPOIDEA.
Of the Primates I only report upon two or three species,
which happen to be remarkable in various ways. In the Gelada
Baboon (Theropithecus gelada, sometimes called Gelada rueppellr)
the intestinal tract as well as other details in its anatomy have
been described by the late Mr. A. H. Garrod*. In a male
and female dissected by him, the proportions between the
small and large intestines differed greatly. In the male the
colon was 3 of the length of the small intestine, in the female
the proportion was much less, ¢.é., 7%; nearer to 3. The example
dissected by myself was also a female; but the proportions in
length of the two sections of the gut were much nearer equality,
though I have, I regret to say, no exact measurements. The
ascending colon from its very beginning (7. e., opposite to the
entrance of the ileum) and a large portion of the transverse
colon were attached to the great omentum. Moreover, the
greater part of the ascending colon was bound down by a
mesentery to the dorsal parietes. The colon had of course no
fixed loops, which indeed do not occur among the Anthropoidea.
In Semnopithecus melalophus (a species of which the Society
has possessed no previous examples) the small intestine was
thrown into few wide and more or less fixed coils owing to the
shortening of the mesentery. The colon was long, about double
the length relatively of a Cynocephalus porcarius examined for
purposes of comparison on the same day. Its arrangement was
remarkable. The ascending colon and a portion of the transverse
colon were sacculated in the usual way along three bands; and
the greater portion of the descending colon was similar in its
sacculation. Between the two, and corresponding to the greater
part of the transverse colon, was a tract of uniform and small
calibre entirely without sacculations. The omentum was attached
to the mesocolon of the anterior sacculated region of the colon,
but at a considerable distance from the colon.
The cessation followed after an interval by the resumption of
a sacculation in the course of the colon, recalls a quite similar
state of affairs in the colic loop of the Rhinoceros sondaicus,
figured some years ago by Sir Frederick Treves and myself in
that animal. Although the colon has not, as in Theropithecus,
* P.Z.S. 1879, p. 451.
+ Trans. Z.S. vol. xii. pl. xxxiv. figs. 1, 2.
1908. | ANTECHINOMYS AND OTHER MARSUPIALS. 577
a special mesentery not to be confused with the mesocolon, and
attaching it to the parietes of the right side of the body, there
is something of the same kind present. For the cecum of this
Semnopithecus, which is quite blunt at the apex, is fixed by a
membrane to the parietes in the inguinal region on the right
side. This had not the look of a former pathological adherence,
which would, I think, have presented a more irregular appearance.
There were of course no fixed loops in the colon. This special
attachment of the cecum is also found in some other Monkeys.
The general anatomy of the Squirrel Monkey, Chrysothrix
sciureus, has been described by Martin*, who pointed out the
existence of an extremely short colon, which he found to be
only 63 inches in length. I can confirm this and add some
details which bear upon the subject of the present investigation.
The colon, rectum, and short cecum presented almost exactly
the appearance of those viscera in a Viverrid, the large intestine
being slightly curved to the right, and thus showing as in many
Viverrids a rudimentary transverse colon. It is an exaggeration
of the condition observable in Ateles melanochir, where the whole
of the colon is disposed in one bold curve rather more than
semicircular and precisely like that of Armadillos. In Ateles,
in fact, there is a well developed transverse colon, but hardly
an ascending colon. The whole gut appears at first sight to be
suspended on a continuous mesentery, for it can be laid out in
a continuous curve either on the right or left side without
removing it from the body, just as in such types. A closer
imspection, however, shows the presence of a ligamentwm cavo-
duodenale, so that the conditions obtaining in this Monkey are
just like those which have been described above in the Carnivora.
This simplification can hardly be due to reduction in size. For
in the smaller Hapale penicillata the gut cannot be turned over
freely to right or left, and has the normal syphon shape.
LEMUROIDEA.
The ansa coli of the Lemurs offers some particularly interesting
modifications.
The simplest form of colon of those which I have examined
is shown in Microcebus smithti. Of this Lemur the general
anatomy has been described by Martin. The colon shows no
special ansa or anse, but is comparatively short and reaches
the terminal straight portion by a boldly curved tract in which
there are no permanent folds. This arrangement was identical
in two examples of the Lemur which I dissected. This is very
similar to the conditions figured in Cheirogaleus coquereli by
Dr. Mitchell, though I am not quite certain from his description
and figure taken together, whether there is or is not a well defined
colic loop such as occurs in the genus Lemur. Dr. Mitchell
speaks of “a colic loop... . relatively much shorter in Cheiro-
* P.7Z.S. 1833, p. 88. + P.Z.S. 1835, p. 126.
Bre
578 MR. F. E, BEDDARD ON THE ANATOMY OF [May 26,
galeus,” but figures a state of affairs like, for instance, Dasypus
where there is no defined colic loop. It would be, as I think,
convenient to restrict the term colic loop to such a defined loop
as occurs in the genus Lemur, to which I shall now refer,
Sir W. Flower has figured the single ansa coli in the genus
Lemur (in L. flavifrons) and Dr. Klaatsch has figured an
identical loop in Hapalemur, and Dr. Mitchell (without referring
to Flower’s figure or to that of Klaatsch) has figured an identical
loop in LZ. mongoz, var. nigrifrons. They are indeed obviously
identical. J can confirm from my own dissections the existence
of this loop in Z. mongoz, which seems to me to correspond to
the ansa coli dextra of Tullbervg, but of which the constituent
limbs were closer together than is figured by Mitchell, agreeing
therefore more closely with the figure of Flower. In L. macaco
Text-fig. 114.
Caecum, colic spiral, &c. of Galago garnetti.
c.d. Ligamentum colico-duodenale, which lies opposite to a ligament binding,
extremity of colic spiral to commencement of colon. O.Omentum. sp. Colic
spiral. The cut ends of the small intestine are joined by dotted lines.
was the same ansa coli. In ZL. albifrons again the same: {but
the two limbs were a little further apart in both of two
examples. Lemur sclateri was the same. The genus Galago
shows an interesting further development of this simple ansa of
Lemur. I have ascertained that this loop has been fieured
previously in Galago crassicaudata by Flower ; but I cannot accept
the figure given by Dr. Mitchell as representing the facts in even
an approximate fashion. I have lately examined two examples of
Galago garnetti (the species described by Dr. Mitchell), and some
1908. | ANTECHINOMYS AND OTHER MARSUPIALS. 579
years ago a single specimen of G. maholi. In all of these the
ansa coli dextra (if I am right in so identifying the loop) is
coiled as is shown in the accompanying figure (text-fig. 114) into
a short spiral exactly like that of certain Rodents and most
if not all Artiodactyles, though much simpler than the spiral
of the vast majority of the latter. This characteristic spiral
arrangement is entirely lost in the figure given by Dr. Mitchell*,
to which I here refer, though accurately represented by
Flower. The matter is of special interest because the same
spiral arrangement of the ansa coli dextra is to be seen in the genus
Nyeticebus of the subfamily Lorisine, as is abundantly shown
in the figures and descriptions of Vrolik ft and Schroder van der
Kolkt, which I am able to confirm by the dissection of two
examples of Vycticebus tardigradus. The resemblance to Galago
is exact. J am disposed to think that Dr. Mitchell’s figure of
the Potto requires revision, in which animal he represents
two adjacent ans coli disposed like those of many Rodents.
Dr. Mitchell has not referred to the papers cited below.
The dissection of an example of /rdris enables me to confirm
the figures of Milne-Edwards§ as to the existence of a colic
spiral in the Indrisine which is more elaborate than in the
other types just dealt with. I may be permitted to point out
another relation between these Lemurs with a spirally twisted
ansa coli|| which has not been insisted upon. In the genera
where this occurs, 7. e., in Galago, Loris, Nycticebus, Indris, and,
I imagine, Perodicticus, the characteristic carpal vibrisse are
absent; they are present in the remaining Lemurs with no
ansa coli or only a simple one. Specialisation of structure has
occurred concurrently in two features of thelr organisation. It
is important to notice that in these Lemurs with a simple or a
spirally twisted ansa coli, the omentum is attached to the loop
or spiral. Furthermore, the opposite side of the spiral is
attached to the cecum, or to the colon just where it leaves the
cecum, by a ligament, and the duodenum is attached to the
colon just opposite to this latter ligament by a colico-duodenal
ligament.
HYRACOIDEAS
I have examined several examples of Hyrax capensis and have
more particularly studied two specimens of which one was
specially favourable for study. This example was not more
* Loe. cit. fig. 41, p. 506.
+ N. Verhandel. Nederlandsche Inst. Amsterdam, vol. x. 1844, p. 75. fe
ft Tijdschr. Natuurk. Gesch. Leiden, viii. 1841, p. 277, pl.v. figs. vi., vil.; and ajoint
paper by these two anatomists in Bijdragen tot de Dierk. i. 1848-1854, p. 29.
Gegenbaur also (Vergl. Anat. Wirbelth. ii. 1901, p. 178) figures the spiral in
Loris gracilis. So also does Klaatsch, though not, as I think, quite accurately
in Nycticebus.
§ Hist. Nat. Phys. et Polit. de Madagascar. :
|| I cannot understand Klaatsch’s statement that “die rechte Colonflexar ist . .
bei Lemur relativ noch miachtiger enttaltet als bei Stenops.”
580 MR. F. E. BEDDARD ON THE ANATOMY OF [May 26,
than half grown, measuring not more than a foot in extreme
length. The alimentary tract of this animal has been described
at considerable length by Dr. Mitchell, but I find myself in
grave disagreement with him, the disagreement extending to
matters of fact as well as to interpretations of the value of
certain structures. JI furthermore deal with certain points
which are not dealt with by Dr. Mitchell.
When the body-wall is cut and the halves reflected, the intestinal
tract is seen to be represented by the large caecum which occupies
the greater portion of the left moiety, and by the paired ceca
and the immediately preceding and succeeding sections of the
colon which occupy the right moiety of the superficial part of
the abdominal cavity thus disclosed. Above, a few coils of the
small intestine are visible, but very few. As Dr. Mitchell has
correctly stated, the duodenal loop is longer than he has re-
presented in his figure *. It extends in the very general fashion
down to about the middle of the lumbar region below the kidneys,
and is there attached by the usual ligamentum cavoduodenale.
This loop of the duodenum shows on the opposite side another
remarkable mesenteric fold. When the single cecum is turned
forwards, it is seen that a mesentery with a free edge directed
forwards runs over the duodenal loop, being attached on the left
to the colon where it emerges from the single cecum, and on the
right to the colon where it passes towards the paired ceca. As
this fold has a free edge, a pocket is formed which appears to
be imperforate at the bottom. I have no facts to offer for the
purposes of a comparison of this mesenteric recess with possibly
similar structures in other mammals. The coils of the small
intestine, with the exception of the duodenal loop, are, as usual,
temporary coils, and the intestine can be straightened bit by
bit as it is passed through the fingers. The ileum opens inte
the single ceecum, the resemblance of which to the cecum of
the Perissodactyles is apparent from the descriptions of others
and from an inspection of this portion of the gut which has been
somewhat confused by Dr. Mitchell’s figure. He represents it as
a bilobed dilatation on the course of ‘‘ Meckel’s Intestine.” Jt is,
in fact, almost a facsimile of the cecum of a Rhinoceros, the chief
difference being that it has a much blunter termination than in the
Perissodactyle. It is moreover sacculated, and the relations to it of
the small and large intestines respectively are exactly as in the
Rhinoceros’ cecum. Moreover, the large intestine which emerges
from it is of greater calibre than the small intestine which enters it.
Finally Dr. Mitchell has taken no account of certain mesenteries
related to this cecum which unquestionably suggest its homology
with the usual unpaired cecum of mammals. That the paired
appendages of the gut which arise further down may be the
equivalents of the Edentate paired cecum is quite possible. But
if so, it is only in my opinion further evidence that the latter are
not the equivalents of the usual unpaired cecum of other
* Trans. Z. S. 1905, xvii. p. 461, fig. 14.
1908. ] ANTECHINOMYS AND OTHER MARSUPIALS. 581
mammals. The paired ceca of Hyrax have no mesenteries of any
kind attached to them. They hang perfectly free of peritoneum
folds into the body-cavity. I do not think that any true unpaired
cecum, even the small one of the Carnivora, is thus free of
mesentery.
The only other Mammals known to me in which the cecum
or ceca are thus entirely free of peritoneal folds, are the American
Edentates. On the other hand, the unpaired cecum of Hyrax
has peritoneal folds connected with it. These are not mentioned
by Dr. Mitchell, and they have a very important bearing upon the
nature of the unpaired cecum of Hyrax. It will be seen from the
accompanying figure (text-fig. 115, p. 582), which may be com-
pared with that given by Dr. Mitchell to which reference has
already been made, that the colon where it leaves the cecum is
bent sharply upon itself, and then forms a more or less L-shaped
curve. The whole of this part of the colon is attached to the
cecum by a mesentery which runs to the very tip of the cecum.
When the cecum is examined more closely, the czeco-colic igament
is seen to arise from the cecum along a straight line distant from,
but continuing the line of, the ileum where it enters the cecum.
This fixed loop of the colon is surely to be compared to the single
colic loop of the Perissodactyles, where indeed, as I myself * and
others have pointed out, the same ligament occurs but is much
shorter. The slightly twisted commencement of this ansa para-
cecalis, as I interpret it, is reminiscent of that of many Artio-
dactyles and Rodents (e. g. Arvicanthis: for which see below,
p. 589), but the length of the entire ansa is obviously more like
that of the Perissodactyles. This marked loop is not figured by
Dr. Mitchell, who only represents an alteration in the direction
of the “small intestine ” (as he regards it) where it leaves the
unpaired cecum. There is no mention in the text of any such
ansa coli; the author to whom I am referring contenting himself
with remarking that “the hind gut is subdivided into a distinct
colic loop (C.L.) and a long straight rectum.” The “loop” in
question is not comparable to the ansa coli described by myselt
in this paper, and by others. It is merely the wavy transverse
colon in which there are asaruleno anseatall. It is regrettable
that Dr. Mitchell has used indifferently the same term (‘colic
loop”) for the fixed ansz coli which are permanent structures,
and for alterations in the direction of the gut. The colon of
Hyraa in fact, after leaving the ansa paracecalis referred to, has
no further anse coli. It has an irregularly looped course owing
to its great length, and passes upwards giving off the paired czeca
in an ascending colon, then runs across the body-cavity as the
transverse colon, and descends in a series of wavy convolutions
as the descending colon into the rectum. As is the case in all
of the more specialised Mammalia, the omentum is attached
to the transverse colon. And the mode of its attachment is
* “The Anatomy of the American Tapir,” P. Z.S. 1889, p. 254.
582 MR. F. E. BEDDARD ON THE ANATOMY OF | May 26,
very interesting. The colon where it leaves the region of the
two paired ceca is attached to the stomach by a fold of the
omentum. After this follows a section of the colon to which
there is no such omental attachment, and again at the com-
mencement of the descending colon the omentum is for a second
time inserted upon it. This arrangement of the omental
Text-fig. 115.
Al:mentary tract of Hyrax capensis.
Ce. Cxcum. C. Region of colon in which fixed loops occasionally occur. e.d.
Ligamentum colico-duodenale. 7. Ligament uniting cecum and commencement
of colon. O.Omentum. p.a. Ansa paracecalis.
attachments is precisely like that of Orycteropus as I have
described on p, 573. And, as I point out in describing this
feature in Orycteropus, there is a likeness with the fcetal condition
im man. There is furthermore, as the figure cited shows, a
well-marked colico-duodenal ligament attaching the duodenum
1908. | ANTECHINOMYS AND OTHER MARSUPIALS. 583
immediately after it has left the stomach to the colon opposite
to the attachment of the first part of the omentum. It may be
that the attachments respectively to the colon immediately after
it has left the paired ceca, is an argument in favour of regarding
those ceca as the equivalents of the unpaired ceca in other forms.
For in Rodents the attachments of the membranes in question
are sometimes to the colon immediately after it has emerged
from the cecum in those animals, and the same relations are to be
found in Lemurs. But against this resemblance may be placed
the facts of the attachment of the membranes in question in
Dasyprocta. In that Rodent, as I point out, the omentum and
the duodenum are inserted upon the ansa coli, which hes at
a considerable distance from the point of emergence of the colon
from the cecum. The evidence therefore cannot be regarded
as very strong. Whereas the evidence already dealt with against
the identification is very strong.
It will be clear from the foregoing that Dr. Mitchell’s statement
that “the hind gut divided into a simple colon and rectum merely
conforms to the fundamental mammalian plan” is not correct.
Nor can I agree with him in the further observation that “ the
general pattern of the intestinal tract in Hyrax, however, sug-
gests no affinity with the patterns exhibited by Rodents and
Ungulates.” It appears to me to resemble both. But this is
of course a matter of opinion. It is not without importance to
observe that Hyrax shows some variation in certain of the
features described above. Since writing the account which I
have just given of this Ungulate I have dissected three other
specimens, all of small size like that from which the above
account has been practically entirely drawn. In one of them,
the colon at the end of the transverse section at the further
attachment of the omentum is fixed into a short ansa coli which
is not very narrow, 7. e., the two limbs are not closely approxi-
mated. This corresponds in position to the splenic flexure of
human anatomy, and is very like the ansa coli sinistra of Zamias
striata described under my account of the anse coli of Rodents.
The two other specimens did not show this specialised loop.
Furthermore, the mesocoion undergoes some variations in its
region of attachment to the dorsal parietes. I did not observe
the exact arrangement in the first example dissected. But in
the three now under consideration there were three different
modes of attachment, thus clearly showing a great variation.
In the individual just referred to, this mesentery was attached
altogether outside of the left kidney. In a second individual, the
left kidney lay for the greater part to the left of, z. e. outside of, the
mesocolon, but the attachment of the latter was in part to the
kidney and cut off an angle of that viscus anteriorly and to the
right. In the third example, the line of attachment of the meso-
colon divided the kidney into two longitudinal areas, of which
the inner lay within the mesocolon area and the outer lay outside
of the mesocolon. The pocket of peritoneum referred to was
584 MR. F. E. BEDDARD ON THE ANATOMY OF [May 26,
present in at least two of the subsequently dissected examples
of Hyrax capensis, but the actual excavation to form a pocket
was hardly at all marked. I may further remark that the
cavoduodenal ligament ended some little way in front of the
posterior angle of the duodenum*. In all the other points
treated of in the above description of the intestinal tract of
Hyrax, 1 found these three examples precisely like those
originally dissected. It will be observed that my account sub-
stantially bears out that of George f, though adding some details.
RODENTIA.
As I have dissected a considerable number of Rodents with
a view to the accurate mapping of their intestinal resemblances
and differences, I may be permitted the following remarks,
which, however, contain observations that are partly in accord
with those stated at first hand, or as a result of agreement with
others, by Tullberg. ;
Among the Sciuroidea I have examined the following species,
viz.: Cynomys ludovicianus, Tamias striatus, Sciurus maximus,
Sciurus macrurus, and Sciurus cinereus (text-fig. 116), of which
the second has been also examined by Tullberg as well as
the Common Squirrel and some genera which I have not
seen. Tullberg has not described or figured the ansze coli of
Oynomys. They are typically like those of other Squirrels.
That is, there are two permanent loops, of which the first
is very much the longer. In Zamias striata I find conditions
rather different from those described and figured by Tullberg.
He describes “nur ein rechte parallelschlinge,” but figures two
such loops of which the first, 7. e. that nearest to the cecum,
is the shorter, though both are not so marked as in his
figure of Sciwrus vulgaris. I find in an example of this species
two loops, the usual arrangement in Squirrels. Of these the
first, that nearest to the cecum, is of considerable length;
and the second is much shorter and also much wider, its
constituent tracts of colon being further away from each
other than is the case with the anterior ansa coli. Apart
from the shortness of both ansz, especially the second, the
colic loops of this Squirrel are like those of Sciurus maximus,
where there are two; the longer of these next to the cecum is
actually six inches in length, with the two lengths of intestine
closely approximated; the second loop is very much shorter but
much wider; thereafter the rectum is nearly straight. In
Sciurus macrurus the arrangement is nearly identical, but the
shorter loop is composed of more closely approximated limbs ;
so too with Sewwrus tennanti (W. A. Forbes MS.). In Sciwrus
* This state of affairs would seem to be impossible until it is recollected that
the ascending limb is tightly fastened down to the right kidney as it passes over
the viscus. (See above, p. 580.)
+ Ann. Sci. Nat. (6) i. 1874.
1908. | ANTECHINOMYS AND OTHER MARSUPIALS. 585
vulgaris Tullberg shows a somewhat different state of affairs;
there are two well developed and narrow loops close together
which are, however, of nearly the same length. This agrees
with a figure given by Mitchell of Xerws capensis, and by
Gegenbaur of Cynomys ludovicianus*, and W. A. Forbes (M8.)
of Arctomys marmotta.
Text-fig. 116.
Part of alimentary tract of Sciwrus cinereus.
a.c.d, Ansa coli dextra. a.c.s. Ansa coli sinistra. Co. Cecum. O. Omentum.
One of the Hystricomorph Rodents not figured by ‘Tullberg
is the Vizcacha, Lagostomus trichodactylus. It is stated,
however, to resemble Chinchilla in possessing an ansa coli deatra
and an ansa paracecalis, which however is more free from
the cecum than in Chinchilla.
In the two examples which ! have dissected there are
differences from each other, but I am not clear how far—if at
all—either of them differs from the example described by
Tullberg. In the one, a female, the ansa paracecalis was a very
short, but rather wide loop, totally unlike that figured by
Tullberg for Chinchilla. The ansa coli dextra was long and
* Verel, Anat. d. Wirbelthiere.
586 MR. F. E. BEDDARD ON THE ANATOMY OF [May 26,
narrow, and beyond the colon and rectum lay in loosely coiled
folds which could of course be straightened out. In the other
example, a male, the ansa paracecalis was very large and wide
and commenced immediately after the ceecum, ending only with
the commencement of the ansa coli dextra; the latter was long
and narrow as usual, and the rest of the colon, instead of being
gathered into temporary folds in a comparatively limited space,
ranged, so to speak, throughout the entire colon in a series of
broad loops.
Text-fig. 117.
Part of alimentary tract of Aulacodus.
ed. Ligamentum colico-duodepale. O, O'. Omentum.
Aulacodus swindernianus is one of the types of Hystricomorphs
not dealt with by Tullberg as regards the points under discussion.
I may therefore record the results of a dissection of two examples
of this Rodent. Immediately after the cecum there is a single
enormous loop measuring quite a foot long, which is followed
by a nearly straight colon and rectum. IT am = uncertain
whether this fold is to be looked upon as the paracecalis or
one of the anse coli, dextra or sinistra. Garrod does not seem to
have mentioned it in his account of the viscera of this animal *.
_ I figure this loop of dwlacodus (text-fig. 117) since it differs
in various details from that of any other Rodent known to me.
It will be observed that the two limbs of the loop running
parallel to each other and at no great distance are fringed on
* P Z.S, 1873, p. 786.
1908. | ANTECHINOMYS AND OTHER MARSUPIALS. 587
both sides by a membrane with a free edge. On the one side,
the membrane has a clearly marked perfectly straight edge, and
traced up to the beginning of the loop this is seen to be
continuous with the great omentum. The omentum is thus
attached to the whole of the ansa coli. On the other side is an
equally free fold of membrane, but here the membrane has not
a clear cut edge. It ends raggedly and unequally as is shown
in the figure. I should say that the figure is made without
any cutting of the membranes, which preserve the condition
they showed while the gut lay undisturbed within the abdominal
cavity. I take it that the ragged edge is really the actual free
edge of the omentum, which does not therefore actually end upon
the ansa coli but is continued beyond it.
Text-fig. 118.
Part of alimentary tract of Hydrocherus capybara.
a.c.d. Ansi coli dextra. Cc. Caecum, cut through at about the middle of its
course. O. Omentum. S¢. Stomach.
Although Grant* and Martin? as well as Tullberg have
described the gut of the Paca, Cologenys paca, I think it worth
* Trans. Wern. Soc. loc. cit. + P.Z.S. 1888, p. 54.
588 MR. F, E. BEDDARD ON THE ANATOMY OF [May 26,
while to record my own notes as a contribution towards possible
variations in the nature of the spirally coiled ansa paraczcalis.
The example which I dissected was coiled in a rather compli-
cated fashion. There are altogether eight limbs in the spiral,
and the figure given by Tullberg appears to me to represent rather
fewer. That is to say, the loop originally a straight loop has
been folded upon itself five times as will be seen from the
number of “ limbs.”
Text-fig. 119.
Part of alimentary canal of Dasyprocta punctata.
a.p. Ansa paracecalis. a.c.d. Ansa coli dextra. c.d. Colico-duodenal ligament.
With reference to this characteristic Hystricomorph spiral,
I may mention that in Hydrocherus it is apparently a late
growth. I have dissected two examples of about half the full
size to which the animal attains, and in both of them there
was no complete spiral but merely a loop with a slight twist
upon itself at the distal extremity (see text-fig. 118, p. 587).
Dasyprocta aguti possesses, according to Tullberg, a limited
spiral. I found the same state of affairs in a newly born spe-
cimen, where the spiral was only twisted upon itself once, the
1908. | ANTECHINOMYS AND OTHER MARSUPIALS. 589
condition being therefore much as in the half-grown Hydrocherus
just referred to.
Tullberg’s figure of the Agouti is not quite so satisfactory as
are the majority of his figures. It is so small that various details
are left out. I therefore venture to supplement him by another
figure of an allied species Dasyprocta punctata (text-fig. 119). In
this specimen the colic loop is, as shown, rather larger than in
D. aguti. It is precisely as in the Lemurs G'alago and NVycticebus
(see p. 578). The first part of the colon, as correctly shown by
Tullberg, runs parallel with and very close to the cecum, to which
it is attached by a mesentery. There is a kind of attempt—so to
speak—at its origin of an ansa paracecalis like that of the Murines.
The duodenum is attached both to the colic spiral and to the
commencement of the cecum, which bends back upon itself at
its free end as shown in the figure. The great omentum is
attached to the colon where it emerges from the spiral and also
to the contiguous part of the spiral itself. It is not, however,
attached to the whole left border of the spiral as in the Lemurs
mentioned. I should mention that the ceeco-colic ligament is
attached along one of the two muscular bands upon the cecum,
the other being on the opposite side.
The small Barbary Mouse, dArvicanthis pumilio, has the
simplest colon of any Rodent which I have had the opportunity
of examining *, and the conditions characterising this genus have
net been dealt with by Tullberg. The colon itself is relatively
short and thrown into no temporary folds. The cecum lies on
the right side of the body rather low down, and the colon ascends,
shows a transverse region, and then forms the descending colon.
There is only one ansa coli present, and that is just where the
colon emerges from the cecum. It is there twisted into a short
spiral. This ansa paracecalis is in principle like that of other
Rodents suchas Cricetus. But it is the only loop present in Arvi-
canthis. It is noticeable that it has the characteristically Murine
form. I could find no trace, at the angles formed by the bending
of the colon, between the transverse and descending regions,
of even so rudimentary a persistent loop as there is in Cricetus.
Tullberg has described various points in the anatomy of the two
species of Otomys, viz. O. wnisulcatus and O. bisulcatus, but has
not dealt with the gut except to give the proportions of the
several regions in the latter of those two species. Having had the
opportunity of dissecting O. irroratus, I am able to fill in that
lacuna in our knowledge of the Rodentia. In this Rodent we
find almost exactly the same characters as in Jus. That is, there
is only one colic loop and that is situated just at the commence-
ment of the colon where it emerges from the cecum. This ansa
coli is doubled upon itself once, and this forms an “ N ” which is
bound down to the cecum. There are no other anse along the
course of the colon. The great omentum seems to be not present
* Mus rattus is equally simple, and like MW. decumanus figured by Mitchell—with
perhaps even a less marked ansa coli. :
590 MR. F. E. BEDDARD ON THE ANATOMY OF [May 26,
at all as a free fold or as attached to the colon. This is an
exaggeration of the characters to which Klaatsch has called
attention in Jus and Myorus. The splenic omentum (ligamentum
recto-lienale) is attached to the colon direct as in Jus, and not to
the mesorectum as is often the case in Mammals.
Although the loops of the colon in the Beaver have been figured
by Tullberg, I do not find myself entirely in agreement with that
figure. My own observations refer to the parts of the intestine
to be dealt with as seen from the ventral surface—the surface
exposed on dissection. I gather that the same view has been
taken by Dr. Tullberg, since he represents the rectum as lying
dorsally to those folds. If this be the case, the Beaver is another
example of a Rodent showing some variation from specimen to
specimen as is shown in the Vizcacha. The colon where it leaves
the ceecum bends to the left and runs forward in close proximity
to itself, being bound here bya mesentery. It then curves round
and passes back again parallel to the beginning of the cecum and
large intestine, but on the opposite, 7. e. the right, side, being here
also attached closely to the gut in question by mesentery. The
tube then bends upon itself and runs again parallel to itself for
a little distance, being still attached by mesentery. The colon
diverges to the right and forms the first of two ansz coli like
those of Sciwrus &c. The first of these is attached by its left-
hand limb to the ascending portion of the colon. This loop is
slightly twisted to the left, and is indeed a rudimentary spiral
like that of the Capybara &c. Immediately after the end of this
loop the colon is modified into a similar loop of about equal length.
Thereafter the colon runs in a broad curve to the rectum. It
will be observed that the direction of the colon according to my
observations is at first 1m the opposite direction as described by
Tullberg.
The duodenum has varied attachments to the neighbouring
regions of the colon and to the parietes, the latter of which I am
not able to describe accurately. It is, however, important to note
that the duodenum follows the curve of the first ansa coli and is
attached to it by mesentery; there is also the usual ligamentum
cavoduodenale.
(3) Coils of Small Intestine—As a general rule the festooned
coils of the small intestine are not at all permanent coils; the
intestine can be passed through the fingers ina perfectly straight
line without tearing or in any way distorting the mesenteron.
At the same time of course the entire gut cannot be laid out in a
circle or a portion of a circle owing to the shortness of its sup-
porting mesentery. One portion can thus be freed from the rest,
which in correspondence become closer and denser elsewhere.
There are, however, exceptions to this general rule which I have
observed among the Marsupials. In a specimen of Pseaudochirus
peregrinus it was impossible to straighten out the coils of the
small intestine which were quite fixed. This characteristic,
1908. ] ANTECHINOMYS AND OTHER MARSUPIALS. 591
however, appears to be individual and not to pertain to the
species or genus; for in another example the small intestine
was “normal.” In another Marsupial, “Zpyprymnus rufescens,
the small intestine, considerably shorter than that of Pseuwdochirus,
was also thrown into permanent coils. Here, however, I can only
report upon a single individual.
Among a considerable number of lower Mammals whose alimen-
tary tracts I have recently studied, only Dasypus vellerosus (of
which I have dissected a single example only) shows the same
fixation of the numerous coils of the small intestine.
It is evident that this phenomenon is not a common one among
the Mammalia, and it is at present doubtful how far it is charac-
teristic, in the rare cases where it does occur, of a given species or
genus.
General Considerations.
From the foregoing considerations it is clear that we can trace
a number of stages of evolution of the intestinal part of the
alimentary tract in the various groups of Mammalia which are
not shown in their complete entirety in any one group.
Text-fig. 120.
\\ \
5S
\|
Hl
|
yi
>
A, Diagram of primitive Mammalian gut, Stage I—B, Stage II.
Ce. Cecum. Cav. Lig. cavoduodenale. m. Dorsal mesentery.
In Stage J. the intestine is suspended upon a continuous
mesentery and is not rotated upon itself to form the primary loop.
Proc. Zoou, Soc.—1908, No. XX XVIII. 38
592 MR. F. E. BEDDARD ON THE ANATOMY OF [May 26,
This is met with as an exception in the Marsupialia (e. g.
Antechinomys); in the two Edentate American Anteaters,
Myrmecophaga jubata and Tamandua tetradactyla ; in Centetes
(occasionally), at any rate, as representing the Insectivora ;
in the Elephant, so far as can be gathered from Flower’s
description *, as representative of the Ungulates. In the Odon-
toceti (vide Max Weber 7) this arrangement of the intestine
would appear to be the rule; but not in other Whales. Tarsius
is the only Lemurine Mammal known to form an intestine slung
upon a continuous mesentery <. i
In Stage J/. we have an intestine showing only the single
rotation upon itself without any further specialisation, so far as
concerns the gut. The suspensory mesentery, however, has.
Vext-fig. 121.
Diagram of Mammalian gut, Stage II.
Lettering as in text-fig. 120.
naturally divided into two, the additional one being what Klaatsch
terms the “ligamentum cavo-(or recto-)duodenale.” We find
this state of affairs in Marsupials (e. g. Didelphys, Macropus), in
Edentates (e. g. Dasypus, Orycteropus), in Carnivora (2? without
exception), in Lemurs (e. g. Microcebus),in Primates (? without
* Med. Times & Gazette, loc. cit.
+ DieSiugethiere. See for a particular instance (Porpoise) Hepburn & Waterston,
Trans. R. Soc. Hd. xl. pt. 11. 1902.
{£ Klaatsch, loc. cit.
1908. ] ANTECHINOMYS AND OTHER MARSUPIALS. 593
exception *), A slight modification of this stage is seen in many
Carnivora and in the Primate Chrysothrix, where the intestine
can be laid out flat without tearing the mesenteries ; this is
achieved by the reduction of the hgamentum cavoduodenale, and
it is possible that the entire disappearance of this mesentery may
account for such cases as Centetes, which therefore show an unreal
primitiveness due to a reversion by degeneration.
This stage may be subdivided into two; of which one,
Stage IT. A, will include those formsin which the intestine is merely
folded over once without further specialisation, and will include
some Marsupials (e. g. Didelphys, Hypsiprymnus), Edentates (e. g.
Dasypus), Carnivora (? all genera), Lemurs (e. g. Wicrocebus) ; and
Stage IT. B, which will be characterised by the further specialisation
caused by the attachment to the colon of the omentum :' this
second group will contain many Marsupials (e. g. Macropus),
Edentata (Orycteropus), Primates (division of Anthropoidea
without exception).
Text-fig. 122.
Ply
ZZ
Coe. ; =
Diagram of Mammalian gut, Stage IT. A.
O. Omentuz. Si. Small intestine. Other letters as in text-fig. 120.
In Stage I77. the essential difference from the two earlier
stages is the formation of fixed permanent loops of the colon,
termed ansc coli. Combined with this is always an intestine
* Kilaatsch’s figure (Joc. cit. fig. 12, p. 671) of the young Hapale with an ansa coli
absent in the adult (fig. 13, p. 672) suggests the possibility of the Primate simplicity
being due to reversion.
38%
594 MR. F, E. BEDDARD ON THE ANATOMY OF [May 26,
with the usual rotation of Stage II., and there is also invariably
a connection of the omentum with the colon, and furthermore,
always a secondary connection of the duodenum with the colon at
its commencement. This stage is represented by Lemurs (the
majority), all of the Hyracoidea, Perissodactyla * and Artio-
dactyla T, and, finally, Rodents.
Text-fig. 123.
YH yy / | mT Was AWN O
GAMA Maa Tis K
20.2. NN
a.C.8.
Diagram of Mammalian intestine, Stage ILI.
a.c.d. Ansa coli dextra. a.c.s. Ansa coli sinistra. a.p. Ansa paraczcalis.
e.d, Colico-duodenal ligament. Other letters as in text-fig. 120.
This Stage is hardly divisible into different grades. There are,
it is true, simpler forms and more complexly convoluted colons.
It might perhaps be permissible to place at the base of the series
the Perissodactyla and Hyracoidea where there is but one ansa
coli, and that apparently the ansa paracecalis of other types. If
Klaatsch be right in regarding the coil of Ruminants and Swine
as an ansa paraceecalis, this group would be added. In this case
we arrive at the interesting conclusion that the colic spiral is not
strictly homologous through those groups which show it. For in
the Lemurs it cannot be doubted that the spiral is the ansa coli
dextra. It is only the Rodentia which show the maximum of
coiling of the colon, and in the more differentiated genera of that
order (e. g. Castor) there are three anse coli, though not more,
* T have not dealt above in detail with the single simple ansa coli of Horses, Tapir,
and Rhinoceroses, since it has been so often described.
+ See especially Lonnberg, Acta Ac. Upsala, 1903, K. Vet.-Ak. Handl. 1901, and
some other memoirs.
1908. ] ANTECHINOMYS AND OTHER MARSUPIALS. 595
But other genera, e. g. Mus and Arvicanthis, with one ansa
paracecalis, and Sciwrus with two anse, dextra and sinistra, and
Aulacodus with only the ansa coli dextra, render it impossible to
make hard and fast lines of division.
It will be noticed from the above account of the several stages
seen in the Mammalian gut, that the Lemurs are the only group
in which every stage but one is to be seen in a well-marked fashion.
This fact of itself is enough to negative any accurate classificatory
results to be deduced from the series of facts brought together in
the present communication, though I offer later (p. 596) some
observations upon the affinities of different groups as judged by
the varied modifications of the intestinal canal.
Furthermore, it will be gathered from what has been brought
forward in the present communication that the Marsupials,
although their intestinal tract shows in a well-developed fashion
only two stages, show indications as it were of the third stage.
Occasionally an ansa coli exists, while secondary connections
between the duodenum and commencement of the colon are found
in Trichosurus. Arising, as is now believed, from some early
Kutherian type, the Marsupials seem to have retained the poten-
tialities of intestinal development exhibited in the later Eutheria.
It is important moreover to note that the attachment of the
omentum to the colon in Trichosurus is to the extreme right of
the latter ; for it is on this side that the attachment commences
in Man (Johannes Miiller quoted by Klaatsch).
Simplification of structure does not always imply an archaic posi-
tion with reference to allied forms which show a less simple anatomy.
It has been again and again pointed out that size is an element
which is not to be left out of consideration in weighing such
apparently archaic structural conditions. For example, the smooth
brain of many small mammals is not to be interpreted as evidence
of the lowly position in the series of such smooth-brained types.
On the other hand, the simple organisation of a Naiid Oligochete
as compared with that of a large earthworm, may be at least
partly interpreted as simplicity not altogether due to reduced
size. It is important therefore to note that the simple intestine
of Antechinomys slung upon a single continuous mesentery is not
a feature confined only to such small mammals. In comparing
this form with Arvicanthis, which is even smaller, we find in the
latter the usual rotated intestine with even a fixed colic loop.
And other examples will be apparent from the foregoing pages as
well as from previous writings on the subject. Atthe same time it
is not to be forgotten that other instances may be due to simpli-
fication, and to belong therefore to a different category though
apparently quite similar. :
Although it is true that the specialised loops of the colon are
often associated with a colon of great length, there-is as 1t would
appear no necessary connection between length and complexity,
or even occurrence, of these special loops. The existence of such
coils is in fact a character of given orders of mammals. Amony
596 MR. F. E. BEDDARD ON THE ANATOMY OF [ May 26,
those orders where they occur, that is to say the Rodentia,
Perissodactyla, Artiodactyla, Hyracoidea, and Lemurs, there is no
relation between complexity of coils and length of gut. The
small Rodent Arvicanthis has a short colon not longer than that
of many mammals of other groups without any trace of coils.
And yet it possesses one ansa, the postcecalis. The existence of
three ansze in the Beaver does not argue a longer colon than in
the Agouti, where there is only one ansa, the ansa coli dextra.
The complex spiral of Zndris is not associated with a markedly
longer colon than that which bears the one loop of the genus
Hyrax. \t is therefore clear that we must seek for the origin of
these fixed loops of the colon in some other way than need for
packing away a large tract of gut in a limited space. And it has
been already suggested that this may be found in the attachment
of the omentum.
Value of Intestinal modifications in Classification.
Apart from certain facts given by Weber and others as dis-
tinctive of various groups of Mammals, Dr. Mitchell seems to be
the only person who has attempted to discuss in detail the’
classificatory results to be obtained from a consideration of the
varying characters of the intestinal tract. In a preliminary
criticism of resemblances—an attempt to differentiate those upon
which weight should be laid from those which cannot be admitted
as of classificatory importance, this author has committed himself
to a statement that will not receive the agreement of zoologists.
“« Likenesses ” he writes (on p. 528) “ which are due to the common
possession of primitive features cannot be regarded as evidence
of near relationship; that certain members of a group have
retained what was once the property of all the members of that
group can be no reason for placing such creatures close together
in a system if that system is to be based on blood-relationship.”
It is, I imagine, by absolutely universal consent that Hehidna
and Ornithorhynchus ave placed together in one order, Mono-
tremata, and mainly by virtue of the facts that in both there
is a large coracoid and a generally “ primitive ” shoulder-girdle :
that in both the egg is large-yolked and meroblastic with a
follicle of at most two layers of cells: that in both the anterior
abdominal vein is either present or indicated by a large ventral
mesentery: that in both the heart valve of the right auriculo-
ventricular ostium has retained the partly muscular structure of
that of lower types; and by other features all of which are
prinitive.
One can of course accede to Dr. Mitchell’s assertion that the loss
of a particular character in two groups is no reason for placing them
in proximity, and that a new structural acquisition is evidence of
relationship in proportion to the anatomical complexity of that
structure ; this latter is a perfectly correct restatement of Sir E.
Ray Lankester’s use of the Molluscan Odontophore as a test for
1908. | ANTECHINOMYS AND OTHER MARSUPIALS, 597
the inclusion of a particular type in that phylum. Dr. Mitchell’s
sketch of the “ archecentric ” condition of the mammalian gut
agrees absolutely with the figures given in any text-book of
Human Anatomy * of the early human gut, and any mammalian
gut.
There is in fact no doubt whatever that the primitive
Mammalian gut was in all essentials a gut like that of the Reptilia,
i. e., a tube of no great length, and therefore with but few
convolutions suspended by a contmmuous mesentery and with no
permanent folds of any part. I leave undecided whether a cecum
or ceca are necessary adjuncts of this archetypal intestine, or
whether they or it should lie about halfway down the intestinal
tube.
Greater or less length is clearly of no importance inasmuch as
that feature has been shown to vary in individuals (see above
p. 585). Viewing the matter from this point of vantage, we ought
to regard as most primitive in position any groups or group in
which the alimentary tract has retained this Reptilian character
throughout ; which in fact are so far not one generation removed
from the entire group of Lacertilia (including Hatteria), where no
other conditions are, so far as anatomical investigation has gone,
to be found.
So far as I can say from my own knowledge and from reliable
statements published upon the matter, the only groups in which
this primitive gut exists obviously are the Polyprotodontia
(excluding the American forms), the Xenarthra (excluding
Armadillos), the Proboscidea, the Odontoceti, and the Insectivora.
But with regard to the latter the case of Centetes described above
rather suggests a reversion. The Lemurs can hardly be added,
since Zarsius is the only form which shows this straight
mesentery unfolded anywhere; and as that genus is so minute in
size the feature may be the result of degeneration.
Why Dr. Mitchell should remove from such an assemblage T
the Insectivora, Proboscidea, and Odontoceti, and add to it the
Tubulidentata and Diprotodont Marsupials, is not altogether
easy to understand. His arrangement appears to me to be
so far purely capricious, and to be based upon no facts. More-
over, I would point out that very nearly all zoologists would agree
in regarding the groups which I have thus placed in juxtaposition
as being ancient groups.
Dr. Mitchell, however, appears to me to be perfectly right in
asserting that the Carnivora have not moved far from the common
centre; though why this statement should be qualified by the
suggestion that the reduction of the hind gut is a specialisation
is notso apparent. As Dr. Loénnberg has well pointed out in the
case of certain Marsupials ¢ (and others have pointed out im other
* Of. e.g. D. J. Cunningham’s Text-book. ee
+ According to him the “ancestral group” contains Marsupialia, Xenarthyra,
Tubulidentata, and Galeopithecide. -477 Marsupials, I presume, are included.
ft P. Z. S. 1902, vol. i. p. 12.
598 MR. F. E. BEDDARD ON THE ANATOMY OF [May 26,
groups), there is a close association between the relative lengths of
the regions of the gut and the food, a relation which is by no
means ignored by Mitchell, though he does not quote any previous
memoirs in discussing this matter.
Surely the American monkey Chrysothrix (see above p. 577),
with a very short straight intestine, cannot be considered to
differ importantly by this character from e. g. Hapale with the
usual three-sided Primate large intestinal loop. On the eontrary,
T should be disposed to assert that the short colon of the Carnivora,
persisting as it does through the whole order, differing as they do
widely in their food, is rather evidence of an ancient state of
affairs.
Moreover, a glance at the earliest Mammalia known would seem
to suggest that a carnivorous, insectivorous, or at most omnivorous
way of life was the primitive mammalian mode of life, a view
which is strengthened by reflections upon the origin of the group,
whether from Reptilian or Amphibian like forms. Otherwise it
might be pointed out that on the whole the simple form of gut
was associated with a shortness of gut associated in its turn
with a carnivorous habit. The Elephant however (if I rightly
interpret the investigations and statements of others) seems to
possess a simple gut supported upon a continuous mesentery.
This, however, is by no means saying the same thing as to
assert that the five groups mentioned are to be combined into a
superorder and contrasted with the remaining Mammalia which
stand in various relations to them. On the contrary, it appears
to be totally impossible to classify the mammals by the form of
their intestine, the chief reason for this being that so many
grades are seen in the same group. On the other hand, it may be
confidently said that the Ungulates and Rodents are some way
removed from the base of the mammalian series; for in none of
them are primitive conditions to be seen. These have, it would
appear, become entirely lost.
It is particularly noteworthy that the Anthropoidea (under-
standing by this term the “ apes” and “ monkeys ”), as contrasted
with the Lemuroidea, exhibit primitive characters *, though not so
primitive as the five groups with which we commenced this survey.
There are no fixed loops to the colon,and there are the same
fluctuations in the relative lengths of the small and large intestine
that we find in e. g. the Marsupials. But special mesenteric
connections render complex the coils of the gut, though not so
numerous as we meet with in Rodents. On the other hand, the
Lemurs present us with no particular likeness to the other
Primates. The path pursued by these animals is really much the
same as that pursued by the Rodents and the Ungulates. But
this does not in my opinion imply affinity; it means no- more
than that there is a definite line of increasing complexity of the
gut which is followed in all.
Tn fact, on the whole a study of the intestinal coils of Mammalia
Cf. however Klaatsch, p. 671, fig. 12 for “lemurine” stage in young Hapale.
1908. ]
“eqgoprpoyd ‘eproyyidosrey ‘exegdoaryy ‘erueatg “ztA
YVULIOFUL UTY}.1e9 OU PU Uvo T YoIyM surutedUCD sdnozs aso} 4IUIO T
“TLOT
ANTECHINOMYS AND OTHER
MARSUPIALS. 599
seeercece TIL ase4a
q'1I 9se4g
‘DTT 90R4¢
‘T asnyg
ee eee ree
|
- Marsvuprania.
| cal
XENARTHRA.
| PROBOSCIDEA.
|
_CETACEA.
|
LEMUROIDEA.
INSECTIVORA.
MonovrREMATA.
| CARNIVORA.
| |
| TUBULIDENTATA. |
| ANTHROPOIDEA.
-HyYRAcorpra.
RoDENTIA.
|
UNGULATA,
600 MR. F. E. BEDDARD ON THE ANATOMY OF | May 26,
seems to lend support to the view that existing mammals have
radiated out separately in many directions from a common stem,
and that no two groups are very markedly nearer to each other
than any others.
This view is in accord with certain opinions expressed by
Prof. Osborn. The relative positions of the various groups
(excluding those of which I have not sutticient knowledge derived
from memoirs or from my own observations) can be expressed in
some such diagram (p. 599) as that which accompanies these
remarks. The stages referred to are described on a previous page.
IV. Note on the Heistence of a Suprarenal Portal
System in Marsupials.
Although it is possibly true that physiologically there is no
renal portal or suprarenal portal system *, it is plain that among
Reptiles, as contrasted with Mammals, there are veins entering the
kidneys and the suprarenal bodies from the parietes and hind
limbs forming afferent veins to those glands, and thus contrasting
with efferent veins which convey the blood from the glands in
question directly to the postcaval system. Whether the capillaries
intervening between the afferent and efferent veins are real
capillaries or sinusoids does not affect the anatomical facts just
stated, though naturally of great embryological and physiological
importance. Neither does my own suggestion t that the supra-
renal portal system is really due to the disappearance of that
section of the postcardinal vein which runs over or near to that
organ on each side and the consequent opening of its (the post-
cardinal vein’s) affluents into the substance of the gland. It is
still a fact that among Reptiles the suprarenal organs send a vein
or veins to the postcaval and receive a vein or veins from the
adjacent parietes. And this feature, whatever may be its physio-
logical importance or want of importance, is an anatomical feature
in which the lower Vertebrata differ from the Mammalia.
This being the case, attention is directed to the accompanying
drawing (text-fig. 124, B) which represents the veins in the
immediate neighbourhood of the kidneys of an example of the
Marsupial Dasyurus mauger. It willbe seen that the suprarenal
body of the left side of the body lies anteriorly to the left renal
vein and not in contact with it. The vein from this body opens
directly into the postcaval vein between the points of entrance
thereinto of the two renal veins, but nearest to the left renal
vein. In addition, however, to this vein connected with the supra-
renal body, another vein is depicted in the figure which arises by
more than one veinlet from the adjacent muscles. This vein runs
in a direction parallel to that of the postcaval and ends in the
suprarenal body, which it enters at the end remote from that
whence the suprarenal affluent of the postcaval emerges from the
* Cf. Woodland, P. Z. S. 1906, p. 886. 7+ P.Z.S. 1906, p. 24.
1908. | ANTECHINOMYS AND OTHER MARSUPIALS, 601
gland. Dasyurus viverriiius was exactly the same. In an example
of Macropus agilis the same vein is present, but there are slight
differences from the conditions observed and just described in
Dasyurus maugei. In the Macropus the suprarenal vein enters
the left renal, and the suprarenal portal, as I venture to term the
Text-fig. 124.
Suprarenal veins in various Mammals.
A. Thylacinus. B. Dasyurus maugei. C. Nasua rufa.
D. Peragale lagotis. E. Trichosurus vulpecula. FE. Macropus dorsalis.
G. Dasyurus viverrinus. HH. Bettongia ogilbyi. I. Phascologale penicillata.
p.c. Posteaval vein. K. Kidney. +. Renal vein. Sr. Suprarenal body.
s.7.p. Suprarenal portal. s.7.v. Suprarenal vein.
vein to which attention is directed in the present paper, enters as
in Dasyurus at the opposite end of the suprarenal body. The
suprarenal vein differs from that of D. maugei in that it receives
a tributary from the parietes. The left suprarenal veins of
602 MR, F. E. BEDDARD ON THE ANATOMY OF {| May 26,
Macropus ualabatus were much the same, but I did not observe
the parietal branch of the suprarenal vein.
This state of affairs may now be compared with that observable
in the higher Mammalia. In an example of Vasua rufa the
veins in question had the disposition shown in text-figure 124, C.
There is, asin the Marsupials already dealt with, a vein arising
from the parietal musculature anteriorly. It arises by two main
branches. But the vein formed by the junction of these passes
ultimately to the left side of the suprarenal body, and receiving
from it the suprarenal vein opens into the left renal vein. The
parietal vein in fact does not touch the suprarenal body ; it is
merely an affluent of the suprarenal vein. Jt may be, however,
the homologue of the vein in the Dasyure which enters the supra-
renal body. On the right side of the body there was much the
same disposition of veins observable. But the suprarenal vein
poured its contents into the parietal vein which passes over the
suprarenal body and was directed outwards to the right at right
angles to the longitudinal axis of the body. These details were
worth recording inasmuch as in a second example of Vasua rufa,
and of the opposite sex, the same arrangements were met with and
were alike in every detail.
Reverting to the Marsupials, an example of Peragale lagotis
showed essentially the same disposition of the vein running into
or from the suprarenal body that. has been described in
Dasyurus. In this Marsupial the suprarenal vein entered the
postcaval independently of and anterior to the renal vein on the
left side of the body. The suprarenal body received two afiluents
from the parietes. The first of these was a vein formed by the
union of two branches which entered the gland anteriorly, and
evidently is to be compared to the vein described above in
Dasyurus maugei. The second vein passes by the anterior end
of the left kidney and enters the suprarenal body at about the
middle of its length on the left side. In the Common Phalanger
(Trichosurus vulpecula) the same evidence of a suprarenal portal
system was obvious. In this case also the anterior suprarenal
portal vein was present, and no other. The suprarenal vein
entered the left kidney vein. JMJacropus dorsalis had but one
suprarenal portal, the anterior vein, which is apparently so constant
among the Marsupials and which arose from two affuents. In
the features described in the present communication, Petrogale
penicillata is exactly like Trichosurus vulpecula. Phascologale
penicillata is slightly different from any of the Marsupials as yet
described. The suprarenal vein opens, as is so frequently the case,
into the renal vein on the left side of the body. Exactly opposite
to it the suprarenal portal opens into the suprarenal body. But
this vein before entering the gland receives a branch running
transversely and just skirting the anterior margin of the left
kidney, the conditions being therefore slightly like those of
Peragale just described, and indeed intermediate between the
condition observable in that Marsupial and in those where the
1908. ] ANTECHINOMYS AND OTHER MARSUPIALS. 603
one vein bifurcates anteriorly. ettongia ogilbyi is like many
other Marsupials; the suprarenal portal is single and anterior in
position, being formed by the union of two afiluents which join at
an acute angle. As Thylacinus is a scarce type and not likely to
be much dissected in the future, I venture to give a particular
account of the veins connected with the suprarenal body, which
I noted during the dissection of a specimen which died in the
Society's Gardens in January 1906. The suprarenal vein enters
the postcaval (as is shown in text-figure 124, A) a little anteriorly
to the entrance of the left renal vein. It emerges from the
suprarenal body some little way in front of the posterior border
of the gland, The suprarenal portal vein enters the gland on
the right side a little way behind the anterior border. It is
formed of three afiluents. The middle one arises from the dia-
phragm and joms a branch arising from the parietal musculature
to the right of the suprarenal body. Just before entering
the suprarenal body the trunk formed by the union of these
two vessels is reinforced by a vein arising to the left of the supra-
renal body. The common trunk is thinner than the suprarenal
vein.
I have examined a number of Mammals belonging to orders
other than the Marsupialia, but have not found anything at all
resembling this apparently characteristic Marsupial feature in the
blood-supply of the suprarenal bodies. I believe myself at present
justified in asserting that this character, whether or not it be held
to be a persistence of a condition to be met with among Reptiles
and other lower Vertebrata, is distinctive 6f the Marsupialia.
V. Resumé.
I extract from the foregoing pages the principal new facts
which I have been able to add to our knowledge of the intestinal
tract of mammals and to certain features in the anatomy of
the Marsupialia.
(1) The most important features in the visceral anatomy of
Antechinomys are: the intestine borne upon a _ continuous
mesentery, the absence of a Spigelian lobe in the liver, the wide
dilatation of the uteri at their junction with the Fallopian tube,
the development of a short unpaired cecal chamber at the junction
of the uteri.
(2) A specimen of Phascologale macdonellensis showed a
persistent umbilical membrane (proving an umbilical placentation
in this species), which passes between the fibres of the rectus
muscle divided for its passage, and is continuous with the great
and splenic omentum. The umbilical membrane is also attached
to smail intestine. The intestinal canal is short and carried
on a continuous mesentery. The liver in this species, as in
P. penicillata, has a Spigelian lobe, also present in the genus
Sminthopsis.
(3) In many (? in all) Marsupials the suprarenal bodies receive
604 ON THE ANATOMY OF ANTECHINOMYS ETC. [May 26,
a vein from the parietes as well as emit one to the renal vein or
postcaval as the case may be; there is thus a rudiment of a supra-
renal portal system in these animals, not found in at least many
Eutherian Mammals.
(4) Though the intestinal tract of Marsupials is on the whole
simple, there are traces (Didelphys, Trichosurus) of the anse
coli and (Vrichoswrus) of the colico-duodenal ligament of more
differentiated forms.
5) A gut suspended upon a continuous mesentery is described
for the first time not only in Antechinomys, but in Tamandua ;
on the other hand, a number of genera of Carnivora
are described and the alleged continuous mesentery in Ursus
is shown to be only apparent and due to the reduction of the
ligamentum cavoduodenale. The continuous mesentery of Centetes
is shown to be not universal in the species and is therefore
probably to be looked upon as a reversion.
(6) To the numerous descriptions and figures of Rodents’
alimentary tracts gathered together or published for the first time
by Tullberg, a description of the colon and anse coli of Otomys,
Aulacodus, and some other forms is added. The enormously long
ansa coli dextira of the latter shows that the spiral found in certain
Rodents is not necessarily to be looked upon as due to the need
for packing away such a long loop. The spiral of Hydrocherus is
shown to bea late development since it does not occur in half-
grown examples. The colon of the minute Arvicanthis (with one
ansa only, the a. paraceecalis) shows that in this group reduction
of size is not necessarily accompanied by entire simplification of
the gut.
(7) The older descriptions of the spiral coil in certain Lemurs,
e.g. Vycticebus, are shown to be correct as against more recent
statements. Jicrocebus is shown to possess a simple colon without -
anse. Galago (2 spp.) is Shown to possess a spiral like Vycticebus.
&e. And it is pointed out that all the forms with a specialised
gut, i.e. with this spiral, are also specialised in the loss of the
elsewhere characteristic carpal vibrisse.
(8) Some account is given of the alimentary tract of the little
known species Theropithecus gelada and Semnopithecus melalophus
and the American Chrysothria sciureus.
(9) The intestinal tract of Hyrax, contrary to some statements,
has been shown to possess an ansa paracecalis which may perhaps
be compared to that of the Perissodactyla, and to possess the
ligamentum colico-duodenale of more differentiated forms.
(10) As a very general rule the loops of the small intestine
are loose folds not in any way fixed. Rarely, however (e.. g.
Dasypus vellerosus), | have found them to be fixed.
(11) That the colic loops vary is shown by the instance of
Lagostomus trichodactylus, in which each of the three individuals
dissected by myself or Tullberg is slightly different im the
proportions of those loops, and by Hyrax capensis.
(12) It has been pointed out that in man the omentum is at
1908. ] ON THE ARMOUR OF EXTINCT REPTILES. 605
first attached to the right side of the transverse colon and
subsequently to the left side, the intermediate space being filled
up later. The two earlier stages are represented in lower
mammals ; in 7richosurus the omentum is attached to the colon
only on the extreme right of the transverse bend, and in
Orycteropus and Hyrax the attachment is double, to the early
part of the colon and to a more distal region—the intervening
tract being free of the omentum.
(13) The view, deducible from previous investigations, that
four stages of advancing complexity are shown in the Mammalian
gut, is strengthened by fresh facts ; the Lemurs are shown to be
the only group in which all but one of these four stages occur.
4. The Armour of the Extinct Reptiles of the Genus
marcacounus. by CieG. SEELEY, F.R.S., EZ.8)
King’s College, London.
[Received April 29, 1908.
Text-figures 125-129.
g
In “ Further Observations on Pareiasaurus,” Phil. Trans. B.
Royal Society, 1892, I gave a short account of the dermal armour,
pp. 345-6. It is limited to the dorsal region, and is figured in
plate 17, and indicated by the letters ds in the description of the
plate, p. 368. The scutes are only known in this example of
Pareiasaurus baini, extracted from the rock by myself. They
were originally covered with matrix. Their existence was not
suspected, and it is possible that the more anterior scutes may
have been partly lost in removing the intractable rock; and those
seen in the British Museum specimen were preserved by great
skill m chiselting. The ossifications are flat and inconspicuous,
‘except where the lateral plates overhang the neural spines.
In the small figure of the skeleton given in the ‘Story of the
Karth,’ 1895, text-fig. 18, p. 126, the scutes were made more evident
by dark outlimes. Each scute is about 2 inches wide by 1? inch
long. There is a median row extending down the back, which as
preserved now rests upon the summits of the neural spines of the
dorsal vertebre and the interspaces between them. There are
also two lateral rows, one of which flanks each side of the median
row. These are arranged symmetrically im pairs, and extend
transversely outward from their contact with the median row,
but alternate with them by being placed at the junction between
each two median scutes. The lateral scutes in Pareiasawrus baini
are not flat but convexly curved as they extend outward, giving
some support to the idea that this armour formed an elevated
ridge on the back. In the present condition of the specimen
this armour is only seen on seven consecutive later dorsal vertebrae
and one or two earlier dorsals; and there is no evidence that
it was present over more than twelve vertebre.
[May 26,
PROF. H. G. SEBLEY ON THE ARMOUR
606
Dorsal armour of Pareiasaurus steenkampensis ; the scutes are arranged
as in P. baini.
1908. } OF A GENUS OF EXTINCT REPTILES. 607
Therefore the restorations which show elevated scutes extending
from the skull to the extremity of the tail, or three parallel rows
of scutes on the back entirely separated, and those which show
the body clustered over with rows or groups of scutes, are entirely
imaginary, for the only evidence for the armour is the skeleton
in the British Museum.
Some writers in this country, and in Germany, have denied
that any armour at all is present. The British Museum skeleton
is sufficient evidence of its characteristics. If it had been more
extensively developed over the body it is improbable that it would
have escaped detection in the careful removal of the matrix
during the two years that I watched the development of the
skeleton; and there is no reason to modify in any way the
original description or figure.
That evidence may now be added to by a short account of
specimens of scutes already referred to (l.c. pp. 315, 346) as
collected by Mr. J. van Renen, R.N., at Steenkamps Poort, south
of Fraserberg. I had just collected the Pareiasaurus and was
passing north, when this gentleman showed me a series of badly
preserved bones collected as weathered, and invited me to select
any example which might be necessary. I had no doubt they
were Pareiasaurian, though the essential characteristic parts of
the skeleton were not preserved. I accepted one caudal vertebra,
and a series of nine scutes as giving evidence of armour, which
I had not seen at that time.
The scutes are free from matrix, vary greatly in size, and
belong to a different species from P. baini, which I propose to
indicate on the evidence of these scanty materials as Pareiasaurus
steenkampensis. The scutes can only be supposed to have been
arranged as in P. baini; that is, in a single longitudinal row
down the back, with lateral scutes directed transversely outward
on each side from the union between each two successive scutes of
the linear series. All the ossifications are irregular, and about
half are broken (text-fig. 125). It is possible that all of those
preserved belong to the median series only, for none show the
eurved convex forms of the lateral scutes of P. bain, and this
difference may be a specific character. Four or five can be recog-
nised as median by their elongated forms; and the remainder
may be median or lateral, if lateral scutes were present, as I think
the evidence of the surface characters indicates. They are smooth
on the under side, marked on the upper surface with a central conical
blunt boss, from which numerous short grooves radiate irregularly
to the margin, which is commonly thick and rough, as though
the plates were imbedded in the skin. Behind the central boss,
which is more or less flattened above, and less than half an inch
in diameter, is a distinct pit nearly as wide, which is seen in half
a dozen examples. The radiating ridges are more or less pitted,
and all the surfaces, superior and inferior, are pierced with fine
vascular markings. The largest plates are about 2+ inches long
by 14 inch wide, and fully half an inch thick at the central boss,
In form they are irregularly ovate; some appear to be trans-
versely ovate and have the central boss less conspicuous.
Proc. Zoou. Soc.—1908, No, XX XIX, 39
608 PROF, H. G. SEELEY ON THE ARMOUR [May 26,
The smallest is 1,45 inch long, 1 inch wide, and half an inch thick.
Its inferior surface is slightly convex, and the external margin is
a sharp edge. The central part of the plate being occupied by
the boss, the radiating ornament is very short and is a marginal
fimbriation.
The second plate, slightly broken in front, is 2} inches long as
preserved, and just over an inch wide in front of the boss, but is
narrower posteriorly, though the lateral margins are weathered.
The boss, 7°; inch in diameter, is shield-shaped, its hinder border
being concave, with the excavation of the pit behind it. The
substance of the plate is fully + inch thick, both in front and
behind the boss. The radiating ornament is chiefly seen
anteriorly, and is irregularly pitted and corrugated. The con-
vexity of the base made the lateral margin sharp, but the edge is
almost removed by weathering.
The third plate is an elongated irregular pentagon with the
base in front, about as long as the second plate, but wider. It is
an inch and a half wide as preserved, but the margin appears to
be worn. The boss is somewhat smaller but not less elevated,
and the excavation of the pit behind it gives the aspect of a
posterior position. The radiating ornament is similarly irregular,
and like that on the second plate; but the base also develops
in a less degree some short-ribs, especially towards ‘the hinder
margin.
Text-fig. 126,
i Ds
=< il
Dorsal scute of Pareiasawrus steenkampensis.
The fourth and fifth median plates are both imperfect. The
central boss is rather less elevated, but the posterior pit continues
to be a marked characteristic.
Of the remaining plates, three are wider than long on the
hypothesis that the pit below the central boss is always posterior
in position ; and these ossifications are regarded as being placed
laterally. They are rather large plates, like the lateral plates in
1908. ] OF A GENUS OF EXTINCT REPTILES. 609
P. bain, and may have been in lateral contact with each other (text-
fig. 126). The best preserved is thick at the margin, concave on
Text-fig. 127.
Anterior aspect.
Text-fig. 128. Textfic. 129. / \
Posterior aspect. Lateral aspect.
Anterior, posterior, and lateral views of caudal vertebra of
areiasaurus steenkampensis.
the under side. They may have been inclined obliquely backward.
Their external surfaces have the same type or ornament as the
39*
610 ON THE ARMOUR OF EXTINCT REPTILES. [May 26,
median plates. The last specimen is.a fragment about 2 inches
wide and half an inch thick at the margin, much thinner at the
fracture, and without indication of boss or pit, so that its position
cannot be located.
We can only regret the imperfections of this evidence of
dermal armour, but when I saw the remains they had already
been removed for some time from the rock, and it is certain that
they would have been carried away by the torrential drainage in
the wet season, but for the interest taken by Mr. Van Renen in
their preservation.
This armature differs from that of Pareiasaurus baini, first
in the elongated form of the median scutes, secondly in the
presence of the central truncated boss with the depression behind
it, and thirdly in the radiated ornament—features which are
absent from Pareiasaurus baini. In that species the anterior
median scutes appear to be subcircular, or subquadrate, with a
few circular vascular openings or small pits; but in the later
plates no ornament is recognised, and the plates appear to be
thinner and arched outward.
The caudal vertebra is from a position between the fifth and
tenth in the tail. This early position is indicated by the trans-
verse width of the anterior face of the centrum exceeding its
vertical depth (text-fig. 127); by the strong vertically compressed
lateral ridges above the transverse processes for the caudal ribs,
which are directed outward and slightly downward; by the large
size of the transverse posterior facet for the chevron-bone, which
gives the centrum the aspect of being obliquely crushed from front
to back (text-fig. 129); and by the vertical position of the pre-
zygapophyses, with the facets looking inward and upward (text-
fig. 127).
When this centrum is compared with the earlier tail-vertebree
of Pareiasaurus baini the centrum is shorter from front to back,
for it only measures one inch; and in the species referred to, the
measurement is always longer when the anterior face of the
centrum is wider than deep. The neural canal is smaller (text-
fig. 128), as in later caudals of Pareiasaurus baini. These dif-
ferences would indicate a shorter tail with less lateral movement.
The neural arch is not distinctive. The neural spine is broken
away, but its base has the usual triangular form. The anterior
articular face of the centrum is roughly hexagonal with the
margin slightly rounded, and a moderate central concavity. It is
1,8, inch wide and 13 inch deep. The posterior face is rather
smaller and rather more concave. It is roughly four-sided, with
the lateral margins approximating superiorly (text-fig. 128).
It is 1,4; inch deep, 1,5, wide above the chevron articulation, and
1 inch wide on the neural canal. The oblique surface for the
articulation of the chevron-bone measures half an inch from
front to back, and extends over the width of the vertebre which
it truncates.
1908.] ON THE SKULL OF THE FOSSIL REPTILE DIADEMODON. 611
5. Additional Evidence as to the Dentition and Structure of
the Skull in the South African Fossil Reptile Genus
Diademodon. By H.G. Srutey, F.R.S., F.Z.8., King’s
College, London.
[Received May 26, 1908. |
(Text-figure 130.)
The genus Diademodon was founded on the molar teeth and
imperfect middle portions of small skulls. Four species were
figured in Phil. Trans. Royal Society, 1894, B, pl. 89, referred to
D. tetragonus, D. brachytiara, D. mastacus, and D. brown. They
were the most remarkable evidences of dentition of mammalian
type in extinct reptiles which have been found in South Africa.
There would have been grounds, had the remains been mammalian,
for referring them to three genera; and in the description of
plate 89, figure 11 is described as the left maxillary region of
Diademodon (or Gomphognathus) mastacus. And in the original
description of D. browni (I. c. p. 1039) it is observed, “it is
probably the type of a distinct genus.” Later in the same year
the group Gomphodontia was defined as comprising animals with
a Theriodont type of dentition, in which the molar teeth are
expanded transversely, and as having more or less tuberculate
crowns, of the type shown in Diademodon. In that group the
genus Diademodon was included (J. c. 1895, B, p. 3). The types
of Gomphognathus had the crowns of the molar teeth well worn,
but the elevation of the external cusps or ridge in G'. polyphagus
made a suggestive resemblance to Diademodon mastacus ; while
the condition of the single well-preserved crown in Diademodon
browni makes an equally suggestive approximation to Diademodon
brachytiara. In 1896, in a short communication to the British
Association at Liverpool, I briefly noticed another skull discovered
by Dr. D. R. Kannemeyer. I have removed the matrix in the
laboratory of King’s College, so as to demonstrate the sutures in
the middle part of the skull and to expose the palate. The
specimen is slightly squeezed so as to have a lateral obliquity
towards the right side, from which the similar example of
Diademodon browni is not free. There is a coincidence in the
anterior and posterior fractures being in identical positions in both
specimens, favouring comparison. ‘They are closely related species,
but the snout in the new example is narrower and rather smaller,
and the dentition being unworn favours the idea of specific
difference, though the forms of the transversely ovate sections of
the molar and premolar teeth are almost identical.
As preserved the specimen is 22 inches long. It extends
between an anterior transverse fracture through the two concave
pits on the snout, which lie at the junction of the maxillary and
nasal bones, which in Gomphognathus are situated midway between
the orbits of the eyes and anterior nares, and a posterior fracture
612 PROF. H. G. SEELEY ON THE SKULL [May 26,
behind the orbits, just behind the post-frontal bones, which are
imperfectly preserved. The lateral margins converge forward in
a wedge-like outline similar to the corresponding part of the
skull of Gomphognathus. Owing to the lateral compression the
anterior transverse measurement is narrowed by one or two tenths
of an inch. As preserved it is one inch wide, and the vertical
height from the median longitudinal ridge on the palate to the
nasal bones is the same. The corresponding measurements in
Diademodon browni are: vertical 15, transverse 1;4 inch. The
pre-orbital lateral areas of this, formed chiefly by the maxillary
bones, are inclined towards each other, are gently convex from the
alveolar border to the nasal region, longitudinally furrowed by two
shallow concavities on each side, and then round with a gentle
convexity into the upper surface formed by the nasal bones.
Diademodon brownt distinctly suggests an angle between the
sides of the face, which are more vertical, and the roof of the
snout; but the difference between the specimens is one of degree.
In both there is some lateral concavity of the pre-orbital region
from front to back.
The head widens backward to the posterior fracture, which
passes through the back of the frontal bone, the post-frontal
bones, and the pterygoid bones. Thespecimen is about 2,3, inches
wide behind the dentary tract, which is in a line with the middle
of the orbits, and 1,4, inch high at the back of the frontal
bones. :
The bones seen on the superior aspect of the skull are the
frontal, post-frontal, pre-frontal, nasal, lachrymal, and maxillary.
The naso-maxillary region is convex from side to side, but as the
nasal bones extend between the orbits their flattened upper
surface merges in the flattened frontal region. The orbits are
inclined so to look outward, and to a less extent upward and
forward. The vertical measurement of the rounded cavity is one
inch, and the transverse measurement between them over the pre-
frontal bones is 1,2, inch. In D. browni it appears to have been
144 inch.
Only the part of the post-frontal bone which is above the back
of the orbit is preserved. It is rather less than half an inch wide
between the temporal vacuity and the suture with the pre-frontal
bone. It is transversely channelled owing to elevation of its front
and back borders. These bones are slightly raised above the
frontal bones, which extend longitudinally between them. As
preserved the frontal bones are oblong, 38; inch long, and more
than half an inch wide towards the middle at the suture between
the post-frontal and pre-frontal bones, and narrow anteriorly to
the transverse suture with the nasal bones. Both bones are
longitudinally concave with the median sutural line raised. This
feature is absent in Diademodon browni. The lateral sutures with
the post-frontal and pre-frontal bones are similarly raised as slight
ridges, but there is no ridge between the frontal and nasal bones.
The pre-frontal bone forms much of the superior border of the
1908. ] OF THE FOSSIL REPTILE DIADEMODON. 613
orbit where the margin is compressed posteriorly and rounded in
front. The bone is in front of the post-frontal, external to the
frontal, makes an oblique suture with the nasal, and a narrow
junction with the lachrymal bone, as its sutural junctions diverge
outward and forward. It is ? inch long from the post-frontal to
the lachrymal and 3, inch wide at the fronto-nasal suture, where
it 1s widest in about its middle length. The inner short border
next the frontal is parallel to the longer external border above the
orbit and lachrymal. The pre-frontal bone forms a large part of
the internal anterior wall of the orbit.
The nasal bones, somewhat lanceolate in form, are imperfect
anteriorly. They extend from the frontal bones forward as
preserved to between the pair of pits on the front of the snout,
which are not seen in Diademodon browni, with a length of
1,5 inch, and in this length they are not in contact with the pre-
maxillary bones. They are separated from each other by a fine
straight suture, and widen from the frontal suture anteriorly, with
the lateral divergence of the sutures dividing them from the pre-
frontal and lachrymal bones, to 1} inch at the front of the
lachrymal bones ; and anteriorly the sutures between them and
the maxillary bones converge forward, to a transverse width over
the nasal bones of half an inch, at the anterior fracture through
the lateral-nasal pits. The bones are smooth, convex from side
to side, and slightly raised posteriorly, with a partial prolongation
forward of the median frontal sutural ridge.
The lachrymal bone is best exposed on the left side, where I
have partially removed the matrix from the orbit. It is at the
front of the orbit between the maxillary bone below and the nasal
and pre-frontal bones above. Externally it is of irregular sub-
quadrate form, half an inch in each measurement. It has a
considerable extension in the front of the orbit internally, below
the pre-frontal bone. On the lower part of the inner front border
the bone is pierced by two circular canals placed one below the
other.
The maxillary bones form the sides of the face from the hinder
fracture at the back of the alveolar tract below the orbit, where
the bone is ;4, inch deep, forward to the anterior fracture, where
the depth is 1,4, inch. The ascending orbital border below the
orbit is compressed, rounded, and slightly reflected outward.
Below the lachrymal canal the depth to the alveolar border is
‘inch. A slight wide shallow concavity extends longitudinally
forward, from the orbital junction between the lachrymal and
maxillary bones; but on the right side the bone appears to be
accidentally impressed in this region. The lower part of the
maxillary bone is moderately concave in length, and markedly
convex downward owing to the compression of the bone imme-
diately above the molar teeth. On the convex ridge above are
two ovate foramina above the teeth, such as occur in many fossil
reptiles.
The palate has shared in the side to side compression and
614 PROF. H. G. SEELEY ON THE SKULL [ May 26,
distortion of the specimen and is probably narrowed by a tenth
of an inch. The teeth extend in diverging curves as they range
backward ain outward. The transverse width over the pr emolats
in front is $4 7 inch; over the last molars it is about 235 inches.
These aeectiimen's are less than in Diademodon brown. Ten
teeth are indicated or preserved, of which the two in front, with
small circular fractured bases to the crowns, are classed as pre-
molars, and the eight succeeding teeth are molars. They have
the crowns transversely ovate, each with its axis at right angles.
with the concave external alveolar border, except the Tast tooth,
which is parallel to the alveolar border. The crowns increase in
width to the fifth molar and then become smaller, the seventh
and eighth rapidly narrowing acquire a triangular nor comma
shape. The length occupied by the eight molars is 142 inch.
The anterior ‘teeth are separated by the hard palate between
them. They rise with a vertical inner alveolar border corre-
sponding to ube compressed external border. As preserved the
hard palate is }} inch wide between the last premolar teeth, and
3 inch wide where it terminates between the fifth pair of molars.
It is narrower than in D. br ownt, in which the fourth molar
appears to be the largest. The hard palate as preserved is made
by the maxillary bones, which extend behind the second molar
teeth, and unite by a transverse suture with the palatine bones,
so that the suture is in about the same position as in Gompho-
gnathus polyphagus (Phil. Trans. 1895, B, p. 16, fig. 7). (1 Tts
distinctive feature is a strong elevated median ridge dividing the
palate into two concave channels. This ridge is continued bool
ward by what I regard as the vomerine bone, dividing the
posterior nares, extending upon the median union of the posterior
palatine bones. This ridge on the hard palate is absent from
D. browni ; its presence makes the transverse hinder border of
each half of the hard palate concave, instead of both bones
combining to form one concave posterior surface. The back of the
palate behind the posterior nares has a close general resemblance:
to the corresponding region of Gomphognathus. ‘There are the
same pair of convex rounded tumid areas behind the hard palate
converging backward from the hinder cheek-teeth to terminate in
a pair of hemispherical convexities which were just in front of
the median post-palatal ridge in that genus, flanked externally by
the broken bases of the pair of transverse processes which
descended between the rami of the mandible (compare J. c. p. 24,
fig. 11). The transverse width over these processes in this speci-
men is 1,4 inch. Those processes are regarded as being made:
chiefly by the transverse bones and as defined by sutures which
converge inward from behind the maxillary bones backward to the
hemispherical tubercles at the posterior fracture.
The teeth have been more or less broken, possibly by strain or
compression. Small parts of the enamelled surfaces of the tuber-
culate crowns remain in the first and second molars of the right:
side, The first shows a marginal external rim behind the crown
1908. ] OF THE FOSSIL REPTILE DIADEMODON. 615
and laterally, and a small central tubercle in front. The second
indicates two lateral external tubercles. None of the crowns show
the slightest trace of wear by the apposition of the mandibular
teeth, in this respect being in striking contrast to Gomphognathus,
in which the crowns of all the molar teeth are always worn so that
nothing remains of tuberculate structure except the external cusp.
The fifth, sixth, seventh, and eighth crowns are preserved on both
sides.
Text-fig. 130.
Middle molar tooth
crown, enlarged.
Restoration of the skull of Diademodon entomophonus. About 3.
The fifth and sixth crowns are transversely ovate, less than
7p Inch wide and 2 inch from front to back. They have a strong
616 ON THE SKULL OF THE FOSSIL REPTILE DIADEMODON. | May 26,
external crenulate border and a median crenulate transverse ridge,
dividing the concave posterior half of the crown,which has acrenulate
external margin, into larger external and smaller internal concave
spaces (see text-fig. 130). In front there is a sharp or crenulate
marginal border, with transverse crenulations or cusps; on the
middle of the crown two small anterior cusps and two posterior
cusps. These crowns are essentially of the type of the described
species of Diademodon. The last tooth of D. mastacus shows a ten-
dency to develop a posterior talon (J. c. 1894, B, pl. 89. figs. 11, 12).
The penultimate tooth of this specimen has the posterior talon so
developed as to make the form of the crown almost triangular.
The crown is only a quarter of an inch wide, and slightly shorter
from front to back externally. The strong external anterior cusp
is broken, but a small external cusp rises from the talon. On the
inner border of the crown are two or three cusps or crenulations
like those similarly placed on the fifth and sixth molars. The last
molar is compressed from side to side, ;2, inch long by 345 inch
wide, broader in front than behind, with small tubercles back and
front. The small size of these teeth gives the molars the aspect
of exceptional divergence posteriorly. The transverse internal
measurement between the last pair of molar teeth is 1,5, inch ;
between the fifth pair it is ? inch, and between the first pair of
molars about 4} inch. From front to back the crowns form a
convex curve.
The dentition is imperfectly preserved, but not more than two
or three premolar teeth appear to be lost. From the resemblances
of the skull to allied types I infer that there was a toothless
diastema between the first premolar and the canine, where the
jaw contracted from side to side. J should expect four incisors
as in Gomphognathus. The missing extremity of the snout would
be about 1,8, inch long; the missing hinder part of the head was
about 24 inches long, giving the complete skull a length of
64 inches. The skull may be restored on the type of Gompho-
gnathus (text-fig. 130).
The most remarkable feature of the dentition is the unworn
condition of the crowns of the teeth, also seen in other species of
the same genus. The transversely ovate forms of the molar
crowns acquire new interest from the teeth of Procolophon
having this form, with inner and outer cusps recalling the tooth
ot Diademodon browni. But while Procolophon is typically
reptilian in its dental armature (Proc. Zool. Soc. 1905, vol. 1.
p- 225), in this fossil the teeth suggest mammalia. The trans-
versely ovate form of the crown, with the sight cingulum,
approaches the condition in lemurs, but the molars are more
numerous and the other dental characters unlike. The diastema
occurs among mammals as various as marsupials, tapirs, rodents,
but is never associated with a transversely ovate molar, and full
series of incisor and strong canine teeth as in these fossil reptilian
types. Mammals of various groups have the molar teeth
progressively increasing and afterwards decreasing in size, as
P ZS§ 1908, Pl XXVIII,
6.
Lith Anstv. K Wesser dena.
K.dordan del.
New Siphonaptera.
K. Jordan del.
New Siphonaptera.
PZ.S. 1908, Pl. XXX.
Tath Anst v.KWesser dena.
" aS = :
ae en Shee oe
Vist.
K.dordan del,
New Siphonaptera.
PZ.S. 1908, Pl. ®
Lith Anst vKWesser
K. Jordan del.
New Siphonaptera.
1908. | ON NEW SPECIES OF FLEAS. 617
among lemurs, insectivora, carnivora. But the feature of this
reptilian type is its generalised mammalian resemblances in dental
characteristics, which are highly specialised distinctions among
mammals, so that the teeth have undergone an evolution of
mammalian type. It is not to be anticipated that a complete
skeleton of Diademodon will make a closer approximation to that
of a mammal than is already evidenced by other Theriodont
reptiles; but the dental characters emphasise the mammalian
approximations which have been found in the shoulder-girdle,
pelvic arch, and limb-bones.
I propose to distinguish this species, characterised by the median
ridge on the palate, the ovate unworn multituberculate crowns of
the middle molars, the moderate interspace between the orbits,
and slender snout rounded above, as Diademodon entomophonus.
The absence of wear to the crowns is only consistent with a diet
which did not involve trituration. It is in contrast with the con-
dition in Diademodon browni, which it approximates in general
characters.
6. New Siphonaptera.
By the Hon. N. Cuarzes Roruscuixp, M.A., F.Z.8.
[Received May 1, 1908. |
(Plates XX VIIT.—XXXT. *)
Genus PyYGIoPsyLia.
Pygiopsylla Rothschild, Ent. Mon. Mag. (2) xvii. p. 221 (1906)
(type: hilli).
The species belonging to this genus are easily distinguished
from Ceratophyllus Curtis (type of name: hirundinis) by the
sensory plate (so-called pygidium) of the ninth abdominal tergite
being strongly convex (ef. Pl. XXX. fig. 14). All the species
are very hairy. They are inhabitants of the Old World, being
known both from the Oriental and AXthiopian Regions. Nine
species are known, namely: Ailli Rothschild (1904), nove-
guinee Rothschild (1904), robinsont Rothschild (1905), colossus
Rothschild (1906), echidne Denny (1843), ahale Rothschild (1904),
torvus Rothschild (1908), woodwardi Rothschild (1904), and
rothschildi Rainb. (1905), the last two appearing to me to be but
doubtfully distinct from each other. In the present paper six
more species are described—-namely, two from Australia, two from
New Guinea, one from Ceylon, and one from West Africa
(Angola),—making in all 15 species of this genus. The wide
distribution of the genus renders it probable that these fifteen
forms are only a small percentage of the actually existing species
of Pygiopsylla.
* For explanation of the Plates, see p. 629.
618 THE HON. N. C. ROTHSCHILD ON [May 26,
P. echidne is a more specialised species than the others. The
genal edge of the head is produced into a broad tooth-like lobe,
corresponding to the lobe found in the Sarcopsyllide and in
Pariodontis Rothschild (1908) and Lycopsylla Rothschild (1904),
and the comb of the pronotum is reduced to a few spines (4 to 6),
which ave dorsal. The head of P. echidne, moreover, is short,
bearing two regular rows of three bristles each on the frons, there
being no row of small bristles between the anterior corner of the
frons and the base of the antennal groove, as is the case in all
the other species of Pygiopsylla (cf. Pl. X XIX. fig. 7). It may
possibly become necessary to move echidne from Pygiopsylla and
place it in a new genus. For the present, however, there is no
necessity for this change.
1. PYGIOPSYLLA AFER, sp.n. (Plate X XIX. figs. 7, 8.)
Head.—The head (Pl. XXIX. fig. 7) is gradually rounded,
the lower part of the frons not being curved backwards (7. e.
towards the fore cox). The rostrum does not quite reach the
apex of the fore coxa.
Thorax.—The pronotum bears a comb of 23 spines and two
rows of bristles. On the mesonotum there are 4 rows of bristles,
the anterior row being abbreviated and there being also some
dorsal bristles in front of this row. The mesopleura have 8 long
bristles and a few short ones. The metanotum bears 4 rows of
bristles, the first row consisting of but a few bristles situated on
the back. The epimerum of the mesothorax bears an irregular
anterior row of 7 or 8 bristles, a central row of 3, and a posterior
row of 3 or 4 long and some short bristles.
Abdomen.—The first tergite is practically hairy all over. The
other tergites bear 4 rows of bristles, the first row being repre-
sented only by a few bristles on the fifth to seventh segments.
There is a stout apical spine on each side of tergites 2 to 5. The
basal sternite bears an oblique patch of short bristles on the side,
consisting of 3 irregular rows. On the sternites of the third to
seventh segments there is a subapical row of 4 or 5 long bristles,
proximally to which are numerous small bristles.
Legs.—The mid and hind femora bear on the outer side three
subapical ventral hairs, which are of nearly equal size, there being
no other ventral hairs between these three and the widest point
of the femora. The hind tibia bears about 20 bristles on the
outside, arranged in three irregular rows, besides a number of
smaller bristles situated at and near the anterior edge of the tibia.
The first fore-tarsal segment is longer than the second and has
four thin and long bristles on the hinder side. The first
mid-tarsal segment is much longer than the second. The hind
tarsus 1s long, especially the first and second segments, the third
segment being longer than the fifth. The first and third pairs of
lateral bristles of the fifth segment are moved towards the mesial
line, especially in the fore and mid tarsi.
1908. ] NEW SPECIES OF FLEAS. 619
Modified segments.—The seventh abdominal sternite has a deep
triangular sinus (Pl. X XIX. fig. 8), the upper loke being broad, but
tapering to a point. The bristles on the eighth tergite are more
numerous than in P. robinsoni, to which the present species is
allied. At the apical margin of this segment, there is one long
bristle, and aboveit are situated two short ones and beneath it one
moderately long one. There is a row of 5 or 6 long bristles
along the ventral edge, the most distal bristle being the longest
of all. Above this bristle there are two more long ones, and
further proximad about 16 short ones. The anal sternite is
rounded beneath near the base, bearing on this rounded portion a
row of 4 bristles on each side, there being a further pair of bristles
on each side close to the apex.
Length: 3°6 mm.
One 2? from Benguella, Angola, 200 miles from the coast,
found at an altitude of 4780 ft. by Dr. F. Creighton Wellman
the host not being stated; received from Mr. Oldfield
Thomas, F.R.S.
2. PYGIOPSYLLA RAINBOWI, sp. n. (Plate XXVIII. fig. 5;
Plate XXX. fig. 13.)
The present species apparently agrees in all details, except the
modified abdominal segments, with P. colossus Rothschild 1906,
of which only one 2 is known.
3. The small eighth tergite bears about 8 short bristles above
the stigma. The eighth sternite (Pl. XXX. fig. 13), on the other
hand, is very large, being covered with numerous bristles, of which
those placed near the dorsal and apical edges are longest and thickest.
The ventral margin of this segment (in lateral view) is incurved
twice, the segment being incised in the mesial line from the apex
to the point where the ventral margin bulges out. The clasper (Cl.)
is distaily produced into a thumb-like process, which is shorter
than the pointed and slightly curved movable process (F). The
manubrium (M) is triangular, ending in a short process. The
vertical arm of the ninth sternite (IX. st.) is club-shaped, and at
the apex truncate, with the distal margin of the widened portion
rounded. The horizontal arm is of nearly even width, its upper
margin being twice incurved. This arm bears numerous small
hairs on the apical as well as proximal portions, there being in
addition on each side a row of five ventral bristles, of which the
most proximal one is the longest and thickest. The penis ends
in a short and sharp hook, which points downwards. The anal
tergite (X. t.) is triangular in side-view, being about twice as long
as it is broad at the base. The tenth sternite is much slenderer
than the tergite, bearing two long apical bristles on each side.—
Q@. The apical margin of the seventh abdominal sternite is
rounded, being ventrally obliquely truncate and bearing a small
sinus in the centre (Pl. XXVIII. fig.5). The eighth sternite has
fewer bristles than in P. colossus at and near the apical and
620 THE HON. N. C. ROTHSCHILD ON [May 26,
ventral margins. The ninth and tenth segments resemble those
of P. colossus.
Length: ¢ 3°3 mm., 2 5 mm.
We have a long series off Mus assimilis from Emerald, Victoria,
Australia, collected by Mr. Edw. Jarvis during 1907.
3. PyGIOPSYLLA GRAVIS, sp.n. (Plate XXX. fig. 14.)
This species closely resembles P. rainbowi, except in the
genitalia. We have only one ¢.
3g. The eighth abdominal sternite (Pl. XXX. fig. 14) is very
large, as ib is in rainbowi, and is densely covered with bristles as
in that species. But the long bristles which are placed along the
apical and dorsal edges of the segment are more numerous and
more slender than in rainbowi. The upper margin of the eighth
sternite is gradually rounded, the ventral margin being straight.
The clasper (Pl. XXX. fig. 14, Cl.) is distally produced into a
finger-like process ( P), which bears a row of thin hairs at the
dorsal margin. The movable flap (F) is very large, being leaf-
shaped, with the pointed tip curved upwards. The manubrium
(M) is curved dorsad, the apical portion being somewhat twisted.
The ninth sternite (Pl. XXX. fig. 14, LX. st.) is very broad. The
horizontal arm bears ventrally at the apex on each side five long
stout spines, of which the most distal one is the longest. The
anal segment (= tenth) is long and slender, the anal sternite
bearing a pair of very long bristles at the apex.
Length: ¢ 4 mm.
We have one ¢ from Emerald, Victoria, off Mus assimilis,
collected on 18th September, 1906, by Mr. Edw. Jarvis.
4, PyG@topsYLLA LACINIosUS, sp.n. (Plate XXIX. fig. 10.)
@. As large as P. rainbow.
Thorax.—The pronotum bears two rows of bristles and a comb
of 19 spines. The pleura of the mesothorax have 7 or 8 bristles,
while the mesonotum bears four rows of bristles and some
additional ones in front of these rows on the back. The
yaetanotum has likewise four rows of bristles, but the first row
contains on each side only about 5 bristles, and there are dorsally
fewer hairs in front of this row than on the mesonotum. The
epimerum of the metathorax has four irregular rows of bristles
(5, 5, 2 or 3, 3), the bristles of the posterior row being the longest.
There are also one or two additional short bristles in front of the
posterior row.
Abdomen.—The first tergite is hairy all over. The other
tergites bear fewer bristles than in P. rainbowi and colossus, the
second and third having four rows and some additional dorsal
bristles, while the sixth and seventh tergites bear three rows and
a few bristles representing a fourth row; the basal sternite has
no bristles on the sides, apart from a few extremely small hairs.
1908. ] NEW SPECIES OF FLEAS. 621
The short bristles on the sternites of the third to sixth segments
are less numerous than in the allied species just mentioned.
Legs.—As in colossus.
Modified segments.— 2 . The seventhsternite (Pl. X XIX. fig. 10)
is bisinuate, closely resembling that segment of P. colossus, but
differing in the lower lobe and the lower sinus being much wider,
in the upper sinus being smaller, and in the bristles being
differently arranged. i
Length : 5 mm.
We have 3 9 from Mt. Albert Edward, British New Guinea,
off Mus mordax ; received from Mr. E. C. Chubb.
5. PYGIOPSYLLA MORDAX, sp.n. (Plate XXVIII. fig. 6; Plate
XXIX. fig. 9.)
Head.—The frons is strongly curved, as is the case in P. ahalee
Rothsch. (1904), the bristles being thick. The rostrum is shorter
than in all the other species, reaching only a little beyond the
middle of the fore coxa.
Thorax.—The pronotum is short. It bears two rows of bristles,
the anterior row being irregular and represented by but a few
dorsal hairs. The comb consists of 16 to 18 spines, which are
longer than the pronotum. The meso- and metanotum each
bears four rows of bristles, the first row not reaching so far
downward as the others. The mesothoracic pleura have about
twelve bristles, of which four or five anterior ones are short.
Abdomen.—The first tergite has four rows of bristles, the
second to seventh tergites three rows. The basal sternite bears
in the ¢ about 6 minute hairs on the side arranged in two oblique
rows. The sternites of the third to sixth segments of the ¢
have on each side a curved row of four subapical bristles and
proximately to this row six or eight smaller bristles. In the 9
the basal sternite has two irregular oblique rows of bristles on the
side, each row containing about ten bristles, the bristles being
more numerous also on the other sternites than in the ¢.
Legs.—The mid and hind femora bear three ventral subapical
bristles, the first being smaller than the others. The mid and
hind tibize have numerous bristles practically all over the outer
surface, the bristles being more numerous in the Q than in the
3. The first mid-tarsal segment is much longer than the second,
but is shorter than in P. ahale. The fifth hind-tarsal segment
is as long as the third.
Modified segments——g. The apex of the large eighth
abdominal sternite is irregularly rounded. There are four pairs
of long bristles below the upper edge of this sclerite, a single long
bristle below the most distal pair, and further down at the ventral
margin two or three more long bristles. Between these long
ventral bristles and the base of the segment there are about 12
shorter bristles, there being also three or four additional bristles on
the lateral surface. The clasper (Pl. XX VIII. fig. 6, Cl.) is distally
622 THE HON. N. C. ROTHSCHILD ON | May 26,
truncate-emarginate, the lower corner being somewhat produced
and bearing a long thin bristle accompanied by a small one. The
manubrium (M) is very broad, the apex being pointed and
curved dorsad. The movable process (F) is very long. It is
pointed and curved, its upper edge being twice incurved. There
is a large number of bristles at the ventral margin of this
process, the distal ones being long. The vertical arm of the ninth
tergite (Pl. XX VIII. fig. 6, IX. st.) is almost evenly curved. The
horizontal arm is shorter than the vertical one and bears two
strong spines ventrally at the apex, there being also a number of
thin bristles along the ventral margin and at the apex, as shown in
the figure.— 2. The seventh sternite is deeply sinuate, the upper
lobe being broad and the lower one narrow (Pl. X XIX. fig. 9).
The eighth tergite bears on each side three or four small. bristles
above the stigma. On the ventral portion of this sclerite there
are about twenty bristles, three or four placed at the apical margin
and five along the ventral edge. The lower apical angle of the
eighth tergite is produced. The anal sternite has on each side
two long bristles near the base, one in the centre and one near
the apex, besides two smaller apical ones.
Length: ¢ 2 2:1 mm.
We have one ¢ and two 2 from Mt. Albert Edward, British
New Guinea, off Mus mordax; received from Mr. E. C Chubb.
6. PYGIOPSYLLA FERINUS, sp.n. (Plate X XIX. fig. 11.)
Nearest to P. mordax.
Head.—The rostrum reaches nearly to the apex of the fore coxa.
The bristles of the anterior row of the frons are a little thicker
than in P. mordax.
Thorax.—The pronotum has one row of bristles and a comb of
17 spines: The meso- and metanota have three rows of bristles, a
fourth (anterior) row being represented by a few short dorsal
bristles only. The epimerum of the metathorax has ten bristles
(4, 3, 3), with some small hairs in between the posterior bristles.
Abdomen,—The tergites have three rows of bristles, the
anterior row being represented by but few bristles, except in the
case of the first segment, which bears about four additional
bristles on the two sides together, representing a more complete
fourth row. The bristles on the sternites are less numerous
than in P. mordacx.
Modified segments.— 2. The seventh sternite (Pl. XXIX.
fig. 11) is bisinuate, the upper sinus being smaller than the
lower. The eighth segment is similar to that of P. mordaa.
The anal sternite, however, is quite different. This sclerite has
beneath a prominent tubercle bearing a brush of long bristles,
Length: 2 3 mm.
We have one 2 from Pundaloya, Ceylon, taken off Sorex sp
by Mr. E. E. Green.
1908. | NEW SPECIES OF FLEAS. 623
7. STEPHANOCIRCUS JARVISI, sp.n. (Plate XXIX. fig. 12; Plate
XXXI_. fig. 16.)
Head.—The helmet is rounded, resembling that of S. simsoni
Roths. (1905), but being broader and bearing on each side 17 or
18 spines. The genal comb consists of 11 or 12 spines, which are
obtuse, like those of the helmet, not being pointed as in S. dasyurt
Skuse (1890). The occiput is shorter than in all the other species,
its bristles being thick. The mouth-parts are short, the maxillary
palpus as well as the rostrum being only twice the length of the
spines of the genal comb. The last segment of the rostrum is
broader than it is long. The maxilla is pointed.
Thorax.—The pronotum bears a comb of 30 to 40 spines and
two regular rows of thick bristles: The mesonotum has about
7 rows of bristles, the anterior bristles being small. On the pleura
of the mesothorax there are about 24 bristles, some being short.
The metanotum has three rows of bristles and in front of them a
few additional shorter hairs. The episternum of the metathorax
bears about 6 bristles, while the epimerum has two rows, the first
being irregular and containing 6 or 7 bristles and the second
containing 5.
Abdomen.—The abdominal tergites 1 to 7 bear each two rows
of bristles, the seventh tergite having 2 apical bristles in the
male, and 4 of nearly equal size in the female. On the first
tergite there is a comb of 27 spines in the 6, which are only a
little shorter than those of the pronotal comb, the comb of the 9
containing 34 spines; the second tergite has a comb of 17 shorter
spines in the ¢ and of 22 in the 9, the comb of the third tergite
consisting of 15 spines in both sexes. The fourth tergite bears
on each side 2 or 3 shorter and paler apical spines and the fifth
and sixth tergite one spine.
Legs.—Resembling most those of S. mars. The hind coxa is
longer than in that species. The hind femur bears posteriorly 3
subyentral bristles and between these and the dorsal edge several
more bristles, there being also one or two bristles near the base
on the outer surface. The tibie are very characteristic. In the
fore and mid tibie the outer bristles of the dorsal pairs are shifted
towards the lateral surface, forming a close-set row of thick and
equal-sized bristles. In the hind tibia these bristles are in their
normal position close to the long dorsal bristles. The dorsal
bristles are very long, the fifth being of the length of the tibia.
The hind tibia bears numerous bristles scattered over the outer
surface. The first mid-tarsal segment is twice the length of the
second. The first hind-tarsal segment is only one-sixth shorter
than the hind tibia, its longest apical bristle nearly reaching to
the tip of the second segment, which latter is twice the length ot
the fifth segment (claws excluded).
Modified segments.— ¢. The clasper (Pl. XXIX. fig. 12, Cl.) is
produced into a broad, leaf-shaped apical lobe, which bears three
large bristles placed on the lateral surface. A number of small
Proc. Zoou. Soc.—1908, No. XL. 40
624 THE HON. N. C. ROTHSCHILD ON [ May 26,
bristles are situated along the dorsal edge of this lobe, while
two fairly long ones are placed just beneath the pointed apex of
the lobe. The clasper, moreover, is ventrally widened into an
obtuse triangular lobe, which bears one slender bristle. The
finger (F) is asymmetrical, being leaf-shaped with the apex
curving upwards, bearing two moderately long bristles below the
apex and a few still shorter ones further proximad. The manu-
brium (M) curves upwards, the ventral margin bulging out in the
centre. The ninth sternite (Pl. X XIX. fig. 12, IX. st.) resembles
that of S. simsoni in general structure, but the inner arm is
different in outline, the horizontal arm is longer, and the number,
size, and position of the bristles are different. There are, as in
S. simsoni and dasyuri, two apical spines on each side of the
horizontal arm of the ninth sternite, as shown in the figure.
The anal tergite is very different from that of S. simsoni, bearing
a few thin bristles and on each side a pair of long apical ones.
The anal sternite is divided at the apex, each lobe bearing three
long bristles.— 9. The eighth tergite, which resembles in outline
that sclerite of S. simsonz, bears about 9 short but strong bristles
above the stigma, and from 29 to 34 bristles on the sides and at
the apex, as shown in the figure (Pl. XX XI. fig. 16). The eighth
sternite is elongate-triangular, ending in a long sharp point as
in S. simsoni. The tenth tergite is distinctly separated from the
ninth tergite, as is also the case in S. simsoni. Proximally to
this suture there is a transverse row of bristles on the ninth
tergite. The stylet is very long.
Length: ¢ 27 mm., 2 3:3 mm.
We have examined a pair of this species, the ¢ (the type)
from Emerald, Victoria, found under a rotten tree-trunk in the
virgin forest by Mr. Edw. Jarvis; the 9, from Victoria, off
Phascologale swainsonit, was forwarded to us by Mr. D.
McAlpine.
8. CrENOPHTHALMUS RETTIGI,sp.n. (Plate XXVIII. figs. 3, 4.)
Similar to C. agyrtes, but differing especially in the bristles of
the abdomen and in the modified abdominal segments.
Thorax.—The comb of the pronotum consists of 18 spines.
Abdomen.—The bristles are longer than in C’. agyrtes, especially
the three apical ones of the seventh tergite. The sternites of
segments three to seven in the ¢ have a transverse row of 4 or 5 ©
bristles and generally some small bristles in front of this row.
The basal sternite in the 9 has two or more bristles on the side;
the following four sternites have a row of 6 or 7 bristles and
from 6 to 10 smaller ones in front of the row; on the seventh
sternite the row contains about 10 long bristles.
Modified segments.— ¢ . The eighth tergite (Pl. XX VIII. fig. 3)
has about 6 small bristles above the stigma. The sternite becomes.
gradually narrower towards the apex, which is truncate ; it bears
on its lower portion 14 or 16 bristles (Pl. XXVIII. fig. 3). The
1908. } NEW SPECIES OF FLEAS. 625
clasper (Pl. XX VIIT. fig. 3, Cl.) is produced into a short square
process (P), of which the apical margin is feebly incurved. At the
upper corner of this process there are about half-a-dozen slender
bristles and one which is very stout and long, whereas at the
oblique ventral margin of the process there are two such long
bristles. The manubrium (M) gradually tapers to a point, being
somewhat curved upwards. The movable process (F) is widest
near the base. It is irregularly conical, the ventral margin being
somewhat incurved and proximally strongly rounded. There are
four short broad bristles near its apex at the dorsal edge and
three thin ones near the apex at the ventral margin, there being,
moreover, four thin bristles at this margin on the widest part
towards the base. The horizontal arm of the ninth sternite
(IX. st.) is much shorter than the verticalarm. Thereareabout
17 bristles at and near the ventral margin of this sternite from
before the middle to the apex, the four or five proximal bristles
being close together and longer than the sternite is broad.—
@. The apical margin of the seventh sternite is sinuate below the
centre, the upper lobe being very broad and nearly square
(Pl. XXVIII. fig. 4), while the lower lobe is small and obliquely
rounded. The eighth tergite bears several small bristles above
the stigma and about 16 bristles on the lower portion, there
being also ten or more small bristles near the apex on the inner
surface. The stylet is nearly three times as long as it is broad at
the base.
Length: ¢ 2:1 mm., 9 3:2 mm.
We have a large series collected by Mr. A. Rettig at Malcoci,
Roumania, off Mesocricetus newtoni, Putorius desertorum, and
Spalax typhlus.
9, CrENOPSYLLUS ALLOPHYLUS, sp.n. (Plate XX VIII. figs. 1,2.)
Head.—The frons (Pl. XXVIII. fig. 2) is strongly and evenly
rounded and bears a vertical comb of 6 spines. The first spine
is short and very broad, while the third is placed beneath the
second and fourth. There is anteriorly a row of 8 bristles,
followed by a second row of 3 longer ones; 2 more bristles are
situated in front of the comb, whilst a very long one is placed
about halfway between the insertion of the maxillary palpus and
the anterior row of bristles. The occiput bears 3 rows of bristles.
The rostrum is about one-third shorter than the fore coxa.
Thorax.—The pronotum bears one row of bristles and a comb
of 21 spines. The meso- and metanotum have each 3 rows of
bristles, the anterior row being incomplete. The episternum of
the metathorax has no bristles, while the epimerum bears 6 long
ones (3, 3) and 1 or 2 short ones.
Abdomen.—The tergites bear 2 rows of bristles, there being
one or more additional bristles in front representing a third row.
The second tergite has one or two slender, bristle-like spimes on
each side at theapical edge. The seventh tergite bears | apical
40*
626 THE HON. N. C. ROTHSCHILD ON [ May 26,
bristle, which is short, being about as long as the third post-
median bristle of this segment. 'The sternites of segments 3 to 7
have a row of 4 long bristles on each side, the basal sternite,
however, bearing but 1 bristle.
Legs.—The mid and hind coxe are rather narrow and long,
both having a single bristle posteriorly at the apex. There are
no short spines on the inside of the hind coxa. The fore-femur
has on the outside 1 subapical bristle and 4 or 5 lateral ones.
The mid and hind femora bear 2 subapical bristles on the outside
and 1 on the inside. There are about 18 bristles on the outer
surface of the hind tibia. The hind tibia has 7 dorsal notches.
The longest dorsal bristle of this tibia is hardly twice as long as
the tibia is broad, the longest apical bristle being only about one-
third the length of the first hind tarsal-segment. The tibiz have
a long and a short bristle in most of the notches. The first
hind-tarsal segment is nearly as long as the hind tibia, the bristles
situated at its posterior side being rather long and very thin. The
thick apical bristles of this segment are short, the posterior one
being only one-third the length of the second segment and the
anterior one being about as long as the first segment is broad.
The fifth segment bears in all the tarsi 4 strong lateral bristles
and a subbasal pair which are placed on the ventral surface in
between the first lateral pair.
Modified segments.— ¢. The large eighth sternite bears a row
of 5 long bristles on the side. The clasper (Pl. XX VIII. fig. 1, Cl.)
has a short rounded process (P) bearing two long bristles. There
is 1 long bristle at the insertion of the movable process (F). This
process is very large, being first narrow and curved upwards and
then much widened and curved downwards. It bears a row of
long bristles at the ventral edge, a short, broad, and somewhat
twisted spine at the tip, an irregular double row of bristles on
the side and another row at the dorsal edge, the central bristles
of this dorsal row being flattened and lanceolate. The manu-
brium (M) is widest at the apex. The ninth sternite (IX. st.)
is likewise very peculiar. The internal (=vertical) arm is
broadened at the apex, this widened portion being excised in the
usual way, as Shown in the figure. The horizontal arm has an
almost straight dorsal margin, while the ventral margin is curved,
being evenly rounded in the distal third and bearing here a
number of bristles. This sternite has basally a lateral horizontal
projection crowned with a very dense brush of long thin
bristles, which are curly at the end, and among which is one long
thick bristle. Beyond the middle of the ventral margin there
are 3 short, hook-like spines, and before the apex a large and a
small hooked spine, the large one apparently bifurcating at the
apex. The tenth segment is long, the sternite bearing two long
apical bristles on each side.
Length: ¢ 3 mm.
We have one ¢ from Temuco, Chile, off Dromiciops australis,
collected by Mr. D. 8S. Bullock in November 1906.
1908. | NEW SPECIES OF FLEAS. 627
Siphonaptera collected by Mr. M. P. Anderson in Japan in 1904.
By the Hon. N. Cuaries Rotuscuitp, M.A.
(Plates XXX. & XXXI.)
The collection contains five species, of which three are new.
1, CERATOPHYLLUS MELIS Walk. (1856).
Pulex melis Walker, Dipt. Brit. p.5. n. 14 (1856) (off Badger) ;
Tasch. Die Flohe, p. 73. n. 10, t. 2. figs. 15, 15 a, t. 3. fig. 16 (1880)
(off Badger and Fox).
Five females taken off Meles anakuma at Jinrio, Tokushima
Ken, Shikoku, Japan, on February 17th.
2. CERATOPHYLLUS ARGUS, sp.n. (Plate XXX. fig. 15; Plate
XXXI. fig. 18.)
This species is closely allied to C. sciwrorwm Schrank (1804)
and C. anisus Rothsch. (1907). These insects agree with one
another in almost every detail of the exo-skeleton except in the
modified posterior segments of the abdomen.
The rostrum is somewhat longer in the new species than in the
others mentioned above, reaching to the apex of the trochanter or
a little beyond.
Modified segments.— $. The eighth tergite bears 4 (some-
times 5) bristles along the upper edge from the stigma anad,
there being 3 to 5 additional bristles on the lateral surface,
besides 1 or 2 which are placed near the ventral margin. The
eighth sternite (Pl. XXX. fig. 15) resembles that of C. anisus,
being much longer than in (. sciuwrorwm. The clasper is pro-
duced into a short obtuse process (Pl. XXX. fig. 15, P), which is
much broader and more rounded than in the allied forms. The
finger (F) of the clasper is very slender. The vertical arm of the
ninth sternite (IX. st.) is curved as in C. anisus: The proximal
portion of the horizontal arm is only slightly dilated, and there
are less hairs on this dilated part than in C. anisus— 2. The
seventh sternite of the abdomen (Pl. XXXI. fig. 18) becomes
narrower distally, the upper edge being incurved before the apex,
while the apical margin is slightly emarginate. This sclerite is
as long as it is broad. The eighth tergite resembles that of
C. sciurorum. It bears about 7 small bristles above the stigma
and | long and 2 short ones near the margin below the stigma.
The apex of this segment is slightly emarginate, there being
3 bristles at the lower angle of the apex, 3 short but rather stout
ones proximally to them, 4 along the ventral margin and 4 to 6
dorsally to these. The stylet is more than three times as long as
it is broad at the base.
Length: ¢ 2.4mm., 2 4mm.
We have four ¢ and one 9Q, taken off Petawrista leucogenys,
at Mitai, Miyasaki Kiushiu, Japan.
628 ON NEW SPECIES OF FLEAS. [May 26,
3. CERATOPHYLLUS INDAGES, sp.n. (PI. XXXI. fig. 17.)
We know only the 2. ‘The differences from the 2 of the
preceding species are apparently constant, though sight. This
flea is deeper brown than the preceding one. The seventh abdo-
minal sternite is longer, its upper margin being even more
emarginate distally, while the apical margin is not sinuate at all
(Pl. XXXT. fig. 17).
' Length: 2 3°1 mm.
We have six 9, taken off Sciwrus vulgaris orientis, at Nobori-
betsu, near Moruran, Hokkaido, Japan.
As the differences between the females of closely allied species
in this group of Ceratophyllus are generally slight (quantitatively),
we consider that the above-mentioned characters indicate that
this insect is distinct from C. argus. The discovery of the ¢
will doubtless settle the point.
4, CERATOPHYLLUS ANDERSONI, sp. n. (Plate XXXI. fig. 19.)
Thorax.—The meso- and metanotum and the abdominal ter-
gites 1-7 bear each 2 rows of bristles, the mesonotum having
some additional hairs on the back besides the small hairs situated
at the anterior edge. The metathoracic epimerum has 5 bristles
(1, 3, 1). The long apical bristle of the seventh tergite is as long
as the first hind-tarsal segment.
Legs.—The hind femur bears 2 bristles on the inner surface,
one being subbasal, the other placed subventrally near the apex.
The first mid-tarsal segment is about one-fourth longer than the
second (20:16). The bristles situated at the dorsal edge of the
hind tibia and at the anterior and posterior edges of the hind
tarsus are very deep brown. ‘The first hind-tarsal segment, like
the second to fourth segments, bears 2 rows of bristles on the
outer surface.
Modified segments.— 9. The seventh abdominal sternite is
quite unlike that sclerite of the allied species, being less broad
vertically and much more rounded (Pl. XX XI. fig.19). The eighth
tergite bears a few more bristles than in the preceding species.
The bristles of the tenth sternite are very stout. The stylet is
four times as long as it is broad at its base.
Length: 92 3:1 mm.
We have one @, taken off Putorius itatsi, at Takamori,
Kumamoto Ken, Kiushiu, Japan, on April 6th.
5, CHATOPSYLLA GLOBICEPS Tasch. (1880).
Pulex globiceps Taschenberg, Die Fiche, p. 66. n. 6, t. 2. figs. 10
10a, 11 (1880) (off Fox and Badger).
‘There are five 9 in the collection, which are apparently identical
with European specimens. Taken from Meles anakwma, at Jinrio,
Tokushima Ken, Shikoku, Japan, on February 17th.
1908.] THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 629
EXPLANATION OF THE PLATES.
Prate XXVIII.
Fig. 1. Genitalia of the g of Ctenopsyllus allophylus. Cl = clasper; P =
process of clasper; EF movable process of clasper; M = manubrium;
VILL. st. = eighth abdominal sternite; IX. st. = ninth abdominal
sternite.
Head of Ctéenopsyllus allophylus 3.
. Genitalia of the ¢ of Ctenophthalmus rettigi.
. Seventh and eighth abdominal segments of Ctenophthalmus rettigi 2.
. Sixth and seventh abdominal sternites of Pygiopsylla rainbowi 9°.
. Genitalia of the ¢ of Pygiopsylla mordax.
PLATE XXIX.
7. Head of Pygiopsylla afer 2.
8. Seventh abdominal sternite of Pygiopsylla afer 9.
9. The same of Pygiopsylla mordax 2.
10. The same of Pygiopsylla laciniosus 2.
11. The same of Pygiopsylla ferinus 9 .
12. Genitalia of the ¢ of Stephanocircus jarvisi.
PuatTE XXX.
Fig. 13. Genitalia of the g of Pygiopsylla rainbowi.
14, The same of the ¢ of Pygiopsylla gravis.
15. The same of the g of Ceratophyllus argus.
PLratTE XXXI.
Fig.16. Posterior abdominal segments of Stephanocireus jarvisi 9.
17. Seventh abdominal sternite of Ceratophyllus indages 9.
18. The same of Ceratophyllus argus °..
19. The same of Ceratophyllus andersoni 2.
SP OVE Co bo
June 16, 1908.
Dr. Henry Woopwarp, F.R.S., Vice-President,
in the Chair.
The Secretary read the following report on the additions made
to the Society’s Menagerie during the month of May 1908 :—
The number of registered additions to the Society’s Menagerie
during the month of May was 189. Of these 116 were
acquired by presentation and 46 by purchase, 15 were received on
deposit, 4 by exchange, and 8 were born in the Gardens.
The number of departures during the same period, by death
and removals, was 175.
Among the additions special attention may be directed to :—
One Black-faced Chimpanzee (Anthropopithecus troglodytes),
var. 2, from Sierra Leone, deposited on May 30th.
One Agile Gibbon (Hylobates agilis) 3 , from Sarawak, presented
by the Earl of Crawford, K.T., F.Z.S., on May 8th.
Three Grévy Zebras (Hquus grevyi) 5, 2 2, from Abyssinia,
purchased on May 9th.
Twenty-one Indian domestic Cattle (Gos indicus), representing
five different breeds, and 5 Fat-rumped Dumba Sheep (Ovis aries),
from India, presented by H.G. the Duke of Bedford, K.G.,
President of the Society, on May 13th.
630 DR. FORSYTH MAJOR ON FOREST-BED MAMMALS. [June 16,
Dr. A. Smith Woodward, F.R.S., F.Z.S8., exhibited photographs
and fragments of skin and bone of a Mammoth and a Rhinoceros
discovered in an ozokerite mine at Starunia, Galicia. The
carcases of these animals appeared to have found their way into
an old marsh saturated with petroleum, which had completely
preserved them. The photographs and specimens had been
received from Dr. George von Kaufmann, who intended to present
them to the British Museum.
Jaw of Canadian Beaver, with five lower cheek-teeth.
Dr. C. I. Forsyth Major, F.R.S., F.Z.S., exhibited, and made
the following remarks upon, the lower jaw of a young Canadian
Beaver (B.M. No. 55.3.11.4), in which there was present on each
side a small conical tooth anterior to the deciduous premolar
(text-fig. 131).
Text-fig. 131.
a) it
Ld fe
2
A.
Molar teeth of right lower jaw of Castor canadensis.
A. Side view; B. Upper view of d; and pj. po=supernumerary premolar.
“ No Simplicidentata are known with normally more than four
lower cheek-teeth ; this holds good also with respect to the oldest-
known fossil Simplicidentata. We must, however, postulate still
older fossil predecessors with five lower cheek-teeth. As a transi-
tional stage between the latter condition and that exhibiting four
cheek-teeth, Rodents must have existed, probably in the Lower
Kocene, which had normally exactly such a diminutive anterior
tooth as is abnormally shown in the present specimen. I therefore
consider this supernumerary premolar to be a case of atavism.”
On species of Castor, fossil and recent.
Drawings of some remains of two species of Castor from the
East Runton Forest-bed were also exhibited by Dr. Forsyth
Major.
(1) Text-fig. 132 represents the four cheek-teeth from a right’
mandibular ramus in the British Museum (M 7024), from
Mr. Savin’s collection. They are remarkable for the complex
and elegant plication of their enamel, whereby they agree with
the Beaver from the Upper Pliocene of the Valdarno, Castor plici-
dens Maj., this specific name implying the principal and almost
1908. | DR. FORSYTH MAJOR ON FOREST-BED MAMMALS, 631
the only character by which the fossil remains can be distin-
guished from recent Beavers. Dr. Bosco some years ago* fully
described and figured the Valdarno remains; he points out
another characteristic feature of this species, viz. the considerable
breadth of the incisors. Herein the fossil from East Runton
equally agrees with the Valdarno specimens.
Text-fig. 132.
Castor plicidens Maj. East Runton Forest-bed.
Upper view of right lower molar series.
(2) A second species of a Castor from the Forest-bed (B.M.
M 7025), likewise from the East Runton upper freshwater bed,
is represented by a left mandibular ramus, vertically split from
before backwards, only its outer half, and of the teeth m, alone
(text-fig. 133) being preserved. It doubtless belongs to the same
Text-fig. 133.
m2
Castor sp. East Runton Forest-bed.
Upper view of left mo.
species as the specimens described from West Runton by Mr. E. T.
Newton ; as in the latter, the molar is slightly smaller than the
corresponding one of C.. plicidens ; the incisor, as apparent from
its alveolus, is narrower. The enamel of the molar is consider-
ably less plicated than in the latter species, although slightly
more so than is the rule in recent Beavers ; a moderate plication
of the enamel occurs, however, in old specimens of the recent
form (text-figs. 134, 135 B, 136 A).
Text-fig. 134.
Castor fiber L. Peat deposit, England.
Upper view of right lower true molars.
Mr. Newton has identified the West Runton Beaver with the
* Palezontographia Italica, vol. v. (1899).
632 DR. FORSYTH MAJOR ON FOREST-BED MAMMALS, [June 16,
recent European animal; and, in fact, his specimens, as well as
the specimen in the British Museum Geol. Dept. No. M 7025
from East Runton, share with C. fiber the characters—so far as they
are known in the fossil specimens—which distinguish the recent
animal from C’. plicidens. I expect, however, that more complete
specimens will reveal differences from the recent form or forms.
The European Beaver ranges, or ranged, from Great Britain to
Mongolia, and from Lapland to Spain (according to Strabo) and
Italy. There is therefore every likelihood that more than one
form will have to be distinguished in this vast region; but in
order to arrive at some definite conclusion, we require whole series
(at least of skulls) from the different countries : this will be no
easy task, considering that in most countries the Beaver has
become extinct within historic times.
According to Prof. Matschie*, Desmarest separated the Beaver
of the Rhone under the name of Castor gallic, while Owen pro-
posed the name of (’. ewropwus for the English sub-fossil Beaver.
The German writer restricts the Linnean name Castor fiber to the
Swedish Beaver, known to him only by the figures of the teeth
and the skull in Meves’s Atlas’. From the Swedish Beaver that
of the Elbe is said to differ by some characters of the skull
and the teeth, and is therefore called Castor albicus. The name
C. balticus is given to the Beaver which formerly lived in Pome-
rania and Holstein and is based on two skulls. For a specimen
from Western Poland (drainage system of the Vistula) which
lived at the Berlin Zoological Gardens, the name of C. vistulanus
is proposed, its skull being found to be different from those of
“CO. albicus” and “C. balticus.” The skull of “C. vistulanus”
is declared to differ also considerably from a skull of a Beaver
from the Caucasus and from one from Poland in the Kiew
Museum, both of them figured by Brandt. Lastly, a skull
from Schwerin a. d. Warthe (Posen), although agreeing in most
characters with “ C. vistulanus,” is supposed to be possibly a
distinct species.
If the Rhone Beaver can be shown to be a distinct form, the
name CU. gallie will be available for the same; this was not,
however, Desmarest’s opinion. From the context and the explicit
statement of the latter’s notice § it results that he introduced this
name for the European Beaver in general, which he wished to
distinguish from the American species.
In the same way Owen applied the name C. ewropwus to the
European Beaver generally, he did not mean to restrict it to
the English animal as Matschie assumes. Until further notice
“C. gallie” will therefore remain a synonym of C’. fiber L., and
the same is the case with “ C. ewropeus.”
* Sitzungsber. Ges. Naturf. Fr. Berlin, No. 8, 1907, pp. 215-220, pls. i. & ii,
ie a ofver Skandinaviens Dagedjur, Suppl. pl. ii. figs. 1, 1a, 1b (Stockholm,
“ Mém. Ac. Sc. St. Pétersbourg (6) ix. Sc. Nat. vii. pls. i. & ii. (1855).
§ Desmarest, in Nouv. Dict. d’Hist. Nat. v. pp. 372, 373 (1816).
1908. ] DR. FORSYTH MAJOR ON FOREST-BED MAMMALS. 633
Of the Swedish Beaver Matschie says that in the last upper
molar the internal enamel-fold is absent, ‘“‘ und bei den tibrigen
Molaren legt sie sich nicht an die vorderste Aussenfalte an,
sondern verlauft senkrecht zum Langsdurchmesser der Zahne
und endigt frei zwischen der vorderen und mittleren Aussenfalte.”
Besides, ‘‘ the nasals are obliquely truncated towards the front and
not notched near the intermaxillary, so that their anterior margin,
seen from the side, seems to be angularly notched.” *
Text-fig. 135.
A. Castor fiber L. Sweden. Lower view of right upper molar series. Copied from
Meves’s Atlas ofver Skandinaviens Dagedjur, Suppl. pl. i. fig. la.
B. Castor fiber L. Lapland. Lower view of right upper molar series of an old
specimen. R. Coll. of Surgeons Museum.
Text-fig. 136.
A. Castor fiber L. Peat-deposit, England. Lower view of right upper true molars.
B. Castor fiber L. Lower Rhone (B.M. No. 94.5.30.1). Lower view of right
upper molar series.
An inspection of the figure to which Matschie alludes (see text-
fig. 135 A, which is a copy of the figure in Meves’s Atlas) shows
that the internal enamel-fold is not absent from the last upper
molar, only it is fused with the antero-external fold, a condition
* Op. cit. p. 216.
634 DR. FORSYTH MAJOR ON FOREST-BED MAMMALS. [June 16,
which obtains not unfrequently in younger stages of Beavers’
molars (especially in p' and m*) and sometimes persists in the
adult. ‘Text-fig. 136 B shows the upper molars of a young adult
of the Rhone Beaver, where m® exhibits the pattern of Meves’s
figure. A second, rather old specimen of the Rhone Beaver in
the British Museum (No. 5.3.9.1) shows likewise on both sides
the same conformation of the m* as in the younger specimen.
Text-fig. 135 B, on the other hand, exhibits the upper molar series
of a rather old Swedish Beaver in the Museum of the Royal
College of Surgeons*, kindly placed at my disposal by Prof. Keith.
Here m* presents the normal condition ; and together with the
other cheek-teeth, when compared with Meves’s figure, illustrates
the well-known fact of the considerable individual variation in the
pattern of the enamel, chiefly due to different stages of wear.
Text-fig. 137.
Castor fiber L. Lapland. Side view of anterior portion of skull.
R. Coll. of Surgeons Museum.
The text-figure 137 exhibits the side view of the anterior
portion of the skull in the Royal College of Surgeons, which does
not depart from the normal condition in the European Beaver.
The figure in Meves’s Atlas to which Matschie alludes is appa-
rently taken from the skull of a youngish specimen, and, besides,
may not be quite correct.
As to the characters assigned to Matschie’s new species, C. bal-
ticus, notably the greater elongation of the sagittal crista, they
are simply those of old age. ;
* No. 3102, “from an animal taken in Lapland about the year 1830.”
P.Z.S.1908 Peis
ay
Mies
H.Goodchild del,et lith. Toth imp.
CMERUR ONS IDE OQRUDIL. Y
1908. | ON MAMMALS FROM NORTH CHINA. 635
Dr. Forsyth Major also exhibited photographs of Pliocene
Bovine from specimens in the Florence Museum, stating that
these unpublished figures showed the great variability of the
Pliocene Bovine. He added that he endorsed Falconer’s opinion
that these Pliocene Bovine were nearly related to the primitive
Buffaloes from the Siwaliks.
The following papers were read :—
1. The Duke of Bedford’s Zoological Exploration in Eastern
Asia.—X. List of Mammals from the Provinces of
Chih-li and Shan-si, N. China. By OLpFieLp THomas,
HER Ssh Z.Ss-
[Received May 2, 1908. ]
(Plate XXX.)
During the four months following his excursion to the Mon-
golian plateau +, Mr. M. P. Anderson made collections in different
parts of the northern provinces of China, Chih-li and Shan-si,
and it is an interesting comment on our ignorance of the Fauna
of that part of the world that, in spite of the previous work
of David, Swinhoe, Styan and others, he has obtained quite a
number of new forms.
Throughout the region, the country has proved to be extremely
barren and poor in mammals, and the possible collecting-grounds
few and far between. But this very fact renders such collecting
places as Mr. Anderson has found all the more interesting, for
they almost bear the character of faunistic islands, in which
the original inhabitants have been locally preserved, and which
are separated from each other by a sea of barren treeless plains
where few animals can live. The proper exploration of these
oases of life is therefore peculiarly valuable. No doubt the
difficulties of collecting have been accentuated during the winter
months, and we may hope that during the present spring
Mr. Anderson will find a number of additional forms which in
the winter have been lying dormant.
Of previous literature there is not much to be referred to
beyond the well-known publications of David, Milne-Edwards,
and Swinhoe, and an interesting paper by O. F. von Mollendorff ¢
giving a popular account of the Mammals of Chih-li, with notes
on the Chinese names.
* [The complete account of the new species described in this communication
appears here; but since the name and preliminary diagnosis of one were published
in the ‘ Abstract,’ that species is distinguished by the name being underlined.—
EpirTor. |
+ Supra, p. 104. ‘
{ “he Vertebrata of the Province of Chih-l, with Notes on Chinese Zoological
Nomenclature,” Journ. N. China Branch Roy. As. Soc. (2) xi. p. 41, 1877.
636 MR. OLDFIELD THOMAS ON [June 16,
Prof. Matschie’s work* on the Filchner Mammals deals with
a somewhat different region, further to the west and south, but,
true to his peculiar creed that the animals of different river-
basins must be specifically distinct from each other, the author
gives new names to some of the Chih-h mammals. The material
he worked upon seems to have consisted largely of single pur-
chased skins, mostly without skulls, and the possibility of any
such variation in colour as is found in the Shan-si foxes (see
infra) is entirely ignored.
No less than 19 species are described as new by Prof. Matschie
on undated skins without measurements or skulls, or at least
without mention of the latter, and I cannot refrain from ex-
pressing the opinion that such work is neither worthy of the high
standing of the Berlin Museum nor of the present date, when
pains are being taken in all directions to ensure that mammal
work should be based only on proper and carefully collected
material. The Americans have set us a good example in this
respect, and it is to be regretted that work issuing from the
Berlin Museum should be done in so retrograde a manner.
All the names thus founded will remain an incubus to science
until the time when they can be successively examined and
weighed in the light of complete material, such material, for
instance, as the Washington Museum has received from Dr. W. L.
Abbott, or our own National Museum owes to the Duke of
Bedford, Mr. C. D. Rudd, Mr. W. H. Balston, and many others.
About 100 specimens are dealt with in the present communi-
cation, belonging to 20 species.
The following are Mr. Anderson’s notes on the localities he
collected in :—
CHIH-LI.
“ After returning from my trip to the Mongolian Plateau, I
visited Tung-ling, the forest of the reserve of the Hastern
Imperial Tombs, and on 12th September, 1907, I began work at
Yen-mon, a hamlet about 65 miles north-east of Peking. At
this point I was well within the hills which border the Chih-li
plain on the north, and my hamlet was at an altitude of about
1000 ft., while the surrounding hills rose to 1800 or 2000 ft.
The hills of this region are for the most part very abrupt, rugged
and rocky. Difficult peaks and narrow, almost impassable canyons
are frequent. Tung-ling is forbidden ground to the wood-cutter
and charcoal-burner, therefore woods persist and even grow dense
and old in the remoter parts of the reserve. One finds some fine
oaks, walnuts, chestnuts, and cottonwoods The frequently
occurring open spaces are well covered with deep grass.”
SHAN-SI.
“On 25th October, 1907, 1 reached Tai-Yuen-Fu, the capital
* Wilchner Exped. Zool. pp. 134-244, 1907 (postdated 1908).
1908. | MAMMALS FROM NORTH CHINA, 637
of Shan-si Province, and on 31st October left that city for the
mountain-range some 15 miles west of there.
“At this point, at an altitude of 5400 feet, I found the
mountains sparsely covered with pine trees and the canyon sides
overgrown with dense bushes.
“On 17th November I began work at Chao-Cheng-Shan, a
mountain of 10,000 ft. altitude, situated about 100 miles west-
north-west of Tai-Yuen-Fu. There, at an altitude of 8000 ft.,
1 made my home in a peasant’s hut. Above me extended a
dense forest of spruce and hemlock, below I overlooked the
rugged bare hills and cultivated valleys characteristic of North
China.
“‘T remained in this place till Dec. 6th, 1907. The weather
throughout was fiercely cold, as a north wind blew almost
unceasingly.
“On December 27th I went eastward of Tai-Yuen-Fu about
20 miles to a temple wood among the ‘loess’ hills. But this
proved such a poor collecting-ground, and the weather so very
cold, that on 4th January I thought it advisable to return to
the city.”
1. RHINOLOPHUS FERRUM-EQUINUM NIPPON Temm.
3. 1571. Cave 30 miles W. of Peking. 600".
This and the two following species were obtained in a sacred
cave which Mr. Anderson might not have been allowed to enter,
certainly not to shoot in, had it not been for the kind offices of
Dr. J. H. Ingram of Tung-chou, who persuaded the priest of the
cave to allow him to do so. Great numbers of bats were hanging
from the roof, but besides the one Rhinolophus, which was caught
low down within reach, and the two specimens of J/yotis, all
proved to belong to one species, a Winiopterus.
2. Myoris (LEUCONOE) PEQUINIUS, sp. n.
3S. 1573, 1589. Cave 30 miles W. of Peking. 600’.
A comparatively large Leuwconoe, with a fringed interfemoral
membrane.
In size one of the largest species of the group, exceeding all the
Old-World species of Lewconoe, except M. ricketti. Fur rather
short and velvety, hairs of back about 5 mm. in length. General
colour above uniform ‘drab-grey,” the bases of the hairs slaty.
Under surface whitish grey, the ends of the hairs nearly white,
their bases slaty; under side of hind legs and the anal region
edging the membranes white and practically hairless. Ears of
medium size, rather narrow, concave on their external border ;
tragus about half the length of the ear, narrow, not sharply
pointed, slightly curved outwards above. Wings attached to the
lower end of the tibiew. Feet of average Leuconoe proportions.
Interfemoral membrane fringed posteriorly with pale buffy hairs ;
tip of tail not projecting from the membrane, so far as can be
638 MR. OLDFIELD THOMAS ON [June 16,
determined on skins from which the caudal vertebre have been
pulled out. In colour all the membranes and the feet are dark
drab-grey, except that the terminal half-inch of the interfemoral
is slightly marbled with white.
Median upper premolar minute in one specimen, absent in the
other, but both are very old examples with the teeth much worn
down. Also very minute in the lower jaw.
Dimensions of the type (the starred measurements taken in the
flesh) :—
Forearm 50 mm. (in the second specimen 48-5).
*Head and body 62 mm.; *tail 427; *hind foot (s. u.) 12;
*ear 18; tragus on inner edge (dry) 7; third finger, metacarpal
46, lst phalanx 14°5, 2nd phalanx 14; tibia 18.
Skull—basi-sinual length= (¢c.) 145 mm.; zygomatic breadth
12:2; interorbital breadth 4:9, breadth of brain-case 4:7; front
of canine to back of m’ 6°9.
Hab. China, 30 miles W. of Peking. Alt. 600’.
Type. Old male. B.M. No. 8.8.7.2. Original number 1573.
Collected 11 October, 1907.
By its size and the presence of a fringe on its interfemoral
membrane this very distinct Bat is easily separable from any
known Asiatic member of the genus J/yotis.
The specimens were found hanging in the same cave as the
series of Miniopterws—the association recalling that so frequent
in Europe of Miniopterus with Myotis (Leuconoe) capaccinii.
3. MINIOPTERUS SCHREIBERSI CHINENSIS, subsp. n.
29,132. 1574-1588. Cave 30 miles W. of Peking. 600’.
Size averaging slightly larger than in J/. s. gaponie Thos., the
forearm ranging in length from 47 to 50 mm.
Colour dark, as usual in the Eastern forms, but without the
reddish ‘‘ Prout’s brown ” suffusion found in japonie, the general
tone being markedly more drabby than in that form. The colour
cannot be exactly matched in Ridgway, but is between “seal-
brown ” and ‘“ drab,” with something of each in it according to
the light the specimens are seen in.
This difference in colour is perfectly uniform throughout series
of fifteen specimens of chinensis and a dozen of japonic, so that
it seems necessary to recognise the N, China form as a different
subspecies from that of Japan.
Dimensions of the type, measured in the flesh :—-
Forearm 49 mm.
Head and body 62 mm.; tail 52; hind foot 10:5; ear 12.
Skull—greatest length 15:8 mm.; basi-sinual length 12.
+ Probably below the normal; in the other specimen 49 mm.
{ In describing Bats a name is frequently wanted for the measurement from the
basion to the hinder edge of the anterior palatal notch. As the Latin for notch,
incisio, makes a compound too like one founded on the incisor teeth, I would suggest
the above word, based on sinus, a bay or gulf, with which this deep rounded hollow
may be suitably compared.
1908. | MAMMALS FROM NORTH CHINA. 639
Type. Adult female. B.M. No. 8.8.7.15. Original number
1585. Collected 11 October, 1907.
Bonhote’s JZ, s. fusews from the Liu-Kiu Islands is smaller,
with a forearm about 44 mm. in length.
4, CrocipuRA coREx Thos.
3. 1553. Imperial Tombs, 65 miles E. of Peking.
Closely similar to the typical Korean series.
“Trapped beneath a thick bush among loose rocks in a cold
damp canyon.”—W/, P. A.
5. CHODSIGOA HYPSIBIA de Wint.
3. 1558. 92. 1559. Imperial Tombs, 65 miles E. of Peking.
1000’.
Since Mr. de Winton described his Soriculus hypsibius * from
N.W. Sze-chuen, correctly noticing the absence of the minute
fourth unicuspid found in typical Soriculus, Dr. Kashtchenko +
has made a new subgenus, Chodsigoa, for the species without that
tooth. But while recognising the distinction of Chodsigoa, and
even considering it rather genus than subgenus, I think that the
typical species ‘‘ Soriculus (Chodsigoa) beresowshi” is undoubtedly
identical with de Winton’s animal, coming from practically the
same locality, and having just about the same proportions. Nor
can I at present see any reason to distinguish Mr. Anderson’s
specimens from hypsibia, in spite of their very different locality.
Mr. de Winton’s type was previously the only specimen of this
rare group of Shrews possessed by the British Museum.
“Trapped in a radish garden on a rocky hillside.” —W/. P. A.
6. VULPES VULPES L.
3. 1645, 1646, 1651. ‘Tai-Yuen-Fu. 2700’.
These three skins illustrate the remarkable colour variation
found among Foxes, one of them having a whitish, the second a
slaty-grey, and the third a red under-side.
“ Common.”—W.. P. A.
7. SCIUROTAMIAS DAVIDIANUS M.-Edw.
3d. 1570. Imperial Tombs, 65 miles E. of Peking.
@. 1643. 100 miles N.W. of Tai-Yuen-Fu, Shan-si. 8000’.
The genus Sciwrotanvias was formed by Mr. Gerrit Miller = for
this remarkable Squirrel, which has a skull very like that of a
Chipmunk, with the external appearance of a Squirrel. Prof.
Milne-Edwards had previously noticed its near relationship to
Tanuas §.
“ Not a common species, aS only one was seen besides the
* P. Z. S. 1899, p. 574.
+ Ann. Mus. Zool. Acad. Sci. St. Pétersb. x. p. 252 (1906).
~ P. Biol. Soc. Wash. xiv. p. 23 (1901).
§ Rech. Mamm. p. 160 e¢ seqq. (1874).
Proc. Zoou. Soc.—1908, No. X LI. 41
640 MR. OLDFIELD THOMAS ON [June 16,
present specimens. From the native reports I judge this animal
to be a rock-loving species. Its colour is very like that of the
local rocks.
‘“‘Bvidently rare in Shan-si.”
MM, P. A.
8. EUTAMIAS SENESCENS Mill.
3. 1561, 1562. Imperial Tombs, 65 miles HE. of Peking.
The type of #. senescens was obtained about 15 miles to the
west of Peking.
“Rare. I failed to see any alive, but got these two from
hunters.”— MW. P. A.
9. MreRIoNES PSAMMOPHILUS M.-Edw.
3. 1644. Tai-Yuen-Fu, Shan-si. 2800
This species was discovered by Pere David at Suen-hoa-fur
near Kalgan, but although Mr. Anderson, when in that region,
obtained a good series of the other species, JJ. unguiculatus, he
did not get MW. psummophilus, which we are therefore very glad
to obtain, as the only specimen in the Museum is without a
skull.
“Tnactive in winter, but doubtless conspicuous in warm
weather.”—/, P. A.
10. MERIONES AUCEPS, Sp. n.
3. 1647. East of Tai-Yuen-Fu, Shan-si. 4000’. 31 Dec.,
LOO SBME Non8:82/030) se lape:
A medium-sized species with the immensely large inflated
bullee of W/. erythrurus.
Size about as in MW. unguiculatus, the common species of
Mongolia. General colour above of the usual buffy fawn, very
much as in that species, Sides with a brighter buffy band edging
the white. Belly practically pure white, the hairs mostly white
to their roots, but some few with a little grey at their bases.
Kars bright buffy, contrasting with the general tone. Hands
and feet pure white; claws thin, whitish horn-colour, not blackish
as in WM. wnguiculatus. Tail well-haired, but not specially tufted,
rich ochraceous-bufty throughout, a few hairs at the extreme end
tipped with black.
Skull much more heavily built than that of IZ. wngwiculatus,
with broad interorbital space and heavy muzzle. Bulle ex-
ceedingly large, the part just in front of the external meatus so
swollen as to touch the zygomata, which they surpass in lateral
spread.
Dimensions of the type, a rather young adult :—
Head and body 110 mm.; tail 105; hind foot 31; ear 15-5.
Skull—greatest length 36 mm. ; basilar length 27-5; zygomatic
breadth 19°5; tympanic breadth 20; interorbital breadth 6-2 ;
palatilar length 15-5; diastema 9-2; palatal foramina 7; greatest.
1908. ] MAMMALS FROM NORTH CHINA. 641
horizontal diameter of bulle 14:5; length of upper tooth-row
(alveoli) 5:3.
flab. and Type as above.
Of the other Chinese Meriones described, this very pretty
Species may be distinguished from J, unguiculatus by its whitish
claws, buffy ears, whiter belly, and less blackened tail; from
M. psammophilus by its larger size and buffy ears; and from both
by its enormously larger bulle, which indicate that it is not
really closely allied to either of them.
11. Mus conructanus, M.-Edw.
3.1551. 2. 1567, 1569. Imperial Tombs, 65 miles E. of
Peking.
¢. 1600. 92. 1599. Near Tai-Yuen-Fu, Shan-si. 5300’.
3. 1648. East of Tai-Yuen-Fu. 4000’.
The Imperial Tombs specimens are very similar to the Kuatun
examples considered as typical confucianus by Bonhote, and are
equally distinct from the form found on the Chefoo Peninsula
which I have named J. c. sacer in a previous paper.
*“Not common; trapped among broken rocks and canyon-
bottoms on hill-sides.”—W. P. A.
12. Mus waGNERI MonGcouiuM Thos.
6. 1552, 1566. 9. 1556, 1565, 1568. Imperial Tombs,
65 miles HE. of Peking.
6. 1611, 1617. 2. 1603, 1621, 1629, 1635. 100 miles
N.W. of Tai-Yuen-Fu, Shansi. 8000’.
Although these mice have no trace of an anterior supple-
mentary cusp on their first upper molars, they are not improbably
related to the Kan-su Jus ‘“(Leggada)” gansuensis Satunin, a
form evidently of the musculus group, and not a true Leggada
at all.
“This small mouse seemed to be the commonest species in the
vicinity of the Imperial Tombs, but still it was not met with very
often. It lived under half-buried stones among the grass and
bushes, or along the rocky banks of streams.
‘‘Tn Shan-si it was somewhat common in the fields and about
the peasants’ threshing-grounds at Chao-Cheng-shan, but I did
not see it elsewhere.”—-M/. P. A.
13. APoDEMUS SPECIOosUS Temm.
3. 1560, 1563. @. 1555, 1564. Imperial Tombs, 65 miles
E. of Peking.
3. 1598. ©. 1590, 1594. Near Tai-Yuen-Fu, Shan-si.
5300’.
S$. 1623, 1636. 2. 1608, 1630, 1631. 100 miles N.W. of
Tai-Yuen-Fu. 8000’.
The Shan-si specimens are not unlike the Korean subspecies
A. s. peninsule, to which perhaps the whole series should be
4|*
642 MR. OLDFIELD THOMAS ON [June 16,
provisionally referred until further material is received bearing
on their relationship to A. s. chevrieri and draco, of the 8.W. and
S. of China.
“Ags common as A. agrarius, but usually livmg among the
bushes.” — WU. P. A.
14. APODEMUS AGRARIUS COREE Thos.
go. 1550, 1554, 1557. Imperial Tombs, 65 miles E. of Peking.
Ags with most of the other Imperial Tombs animals the nearest
relationship of this striped rat seems to be with the Korean sub-
species, but in such a variable group the present series is not large
enough for me to be very positive on the point.
“Moderately common ; living in the tall grass which grows in
certain open valleys.”—W. P. A.
15. GRICETULUS TRITON de Wint.
3. 1622 (immature). Chao-Cheng-Shan, 100 miles N.W.
of Tai-Yuen-Fu, Shan-si. 8000’.
“ Brought to me by a farmer, who had caught it im a straw-
stack in his threshing-ground.”—W/, P. 4.
16. CRICETULUS ANDERSONI, sp. 0.
6. 1591, 1592, 1596, 1601) 1602) 2. 1595; ) Veo ease
Near Tai-Yuen-Fu, Shan-si. 5300’.
3S. 1626, 1641. 9. 1604, 1605, 1619, 1620, 1627, 1628, 1642.
100 miles N.W. of Tai-Yuen-Fu. 8000’.
3. 1649. 9. 1650. East of Tai-Yuen-Fu. 4000’.
A small long-tailed species like C. longicaudatus M.-Edw., but
with the belly-hairs grey basally.
General colour above drab-grey, the centre of the back in-
distinctly darker, but without a definite dark line. Sides often
with a vaguely marked buffy area just in front of the hips.
Under surface dull whitish grey, not sharply defined laterally,
the hairs dark slaty for about two-thirds of their length. Lars
blackish, with sharply contrasted white tip and edges. Hands
and feet white; palms naked; soles hairy except in the region
of the pads and on the under sides of the toes. Tail comparatively
long, coloured above like the back, white below, and sometimes
white all round at the tip.
Skull not strikingly different from that of C. griseus.
Dimensions of four specimens, measured in flesh :—
g. Head and body 80 mm.; tail 38; hind foot 15 ; ear 15.
3 (type) ,, Soe ee SOs 5 16. 3; es Milas
2 : ” 84 ” 9 99 44 ; 9 hi 5 9 1,3},
ae %» BD) 93 a 805 7 LOrD ie, ae
Skull of type—greatest length 25°5 mm. ; basilar length 20:3 ;
zygomatic breadth 13:3; nasals 9°23; interorbital breadth 3°7 ;
breadth of brain-case 11‘7; interparietal 2°6x9-2; palatilar
1908. ] MAMMALS FROM NORTH CHINA, 643
length 10-4; palatal foramina 5:3; length of upper molar
series 3°9.
Hab. Shan-si
8000".
Type. Adult male. B.M. No. 8.8.7.71. Original number 1626.
Collected 3 December, 1907.
This Hamster is readily distinguishable from C. longicaudatus
M.-Edw. by its grey-mixed belly, from C. grisews M.-Edw. and
C. dichrootis Sat.* by its longer tail, and from “ Urocricetus ”
kamensis Sat.* by its shorter tail and smaller size. I confess
I fail to see any sufficient reason why the long-tailed forms of
this group should be separated in a special subgenus, even apart
from the evidence given by C. andersoni, the tail of which is of a
more or less intermediate length.
In laying such emphasis on the colour of the ears in C. dichro-
otis, Dr. Satunin does not seem to be aware that particoloured
ears are a characteristic of most of the Far Eastern species,
C. griseus and C’. obscurus both having similar black and white
ears.
I have named this pretty species after its collector, Mr.
Anderson, by whom its distinctness from C. griseus was noticed.
“The common species of Hamster in Shan-si. It inhabits the
neighbourhood of cultivated fields, making many horizontal
burrows just beneath the surface of the earth.”—I/, P. A.
type from 100 miles N.W. of Tai-Yuen-Fu.
17. CrAsEoMys REGULUS Thos.
@. 1549. Imperial Tombs, 60 miles E. of Peking.
I cannot perceive any character by which this Vole can be
distinguished from C. regulus, which was described from Korea.
“‘ Trapped among bushes at the foot of a talus-slide; the only
specimen seen, though I made great efforts to find more.”—
Si gpl er sie
18. CRASEOMYS SHANSEIUS, sp. n.
3S. 1610, 1618, 1625, 1632, 1633, 1634, 1637, 1638, 1640.
©. 1607, 1609, 1616, 1624, 1639. 100 miles N.W. of Tai-
Yuen-Fu, Shan-si. 8000’.
‘Taken in spruce forest.”
A large pale-coloured species with comparatively short tail.
Fur long, soft and loose; hairs of back (in winter coat)
12-13 mm. in length. Upper surface pale greyish Hvotomys-
colour, the reddish more suffused with grey than usual, though
possibly this is not so much the case in specimens in summer
pelage. Face and sides markedly greyer, without rufous suffusion.
Under surface pale cream-bufty, the broad slaty bases to the hairs
showing through. Hands and feet white above. Tail heavily
haired, brown above, whitish or cream-coloured on sides and
below.
* Ann. Mus. Zool. St. Pétersb. vii. pp. 567 & 574 (1902).
644 MR. OLDFIELD THOMAS ON [June 16,
Skull rather smaller than that of C. regulus; on the whole
similar in shape except that the mesopterygoid fossa is unusually
narrow, and the ridges bounding it do not slope upwards (dorsad)
so much as usual, as they pass above (dorsad to) the posterior edge
of the palate; the vertical space formed between the ridges and
the hinder end of the palate is therefore of much less vertical
extent than in other members of the Hvotomys-Craseomys group.
Teeth apparently as usual, the last upper molar with six, and
the first lower with nine salient angles.
Dimensions of four of the largest specimens :—
3 (veryold). Head and body 104 mm.; tail 32; hind foot 18; ear 13.
3 (type) ” te) een Sy 33; ” TS; ” 13.
2. iy 100 Fries aos 33) See
On 93 23) tp yon aeeees 9 yolileghcemmece
Skull of type—greatest length 26-2 mm.; basilar length 23:3 ;
zygomatic breadth 14°6; nasals 7-4; palatilar length 12°6; palatal
foramina 5°5; length of upper molar series (crowns) 6:2.
Hab. As above.
Type. Adult male. B.M. No. 8.8.7.85. Original number 1634.
Collected 4 December, 1907.
This fine species, which was found by Mr. Anderson high up in
the spruce-covered mountains N.W. of Tai-Yuen-Fu, may be
readily distinguished from any of its allies by its unusually short
tail, which barely surpasses that of average members of the
Microtine series of Voles. In addition its pale colour and peculiar
palate are characteristic.
No Red Voles have hitherto been found anywhere near
Shan-si, the nearest being the Chih-li example of C. regulus
referred to above. I continue to use the name Craseomys in a
generic sense for the group of Red Voles which either do not
form roots to their molars at all, or only do so in extreme old age.
Hardly a specimen of the Far Eastern species has been found with
its molar teeth no longer encapsuled, so that it is of interest to
mention that in No. 1625 the capsules have almost disappeared,
and the molars appear to be on the point of forming roots. In
the Scandinavian C. rufocanus roots appear to be formed at rather
an earlier period of life, so that that species, while technically
genotype, is the least typical member of the genus.
‘“‘ Rather common in the brush-covered valley-bottoms at Chao-
Cheng-Shan. Not seen elsewhere.” P. A.
19. Lepus swinHoetr Thos.
2. 1572. Tung-chou, on the Peking plain.
3. 1613. 2. 1606. 100 miles N.W. of Tai-Yuen-Fu, Shan-si.
8000".
‘Said to be common round Peking, but if so is not easily
seen.”
“A common species at Chao-Cheng-shan, and near Tai-Yuen,
1908. | MAMMALS FROM NORTH CHINA. 645
as indicated by the number of tracks, but difficult to secure as
they do not flush till one is close upon them.”—JW/, P. A.
20. CAPREOLUS BEDFORD! Thos. (Plate XXXII.)
Abstr. P. Z.S. 1908, p. 32 (June 16).
3. 1612, 1614 (skulls only). @. 1615. 100 miles N.W. of
Tai-Yuen-Fu, Shan-si. 8000’.
Size rather larger than in the European C. capreolus, therefore
much smaller than in C. pygargus. Horns comparatively small,
therefore not like those of C. tianshanicus.
General colour above of a winter specimen buffy clay-colour,
rather paler than the tone often rather loosely called “red” by
sporting writers, therefore very different from the greyish brown
of C. capreolus. Under surface dull whitish. the hairs grey at
base, then whitish, washed terminally with pale fulvous. Head
rather more rufous. Area behind nostrils blackish, but, at least
in the winter coat, without the marked black band characteristic
of C. capreolus. Lips, both upper and lower, and chin dull white,
without blackish marks laterally. Hairs of throat ‘ drab-grey,”
with whitish tips. Ears grizzled buffy and blackish, with darker
edges, their internal surfaces whitish. Limbs dull buffy or pale
tawny, more fulvous proximally, paler distally.
Skull larger than that of C. capreolws, markedly smaller than
in C. pygargus. Horns comparatively slender, with the usual
three tines.
Dimensions of the type, measured in flesh :—
Head and body 1125 mm.; hind foot 310; ear 130.
Skull of type—condylo-basal length 186 mm.*
Skull of old male—condylo-basal length 207 mm.; greatest
breadth 95; length of nasals 70; interorbital breadth 57; palatal
length 126; length of upper tooth-series 66.
The condylo-basal lengths of two adult males of C. pygargus are
221 and 225 mm., while in a pair of C. capreolus this measurement
is 184 (¢) and 181 (@).
Type. Old female. B.M. No. 8.8.7.99. Original number 1615.
Collected 25 November, 1907.
Although the original description is of a character to make
identification difficult, I have little doubt that this is the Roe
described by Noack t as C.[ervus]£ pygargus var. mantschuricus,
but this name, being preoccupied in the genus Cervus, was invalid
ab initio, and cannot be reinstated (as was done by Lydekker $),
whatever genus the animal is afterwards proved to belong to.
Under these circumstances I have particular pleasure in naming
it after the Society’s President, the Duke of Bedford, K.G., in
* Slightly distorted, probably below the normal size.
+ ‘Humboldt,’ viii. p. 9, 1889.
{ That the C. stands for Cervus and not Capreolus is clear from the fact that in
this paper the whole of the Cervide are included in one genus, the subordinate
genera being barely accorded the rank of “ groups.”
§ ‘ Deer of all Lands,’ p. 231, 1898.
646 MESSRS. RITCHIE AND McINTOSH ON [June 16,
recognition both of his carrying out of the present exploration, by
which our knowledge of the Mammals of the Far Hast is being
steadily revolutionized, and of the fact that his own personal
acquaintance with the Cervide and his wonderful collection of living
Deer at Woburn have been the basis of much of the considerable
nerease in our knowledge of the group which has taken place of
recent years.
“¢ A common deer about the edges of the forest at Chao-Cheng-
Shan. They were to be seen at all times of the day in groups of
two to five. Rarely were they solitary. During one long tramp
T saw fourteen in the day.”—M/. P. A.
2. On a Case of Imperfect Development in Lchinus escu-
lentus. By James Rrrcwmie, M.A., B.Sc., The Royal
Scottish Museum, and D. C. McIntosu, M.A., B.Sce.,
[Received May 7, 1908. ]
(Plate XX XIII. and Text-figures 138-142.)
The description of abnormalities is of special value when these
are of unusual character and occur in a species little lable to
deviation from the type. Moreover, there has not hitherto been
recorded any case of the special degree of abnormality illustrated
by our specimen. Therefore it is that we venture to set down
these observations, in spite of the fact that it seems impossible
to account with certainty for the origin, or even to determine
precisely the status, of the abnormality (whether it should be
regarded as an example of congenital variation, or simply as a
case of arrested development due to functional disturbance of the
organism by some external factor).
The specimen, an example of the most common British Sea-
Urchin (Hchinus esculentus Linn.), for which we are indebted to
Dr. A. Bowman, of the scientific staff of the Scottish North
Sea Fishery Investigations, was obtained by him, in July 1907,
in Basta Voe, Shetland, where it was trawled from a depth of
twenty-four metres. In a note regarding it Dr. Bowman says :—
“The malformed Urchin occurred amongst a number of typical
ones. Unfortunately I took no notice at the time of any pecu-
liarity in the living animal.... The sport was not noticed until
the spines etc. were nearly all cleared off. I thought at first it
was an unusually flat variety.”
DESCRIPTION OF SPECIMEN.
(a) General Description—Shape, Symmetry, &c.
At first sight the specimen appears to be, as Dr. Bowman had
noted, merely a rather flat variety, with a large oral surface and
* Communicated by F. A. Barner, D.Sc., F.Z.S.
+ For explanation of the Plate, see p. 661.
Diz. 8OOS) PIO
yy
Ke fam
1908.] AN ABNORMAL ECHINUS. 647
a depressed apical region. Closer investigation, however, shows
that the general shape has departed considerably from the normal.
Viewing the test in plan, one sees marked divergence from the
apparent radial symmetry characteristic of regular Sea-Urchins.
This is due to a distinct bulging on the side remote from the
madreporite, which has caused the ambitus to assume a bilaterally
symmetrical, almost oval shape (Pl. XX XIII. fig. 1). The same
portion, moreover, viewed in elevation, is seen to be considerably
depressed as compared with the globular form which charac-
terises the other regions (Pl. XX XIII. fig. 2). Further, it is to be
noted that the apical disc has departed from its normal horizontal
position, the madreporite standing at a distinctly bigher level
than the plates on the opposite side of the periproct, for these
appear to have been dragged downwards towards the bulging
portion of the test. On the oral surface the peristomal opening
is excentric, it too apparently having been dragged towards the
bulging portion, for in that region the margin of the opening is
only 29 mm. distant from the ambitus, whereas on the opposite
side the distance is 34 mm.
All those deviations from radial symmetry are due to, or at
least are connected with, the fact that a portion of one of the
ambulacra is absent. Orienting the specimen in the recognised
manner, by placing the aboral surface upwards, with the madre-
porite in the right anterior position, and adopting Lovén’s
notation, we find that the incomplete area is number V, the left
posterior ambulacrum, the tube-foot area of the left division of
the bivium. On the aboral surface this radial area is absent, but
commencing a little above the ambitus, at the bulging portion of
the test, it runs thence to the peristome, being fully represented
on the oral surface.
The most noticeable result of this partial cutting out of the
ambulacrum, apart from the general distortion of the whole
skeleton already described, is that the two sets of interambulacral
plates (areas 4 and 5) come together, four rows of interambulacral
plates thus occurring in close proximity. The two sets are
separated by a zigzag suture, whereas the edges of interambu-
lacral plates abutting against an ambulacral area (that is, the
edges corresponding to those bounding the above zigzag suture) are
normally straight. Less noticeable are such minor distortions
as the divergence of the line of bilateral symmetry, which passes
through the middle of areas IIL and 5, from its normal straightness,
owing toa bending of the interambulacral suture in the latter area
towards the locality of disturbance. Quite distinct as this devia-
tion is on the aboral surface, on the oral surface it does not exist,
the line through areas III and 5 being there perfectly straight.
Again, abnormal distortion occurs in the interambulacral sutures
of areas 4 and 5, and in the junction between areas 5 and LI., all of
these lines bending with gentle curves inwards towards the point
where the ambulacrum has disappeared. These curvatures are
obviously due to an increase in the size of the plates in the
648 MESSRS. RITCHIE AND McINTOSH ON. [June 16,
direction of their long axes, but, marked as the curves are, the
increase in the size of the plates is measurably of small moment.
The following comparative table indicates the differences in length
(i. ¢., in the direction of the long axis) and in depth (¢. e., at right
angles to the long axis) of certain interambulacral plates forming
a band, interrupted by the ambulacra, round the skeleton. The
1 So ge ||
oc | © >See
x | ee
= x x J tale se)
@ (S) j=) ||| =)
al i) aQ es
| 19 = = 8
= ~ we) zz 2
| 3 x x x 2+
a S ao Oo a
oq a a | mS
ie) | el
2 +B) c=
(eo) x ~ 3) 521
Ps) | x |! @w x 3 2
le ll aah 2
| — a ce | 2
| <i bee Ua —| 32
loa) 10 oD mr
co a) io “8
8 x x x eh
co i i=) R
a a a 2
wm
] Day
10 Ney 10 = 8
_ x x x Pos
fon) j=) i=) ae
| ce nN icy) co
cr) | BEERS
| a =
=~ Re) 2 eS eae
| 38 x x x =
jen) i=) im) Pe
4 = a a SB eo
ea 3 i esameahae eee B| ee aal Mh ia ete
< ] oO ie-a) ny.) —] @
= | eceelee FEE
o
no SW x I x |? 2a
19 19 N Si fl
S =) or = S
| ies N ce S a
i Oe Ie — Fs o
. 2
Nieescounl acs x 24
S Xx KP eee E'S 3
ren re) @ wm eS 2
re iors “ey iG} {S|
: +—— fe
© Sa &
xlele| 323
=) 19 x x eee aris
oO (=) S Dio iH
al a a 2422
! D a
as © 4 A
x Wo) Ro} 2
3 19 x x eS) Gs
co = So 8 =
— N eS | is a
= Sys
= = = = = = |
e mn 2 i ns 2
4 a | : : 25 5
<a] I 3 5 BA FS ee
= = = on a Hot
=] 4 co qo ra]
se) =
Sse i
| | “LVI g
missing ambulacrum is truncated opposite the thirteenth inter-
ambulacral plate, the plates being numbered from the peristome.
The thirteenth plate of each interambulacral area has, on this
account, been measured and along withit, forthe sake of comparison,
the plate which bounds it on the proximal and on the distal side ;
1908. ] AN ABNORMAL ECHINUS. 649
so that in the short series of three there are included the plate
formed immediately prior to the truncation (12), that formed
concurrently with the truncation (13), and that immediately
succeeding the truncation (14). The letters @ and 6 refex to the
plate-rows in an area taken in counter-clockwise rotation.
The above measurements indicate that, in those areas (4 and 5)
which bound the centre of disturbance, the plates formed con-
currently with the disturbance are on the whole slightly longer
than the corresponding plates in the other areas, while the plates
immediately preceding and immediately succeeding the disturb-
ance are on the whole shorter. But the depth of the plates in
the affected areas is in every case greater than the average depth
of the corresponding plates in the normal areas. The details here
recorded are of value as showing to what measurable extent
definite portions of the test have been affected in the effort of the
organism to adapt itself to highly abnormal conditions. This
phase of regulation will be referred to later.
The following measurements give some idea of the proportions
of the test in various directions :—Height = 55 mm.; long axis
= 95 mm.; short axis = 89 mm.; circumference at ambitus =
293 mm.
(b) Detailed Description—Abnormalities in Plates.
Examination of the elements which make up the test reveals
additional features of interest. The numbers of the plates in the
various interambulacral series are :—
TABLE IT.
Area ...... 1 2 3
|
Series ...... a | b Pyiecita lomo @| a
————s |! —_ SS SS ——— tl
(No of plates. .|| 23 | 24 || 25 i 24
}
TasBLE showing the number of interambulacral plates in the respective series.
The heavy line indicates the relative position of the area of disturbance.
In those areas which bound the incompletely developed ambu-
lacrum, all the series have suffered reduction of plates. Roughly,
they contain two plates short of the number normal for the
remainder of the test. Amongst the interambulacral plates there
is little abnormality other than the increase in length and breadth
in areas 4 and 5 already recorded, except in the two plates which
between them include the termination of the truncated ambu-
lacral area. These, instead of being rudely rectangular, are
pentagonal, the extra face abutting against the terminal plate of
the ambulacrum; and not only is the aboral half of each longer
650 MESSRS. RITCHIE AND McINTOSH ON [June 16,
than the adoral, but the portion facing the ambulacrum is con-
siderably deeper than that remote from it (text-fig. 142).
In the plates of the ambulacral areas there are more frequent
departures from the usual form. Fully-developed ambulacral
plates are formed by the union of small pore-plates, each bearing
a single pore-pair. In Hchinus esculentus three of these primitive
plates formed near the apical area, under the shelter of the
oculars, are compressed, by the formation of new plates, to form
a compound triad, the ordinary plate of the ambulacral area.
Even in the fused plate the original pore-plates can be distin-
guished by shallow ,boundary grooves; and we are following the
usual terminology in designating the two outer plates, which
are bounded on one side by the interambulacral area and on the
other by the zigzag intra-ambulacral suture, the adoral and aboral
primaries ; while the median plate touching the interambulacral
area but failing to reach the zigzag suture in its own area, is
known as a demi-plate. The three pore-pairs in a compound
plate are arranged, not in a single vertical series, but lie in three
distinct longitudes. These details of plate structure have been
recounted in order to facilitate reference to the abnormalities
which occur, and which consist, for the greater part, of an
imperfect complement, or an incomplete fusion of the primitive
plates which ordinarily go to the formation of a compound ambu-
lacral plate.
In the posterior series, a, of the right posterior ambulacral
area I (text-fig. 138) the twenty-eighth plate, numbered from the
Text-fig. 138.
ABNORMALITIES In AMBULACRAL AREAS oF Echinus esculentus.
Roman numerals beneath the figures indicate the ambulacral area in which the
abnormalities occur. Arabic numerals alongside the figures indicate the
numbers of the plates, reckoned from the peristome. a and 4, series in
ambulacral areas. In text-fig. 142 two interambulacral plates are included,
and are numbered according to their area, series, and position in series.
peristome, consists of only two complete primaries, a demi-plate
being lacking. It is succeeded by a solitary demi-plate, perhaps
the remains of the aboral of twenty-eight, the place of which may
1908. ] AN ABNORMAL ECHINUS. 651
have been taken by a fully developed median. Twenty-nine is also
formed of two primaries; while thirty, complete as regards the
number of plates and pore-pairs, possesses an arrangement
altogether unusual. The apparent adoral plate is an included
plate, for it touches the zigzag suture, but fails to reach the
straight suture between ambulacrum and interambulacrum; the
demi-plate is present, but instead of being median, it is external,
touching plate twenty-nine and lying in the same latitude as the
adoral plate; the aboral primary is normal. The pore-pairs of
the adoral and demi-plates are surrounded by a deep hollow
instead of by the usual faintly-marked peripodal groove. In
series 6 of area I the adoral plate of twenty-nine has no pore-
pair, but the median and aboralare normal. Plate thirty contains
only two single plates, a very deep adoral and an aboral demi-
plate. It is succeeded by a solitary demi-plate, the position of
the pore-pair of which would indicate that it represents the
missing aboral of the preceding plate.
In area IT (text-fig. 139), series a, the twenty-eighth plate is
formed of two simple individuals, probably an adoral demi-plate and
an aboral primary, but the obscurity of the sutures renders certainty
impossible. Plate twenty-nine is a single primary possessing no
pore-pair. It is succeeded by a solitary demi-plate, this again
being followed by a compound plate containing four elements, the
aboral of which lacks a pore-pair. In series b, twenty consists of
Text-fig. 139.
ABNORMALITIES IN AMBULACRAL AREAS OF Hehinus esculentus.
(For explanation of the lettering see text-fig. 138.)
a union of six primitive plates, no suture separating a first normal
triad from a second, twenty-eight consists of two primaries,
twenty-nine of only one, while between twenty-eight and twenty-
nine is wedged in a small insulated poreless individual. Succeeding
twenty-nine come two separate and distinct demi-plates, the distal
about half the size of the proximal; and these are followed by
thirty, possessing only an adoral primary and an aboral demi-plate.
In ambulacrum ITI (text-fig. 140) plate twenty-nine in series a
contains the normal number of plates, but the aboral is faintly
marked and is imperforate. The succeeding plate is also a
normal triad, but the peripode of the median element contains
only one pore, and must in its present condition have been
652 MESSRS. RITCHIE AND McINTOSH ON [June 16,
functionless. It is moreover the nearest to the centre of the
plate, whereas the median pore-pair should lie in the outermost
row. Series 6 contains two abnormal plates, twenty-nine and
Text-fig. 140.
Heat
ABNORMALITIES IN AMBULACRAL AREAS OF Hehinus esculentus.
(For explanation of the lettering see text-fig. 138.)
thirty, each composed of two pore-plates. The absence of distinct
sutures in the first renders identification of plates impossible, but
the second is composed of an adoral primary and an aboral demi-
plate. Thirty is an intercalated plate having no corresponding
individual in the adjoining ambulacral series.
Series a of area IV (text-fig. 141) contains but one abnormal
plate, thirty, which is composed of four primitive plates, three of
which appear to be primaries, only that preceding the aboral being
unmistakably a demi-plate. Of the four the adoral is imperforate.
Tn series 6, twenty-nine is composed of an adoral primary succeeded
by a demi-plate; thirty, of two primaries, the adoral without
Text-fig. 141.
Y 32
} Sla
31
5%) 30
» 29
ABNORMALITIES IN AMBULACRAL AREAS OF Hchinus esculentus.
(For explanation of the lettering see text-fig. 138.)
pores; thirty-one, of a single huge primary; thirty-one a, of a
separate demi-plate, perhaps an isolated portion of thirty-one ;
and thirty-two, of three plates, the adoral a large primary, the
other two, smail demi-plates crushed into the upper corner of the
compound plate. Thirty is an intercalated plate with no corre-
sponding individual in the adjacent row. Thirty-eight is also
unusual, beimg composed of five elements, the adoral and median
of which are primaries, while the remainder are demi-plates.
1908. ] AN ABNORMAL ECHINUS. 653
The aboral, however, almost reaches the zigzag intra-ambulacral
suture.
Ambulacrum V (text-fig. 142) is thearea the development of which
is incomplete, and here also abnormal plates occur. In series a the
twenty-ninth, or last plate of the series, is much deformed, for,
while it contains the usual triad, the median demi-plate is pore-
less ; while the adoral and aboral primaries are much misshapen,
the latter being roughly square and having its pore-pair lying in
Text-fig. 142.
Vv
ABNORMALITIES IN AMBULACRAL AREAS OF Echinus esculentus.
(For explanation of the lettering see text-fig. 138.)
a peninsula-like corner almost without the boundary of the
plate. In series 6 the penultimate plate, twenty-nine, contains
but two elements, an adoral primary and an aboral demi-plate.
It is followed by a large rudely-triangular plate whose apex falls
at the junction-line of the thirteenth plates of the adjoining
interambulacral areas. Hach of the terminal plates of j this
aborted area is bounded on two sides instead of on one by inter-
ambulacral plates.
The following table shows at a glance the relative positions of
these abnormalities with regard to the respective areas in which
they occur :—
Taste ITT.
| Avea..e sc. sees: [ Il ST eal ay en Vi
Series ......... a) Bila |b || a \ 6 | @ Was Nb
|< ee — —______—__| | —____ ~ || ——]——_
| Total no. of |
plates. |
No. of first plate i | | |
in each | 28
abnormal group. | \| | |
Summarising these observations regarding the ambulacral areas,
we find that, of the ten rows of ambulacral plates grouped in the
654 MESSRS. RITCHIE AND McINTOSH ON [June 16,
five ambulacral areas, not a single row is free from more or less
marked abnormality. Further, all the abnormalities, with two
exceptions, are grouped in a band, broken by the interambulacra,
which passes round the test at a definite distance from the
peristome. The exceptions are the twentieth plate in II } and
the thirty-eighth in IV 6.
With regard to the apical disc as a whole there is little worthy
of note. The plates are normal in number and arrangement, but
the whole disc has become slightly elongated as if the part
towards the abnormal area had been dragged downwards by it.
Consequently several of the genital plates have lost the bilateral
symmetry which usually characterises them. ‘The ocular plate
corresponding to the abnormal ambulacrum is of unusual shape,
possessing four, instead of five, sides and presenting an angular,
instead of an almost straight, boundary to the corona. The
ocular pore is absent, but its position is probably indicated by a
minute pin-hole, which fails to penetrate to the inner surface of
the plate, for neither is there any sign of an internal opening,
nor can a strong light pass through.
PROBABLE DEVELOPMENT OF THE SPECIMEN.
To bring those observations into relation to one another, the
most satisfactory way is to trace the probable development of
the shell. It is with the idea of attaining an approximation
to chronological sequence that the plates have throughout been
reckoned from the peristome, and not from the apical termination
of the series to which they belong. This mode of reckoning
has the disadvantage of increasing the difficulty of numerical
determination owing to the excessive compression of plates which
takes place as the peristomal region is approached, but it has the
advantage of following the natural course of development. For
it is evident that, since all the coronal plates are formed around
the margin of the apical disc and are pushed thence down the
sides of the test, the oldest plates will lie around the peristome.
Thus, counting from the oldest plates recognisable towards those
more recently formed, we get a measure of the age of the animal
computed according to a standard, not of time but of develop-
ment *. Thus, instead of saying that when a certain plate was
formed, the test was three months old, a statement which our
ignorance of the growth of the Echinoid imago renders impossible,
we can say that at that time the test was, say, ten plates old, the
actual age of course being indicated by the formula 10+, where
« represents the number of the plates which have been pushed
over the edge of the peristome in any one series. But since we
* Such a measure, it need scarcely be said, is not absolute but comparative, for the
first plates, and we know not how many of their successors, have already been
pushed over the edge of the peristome and are no longer reckonable. Assuming,
however, that in each series the rate of pushing over is approximately the same, we
arrive at a measure sufficiently accurate for all practical purposes.
1908. | AN ABNORMAL ECHINUS. 695
can assume that a is approximately the same for the various
ambulacral plate-rows of the same specimen at any latitude, it is
virtually a constant for a particular latitude and therefore cannot
affect our comparison.
In this Shetland specimen the development, up to a certain
stage, appears to have been normal, Thus on the oral surface
not only is the arrangement of the various rays regular, but
the line of bilateral symmetry is straight; while in the minute
structure of the plates no abnormalities occur, save the single
insignificant deviation on area IT 6, where, although the plates
are normal in number and in arrangement, a suture is missing
between two triads. We are also justified in stating that during
the earlier stages of growth the ocular plate opposite the abnormal
ambulacrum was perforated by an ocular pore, and that this pore
was occupied by the terminal tentacle of the radial water-vascular
system, for in no other way can the presence of the pin-hole
already mentioned be explained, seeing that in the ordinary
course of development the very existence of the pore is due to
the presence of the terminal tentacle *
At acertain stage, when rather more ‘than twenty-eight (28+ 2)
ambulacral plates had been formed, or, judging from young speci-
mens with a similar number of plates, ‘when the test was between
20 and 25 mm. in diameter, some functional derangement took
place. As an immediate consequence ambulacrum V ceased to
grow, no more plates being added to that area after the thirtieth.
But a more general disturbance also occurred, for in each of the
rows of the five ambulacra abnormal plates were formed; and in
these groups of aberrations, containing sometimes a sequence of
as many as five peculiar plates, the first abnormal plate, as a
glance at Table III. will show, is the twenty-eighth or the
twenty-ninth or, in a solitary case, the thirtieth. This approxi-
mation of numbers indicates, as we have already shown, that the
plates were formed approximately at the same stage of develop-
ment; and the significance of the close numerical correspondence
between the commencing points of the abnormal series is not
lessened when we consider the difficulty of counting the number of
plates at the edge of the peristome, and the uncertainty as to the
relative numbers that have been pushed off during development.
In themselves, considered separately, the abnormalities described
are perhaps of little significance, although we have been unable
to find, from examination of other tests, that such abnormalities
are of frequent occurrence. But that abnormalities so distinct
should manifest themselves at all points of the test at practically
the same period is indeed remarkable. There can be but one
explanation, namely, that a general derangement affecting all
* “Die Primordialtentakel] werden endlich von den sich bildenden Radial-
(Ocellar-)Platten deren Rand sie beriithren, umwachsen und sind so zu den
Terminal- oder Endftihlern geworden.... Sie treten dann durch einen Porus der
Platte hindurch.” Hamann, Otto, “Die Echinodermen”; Bronn’s Klassen und
Ordnungen des Thier-Reichs, ii. Bd., 3 Abth, p. 1167.
Proc. Zoou, Soc.—1908, No. X LIT. 42
656 MESSRS. RITCHIE AND McINTOSH ON [June 16,
the ambulacral areas took place about the time that ambu-
lacrum V ceased to grow. The derangement was in most cases,
however, only a temporary one, for in all the areas, save No. V,
the elasticity of the organism appears to have overcome the
functional disorder, and after the formation of a few unusual
plates, the normal srowth was resumed. Nevertheless, a slight
indication of after-eftect is afforded by differences in the size of
the tubercles, for a cursory glance shows that they are smaller
above the affected zone aii, below it. This variation, however,
is somewhat discounted by the fact that even in normal speci-
mens a similar, though less marked, difference in size exists
between the tubercles above and below the ambitus *.
The disappearance of the two rows of ambulacral plates from
area V rendered necessary considerable modifications in the test,
and this regulation was carried out mainly in two ways. The
plates of the adjoining interambulacral areas increased a little
in length and considerably in depth, and closing in around the
truncated area came together in the mid-line, where they were
united by a zigzag suture. As a direct result of the mcrease of
the plates, the sutures in the neighbourhood became distorted.
The increase in the size of the interambulacral plates, however,
was not sufficient to make up for the loss of a double row of
ambulacral plates measuring over 15 mm. across, hence another
modification became necessary in order that the space between
the adjoining areas on each side might be spanned. This was
brought about by the plates passing directly across the space
instead of building a material-wasting globular dome, the result
being evident in the flattened portion of the test which lies
between the truncated ambulacrum and the apical disc.
The increase in the depth (that is, direction of short axis) of
the plates, and the flattening of the surface of the shell have
together had the effect of pushing the incomplete ambulacrum
further from the apical area, so that it has come to form the
centre of a distinct bulge in the outline of the test, while it has.
also given rise to an abnormally flattened area on the oral surface.
It was perhaps at this period of disturbance that the terminal
tentacle disappeared and that fresh deposits of calcareous matter
began to close up the unoccupied ocular pore.
RELATION TO PREvyiousLY DESCRIBED CASES OF A
SIMILAR CHARACTER.
Mr. W. Bateson has brought together the cases of abnormality
in the major symmetries of Echinoids recorded prior to 18947.
The remaining records up to 1902 are mentioned by Hamann ¢.
* F. Jeffrey Bell: ‘Catalogue of British Echinoderms in the Brit. Mus. (Nat.
Hist); p. 152: London, 1892,
eke Bateson, W., ‘ Materials for the Study of Variation,’ pp. 441 e¢ seg., London,
94:
ay Hamann, Otto, 7. e. p. 1293.
1908. ] AN ABNORMAL ECHINUS. 657
We can find no later reference to new descriptions of abnor-
malities of a similar nature in major symmetries, although two
early cases are mentioned by Gauthier * which Bateson appears to.
have omitted. The first, a specimen of Hchinobrissus orbicularis,
is described by Cotteau? as having the anterior ambulacrum
completely atrophied ; the second, a Pyrina ovulum, in which the
right posterior ambulacrum was wanting, has also been described
by Cotteau £.
The majority of the cases cited belong to fossil forms, this
being no doubt due to the greater readiness with which an
abnormality may be detected in a clean fossil test than in a
recent well-preserved specimen, where plate-groupings are
obscured by epiderm and spines, The cases to which the present
example bears closest resemblance are those grouped by Bateson
in his class (2), wherein the specimens are distinguished by
the “ partial or total disappearance of a definite ambulacrum or
interambulacrum.” At first glance the parallel between the
Shetland specimen and the Hchinus melo described by Philippi §$
appears to be almost complete, but in that case, as in the specimens
described by Bell ||, Chadwick 4], and Osborne**, the defaulting
member constitutes a complete morphological system, the homo-
logue of an Asteroid ray, whereas in the present specimen only
the ambulacral portion of a ray has suffered reduction. There
is a much closer resemblance to specimens of Hemiaster, described
by Gauthier 7?, in which only the ambulacral portion of a ray has
disappeared. Of those specimens the case of Hemiaster batnensis,
No. I, appears to show the closest analogy. There the corre-
sponding ambulacrum, the left posterior, has partly ££ disappeared,
having at a certain stage received a check in development,
the stage being indicated by the dying out of the ambulacral
pores and by a slight depression in the test. As a con-
sequence four sets of interambulacral plates follow one another
without interruption, and the posterior interambulacral suture
* Gauthier, M. V., “Sur quelques Echinides monstrueux appartenant au genre
Hemiaster.’ C. R. Assoc. Franc. pour Vavanc. des Sciences, 13th Sess., 1884.
(Paris, 1885), p. 259.
+ Cotteau, G., Echinides nouveaux ou peu connus, 1862, p. 66, pl. ix.
t+ Cotteau, G., 7. c., 1867, p. 133, pl. xviii.
§ Philippi, F. W., Arch. fur Naturg. iii, 1837, p. 241, and plate.
|| Bell, F. Jeffrey, Journ. Linn. Soc. (Zool.), xv. 1881, p. 126, and plate.
@ Chadwick, H. C., “Note on a Tetramerous specimen of Hehinus esculentus’’.
Trans. Liverpool Biol. Soc., 1898, p. 288, and plate.
** Osborne, H. L., “A case of variation in the number of ambulacral systems of
Arbacia punctulata”’: American Naturalist, vol. xxxii. 1898, p. 259, and figs. (The
reference to Lang’s ‘Comparative Anatomy,’ vol. i, p. 321, is evidently a slip for
. 341.
: +r wadek, M. V., 1885, J. c., p. 258, and plate.
tf Bateson seems to err in placing this example among those in which “one
ambulacrum is wholly wanting in the affected radius” (/. c. p. 443); for while the
functional part of the ambulacrum is not represented on the test as found, yet in
the earlier stages of development the ambulacrum apparently did exist, for beyond
the point where the ambulacral groove ought to run “ apparaissent quelques paires
de pores arrondis, qui continuent l’aire ambulacraire de l'autre cété [that is, the
oral side] du fastiole.” Gauthier, 7. c., p. 259. ee
6
658 MESSRS. RITCHIE AND McINTOSH ON [June 16,
has become somewhat distorted. As in our case also, the
complement of apical plates is perfect, while the ocular pore
(Bateson says merely “ ocular”) corresponding to the imperfect
area is absent. There is therefore a remarkable correspondence
between the two examples.
PossIBLE ORIGINS OF THE ABNORMALITIES.
In the majority of such cases as have been described, authors
have made no reference to the probable origin of the abnormality.
Gauthier dismisses the case of Hemiaster batnensis, above mentioned,
with the rather depreciatory remark, “il ne présente qu’une
simple atrophie.” Bateson in his remarks prefacing the summary
of the Echinoderm variations, says that “it cannot be doubted that
the variation[s]seen in Echini... are truly congenital. Similarly,
though in Asterias, &c., reduction in the number of arms might
otherwise be thought to be due to mutilation, it cannot be so in
KEchini.”* But while the majority of the abnormalities appear
to be congenital, so sweeping a statement must be avoided, since
it would preclude our even considering the possibility of reaction
to immediate external influences. For, although in the meantime
we cannot definitely point to any member of a major symmetry
which has demonstrably suffered alteration through external
factors, the occurrence of such alteration is not at all improbable,
considering the extraordinary sensitiveness of Sea-Urchins to
unusual conditions of environment *.
Hamann, realising the difficulty of confining attention to only
one of several possible causes, says, in his résumé of the form-
abnormalities in Echinoids, that, should the aberrancies not be
due to discontinuous congenital variation, their origin might
be set down to loss and subsequent regeneration or to fusion {.
Renamed in accordance with this conception of the potential
influence of external factors, Class (2) of Bateson becomes,
according to Hamann, that of incomplete regeneration (‘ unvoll-
stindige Regeneration”). But even this conception confines
the possibilities within far too narrow limits. ‘ Loss” implies
the previous existence of some part which disappears, and it
* Bateson, 1894, 7. c. p. 433.
+ Lo Bianco states that on the coast-line, where, previous to the 1906 eruptions
of Vesuvius, thousands of Echini had been scattered on the rocks, not a single live
specimen could be found subsequent to the ash showers. None of the other marine
invertebrate groups mentioned by Lo Bianco suffered to the same extent as the Sea
Urchins. Lo Bianco also demonstrates that in the case of the artificial introduction
of ashy material into a vessel containing Hchini, the Echini had already begun to
putrefy on the morning of the third day after the experiment began, while two
days later the organs were completely macerated and the spines had fallen off.
The rapidity with which the Sea Urchins succumbed shows sensitiveness to derange-
ment of function. Lo Bianco, Salvatore, “Azione della pioggia di cenere, caduta
durante Veruzione del Vesuvio dell’ Aprile 1906, sugli animali marini”: in
Mittheil. Zool. Stat. Neapel, Bd. xviii. Heft i. 1906, pp. 91 e¢ seq.
£ “Wenn die Abnormitiaten nicht sprungweise congenitale Varietiiten sind so
kame fiir ihren Ursprung Verlust und nachtragliche regenerative Processe . .
-oder Verschmeltzung in Betracht.” Hamann, O., J. c., p. 1294.
ee
1908. ] AN ABNORMAL ECHINUS. 659
is almost impossible to conceive that, in a Sea-Urchin where the
morphological systems are welded together into a united whole,
a definite area could be lost in part, in the way that the arm of
a Starfish could be lopped off. It is possible to conceive, however,
that damage to the growing point of an area might check,
temporarily or permanently, the growth of that area. More-
over, it is not necessary to suppose that the damage be followed
by regeneration, although test-regulation must be an almost
inevitable consequent. We assume that, since the whole question
is one of major symmetries, the regeneration referred to by
Hamann is regeneration in a major symmetry as a whole (7. ¢., the
equivalent of the regrowth of the arm of a Starfish) and not the
insignificant substitution of new plates for broken ones, a form
of regeneration the comparative triviality of which is better
indicated by the term replacement (the réparation of Prouho).
On account of these difficulties the phrase “‘ incomplete re-
generation ” cannot be taken to comprehend the connotation of
Bateson’s Class (2), for neither loss nor regeneration is an essential
agent in producing such results as are included in that class. As
a wider designation and one which seems to include most of the
possibilities, we suggest arrested development.
Congenital variation—facile phrase—might well account for
the original abnormalities, and test-adaption for the subsequent
and consequent distortions. But as an alternative to congenital
variation, reaction to immediate external influences appears to
offer an explanation as probable and more simple. Although lack
of experimentation renders conjecture somewhat hazardous, it is
possible that some voracious enemy * or some wave-borne rock
fragment might break not only the newly-formed and extremely
delicate plates at the apical end of an ambulacral area, but along
with them might damage, either indirectly, by destroying the
controlling nerve, or directly, that portion of mesenchyme in
which the ambulacral plates are built up. The check to further
development received by the ambulacrum would give an oppor-
tunity to the interambulacral plates on each side to push outwards
and usurp the position hitherto occupied by the ambulacrum ;
and even if new generative mesenchyme were thereafter to be
regenerated in the old ambulacral position, it is conceivable that
by an adaptation of function these new plate-forming cells might,
instead of forming new and independent plates, reinforce the
interambulacral-forming cells, by depositing their calcareous
material along the edges of the interambulacral plates. Such
reinforcing power would account for the greater depth which
characterises the interambulacral plates between the abnormally
truncated ambulacrum and the apical disc. That in such a case
* Pyouho has observed young Mullets not only snatching off the spines but even
raising the epiderm on the surface of Dorocidaris papillata. Such a wound is
sufficient to cause the underlying plates to be thrown off and replaced. Prouho, H..,
“Recherches sur le Dorocidaris papillata et quelques autres Hchinides de la
Mediterranée”: in Arch. Zool. Expér. ser. ii, vol. 5, 1887, p. 250.
660 ON AN ABNORMAL ECHINUS. [June 16,
broken plates would remain to demonstrate the occurrence of an
accident is not to be expected, for the researches of Prouho*
show that, in Dorocidaris papillata at least, broken plates are
rejected and are replaced by new ones. All therefore that one
could expect to find in place of the plates originally damaged
would be plates irregular in outline and in structure, and such
exist in our specimen at the truncated end of the ambulacrum
(vide p. 653).
Tt is more difficult with a particular-accident explanation to
account for the abnormalities which occurred on all sides of the
test simultaneously with the destruction of the ambulacrum. A
serious accident to any animal is followed by a general loss of
vitality, and such a loss is very likely to be reflected in those
parts which are in most delicate equilibrium, that is, in the
parts where growth is taking place. Of the thirty abnormal
plates which occur in the band comciding with the truncated
ambulacrum, twenty-two are deficient in the number of primitive
plates of which they are composed, while of the remaining eight,
four are deficient as regards the possession of pore-pairs. These
deficiencies agree well with the idea of loss of vitatility, for they
seem to indicate that at that period, primitive plates were formed
with less than the usual rapidity, the result being that owing to
the decrease of pressure from newly-formed plates, only two
pore-plates, instead of three, were compressed to form a compound
ambulacral plate.
An explanation of the abnormalities in the Shetland specimen
such as that given above is of necessity largely conjectural.
Whether it altogether meets the requirements of the case must
be left to the decision of experiments, in which the natural
conditions surmised are reproduced as nearly as possible by
artificial methods. .
SUMMARY.
The imperfect development of this: test of Hehinus esculentus
is expressed in :—
(a) General shape.—Flattened, with a bulge at one portion of
the ambitus, and above the bulge a depressed area of the corona.
(b) Major symmetries.—The left posterior ambulacrum does
not exist after its twenty-ninth plate (counting from the
peristome). The corresponding ocular plate is present, but its
shape is unusual and its pore is a mere pin-hole.
(c) Plate details—On the twenty-eighth, twenty-ninth, or
thirtieth plate of each of the ten ambulacral rows a series of
abnormalities begins, the abnormalities consisting, for the
most part, of deficiencies in the number of primitive plates
forming a compound plate.
Less marked divergencies from the normal can be detected in
the sizes of some of the plates and in the irregular courses of
certain of the sutures.
* Prouho, H., 7. ¢. p. 251.
P.Z.S. 1908. Pl 2oScive
Bale & Danielsson, Ltd.
SPICULES OF CALCAREOUS SPONGES.
P.Z.S. 1908. Pl. Xxoaye
is sah la EAE OEE
Bale & Danielsson, Ltd.
SPICULES OF CALCAREOUS SPONGES.
eS
P.Z.S. 1908. Pl. XXXVI.
SPICULES OF CALCAREOUS SPONGES. .
PYZS5 1808, ill, KOORWILL.
18
Bale & Danielsson Ltd.
SPICULES OF CALCAREOUS SPONGES.
1908. | ON THE SPICULES OF CALCAREOUS SPONGES. 661
The stoppage of the growth of the ambulacrum and the plate
abnormalities occurred approximately at the same time, and it is
suggested that they may be due to functional disturbance caused
by some external agent. The distortion of the test subsequent
to the arrested development of the ambulacrum has been brought
about by a process of regulation.
The specimen of Hchinus esculentus above described has been
deposited in the Royal Scottish Museum, Edinburgh.
EXPLANATION OF PLATE XXXIII.
Fig. 1. Test of abnormal Hehinus esculentus viewed in plan, natural size.
Fig. 2. Test of abnormal Hehinus esculentus viewed in elevation, natural size.
Lettering :—m., madreporic plate; ¢.p., plate which terminates Area V; the
remaining symbols indicate the various areas according to Lovén’s system.
3. Observations on the Minute Structure of the Spicules of
Calcareous Sponges. By E. A. Mincuty, M.A., V.P.Z.S.,
Professor of Protozoology, University of London, and
DA Se Ein, Vee Oxvies ZS:
[Received May 12, 1908. ]
(Plates XXXIV.-XXXVIL.*)
Introduction.
The minute structure of calcareous sponge-spicules has been
the subject both of much laborious investigation and of many
contradictory statements. In regard to the structure of siliceous
sponge-spicules investigators are practically agreed upon the
following points :—the mineral matter of the spicule, or spicule-
ray, aS the case may be, forms a hollow tube with a relatively
thick wall; in the lumen of the tube, termed the axial canal, is
lodged an axial filament of organic nature; the siliceous tube
may be homogeneous in structure or may be stratified, that is,
composed of concentric layers of silica alternating with fine layers
of organic material ; and the outer surface of the siliceous tube is
enveloped in an outermost layer of organic substance forming a
sheath to the whole spicule. Thus in siliceous spicules we find,
apparently universally present, the following parts, passing from
within outwards :—(1) the axial filament, a definite structure
that can be isolated by hydrofluoric acid ; (2) the siliceous tube,
stratified or homogeneous; (3) the spicule sheath. Turning now
to calcareous sponge-spicules, it is found that the only point on
which all investigators are agreed is the presence of a sheath
enveloping the surface of the spicule. The following brief
historical summary of the statements that have been put forward
will make this clear.
* For explanation of the Plates, see pp. 675-676.
662 PROF. MINCHIN AND DR. REID ON THE [June 16,
Historical Review of the Question.
Grant (1826 *) pointed out that in certain sponges the skeleton
is calcareous, consisting of carbonate of lime, and exhibiting no
trace of phosphate of lime. In Spongia (Grantia) compressa he
described the rays of the triradiates as ‘“ hollow within, shut at.
their free extremities, and having no superficial openings; but
their internal cavities communicate freely at their point of junc-
tion and form there a small central reservoir.” The monaxons
are described as “ distinctly tubular and shut at both extremities.”
Of the triradiates of S. (Leuconia) nivea he remarks, “their
internal cavities are very distinctly seen.”
Schmidt (1862), on the other hand, included the spicules of
calcareous sponges in that category of spicules in which both
central canal and lamination were entirely wanting. He considered
it beyond doubt, however, that organic substance takes part in
building up calcareous sponge-spicules, since heating produces.
small vesicles in them.
Bowerbank (1864) described the effects of heat on siliceous.
and calcareous sponge-spicules, and concluded that the latter
contained so great a proportion of calcareous matter as to prevent
their disintegration by heat. He stated the concentric stratifica-
tion to be visible in the transverse fractures of any spicule,
calcareous or siliceous.
Kolliker (1864), on the other hand, was unable to find any
stratification or other internal structure in calcareous sponge-
spicules, and considered it doubtful if they contained any organic
matter, since no residue was detected if the spicules were dissolved
with acids. He described the spicule-sheaths in “ Vardoa spon-
giosa” (probably a synonym of Clathrina contorta, vide Minchin,
1898, p. 533, footnote, and P. Z. 8. 1905, ii. p. 17). Im siliceous.
sponge-spicules, however, Kolliker observed and described, in
detail, the axial filament and the stratification of the silica.
Lieberkiihn (1865) observed in the gastral rays of the quadri-
radiates of Lewcosolenia “a fine layer of the contractile substance,
which protrudes between the ciliated cells and either envelopes:
the spicule partially or completely asa fine layer, or only surrounds
the foot of it as a stronger thickening (Anhiufung).” These
sheaths were left behind when the spicule was dissolved with
acetic acid; they were considered by him as retractile. From the
description it is evident that the structures observed by Lieberkiihn
were really the cellular sheaths or gastral actinoblasts enveloping
the projecting gastral rays, and not the true spicule-sheaths.
Carter (1869) stated that the spicules of Grantia ciliata differ
from those of siliceous sponges in lacking a central canal; but in
his later note of the same year, he modified this statement and
admitted that something like a central canal may often be seen
towards the base of the straight arm of a triradiate; while for
the most part there is no trace of a central canal nor of the
* For bibliography, see pp. 674-5.
1908. ] SPICULES OF CALCAREOUS SPONGES. 663
concentric lamination seen in siliceous spicules, although both
siliceous and calcareous spicules break with a conchoidal fracture.
He points out that a central canal is very obvious in siliceous
spicules, “whereas in the calcareous one you can only fancy its
existence here and there.”
Haeckel (1872) described the structure of calcareous sponge-
spicules in a most detailed manner. The spicule-sheaths were
stated by him to be structureless envelopes arising as a thickness
of, and separation from, the ground-substance ; which, it must be
remembered, Haeckel regarded as a syncytial mass of protoplasm
or “sarcodine” formed by fusion of cells, and not as a secreted
gelatinous mesogloeal layer, as it is now universally held to be.
Haeckel contradicted Lieberkiihn’s statement that the sheaths
were retractile. Hach spicule or spicule-ray, according to Haeckel,
consists of a system of numerous very thin concentric lamine,
having the form of hollow cylinders or cones, surrounding a
common axis, a very fine central filament. The stratification and
the axial filament were stated to be universally present, though
the filament was much finer and more difficult to make out than
in siliceous spicules, and sometimes not visible unless the spicule
were moderately heated, whereby the axial filament was caused to
turn brown and then became visible by obliquely transmitted
light. The axial filament was stated to run up to the tip of the
spicule and become continuous with the “ sarcodine,” from which
it scarcely, if at all, differed in chemical nature. At the centre
of triradiates and quadriradiates a small spherical hollow was to
be seen, in which the axial filaments unite.
The spicules were stated by Haeckel to consist of calcium
carbonate and water together with a varying amount of organic
matter, termed by him “ spiculin,” which was left behind as a
colourless and structureless residue when the spicule was dis-
solved by weak acids. The spiculin substance did not stain in
carmine, lodine, &c., and was dissolved by caustic alkalis; its
presence was stated to be best demonstrated by moderately
heating the spicules. By the amount of spiculin present the
spicules could be placed in a series with two extreme types, the
one poor, the other rich in spiculin. The two extremes were
stated to be distinguishable at first sight under the microscope,
the spicules rich in spiculin appearing darker, more refractile than
those poor in spiculin, which were dull and pale in appearance.
The phylogenetically older forms of spicules, that is to say, the
monaxons of simple form and the regular triradiates, were stated
to contain least spiculin; the phylogenetically younger forms of
spicules, such as the sagittal triradiates, possessed most spiculin.
Lendenfeld (1885) stated that spicules of calcareous sponges
consisted of carbonate of lime mixed with organic substance ;
by treatment with gold-potassium chloride the spicule was shown
to consist of “a great number of small prisms, parallel to one
another, radiating from the axis,” which was “a cylindrical cord
of organic matter without lime.” The oldest part of the spicule
664 PROF. MINCHIN AND DR. REID ON THE [June 16,
was stated to contain more organic substance than the younger,
2. €. outer parts.
An entirely new epoch in the study of calcareous sponge-
spicules was inaugurated by the elaborate and exhaustive inves-
tigations of Ebner (1887), by whom and by Sollas, independently,
the peculiar crystalline nature of these spicules was discovered ;
namely, the fact that each spicule, whatever its form, behaves
optically like a single crystal of calcite. As regards the minute
structure of the calcareous spicules, Ebner’s results differ totally
from those of Haeckel. Ebner was unable to find any residue
after dissolving the spicules with various acids, and his attempts
to demonstrate any such organic residue with stains gave negative
results. He explained the optical differences between the two
types of spicules, described by Haeckel as rich and poor in
spiculin respectively, by the fact that in the regular triradiates
the crystalline optic axis is vertical while in sagittal forms it is
inclined or even horizontal, in the facial aspect of the spicules.
Ebner pointed out further that the “ browning” of the spicules
produced by heating is not due to the formation of carbon
through charring of organic substance, but is due to the disen-
gagement of fine bubbles of gas in the substance of the spicule
(compare Schmidt), making it opaque by transmitted light, milk-
white by reflected light; with stronger heating the gas breaks up
the spicule with decrepitation. Ebner found, however, certain
differences between calcareous sponge-spicules and pure calcite,
and analysis showed the presence of magnesium, sodium, and
sulphates, as impurities mixed with the calcite comprising the
spicules.
According to Ebner the axial filament of Haeckel is due to the
axial portion of the spicule having a different composition to the
peripheral portion, rendering the axis more easily attacked by
acids, by the action of heat, &c.; but the difference between
axis and periphery is a gradual and quantitative, not a sharp
qualitative contrast. Ebner found the alleged stratification to
be present only in a few spicules, mostly very large forms. He
studied the stratification of the huge triradiates of Leucaltis
solida, and the large monaxons of Leucandra aspera and L. alci-
cornis, and found it also due to a special distribution of more, or
less, decomposable substances in different parts of the spicule.
Thus the appearance of an axial filament and of stratification are
both due to a similar cause, namely, the periodic deposition of
more, or less, pure calcite in the building up of the spicule.
Ebner concluded from his observations that the spicules of
Calcarea are mixed crystals consisting chiefly of calcite without
organic substance, but containing inorganic impurities (Na, Mg, 8S,
probably also water), and that the conditions of the mixture
differ at different periods of the growth and in different part sof
the spicule.
_Lendenfeld (1891), while quoting Ebner’s results, reiterated
his former statements to the effect that ‘“ Each spicule-ray
1908. ] SPICULES OF CALCAREOUS SPONGES. 665
consists of a thin, somewhat turbid (ériib) axial thread, enveloped
by the hyaline (glashell) spicule-substance. In the latter a con-
centric stratification round the axial filament is to be recognised ”
(2. c. p. 369).
Minchin (1898, p. 569) stated that the spicules of Clathrina
coriacea, 1f treated with picro-nigrosin (saturated solution of picric
acid in water, 9 volumes; 1°/, nigrosin in water 1 vol.), showed
the sheath and the axial filament, left behind after solution of the
calcareous matter, and stained blue by the nigrosin.
Biitschli (1901) published elaborate investigations upon the
minute structure of siliceous and calcareous sponge-spicules,
taking the large monaxons of Leuwcandra aspera as examples of
calcareous spicules. His results were, in the main, confirmatory
of Ebner’s. He found that moderate heating produced a finely
alveolar structure in the spicule. No axial canal or filament was
found to be present; the axial thread, sometimes visible after
moderate heating, was stated to be due to a modification of the
calcareous substance, which at the central part of the spicule was
distinguished from the remaining part by greater refractility and
by being more easily attacked by acids. No trace of an axial
filament or sheath was found when spicules were dissolved by
acetic acid. The stratification was ascribed to the concentric
arrangement of the minute alveoli composing the spicule.
Unlike Ebner, Biitschli found small quantities of organic
substance left when the spicules were dissolved with hydrochloric
acid. The spicule-sheaths were found to be not purely organic
in nature but to contain a certain amount of inorganic matter,
probably calcium carbonate.
Maas (1904) and Weinschenk (1905) agree in assuming that
the calcite of the spicule must be combined with finely divided
organic substance. Maas explains the solvent action of caustic
alkalis on the spicules by the supposition that the caustic attacks
a substance holding together the constituent particles, and so pro-
duces a disaggregation of calcareous elements which were formerly
crystallographically orientated. He considers the phenomena
seen on heating to be explicable also on the assumption of a
finely distributed organic material in the spicule ; and he showed
that when the sponge is grown in water deprived of CaCo, the
organic substratum alone of the spicule is secreted. Weinschenk
dwells on the differences between calcareous sponge-spicules and
pure calcite, and considers these differences, and also the peculiar
form of the spicules, explicable only by the presence of a fine
organic tissue in the spicules.
Biitschli (1906) controverts the statements of Weinschenk
upon certain points which are of secondary importance for the
subject of this memoir, and maintains his former position. He
denies that the opacity and decrepitation produced by heating is
due to the presence of organic matter in the spicules.
It is seen from the foregoing that opinions are greatly divided
upon the question of the structure and composition of calcareous
666 PROF. MINCHIN AND DR. REID ON THE [June 16,
sponge-spicules, and especially with respect to the amount of
organic matter present in them. While Haeckel, Lendenfeld,
Maas, and Weinschenk assert or assume the presence of con-
siderable quantities of organic substance in the spicules, Kollhker
and Ebner allow none at all except in the sheath, and Biitschli
admits the existence only of a trace of organic matter and
considers even the sheath to be largely inorganic. It should be
further pointed out that the two most detailed investigations
upon this question, namely, those of Ebner and Butschli, were
based, so far as the composition of the spicules is concerned, on a
very limited number of forms; Buitschli, in fact, studied only the
large monaxons of Leucandra aspera. Hence there is a possibility
that their investigations do not cover the whole range of varia-
tion that those spicules may present. Both Ebner and Butschli
worked at the largest types of spicule that they could obtain, in
order to facilitate the handling and treatment of the material.
Observations upon Calcareous Sponge-spicules.
Our investigations have been directed towards endeavouring to
demonstrate the existence of a residue after decalcification, by
means of specific stains, as Ebner attempted to do, but without
success. It is obvious that if any organic residue were left after
decalcification, it might be expected to have an affinity for certain
stains and not for others, and might therefore be demonstrable
only by means of particular dyes, so that the negative results
obtained by Ebner would not necessarily disprove the existence
of an organic residue. We obtained in all cases positive results
with nigrosin, as stated previously by one of us (Minchin, 1898),
and also with the allied stain indulin. In our investigation we
have made use chiefly of the spicules of Clathrina contorta, but
we have examined the spicules of several other species: of Clath-
rinide, Clathrina clathrus and Ascandra falcata; of Leuco-
solenude, Leucosolenia lieberkiihnii and ZL. complicata; and of
Heteroceela, Sycon ciliatum, Leucandra aspera, and Heteropegna
nodus-gordii.
Our method of procedure was as follows. A piece of the
sponge taken from a specimen preserved in alcohol was washed in
water and placed in a tube of a small hand-centrifuge with a few
drops of Eau-de-Javelle, and gently shaken. In a short time,
generally about half-a-minute, the sponge is dissolved into a
cloud of spicules. The tube is then filled up with distilled water
and shaken up, and then with the centrifuge the spicules are
driven down to the bottom of the tube. The liquid is then
carefully poured off, taking care not to disturb the spicules, the
tube is filled up again with water, shaken up, and the process
repeated. In this way the spicules can be given three or four
washings in as many minutes, and are freed both from organic
matter of the sponge-body and from the Hau-de-Javelle. The
next procedure was usually to add to the tube containing the
1908. ] SPICULES OF CALCAREOUS SPONGES. 667
spicules in distilled water a few drops of ordinary glycerine and
albumen-solution, such as is used for sticking sections on slides.
The spicules were shaken up in this and then centrifuged down,
after which a drop or two of the fluid, with the spicules, was
drawn up with a pipette, spread out on a slide, and dried off on
the paraffin oven. When dry, the spicules were fixed on the
slide by plunging it into absolute alcohol, whereby the glycerine
is extracted and the albumen coagulated. The spicules can now
be decalcified and stained in any way that is desired. Other
methods of imbedding and fixing the spicules were also tried but
were not satisfactory. By means of the albumen solution,
provided that neither too much nor too little be used, good
permanent preparations of the decalcified and stained spicules can
be made and mounted in Canada balsam.
In addition to this method, spicules in distilled water, without
addition of albumen, were treated with acids and stains and the
effects of them watched under the microscope. Since the spicules
treated in this manner were not fixed to the slide, it was impos-
sible to wash out the stain and mount them permanently in
Canada balsam, but it was possible to observe in detail the effects
of the acids and stains upon the spicules, and there was the
advantage that the results were not complicated by the presence
of the albumen, which is itself stained by both nigrosin and
indulin.
The Spicules of Clathrina contorta *.
We shall begin with an account of the results attained with the
spicules of Clathrina contorta, of which we had a very abundant
material; the differences presented by other species will be noted
subsequently. Spicules fixed to the slide with albumen were
treated for about half-an-hour with a combination of an acid and
a stain in the following proportions :—
2°/, to 1°/, of acid in distilled water, except in
the case of picricacid of which a saturated
SOMONE Was USEC om tect ree soso eh ey 9 vols.
iy) 4 Sietneaiatin Lisl eal WeIeres (oAspoaoccoddouaaepnesdae 1 vol.
In this way, picric, nitric, acetic, and hydrochloric acids were
combined, respectively, with either nigrosin or indulin as a stain.
After staining for 20 minutes or half-an-hour, the preparations
were washed with distilled water, absolute alcohol, oil of cloves,
and mounted in Canada balsam. The results in all cases were
the same. The spicule was completely decalcified, and left behind
a deeply stained sheath, and an axial filament in each ray (figs. 1—
10). The best and clearest results were obtamed with the picric
acid combinations; with the other acids the filaments were
stained just as deeply, but there was a frequent tendency to form
a flaky deposit which obscured the result.
* For an account of the spiculation and nomenclature of this sponge, see Minchin,
P. Z. §. 1905, ii. pp. 3-20.
668 PROF. MINCHIN AND DR. REID ON THE [J une 16,
The same combinations of acids and stains were also applied to
the spicules placed on the slide in water, without any albumen.
When treated in this way, the sheath of the spicule stains so
deeply as to largely obscure the axial filament, either on account
of the spicule being exposed on all sides to the action of the stain,
or because the preparation does not go through the processes of
washing and clearing necessary for a permanent preparation,
processes which probably extract a certain amount of the colour.
The deep colour of the sheath is especially marked in the combina-
tions with nitric, hydrochloric, and acetic acids, more so than
with picric acid. The fact that the sheath stains so intensely is
of interest, since it shows that the action of Hau-de-Javelle in
isolating the sponge-spicules does not destroy the sheath. When
the spicules are fixed to the shde with albumen, each spicule
appears after treatment with the combined acid and stain asa
space or mould in the layer of albumen limited by a deeply-
stained contour (compare fig. 1, Pl. XX XITV., fig. 5, Pl. XXXYV..,
and fig. 8, Pl. XX XVII. especially) ; but since the albumen takes
the stain also, the sheath cannot be distinguished with certainty
as a structure separate from the enveloping albumen, in spicules
fixed in this way. The fact that the spicule-sheath is not dis-
solved in Eau-de-Javelle is in favour of Bitschli’s view that the
sheath is chiefly inorganic in nature; a conclusion founded by
him on the observation that the sheath could be isolated by means.
of caustic potash (35"/,), though this reagent dissolved the rest of
the spicule, and that sheaths so isolated were dissolved by strong
acetic acid.
When the processes of decalcification and staining were watched
under the microscope, it was observed that the picric acid com-
binations did not break up the spicules so much as the other
acids, even when these were used in strengths much lower than
those quoted above. With picric acid and nigrosin combined, the
filament appears first at the tip of the ray, and as the decalcification
goes on, the filament appears as if traced by the tip of the gradually
receding calcite, until it reaches the centre, when decalcification
is complete. With hydrochloric and nitric acid combinations the
decalcification does not go on so regularly; fragments of the
spicules are frequently seen to be cut off from the rest of the
spicule, and when separated, the fragments rush along to the
tips of the raysas if impelled by powerful currents. The violence
of the action of the acid was most marked with hydrochloric,
less with nitric, and least with acetic acid ; it probably accounts.
for the fact that the axial filaments are not, as a rule, so well
shown with these acids as with the picric-acid combinations.
In addition to the combinations of acids and stains mentioned
above, many experiments were made with acids and stains used
separately. When clean spicules, placed in distilled water on the
slide without any albumen, were treated with acids, it was usually
observed, especially when acetic acid was used, that the whole
spicule seemed to disappear, leaving only the axial thread, without.
1908. } SPICULES OF CALCAREOUS SPONGES. 669
any sheath. Biitschli also (1908, p. 317) was unable to find any
remains of the sheath after dissolving the spicules with acid.
When, however, the acid was combined with the stain, both
filament and sheath were left intact and stained. Hence it is
probable that the disappearance of the sheath, when acid alone is
used, is due, not to the destruction of the sheath by acid, but to
its collapsing on the filament. This conclusion is supported by
the fact that when spicules stuck on with albumen are treated
with a combination of acetic acid and nigrosin, many of them
appear to contain unusually thick filaments, which are seen on
closer inspection to consist of the true axial filament together
with the collapsed sheath. Acetic acid would thus seem to have
a solvent or partially softening action upon the sheath. Biitschli
also found (1906, p. 317) that spicule-sheaths isolated by caustic
potash were dissolved completely by strong acetic acid, but were
preserved by very dilute acetic. Attempts to decalcify the
spicules with acid first and then to stain the sheath and axial
filament subsequently with nigrosin or indulin, were successful
when picric, nitric, or acetic acid were used, but not with hydro-
chloric acid. Various other stains were used without any effect
on the filament, for example carmine stains (borax- and alum-
carmine), Kernschwarz, indigo-carmine, &c.
Appearance and Structure of the Axial Filament.
The axial filament occupies a central position in the axis of the
spicule-ray. By focussing carefully the upper and lower surfaces
of the spicule-sheath in a spicule, decalcified and stained, it can
be clearly seen that the filament lies midway between the two
surfaces. In the optical transverse section of a ray, such as can
be easily obtained in the case of the gastral rays of the quadri-
radiates (fig. 4, Pl. XXXV.), the filament appears as a black dot
occupying the centre of the ray, and can be traced up and down
the ray by focussing. The axial filament exhibits a certain
amount of tenacity and strength: this is shown by the fact that.
when the decalcification proceeds irregularly in a spicule im-
bedded in albumen, detached fragments of calcite may be held
still for a time by the filament, until set free either by the
filament giving way under the strain, or by decalcification taking
place at the centre of the fragment round the filament, after
which the loose fragment rushes along inside the sheath of the
spicule. In spicules not imbedded in albumen, the sheath some-
times breaks across the ray, after decalcification and staining,
but the distal part of the sheath is held on by the filament,
which stands a great deal of bending and washing about without
breaking across. In such preparations it is clearly seen that the
sheath is a very delicate structure, much less strong and resistant
than the actual filament.
In the triradiate systems the axial filament appears to start
from the extreme tip of each ray of the spicule, and to be con-
670 PROF. MINCHIN AND DR. REID ON THE [June 16,
tinuous at this point with the sheath (fig. 8, Pl. XXXVIT.).
The terminal portions of the filaments are very slender and
delicate, but they soon become thicker as we pass towards the
centre of the spicule, and in the greater part of the shaft of each
ray the axial filament is a coarse structure very obvious when
stained. At the junction of the rays the filament widens out
very greatly and forms a cobweb-like arrangement, usually of
triangular shape, which may be termed the central triangle,
and occupies the centre of the triradiate system (figs. 2, 5-7,
Pl. XXXV.). By comparing different spicules, it is seen that the
structure of the central part varies. In those spicules which have
developed a fourth ray and become quadriradiates, each of the
three axial filaments of the triradiate system is continued into
the central triangle, but usually not quite to the central point of
the spicule: the filament seems to break up as it were, to form
the triangle (figs. 6, 7). On the other hand, in the triradiates
with no gastral ray, the axial filaments are continued scarcely
diminished to the centre and there become continuous, and the
central triangle is very faint (fig. 2). This arrangement, though
specially characteristic of the trivadiates, is sometimes seen also
in the quadriradiates (fig. 5).
The monaxons of C. contorta were found very difficult to deal
with on account of their huge size. In the albuminised prepara-
tions they are only partly covered by the albumen, hence the
sheath stains very deeply, just as in the triradiates when they are
stained without being imbedded. Further, when they are cleared
and mounted in Canada balsam after staining, the larger mon-
axons collapse. Smaller monaxons, however, give satisfactory
preparations from which good photographs can be taken (figs. 9,
10, Pl. XXXV.). It is seen that the monaxons contain an
axial filament which commences at each extremity of the spicule
asa fine thread, and as it passes towards the middle point of
the spicule the thread widens out so as to be represented by a
doubJe contoured band, which extends through the greater part
of the shaft of the monaxon. In some monaxons the band may
be quite one-third the width of the spicule ; in others it is com-
paratively narrower.
The monaxons of C. contorta require decalcification for not
less than half-an-hour, or even longer.
The Spicules of other Calcareous Sponges.
In the Clathrinide examined, namely C’. clathrus and Ascandra
falcata, we have found the filaments exceedingly distinct, as was
noted by Minchin (1898), in C. coriacea, when stained by the
methods above described; and there is scarcely any difference to
be noted except in minor points, from what has been described
in Clathrina contorta. Ascandra falcata is a very favourable
object for studying the filaments, especially in the characteristic
sickle-shaped monaxons (fig. 12, Pl. XX XIV., and figs. 13, 14,
1908. | SPICULES OF CALCAREOUS SPONGES, 671
Pl. XXXVIT.) which are abundant and easily found, and at
the same time are not so inconveniently large as in C. contorta.
It is seen that in the distal blunt curved portion (fig. 13) the axial
filament is very thick, and forms a broad band showing a
dark double contour enclosing a central lighter portion ; in the
proximal straight pointed portion (fig. 14) the filament appears
as a Single thread, as in the rays of the triradiates. Hence the
monaxons of A. falcata show a noteworthy difference from those
of C’. contorta, a point to which we shall return.
In C. clathrus (fig. 15, Pl. XXXVI, fig. 16, PI. XXXITV.,
and fig. 17, Pl. XX XVII.) it was found that in the majority
of the triradiates the axial filament terminated abruptly at an
appreciable distance from the end of the spicule, and was not
continuous with the sheath, This condition is probably correlated
with a peculiarity in the mode of growth of the spicules of this
species which was pointed out by Minchin (1898), namely, that
the apical formative cell or “founder” does not leave the ray,
but persists and helps, apparently, to secrete the blunt thickened
termination of the ray which characterises this species. Ina few
cases, however, a continuation of the axial filament up to the
sheath could be seen distinctly (fig. 16), but from the shape of
the rays it is probable that in such cases the spicules were not
quite full-grown.
The spicules of Leucosoleniide and Heterocela examined by
us appear, with one exception presently to be described, very
differentifrom those of Clathrinide. The first impression derived
from examination of them is that no axial filament is present.
A more careful study reveals a filament presenting a certain
sinilarity to that of the monaxons of Ascandra falcata, namely,
a broad band towards the base of each ray, which narrows to a
delicate filament towards the tip of the ray (fig. 18, Pl. XX XVII).
It is very difficult to get satisfactory photographs of the filament,
both on account of its feeble staining powers, and of the difficulty
of getting it in focus, due to the frequent curvature of the rays.
In connection with the phylogenetic speculations of Minchin
(1900, p. 109, and 1908), it is of interest to find so great a differ-
ence in the structure of the spicules of Clathrinidz on the one
hand, and of Leucosoleniide and Heterocela on the other hand.
It may be pointed out further that, as stated above, the studies
of Ebner and Biutschli, with regard to the presence of an axial
filament, were based entirely upon examples of the Heteroccela.
An exception, however, to the foregoing statements concerning
Heteroceela is furnished by the remarkable sponge Heteropegma
nodus-gordii Pole}j., of which, by the kindness of Professor Dendy,
we have been able to examine a specimen. In this sponge we
find the axial filaments very distinct, especially in the triradiates
of moderate size *, in which they present the same characters as
in Clathrina contorta, and can be photographed easily at low
* For figures of the spiculation of Heteropegma see Poléjaeff, ‘ Challenger”
Reports, Zool. vol. viii. part xxiv. (1883) pl. iv. figs. 1, a—-d.
Proc. Zoon. Soc.—1908, No. XLITI, 43
672 PROF. MINCHIN AND DR. REID ON THE [June 16,
magnifications. The very large triradiates, however, tend to
collapse when decalcified, like the monaxons of C. contorta, so
that it is difficult to obtain satisfactory preparations of them.
In the minute triradiates also the filaments are clearly seen, and
present no other difficulties to the photographer than such as
are caused by their minute size, and by the fact that the rays
usually lie in different planes. A study of the filaments in this
sponge brings out a point of some morphological interest. The
minute triradiates, as is well known, are of two types of form,
regular and sagittal. An examination of the filaments shows,
however, that in both forms alike the filaments meet at the centre
at angles of 120° (fig. 22, Pl. XXXIV., fig. 23, Pl. XXXVI.)*.
‘Thus the spicules of Heteropegma are distinctly Clathrinid in type, —
and the sagittal forms occurring in this sponge are to be regarded
as arising simply by secondary curvature of the rays of a primi-
tively regular trivadiate; they may be termed pseudo-sagittal.
It has already been pointed out by Bidder (1898) and Minchin
(1900, p. 109), that Heteropegma is a sponge which stands apart
from other Heterocela and approaches the Clathrinide in its
characters ; and the study of its axial filaments certainly supports
these conclusions.
The Nature of the Axial Filament.
If we compare one of the photographs given here (figs. 2, 6-8)
of the axial filaments of the triradiate systems of Clathrina
contorta, magnified 1000 diameters, with the figure of a young
spicule of this sponge given by Minchin (1898, pl. 42. fig. 49) at
the same magnification, we are at once struck by the fact that
the central triangle, as we have termed it, formed by the axial
filaments, is nearly as large as the whole spicule at this early
stage; much larger, in fact, than the earliest stages of the
spicules that can be found.
It was further shown by Minchin (J. ¢. pp. 572-579) that
the spicules in early stages of development, when examined
between crossed nicols, light up scarcely or not at all. Hence in
early stages the spicules must contain very little, if any, erystal-
line substance, that is to say calcite, but must consist chiefly of
non-crystalline substances, perhaps both organic and inorganic.
We have referred above to Ebner’s statements as regards the
inorganic impurities in the spicule. It must be supposed that
the first portions of the spicule formed consist chiefly of these
“impurities,” and that the axis of the spicule, as it grows in
length, is also formed of substance containing very little calcite.
On the other hand, the portion of the spicule formed later is
_* In all the preparations the axial filaments are very liable to become displaced,
since they are entirely unsupported after decalcification of the spicule. In the
spicule photographed in fig. 23, it can be seen that the filament of the left-hand ray
is displaced, but that of the right-hand ray shows the typical regular angle.
1908. } SPICULES OF CALCAREOUS SPONGES, 673
almost pure calcite; but the layer formed last of all, namely the
sheath, is again an “impure” layer.
Having regard to the mode of formation of these spicules
discovered by Minchin (1898 and 1908) and Woodland (1905), it
would be a tempting hypothesis to refer the two substances
secreted to the activities of the two formative cells; the apical
formative cell or “founder” may be supposed to lay down the
“impure ” substance, while the basal formative cell or “thickener ”
secretes the purest calcite. On the other hand, the formation of
the sheath must also be ascribed to the thickener.
The continuity, generally to be observed, of axial filament and
sheath, and the similarity of their staining reactions are points
in favour of considering these two structures to be of similar
nature. We have referred above to Biitschli’s arguments in
favour of regarding the sheath as being chiefly of inorganic
nature, a conclusion for which there is much to be said, and
which may be extended to the axial filament. The fact, however,
that both filament and sheath have an affinity for special stains,
is in favour of their containing a certain amount of organic
matter, and we may regard sheath and filament as consisting of
an organic basis richly impregnated with inorganic non-crystal-
line materials. At this point we must leave the question of the
nature of these structures to receive more exact and definite
solution from more competent observers. We claim merely to
have demonstrated the following proposition :—The spicules of
calcareous sponges leave after decalcitication a residue in the form
of structural constituents, sheath und axial filament, which can be
coloured by special stains.
In conclusion, attention may be drawn to some points relating
to the morphology of the spicules, upon which the axial filaments
throw some light. It is seen that in the rays of the triradiates,
the filament is broad and even band-like at the base, and tapers
to a fine point at the apex. Comparing with this the monaxon
of A. falcata (fig. 12, Pl. XXXIV., figs. 13, 14, Pl. XXXVIL.),
it is seen that the filament is broad and band-like at the blunt
distal end of the spicule, and tapers to a fine thread at the
pointed proximal end. ‘This supports the conclusion, based by
Minchin (1908) upon developmental data, that the distal pro-
jecting ends of the monaxons are homologous with the central
ends of the rays of the triradiates.
Comparing, however, the monaxons of C.. contorta with those of
A, falcata (figs. 9,10, Pl. XXXYV.), it is seen that in C. contorta
the filament is band-like towards the middle of the spicule, but
tapers to a fine thread at each end. This strongly suggests that
the monaxons of this sponge are not really primary monaxons,
but are secondarily derived from triradiates and are to be re-
garded as biradiates as suggested by Minchin (P. Z. 8. 1905, ii.
p. 10). On the other hand, the monaxons of 4. falcata would
appear to be true primary monaxons.
Oe
A DX
e
674 PROF. MINCHIN AND DR. REID ON THE [June 16,
ADDENDUM.
Intracellular Networks in the Gastral Layer.
A point of some interest was observed in a preparation made
in the following manner. A piece of the body-wall of Clathrina
contortw (preserved in alcohol) was stained with picro-nigrosin,
and passed through water and alcohols into oil of cloves; then
She inner gastral surface was brushed gently with a soft paint-
brush to remove the gastral layer of collared epithelium ; finally,
the piece was mounted in Canada balsam with the gastral surface
uppermost. The upper surface of the preparation then showed a
delicate honeycomb-like network, stained blue with the nigrosin,
enclosing irregular polygonal meshes fairly uniform in size, with
here and there a much larger rounded mesh and occasionally a
small, circular mesh. As the preparation was not all in one
plane, only small stretches of the network could be sharply
photographed (fig. 24, Pl. XXXVIT.). The polygonal meshes
represent spaces formerly occupied by collar-cells, many of which
are to be seen still in sitw in the preparation; the large rounded
meshes are spaces left by porocytes; and the small circular
meshes are shown by their relations to underlying triradiate
systems to be the spaces occupied by the gastral rays of quadri-
radiates. The network itself is an extension of the gelatinous
ground-substance between the bases of the collar-cells and gastral
rays and round the inner ends of the porocytes, probably forming
a cementing substance, as it were, helping to keep the easily
detached collar-ceils in their places.
BIBLIOGRAPHY.
Brpper, G. P.—‘‘ The Skeleton and Classification of Calcareous
Sponges.” Proc. Roy. Soc. Ixiv. (1898) pp. 61—76, 10 text-figs.
BowErBANkK, J. 8.—A Monograph of the British Spongiade. I.
Ray Society, 1864.
Birscui1, O,—‘‘ Einige Beobachtungen tiber Kiesel- und Kalk-
nadeln von Spongien.” Zeitschr. f. wiss. Zool. lxix. (1901)
pp. 235-286, pls. xix.—xxi., 2 text-figs.
—— ‘Ueber die Kinwirkung von konzentrierter Kalilauge
und konzentrierter Losung von kohlensaurem Kali auf
kohlensaurem Kalk, sowie tiber dabei entstehende Doppel-
salze von kohlensaurem Kali und kohlensaurem Kalk.”
Verh. Naturhist. Med. Ver. Heidelberg, n. f. viii. (1906)
pp- 277-330, 7 text-figs.
Cartrr, H. J.—‘‘ Description of a Siliceous Sand-Sponge found
on the South Coast of Arabia.” Ann. Mag. Nat. Hist. (4)
ii. (1869) pp. 15-17, 4 text-figs.
— ‘Calcareous Sponges.” Ann. Mag, Nat. Hist. (4) i.
(1869) p. 466.
HpneR, V. v.—‘‘ Ueber den feineren Bau der Skelettheile der
Kalkschwimme, nebst Bemerkungen tiber Kalkskelete tiber-
haupt.” Sitzber. k. Akad. Wiss. Wien, I. Abth. xev.
(1887) pp. 55-149, pls. 1-1v.
1908. |; SPICULES OF CALCAREOUS SPONGES. 675
Grant, R. E.—“ Remarks on the Structure of some Calcareous
Sponges.” Edinburgh New Philosophical Journal, i. (1826)
pp. 166-170.
Harcket, E.—Die Kalkschwiimme. Bd. I. Berlin, 1872, pp. xvi-+
484.
Koutrker, A.—Icones Histiologice. Erste Abtheilung. Leipzig,
1864.
LenDENFELD, R. v.—‘‘ The Histology and Nervous System of the
Calcareous Sponges.” Proc. Linn. Soc. N.S. Wales (1) ix.
(1885) pp. 977-983.
— “Die Spongien der Adria.—I. Die Kalkschwiimme.”
Zeitschr. f. wiss. Zool. liii. (1891) pp. 185-321, & pp. 361-433,
pls. vili.xv., 1 text-fig.
Lizserktun, N.—“ Beitrige zur Anatomie der Kalkspongien.”
Arch. f. Anat. Physiol. und wiss. Med. 1865, pp. 732-748,
ple xix.
Maas, O.—“‘ Ueber den Aufbau des Kalkskeletts der Spongien
in normalem und in CaCo,-freiem Seewasser.” Ver. Deutsch.
Zool. Ges. 1904, pp. 190-199.
Mincuin, KE. A.‘ Materials for a Monograph of the Ascons.—
I.” Quart. Journ. Micr. Sci. n.s. xl. (1898) pp. 469-587,
pls. xxxvill.—xlii.
— Article “Sponges,” in Lankester, A Treatise on Zoology,
part 11. London, 1900.
“* Materials for a Monograph of the Ascons.—II.” Quart.
Journ. Micr, Sci. n.s. li. (1908) pp. 301-355, pls. xvii —xxi.,
5 text-figs.
Scumipr, O.—Die Spongien des Adriatischen Meeres. Leipzig
(W. Engelmann), 1862.
WEINSCHENK, H.—“ Ueber die Skeletteile der Kalkschwaimme.”
Centrbl. Mineral. Geol. Paliontol. 1905, no. 19, pp. 581-588.
Woopiann, W.—‘“< Studies in Spicule Formation.—I. The Devel-
opment and Structure of the Spicules in Sycons; with
Remarks on the Conformation, Modes of Disposition and
Evolution of Spicules in Calcareous Sponges generally.”
Quart. Journ. Micr. Sci. n. s. xlix. (1905) pp. 231-282,
pls. xili.—xv., text-figs. 1-11.
EXPLANATION OF THE PLATES.
All the photographs are from spicules stuck on the slide with albumen, decalcified
and stained with picro-nigrosin, and mounted in Canada balsam.
Figs. 1-10. Clathrina contorta.
Fig. 1, Pl. XXXIV. A triradiate. x 400. FF tae
2, Pl. XXXV. The central portion of another triradiate. X 1000.
3, Pl. XXXIV. A quadriradiate, gastral aspect. xX 400.
4, Pl. XXXV. A quadriradiate, gastral aspect, at a high focus to show the
gastral ray in optical transverse section. X 1000.
5, Pl. XXXV. The central part of a quadriradiate. X 1000.
6, Pl. XXXV. The central part of another quadriradiate. x 1000.
7, Pl. XXXV. The central part of another quadriradiate. X 1000.
8, Pl. XXXVII. The extremity of a ray of a quadriradiate. XX 1000.
9, Pl. XXXV. A small-sized monaxon. X 150.
10, Pl. XXXV. Another small monaxon. X 150.
676 DR. T. A. CHAPMAN ON NEW [June 16,
Figs. 11-14. Ascandra falcata.
Fig. 11, Pl. XXXIV. Central portion of a triradiate. > 1000.
12, Pl. XXXIV. A monaxon. X 250.
13, Pl. XXXVII. Distal extremity of amonaxon. X 500.
14, Pl. XXXVII. Proximal extremity of amonaxon. X
Figs. 15-17. Clathrina clathrus.
ig. 15, Pl. XXXVI. The extremities of two triradiates and a broken ray of ai
third. X 1000.
16, Pl. XXXIV. The extremity of a triradiate. > 1000.
17, Pl. XX XVII. The central part of a triradiate. 1000.
a
Ry
Figs. 18, 19. Leucandra aspera.
Fig. 18, Pl. XX XVII. A triradiate showing the double-contoured filaments. X 250.
19, Pl. XXXVI. A quadriradiate. x 500.
Figs. 20, 21. Sycon ciliatum.
Fig. 20, Pl. XXXVI. A triradiate. > 500.
21, Pl. XXXVI. The same triradiate at a slightly lower focus. xX 500.
Figs. 22, 23. Heteropegma nodus-gordii.
Fig. 22, Pl. XXXIV. A small triradiate (one ray broken). 1000.
23, Pl. XXXVI. A small sagittal triradiate, showing the filaments; on the
left the filament has become displaced. X 1000.
Fig. 24, Pl. XXXVII. Photograph of the gastral surface of the body-wall of
Clathrina contorta, stained with picro-nigrosin, the collar-cells
brushed off; showing the network left between the collar-cells,
porocytes, and gastral rays. Owing to this network not being exactly
in one plane, it is not seen all over the photograph. X 1000.
=
0g
4, Two New Genera (and a New Species) of Indian
Lycenids. By T. A. Cuapman, M.D., F.Z.S.
[Received May 14, 1908. }
(Plate XXX VIIT.*)
In trying to gain some knowledge of the genus Cyaniris by
examining the ancillary appendages, I met with much trouble
over Cyaniris chennellii de Nicév. I obtained specimens from
various sources, and informed various people that they had a Zizera.
or something thereabouts, and not a Cyaniris. Herein I was
right, but so were they, their insect being chennellit de Nicév. I
stuck to my guns unnecessarily, largely because Col. Bingham
found in his collection a specimen that was certainly not a Zizera.
but probably a Cyaniris, and which he had compared with the
type of chennellii and found to agree. I took it therefore that
this was chennellii, but could come across no other specimen.
I also, of course, assumed de Nicéville to know what was and what
was not a Cyaniris, and that he would not call a Zizera-like species.
a Cyaniris. It turns out, however, that this was precisely what.
he did do, and in doing which, succeeding authorities appear to have
* Wor explanation of the Plate see p. 678.
1 Za Sin MGs Tells SO-OOV0Ne
Photo F. N. Clark. Bale & Danielsson, Ltd.
ANCILLARY APPENDAGES OF—
1. BOTHRIA CHENNELLII. x 45.
2. NOTARTHRINUS BINGHAMI. x 45.
ee
ai)
a
1908. | INDIAN LYCHZNID BUTTERFLIES. 677
followed him. Col. Bingham had in his possession (lent from
India) the type specimen of chennellii, and when I saw this.
specimen, it appeared at once that it was the insect I had been
refusing to recognise as chennellit.
chennellit has all the aspect of a Cyaniris, but the neuration is.
not that of Cyaniris but of Zizera. The ancillary appendages.
are very different from those of Cyaniris and also from those of
Zizera, and approach more nearly to those of Lveres.
It appears to require a new genus to contain it, and I therefore
propose for it Boruria *, n. g.
I would characterise the genus as having C. chennellit de Nicé-
ville as type; as closely resembling Cyaniris in facies, but
having the anastomosing subcostal vein as in Zizera; as having
the ancillary appendages very different from Cyaniris and Zizera.
The dorsal portion (¢eguwmen) consists of a central piece, with two:
horns jointed to it, both of very similar form and structure to:
those of Hveres (Cyaniris and Zizera have the dorsal armature in
two lateral portions, the actual dorsum being merely part of the
chitinous ring of the 9th abdominal segment with no armature).
The claspers have the two processes (characteristic of Lycznids) of
nearly equal size, each being very long and slender but reminding
one a good deal.of Hveres. In Cyaniris and Zizera the ventral,
soft (7. e., unarmed with spicules or teeth) process is nearly or
quite obsolete.
It remains to deal with Col. Bingham’s Cyaniris. As a
specific description, de Nicéville’s description of chennellii is
all but accurate for this species, which is however a Cyaniris (and
chennellii, as we have just seen, is not).
As a Cyaniris it belongs, however, as an extreme instance, to
a division that requires a separation generically. This section
is especially characterised by each lateral piece of the dorsal
armature of the ancillary appendages having a jointed horn or
process (as in so many Lycenids) and not merely a simple process,
usually rounded and soft but in many species with a more or less
developed chitinous point or spike.
I would call this new genus Wotarthrinus.
NovarTHRINUS 7, 0. g.
Differs from Cyaniris in each portion of the dorsal armature of
the ancillary appendages having the separate jointed spine (as in
Lycena). In Cyaniris the spine, when there is one, is merely a
chitinous process continuous with its base.
Type, binghami, sp. n.
musina (and corythus, which seems to be identical with
musina) and catrews probably belong to this genus.
* BoOpos a pit, in allusion tothe pitfall into which it led de Nicéville. For those
who like a little canine flavour to their classics, it may suggest that it is like both
Cyaniris and Zizera without being either, and has perhaps some allusion to the
bother which de Nicéville and others did not suffer but passed on to me.
+ voros apQpos.
678 MR. F. E. BEDDARD ON THE [June 16,
NoraRTHRINUS BINGHAMI, Sp. 0.
Agrees with de Nicéville’s description of Bothria chennellit,
except that it is without the discal lme on the upper side of the
fore wing, and the 6th spot of the row on the under side is all but
obsolete. A more important difference is that in this row of spots,
the first one in chennellii is in line with the others, in binghami
it is markedly moved inwards, as in argiolus and many other
Cyanirids.
I cannot help a suspicion that de Nicéville had these two
species, chennellii and binghami, mixed (Col, Bingham, also, had
not separated them), and when referring chennellia to Cyaniris
had binghami in view.
Type in Col. Bingham’s collection. A cotype in somewhat finer
condition is in the museum at Tring. Col. Bingham’s specimen
is from Shillong; the Tring specimen is labelled ‘‘ Khasia Hills,
Assam.”
EXPLANATION OF PLATE XXXVIII.
Fig. 1. Ancillary appendages of Bothria chennellii X 45.
Fig. 2. Ancillary appendages of Notarthrinus binghami X 45. The Aidceagus was
unfortunately mounted on the slide too far off to come into the photo-
eraph. I have not based any characters of genus or species on it.
Both photographs by Mr. F. N. Clark.
5. A Contribution to the Knowledge of the Batrachian
Rhinoderma darwin. By Frank E. Bepparp, M.A.,,
WaRESe, EAS:
[Received May 26, 1908. ]
(Text-figures 143-149.)
I have examined a number of examples of this small Chilian
Engystomatid Frog which were presented to the Society by
Dr. EK. P. Reid some months since. This examination enables me
to add something to what is already known concerning this Batra-
chian, the breeding-habits of which are so remarkable and now so
well known through the investigations of Gay, Espada, Howes and
others, especially and most recently Prof. Biirger, for the loan of
whose memoir written in Spanish I am indebted to Dr. Reid.
Dr. Birger* and Prof. Howest give lists of the memoirs
relating to this species, which I need not therefore recapitulate
here. The contributions made respectively by the two
authorities mentioned deal with the gular sacs, their structure
and development. Incidentally some information is given in
these papers concerning the visceral and muscular anatomy with
which I have occupied myself for some time past; and so far as
the facts go I am able to confirm those authors. As, however,
* La Neomelia de la Rhinoderma darwini D. & B., por el Dr. Otto Burger.
Santiago de Chile, 23 pp., 3 pls.
Le Notes on the Gular Brood-pouch of Rhinoderma darwini,’ P. Z.S. 1888,
p. sol. ‘
1908. | BATRACHIAN RHINODERMA DARWINI. 679
their object was not to enter into the general anatomy of
Rhinoderma, but only to describe structures associated with the
breeding-pouches, there is naturally some room left for a fuller
account of this Engystomatid frog, which I specially compare with
its African relative the genus Greviceps, dissected by myself some
time since and referred to in a paper communicated to this
Society lately*. Rhinoderma is an Engystomatid frog without a
narrow mouth. The mouth is not far from being of the ordinary
Batrachian capacity. The general aspect also of the species is
widely removed from that of Sreviceps, with which it would
certainly not be associated were external characters alone taken
into consideration. Externally, in fact, Rhinoderma is a typical
frog, except indeed for the ‘considerable projection of the upper
beyond the lower jaw, and the presence of a narrow projection of
the snout region of the head which has nothing to do with the
nostrils. This process is not always present, but its presence or
absence has nothing to do with sex, for I found the process well
developed in one female, and absent in another female, both speci-
mens being of about the same size. I have not, however, examined
with care a sufficient number of examples to enable me to
contribute any material facts with reference to the development
of this ‘‘ snout” f.
§ Visceral Anatomy.
The viscera show a number of differences from those of
Breviceps.
The liver in Rhinoderma has not the peculiar form and relations
to the heart which I have described in Greviceps. It is consti-
tuted more on the plan of that of Rana. That is, there is a
primary division into right and left lobes of which the right is
much the larger and is again divided into two lobes. The whole
mass of the liver lies entirely behind the heart, which is not in
the least hidden by it in the remarkable fashion which I have
noticed in Breviceps. That this is the case is shown by the
attachment to the posterior border of the pericardium of a
peculiar muscle, which passes from the body-wall under the lobes
of the liver without being attached to them and ends on the
pericardium ¢.
The alimentary tract is proportionately and roughly of the same
general appearance and length as that of Rana. There are, how-
ever, some differences, especially when the gut is slit up and the
characters of the lining membrane in different regions compared
with each other and with the corresponding or apparently corres-
ponding sections of the gut of Rana. The stomach itself has a
tendency to be more spherical in shape than in Rana. The part
which ensues and corresponds in its position to the duodenum of
* P. Z.S. 1908, p. 11.
+ The skull, showing the cartilaginous basis of this process, is figured by W. K.
Parker (Phil. Trans. 1881, pl. 39, figs. i-iv).
t This is described below (on p. 684).
680 MR. F. E. BEDDARD ON THE [June 16,
Rana demands attention before its homologies can be thus rapidly
disposed of. Where this tube arises from the undoubted stomach
there is no valve or change of a sudden character in its lining
membrane. Furthermore, the pancreas does not extend down
into the V-shaped loop which it makes with the stomach, and the
ducts of liver and pancreas open into the alimentary tract farther
up this ascending limb of the gastric U than they are represented
to open in Rana*. Iam disposed in fact to assign the greater
part of this ascending limb—the first deflection from an antero-
posterior course which the whole alimentary tract shows—to the
stomach. Its lining membrane has quite the characters of that of
the lining membrane of the latter half of the undoubted stomach.
Text-fig. 143.
Alimentary tract of Rhinoderma darwini partly opened to show folds of
lining membrane.
S¢. Posterior boundary of stomach. JZ/. Ileum.
It is very thick and arranged in close longitudinal folds which
undergo no change where the tube suddenly lessens in calibre.
Later on, a tendency to a reticulate arrangement also observable
anteriorly becomes rather more marked. This thick layer
suddenly ends near the top of the ascending limb of the V already
referred to as characteristic of this and (? all) other frogs. There-
after the walls of the gut are thin for a considerable distance and
the lumen is perhaps slightly wider. The inner surface is very
definitely reticulate in a honeycomb fashion. The break between
* Haslam’s Translation of Ecker’s ‘Frog,’ p. 296, fig. 195, Dc?.
1908.} BATRACHIAN RHINODERMA DARWINI. 681
this section of the gut and that which precedes is rather in the
abrupt thinning of the lining membrane than in anything else.
Later on the reticulate arrangement is still retained; but there
is a tendency towards emphasising the transversely running folds
of the reticulum, but to nothing like the extent that is figured in
Rana*, These facts are well shown in the accompanying figure
(text-fig. 143). The small intestine opens very abruptly into the
short dilated colo-rectum. A little way in front of the junction
of the two the ileum, as we may term it, becomes somewhat.
narrower in calibre and it has been for some distance thicker-
walled. The end of the small intestine in fact is as thick-walled
as that part of the tube (whatever its homologies may be) which
immediately succeeds the dilated stomachal chamber. Both these
regions contrast very markedly with the thin-walled middle
section of the gut. On cutting open, these differences were very
apparent. The colo-rectum is also thin-walled—at any rate in
comparison with its calibre. The end of the ileum actually
projects into it for some distance, like the uterus into the vagina.
The figure which I give here of the intestinal tract of Rhinoderma
(text-fig. 143) may be compared with that of Breviceps t, although
the former is represented as seen when cut open and the latter is
not. I have pointed out in Breviceps ~ that the stomach does not
end where it suddenly diminishes in calibre, but that it is clearly
continued for a short distance along the upward limb of the U
which it forms with theduodenum. I believe that in Rhinoderma
this extension of the stomach is still greater.
Text-fig. 144.
/
Alimentary tract of Rhinoderma darwini, to illustrate the shortness of that of the
male (upper figure) and the greater length of that of the female (lower figure).
The accompanying drawings (text-fig. 144) show the different
appearance of the gut in the male and in the female of this frog.
* Haslam, foc. cit. p. 288, fig. 189.
+ P.Z.S. 1908, p. 32, text-tig. 10. ft Loe. cit. p. 31, text-fig. 9.
682 MR. F, E, BEDDARD ON THE [June 16,
When the body of the male is cut open, the whole of the alimentary
tract is displayed and may be seen without moving that tract or
adjacent organs. On the other hand, in the female the coils of
the gut are rather more complex. The difference, as will be seen,
is due to the greater accentuation in the female of the loops of
the intestine. This is obviously associated with a considerable
difference in the length of the tube in the two sexes. The
measurements of two individuals were as follows:—In a male
measuring 22 mm. from snout to anus (the anterior process of
the snout being omitted) the gut from the commencement of
the ascending limb, which may or may not belong really to the
stomach, to the point of entrance into the colon of the ileum was
only 18 mm. Ina female measuring 28 mm. the gut was 35 mm.
In the former, therefore, the gut is actually shorter than the body
length. It is rather longer in the female.
§ Uro-Genital Organs.
The kidneys have the flattened leaf-like form that characterises
those organs in Rana. They were, however,—in an example in
which I measured them—proportionately very much larger than
in an example of Rana esculenta, of which I made measurements
for the purposes of comparison. The specimen of Rhinoderma
darwint measured from the extremity of the snout (this example
had not the anterior prolongation so characteristic of the species)
to the anus 33 mm. ‘The left kidney measured 8°5 mm., being
thus very nearly one quarter of the length of the body—an
extraordinary size. In correlation with this great size was the
fact that the anterior extremity of the kidney nearly reached the
anterior wall of the pleuro-peritoneal cavity, and the fat-bodies
were so thrust against that anterior wall by the growth of the
kidney that they lay back over it, being directed towards the vent.
Ina Rana esculenta measuring between the same points 175 mm.,
the length of the corresponding kidney was only 14 or 15 mm.
Thus in this Batrachian (possessing a kidney of the usual size—
in Kcker’s ‘ Frog’ 16 mm. is the length given) the kidney was
only one-eleventh to one-twelfth of the body length. The
difference is enormous.
The testes are spherical, much pigmented, and have the
mulberry-like form of those of Rana. The fat-bodies in the one
male which I dissected were much smaller than in both of two
females which I also dissected. And moreover, in all three cases
the left fat-body was larger than the right. In view of the
peculiarity of the testes in Breviceps in possessing only one vas
efferens, I was surprised to find that Rhinoderma is more normal
in that each testis has four or five slender vasa efferentia.
The oviducts are long and thick and much coiled, and thus
differ from those of Breviceps, presuming that the latter were
fully advanced in development in the specimen which I dissected.
It is remarkable that the proximal part of the oviduct (i. e. that
1908. ] BATRACHIAN RHINODERMA DARWINI, 683
section immediately following upon the funnel) in Rhinoderma is
very short and quite straight, much shorter than is, according to
my experience, the rule among frogs. The funnel itself is
attached sideways to the surface of the obliquus internus, where
that muscle forms the anterior wall of the abdominal cavity, and
is of an elongate form, the orifice being a comparatively narrow
and terminal slit. The lining membrane is grooved at the mouth
of the funnel. The oviducts open into the cloaca by a single
common orifice.
§ The Musculature of the Ventral Surface.
The rectus abdominis shows no great peculiarities of structure.
It has five divisions, visible when the skin is removed and no
further dissection made. These are divided by four inscriptiones
tendinee. This contrasts in the most marked way with Breviceps,
where there is but a single inscriptio tendinea. These are all
behind the sternum, where the muscle appears to end. The
anterior abdominal vein is visible from the last inscriptio tendinea
up to just behind the sternum, where it dips down and dis-
appears from view. The abdominal section of the pectoralis
muscle arises from the first three poststernal masses of the rectus.
The sternal portion of the pectoralis is hardly distinguishable
from a sterno-radialis anteriorly; but the latter—if it exists
as a separate muscle—is quite plainly divided off from the
adjacent slender head of the deltoid. The posterior part of the
pectoralis sternalis is distinct from the anterior region in that it
comes closer to its fellow of the opposite side in the middle line
than does the anterior part of the same muscle.
The obliqui externus et internus have not the extraordinarily
complicated and specialised disposition of their bundles that I
have described in Lreviceps verrucosus*. The obliquus externus is
a tolerably stout muscle the fibres of which run at right angles to
the long axis of the body in the ordinary way, and which forms as
usual a continuous sheet covering the sides of the body. Opposite
to the second inscriptio tendinea of the rectus abdominis it is
overlapped by the pectoralis abdominalis, and in this region arises
(or is inserted) from that muscle, or rather from the septum
between the two. In front of this area of overlap the obliquus
externus is seen—when the here superjacent pectoralis abdominis
is dissected away—to end abruptly at the septum between itself
and the several compartments of the rectus abdominis. The
fibres of the two muscles where they thus nearly come into
contact are absolutely at right angles. Anteriorly and much at
the same point, or rather along the same line, as in the Common
Frog the obliquus externus ends definitely in a straight anterior
border. There is not, however, in Rhinoderma darwini any trace
that I could discover of an omo-abdominalis muscle, such as is
* P.Z.S. 1908, p. 22
684 MR. F. E, BEDDARD ON THE [June 16,
well developed and quite obvious in Fana and very greatly
developed in Breviceps. It is rather remarkable that Khinoderma
not only shows no likeness to its ally Breviceps, but 1s even more
simple than Rana. For some distance in front of the end of the
obliquus externus the obliquus internus becomes obvious, its fibres
running at an angle with those of the externus. These fibres end
at the edge of the sternohyoideus just as the fibres of the obliquus
externus end at the edge of the same muscle and of the rectus
abdominis further back. This ending, however, is apparent rather
than real. When the rectus is cut across, the obliquus externus is
seen really to end abruptly at its outer boundary. On the other
hand, the obliquus internus anteriorly dips into the body and
forms a portion of the anterior partition between the neck and the
trunk in a way which will be described immediately in connection
with the transversus portion of the obliquus internus complex
which was originally compared by Huxley to the Mammalian
diaphragm, and which I have named accordingly in the following
paragraph.
The diaphragmatic muscle.—This muscular sheet arises as 1s
usual among the normal Batrachia Salientia—that is, the muscle
has not the large extent and backwardly prolonged origin that it
has among the Pelobatide*. It overlaps the kidney on its way
to be inserted on to the cesophagus, which shows the enormous
extension of that viscus forwards, upon which I comment else-
where in this paper. The muscle is entirely inserted on to the
esophagus from the point where the latter enters the pleuro-
peritoneal cavity up to nearly its junction with the stomach. The
muscle also shares in the formation of the anterior wall of the
pleuroperitoneal cavity; for it merges completely into the
obliquus internus, the two meeting (though the exact line of
junction cannot in the least be distinguished) at about the centre
of the concave wall which they together form and which bounds
the pleuroperitoneal cavity anteriorly, as has been said. It is
difficult to say for this reason whether the muscle does or does not
supply fibres to the root of the lung. Im any case strands of the
obliquus internus end upon the root of the lung.
Pericardial Muscle——In the Common Frog a ‘portion of the
obliquus internus has been described as having the following
relations to the pericardium and in the following words { :—‘“ A
third portion [of the muscle in question], placed behind the
preceding, runs from the pharynx over the pericardium and is
attached to this nearly as far as the middle line, resting on the
sternum, the m. rectus and m. sternohyoideus. The lines of inser-
tion of the muscles of opposite sides form an angle open in front.”
This is illustrated by a figure; but neither the figure nor the
description appears tome tobe clear. The portion of the obliquus
internus thus referred to is perfectly continuous with the rest of
* Beddard, P. Z.S. 1907, p. 346, text-fig. 98; and p. 886.
+ Swpra, p. 682.
{ Ecker’s Frog, Engl. Transl. p. 71.
1908. ] BATRACHIAN RHINODERMA DARWINI. 685
the muscle (I distinguish the cesophageal muscle as distinct) which
forms the internal sheet of the abdominal musculature. The
region, however, now under consideration is that portion of the
obliquus externus which is uncovered anteriorly by the obliquus
externus. The latter muscle ceases at about the level of the apex
of the heart, its most anterior region being separated off as the
omo-abdominal muscle. in front of this the lateral wall of the
pleuroperitoneal cavity is formed by the obliquus internus only,
and this muscle (of course with its liming peritoneum) limits the
abdominal cavity antero-laterally and constitutes the muscular
wall of the “cervical limiting membrane of abdominal cavity”
(Keith*). Ventrally this section of the internal oblique muscle
becomes divided into two insertions. The most posterior ends in
a delicate aponeurosis which passes outside of the sternohyoid
muscle and is attached to the coracoid and sternum. The anterior
Text-fig. 145.
Sth.
A portion of the musculature of the ventral surface of Rhinoderma darwini.
Ant.abd. Anterior abdominal veins H. Heart. .¢. Tendinous intersection of
rectus abdominis. O6/. Obliquus externus. P. Pericardial muscle. Pect.
Pectoralis. #. Rectus abdominis. St.h. Sterno-hyoid.
section of the muscle is attached by aponeurosis to the pericardium
quite anteriorly (in the region of the emergence of the conus
arteriosus), and to the roots of the lungs. The whole of the
ventricular region of the pericardium is quite free from any
muscular insertions. This fact (the freedom of the whole
ventricular region of the pericardium from connection with the
obliquus internus) is not plainly shown in the figure of Gaupp
reproduced by Keith?. In addition to the insertion of obliquus
internus fibres upon the anterior and lateral regions of the peri-
cardium, Dr. Keith has remarked that some of ‘the deep fibres
[of the rectus abdominis, behind the sternohyoid| may end on the
* J. Anat. Phys. xxxix. 1905, p. 244, fig. 1, explan. of figure.
+ Loe. cit. p. 259, fig. 14.
686 MR. F. E. BEDDARD ON THE [June 16,
pericardium.” These will then form the ventral part of the
diaphragm. I do not find after a careful dissection of an average-
sized example of Rana esculenta any evidence of a deflection of
rectus fibres to the pericardium. Iam the more confident in the
accuracy of my observation in that I have discovered such a
muscle in the small Rhinoderma darwini. This I figure in the
accompanying illustration (text-fig. 145). The muscle is thin and
wide and flat, and its fibres run accurately in a direction parallel
with the long axis of the body. It is attached to a good deal of
the posterior and lateral margins of the pericardium. It underlies
the obliquus externus behind the sternal region (7. e. is dorsal to
it), and is therefore perhaps to be regarded as a portion of the
obliquus internus.
The relations of the swbmawillaris are different to those which
obtain in some other frogs. The muscle is, however, similar in
that its posterior region is separated off as a distinct muscle, the
Text-fig. 146.
Sm.
Musculature of floor of mouth of Rhinoderma darwini.
a. Genioglossus (?). 6, ¢. Differentiated portions of submaxillaris.
Sh. Subhyoideus. Sm. Submentalis.
subhyoideus. The latter is only plainly differentiated from the
former near to the edge of the lower jaw, where it dips down toa
lower plane. Its relative dimensions appear to be very much those
of the muscle in Rana. Nor is there anything in the structure or
size of the swhmentalis that calls for particular comment. The
muscle appears to be exactly like that of Rana. It will be noticed,
however, in the accompanying drawing (text-fig. 146, a) that two
large triangular muscles, one on each side, underlie the submaxil-
laris, anil muscles are not visible in a corresponding dissection
of Rana. Nor have I seen them in the same place in such
Pelobatidz: as I have dissected *. These two muscles, as will be
seen in the text-figure referred to, are closely approximated in the
middle line; and into the space left by their divergence in front
fits exactly the submentalis.
* On Megalophrys nasuta, P.Z.S. 1907, p. 338, &c., and on Pelobatide, ib. ¢. ¢
p- 871.
1908. } BATRACHIAN RHINODERMA DARWINI. 687
These two muscles appear to me to be quite possibly the genio-
glossi, which are thus in the species Rhinoderma darwin not only
of very large size but rather abnormal in position. They are
normal only in that they arise on either side from the mandible;
they are abnormal in that they have intruded into an area
belonging to the submaxillaris which ceases to exist as a separate
layer at the margin of the genioglossi. Furthermore, the intru-
sion of the genioglossi on to the superficial area of the throat has
caused another peculiarity in the arrangement of the fibres of the
submaxillaris. It will be seen from an inspection of text-fig. 146
that the fibres of the submaxillaris run in different directions in
different portions of this muscle. Laterally to the possible genio-
glossi the fibres of the submaxillaris run obliquely to each ramus
of the lower jaw; posteriorly to them the fibres are at right angles
to the longitudinal axis of the head and run therefore accurately
across the throat, with no obliquity of direction like the anterior
part of the muscles. If I am right in identifying the triangular
pair of muscles just described with the genioglossi of other Batra-
chians, they certainly differ in not being inserted on to the tongue
(which of course does not necessarily do away with this suggested
homology), for they can be easily dissected away with the sub-
maxillaris, displaying the hyoglossus and geniohyoidei beneath.
In any case, whatever be the nature of these muscles, it is clear
that the muscular floor of the mouth in Rhinoderma darwini is
peculiar and unlike that of other frogs. JI should mention,
furthermore, that there is no vestige in this frog of the small
muscles at the side of the mouth lettered “«” in my figure of
the muscular system of the ventral surface of the body in
Breviceps verrucosus*. It may be that these muscles are
the homologues of the peculiar muscles which I describe in
Rhinoderma (text-fig. 146, a). If so, they have undergone in the
one or in the other genus a very considerable shifting of position.
It must be borne in mind that the species the anatomy of which
forms the subject of my present communication to the Society
is hardly to be described as ‘‘ engystomatous.”
§ The Musculature of the Back.
The latissimus dorsi (text-fig.147, Z.d.) is distinctly different from
that of Rana, as figured. It is a distinctly narrow muscle, being
about one half of the diameter of the underlying infraspinatus.
Its course is straight and is exactly at right angles to the long axis
of the body. It has no obliquity of direction as in other frogs.
It is therefore also exactly parallel to the partly underlying infra-
spinatus. It arises from the middle line of the back behind the
scapula. It did not appear to me to arise from the dorsal fascia ;
but as this pigmented membrane was so delicate it had to be
picked away in little bits, and its relations were therefore rather
* P,Z.S. 1908, p. 16, text-fig. 3.
Proc, Zoou. Soc.—1908, No. XLIV. 44
688 MR. F. E. BEDDARD ON THE [June 16,
obscure. Still it seemed to be free of the underlying musculature
including the longissimus dorsi except Just at the spinous processes
of the vertebre.
The rhomboideus (or retrahens scapule) differs, as I have
pointed out, in Rana guppyr™ and R. esculenta; tor in the former
species it arises from the spine of a vertebra and is thus a true
rhomboideus and not a serratus.
So also is this muscle in Rhinoderma darwini. It arises in front
of the latissimus dorsi and is partly overlapped by it. It is at
first rather broad, but narrows rapidly when it passes under the
cucullaris into a narrower but still flat and strap-shaped muscle
to be attached to the scapula.
The depressor mandibule is partly absent in this frog. There
ig no trace that I could discover of the dorsal part arising near to
the latissimus dorsi. Nor do I think it possible to have missed
this part of the muscle which is so obvious in those frogs where
T have iooked for it. It is important to notice that in this
particular Rhinoderma agrees with its ally Breviceps. ‘The other
portion of this muscle, however, that arising from the skull-wall,
is very large and passes as usual behind the tympanum, its
antithesis, the éemporalis, passing in front of the tympanum.
Both these muscles are large and about equally developed.
The infraspinatus appears to cover the dorsal surface of the
scapula nearly entirely from the extreme edge of its ventral
margin. Only a thin edge of the scapula (indicated by dots in
text-fig. 147) is left exposed, not so much as in Breviceps.
The cucullaris, or levator scapule, is a large and well-developed
muscle as is shown in the figure (text-fig. 147). Its direction is
oblique, the obliquity bemg towards the dorsal median line postero-
anteriorly. In this region it covers a part of the anterior section
of the longissimus dorsi, that inserted on to the head.
The longissimus dorsi is quite like that of Rana, and is shown
together with the adjacent muscles in the accompanying text-
figure (text-fig. 147). They all agree on the whole with those of
Rana, and a reference to my figure will show this agreement in
detail. The iio-lumbaris is rather more marked on the dorsal
surface. The view of this muscle from the inside of the body is
rather different from that of Rana. It runs uninterruptedly
over the transverse processes of four vertebrae (beginning with the
sacral £), and was finally inserted on to the one in front, 7. e. the
fifth reckoning from behind forwards. This is also the fifth
vertebra of the series commencing with the atlas. From this
vertebra to the fourth a band of muscles runs in the same straight
line as the ilo-lumbaris, but not in continuity with its fibres.
This transverse process (that of the fourth vertebra) gives origin
* Anatomy of Pipa, P. Z.S. 1895, p. 835. I am able to confirm this statement
after a re-examination of that frog.
+ Beddard, P.Z.S. 1907, p. 333, text-fig. 94 (Rana guppyi).
‘{ I may note that this vertebra was not fused with the coccyx.
1908. | BATRACHIAN RHINODERMA DARWINI. 689
to the transverso-scapularis muscle *, which is best seen on the
view presented by the dissection now being described, and which
is therefore dealt with here and not in its proper place. The
ilio-lumbaris gives off slips to the transverse processes lying between
its origin from the ilium and its insertion on to the fifth
vertebra ; but these are concealed from view when the body-cavity
is opened and the muscle exposed, for a considerable mass of its
fibres runs directly and without interruption between the extremes
of origin andinsertion. Itsuggests, therefore, the long slip of the
Text-fig. 147.
Dorsal musculature of Rhinoderma darwini.
Cu. Cucullaris. isp. Infra-spimatus. l.coce. Lio-coccygeus. L.d. Latissimus
dorsi. Obl.int. Fan-shaped tract of obliquus internus attached to ilium.
Py. Pyriformis. rh. Rhomboideus.
ilio-lumbaris in Vegalophrys nasuta tT, which appears, however, in
that frog, and in the Pelobatide generally £, to be more completely
differentiated from the rest of the ilio-lumbaris than is the case
with Rhinoderma darwini. More noteworthy is the exact likeness
which this muscle shows to the corresponding muscle in Breviceps.
The coccygeo-sacralis and ilio-coccygeus are present and obvious
as is shown in the figure (text-fig. 147), but have no special
features of interest. On this view the gluteus, the rectus femoris
* It is noteworthy that only one pair of these muscles appears to exist in Rhino-
derma ; for there are two in Rana. The above dissection also showed plainly the
Levator anguli scapule arising from the skull quite as in Rana.
+ P. Z.S. 1907, p. 332, text-fig. 93, I7. lumb.
TPZ. Ss: 1907; pa sil:
AA*
690 MR. F, E. BEDDARD ON THE [June 16,
anticus, and the pyriformis are exposed and very prominent. It
will be seen from the drawing (text-fig. 147) that there is no
possible room for an enormous lymph-heart like that which I have
described and figured in Breviceps on either side of the spine.
Nothing of the kind can, I believe, have been overlooked by me.
A peculiarity of the obliquus internus is shown on the view of the
dorsal musculature which is illustrated in text-fig. 147, obl.int.
A fan-shaped origin of this muscle from the ilium is to be seen
underlying the obliquus externus. The origin is by a head of very
limited extent from the ilium just above the origin of the gluteus
muscle, that is, about halfway down the bone. An iliac origin of
this muscle is of course known in Lana.
§ Musculature of Hyoid.
T have dealt at some length in my paper upon the anatomy of
Breviceps with the hyoid musculature of that frog. This muscu-
lature presents, it will be recollected, more than one peculiar
feature. The corresponding musculature of Rhinoderma does not
present many peculiar features, and agrees on the whole with that
of Rana, differing therefore from its near ally Breviceps, to which
however it presents some likeness, as I have identified, and have
no particular comments to make upon, the following muscles,
which appear to me to be like those of Rana, viz., geniohyordeus and
omohyoideus. The sternohyoideus, on the other hand, appears to
me to be like that of Breviceps in that it consists of two portions
with a quite separate insertion on to the hyoid. The larger and
more superficial half of the muscle has not the same origins as the
sternohyoid of Rana, for there is no sternum to arise from, and it
is simply a continuation of the rectus and completely conceals
(when the animal is viewed from the ventral surface in the usual
position of dissection) the underlying portion of the muscle.
This is very slender, and is connected only in its origin with the
abdominal musculature. It is inserted on. to the hyoid a
considerable distance behind the insertion of the larger half of
the muscle, and the insertions are not continuous. Just below this
muscle lies the pericardial muscle which I describe on another
page (see p. 684). It is interesting to note that the sternohyoid
is complétely free from the shoulder-girdle, and is merely a con-
tinuation of the rectus abdominis.
The petrohyoidet ave as in most other frogs; 7. ¢., there is an
anterior and a posterior petrohyoideus and the latter is divided
into three slips. The latter muscle is entirely inserted upon the
bony thyrohyals, and the three slips of which it is composed form
a continuous mass of muscles which are broad and leave no inter-
spaces. The last of the series, as in some other frogs, lies rather
superficially to the rest and is attached to tle tip of the thyrohyal
bone ventrally. Italso seems to pass beyond it as in Yenophrys*,
and to be therefore associated with the vocal apparatus.
* See Beddard, P. Z. S. 1907, p. 898, text-fig. 238, p.h. 3.
1908. } BATRACHIAN RHINODERMA DARWINI. 691
§ Muscles of the Thigh.
When the skin is removed and the muscles of the thigh inspected
from the inner aspect (text-fig. 148), some difference is visible
from the appearances observable in Rana under similar conditions.
For figures of Rana I may refer to Ecker’s ‘ Anatomy of the
Frog,’ and to a figure of the muscles of the inside of the thigh in
the large Rana guppyi*, illustrating a paper by myself on the
structure of the Pelobatide. I have also figured the correspond-
ing muscles of Breviceps t,an African genus belonging to the same
family (Engystomatide) as that which contains the subject of
the present communication to the Society. Although there are
Text-fig. 148.
Thigh-muscles of Rhinoderma darwini exposed from the inside.
R. Rectus abdominis. v.%.m. Rectus internus minor, where it is attached to
the skin. V.i. Vastus internus. Sa. Sartorius.
differences in detail between the thigh muscles of Rhinoderma
and those of Rana, the general aspect of the muscles—with one im-
portant exception, to be mentioned presently—is much like that of
Rana,and is even definitely more like Rana esculenta than hk. guppyt.
It is easy to recognise the vastus internus, adductor longus, sartorius,
adductor magnus, adductor brevis, rectus internus major and rectus
internus minor, lying in the order named (and commencing of
course at the anterior border of the thigh) and having roughly
much the same proportions as the corresponding muscles of Kana
esculenta. Moreover, it will be noticed that the smallest adductor
muscle lies after the adductor magnus as in Rana esculenta, and
not in front of it as in Rana guppyi. There is, however, visible,
as is well shown in the above figure (text-fig. 148), an important
* P. Z.S. 1907, p. 887, text-fig, 234, + PB. Z.S. 1908, p. 25, text-fig. 6,
692 MR. F. E, BEDDARD ON THE [June 16,.
difference between Rana and Rhinoderma, which is at the same:
time a point of agreement between Lhinoderma and Bbreviceps.
The rectus internus minor arises in Rhinoderma by a number of
more or less separate origins from the skin of the thigh, which
origins are very far from reaching the middle line of the abdomen..
The muscular strands which combine to form this muscle are
spread out upon the skin, when the latter is cut through and
reflected, in a divergent fashion. There are four or five of these
strands, which are naturally flat bands. In Sreviceps such an
extra-skeletal origin of skeletal muscles in the femoral region is
more largely developed than in Lhinoderma; but, as will be seen
by a reference to my paper already quoted, the rectus internus is.
similarly involved in this system of skin muscles.
Viewing the thigh muscles from the dorsal aspect (see text-
fig. 149) and their origins (in some cases) from the back, one-
Text-fig. 149.
Thigh-muscles of Rhinoderma darwini exposed from the outside.
Obl.int. Fan-shaped portion of obliquus internus attached to ilium. Ry.
Pyriformis. R.a. Rectus anticus. v.i.m. Rectus internus minor,
V.i. Vastus internus, lying to the outside of the vastus externus.
obvious and striking difference from Breviceps is to be seen. The
coccyx runs very nearly to the extremity of the body; there is.
not the lengthy exposure of the cloaca with certain muscles
attached thereto and accompanying it that I have figured in
Breviceps*. To the tip of the coccyx is attached the pyriformis,
which in the usual way reaches the femur by passing between the:
vastus externus and the semimembranosus. The disproportion
* P.Z.S. 1908, p: 35, text-fie. 11.
1908. ] BATRACHIAN RHINODERMA DARWINI. 693
between the two last-named muscles is greater than it is represented
tobe in Rana. The figure of Rhinoderma (text-fig. 149) shows that
in that frog the vastus is more than twice the breadth of the semi-
membranous. The rectus anticus of Rhinoderma is peculiar in
that it is a very small and slender muscle covered at its origin
from the ilium by just the front end of the gluteus. It is thus
an inappreciable portion of the triceps femoris complex. The
biceps femoris is much hidden by the vastus externus and semi-
membranosus, between which it lies. Indeed it is only visible for
a very short distance at its insertion. In this the genus agrees
with breviceps.
The semitendinosus is not shown in the two figures (text-figs. 148,
149), which illustrate the musculature of the thigh, since it is
completely hidden on the inner aspect of the thigh by the rectus
internus major. When the latter is cut through and reflected the
semitendinosus is brought into view. It is formed by the union
of two heads as in Rana; but these do not unite until more than
halfway down the thigh. They are moreover fleshy through-
out and roughly speaking of equal size. One head arises, as the
thigh is seen dissected from the ventral aspect, superficially to the
other. It arises from the symphysis pubis in close apposition to
the great adductor and the two rectiabdominis. The second head
is better shown when the first head is cut through and reflected,
since it is distinctly deep of it. It is then seen to run back to
its origin in close apposition to the rectus internus major and to
arise from the pubis very close to it. I observed no tendinous
origin of this head, and no such connection with its head as is
figured and described in the Common Frog.
§ Resumé of facts and Systematic Position of Rhinoderma.
As might be expected from their very different way of life, the
genus ’hinoderma presents, as we have seen, numerous anatomical
differences from its ally Breviceps. Several of these are already
known, and are described in such works as Mr. Boulenger’s
‘Catalogue of the Batrachia Salientia,’ and in Dr. Gadow’s treatise
on Amphibia and Reptiles in the ‘ Cambridge Natural History.’
I leave aside in the present enumeration those external and osteo-
logical features which are dealt with in those and other works.
A general survey of the structure of the muscles shows plainly that
Breviceps has departed much further from the more usual structure
of the Batrachia Salientia than has Rhinoderma. And this state-
ment applies also to the viscera. The extraordinarily enlarged
posterior lymph-hearts of Breviceps are not found in Rhinoderma A
the liver of the latter has the more normal form of that of Rana.
It is, however, in the musculature that the most numerous
divergences between the two types are to be met with. The
remarkable specialisation of the obliquus muscles, which I have
described in detail in Breviceps, does not occur at all in Khino-
derma, which is broadly speaking like Rana in this respect.
694 MR. F. E. BEDDARD ON THE {June 16,
Partly in consequence of this the hyoid musculature of Rhino-
derma is closely like that of Rana, the omohyoid being present,
which muscle has disappeared in Breviceps. On the other hand,
the musculature of the floor of the mouth is quite specialised in
Rhinoderma, and different from that of any other frog the
anatomy of which has been described.
On the other hand, there are a few points in which Rhinoderma
does resemble Breviceps and departs so far from the structure of
Rana. The sternohyoid seems to be a double muscle in both,
though the duplicity of the muscle is not so strongly marked in
Rhinoderma. The attachment and general appearance of the ilio-
lumbaris of Rhinoderma is distinctly like that of Breviceps. In
both, the rectus internus minor of the thigh arises partly from the
skin, and in neither frog is there the dorsal part of the depressor
mandibule muscle present. In my paper upon Breviceps I have
selected 17 characters of importance to distinguish that frog from
Rana. It is only in four of these characters that Rhinoderma
agrees with Breviceps to ditter from Rana.
Nor are there any special points of likeness between the two
genera here considered in any other features not mentioned in the
list of the seventeen principal characters referred to, except, of
course, such general features as both Rhinoderma and Breviceps
share with Rana.
The divergences are most remarkable; and yet there are at
least two equally remarkable points of resemblance, 7. e. the origin
of the rectus internus femoris and the absence of the dorsal part
of the depressor mandibule. There can be no doubt, however,
that, whatever may be the value of these points of resemblance,
the two genera are quite as far removed from each other within
the limits of family relationship as diversity of geographical
position would lead us to expect. A wider knowledge of this
order of animals may reveal surer bases for anatomical criteria.
6. Some Notes upon the Anatomy of Chiromys madagascari-
ensis, with references to other Lemurs. By Frank EH.
Brpparp, M.A., F.R.S., F.Z.8.
[Received May 26, 1908. ]
(Text-figures 150-153.)
The opportunity of examining three specimens of the Aye-Aye
(Chiromys madagascariensis) has enabled me to add a few new
facts to what is already known concerning the structure of this
remarkable Lemur. The three principal Memoirs dealing with
the structure of Chiromys are (in order of appearance) those of
Owen*, Peterst, and Oudemans%. These authors have dealt
with the preceding literature relating to the animal. The
* Trans. Zool. Soe. vol. v. + Abhandl. k. Akad. Wiss. Berlin, 1865.
~ Verh. Akad. Amst. 1890. See also Chapman, P. Ac. Philad. 1900, p. 419.
1908. j AYE-AYE AND OTHER LEMURS. 695
structure of the brain (with which IT am not concerned here,
since the specimens were sold for museum purposes which
rendered the extraction of the brain inadvisable) has been quite
recently dealt with by Dr. Elliot Smith*, who quotes previous
memoirs.
Text-fig. 150.
A portion of intestinal tract of Chiromys.
a.c. Ansa coli. O. Hdge of omentum. S.i. Small intestine. sf. Opening of
duodenum into stomach.
The greater part of the small intestine has been removed : the two cut ends
are connected by a dotted line.
These authors have dealt at length with the external characters,
osteology, visceral and muscular anatomy of Chiromys, and to the
general descriptions given by them of the different organs I have
nothing to add. There are, however, some facts, and these not
altogether without importance, which have not met with much
* Cat. Mus. Roy, Coll. Surgeons, vol. ii. 1902, p. 359.
696 MR. F. E. BEDDARD ON THE [June 16,
attention or have been altogether passed over by the authors
mentioned, largely doubtless by reason of the apparent unim-
portance of those facts at the time when the memoirs in question
were written. Sir Richard Owen found himself obliged to
vindicate the Lemurine affinites of Chiromys from assertions of
its Rodent affinities by comparing it definitely with Sciurus.
Though this is no longer necessary, certain obviously Lemurine
characteristics of Chiromys have not been sufficiently emphasised
either by Owen or by his successors. With these and with some:
other points I propose to deal in the present communication.
Intestinal Tract.—The gut is figured by both Owen* and
Oudemans7, but neither of these figures is at all satisfactory.
I therefore take the present opportunity of refiguring (in text-
fig. 150) a portion of the gut which shows not merely the charac-
teristic ansa coli (flexura coli, colic loop) of Chiromys madagas-
cariensis, but certain mesenteric attachments which are of
importance in the morphology of the intestinal tract of mammals.
Divergent in its general structure from other Lemurs though
Chiromys may be, the intestinal tract points unmistakably to its
affinity with the genera Lemur and Hapalemur, probably with
the subfamily Lemurine. Chiromys possesses in fact, as do those
genera {, a single ansa coli which is a flexure of the colon shortly
after its emergence from the cecum. The two limbs of this loop
were, as In Lemur, closely applied to each other and the loop as a
whole was fully as long as—perhaps even rather longer than—the
loop in the genus Lemur. The loop was perfectly simple and
U-shaped, as is shown in the figure (text-fig. 150), and there was.
no approach to the spiral of the Galaginine, Lorisine, and
Indrisine.
There is some indication in Oudemans’ figure of this loop; but
it is not properly represented; and the various mesenteric
attachments which are of importance from the point of view of a
comparison with other forms are omitted altogether.
As text-figure 150 shows, the omentum is attached to the
region of the colic loop where it bends to the left to become
the straight portion of the transverse colon. Furthermore,
as in some other Lemurs at any rate (there is not at present:
accurate information with regard to the simple forms Micro-
cebus and Cheirogaleus), there is an attachment between the
duodenum where it leaves the stomach and the colon where it.
dips down to form the proximal limb of the ansa coli. This is
also shown in my figure to which I have just referred. The
duodenal attachment is of limited extent, and the omentum is
like that of some other forms in that it is only modified as a
bridge between the stomach and the colon for a portion of the
duodenal region of the former, and as already said for a very
* Loe. cit. pl. 26. figs. 1, 2. + Loe. cit. pl. iii. fig. 14.
{£ For Lemur see Flower (Med. Times & Gazette, 1872), Mitchell (Trans. Zool.
Soc. vol. xvii.), Beddard (P.Z.S. 1908, p. 577); for Hapalemur, Klaatsch (Morph..
Jahrb. xviii. p. 667), :
°
1908. | AYE-AYE AND OTHER LEMURS. 697
limited tract of the other. In my recent memoir upon the
intestine in several mammals*, J have referred to more than one
species in which the attachment of the omentum to the large
intestine is of the same limited extent as in Chiromys. But it is.
greater in the genus Lemur (see text-fig. 151). I may take this
opportunity of remarking that the attachment of the omentum
Text-fig. 151.
A portion of the intestinal tract of Lemur rufifrons corresponding to that of
Chiromys as displayed in text-fig. 150.
Lettering as in text-fig. 150.
to the colon in Hapale penicillata is hardly if at all greater than
in Chiromys. I find myself therefore in disagreement with
Klaatsch, who represents a more lengthy base of insertion of the
omentum upon the colon (in Hapale albicollis).
* “On the Anatomy of Antechinomys, &e.,” P.Z.S. 1908, p. 561.
+ At any ratein L. albifronsand L. rufifrons, where it is attached all over the colic
* loop, and in LZ, branneus, in which species it is attached to halfway down the loop.
698 MR. F. E. BEDDARD ON THE [June 16,
It is interesting to note that, so far as the intestine is concerned,
Chiromys comes closer to the Lemurine than to any of the other
subfamilies of the Lemuroidea. I have pointed out that in the
more specialised Lemuroidea, so far that is to say as concerns the
intestine, the elsewhere characteristic carpal vibrisse have dis-
appeared. This conclusion will require amending since they are
undoubtedly absent in Chiromys, a fact which my colleague
Mr. Pocock observed independently of myself. Nevertheless it
cannot be doubted that in other respects Chiromys is a specialised
Lemur, so that after all the statement may still hold.
The rest of the colon is disposed in a curved course to the
rectum, and there are no traces of any further ans coli.
There is in Chiromys the usual cavo-duodenal ligament, which
was not so clearly a single sheet of membrane as is generally the
case. In one specimen it was single ligament of the usual type:
in the two others a duodeno-renal portion could be differentiated
off, attaching the end of the loop of the duodenum to the right
kidney. An hepato-caval ligament was presenti also in the same
two specimens. On the left side of the body the lieno-rectal
ligament was plain, and also the lieno-renal. J may add that the
right lateral lobe of the liver was attached—naturally on the
right side—by a ligament to the parietes just at the origin of
the dorsal part of the diaphragm. I identified this ligament in
all three individuals. In an example of Lemur brunneus the
right lateral lobe of the liver was also attached to the parietes by
aligament. IJalso recognised in this Lemur the hepato-caval and
hepato-renal ligaments. On the ieft side of the body of this
species of Lemur the lieno-renal and lieno-rectal ligaments were
also very plain.
The vascular system has not been much dealt with by my
predecessors. As is already known the aortic arch gives rise to
two trunks. The intercostal arteries of mammals show some
variation in their mode of origin from the aorta. Here again,
however, there are not sufficient facts known to deduce any
results of classificatory importance. I take this opportunity of
comparing the mode of origin of these little arteries in Chiromys
with those of some other mammals.
In Chiromys the first pair of intercostals arise symmetrically
and very close to each other. The next four are also symmetrical
but a trifle further apart. Then follow two pairs which are as
displayed in the accompanying figure (text-fig. 152), the artery
of the one side being much in advance upon its fellow. The
remaining pairs are symmetrically paired as are the first of
the series, but the distance between the orifices of each pair
of arteries differs. In an example of Pseudochirus peregrinus, of
which I cut open the aorta and examined the mouths of the inter-
costal arteries, they were strictly paired and quite regular. In
Hystriz cristata I counted seven intercostals in front of the
ciaphragm, which however did not commence until the ninth
rib. These arteries were single at their origin from the aorta.
1908. | AYE-AYE AND OTHER LEMURS, 699
In another specimen, however—and this is important as showing
the variation of these structures—the first intercostal was single,
the next two were paired but the left-hand arteriole was smaller
than, and lay behind, the right. Then followed a strictly paired
and equisized couple rather far apart, and after this another pair
closer together. In a Beaver (Castor canadensis) the intercostal
series commenced with a single vessel on the left side; then
followed three pairs the orifices of each pair getting closer
Text-fig, 152.
ee
4
“1
AxX..,
Right-hand figure—The commencement of the aorta of Chiromys cut open to
show origin of intercostals.
a. Commencement of descending aorta. D. Position of diaphragm.
Left-hand figure.—Azygos (Az.) of Chiromys.
Ao. Aorta with some of intercostal branches indicated.
together, then a single median intercostal, followed by three
pairs, and these again by a single median artery. After this
oint the artery traversed the diaphragm, and the intercostals
behind the diaphragm arose singly, and were median in position.
In a Chinchilla (Chinchilla lanigera) the first intercostal was
median and unpaired. Then followed a single vessel on the right
side and then two pairs. The remaining intercostals were median
700 MR. F, E. BEDDARD ON THE [June 16,
and unpaired. In an example of Dasywrus mauget the anterior
series of intercostals were all paired at their origin excepting the
last three in front of the diaphragm which arose by a single
median trunk, each trunk dividing of course into the right and
left intercostal of its segment. After the diaphragm the intercostals
were at least chiefly paired in origin. I have figured these
arteries in the genera of Carnivora*, Helictis, Galictis, and
Suricata, where they arise as paired trunks. ‘There exist,
naturally, descriptions of these arteries in many mammals 7;
but there is need for further collection of facts before they can be
utilised for systematic purposes, for which purposes, however, it is
obvious from what has already been said that their variability
will have to be taken into careful consideration. In the mean-
time I venture to record such facts as I have happened to
ascertain myself, without attempting anything like a revision of
the existing knowledge of these arteries.
I have also ascertained some facts with regard to the venous
system of Chiromys, and I take this opportunity of incorporating
some facts concerning the venous system of MJicrocebus smithii, of
which there is apparently no published information so far as I
can discover. I dissected the postcaval vein (text-fig. 153) in both
of the adult specimens. It was a single vein throughout and lay
as usual to the right of the aorta. The renal veins as is also usual
were asymmetrical, the left lyimg some way behind theright. The
mode of origin of the ovarian veins varies somewhat in mammals.
In Chiromys the vein supplying the right side arose from the
posteaval not far in front of the posterior bifurcation of the
latter. The left vein, on the other hand, arose from the left
renal, as is often the case. This particular asymmetry is very
general in mammals, but it is not always the case that the right
ovarian vein flows into the postcaval so far down. In the male
specimen there was an interesting difference in the place of
influx of the two spermatic veins. That of the right side
corresponded on the whole with the right ovarian vein, But
that of the left side did not arise from the left renal vein but
from the postcaval behind that vein, though some way in front of
the point of origin of the right spermatic vein. The homologue
of the ovarian vein of the female was, however, also present on
the left side. The renal vein, in fact, received a branch at a
point exactly corresponding with the entrance of the left ovarian
vein in the female example. But this vein ended in the muscular
parietes and was evidently concerned alone with the venous
system of the lumbar parietes. These facts are illustrated in the
accompanying text-figure. .
In Microcebus the arrangement of the corresponding veins was
* “On the Anatomy of Helictis personata,’ P.Z.S. 1905, vol. ii. pp. 27, 28,
text-figs. 11, 12.
+ E.g. in Ornithorhynchus, Manners-Smith, P.Z.S. 1894, p. 714; Manatee,
Munie, Trans. Zool. Soc. vol. viii. pl. 26, fig. 44; Horse, Chauveau & Arloing, Traité
WVAnat. Comp. Anim. domest. 1871, f. 555, and many special treatises.
1908. | AYE-AYE AND OTHER LEMURS. 701
somewhat different. In the specimen which I dissected the right
renal vein, which poured its contents into the postcaval headward
of the left renal, was formed by two trunks which, however,
united before entering the postcaval. This may well be a mere
variation. But it is worthy of note that double renal veins
particularly on the right side are very usual among Armadillos.
It is also very general in Tragulus*. And here, again, it is on
the right side that the anomaly occurs.
Text-fig. 153.
—Sp.ve
Postcaval vein of Chiremys, the right-hand figure of a male, the left of a female.
K. Kidney. o.v. Ovarian vein. 7.v. Renal vein. sp.v. Spermatic vein.
The primitive nature of this little Lemur was also shown by
the mode of connection of the ovarian veins with the postcaval.
There was no such asymmetry as has been described in Chiromys,
and which is so general among mammals. The veins in question
are opposite to each other not far from the posterior bifurcation
of the single postcaval. On the left side the vein was more
complex than on the right side. It divided at once into three
vessels. This, however, is not the only peculiarity of these veins
in Microcebus. From each renal vein a slender vessel ran back-
wards parallel with the postcaval trunk, and in the case of that
* McClure, Anat. Anz. Bd. xxix. 1906, p. 875; Beddard, Am. Journ. Anat. 1907,
p. 112.
702 ON THE AYE-AYE AND OTHER LEMURS. [June 16,
of the left side poured its contents into the ovarian vein at its
origin. I did not succeed in tracing the corresponding vein of
the right side all the way; but in any case it commenced
anteriorly in exactly the same way as the other vein. It will be
noted, therefore, that the conditions obtaining in Chiromys could
be arrived at by an obliteration of the connection of the ovarian
vein of the left side with the postcaval, and the retention only of
its embouchure into the renal.
I may observe that in an example of Galago garnetts the origin
of the ovarian veins was quite as in Chiromys. On the other
hand, ina male of Vycticebus tardigradus, the arrangement was
slightly different from, though not in great disagreement with,
Chiromys and Galago. In Nycticebus in fact the left spermatic vem
poured its contents into the left renal. The right, on the other
hand, opened into the postcaval, though very near to the point of
origin of the here more or less symmetrically arising renal veins.
The internal mammary veins of Chiromys run one on each side
and in association with the artery at some distance from the
middle line of the sternum. The two veins are connected with
each branch of the single precaval vein where it bifurcates at the
front end of the thoracic cavity. This is quite the normal
arrangement for these veins, but I mention the facts since there
are sometimes differences. For example, in Lutra vulgaris I
found the internal mammary a single vein on the right side,
but with several branches supplying the left side of the middle
ventral line. Moreover, it is also desirable to note that each of
these veins runs closely accompanied by its artery strictly parallel
to the middle line of the sternum but at some distance from it.
The azygos vein is not figured by previous investigators of the
anatomy of Chiromys. I have examined this vein (text-fig. 152) in
each of the three specimens which I dissected, and most carefully
in the last example which was a young female. It is developed only
upon the right side of the body, as in all Lemurs hitherto studied.
It is a well developed vein and extends backward to nearly the
diaphragm. The first branch supplies two intercostal spaces, and
the last branch of the series but two is also divided in the same
way. There was no trace that I could discover of any corre-
sponding vein upon the left side, neither was there a hemiazygos.
The point at which the intercostal arteries pass outside of the
azygos vein is a matter which varies much among mammals.
But the material does not as yet exist for a use of the facts for
systematic purposes. It is, however, obviously permissible to
state the conditions observed in Chiromys with a view to future
generalisations. I found, in fact, that in this Lemur the first of
the series of intercostal arteries to cross over the azygos vein,
as viewed in the ordinary position of dissection, was that lying
behind the tenth rib. Thereafter all the intercostal arteries
had the same position. In front of this point they underlay
the vein.
ALP. del. West, New na
we. LEUE OCYTOZAO ON: MUSCULI.
1908. ] ON A BLOOD-PARASITE OF WHITE MICE. 703
7, Leucocytozoén musculi, sp. n., a Parasitic Protozobn from
the Blood of White Mice. By Annim Porrer, B.Sc.
Lond., Zoological Research Laboratory, University
College, ben one
[Received May 26, 1908.]
(Plate XX XTX.+ and Text-figure 154.)
CONTENTS. Page
eto WG hom essen ere ee aie aoh ee het saea ee tee. Bao bene TOS
II. Materials ....... Si Resse gaats Ae Ret neha era mame See ne cael eee an OA
III. Occurrence of Parasite Saale sacs eines drach chs oa seem oat eens OA
IV. Methods _...... Les Sod A HRM oe ete Ream ae eames pred (O35
(a) Fresh material — Sea dais Sasa wae tee Se asces Me eeEA OD
(Obixedimaterial 1221.4 SF a eR a SP 706
Weg GenerallS truretume: rh 5: scan. «Soe «sd sne x deitane sae We cee 706
VI. Movements ..... ee dascl cheers ane ei ume OL
(a) Movements of trophozoites .. nica wearer ee eer noe an OD
(6) Movements of merozoites..............0..0600.000.... 709
Wiley DetarledsMiorphologiyes.. .s4.244c-eas4s 20. 5-. eo scsenee sessase - eee 9 LO
(GYR Miro pWozoitess Wee cs gitacccccacae sapcc ree sauce nada ieaen ee 710
(OO) Meri cules sence viese cee Re sea eee eee eee LMR
VILLI. Multiplication .. Spon ndoaRr DORERGReAAe Rar cueet ics a ane RO PRN |
(a) Schizogony .. EARN as cee tera ee aed, ath ger Galak
(6) Possible Meccatione Cl ee AT
IX. Life-History ...... patented: «ees Res
X. Affinities of the Parasite, and Summary... Bo. ES ST 712
XI. References to Literature ............... cc cce sce eee cee cee eee een ees 714
XII. Explanation of Plate XX XIX... 0.0... cceececcee cece cence 715
I. IytrRopuction.
The microscopic organism described in this memoir was found in
the leucocytes or white blood-corpuscles of white mice. It is a
parasitic Protozoon belonging to the class Sporozoa, and being a
blood parasite, it is included in the order Heemosporidia. As it
occurs in leucocytes, it should be placed in the genus Lewco-
cytozoon. Similar parasites have been described during the last
three or four years from the leucocytes of dogs, rats, palm-
squirrels, and cats among mammals, and a few from other
Vertebrates. These will be briefly considered at the end of this
memoir. So far as I know, sucha parasite has not been recorded
previously from the mouse.
As the parasite is shaped like a Gregarine, it is closely related
to the genus Hemogregarina, and some authorities, as Laveran
and Mesnil, would place the Leucocytozoa in that genus. I
prefer, however, to retain the generic name Leucocytozoén, and
since the parasite occurs in the mouse, ws musculus, I propose
the specific name “ musculi” for it. This creation of a new
species is made, not with the intention of merely multiplying
species, but to avoid confusion with. the parasites found in rats.
The nomenclature of these parasites of rats is In a very confused
* Communicated by H. B. FanrHam, D.Sce., A.R.C.Sc., F.Z.S.
+ For explanation of the Plate, see p. 715.
Proc. Zoot. Soc.—1908, No. XLV 45
704 MISS ANNIE PORTER ON A [June 16,
state. The specific name “ muris” has been applied by Balfour
(2, 24]* to the parasite he described from Mus decumanus at
Khartoum. A similar parasite has been described by Adie [1]
from Mus rattus in the Punjaub, under the name of L. ratti, while
a third has been described by Cleland [7] from Perth, Western
Australia.
I think it is very probable that this Leucocytozoon is different
from that in rats. In this connection it is worthy of note that
the Trypanosomes of rats and mice belong to different species.
The name that I propose for this parasite from the leucocytes
of white mice is, then, Lewcocytozoon musculi.
Il. MATERIALS.
The first infected animal received was a white mouse which
came from a stock kept at St. Mary’s Hospital Medical School.
This white mouse, when in an almost comatose condition, was
brought to the notice of Dr. Fantham. Some symptoms were
feebleness of movement (constantly moving slowly, more or less
in a circle, with a tremor or waltz, though the animal did not
appear to be related to Japanese waltzing mice), partial closure of
the eyes with discharge therefrom, and incontinency of urine.
A few parasites were found in the peripheral blood of this
mouse, but the rodent soon died. As soon after death as possible,
smears were made of the heart, spleen, kidney, liver and gut-
contents. There was an extravasation of blood into the gut of
the mouse and in this parasites were found. However, in the
gut many Bacteria occurred, and it is not suggested that the
Leucocytozodn was the sole cause of the death of the mouse.
A second white mouse, from the same stock as the first one,
was procured and kept under observation for nearly three weeks.
A few parasites only occurred in its peripheral blood. Accidental
death overtook this mouse and immediately smears were made of
its internal organs. In these parasites were seen but in much
fewer numbers than in the first specimen. A third mouse from
the same stock was found to be infected to a greater extent than
the preceding one.
The first two mice were scarcely, if at all, verminous, but on
the third one, many lice, Hematopinus spinulosus, were seen.
‘These were dissected and examined for stages in the life-history
of the parasite, as stated in the sequel.
The material was examined as far as possible in both the living
and fixed condition, as the examination of living material is most
important and tends to be overlooked.
The above-mentioned material was kindly given to me by
Dr. Fantham.
IIL. OccuRRENCE OF PARASITE.
The blood of the mouse contained parasites of two forms :—
(1) large,.vermiform organisms, free-living in the plasma; and
* The numbers in square brackets refer to the: list of Literature at the end of
this paper. wae : Satie BLO Ula een
1908. | BLOOD-PARASITE OF WHITE MICE. 705
(2) smaller forms, which are cytozoic. At first the latter are free
in the plasma, then later, they penetrate usually into mononuclear
leucocytes where they feed and grow, finally assuming again
the free form. Very rarely do they occur in polymorphonuclear
corpuscles, but specimens were seen in transitional corpuscles,
No parasites were seen within tissue-cells, whether of liver,
heart, spleen, lung, kidney or gut. They seem purely blood
parasites, though their presence appears to cause enormous hyper-
trophy of cells in their neighbourhood, This was especially well
seen in the liver smears. The parasites were most abundant in
these smears and in the portal blood, were fairly numerous in
heart and kidney smears, but very few occurred in either lung or
spleen preparations, though the latter organ (spleen) was enlarged.
Bone-marrow preparations were also made, and schizogony was
found to occur therein.
In the case of the first mouse, extravasation of blood into the
gut had occurred and the gut-contents showed free parasites in
this blood. live parasites were studied usually from freshly
shed peripheral blood.
Examination of the organs of the lice showed vermicule stages
of the parasite in the gut and Malpighian tubes.
The Leucocytozoa were never associated with Trypanosomes in
these mice, though such an association has been described for
other Leucocytozoa [1] [7].
The lice appear to act as mechanical agents in propagating
the disease, for lice removed from the third mouse and placed on
another resulted in a very slight infection of the hitherto
unaffected one. In the case of L. canis, Gerrard [9] reported
that puppies, which were placed together, were cross-infected by
the agency of ticks.
IV. Meruops.
(a) Fresh material,
Freshly drawn blood, usually taken from the tip of the tail of
the mouse under examination, was mixed with a small quantity
of normal saline solution, to which in most cases a little alkaline
methylene-blue was added. A drop of the mixture was examined
in the well of a micro-slide provided with such a depression, or
else on the slide or on the cover-slip, forming a hanging drop in
the latter case. The cover-glass was always vaselined round the
edges and so air in quantity was excluded from the preparation.
In this way, living parasites could be observed for several hours.
Intra vitam staining with methylene-blue could also be thus
accomplished. Much time was spent in examining the parasites
in the fresh state.
Lice found on the third mouse were carefully examined for
probable stages in an Invertebrate host. Hemiptera removed
‘from the mouse were at once dissected-in normal saline solution.
Especial attention was paid to the alimentary canal, Malpighian
45*
706 , MISS ANNIE PORTER ON A [June 16,
tubules, salivary glands, reproductive organs, and body-cavity of
the lice. Smears of these organs, fixed wet with osmic vapour,
were afterwards stained and examined microscopically, but fresh
preparations in normal saline to which a little methylene-blue had
been added, were also examined in this case.
(b) Fixed material.
The blood smears were usually fixed wet with osmic vapour o1
with osmic vapour and alcohol. Occasionally they were allowed
to dry quickly and afterwards treated with methyl alcohol before
staining. The chief stains used were Giemsa’s mixture of azur 11.
and eosin, Loeftler’s alkaline methylene-blue, and Delafield’s
hematoxylin, the latter used alone or sometimes followed by
safranin. Azur 1. followed by lichtgriin was tried occasionally,
and safranin alone—suggested by the presence of a refractile cyst-
like envelope round some of the parasites—was tried, but found
to be too transparent and diffuse. The best results were obtained
with Giemsa’s stain, alkaline methylene-blue, and hematoxylin.
The same stains were used for the organ smears of lice.
The preparations were usually mounted in Canada balsam,
sometimes left uncovered.
The various methods outlined above were tried for the purpose
of correlation and corroboration, and to eliminate possible errors.
V. GENERAL STRUCTURE.
The general shape of this parasite, which occurs either inside a
leucocyte or free in the blood-plasma, may be described as vermi-
form or gregariniform, in fact, that of a vermicule (Pl. XXXIX.
figs. 1,2). However, there is very often no marked difference
between the ends, which are then somewhat rounded; and
further, as some of the parasites are comparatively broad for
their length, they may be quite accurately described as bean-
shaped or reniform. The size varied from 17 to 7p in length
and 5°9u to 4 in breadth.
The free parasites, averaging 10°94 long by 5:1 broad, are
usually surrounded by a cytocyst which is very refractile and
does not stain at all easily (Pl. XX XIX. fig. 8). Their cytoplasm
is rich in granules which react vigorously toward stains and -
thereby obscure the oval nucleus lying beneath them. The
distribution of the granules varies, and this accounts for the
differences in the nuclear apparatus as seen in the figures. The
nucleus is generally nearer one end in position.
Differences between the ends of the organism occur in cytozoic
forms. One end may be larger than the other. This may be
the natural result of the cytozoic habit, the organism assuming
the form most suitable to a limited space, or it may be due to the
twisting of the body on itself and within the spherical leucocyte,
which results in the “thin edge” or ‘ tailed” appearance of
some of the parasites (text-fig. 154 D-K). U-shaped forms as
1908. | BLOOD-PARASITE OF WHITE MICE. 707
described by Wenyon [14] were not seen. The average size of
endo-globular forms was 8 long by 5 broad.
Among the free forms, some are relatively shorter and broader
than the others (Pl. XX XIX. figs. 1, 6). Some writers [12, 13]
would consider these broader forms as female Leucocytozoa, while
the longer thinner ones (Pl. XX XIX. figs. 5, 8) would be regarded
as male. Ihave no evidence to support this view, but regard
them as extreme forms of a continuous series.
VI. Movements.
(a) Movements of Trophozoites.
When a parasite has penetrated a leucocyte, it remains at first
near the periphery and so directs its movements that it ultimately
comes to surround the nucleus to a very great extent.
Osmotic diffusion between host cell and parasite occurs and
produces movement within the cell. When this is very vigorous,
the oscillation produced may be so great as to cause semi-rotation
of the leucocyte, even to the extent of 180°.
The movement of the parasite is more noticeable at one end.
This appears to advance steadily by an outflow of the cytoplasm.
This outflow is easily seen, for the protoplasm is richly granular
and stains readily intra vitam with methylene-blue. The para-
site lies near to the nucleus of the leucocyte, and its presence
causes a movement of the nucleoplasm which appears in a state
of agitation. Osmosis seems to be taking place from the leucocyte
nucleus to the parasite, and the latter rapidly grows during this
period (text-fig. 154, A—E).
The gliding movement continues, and the nucleus of the
leucocyte, which was originally globular, becomes much altered in
shape. Its nuclear membrane becomes less and less distinct, and
at times it resembles a somewhat lens-shaped mass lying within
the horns of a crescent formed by the parasite (text-fig. 154, D, E).
The organism continues its gliding movements and one end
becomes much larger than the other. This is the more obvious
and may be termed the anterior end. A comma-like appearance
then results (text-fig. 154, G-J). The posterior end, being
thinner, might be termed a tail, though this is not an accurate
description. It certainly appears very filamentous at times, but
this is because the organism has turned on its side and so exposed
an edge to view.
On examining the surface of the parasite, numerous granules
are evident. These are usually arranged in more or less regular
rows. During movement of the organism as a whole, movements
of the rows occur, and this suggests that the arrangement of the
granules in rows is due to myonemes upon the body. Stained
preparations show that such is the case (Pl. XX XIX. fig. 7).
The slow gliding movement would be due then to contractions
of these myonemes, and, further, the bowed appearance of the
parasite within the cell could be explained as being the result
708 MISS ANNIE PORTER ON A [June 16,
of the stronger contractions of the myonemes at the inner edge
of the organism with successively smaller contractions of the
myonemes toward the outer (greater) curvature, where there
would bearing of relaxed myonemes. The axial line of the body,
where, judging from the action of the granules, the movement is
least, would be, according to this arrangement, a neutral area,
neither contraction nor relaxation of the myonemes occurring
within it.
Text-fig. 154.
Living Leucocytozoén musculi.
Diagrams of a young intra-corpuscular parasite observed in the living conditiom
for seven hours.
In these diagrams the cytoplasm of the parasite is represented by small dots, more
closely aggregated where the protoplasm stained more deeply. Vacuoles are left
clear. The chromatin of the parasite is represented black in G-K, and the nucleus
of the leucocyte is shown by shaded lines. The outline of the leucocyte is indicated.
by a circular area. :
The nucleus of the leucocyte is not yet much displaced by the parasite, and no:
marked hypertrophy of the host-cell has yet occurred.
The time required for the assumption of the various forms.
1908. ] BLOOD-PARASITE OF WHITE MICE. 709
mentioned varies with the specimen and with the stage of develop-
ment at which the parasite may be. At the initiation of the
trophic, intracellular phase of the life-history, the activity of the
organisms is very great, and, correlated with this, there is the
phase of most vigorous motion. The infected leucocyte figured
in text-figure 154, A—K, was observed for a period of seven hours,
and even then the exit of the parasite from the leucocyte was
not seen, owing to the death of the leucocyte.
' An early stage is seen in text-figure 154, A, where the parasite
was lying in the leucocyte so that the nucleus of the latter
was practically in full view. About twenty-five minutes later,
the organism had spread itself out and become crescentic in
form (text-fig. 154, B, C). The movements till then were very
vigorous, and much oscillation of granules in both parasite and
jeucocyte nucleus was observed. Half an hour after the obser-
vation was commenced, the forward movements of the parasite
slowed very much, though the movements of the granules remained
much the same (text-fig. 154, D). About the same time the
protoplasm of the parasite began to move from the periphery
towards the anterior end, which became much more globose and
began to curve (text-fig. 154, EH, F). An anterior end could be
distinguished definitely as such at the end of 55 minutes (text-
fig. 154, F). Two hours after the first observation was made the
parasite had assumed the “comma” form and a vesicle began to
make its appearance (text-fig. 154, F—J). It remained in this
condition for along time, the only change that occurred being that
the protoplasm became much more granular and there was a
slight retraction of the posterior end, while the vacuole increased
in size and chromatin masses appeared in it (text-fig. 154, G—K).
Death .of the leucocyte prevented further reliable observations
being made on this parasite, but the exit of the organism from its
host-cell was seen in other specimens (Pl. XX XIX. figs. 18, 19).
Extrusion of the parasite is brought about by internal pressure.
The Leucocytozo6n moves forwards with a slow, gliding movement.
This continues steadily until extrusion is completed. There is
slight resistance at the periphery of the leucocyte, but on the exit
of the parasite, the protoplasm of the host-cell closes up and the
point of exit is invisible (Pl. XX XIX. figs. 18,19). The freed
parasite remains quiescent for a short time after leaving the
corpuscle,
Stages of endoglobular parasites are figured in Pl. XX XIX.
figs. 15, 16, 17.
(b) Movements of the small, free Merozortes. )
In some of the peripheral blood mixed with: normal saline to
which a little methylene-blue was added, sausage- or bean-shaped
bodies occurred (Pl. XX XIX. fig. 13). These moved fairly
actively, the movements being much more energetic if the
organism were in the neighbourhood of a leucocyte. The body
of the parasite would seem to be somewhat flattened, for it is
710 _ MISS ANNIE PORTER ON A [June 16,
able to turn freely somewhat in the fashion of Myctotherus.
The anterior end remains still, but the posterior part of the
body turns over so that the upper surface becomes folded over
the under, which, at the distal end, is now uppermost. The line
of folding is somewhat oblique. Reversal of this movement
occurs, and the organism appears to roll from side to side as a
result of the combined movements.
Accompanying this movement there is a second. One sur-
face of the body of the parasite contracts, and as a result, the
ends of the body approximate somewhat more closely to one
another than before. Relaxation follows, and the body straightens
with a jerk which has the effect of forcibly propelling the
organism forwards.
The path of the organism is never straight. Movement appears
to be initiated at either end indifferently. The path is often very
restricted and the organism remains for long periods at practically
the same spot, though one of its ends may have vibrated in
practically every direction.
VII. Derattep MorpHoioey.
(a) The Trophozoite.
Ectoplasm.—In the trophozoite this isnot markedly differentiated.
It appears as a somewhat clearer portion in some specimens.
Contractile elements or myonemes are present, arranged in longi-
tudinal rows. These myonemesare very evident in some specimens
(Pl. XX XIX. figs. 7, 11) and often are well seen in the region of
the nucleus. A refractile cytocyst is often present, and when
this is well marked, a clear space usually intervenes between
it and the ectoplasm (Pl. XX XIX. figs. 8, 9,10, 11,12). The
cytocyst is always thin and membranous.
Endoplasm.—This is richly granular (Pl. XX XIX. figs. 1, 4)
and the granules react vigorously towards stains. In some speci-
mens the endoplasm appears almost alveolar (Pl. XX XIX. fig. 12)
owing to the disposition of the granules in regular networks.
There are concentrations of granules beneath the myonemes
and also in the region of the nuclear membrane. Frequently a
relatively clear space appears near one end, almost suggestive of a
large vacuole (Pl. XX XIX. figs. 3, 4,7,21). In a few specimens
isolated chromatic granules are present (P]. XX XIX. fig. 8).
The latter do not seem to have any direct connection with the
nucleus.
Nucleus.—This is circular or oval, approximately central in
position or sometimes nearer to one end and possesses a definite
nuclear membrane. Its chromatin is diffusely spread within and
much achromatic substance is present. The structure of the
nucleus may best be described as vesicular. There does not
appear to bea karyosome as described by James [10] in one of his
forms of Leucocytozodn canis. In its general structure and
behaviour towards stains, the nucleus of this parasite is very
Suggestive of that of Trypanosoma raje.
1908. | BLOOD-PARASITE OF WHITE MICE. 711
The nuclear membrane is definite. It has a somewhat beaded
appearance in some stained preparations and that can also be seen
in living material (Pl. XX XIX. figs. 2, 4, 6, 16). Extra-nuclear
chromatin also occurs, for in favourably stained specimens, a
chromatic cap can be distinguished at either end (Pl. XX XIX.
fig. 15), and isolated granules also occur (Pl. XX XIX. fig. 8).
(b) The vermicules of Hematopinus spinulosus.
The vermicules of the lice present the following appearance :—
Their ectoplasm is not well differentiated from the endoplasm,
but indications of myonemes are present and are best seen at the
ends of the organism. ,
The endoplasm is granular and much as in the trophozoites, but
marked concentrations of granules do not occur.
The nucleus in some appears to lie In a vacuole, and in most
specimens there is a clearer portion in the neighbourhood of
the nucleus. Its chromatin is more abundant than in the blood-
inhabiting forms, and the nuclear membrane is fairly distinct
(Pl. XX XIX. figs. 23, 241.
These vermicules were about 8°8u long and 1:4u broad.
VIII. Mutrirerication.
(a) Schizogony.
Examination of bone-marrow showed the presence of small,
oval cysts (cytocysts) about 13-1u long by 8°9u broad (Pl. XX XIX.
fig. 20). These contained relatively few (about 12) but distinct,
small forms with a definite vermicular or reniform contour.
Each of these small vermicules was about 4:4 long by ‘8p broad
(Pl. XX XIX. fig. 20). They are the merozoites, produced by
the multiple or asexual fission of a schizont inside the cytocyst.
The general protoplasm was granular. Remains of the nucleus
of the leucocyte host were seen on the side of the cyst. Inside
the cyst were also some remains of the residual protoplasm of the
schizont. By the dehiscence of the cyst these merozoites are set
free in the blood-plasma, where they become vermicules or young
trophozoites.
(b) Possible Association of Trophozoites.
Two parasites lying in one corpuscle were observed (Pl. XX XIX.
fig. 21), or two which had just left the corpuscles (Pl. XX XIX.
fig. 22). One such case, of two parasites lying in the remains of
a leucocyte, suggested possible association (Pl. XX XIX. fig. 21).
Here the two forms, partially free from the host-cell or leucocyte,
came in contact with one another and became enveloped in a
common cytocyst. The nucleus of one appeared to come nearer the
area common to the two than the other, and a chromatin mass
was seen in the common area of the couple. This suggested that
transference of chromatin takes place from one parasite to the
712 MISS ANNIE PORTER ON A [June 16,
other. However, the formation of a definite zygote from these
associated forms was, unfortunately, not seen.
Again, in Pl. XX XIX. fig. 22, those authorities who believe in
differentiation into male and female forms, would see a female
form (macrogametocyte) in the upper, broader and granular
parasite, and a male form (microgametocyte) in the lower, longer
and narrower form. I do not go quite so far personally, as I have
doubts of sexual differentiation, preferring the series view (see
page 707). However, in the present state of our knowledge, the
suggestion of sexual forms is worthy of note.
TX. Lire-History.
Commencing with the free vermicule which may be either a
sporozoite or a merozoite, the following sequence probably occurs.
The parasite lives awhile in the plasma as a small, active form
(Pl. XX XIX. fig. 13). This ultimately penetrates a leucocyte
and grows actively there as an endoglobular trophozoite. <A
cytocyst may or may not form around it (Pl. XX XIX. figs. 15,
16,17). After a time, it becomes free in the plasma, rupturing
the host-celi as it issues, but leaving little or no trace of its
presence behind. In the plasma it assumes the free trophozoite
form. Association between trophozoites may perhaps occur (cf.
Pl. XXXITX. figs. 21, 22), and-the result of this is probably a
zygote, which one would expect to find in the louse, on homology
with the malarial parasite—but more definite information is
lacking.
Other free forms may reach the bone-marrow. There encyst-
ment occurs, and a schizont, ina thin but distinct cytocyst, is
produced. This schizont by multiple fission gives rise to relatively
few but definite merozoites. By the rupture of the cyst, these
are set free into the blood-stream where growth again occurs,
leading to adult trophozoites. This is the schizogonic cycle of the
parasite.
Ecto-parasitic on the mouse was the louse, Hamatopinus
spinulosus. By the bite of this louse infected blood passes from
the mouse to the mouth and gut of the invertebrate host, and so.
we find the small, gregariniform vermicules shown in Pl. XX XIX.
figs. 23, 24. These ultimately reach the saliva of the louse and:
by this insect are probably transferred to another mouse. Perhaps
a sexual cycle of the Leucocytozoén occurs in the louse, but of
this I have, unfortunately, no definite evidence. Probably the
louse is merely a mechanical agent in spreading the infection. —
X. AFFINITIES OF THE PARASITE, AND SUMMARY.
A finities of Parasite.
The term Leucocytozoén was used by Danilewsky [8] in 1890.
for vermiform parasites stated to occur in the leucocytes of certain,
birds. Danilewsky confined his observations to fresh preparations.
1908. | BLOOD=PARASITE OF WHITE MICE. 713
Similar parasites were afterwards studied by Berestneff, Sacharoff,
Ziemann, and Laveran. The latter states that Danilewsky’s
parasites really occur in immature erythroblasts. The parasite
was stated later by Schaudinn [12] in 1905 to be a stage in the
life-history of a Spirochete. The matter is fully discussed by
James [10], and as it is very controversial, need not be dwelt on
further here.
Bentley [3] and James |10] independently described a parasite
from the leucocytes of pariah dogs in India in 1905. This was a
true Leucocytozodn and is known as Z. canis. A memoir by
Christophers [5] on the same parasite appeared in 1906, and the
following year he worked out the sexual cycle in the tick [6].
Other Leucocytozoa have been described in mammals by Patton
[11] in the Indian palm-squirrel, Yunambulus pennantii, under
the name of LZ. funambuli; in Malay dogs by Gerrard [9] and
Wenyon [14]; and in rats in various parts of the world by
Balfour [2] from leucocytes of Mus decumanus at Khartoum,
by Adie [1] from IMus rattus in the Punjaub (as ZL. ratti), and by
Cleland [7 | from rats in Perth, Western Australia. These para-
sites in rats are probably best known as L. muris (cf. page 704).
A form known as ZL. felis has been described, I believe, from the
Indian bazaar cat in Madras by Christophers and Patton.
Outside mammals, from other vertebrates, we have recorded
L. ranarum from the Amphibian Leptodactylus ocellatus by
Carini [14]; and quite recently Z. lovati, a form from the leuco-
cytes of grouse by Seligmann and Sambon [13].
These are, I think, all or nearly all the Leucocytozoa recorded
to date (May, 1908). Whether they have any intimate connection
with Flagellates, such as that suggested by Schaudinn [12] remains
to be seen—probably they have not.
These parasites are all gregariniform. Those in mammals are
much about the same size, and occur both free in the plasma or
endoglobular in the leucocytes. Schizogony is known in the bone-
marrow [5] and liver.
_ Laveran suggested that the Leucocytozoa should be included in
the genus Hemogregarina. However, as they occur in a non-
hemoglobin-containing host, viz. leucocytes, while strict Hemo-
gregarines are found in erythrocytes, I think the generic name
Leucocytozoén should be retained. The name for the parasite
described in this memoir for the first time in the leucocytes of the
mouse is L. muscult.
Summary.
(1) The forms of Z. mwsculi here described occur in the
mononuclear and transitional leucocytes of white mice.
(2) The free trophozoites in the plasma are gregariniform or
reniform vermicules, the average size being 10°94 long and 5:lu
broad (cf. Pl. XX XTX. figs. 1-12).
(3) The free trophozoites sooner or later enter leucocytes of the
host and grow at the expense of the nucleus of the leucocyte
714 MISS ANNIE PORTER ON A [June 16,
(of. Karyolysus among the Heemogregarines). A thin cytocyst is
formed probably by the host-cell around the parasite (Pl. XX XTX.
figs. 16, 17).
(4) Endoglobular forms are on the average 8p long by 5p broad
(cf. Pl. XX XIX. figs. 15, 16, 17).
(5) Schizogony takes place in the bone-marrow. An endo-
globular trophozoite rounds itself off and becomes a schizont,
breaking up into merozoites, each about 4°44 long and ‘8u broad
(Pl. XX XIX. fig. 20). This again suggests affinities with Karyo-
lysus, judging by Labbe’s figures of schizogony in Karyolysus.
(6) Two parasites may sometimes occur within one host-cell.
Two such forms suggesting the beginning of association are shown
in Pl. XXXIX. figs. 21, 22.
Differentiation into male and female forms could not be made
out with certainty, though some parasites were shorter, broader,
and more granular than others.
(7) Vermicules were found in the gut and Malpighian tubules
of lice, ectoparasitic upon the mice, but unfortunately no evidence
of a sexual cycle in the louse was obtainable. Perhaps the lice
merely act as mechanical agents in the transfer of the parasites
among the mice.
(8) The parasites are found in smears from the heart and liver
in abundance. They are less numerous in spleen and kidney
smears, also in the bone-marrow and peripheral circulation. They
were not abundant in the latter.
(9) No Trypanosomes were seen in the infected mice.
(10) The movements of the vermicules or trophozoites of this
parasite in the blood-plasma of its Vertebrate host are fully
described in section VI. of this memoir.
XI. RererENces To LITERATURE.
f1] Avie, J. R. 1906.—“ Note on a Leucocytozoén found in
Mus rattus in the Punjaub.” Journ. Trop. Med., vol. ix.
pp. 325-326.
[2] Batrour, A. 1905-6--“A Hemogregarine of Mammals
. (HZ. jacult).” Journ. Trop. Med., vol. viii. pp. 241-244, and
vol. 1x. p. 82.
[24] Baurour, A. 1906.—Second Report of the Wellcome
Research Labs. at Gordon College, Khartoum. Lewco-
cytozoon muris, pp. 110-111.
[3] Benritey, C. 1905.—‘“ Preliminary Note on a Leucoeytozoon
of the Dog.” Brit. Med. Journ., 1905 (1), p. 988.
[4] Carin1, A. 1907.—“Un_ leucocitozoo del Leptodactylus
ocellatus,” Rev. da Soc. Scient. de Sio0 Paolo, no. 8.
[5] Curistopuers, S. R. 1906.—‘ Leucocytozoén canis.” Sci.
Mem. Govt. India, No. 26.
{6] CuristopHers, 8S. R. 1907.—“ The Sexual Cycle of Leuwco-
cytozoén canis in ‘the Tick.” Sci. Mem. Govt. India,
No. 28.
1908. ] BLOOD-PARASITE OF WHITE MICE, 715
[7] Cuntanp, J. B. 1906.—“ The Hemogregarine of Mammals
(H. balfouri), and some notes on Rats.” Journ. Trop.
Med., vol. ix. p. 296.
[8} Danitewsky, 1890.—“ La développement des parasites
malariques dans les leucocytes des Oiseaux (Leuco-
cytozoaires).” Ann. Inst. Pasteur, 1890, p. 427.
[9] GerrarD, P. N. 1906.—“ On a Protozoén Parasite found in
the Polymorphonuclear Leucocytes of a Dog.” Journ.
Hygiene, vi. pp. 229-230.
[10] James, 8. P. 1905.—“ On a Parasite found in the white
corpuscles of the Blood of Dogs.” Sci. Mem. Govt. India,
No. 14.
[11] Parron, W. 8. 1906.—“ Ona Parasite found in the white
corpuscles of the Blood of Palm Squirrels.” Sci. Mem.
Govt. India, No. 24.
[12] Scuaupiny, F. 1904.—“ Generations- und Wirtswechsel bei
Trypanosome und Spirochete.” Arb. a. d. Kaiserlichen
Gesundheitsamte, xx. p. 387.
[13] Senremann, C. G., and Samson, L. W. 1907.-—“ Preliminary
Note on a Leucocytozoén found in the blood of Red Grouse
(Lagopus scoticus).” Lancet, Sept. 21, 1907, p. 829.
[14] Wenyon, C. M. 1906.—“‘ Report on Dr. P. N. Gerrard’s
preparations....” Journ. Hygiene, vi. pp. 231-236,
1 plate.
XII. EXPLANATION OF PLATE XXXIX.
The figures on the Plate XXXIX. were outlined with camera lucida (Abbé),
using Zeiss =; inch achromatic and 2 mm. apochromatic oil-immersion objectives
with compensating oculars 4 and 8. Zeiss E objective was also used for fresh
preparations.
Fig. 1. Shows a free trophozoite which is broad and short. Stained Giemsa.
1700. Liver smear of 1st mouse.
2. Free trophozoite, rather large. Shows one chromosome. Giemsa. > 1700.
Liver smear of 2nd mouse.
3. Smaller, possibly younger, form. It has one patch of chromatin. Giemsa.
< 1700. Liver smear of 1st mouse.
4, Form showing dotted ‘nucleus, aiso dots along the nuclear membrane.
Giemsa. 1700. Heart smear of 1st mouse.
5. Parasite showing central chromatin filaments. The Leucocytozoén is not so
markedly granular as some. Giemsa. X 1700. Liver smear of 1st mouse.
Fig. 6. Cf. fig. 1. The nucleus is vesicular and chromatin is distributed round the
nuclear membrane. Giemsa. > 1700. Liver smear of 2nd mouse.
7. Shows pale, vesicular nucleus, also myonemes (my) are well seen. The ends
are somewhat pointed. Giemsa. x 2250. Liver smear of 2nd mouse.
8. Parasite in a clear cytocyst (cy). It has a patch of extra-nuclear chromatin
in the endoplasm. Giemsa. %X°1700. Liver smear of 1st mouse.
9. Parasite in cytocyst. Capsof chromatin present. Giemsa. X 1700. Kidney
smear of 1st mouse.
Fig. 10. Parasite in cytocyst with deeper staining nucleus; also very granular
protoplasm. Giemsa. 1700. Liver smear of 1st mouse.
Fig. 11. Shows pale-staining vesicular nucleus and definite myonemes (my). In
cytocyst (ey). Giemsa. X 1700. Liver smear of 1st mouse.
Fig. 12. Broad parasite within cytocyst (cy), showing alveolar protoplasm. Giemsa.
1700. Spleen smear of 1st mouse.
Fig. 13. Free forms in plasma. Methylene-blue. Peripheral blood of 3rd mouse.
Fig. 14. Free vermicule penetrating mononuclear leucocyte. Methylene-blue.
X 1200. Peripheral blood of 3rd mouse.
716 MR. E. MEYRICK ON [June 16,
Fig. 15, Young intra-corpuscular stage. Parasite has a chromatin cap at either end.
Giemsa. 1700. Liver smear of 2nd mouse.
Figs. 16, 17. Endoglobular parasites in mononuclear Leucocytes. Giemsa. X 1700.
Liver smear of 1st mouse.
. 18, Shows egress of parasite from its host-cell. Intra vitam staining with
methylene-blue. X 1200. Peripheral blood of 3rd mouse.
Fig. 19. Egress of parasite shown in fig. 18 from leucocyte just completed, after
rotation of leucocyte. Intra vitam. Methylene-blue. Peripheral blood
of 3rd mouse.
Fig. 20. Schizont within cytocyst in bone-marrow. ‘Twelve merozoites are present,
together with residual protoplasm (7.pl.). Methylene-blue. XX 1600.
Bone-marrow of 2nd mouse.
Fig. 21. Possible association of the two Leucocytozoa. A common cytocyst is
present. Chromatin is showing passing across from one to the other.
One Leucocytozoén is not entirely free from its host-cell. Hach of the
couple has a vacuole. Giemsa. 1700. Liver smear of 1st mouse.
Fig, 22. Possible association. Both of the Leucocytozoa are free from their host-cell
or cells. Liver smear of lst mouse. Giemsa. 1700.
Figs. 23, 24. Vermicules from the midgut of Hematopinus spinulosus. Giemsa.
x 1700.
Fi
gg
8. Descriptions of African Micro-Lepidoptera.
By H. Mryricr, B.A., F.R.S., F.Z.8.
[Received June 18, 1907. ]
This paper is a contribution towards the knowledge of the Tor-
tricina and Tineina of the African region, which are at present
very little known. The material for it was received from various
collectors, but for the greater part of it I am indebted to Mr. A.
J. T. Janse, of Pretoria, whose specimens are also particularly
notable for their fine condition. Mr. Janse further furnished
notes on localities and time of appearance, based on additional
examples retained by himself besides those which he sent me.
In addition to some known species recorded from Africa for the
first time, 108 species and 10 genera are described as new.
EuUCOSMIDA.
LopesiA AEOLOPA Meyr.
This species, described from India and Ceylon (Journ. Bomb.
N. H. Soe. xvii. p. 976), I possess also from Grahamstown, Cape
Colony, and the island of Réunion.
POLYCHROSIS HARMONIA, Sp. Nn.
3 2. 10-13 mm. Head, palpi, and thorax light ochreous, face
whitish-suffused. Abdomen whitish-ochreous, sometimes suffused
with grey. Fore wings elongate, slightly dilated posteriorly,
costa slightly arched, apex obtuse, termen obliquely rounded; pale
brownish-ochreous ; markings deep yellow-ochreous; an inwardly
oblique spot beneath fold before middle (representing lower portion
of angle of a strongly angulated basal patch, of which remainder is
obsolete); central fascia rather narrow, sometimes with a few
1908. ] AFRICAN MICRO-LEPIDOPTERA, 117
black scales, posterior edge excavated near dorsum; a blotch
before middle of termen, connected by a narrow projection with
apex; three short oblique dark fuscous strigule on costa poste-
riorly, beneath first a smal! deep ochreous spot; a fine line of
blackish scales along termen ; cilia yellow-ochreous, on costa paler
spotted with fuscous. Hind wings grey ; cilia whitish-grey, with
grey subbasal shade.
TRANSVAAL, Pretoria district, from November to March (Janse) ;
nine specimens.
PoLycHROSIS SCORPIODES, sp. 0.
3 2. 17-18 mm. Head whitish-ochreous, face more whitish,
Palpi ochreous slightly sprinkled with dark fuscous, second joint
whitish towards apex and beneath. Thorax light ochreous mixed
with brown. Abdomen grey, anal tuft pale ochreous. Fore wings
elongate, posteriorly dilated, costa slightly arched, apex obtuse,
termen obliquely rounded ; ochreous-whitish, with some scattered
ochreous-brownish strigule, towards dorsum and _ posteriorly
more or less marbled with pale bluish-silvery-grey ; costa directly -
strigulated with blackish; a basal patch of suffused ochreous-
brown strigulation, marked with black on fold, outer edge obtusely
angulated below middle; central fascia ochreous-brown, irregular,
broadly interrupted below middle so as to leave beneath only a
triangular spot before tornus, marked in middle with several small
irregular black spots, and followed beneath costa by a suffused
blue-grey patch ; a roundish ochreous-brown patch before middle
of termen, marked with black on each side; a small ochreous-
brown apical spot marked with black. Hind wings grey; cilia
whitish-grey, with grey subbasal shade.
TRANSVAAL, Pretoria district, Pietersburg, from August to
December (Janse); two specimens.
Kccorsis Z.
Assuming for the present that this genus is tenable, and dis-
tinguished by the peculiar thickened dorsal lobe of hind wings,
then the following species is referable to it.
EiCCOPSIS ACROCOSMA, sp. 0.
6.15 mm. Head and thorax fuscous irrorated with pale
specks. Palpi rather long, porrected, second joint triangularly
scaled, terminal joint moderately long, pointed ; second joint with
base and lower edge whitish, upper edge with a metallic-bluish
stripe, intermediate space orange crossed by an oblique black line
and edged beneath with blackish, terminal joint blackish with
upper edge metallic-bluish, Abdomen fuscous. Fore wings
elongate, posteriorly considerably dilated, costa gently arched,
apex obtuse, termen sinuate, somewhat oblique; rather dark
fuscous, tips of scales whitish, appearing to form a very fine and
regular transverse striation ; costa marked with very short, oblique
718 MR. E. MEYRICK ON [June 16,
dark fuscous strigule, between and beneath which are some leaden-
grey scales; a small irregular black spot in dise at 2, irregularly
centred with metallic-blue ; a triangular orange apical spot, marked
with three blue-leaden-metallic streaks, uppermost short, slender,
two lower stronger and partly edged with black, converging to
apex : cilia fuscous sprinkled with whitish, with darker subbasal
shade. Hind wings rather dark fuscous; cilia fuscous.
Nyassa-LANnb, Songwe Valley; one specimen.
The singular apparently corresponding coloration of the palpi
and tips of fore wings has doubtless some explanation in local
circumstances.
ARGYROPLOCE ORTHACTA, Sp. 0.
2.18mm. Head whitish-fuscous tinged with reddish, crown
irrorated with blackish. Palpi moderate, porrected, triangularly
scaled, ochreous-whitish tinged with fuscous. Thorax reddish-
fuscous mixed with blackish. Abdomen grey. Fore wings
elongate-triangular, costa gently arched, apex obtuse, termen
- younded, somewhat oblique; greyish-olive, with irregular grey-
whitish strie rismg from pairs of whitish strigule on costa; a
large sharply-marked dark fuscous basal patch mixed with dark
red-brown, outer edge straight, oblique, finely edged with white,
reaching on dorsum to beyond middle; a round patch of ground-
colour towards termen above middle outlined by pale striz, and
broadly suffused posteriorly with dark reddish-fuscous ; a streak
of reddish suffusion mixed with blackish along termen; cilia
ochreous-grey irrovated with whitish, indistinctly barred with
darker grey, and with a dark grey subbasal line. Hind wings
fuscous, suffused with dark fuscous towards termen ; cilia fuscous,
paler towards tips.
TRANSVAAL, Pretoria and Zoutpansberg districts, in September
and from December to March (Janse); one specimen.
PAMPLUSIA SARDONIA, Sp. 0.
¢ @. 15-18 mm. Head, palpi, and thorax pale grey, irrorated
with grey-whitish, and more or less suffused with pale crimson,
palpi rather long. Abdomen rather elongate, dark grey. Fore
wings elongate, gradually dilated, costa slightly arched, apex
obtuse, termen almost straight, oblique; dark leaden-grey, with
numerous irregular dull crimson-reddish transverse strige, some-
times largely suffused together; costa obliquely strigulated with
dark fuscous and whitish; cilia fuscous mixed with reddish, with
a whitish basal line on termen, apical third light reddish. Hind
wings dark fuscous; cilia fuscous, with dark fusecous subbasal
shade, tips pale brownish round apex.
TrANsvAAL, N.E. Pretoria district, in December and January
(Janse) ; three specimens.
E,NARMONIA BATRACHOPA, Sp. 0.
3 2. 18-22 mm. Head pale ochreous, face and sides suffused
1908. } AFRICAN MICRO-LEPIDOPTERA. 719
with dark fuscous. Palpi dark fuscous. Thorax with double
posterior crest, pale ochreous, broadly suffused laterally with dark
fuscous, sometimes mixed with ferruginous. Abdomen rather
dark fuscous, in ¢ with large expansible pale greyish-ochreous
genital tuft. Posterior tibie in ¢ with very large dense brush
of greyish-white hairs above, including a blackish-grey tuft towards
apex. Fore wings elongate-triangular, costa moderately arched,
apex obtuse, termen almost straight, somewhat oblique; in ¢ with
rather strong dorsal scale-projections towards base and beyond
middle; light brownish or brownish-ochreous, obscurely strigu-
lated with whitish, especially in ¢ towards dorsum and termen ;
most of wing except dorsum and termen in ¢ irrorated with
fuscous-crimson and strigulated with black, in Q largely suffused
with dark fuscous sometimes mixed with dull crimson, posterior
edge of this area in Q well-defined and forming a blunt wedge-
shaped projection extending downwards from upper half to near
termen before middle, in ¢ indicated but inconspicuous ; costa
marked with some pale strigule, and in ¢ with several small
fuscous-crimson and black spots; sometimes a distinct pale
greenish discal dot beyond middle; a semioval olive-greenish or
dark reddish-fuscous patch mixed with black occupying anterior
half of dorsum, its central area more or less suffused with whitish-
ochreous, especially in g; an acute-triangular olive-greenish spot
mixed with black on dorsum before tornus: cilia whitish-fuscous
irrorated and barred on upper half of termen with ferruginous,
on lower half with blackish. Hind wings in ¢ with a deep semi-
circular excavation on termen above tornus; dark fuscous, darker
posteriorly ; in ¢ vein | 4 clothed with rough whitish hairs; cilia
whitish-grey, with grey subbasal shade.
Cars Cotony, Hastern portion; NataL; TRANSVAAL; RHODESIA ;
bred in July and August from larve feeding on citrous fruits,
stone-fruits, and guavas, sometimes doing extensive damage
(Lounsbury) ; four specimens.
EucosMA LEUCOPETRA, sp. n.
6. 12-14 mm. Head whitish, sides of crown suffused with
ochrecus. Palpi moderate, porrected, second joint with long rough
projecting scales above and beneath; white, second joint tinged
with pale brownish-ochrecus. Thorax whitish, irregularly tinged
with brownish and spotted with dark fuscous. Abdomen fuscous,
more or less suffused with yellowish or whitish towards base, anal
tuft whitish-ochreous. Fore wings elongate, somewhat dilated
posteriorly, costa slightly arched, fold extending to 2, apex obtuse,
termen slightly indented above middle, rather oblique; ferruginous-
ochreous, often more or less suffused with fuscous, variably and
irregularly strigulated with white and dark fuscous ; posterior *
of costa with five pairs of oblique white strigule, whence proceed
short irregular leaden-metallic strigee; a large irregular suffused
white median dorsal blotch, including several grey strigule ;
ocellus large, white, irregular, containing several pale metallic-
Proc. Zoou. Soc.—1908, No. XLVI. A6
720 MR. E. MEYRICK ON [June 16,
grey spots in two transverse series, and two black dots between
those; sometimes several other black dots scattered round this:
cilia dark reddish-grey finely irrorated with white, on tornus with
a white patch. Hind wings with 3 and 4 stalked; grey; cilia
pale grey or whitish-grey, with faint darker subbasal line.
TRANSVAAL, Pretoria district, in December, January, and July
(Janse) ; seven specimens.
CROCIDOSEMA PLEBEIANA Z.
TRANSVAAL, Pretoria, Pietersburg, January to May (Jazse).
I take the opportunity of noting that Steganoptycha obscura
E. Woll., described from St. Helena, is certainly a synonym of
this species, which is now common in suitable localities in all
quarters of the globe, doubtless artificially mtroduced with its
food-plant in gardens. The genus is hardly worth separating from
Oydia, the single species agreeing in all respects except the peculiar
tuft of hind wings in ¢.
CYDIA TUMULATA, Sp. 0.
@. 16mm. Head and thorax pale brownish-ochreous, crown
with a dark fuscous central stripe. Palpi pale brownish-ochreous
sprinkled with fuscous. Abdomen grey. Fore wings elongate,
posteriorly dilated, costa gently arched, apex obtuse, termen
sinuate, little oblique ; pale brownish-ochreous, irregularly mixed
and spotted with grey, with scattered dark fuscous and black
scales; costa strigulated with dark fuscous, between these whitish-
tinged, posteriorly giving rise to three oblique brown strige
alternating with leaden-grey strigee; two dark fuscous spots on
dorsum, first slightly oblique at 4, second larger and somewhat
elongate-triangular at 3, space between these suffusedly striated
with whitish ; ocellus indistinctly margined with leaden-metallic
a small brown apical spot ; some minute black dots on termen :
cilia brownish sprinkled with whitish, round apex and on upper
half of termen suffused with blackish. Hind wings with 3 and
4 stalked ; grey, dark posteriorly ; cilia grey.
TRANSVAAL, Pretoria district, in December and January (Janse) ;
one specimen.
CYDIA ISOGRAMMA, Sp. n.
3 2. 12-16 mm. Head fuscous, sides and face more or less
mixed with ochreous. Palpi fuscous, second joint usually with
a central ochreous spot. Thorax fuscous, patagia pale ochreous.
Abdomen whitish-ochreous irrorated with grey. Fore wings
elongate, slightly dilated posteriorly, costa gently arched, apex
obtuse, termen sinuate-indented, little oblique ; whitish-ochreous,
becoming yellow-ochreous towards costa posteriorly and termen ;
margins of cell, internal veins, and veins between cell and termen
more or less lied with blackish-fuscous or sometimes partially
with pale leaden-grey, with some additional interneural lines,
1908. } AFRICAN MICRO-LEPIDOPTERA. 721
ground-colour more or less whitish-suffused between these; dorsal
area marked with irregular pale leaden-grey lines, and some
scattered blackish-fuscous marks, with a narrow blackish-fuscous
patch or streak along dorsum from near base to about 2, anda
small irregular ill-defined blackish-fuscous dorsal spot at 3; costa
blackish-fuscous, with more or less paired silvery-whitish strigule
throughout, on posterior 4 emitting oblique silvery-grey strigz ;
a slender ferruginous-ochreous streak runs from middle of costa
through posterior margin of cell to posterior dorsal spot ; a silvery
mark along lower half of termen: cilia whitish-ochreous, with a
dark grey bar marked with dark fuscous in middle of termen, and
base more or less marked with dark fuscous round tornus and
above apex. Hind wings with 3 and 4 connate; grey; cilia
whitish, with grey subbasal line.
TRANSVAAL, Pretoria, in March (Janse); Cryton, Patipola,
Maskeliya, and Diyatalawa, from March to September (Green,
Pole, Alston, Fletcher); eight specimens.
CyDIA PSAMMACTA, Sp. n.
3 Q. 17-19 mm. Head whitish-ochreous. Palpi with long
rough scales, pale fuscous, towards tip of second joint above
ochreous-whitish. Thorax whitish-ochreous, partially tinged with
brownish. Abdomen light fuscous. Fore wings elongate, costa
gently arched, apex obtuse, termen somewhat indented-sinuate,
rather oblique ; whitish-ochreous, suffusedly strigulated with
fuscous suffused with brownish-ochreous, except on a large clear
Space occupying most of dorsal area towards middle, posterior
half suffused with brownish-ochreous; costa and dorsum shortly
strigulated with blackish, posterior half of costa with five pairs
of whitish strigule, whence proceed oblique leaden-grey strigze ;
ocellus represented by an irregular silvery-grey whitish blotch,
cut by a slender transverse streak of ground-colour containing
several irregular black marks, and with some other scattered
black marks before and above it: cilia pale ochreous irrorated
with whitish, round apex and upper half of termen suffused with
dark fuscous. Hind wings with 35 and 4 stalked; grey; cilia
whitish-grey, with darker subbasal line.
TRANSVAAL, Pretoria district, in December and January (Janse) ;
three specimens.
LASPEYRESIA DELINEANA Walk.
(Grapholitha delineana Walk. Cat. xxvill. 389; G. apicatana,
ib. 390; Laspeyresia isacma Meyr. Journ. Bomb. N. H. Soc. xviii.
144.)
TRANSVAAL, Pretoria, in January (Janse); Mavririus, Port
Louis, in May. Occurs also in India and China.
LASPEYRESIA TRICENTRA Meyr.
TRANSVAAL, Pretoria, Pietersburg, from December to February
(Janse).
46*
(22, MR, E, MEYRICK ON [June 16,
A common Indian species, probably imported into Africa with
its food-plant (Crotalaria). African specimens show some indi-
vidual variation, and the hind wings are generally rather lighter,
but I can find no reliable distinction. In India there are several
very closely allied species,
TORTRICGIDS.
CACOECIA HEDRASTIS, sp. n.
3. 12-13 mm. Head whitish-ochreous. Palpi moderately
long, whitish-ochreous irrorated with dark fuscous. Thorax
whitish-ochreous irrorated with fuscous. Abdomen grey, anal
tuft ochreous-whitish. Fore wings suboblong, anteriorly rather
strongly, posteriorly slightly arched, apex obtuse, termen straight,
rather oblique, costal fold narrow, extending from base to beyond
1; whitish-ochreous, with some scattered “dreams scales; basal
patch indicated by increased fuscous irroration but not defined,
on costal fold dark fuscous; central fascia moderately broad, dark
fuscous, lighter and more ill-defined on lower half of posterior
edge; costal patch flattened-triangular, dark fuscous, its apex
connected by a dark fuscous striga with tornus; a dark fuscous
striga along upper part of termen ; cilia whitish-ochreous. Hind
wings grey; cilia ochreous-grey-whitish, with a grey shade.
TRANSVAAL, Pretoria district, from January to March (Janse) ;
two specimens,
ToRTRIX CEDROTA, Sp. 0,
g.12mm. Head and thorax pale yellowish-ochreous. Palpi
moderate, porrected, ochreous-yellowish, basal half of second joint
irrorated with blackish. Abdomen grey, anal tuft pale yellowish.
Fore wings elongate, posteriorly dilated, costa gently arched, apex
obtuse, termen obliquely rounded ; glossy whitish-ochreous ; basal
patch suffused with ochreous-yellowish, costal edge and dorsal half
irrorated with black, outer edge obtusely angulated in middle ;
central fascia moderately broad, suffused with ochreous-yellowish
and irrorated with blackish, anteriorly edged with ochreous-yellow;
costal patch triangular, dark grey mixed with black, edged with
ochreous-yellowish, and connected with tornus by an ochreous-
yellow striga sprinkled with blackish ; some yellowish strigule
irrorated with blackish along termen: cilia whitish-yellowish.
Hind wings grey; cilia yellowish-grey-whitish, with a grey line.
TRANSVAAL, Pretoria, in March (Janse); one specimen.
TorvTRIX AFRICANA Wals.
(Conchylis africana Wals. Trans. Ent. Soc. Lond. 1881, 227,
pl. x. 6.)
This species, which I have from the Transvaal and Natal, I
refer here.
1908. | AFRICAN MICRO-LEPIDOPTERA, 723
TORTRIX CAPENSANA Walk.
(Teras capensana Walk. Cat. xxviii. 295; 7’. reciprocana, ib.
295; 7. meridionana, ib. 295; Tortrix capitana Feld. Reis. Nov.
pl. exxxix. 48, 49; Cacoecia adustana Wals. Trans. Ent. Soe.
Lond. 1881, 222, pl. x. 1; Lozotenia dorsiplagana, ib. 223,
ple)
I am of opinion that all these names represent only one variable
species, which is common and generally distributed in Transvaal,
Natal, and Cape Colony, and is a true Tortrix.
ToRTRIX IOCOMA, sp. n.
3. 18-22 mm. Head orange. Palpi moderate, porrected,
ochreous-orange, sometimes sprinkled with fuscous. Thorax pale
ochreous-yellowish, anteriorly suffused with orange. Abdomen
light ochreous-yellowish. Fore wings elongate, hardly dilated,
costa gently arched, apex obtuse, termen nearly straight, rather
oblique; light ochreous-yellowish, costa slightly yellower: cilia
pale yellowish. Hind wings very pale grey, suffused with whitish-
yellowish towards costa and apex; cilia pale yellowish.
Transvaal, N.K. Pretoria district, in December and January
(Janse) ; four specimens.
TORTRIX AGROECA, Sp. n.
36. 20mm. Head, palpi, and thorax brownish-ochreous, palp
moderate, porrected. Abdomen dark grey, anal tuft light yellowish.
Fore wings suboblong, slightly dilated posteriorly, costa gently
arched near base, thence nearly straight, apex obtuse, termen
nearly straight, rather oblique; brownish-ochreous: cilia light
brownish-ochreous. Hind wings fuscous-grey finely irrorated
with blackish ; cilia pale brownish-ochreous.
TRANSVAAL, Pretoria district, im December (/anse); one
specimen.
GELECHIAD2.
PALTODORA PSACASTA, Sp. n.
3 2. 12-14 mm. Head and thorax whitish-yellowish. Palpi
yellow-whitish, lower half of second joint fuscous, tuft short,
spreading. Antenne fuscous. Abdomen yellow-whitish, in g
with a central black spot. Fore wings elongate, very narrow,
costa gently arched, apex pointed, termen very “obliquely rounded ;
whitish-yellow, with brassy reflections; plical and second discal
stigmata black; some light brown suffusion, sometimes sprinkled
oh darker fuscous, forming usually a streak along basal 2 of
costa, a spot resting on plical stigma, a smaller spot obliquely
above and beyond this, a small spot before tornus, a larger spot
on costa at 2, a patch between second discal stigma and termen,
and an apical patch, but these vary somewhat in development :
cilia whitish-yellowish, above apex with a fuscous spot, sometimes
with two or three cloudy fuscous antemedian dots on termen.
124 - -‘MR. E, MEYRICK ON [June 16,
Hind wings with 6 and 7 stalked ; grey-whitish ; cilia pale whitish-
yellowish.
TRANSVAAL, Pretoria district, from November to February
(Janse); five specimens.
ARISTOTELIA SIROTA, Sp. 1.
6. 15mm. Head and thorax ochreous-whitish sprinkled with
brown. Palpi moderately long, rather thickened with scales,
whitish, second joint light brownish. Antennz whitish-ochreous.
Abdomen light grey, three basal segments ochreous-yellowish.
Fore wings elongate, narrow, costa gently arched, apex acute,
termen extremely obliquely rounded; 6 out of 7 near base ;
ochreous-whitish, faintly streaked longitudinally with pale
brownish-ochreous ; some dark fuscous irroration tending to form
indistinct streaks on veins; plical stigma small, fuscous, second
discal formed by a small round spot of dark fuscous irroration :
cilia whitish-ochreous tinged with brownish. Hind wings grey-
whitish ; cilia whitish-ochreous.
TRANSVAAL, Pretoria district, in August, September, and
January (Janse); one specimen,
ARISTOTELIA ACRENA, Sp. 0.
©.12mm. Head and thorax whitish-ochreous mixed with
deep ochreous-yellow. Palpi moderately long, loosely scaled,
whitish, second joint mixed with yellowish and sprinkled with
dark fuscous, terminal joint sprinkled with ochreous. Antenne
pale ochreous. Abdomen ochreous-yellowish, mixed with dark
fuscous on sides and posteriorly. Fore wings elongate, very
narrow, costa slightly arched, apex acute, produced, termen ex-
tremely oblique, sinuate ; 7 and 8 out of 6; whitish-ochreous some-
what sprinkled with fuscous; some deep ochreous-yellow suffusion
forming streaks beneath costa and along submedian fold, and a
broad patch occupying most of postmedian area except a narrow
terminal fuscous fascia sprinkled with dark fuscous ; a blackish
dot at base of costa, two beneath costa at + and 2, two on fold
obliquely bey ond these respectively, one in disc beyond middle,
and one at 2: cilia ochreous-yellowish, round apex with a few dark
fuscous specks. Hind wings grey; cilia whitish-ochreous tinged
with grey.
TRANSVAAL, Pietersburg, in April (Janse) ; one specimen.
ARISTOTELIA PELTOSEMA Low.
TRANSVAAL, Pretoria, in August and January (Janse).
I imagine this wide-ranging species, which occurs in Ceylon
and Australia, is attached to some garden plant.
EPHYSTERIS, n. g.
Head smooth; tongue developed. Antenne 4, in ¢ simple,
basal joint moderately long, without pecten. Labial palpi mode-
rately long, recurved, second joint beneath with rough projecting
1908. | AFRICAN MICRO-LEPIDOPTERA, 725.
scales, terminal joint shorter than second, loosely scaled, acute.
Maxillary palpi rudimentary. Posterior tibie clothed with long
hairs above. Fore wings with 2-5 parallel, 7, 8, and 9 out of 6,
7 to costa, 11 from middle. Hind wings under 1, elongate-
trapezoidal, apex strongly produced, termen emarginate, cilia 3;
3 and 4 connate or stalked, 5 rather approximated, 6 and 7
tolerably parallel.
Apparently a development of Gnorimoschema.
EPHYSTERIS CHERSHA, Sp. 0.
3 2. 10-12 mm. Head ochreous-whitish, more or less mixed
with dark fuscous. Palpi whitish mixed with dark fuscous. An-
tenne dark fuscous. Thorax light brownish-ochreous, more or
less irrorated with dark fuscous. Abdomen dark grey, second
and third segments more or less suffused with ochreous-yellowish
dorsally. Fore wings lanceolate, apex produced, acute ; brownish-
ochreous more or less irrorated with dark fuscous ; an indistinct
dark fuscous spot on fold towards base; stigmata cloudy, dark
fuscous, sometimes large, first discal little before middle, plical
obliquely before first discal ; sometimes an indistinct pale ochreous
spot on costa at 2: cilia grey, with some black specks. Hind
wings rather dark grey ; cilia grey.
TRANSVAAL, Pretoria, from September to November (Janse) ;
four specimens.
GELECHIA AROTRIAS, sp. 0.
@. 21mm. Head and palpi pale ochreous-yellowish, base of
palpi fuscous, terminal joint almost as long as second. Antenne
dark fuscous. Thorax dark fuscous, with broad central pale
ochreous-yellowish stripe. Abdomen fuscous. Fore wings
elongate, rather narrow, costa gently arched, apex round-pointed,
termen very obliquely rounded; dark purplish-fuscous, lighter
and crimson-tinged towards base of costa, darkest above dorsal
stripe; a pale ochreous-yellowish dorsal stripe from base to near
tornus, rather broad towards middle but narrowed to extremities,
before posterior extremity emitting an oblique bar to 2 of disc;
(cilia imperfect). Hind wings rather light fuscous, somewhat
darker posteriorly ; cilia pale fuscous.
Nata, Weenen, in October (Spiller) ; one specimen.
GELECHIA TRISIGNIS, sp. n.
[email protected] mm. Head ochreous-white, sides narrowly blackish-
fuscous. Palpi white, from base to above middle of second joint
dark fuscous, terminal joint as long as second. Antenne black.
Thorax dark bronzy-fuscous, with broad white central stripe.
Abdomen grey. Fore wings elongate, narrow, costa gently arched,
apex round-pointed, termen hardly rounded, very oblique; very
dark bronzy-fuscous ; a rather broad ochreous-whitish costal stripe
from base to near apex, narrowed posteriorly; a rather narrower
ochreous-white dorsal stripe from base to tornus, narrowed to
726 MR. E. MEYRICK ON [June 16,
extremities: cilia bronzy-fuscous, becoming whitish-fuscous to-
wards tornus, on costal streak ochreous-white. Hind wings light
grey ; cilia whitish-fuscous, darker round apex.
TRANSVAAL, Pretoria district, from October to December (Janse) ;
one specimen.
PHTHORIM#A OPERCULELLA Zell.
TRANSVAAL, Pretoria and Pietersburg districts, from September
to May (Janse); common.
This destructive potato-feeding species has not previously been
recorded from South Africa, though now introduced into many
other countries.
LEcrrHoceRA CHOLOPIS Meyr.
TRANSVAAL, Pretoria and Pietersburg districts, from September
to April (Janse); Narau; NyassaLanD, Fort Johnston. Widely
distributed in India.
PrILOTHYRIS PURPUREA Wals.
The female, which is not described by Lord Walsingham, is
without the white patch which characterises the hind wings of
the male, these wings being wholly blackish.
DRAGMATUCHA, 0. g.
Head with appressed hairs; ocelli absent; tongue small. An-
tenn +, basal joint moderately elongate, without pecten. Labial
palpi very long, recurved, second joint thickened with dense
scales, forming a loose spreading tuft towards apex beneath,
terminal joint longer than second, slender, acute. Maxillary palpi
rudimentary. Posterior tibize clothed with very long rough
spreading hairs. Fore wings with 2 and 4 short-stalked from
angle, 3 absent, 5 approximated, 7 to costa, 8 and 9 out of 7,
11 from beyond middle. Hind wings over 1, trapezoidal, apex
obtuse, termen hardly sinuate, cilia 3: 3 absent, 5 parallel, 6 and
7 stalked.
Apparently related to some extent to 7Zimyra.
DRAGMATUCHA PROAULA, Sp. 0.
©. 30 mm. Head dark fuscous, face and sides of crown
ochreous-yellow. Palpi light ochreous-yellowish, second joint
suffused with orange-yellow, terminal joint sprinkled with dark
fuscous. Antenne whitish-ochreous, basal joint and a short
streak towards base above blackish. Thorax dark purplish-
fuscous, posterior margin narrowly ochreous-yellowish. Abdomen
fuscous, dorsally coppery-tinged, segmental margins and apex
ochreous-yellow. Legs yellow banded with blackish, hairs of
posterior tibize mixed with whitish. Fore wings elongate, rather
narrow, costa gently arched, apex. obtuse, termen somewhat
obliquely rounded; dark purplish-fuscous; a small irregular pale
1 908. | AFRICAN MICRO-LEPIDOPTERA. 727
ochreous-yellow spot at base; two narrow irregular whitish-
ochreous transverse fascie, dilated towards costa and becoming
deep ochreous-yellow on costal edge, at about 2 and + re spectively :
cilia grey mixed with blackish, on upper half of ter men ochreous-
yellow. Hind wings rather dark grey ; cilia ochreous-yellowish,
at apex with a grey patch, towards middle of termen with an
indistinct grey shade.
TRANSVAAL, Pietersburg, in September, October, and March
(Janse) ; one specimen.
BRACHMIA MUSICOPA, Sp. 0.
3 2. 13-14mm. Head and thorax ochreous-whitish, patagia
pale brownish. Palpi ochreous-whitish, second joint flatly com-
pressed, loosely scaled beneath and with scales somewhat expanded
towards apex above. Antenne dark grey. Abdomen pale fuscous,
suffused with whitish-ochreous above towards base, anal tuft pale
yellowish. Fore wings elongate, narrow, costa gently arched,
apex obtuse, termen very obliquely rounded; 7 and 8 stalked,
7 to apex, 9 separate; purplish-grey, veins marked with well-
defined lines of black and brown irroration; a fine costal streak
from base to #, another subcostal from base to middle, and a more
or less broad dorsal suffusion from base to # ochreous-whitish,
sometimes partially tinged with yellowish; second discal stigma
represented by a round whitish-yellowish spot, marked beneath or
almost wholly suffused with ochreous-brown, sometimes centred
with dark fuscous: cilia purplish-fuscous finely irrorated with
whitish, basal third dark fuscous spotted with ochreous-whitish.
Hind wings whitish-grey ; cilia yellow-whitish tinged with grey
round apex, with a faint grey shade.
TRANSVAAL, Pretoria district, in January (Janse); two
specimens.
BRACHMIA STERICTIS, Sp. n.
©. 15-16 mm. Head, thorax, and abdomen pale ochreous-
yellowish. Palpi whitish-ochreous, second joint infuscated except
towards apex. Antenne whitish-ochreous, basal joint partly
infuscated. Fore wings elongate, costa gently arched, apex
obtuse, termen obliquely rounded; 7 and 8 stalked, 7 to apex,
9 separate; whitish-ochreous tinged with yellow-ochreous; ex-
treme base of costa dark fuscous; stigmata small, blackish, plical
obliquely before first discal, second discal larger ; a row of ill-
defined blackish dots immediately before termen and apical portion
of costa: cilia whitish-ochreous tinged with yellow-ochreous.
Hind wings pale whitish-grey ; cilia pale whitish-ochreous tinged
with grey.
TRANSVAAL, Pretoria district, in January and February (Janse) ;
two specimens.
BRACHMIA SERIALIS, gp. 0.
@.14 mm. Head and thorax greyish-ochreous mixed with
728 MR. E. MEYRICK ON [June 16,
fuscous. Palpi whitish-ochreous, second joint suffused with
fuscous. Antenne pale ochreous suffusedly ringed with fuscous.
Abdomen fuscous. Fore wings elongate, costa gently arched,
apex obtuse, termen rounded, rather oblique; 7 and 8 stalked,
7 to apex, 9 separate; pale ochreous suffusedly irrorated with
fuscous; blackish basal dots on costa and in middle; stigmata
blackish, plical rather obliquely before first discal; a row of unde-
fined blackish dots immediately before termen and apical part of
costa, terminating in a small suffused dark fuscous pretornal spot :
cilia pale ochreous irrorated with fuscous, Hind wings grey ;
cilia light grey.
TRANSVAAL, Pretoria, in January (Janse); one specimen.
POLYHYMNO TROPA, sp. 0.
@. 8mm. Head white, posterior edge of crown dark fuscous,
collar white. Palpi white, terminal joint grey except apex. An-
tenn white, with a blackish line above. Thorax white, patagia
mixed with fuscous. Abdomen grey, apex white. Fore wings
elongate, rather narrow, costa slightly arched, apex acute, pro-
duced, termen sinuate, rather strongly oblique ; bronzy-fuscous
irrorated with dark fuscous; markings white; a median longi-
tudinal streak from base to middle, thence bent to meet at a very
acute angle a narrow very oblique streak from middle of costa,
the bent portion closely followed by a similar parallel streak
meeting the same costal streak produced ; two shorter less oblique
costal streaks posteriorly, second mostly in cilia and edged with
blackish ; a narrow irregular streak along posterior part of fold ;
a narrow almost marginal streak along lower half of termen,
extended round tornus: cilia grey, above apex with a white spot
margined posteriorly by an oblique dark fuscous line, beneath
apex with a white patch, above and below which are fragments of
a black antemedian line. Hind wings grey; cilia light grey.
TRANSVAAL, Pretoria district, from September to November and
in February (Janse); one specimen.
XYLORYCTIDAE.
Eporycta, n. g.
Head with appressed scales, side-tufts somewhat spreading ;
ocelli and tongue apparently absent. Antenne 2, in ¢ shortly
unipectinated (1) and ciliated, basal joint moderate, without.
pecten. Labial palpi very long, recurved, second joint thickened
with appressed scales, terminal as Jong as second, slender, acute.
Maxillary palpi rudimentary. Posterior tibis clothed with hairs.
above. Fore wings with 2 from 2, 3 from angle, 7 and 8 stalked,
7 to apex, 11 from middle. Hind wings over 1, trapezoidal-ovate,
cilia 2; 3 and 4 connate, 5 parallel, 6 and 7 short stalked.
Clearly related to the Australian genus Xylorycta.
Ld
1908.1] AFRICAN MICRO-LEPIDOPTERA. 129:
EporycTA TARBALBEA, Sp. 0.
3. 25 mm. Head, palpi, antenne, thorax, and abdomen
ochreous-whitish, palpi with second joint and anterior edge of
terminal mixed with fuscous. Fore wings elongate, rather narrow,,
costa gently arched, apex obtuse, termen slightly rounded, rather
strongly oblique ; ochreous-white ; costa, apex, and termen some-
what suffused narrowly with pale yellow-ochreous: cilia yellow-
whitish. Hind wings pale whitish-grey, slightly ochreous-tinged 3.
cilia ochreous-whitish.
TRANSVAAL, N.E. Pretoria district, in January (Janse); one
specimen.
NEPHANTIS XYSTOPALA, sp. 0.
6. 24-26 mm. Head and thorax ochreous-whitish. Palpi
whitish-ochreous, second joint white above towards apex. An-
tenn dark fuscous, towards base white. Abdomen with ochreous-
whitish scales overlying ferruginous-brown surface, which tends
to appear in longitudinal lines. Fore wings elongate, costa
moderately arched, apex tolerably pointed, termen rather sinuate,
oblique; white; a narrow light brownish-ochreous stripe above
middle from base to apex: cilia white. Hind wings light grey,
paler and tinged with whitish-ochreous anteriorly ; cilia whitish-
ochreous, tips white.
TRANSVAAL, Pretoria district, in December and January (Janse) ;
eight specimens.
ODITES CITRANTHA, sp. n.
3.21 mm. Head dark fuscous, face pale yellowish. (Palpi
broken.) Antenne pale fuscous, ciliations 14. Thorax light
yellow. Abdomen ochreous-yellowish. Fore wings elongate, costa
gently arched, apex obtuse, termen rounded, hardly oblique; 2
from near angle; clear yellow; discal stigmata minute, blackish :
cilia yellow. Hind wings ochreous-whitish ; cilia whitish-
yellowish.
Nara, Durban, in August (Leigh); one specimen.
ODITES PROCELLOSA, sp. 0.
3 @.17-18mm. Head, palpi, antenne, thorax, and abdomen
whitish-ochreous tinged with yellowish; palpi with lower = of
second joint and base of terminal joint dark fuscous; antennal
ciliations 2. Fore wings elongate, slightly dilated posteriorly,
costa gently arched, faintly sinuate in middle, apex obtuse, termen
rounded, hardly oblique; 2 from near angle; whitish-ochreous
tinged with yellow, thinly and irregularly sprinkled with fuscous ;
towards base some fuscous suffusion sprinkled with dark fuscous,
especially on costa and dorsum ; stigmata dark fuscous, first discal
enlarged into a round cloudy rather dark fuscous spot, obliquely
above and before which is another dark fuscous dot, plical obliquely
beyond first discal; a suffused fuscous spot on costa beyond
730 MR. E. MEYRICK ON [June 16,
middle ; a curved posterior series of cloudy dots of fuscous and
dark fuscous scales: cilia whitish-yellowish. Hind wings
ochreous-whitish, faintly fuscous-tinged ; cilia whitish-yellowish.
S. Nigeria, Wari, in May and August; three specimens.
EPIMACTIS METAZONA, Sp. 0.
Q. 25-26 mm. Head and thorax white. Palpi white, lower
half of second joint fuscous. Antenne grey, base white. Ab-
domen light yellow-ochreous. Fore wings suboblong, costa
moderately arched, apex obtuse, termen nearly straight, vertical ;
white; second discal stigma grey; a narrow silvery-grey fascia
close before termen, dilated towards costa but not quite reaching
it: cilia white. Hind wings and cilia white.
SreRRA LEONE; two specimens.
Procometis Meyy.
To this genus should be referred acutipennis Wals., described
under Apiletria, as well as the three following species. All four of
these agree with the single known Indian species, P. trochala, and
differ from all the Australian species (ten in number) in the
character of the labial palpi, which in these have the second joint
loosely haired above and the terminal joint relatively very short
(3 or less), whilst in the Australian forms the second joint is
clothed with appressed scales and the terminal joint is as long as
second. Notwithstanding this distinction, the two groups are so
obviously closely related in all other characters, structural and
superficial, and connecting forms are so likely to be found eventu-
ally in intermediate regions (the differences being only com-
parative), that I think it best to treat them as congeneric ; but the
group to which the African and Jndian species belong seems
worthy of a subgeneric name, and I therefore give it the name of
Hypostola.
PROCOMETIS OXYPORA, Sp. 0.
$. 33mm. Head, thorax, and abdomen pale ochreous. Palpi
with second joint loosely haired above, terminal joint 4 of second ;
pale brownish-ochreous mixed with whitish. Antenne whitish.
Fore wings elongate, rather narrow, costa moderately arched, apex
very acute, produced, termen sinuate, extremely oblique; pale
yellowish-ochreous, tinged with brown-reddish posteriorly ; a
broad very undefined streak of pale fuscous suffusion beneath
costa from before middle to apex, and some undefined pale fuscous
suffusion towards dorsum: cilia pale ochreous-yellowish. Hind
wings narrower than fore wings, with very long tornal cilia, and
large light ochreous-fuscous costal hair-pencil reaching to apex ;
grey, paler and whitish-tinged towards base ; cilia light ochreous-
fuscous, becoming light yellowish- ochreous towards tornus.
Navan, TWeemen, | in January ; one specimen.
1908. ] AFRICAN MICRO-LEPIDOPTERA. el
PROCOMETIS ACHARMA, Sp. n.
3. 32 mm., 9 46 mm. Head and thorax pale brownish-
ochreous. Palpi with second joint loosely haired above, terminal
joint 4 of second; brownish-ochreous sprinkled with whitish.
Antenne ochreous-whitish indistinctly ringed with fuscous. Ab-
domen light greyish-ochreous. Fore wings elongate, rather
narrow, costa moderately arched, apex in ¢ round-pointed, in
obtuse, termen in ¢ straight, very oblique, in ? slightly rounded,
rather strongly oblique; light fuscous suffusedly irrorated with
whitish, suffused with pale brownish-ochreous towards costa
anteriorly, with a few black scales on veins on costal half; discal
stigmata faintly indicated, the white suffusion more pronounced
on a longitudinal streak traversing these and towards dorsum:
cilia whitish-yellowish, in 2 sprinkled with whitish. Hind wings
in ¢ narrower than fore wings, with very long tornal cilia, and
grey costal hair-pencil reaching to+; grey,in 2 somewhat lighter ;
cilia light grey mixed with whitish, in ¢ becoming pale ochreous-
yellowish towards tornus, in Q with grey subbasal line.
Narat, Weenen, in December ; two specimens.
PROCOMETIS TERRENA, Sp. 0,
2.3/7 mm. Head, palpi, antenne, and thorax rather dark
fuscous ; palpi with second joint loosely haired, terminal joint
half second. Abdomen light fuscous. Fore wings elongate,
rather narrow, costa moderately arched, apex obtuse, termen
rounded, oblique; rather dark ashy-fuscous, lighter posteriorly ;
a broad ochreous-brown median stripe from base, becoming
suffused and obsolete beyond middle, edged above by groups of
scattered black scales beyond + and about middle, and followed by
two minute biack dots transversely placed at 2: cilia fuscous.
Hind wings light fuscous; cilia whitish-fuscous, with darker
subbasal line.
NyassaLAnD, Mpeta, on Loangwa River, in December; one
specimen.
(HcoerHORID..
CoESYRA BALANTIAS, Sp. 0.
2. 20mm. Head, palpi, and thorax bright yellow. Antenne
grey, becoming yellowish towards base. Abdomen ochreous-
yellowish. Fore wings elongate, costa gently arched, apex obtuse,
termen nearly straight, oblique; bright yellow; a triangular
whitish-fuscous spot irrorated with dark fuscous on dorsum beyond
middle, its apex sending an irregular outwardly oblique projection
edged with white above to disc at 2; two or three small fuscous
dots in disc posteriorly, representing part of a curved series ;
several minute indistinct fuscous dots on termen and round apex :
cilia yellow. Hind wings and cilia light ochreous-yellowish.
Narat, Weenen, in February (Spiller) ; one specimen.
732 MR. E. MEYRICK ON [June 16,
EPIPHRACTIS, 1. g.
Head with appressed scales, side-tufts spreading; tongue de-
veloped. Antenne 4, basal joint moderately elongate, without
pecten. Labial palpi very long, recurved, with appressed scales,
terminal joint shorter than second, acute. Maxillary palpi rudi-
mentary. Posterior tibiz clothed with hairs above. Fore wings
with 2 from angle, 7 and 8 stalked, 7 to termen, 11 from before
middle. Hind wings 1, elongate-ovate, cilia 3; 3 and 4 connate,
5-7 parallel, 7 connected with 8 by a bar beyond cell.
Probably allied to Cryptolechia.
EPIPHRACTIS PHG@NICIS, sp. .
@. 23 mm. Head and thorax light rosy-ochreous. Palpi
whitish-ochreous. Antennze pale ochreous tinged with crimson.
Abdomen light grey, apex light ochreous. Fore wings elongate,
somewhat dilated posteriorly, costa gently arched, apex obtuse,
termen faintly sinuate, rather oblique; ochreous-crimson, deeper
purplish-crimson towards dorsum, lighter and more ochreous
towards costa; costal edge whitish except towards apex, extreme
costal edge blackish towards base ; second discal stigma represented
by a smal] suffused dark grey spot: cilia rosy. Hind wings grey ;
cilia whitish-grey.
AnGoLa, Bihe; one specimen.
EruMiA BALLISTIS, sp. 1.
29-31 mm. Head and thorax shining grey irrorated with
grey-whitish ; head with a triangular blackish spot on back of
crown, thorax with three black dots in a dorsal triangle, two
others posterior, and two on shoulders. Palpi blackish-grey,
towards base whitish. Antenne blackish, whitish in front to-
wards base, and on lower half of basal jot. Abdomen bright
deep yellow, beneath blackish towards base of segments. Legs
blackish ringed with white, posterior tibie yellow. Fore wings
elongate, rather narrow, costa moderately arched, apex obtuse,
termen rounded, rather oblique; bronzy-grey; base of costa
blackish ; two small black spots beneath costa near base, surrounded
with some whitish suffusion ; black dots on fold at base, near
base, and at +; stigmata black, discal large, round, first little
befere middle, plical small, obliquely beyond first discal: cilia
bronzy-grey. Hind wings bright deep yellow ; a dark grey apical
patch, covering about t of wing; cilia yellow, round apex dark
grey.
~ Grrman East Arrica, Dar-es-Salaam ; two specimens.
BORKHAUSENIA GALACTAA, Sp. 0.
36.16 mm. Head, palpi, antenne, thorax, and abdomen
ochreous-whitish ; palpi with median bands of fuscous irroration
on second and terminal joints; antenne serrate. Fore wings
elongate, rather narrow, costa moderately arched, apex obtuse,
1908. ] AFRICAN MICRO-LEPIDOPTERA. 733
termen extremely obliquely rounded ; ochreous-whitish ; a small
blackish dot beneath costa near base; stigmata black, plical
obliquely beyond first discal ; an almost marginal series of unde-
fined dark fuscous dots round posterior third of costa and termen :
cilia ochreous-whitish. Hind wings light grey; cilia ochreous-
whitish.
TRANSVAAL, Pretoria district, in December and January (Janse) ;
one specimen.
ELACHISTIDS.
STATHMOPODA XANTHOPLITIS, Sp. n.
$6.15 mm. Head and thorax orange, face shining whitish-
ochreous. Palpi whitish-ochreous. Antenne grey, basal joint
orange. Abdomen whitish-ochreous, base of segments golden-
ferruginous. Fore wings narrow-lanceolate, very acutely pointed ;
pale greyish-ochreous ; basal third orange: cilia pale yellowish-
ochreous. Hind wings light grey; cilia pale yellowish-ochreous.
TRANSVAAL, Pretoria, from December to February (Janse) ;
one specimen.
EXLACHISTA CROCOGASTRA, Sp. Nn.
3. 6-7 mm. Head, palpi, antenne, and thorax ochrecus-
whitish. Abdomen whitish, tinged and sometimes suffused with
yellow. Fore wings lanceolate, ochreous-whitish: cilia whitish.
Hind wings and cilia ochreous-whitish.
TRANSVAAL, Pretoria district, in August, December to February,
and April (Janse); six specimens.
CosMOPTERYX BACTROPHORA, sp. n.
2.11 mm. Crown of head and thorax dark fuscous with
three fine longitudinal white lines, face light fuscous. Palpi
white, terminal joint with a black iine on each side. Antenne
white lined with black. Abdomen rather dark grey, beneath pale
ochreous. Fore wings narrow-lanceolate, apex very long-produced
and acute; dark fuscous; an oblique series of three fine white
longitudinal lines about 1, first reaching base of costa, others
detached; a broad ochreous-yellowish transverse fascia beyond
middle, anterior edge somewhat oblique and marked with two
small silvery-metallic spots, first followed by two or three black
scales, and extended anteriorly on costa as a short fine white
streak, posterior edge limited by two small silvery-metallic spots
edged with dark fuscous anteriorly, between which the yellow
colour forms a triangular median projection, whence proceeds a
sinuate fine white line to apex: cilia dark fuscous, with a white
bar on apical line. Hind wings and cilia dark grey.
TRANSVAAL, Pretoria, in February (/anse); one specimen.
CosMOPTERYX TABELLARIA, Sp. N.
¢. 10mm. Crown of head and thorax dark brown with three
fine longitudinal white lines; face whitish-fuscous. Palpi whitish,
734 MR. E, MEYRICK ON [June 16,
terminal joint with a blackish line on each side. Antenne white
lined with black. Abdomen pale yellowish-ochreous. Fore wings
narrow-lanceolate, apex very long-produced and acute; dark
brown; an oblique series of three fine white longitudinal lines
about 4, first reaching base of costa, second reaching base and also
connected with projection of postmedian band, third connected
with same projection beneath ; a broad ochreous-yellow transverse
fascia beyond middle, anterior edge produced as a short white
streak on costa, below middle with a triangular projection, above
this marked with a round golden-metallic spot followed by a black
dot, obliquely beyond this within fascia is a round golden-metallic
spot near dorsum, posterior edge of fascia marked with two
opposite pale golden-metallic spots, between which is an elongate
projection of the yellow colouring, whence proceeds a sinuate
white line to apex: cilia dark fuscous. Hind wings and cilia
orey.
TRANSVAAL, Pretoria, in March (Jase); one specimen.
STAGMATOPHORA SEMNOSTOLA Meyr.
TRANSVAAL, Pretoria district, in August, September, and from
December to April (/anse).
Described from Australia, where the larva feeds amongst spun
leaflets of Acacia decurrens ; I have not yet obtained the species
from intermediate regions.
LIMNa@cIA ICHNOGRAPHA, Sp. ni.
g. 17-18 mm. Head, palpi, and thorax blackish, sprinkled
with white. Antenne blackish. Abdomen dark grey, sides of
back ochreous-whitish towards base, anal tuft whitish mixed with
yellow. Fore wings elongate-lanceolate, acute; dark purplish-
grey, suffusedly streaked longitudinally with black ; a fine white
line above fold from base to 2, beneath which is another on
posterior portion ; a short white oblique mark from beneath costa
at 1; a white mark on middle of costa, beneath which are two or
three small undefined white marks longitudinally arranged in dise
and one on fold; a rather inwardly oblique white streak from
costa beyond #, reaching half across wing; an irregular undefined
line of more or less scattered white scales along dorsum from near
base and lower half of termen: cilia dark grey, with a small basal
spot of white scales beneath apex, and a faint median whitish
shade on upper half of termen. Hind wings dark grey; cilia
grey, basal line pale greyish-ochreous.
TRANSVAAL, Pretoria and Pietersburg districts, from September
to January (Janse); two specimens.
GRACILARIADA.
KiPIcEPHALA PYRRHOGASTRA, Sp. nN.
6 2. 9-10 mm. Head white, somewhat mixed with grey
on forehead. Palpi pale grey, becoming white towards apex.
1908. | AFRICAN MICRO-LEPIDOPTERA. 735
Antenne grey. Thorax grey, sometimes whitish-mixed. Abdomen
light ochreous-reddish, dorsally suffused with grey, sides with
series of oblique dark grey stripes. Fore wings narrowly
elongate-lanceolate, apex blunt-pointed ; ochreous-grey ; three fine
white very oblique streaks from costa at 1, beyond middle, and
at ?, reaching nearly half across wing, dark-edged anteriorly,
sometimes dilated on costa, between these are more or less distinct
whitish marks beneath costa; an irregular white streak along
dorsum throughout, above which is a short white oblique mark
beneath middle of wing, and two approximated fine oblique white
lines above tornus ; a fine silvery-metallic curved transverse line
at 2; a small round black anteapical spot, above and beneath
which are white wedge-shaped spots on margins: cilia white,
towards tornus light ochreous-grey, elsewhere with a blackish
subbasal line, on costa also with apical third blackish. Hind
wings dark grey ; cilia grey.
Transvaal, Pretoria, from December to February (Janse); four
specimens.
ACROCERCOPS DASMOPHORA, Sp. 0.
2. 89mm. Head and palpi shining white. Antenne grey,
beneath whitish. Thorax pale ochreous. Abdomen whitish-
ochreous. Fore wings narrowly elongate-lanceolate; ochreous-
orange ; markings shining white, edged with rather dark fuscous ;
three semioval dorsal spots, first two large, first reaching base,
second rather obliquely placed, third smaller; an elongate-
triangular blotch extending along costa from near base to 2
broadest posteriorly, posterior edge oblique, almost united with
second dorsal spot; three wedge-shaped costal spots posteriorly,
first oblique, elongate, almost united with third dorsal spot,
second and third not dark-margined posteriorly, second extended
as a narrow streak to termen, third small, anteapical, its apex
almost touching this streak ; a small elongate black apical spot :
cilia white, on costa with three dark fuscous bars, round apex
and termen with a dark fuscous subbasal line, and a dark fuscous
bar beneath apex, beneath termen tinged with greyish-ochreous.
Hind wings grey ; cilia pale greyish-ochreous.
TRANSVAAL, Pretoria, in November, January, and February
(Janse) ; two specimens.
MACAROSTOLA ONYCHOTA, sp. 0.
$¢ 2. 8-9 mm. Head white, sides of crown fuscous-tinged.
Palpi loosely scaled, white, lower 2 of second joint and a median
ring of terminal joint fuscous. Antenne whitish ringed with
fuscous. Thorax ochreous-bronze, with two white stripes. Ab-
domen grey. Fore wings elongate-lanceolate, acute; ochreous-
bronze ; markings white, edged with black irroration ; four streaks
from costa, reaching nearly half across wing, first three outwardly
oblique, first at 2, extended as a fine line along costa to near base,
fourth inwardly oblique; four streaks from dorsum reaching half
Proc. Zoou. Soc.—1908, No. XLVII. A7
736 MR. E. MEYRICK ON [June 16,
across wing, first three outwardly oblique, first from rather near
base, fourth inwardly oblique: cilia white, with two posterior
lines of black irroration, on costa grey with a white bar, at apex
with a black hook, beneath this on basal half ochreous-grey
speckled with black and barred with white, beneath tornus
greyish. Hind wings and cilia grey.
TRANSVAAL, Pretoria, in January, February, and May (Janse) ;
four specimens.
PLUTELLIDA,
TRIOTHYRSA, nh. g.
Head with appressed hairs; ocelli absent; tongue short. An-
tenne 2,in ¢ simple, basal joint elongate, with pecten. Labial
palpi very long, porrected, second joint very long, widely diverging,
clothed with dense scales, projecting roughly above, terminal joint.
half second, bent inwardly at right angles to it, loosely scaled,
acute. Maxillary palpi obsolete. (Posterior legs broken.) Fore
wings with 2 from angle, 3 absent, 4 and 5 connate, 6 and 7 out
of 8, 7 to termen, 9 out of 8 before 6, 10 from near end of cell,
11 from beyond middle. Hind wings 2, narrow-lanceolate, cilia
3; 2 separate, 3 and 4 connate, 5 and 6 stalked, 7 separate.
Intermediate in character between Plutella and Coleophora.
IRIOTHYRSA MELANOGMA, Sp. n.
Sg. 21 mm. Head and thorax ochreous-whitish. Palpi
whitish, second joint sprinkled outwardly with fuscous and
blackish. Antenne ochreous-whitish. Abdomen ochreous-
whitish mixed with grey. Fore wings elongate-lanceolate,
termen faintly sinuate ; pale brownish-ochreous, suffusedly mixed
with white, especially towards base ; costal edge blackish towards.
base; some scattered black scales, and posterior # suffusedly
streaked longitudinally with fuscous suffusion irrorated with
black: cilia whitish-ochreous. Hind wings rather dark grey,
anteriorly pale greyish-ochreous towards costa; cilia whitish-
ochreous.
TRANSVAAL, Pietersburg, in March (Janse) ; one specimen.
YPONOMEUTA GLAPHYROPIS, sp. N.
$6 2. 25-28 mm. Head white or whitish, with two black
spots on crown and two or four on face. Palpi blackish, apex of
all joints whitish. Antenne light grey. Thorax grey-whitish,
with two black marks on each shoulder, a dot on each patagium,.
and two on back. Abdomen dark grey or blackish-grey. Fore
wings elongate, narrow, costa gently arched, apex obtuse, termen
-nearly straight, oblique, cell very long, reaching to + of wing;
light grey or sometimes darker grey, with whitish reflections ;.
usually some whitish suffusion on fold, especially on basal half;
five or six black dots immediately beneath costa on basal 2,
a series of about six near costa from 4 to apex, a series of about.
1908. ] AFRICAN MICRO-LEPIDOPTERA, 137
three from 4 of disc to apex, a submedian series of eight or nine
from near base to termen, and a subdorsal series of six to eight
from near base to termen: cilia light grey. Hind wings with
basal transparent fovea narrow, deep; blackish-grey; cilia dark
grey.
Narar, Weenen, from October to December ; six specimens.
TINEIDA.
BUCCULATRIX PORTHMIS, Sp. n.
36 9.6 mm. Head and thorax yellow-ochreous. Antenne
pale ochreous, suffusedly ringed with dark fuscous. Abdomen
rather dark grey. Fore wings rather broad-lanceolate, apex
rather produced, acute ; yellow-ochreous; an undefined patch of
blackish irroration on costa beyond middle, appearing to be
margined laterally by faint oblique marks of whitish suffusion ; a
patch of blackish irroration on dorsum slightly before costal :
cilia grey, with basal and median lines of black irroration. Hind
wings rather dark grey ; cilia grey.
TRANSVAAL, Pretoria, in February (Janse); two specimens.
OPoGONA PHHOCHALCA, Sp. Nn.
[email protected]. Head and palpi shining whitish-ochreous; back
of crown and thorax purplish-bronzy-fuscous. Antenne whitish-
ochreous. Abdomen light grey. Fore wings lanceolate, acute ;
purplish-bronzy-fuscous: cilia bronzy-greyish-ochreous. Hind
wings light grey, with very strong brassy reflections; cilia light
bronzy-greyish-ochreous.
Reunion, St. Denis, in April; one specimen.
OPOGONA CHLOROPHANES, sp. n.
$6 Q. 11-12mm. Head, palpi, antenne, thorax, and abdomen
shining whitish-ochreous. Fore wings lanceolate, very acute ;
shining whitish-ochreous, with a brassy or bronzy tinge: cilia
whitish-ochreous. Hind wings grey, with strong brassy reflections ;,
cilia whitish-ochreous.
Transvaal, N.E. Pretoria district, in August, December, and
January (Janse) ; four specimens.
Mownopis MEGALODELTA, Sp. 0.
g.13mm. Head ochreous-whitish. Palpi ochreous-whitish,
externally suffused with dark fuscous except at apex. Antenne
whitish-ochreous, above suffused with dark fuscous towards base,
basal joint ochreous-whitish. Thorax dark brown mixed with
blackish, shoulders with an ochreous-whitish spot. Abdomen
brownish-ochreous. Fore wings elongate, costa gently arched,
apex obtuse, termen obliquely rounded; 7 and 8 stalked, discal
impression very large, thinly scaled but not transparent ; ferru-
ginous-brown irrorated with blackish, and strewn with small dark
slaty-fuscous spots of more or less raised scales ; a large triangular
47*
738 MR. E. MEYRICK ON [June 16,
ochreous-whitish blotch extending on costa from 4 to #, and
reaching # across wing; disc beyond this blotch more coppery-
ferruginous: cilia ferruginous-fuscous irrorated with blackish.
Hind wings grey, with golden reflections ; cilia ochreous-whitish.
S. Niemria, Ogrugu ; one specimen.
MALACYNTIS, n. g.
Head roughly tufted; ocelli absent; tongue obsolete. An-
tenne over 1, in ¢ stout, simple, basal joint thick, concave
beneath. Labial palpi moderate, porrected, second joint with
dense scales projecting beneath at apex and two or three apical
bristles, terminal joint moderate, densely scaled. Maxillary palpi
obsolete. Posterior tibiz clothed with long hairs above. Fore
wings with 2 from towards angle, 7 and 9 stalked, 7 to costa,
8 absent, 11 from before middle. Hind wings under 1, elongate-
ovate, cilia £; 2-7 tolerably parallel.
A development of Z%ineola.
MALACYNTIS STIBARODES, Sp. 0.
$6.15 mm. Head light yellow-ochreous. Palpi rather dark
fuscous, beneath ochreous-suffused, apex whitish-ochreous. An-
tenne whitish-ochreous. Thorax and abdomen pale brownish-
ochreous. Fore wings elongate, costa moderately arched, apex
obtuse, termen very obliquely rounded ; pale brownish-ochreous ;
base of costa infuscated: cilia pale yellowish-ochreous, tips paler.
Hind wings bronzy-grey ; cilia ight brownish-ochreous.
SrERRA LEONE; one specimen.
TINEOLA LEUCASTIS, Sp. n.
3. 24 mm. Head light ochreous-orange. Palpi moderate,
whitish-ochreous. Antenne grey. Thorax white. Abdomen
ochreous-yellowish. Fore wings elongate, rather narrow, costa
gently arched, apex obtuse, termen very obliquely rounded ;
white ; costal edge finely blackish towards base, elsewhere pale
ochreous-yellowish : cilia pale ochreous-yellowish. Hind wings
light yellowish-grey ; cilia whitish-ochreous.
TRANSVAAL, N.K. Pretoria district and Pietersburg, in October,
December, and March (Janse); one specimen.
TINEOLA XANTHASTIS, sp. n.
3$.19mm. Head and thorax bright ochreous-yellow. Palpi
moderate, grey, upex whitish-yellowish. Antenne grey. Ab-
domen whitish-ochreous. Fore wings elongate, narrow, costa
gently arched, apex obtuse, termen very obliquely rounded ;
bright ochreous-yellow ; costal edge finely blackish towards base:
cilia ochreous-yellow. Hind wings grey ; cilia whitish-ochreous.
TRANsvAAL, Pretoria, in December, January, and March
(Janse); one specimen.
1908. ] AFRICAN MICRO-LEPIDOPTERA. 739
TINEOLA OCHROPSAMMA, Sp. Nn.
$.18 mm. Head fulvous-ochreous. Palpi moderate, pale
ochreous, second and terminal joints suffused with dark fuscous
on basal half. Antennz pale ochreous. Thorax and abdomen
yellow-ochreous. Fore wings elongate, costa moderately arched,
apex tolerably pointed, termen very oblique, almost straight ;
yellow-ochreous : cilia yellow-ochreous, paler towards tips. Hind
wings purplish-grey suffused anteriorly with light Dbrassy-
ochreous; cilia as in fore wings.
Narat, Durban, in September (Leigh) ; one specimen.
TINEOLA MARCESCENS, Sp. 0.
6. 18mm. Head brownish-ochreous. Palpi moderate, dark
fuscous. Antenne pale brownish-ochreous, {infuscated at base.
Thorax and abdomen light brownish-ochreous. Fore wings
elongate, rather narrow, costa moderately arched, apex obtuse,
termen very obliquely rounded; pale brownish-ochreous, some-
what more brownish-tinged anteriorly ; costal edge dark fuscous
towards base: cilia pale brownish-ochreous. Hind wings and
cilia pale ochreous.
British East Arrica, Mombasa, in October ; one specimen.
TINEOLA HOLOPSAMMA, Sp. Nn.
36.19 mm. Head ochreous-yellowish mixed with fuscous.
Palpi moderate, ochreous-yellowish, terminal joint and base of
second suffused with fuscous. Antenne pale ochreous, base in-
fuscated. Thorax and abdomen yellow-ochreous. Fore wings
elongate, rather narrow, costa gently arched, apex obtuse, termen
very obliquely rounded ; yellow-ochreous ; costal edge infuscated
towards base: cilia yellow-ochreous. Hind wings light ochreous-
grey ; cilia whitish-ochreous, more ochreous towards base.
TRANSVAAL, Pietersburg, in September (/Janse); one specimen.
TINEOLA MELANOSTOMA, Sp. n.
S$. 19 mm. Head fulvous. Labial and maxillary palpi
blackish, labial short, maxillary stout. Antenne pale ochreous,
becoming whitish towards apex. Thorax deep yellow-ochreous
tinged with lilac. Abdomen golden-ochreous. Fore wings
elongate, rather narrow, costa moderately arched, apex obtuse,
termen very obliquely rounded ; deep yellow-ochreous tinged with
brownish and suffused with pale lilac; costal.edge dark fuscous
towards base: cilia concolorous. Hind wings grey with lilac-
bronze reflections ; cilia pale golden-ochreous.
British East Arrica, Mombasa, in October ; one specimen.
TINEOLA MERETRIX, sp. n.
$. 15mm. Head fuscous. Palpi moderate, whitish-ochreous,
terminal joint fuscous. Antenne stout, somewhat compressed
740 MR. E. MEYRICK ON [June 16,
laterally, ochreous-whitish. Thorax light ochreous. (Abdomen
damaged.) Fore wings elongate, rather narrow, costa moderately
arched, apex obtuse, termen very obliquely rounded ; purple-
brownish-ochreous : cilia ochreous sprinkled with purplish towards
base. Hind wings grey, with purplish reflections; cilia pale
yellowish-ochreous.
Carr Cotony, Grahamstown ; one specimen.
TINEOLA PHOCINA, sp. 1.
g.12mm. Head pale ochreous-yellowish. Palpi moderate,
dark fuscous, apex ochreous-whitish. Antenne pale greyish-
ochreous, towards apex more whitish, towards base infuscated.
Thorax bronzy-fuscous, dorsally purplish-tinged. Abdomen
greyish-ochreous. Fore wings elongate, rather narrow, costa
moderately arched, apex obtuse, termen very obliquely rounded ;
shining fuscous-bronze: cilia pale bronzy-ochreous. Hind wings
grey ; cilia whitish-ochreous, tips whitish.
Transvaal, N.E. Pretoria district, in December (Janse); one
specimen.
TINEOLA (?) CHLORISTIS, Sp. n.
3. 11mm. Head ochreous-whitish. Palpi whitish-ochreous,
infuscated above. Maxillary palpiapparently rudimentary. An-
tenne, thorax, and abdomen whitish-ochreous. Fore wings
elongate, costa moderately arched, apex obtuse, termen very
obliquely rounded; 7 and 8 stalked; shining whitish-ochreous :
cilia whitish-ochreous, more whitish towards tips. Hind wings
and cilia pale whitish-ochreous, with a bronzy tinge.
TransvaaL, N.E. Pretoria district, in February (Janse); one
specimen
TINEA CNOPIS, sp. 1.
S.17 mm. Head ochreous-yellowish. Palpi dark fuscous,
apex paler. Antenne 1, ochreous-whitish. Thorax dark purplish-
fuscous. Abdomen light brownish-ochreous, segmental margins
whitish-ochreous. Fore wings elongate, narrow, costa gently
arched, apex obtuse, termen very obliquely rounded ; rather dark
purplish-bronzy-fuscous : cilia pale bronzy sprinkled with fuscous.
Hind wings pale bronzy, with a pale grey terminal line; cilia
pale grey.
TRANSVAAL, Pretoria, in December (Janse); one specimen.
TINEA PARAXENA, Sp. 0.
3.14mm. Head bright yellow. Palpi pale ochreous, suffused
above with dark fuscous. Antenne 1, ochreous-whitish. Thorax
and abdomen pale shining ochreous. Fore wings elongate, narrow,
costa gently arched, apex round-pointed, termen extremely
obliquely rounded; pale shining ochreous; costal edge blackish
towards base: cilia pale ochreous, more whitish towards tips.
Hind wings grey with bronzy reflections ; cilia whitish-ochreous.
lod
1908. | AFRICAN MICRO-LEPIDOPTERA. 741
TRANSVAAL, Pietersburg, in September (Janse); one specimen.
Distinguished from 7’. tanystis by the much narrower fore
wings,
TINEA OTHELLO Meyr.
TRANSVAAL, Pretoria and Pietersburg districts, from August to
December (Janse): four specimens, apparently not differing from
Indian examples.
TINEA HOMESTIA, Sp. 0.
S.12mm. Head yellow-ochreous. Palpi whitish-ochreous,
upper edge blackish except towards apex of joints. Antenne 1,
grey-whitish, greyer above, especially towards base. Thorax and
abdomen pale ochreous. Fore wings elongate, rather narrow,
costa gently arched, apex round-pointed, termen very obliquely
rounded; pale ochreous, with a faint brownish tinge: cilia
concolorous. Hind wings very pale greyish ; cilia pale ochreous.
TRANSVAAL, Pretoria, in January (Janse); one specimen.
TINEA TANYSTIS, Sp. 1.
$. 16-17 mm. Head bright yellow. Palpi pale ochreous,
second joint mixed with blackish. Antenne 1, stout, compressed,
ochreous-whitish. Thorax light yellow-ochreous, anterior margin
infuscated. Abdomen yellow-ochreous. Fore wings elongate,
rather narrow, costa gently arched, apex round-pointed, termen
very obliquely rounded; pale yellow-ochreous; costal edge
blackish towards base: cilia pale yellow-ochreous. Hind wings
grey, with a slight bronzy tinge; cilia whitish-ochreous.
TRANSVAAL, Pretoria and Pietersburg districts, from September
to February (Janse); five specimens.
PSEUDURGIS, n. g.
Head rough-sealed ; ocelli and tongue absent. Antenne 3, in
3S unipectinated, apex simple. Labial palpi rather long, porrected,
densely clothed throughout with loosely projecting scales, terminal
joint moderate. Maxillary palpi absent. Posterior tibiz loosely
scaled. Fore wings with 2 from #, 3 from angle, 4 and 5 di-
vergent, 7 to termen, 8—1(0) approximated at base, 11 from middle.
Hind wings 1, elongate-ovate, cilia 2; 2-7 tolerably parallel.
A development of MWelasina, principally distinguished by the
unipectinated antenne, but of peculiar facies.
PSEUDURGIS TECTONICA, Sp. 0.
6. 21-22 mm. Head light brown mixed with whitish and
sprinkled with dark fuscous. Palpi brown sprinkled with dark
fuscous. Antenne whitish, stalk and pectinations lined with
blackish, pectinations 6. Thorax brown mixed with whitish.
Abdomen light brown mixed with whitish. Fore wings elongate,
rather narrow, posteriorly somewhat dilated, costa nearly straight,
742 MR, E. MEYRICK ON [June 16,
slightly sinuate, apex obtuse, termen nearly straight, oblique ;
white, towards costa and dorsum more or less irregularly irrorated
with fuscous and strigulated with dark fuscous and blackish; a
dark fuscous spot marked with blackish on base of costa, one on
middle of dorsum, and one in dise beyond middle; an elongate-
transverse fuscous spot suffusedly mixed with blackish near
termen, not reaching margins, terminal area beyond and beneath
this mixed with light ferruginous and with some dark fuscous
strigule; apical and terminal margins marked with small dark
fuscous and black spots: cilia white, inner half irrorated with
fuscous and limited by a line of dark fuscous irroration, outer half
barred with dark fuscous suffusion. Hind wings grey; cilia
whitish-grey.
TRANSVAAL, Pretoria, in November and December (Janse) ;
two specimens.
STRUTHISCA OMICHLODES, sp. 0.
3 @. 15-20 mm. Head and thorax fuscous, face sometimes.
paler and whitish-tinged. Palpi very short, fuscous. Antenne
grey, pectinations in ¢ 23. Abdomen light greyish, rather hairy
posteriorly, apex in Q whitish-ochreous. Fore wings elongate,
rather narrow, costa moderately arched, apex obtuse, termen very
obliquely rounded; 7 absent: light grey, very indistinctly strigu-
lated with darker: cilia light grey. Hind wings thinly scaled,
light grey; cilia whitish-grey.
TRANSVAAL, Pretoria, in August and September (Janse); three
Specimens.
In my original characterisation of this genus the absent vein of
fore wings is stated to be 9; I am now of opinion that it is really
7 which is missing normally; in S. hormotris, described below,
all veins of the fore wings are present, but 7 and 8 are stalked,
and this species is therefore probably an early form, and indicates
the true homology,
STRUTHISCA AREATA, Sp. Nn.
3. 17-19 mm. Head and thorax ochreous-fuscous. Palpi
very short, fuscous. Antenne fuscous, pectinations 4. Abdomen
rather hairy, pale brownish-ochreous. Fore wings elongate,
rather narrow, costa gently arched, apex obtuse, termen rounded,
rather strongly oblique; 7 absent; light brownish-ochreous: cilia
concolorous. Hind wings and cilia very pale ochreous-grey.
TRANSVAAL, Pietersburg, in September (Janse) ; three specimens.
STRUTHISCA HORMOTRIS, sp. 0.
3.16 mm. Head whitish-ochreous. (Palpi broken.) An-
tenn fuscous, pectinations 4. Thorax and abdomen fuscous.
Fore wings elongate, costa moderately arched, apex obtuse, termen
very obliquely rounded ; 7 and 8 stalked ; grey-whitish, irregularly
strigulated with fuscous; a patch of fuscous suffusion on base of
costa ; a moderate slightly ineurved fuscous fascia from middle of
1908. | AFRICAN MICRO-LEPIDOPTERA. 743
dorsum, reaching 2 across wing, darkest anteriorly ; an undefined
fuscous spot beneath costa at 2: cilia pale whitish-ochreous. Hind
wings grey ; cilia whitish-ochreous-grey.
NyassaLANnD, Zomba, 3000 feet, in December ; one specimen.
THRANITICA, n. g.
Head with loosely appressed scales; ocelli present; tongue
absent. Antenne 3, in ¢ moderately biciliated. Labial palpi
moderately long, subascending, with appressed scales, terminal
joint moderate, obtuse. Maxillary palpi absent. Posterior tibie
smooth-scaled. Fore wings with 2 from angle, 5 absent, 7 to
apex, 8 absent, 11 from middle. Hind wings 1, ovate, cilia 4;
4 absent, 2-7 nearly parallel.
THRANITICA HEMICOPA, Sp. n.
3.20 mm. Head whitish-ochreous tinged with yellowish.
Palpi fuscous. Antennz whitish-ochreous. Thorax pale whitish-
ochreous mixed anteriorly with brownish and dark fuscous. Ab-
domen whitish-ochreous. Fore wings elongate, moderately broad,
costa moderately arched, apex rounded, termen obliquely rounded ;
pale whitish-ochreous, irregularly strewn with: fuscous and dark
fuscous strigule ; base of costa dark fuscous; a narrow erect dark
fuscous fascia from dorsum at 4, reaching more than half across
wing; a dark fuscous dot in disc at 2; three very small dark
fuscous spots on posterior half of costa: cilia whitish-ochreous.
Hind wings and cilia pale whitish-ochreous.
NATAL; one specimen.
MELASINA HALIEUTIS, sp. n.
3. 22mm. Head and thorax white, partially fuscous-tinged.
Palpi short, fuscous mixed with whitish, loosely haired. Antenne
whitish, pectinations 4. Abdomen hairy, ochreous-whitish. Fore
wings elongate, somewhat dilated posteriorly, costa moderately
arched, apex obtuse, termen obliquely rounded ; 7 absent; whitish,
irregularly strigulated throughout with fuscous; costa with several
small darker fuscous spots ; a suffused fuscous spot above middle
at 2: cilia whitish, outer half more or less brownish. Hind wings
grey; cilia grey-whitish.
Nara; one specimen.
MELASINA STELITIS, sp. 0.
3S. 23mm. Head and thorax grey mixed with white, forehead
and upper edge of face white, rest of face dark grey. Palpi short,
slender, loosely scaled, grey. Antenne light grey, pectinations 5.
Abdomen grey. Fore wings elongate, somewhat dilated posteriorly,
costa moderately arched, apex obtuse, termen obliquely rounded ;
7 absent; grey suffusedly mixed with white, and strigulated
throughout with blackish; three small indistinct dark fuscous
spots on posterior half of costa; absence of white suffusion and
744 MR. E. MEYRICK ON [June 16,
increase of dark strigulation forms an undefined narrow fascia
from middle of dorsum reaching 2 across wing : cilia pale fuscous
mixed with whitish. Hind wings grey; cilia grey, towards tips
whitish-tinged,
Transvaal, N.E. Pretoria district, in January (Janse); one
specimen.
MELASINA £DIFICA, Sp. 0.
3. 20-24 mm. Head ochreous-yellowish, sometimes tinged
with fulvous. Palpi moderate, densely scaled, ochreous-yellowish,
basal half suffused with dark fuscous. Antenne ochreous-whitish,
pectinations 5. Thorax ochreous-whitish, anterior margin suffused
with blackish-fuscous, tips of patagia and thoracic crest sometimes
blackish. Abdomen pale greyish-ochreous, anal tuft yellowish.
Fore wings elongate, moderate, costa gently arched, apex obtuse,
termen obliquely rounded; all veins separate; whitish, with
scattered small blackish-fuscous strigule; costal edge whitish-
ochreous; irregular markings formed of confluent cloudy blackish-
fuscous strigule, viz., a small, sometimes partially obsolete, basal
patch, a moderate fascia from 4 of costa to middle of dorsum,
another from 2 of costa to tornus, and a transverse spot from 2 of
costa, reaching half across wing: cilia whitish-ochreous, barred
with fuscous and dark fuscous irroration. Hind wings ochreous-
whitish, more or less tinged with grey: cilia whitish-ochreous,
sometimes with a grey line.
TRANSVAAL, Pretoria, from November to January (Janse) ; five
specimens.
MELASINA PARAPHRICTIS, sp. nl.
S.18 mm. Head pale yellow-ochreous. Palpi moderate,
densely scaled, pale ochreous-yellowish suffused with fuscous except
towards base and apex. Antennze fuscous mixed with ochreous-
whitish, pectinations 5. Thorax fuscous, posteriorly suffused with
whitish-ochreous. Abdomen fuscous, anal tuft ochreous-yellowish.
Fore wings elongate, moderate, costa moderately arched, apex
obtuse, termen obliquely rounded; all veins separate; light
fuscous, suffusedly strigulated with dark fuscous; costal edge
whitish-ochreous ; indistinct markings outlined by cloudy blackish-
fuscous partly confluent strigule, viz., an angulated fascia near
base, a fascia from 4 of costa to middle of dorsum, another from 2
of costa to tornus, and a transverse spot from costa at 2 reaching
half across wing: cilia fuscous mixed with whitish-ochreous and
dark fuscous. Hind wings dark fuscous; cilia whitish-fuscous,
with darker fuscous subbasal shade.
TRANSVAAL, Pietersburg, in December (Janse); one specimen.
MELASINA MORBIDA, sp. 0.
$. 23mm. Head whitish-ochreous with a few fuscous hairs.
Palpi moderate, loosely scaled, ochreous-whitish mixed with dark
1908. | AFRICAN MICRO-LEPIDOPTERA. 745
fuscous except towards apex. Antenne ochreous-whitish, pecti-
nations 4. Thorax whitish, anteriorly mixed with dark fuscous.
Abdomen whitish-brownish. Fore wings elongate, costa gently
arched, apex obtuse, termen little rounded, oblique; all veins
separate; white, rather closely strigulated with light fuscous, with
a few blackish strigule; an irregular angulated blackish-fuscous
transverse streak near base, not quite reaching dorsum; a rather
large blackish-fuscous subquadrate spot beneath middle of disc,
with undefined blackish strigule diverging from its upper angles ;
a series of blackish-fuscous strigule at about 2 parallel to termen :
cilia whitish tinged with fuscous, with fuscous antemedian line
and broader apical interrupted fuscous shade. Hind wings
fuscous-whitish, more fuscous-tinged posteriorly ; cilia whitish,
with light fuscous subbasal shade.
German Hast Arrica, Dar-es-Salaam ; one specimen.
MELASINA SAUROPA, Sp. 0.
3g. 26-27 mm. Head, palpi, and thorax pale ochreous more
or less mixed with dark fuscous; palpi moderate, loosely scaled.
Antenne whitish-ochreous somewhat sprinkled with dark fuscous,
pectinations 5. Abdomen light ochreous sprinkled with fuscous.
Fore wings elongate, rather narrow, costa gently arched, apex
obtuse, termen rounded, rather strongly oblique; all veins sepa-
rate; whitish-ochreous strewn with undefined dark fuscous
strigule or suffusedly irrorated throughout with fuscous and dark
fuscous; a dark fuscous spot on costa at 4, and five on posterior
half of costa, becoming smaller towards apex; a very undefined
basal patch of dark fuscous suffusion; a thick irregular suffused
dark fuscous streak proceeding from dorsum near base to 4 of
dise, whence it sends a branch to first costal spot, thence curved
downwards beneath middle of disc to 2, whence it sends a branch
to tornus, and a longitudinal arm beneath costa which is connected
more or less distinctly with all five posterior costal spots: cilia
ochreous-whitish, broadly barred or almost wholly suffused with
dark fuscous. Hind wings fuscous; cilia pale fuscous, with
darker subbasal line and anteapical shade.
NyassaLanp, Mpeta, on Loangwa River, in December; two
specimens.
MELASINA STABULARIA, Sp. 1.
$6. 13mm. Head pale yellowish-ochreous. Palpi moderate,
with dense projecting scales, whitish-ochreous, basal half suffused
with fuscous. Antenne whitish-ochreous, pectinations 5. Thorax
pale yellowish-ochreous sprinkled with fuscous. Abdomen
whitish-ochreous. Fore wings elongate, moderate, costa gently
arched, apex obtuse, termen obliquely rounded; 9 absent; pale
yellowish-ochreous sprinkled with fuscous, margins strigulated
with fuscous; stigmata indicated by spots of fuscous suffusion,
plical beyond first discal: cilia pale yellowish-ochreous, apical
746 MR. E. MEYRICK ON [June 16,
third paler and barred with fuscous irroration. Hind wings
whitish-grey ; cilia whitish-ocbreous.
British East ArricA, Mombasa, in October; one specimen.
MELASINA CYCLATMA, sp. n.
S. 28 mm. Head, palpi, and thorax fuscous sprinkled with
whitish and dark fuscous; palpi rather short, densely scaled,
pointed. Antenne whitish-fuscous sprinkled with dark fuscous,
pectinations 5. Abdomen grey, anal tuft greyish-ochreous. Fore
wings elongate, costa moderately arched, apex obtuse, termen
littie rounded, rather strongly oblique; all veins separate; fuscous,
somewhat sprinkled with whitish and irregularly and suffusedly
irrorated with blackish-fuscous, the confluence of irroration form-
ing several irregular broken longitudinal marks, and three or
four spots on posterior half of costa; a rounded blotch of whitish
suffusion on dorsum before middle, and an irregular streak of
whitish suffusion along posterior third of dorsum and termen to
apex: cilia pale fuscous, with a dark fuscous antemedian shade,
outer half sprinkled with whitish and indistinctly barred with
dark fuscous suffusion. Hind wings grey; cilia whitish-fuscous,
with dark fuscous subbasal shade.
Transvaal, N.E. Pretoria district, from September to December
(Janse); one specimen.
MELASINA ISOSPILA, sp. n.
3. 25 mm. Head pale ochreous. Palpi moderate, with
appressed scales, pale ochreous mixed with fuscous. Antenne
ochreous-whitish spotted with dark fuscous, pectinations 6, lined
with dark fuscous. Thorax fuscous mixed with dark fuscous and
whitish. Abdomen grey. Fore wings elongate, costa moderately
arched, apex obtuse, termen obliquely rounded ; all veins separate ;
light fuscous, irregularly mixed with white and strigulated with
dark fuscous ; a series of irregular dark fuscous spots along costa,
a larger suffused spot beneath middle of disc, and the confluence
of dark strigulation appears to form other irregular markings,
especially an angulated fascia from 2 of costa to tornus, but these
are hardly definable: cilia ochreous-whitish indistinctly barred
with fuscous. Hind wings light fuscous; cilia whitish-fuscous.
ANGo.LA, Bihe; one specimen.
MELASINA IMMANIS, sp. n.
3S 22-26 mm., 2 35 mm. Head light greyish-ochreous
mixed with dark fuscous. Palpi moderate, densely scaled,
tolerably pointed, whitish-ochreous mixed with dark fuscous.
Antenne whitish-ochreous sprinkled with dark fuscous, pecti-
nations in g¢ 6. Thorax whitish-ochreous irrorated with dark
fuscous. Abdomen light fuscous, anal tuft pale ochreous. Fore
wings elongate, more so in @, costa gently arched, apex obtuse,
termen obliquely rounded ; all veins separate; whitish-ochreous,
more or less irrorated with fuscous, and strigulated throughout
1908. ] AFRICAN MICRO-LEPIDOPTERA. T47
with blackish-fuscous ; second discal stigma distinct, moderate,
dark fuscous; an elongate suffused dark fuscous mark beneath
middle of disc, whence proceeds a slightly curved series of shorter
similar marks to apex, sometimes obscured by the fuscous irro-
ration which tends to form a cloudy patch or suffusion in
posterior part of disc: cilia whitish-ochreous mixed with fuscous
and indistinctly barred with dark fuscous suffusion. Hind wings
whitish-ochreous suffusedly irrorated with fuscous except towards
base ; cilia whitish-ochreous, with a fuscous subbasal line.
S. Nigerra, Ogrugu; Gamera, Bathurst; thirteen specimens.
MELASINA DISSOLUTA, sp. n.
¢ 22-27 mm., 9 35-42 mm. Head light yellowish-ochreous,
with a few dark fuscous hairs. Palpi moderate, densely sealed,
pointed, pale ochreous sprinkled with dark fuscous. Antenne
whitish-ochreous sprinkled with dark fuscous, pectinations in ¢ 4.
Thorax whitish-ochreous more or less suffused with fuscous and
sprinkled with dark fuscous. Abdomen fuscous. Fore wings
elongate, more so in 2, costa gently arched, apex obtuse, termen
obliquely rounded ; all veins separate ; whitish-ochreous, more or
less wholly irrorated with fuscous and strigulated with blackish-
fuscous; the confluence of dark strigulation produces irregular
undefined markings, viz., a basal patch with angulated edge, a
fascia from 4 of costa to middle of dorsam, another from 2 of
costa to tornus, connected with preceding in disc, and an inwardly
oblique patch from costa towards apex: cilia whitish-ochreous
mixed with fuscous and indistinctly barred with dark fuscous
suffusion. Hind wings fuscous, rather darker in 3; cilia whitish-
fuscous, with darker fuscous subbasal line.
NyassaLanp, Zomba, 3000 feet, in December ; six specimens.
MELASINA INIMICA, Sp. Nn.
3. 25mm. Head and palpi pale greyish-ochreous mixed with
dark fuscous; palpi moderate, densely scaled. Antenne pale
ereyish-ochreous, pectinations 6, lined with dark fuscous. Thorax
fuscous mixed with dark fuscous. Abdomen fuscous. Fore
wings elongate, moderately broad, costa moderately arched, apex
obtuse, termen obliquely rounded; all veins separate; fuscous
suffusedly strigulated with dark fuscous ; obscure oblique median
and postmedian fasciz indicated by confluence of strigulation,
former marked with a patch of darker suffusion about fold: cilia
fuscous mixed with darker. Hind wings fuscous; cilia whitish-
fuscous, with fuscous subbasal shade.
ANnGoLA, Bihe; one specimen.
An obscure species, but differs from immanis and its allies by
the obviously broader fore wings.
MELASINA SYSTOLAA, Sp. 0.
3 17-19 mm., 2 23-24 mm. Head pale fulvous; tongue
very short. Palpi very long, densely scaled, pale ochreous suffused
748 MR, E. MEYRICK ON [June 16,
with blackish, Antenne in d¢ ochreous-whitish spotted with
dark fuscous, pectinations 5, lined with black, in Q shortly
pectinated, wholly clothed with dense loose dark fuscous scales.
Thorax brownish irrorated with blackish. Abdomen dark fuscous.
Fore wings elongate, more so in 2, costa gently arched, apex
obtuse, termen obliquely rounded; all veins separate; fuscous
sometimes somewhat mixed with whitish and bluish-grey, and
sprinkled with blackish ; costal edge whitish-ochreous ; markings
ochreous-brown irregularly irrorated with black, viz.,an undefined
basal patch, a fascia from 4 of costa to middle of dorsum, another
from 3 of costa to tornus, and some undefined posterior streaks
rising from small spots on costa: cilia dark fuscous, with several
whitish-ochreous bars, sometimes partly obsolete. Hind wings
blackish-fuscous ; cilia dark purplish-bronzy-fuscous, with blackish
subbasal line.
Narat, Durban and Mooi R., in September and October
(Leigh); seven specimens.
This and the following species, which possess a very short
tongue (absent in the rest), are apparently primitive.
|
iS)
MELASINA AMICA, Sp. N.
3g 23 mm., 9 30-34 mm. Head and palpi light ochreous-
orange, palpi short, rough-scaled; tongue very short. Antenne
light ochreous (in ¢ broken). Thorax yellow-ochreous. Ab-
domen light ochreous. Fore wings elongate, more so in @, costa
gently arched, apex obtuse, termen obliquely rounded ; all veins
separate; yellow-ochreous; cilia light yellow-ochreous. Hind
wings in ¢ light greyish-fulvous, in 9 grey; cilia pale ochreous,
with a faint fuscous subbasal line.
NyassaLAnb, Zomba, 3000 feet, in December ; three specimens.
MELASINA LIOCHRA, Sp. N.
gd. 21-22 mm. Head and palpi light fulvous-ochreous ; palpi
moderate, densely scaled. Antenne grey, pectinations 23, lined
with black. Thorax pale ochreous, anteriorly tinged with fuscous.
Abdomen pale yellowish-ochreous. Fore wings elongate, rather
narrow, costa gently arched, apex obtuse, termen slightly rounded,
rather strongly oblique; all veins separate; light yellowish-
ochreous; base of costa blackish ; a blackish dot in disc at 2: cilia
light yellowish-ochreous. Hind wings dark grey; cilia whitish-
ochreous tinged with fuscous.
TRANSVAAL, Pretoria, in October, December, and January
(Janse) ; four specimens.
MELASINA MYLICA, Sp. 0.
g. 22-23 mm. Head light yellowish-fulvous, face yellowish-
white; tongue very short. Palpi moderate, densely scaled,
yellowish-fulvous mixed with fuscous. Antenne whitish-ochreous,
pectinations 3, lined with dark fuscous, Thorax whitish irrorated
1908. } AFRICAN MICRO-LEPIDOPTERA, 749
with pale ochreous. Abdomen ochreous-whitish, anal tuft large,
pale yellowish. Fore wings elongate, narrow, costa slightly arched,
apex obtuse, termen very obliquely rounded; all veins separate ;
white, thinly and finely sprinkled with brown: cilia ochreous-
whitish, finely sprinkled with brown. Hind wings whitish-
ochreous-grey ; cilia whitish-ochreous.
TRANSVAAL, Pietersburg, in October and November (Janse) ;
two specimens.
MELASINA ABACODES, sp. 0.
3S. 20mm. Head pale bright fulvous, face whitish-suffused ;
tongue very short. Palpi moderate, densely scaled, fulvous-
yellowish, suffused with fuscous towards base. (Antenne broken.)
Thorax white, tinged with ochreous anteriorly, Abdomen whitish-
ochreous, anal tuft large, expansible. Fore wings elongate, narrow,
costa gently arched, apex obtuse, termen very obliquely rounded ;
all veins separate; white, with very numerous pale fuscous strigule
or small spots arranged in longitudinal series between veins,
obsolete in anterior half of cell and absent on anterior half of
costal area; a dark fuscous dot on base of costa; costal edge
ochreous-yellowish to origin of cilia: cilia whitish-yellowish, on
termen with two rows of dark fuscous points. Hind wings grey :
cilia whitish-yellowish.
TransvaaL, N.E. Pretoria district, in November (Janse); one
specimen.
HAPSIFERA PARDALEA, Sp. n.
3 9, 20-30mm. Headand palpi pale yellowish-ochreous. An-
tenn and abdomen pale ochreous. Thorax pale ochreous tinged
with brownish and lilac, and mixed anteriorly with dark fuscous.
Fore wings elongate, narrow, costa gently arched, apex obtuse,
termen very obliquely rounded; 9 absent; whitish-ochreous,
irregularly strigulated with ferrugimous, with scattered black
strigule on margins, all these strigulz more or less raised; larger
tufts near base, and an antemedian fascia near dorsum; very
indefinite markings of irregularly mixed ferruginous, lilac-fuscous,
and black scales, forming a basal patch, an oblique fascia before
middle, and a large posterior patch in dise almost reaching apex :
cilia whitish-ochreous, with two indistinct fuscous lines. Hind
wings pale grey tinged with whitish-ochreous; cilia whitish-
ochreous, with more or less indistinct fuscous line.
Natau, Camperdown (2500 feet) and Northdene, in March and
April (Leigh): ten specimens.
The only known species in which vein 9 is absent (coincident
with 7 instead of stalked), but normal in all other respects.
HAPSIFERA OCHROPTILA, Sp, n.
3 2. 23-27 mm. Head, palpi, and thorax ochreous-whitish.
Antenne and abdomen whitish-ochreous. Fore wings elongate,
narrow, costa gently arched, apex obtuse, termen very obliquely
750 MR. E. MEYRICK ON _June 16,
rounded ; 9 out of 7; ochreous-whitish, sometimes strewn with
small yellow-ochreous strigule slightly sprinkled with blackish,
but these are often obsolete except on costa, where they are
distinct; numerous undefined blackish strigule arranged in two
or three longitudinal series in disc from near base to termen;
strong yellow-ochreous tufts as follows, viz., one at base, two
beneath fold, one in dise before middle, three in an oblique series
beyond middle, one towards costa posteriorly, and a series of
smaller ones round posterior part of costa and termen: cilia
ochreous-whitish, sometimes tinged with yellow-ochreous. Hind
wings ochreous-whitish ; cilia whitish-ochreous.
TRANSVAAL, Pretoria, from December to April (Janse); five
specimens.
HAPSIFERA SEPTICA, Sp. 0.
g. 21-23 mm. Head and antennz ochreous-whitish. Palpi
ochreous-whitish tinged with yellowish, terminal joint with a
faint darker subapical ring. Thorax ochreous-whitish spotted
with yellow-ochreous suffusion. Abdomen grey. Fore wings
elongate, narrow, costa gently arched, apex obtuse, termen very
obliquely rounded ; 2 and 3 short-stalked, 9 out of 7; whitish,
with numerous yellow-ochreous strigule or small spots arranged
in longitudinal series; on a submedian streak from near base to
termen, continued along termen to apex, these spots or strigulz
are blackish-grey ; strong yellow-ochreous tufts as follows, viz.,
one at base, two beneath fold, one in dise before middle, three in
an oblique series beyond middle, one towards costa posteriorly,
and aseries of smaller ones round posterior part of costa and
termen: cilia yellow-ochreous, mixed with whitish towards tips.
Hind wings rather dark grey; cilia whitish-ochreous tinged with
grey.
NyYAssALAND, Fort Johnston, in February ; two specimens.
HAPSIFERA MELICERIS, Sp. n.
g. 25mm. Head and palpi whitish-ochreous partially suffused
with ochreous-yellowish. (Antenne broken.) Thorax whitish-
ochreous spotted with ochreous-yellowish suffusion and anteriorly
with blackish. Abdomen light yellow-ochreous. Fore wings
elongate, narrow, costa gently arched, apex obtuse, termen very
obliquely rounded; 9 out of 7; whitish-ochreous strigulated
throughout with blackish ; some irregular undefined streaks and
patches of pale ochreous-orange suffusion; an ochreous-orange
patch in disc before middle, and an oblique transverse patch
beyond middle, containing raised scales ; an ochreous-orange tuft
at base, and two beneath fold ; some small ochreous-orange spots
round posterior part of costa and termen: cilia whitish-ochreous
suffusedly barred with ochreous-orange, with a few dark fuscous
scales. Hind wings blackish; cilia ochreous-yellowish.
Transvaal, N.E. Pretoria district; Natat; from January to
March (Janse); one specimen.
1908. ] AFRICAN MICRO-LEPIDOPTERA. 751
HAPSIFERA GLEBATA, Sp. 0.
dg. 15-19 mm. Head pale greyish-ochreous, with a few dark
fuscous scales. Palpi whitish-ochreous, basal 2 of second joint
and a median band of terminal joint suffused with dark fuscous.
Antenne pale greyish-ochreous, more whitish towards apex.
Thorax pale greyish-ochreous, more or less suffused with dark
fuscous anteriorly and at posterior extremity. Abdomen whitish-
grey. Fore wings elongate, narrow, costa gently arched, apex
obtuse, termen extremely obliquely rounded; 9 out of 7; pale
greyish-ochreous, sprinkled and irregularly strigulated throughout
with dark fuscous; six moderate dark fuscous costal spots ;
stigmata represented by small somewhat raised dark fuscous
spots, plical slightly beyond first discal, second discal larger; a
series of small dark fuscous spots round apex and termen: cilia
whitish-ochreous sprinkled with dark fuscous. Hind wings pale
grey; cilia whitish-ochreous tinged with grey.
TRANSVAAL, Pretoria (Janse); UGANDA, Kampala ; from October
to March ; five specimens.
PITHARCHA, N. g.
Head with dense loosely appressed scales; ocelli present ; tongue
absent. Antenne #,in d simple. Labial palpi moderately long,
curved, ascending: second joint clothed with dense projecting
scales beneath, forming a rough tuft, with two or three long
bristles externally ; terminal joint shorter, loosely scaled, obtuse.
Maxillary palpi short, drooping, filiform. Posterior tibiz clothed
with very long dense hairs. Fore wings with tufts of scales on
surface ; 2 and 3 stalked, 7 and 8 stalked, 7 to apex, 11 from +.
Hind wings 1, elongate-ovate, cilia 2; 2-7 tolerably parallel.
PITHARCHA CHALINAA, Sp. Nn.
5 2. 20-28 mm. Head pale greyish-ochreous. Palpi
whitish-ochreous, second joint except towards apex, and sometimes
two indistinct bands of terminal joint, suffusedly irrorated with
dark fuscous. Antenne pale ochreous suffusedly spotted with
fuscous. Thorax pale greyish-ochreous more or less irrorated
with fuscous and dark fuscous. Abdomen pale greyish-ochreous.
Fore wings elongate, costa gently arched, apex rounded, termen
obliquely rounded; pale whitish-ochreous, sometimes partially
suffused with fuscous, irregularly and suffusedly strigulated with
dark fuscous, more distinctly on margins; a narrow irregular
blackish-fuscous streak from costa at 2 obliquely outwards, sharply
angulated in middle and not reaching dorsum : cilia pale whitish-
ochreous sprinkled with dark fuscous. Hind wingsin ¢ greyish-
ochreous, in 2 grey; cilia whitish-ochreous, in 2 more or less
tinged with fuscous.
S. Nigeria, Wari; Sterra Leone; NyassaLtanD, Zomba and.
Blantyre; in May, four specimens.
Proc. Zoou. Soc.—1908, No. XLVITII. 48
752 MR. E. MEYRICK ON [June 16,
AMYDRIA Clem.
Head rough-haired; ocelli and tongue absent. Antenne 3,
in ¢ stout, simple, basal joint without pecten. Labial palpi
moderate, curved, ascending, second joint clothed with dense
loose scales projecting beneath towards apex, with a row of long
projecting bristles externally or sometimes with numerous
spreading bristles beneath, terminal joint moderate or short, some-
what pointed. Maxillary palpi short. Posterior tibiz loosely
haired. Fore wings with 2 from towards angle, 7 to apex, 11 from
before middle. Hind wings 1, elongate-ovate, cilia $; 2-7 toler-
ably parallel.
AMYDRIA OPTANIA, Sp. n.
3.18 mm. Head, palpi, antenne, and thorax deep yellow-
ochreous; second joint of palpi with external bristles. Abdomen
pale ochreous. Fore wings elongate, rather narrow, costa gently
arched, apex obtuse, termen obliquely rounded; deep yellow-
ochreous, very faintly strigulated with brownish: cilia yellow-
ochreous, tips blackish on termen. Hind wings light fuscous
tinged with yellowish ; cilia whitish-ochreous.
S. Nigeria, Ogrugu ; two specimens.
AMYDRIA LEONTOPA, Sp. 0.
36 17-18 mm. Head and thorax brownish-ochreous, head
fulvous-tinged. Palpi pale ochreous, second joint and a median
ring of terminal joint suffused with dark fuscous, second joint
with external bristles. Antenne greyish-ochreous, more or less
ringed with dark fuscous. Abdomen pale ochreous. Fore wings
elongate, rather narrow, costa gently arched, apex obtuse, termen
very obliquely rounded; brownish-ochreous: cilia concolorous.
Hind wings grey; cilia pale ochreous, more or less sprinkled with
fuscous.
TRANSVAAL, Pretoria, in October and November (Janse); three
specimens.
AMYDRIA OPIFICA, Sp. 1.
3.13 mm. Head and thorax greyish-ochreous mixed with
dark fuscous. Palpi whitish-ochreous, second joint and median
band of terminal joint suffused with dark fuscous, second joint
with external bristles. Antennz fuscous. Abdomen light grey.
Fore wings elongate, rather narrow, costa gently arched, apex
obtuse, termen very obliquely rounded; pale greyish-ochreous
irrorated with fuscous and dark fuscous: cilia whitish-ochreous
irrorated with dark fuscous. Hind wings light grey; cilia pale
greyish-ochreous, sprinkled with grey.
TRANSVAAL, Pietersburg, in September (Janse); one specimen.
1908. ] AFRICAN MICRO-LEPIDOPTERA. 753
AMYDRIA PERCASTIS, sp. n.
3 2. 17-21 mm. Head, palpi, antenne, and thorax dark
fuscous ; second joint of palpi with numerous spreading bristles
beneath and externally, apex of terminal joint whitish-ochreous.
Abdomen in ¢ fuscous, in 2 light greyish-ochreous. Fore wings
elongate, narrow, costa gently arched, apex round-pointed, termen
extremely obliquely rounded; bronzy-fuscous, suffusedly and
indistinctly strigulated with dark fuscous: cilia greyish-ochreous
mixed with dark fuscous. Hind wings light grey; cilia pale
greyish-ochreous or greyish.
Transvaal, Pretoria, from August to October (Janse); three
Specimens.
AMYDRIA TROPHIAS, Sp. 0.
S6.19mm. Head and thorax pale greyish-ochreous irrorated
with dark fuscous. Palpi whitish-ochreous mixed with dark
fuscous, second joint with numerous projecting bristles on both
sides, terminal joint with basal and median dark fuscous bands.
Antenne pale fuscous suffusedly ringed with darker. Abdomen
fuscous. Fore wings elongate, moderate, costa moderately arched,
apex obtuse, termen obliquely rounded ; whitish-ochreous suffused
with pale fuscous and finely irrorated with dark fuscous, irregu-
larly and coarsely strigulated with dark fuscous ; plical and second
discal stigmata forming suffused dark fuscous spots, preceded and
followed by paler spaces; costal edge whitish-ochreous from 3 to
apex: cilia whitish-ochreous mixed with dark fuscous. Hind
wings light fuscous; cilia whitish-ochreous with two fuscous
shades.
TRANSVAAL, Pietersburg and N.E. Pretoria district, from
‘October to December (Janse); one specimen.
SCARDIA PARACOSMA, Sp. 0.
3. 16-17 mm. Head, palpi, and thorax whitish-ochreous,
yellowish-tinged. Antenne grey, blackish-sprinkled. Abdomen
light grey, anal tuft whitish-ochreous. Fore wings elongate,
rather narrow, costa moderately arched, apex obtuse-pointed,
termen very obliquely rounded; 7 and 8 stalked; whitish-
ochreous, slightly yellowish-tinged; base of costa slightly in-
fuscated: cilia whitish-ochreous. Hind wings grey; cilia
whitish-ochreous.
TRANSVAAL, Pretoria, in December (Janse); two specimens.
HYopPRoRA, 0. g.
Head densely rough-haired; ocelli present; tongue absent.
Antenne 4, in ¢ shortly ciliated, basal jomt with pecten of long
seales. Labial palpi moderately long, straight, porrected, second
joint rough-scaled above and densely tufted beneath, terminal joint
short, slender, pointed. Maxillary palpi absent. a as tibia
704 MR. E, MEYRICK ON [June 16,
clothed above with long hairs. Fore wings with 2 from towards
angle, 7 absent, 11 from before middle. Hind wings somewhat
under 1, ovate- lanceolate, cilia 1; 2—7 nearly parallel.
HyYoprRoRA CRYMODES, Sp. Nn.
g.13 mm. Head, palpi, and thorax white. Antenne grey,
base white. Abdomen pale yellow-ochreous. Fore wings elongate,
rather narrow, costa moderately arched, apex pointed, termen
extremely obliquely rounded; white, with a very few scattered
slightly raised black specks; small undefined yellow spots arranged
as under, viz., two beneath costa antericrly, two on fold, one in
dise before middle, one on costa before middle, one in disc at #, and
a series of very indistinct ones round posterior part of costa and
termen: cilia white. Hind wings pale whitish-grey with a faint
yellowish tinge; cilia ochreous-white.
TRANSVAAL, Pretoria, in January (Janse); one specimen.
A DELID &.
Crromi1iA Zell.
This genus differs essentially from Memophora only in the maxil-
lary palpi being short, loosely scaled, drooping, instead of long,
filiform, folded.
CEROMITIA SPILODESMA, Sp. 0.
¢g. 15-18 mm. Hairs of crown white, face fuscous mixed with
whitish. Palpishort, slender, fuscous. Antenne whitish, towards
base ringed with fuscous. Thorax white, shoulders fuscous.
Abdomen rather dark fuscous, apex pale ochreous. Fore wings
elongate, rather narrow, costa moderately arched, apex obtuse,
termen very obliquely rounded; 8 and 9 usually stalked; white,
more or less mixed with pale fuscous, with some scattered dark
fuscous strigulee, especially towards costa; markings dark fuscous ;
an elongate-triangular spot along basal fifth of costa, broadest
posteriorly ; an elongate spot on costa about 3 _ beneath which is
an inwardly oblique bar in disc, indistinctly soamnacer with it; a
rather thick fascia from beyond middle of costa to 2 of dorsum ; suc
large transverse oval spot in disc at #; a series of small spots round
posterior part of costa and termen: cilia whitish-ochreous. Hind
wings grey; cilia whitish-ochreous.
TRANSVAAL, Pretoria, from November to January (Janse); two.
specimens.
CEROMITIA STATHMODES, Sp. 0.
So. 14 mm. Head white, hairs of crown brownish between
antenne. Palpi short, slender, fuscous mixed with white.
Antenne whitish, with fuscous rings becoming indistinct towards.
apex. Thorax whitish, shoulders fuscous. Abdomen rather dark
fuscous, apex pale ochreous. Fore wings elongate, rather narrow,
1908. | AFRICAN MICRO-LEPIDOPTERA. 755
costa gently arched, apex obtuse, termen very obliquely rounded ;
8 and 9 separate; whitish, partially tinged with very pale fuscous,
and sprinkled with dark fuscous; markings dark fuscous; an
elongate spot along basal fifth of costa; three moderate undefined
fascie, first from 4 of costa to 3 of dorsum, second from beyond
middle of costa to # of dorsum, third from ? of costa to tornus,
indented beneath costa; a series of small spots round posterior
part of costaand termen : cilia ochreous-whitish, basal half whitish
barred with fuscous. Hind wings and cilia rather dark grey.
TRANSVAAL, Pretoria, in January (Janse); one specimen.
CEROMITIA LIBROPIS, Sp. Nn.
@.14 mm. Head whitish, forehead suffused with brownish-
ochreous. Palpi short, loosely scaled, whitish. Antennz whitish,
indistinctly ringed with fuscous. Thorax white, sprinkled with
brownish. Abdomen grey. Fore wings elongate, rather narrow,
costa moderately arched, apex obtuse, termen very obliquely
rounded; 8 and 9 stalked; light fuscous, suffusedly mixed with
white, with a few scattered dark fuscous scales ; an undefined spot
of dark fuscous suffusion in middle of disc: cilia pale fuscous
mixed with whitish. Hind wings grey; cilia whitish-grey.
TRANSVAAL, Pretoria, in December (Janse); one specimen.
CEROMITIA PALYNTIS, Sp. n.
3. 16-17 mm. Head white, sides of face and a frontal bar
dark fuscous. Palpi very short, loosely scaled, white. Antenne
whitish. Thorax white, shoulders fuscous. Abdomen fuscous, apex
pale ochreous. Fore wings elongate, rather narrow, costa gently
arched, apex obtuse, termen very obliquely rounded; 8 and 9
separate; white, with a few scattered dark fuscous scales and
strigule ; a slender dark fuscous streak along basal fifth of costa ;
five dark fuscous dots or small spots, viz., two beneath fold at + and
middle, two in dise at 2 and 4, and one towards costa at 3, and
sometimes one or two small additional dots ; a row of dark fuscous
dots round posterior part of costa and termen : cilia ochreous-white.
Hind wings grey; cilia grey, towards tips whitish-suffused.
TRANSVAAL, Pretoria, from December to February (Janse) ;
three specimens.
CEROMITIA AMPHICHROA, Sp. nN.
3. 20mm. Head ochreous-yellow, crown posteriorly whitish.
Palpi short, whitish. Antenne whitish. Thorax white; shoulders
with a dark fuscous spot. Abdomen pale yellowish-ochreous.
Fore wings elongate, rather narrow, costa moderately arched, apex
obtuse, termen very obliquely rounded; 8 and 9 separate, white ;
a thick dark fuscous streak along basal fifth of costa, beneath which
is a short subcostal line; three irregular dark fuscous dots longi-
tudinally arranged in disc, three others nearer costa obliquely
before them respectively, two on fold beneath the first two sub-
756 ON AFRICAN MICRO-LEPIDOPTERA. [June 16,
costal, one between second of these and second discal, and two or
three other irregularly placed specks; an irregular submarginal
series of small dark fuscous dots, and another series round posterior
part of costa and termen: cilia ochreous-whitish. Hind wings.
grey; cilia whitish-ochreous.
TRANSVAAL, Pretoria, in January (Janse); one specimen.
CEROMITIA GLANDULARIS, sp. n.
36.19 mm. Head ochreous-yellowish. Palpi very short:
(defaced). Antennze ochreous-whitish. Thorax ochreous-whitish,
more ochreous anteriorly. Abdomen fuscous, apex pale ochreous..
Fore wings elongate, rather narrow, costa gently arched, apex
obtuse, termen very obliquely rounded; 8 and 9 separate;
ochreous-whitish, towards costa tinged with ochreous-yellowish ;
markings blackish ; a slender streak along basal fifth of costa; five
roundish small spots, viz., two beneath fold at + and 4, two in dise
beyond these respectively, and one towards costa at # in a line
with the posterior pair ; three or four smaller spots or dots between -
these and termen; a series of dots round posterior part of costa
and termen: cilia whitish-ochreous, tips infuscated. Hind wings.
grey; cilia whitish-ochreous, tips infuscated.
NyassaLanD, Zomba, in December; one specimen.
CEROMITIA SPORHA, Sp. n.
¢.16mm. Head white, between and round antenne brown.
Palpi short, white, with loose projecting scales. Antenne whitish
indistinctly ringed with pale fuscous. Thorax whitish, with pale
brownish subdorsal and lateral stripes. Abdomen whitish-ochreous.
Fore wings elongate, rather narrow, costa gently arched, apex
obtuse-pointed, termen very obliquely rounded ; 8 and 9 separate ;
pale brownish, suffusedly mixed with white, with a few scattered
black scales ; extreme costal edge blackish near base: an undefined
subcostal line of black scales from base to about 1; two or three
undefined dots of black scales in disc before middle: cilia pale
brownish mixed with whitish. Hind wings pale grey ; cilia grey-
whitish.
TRANSVAAL, Pietersburg, in March (Janse); one specimen.
CEROMITIA TYROCHLORA, Sp. 0.
¢.14mm. Head light yellow, lower part of face and back of
crown whitish-suffused. Palpi short, yellow-whitish. Antenne
whitish. Thorax yellow-whitish, shoulders suffused with pale
yellowish. Abdomen whitish-ochreous (partly defaced). Fore
wings elongate, rather narrow, costa gently arched, apex obtuse,
termen very obliquely rounded; 8 and 9 separate: pale whitish-
ochreous tinged with yellowish: cilia concolorous. Hind wings
pale grey; cilia pale whitish-ochreous.
Transyvaat, Pretoria, in January (Janse); one specimen.
abe
mace nr,
PZ.,.S, 1908: Buvsis
The 5.
Huth lith. et imp.
CAPE VERDE ISLANDS CAs @ AEiaros
1908. ] ON CAPE VERDE ISLANDS SPONGES. 757
9. On Collections of the Cape Verde Islands Fauna made by
Cyril Crossland, M.A. (Cantab.), B.Sc. (Lond.), F.Z.8.,
(late of the Gatty Marine Laboratory, St. Andrews
University), from July to September 1904.—The Cal-
careous Sponges. By A. G. Taacker, A.R.C.S. (Lond.),
Research Scholar in Zoology at the Royal College of
Science *.
[Received June 15, 1908. |
(Plate XL. f and Text-figures 155-166.)
It is somewhat remarkable that, although a considerable
number of Calecareous Sponges has been described from the
Azores and from the Canary Islands, only one species, Grantia
tuberosa, dredged off St. Vincent by the ‘Challenger,’ has been
hitherto recorded from the Cape Verde Islands. It was therefore
to be expected that the dredging operations which were carried
out by Mr. Crossland, with the aid of a grant from the Carnegie
Trustees, in the summer of 1904, would yield some interesting
results; and such has in fact been the case, for of the twelve
species contained in the present collection, six are altogether new
to science, and several of the remainder are of interest either from
the zodgeographical or systematic point of view.
The twelve species are as follows :—
Leucosolenia panis (Haeckel).
Leucosolenia atlantica, sp. nu.
Leucosolenia canariensis (Miklucko-Maclay).
Sycon quadrangulatum (Schmidt).
Sycon caminatum, sp. n.
Grantia intermedia, sp. 1.
Leucandra verdensis, sp. n.
Leucandra rudifera (Poléjaeff).
Leucandra sericata (Ridley).
Leucandra typica (Polejaeft).
Leucandra crosslandi, sp. n.
Leucandra gemmipara, sp. 0.
Perhaps the most interesting of these species is the remarkable
sponge, Leucandra rudifera, of which only a few fragments have
been previously found and which is characterised by the possession
of some curious and unique spicules in its gastral cortex.
Another fact, which is of some systematic interest and to which
I would call attention, is the comprehensive sense in which I have
been obliged to use the name Leucosolenia canariensis. From the
great variability of certain characters in some of the Ascons in
this collection, it seems probable that the characters by which
certain sponges (notably Leuwcosolenia nanseni Breitfuss, and
* Communicated by Professor ARTHUR DrEnpDy, D.Sc., F.R.S., F.Z.S., Sec.L.S.
+ For explanation of the Plate see p. 782.
758 MR. A. G. THACKER ON [June 16,
Leucosolenia tenuipilosa Dendy) have been supposed to be speci-
fically distinguished from Haeckel’s original Ascaltis canariensis
are really quite inconstant and not of specific value. As used by
me, therefore, the name Leucosolenia canariensis includes sponges
which have been hitherto considered to be specifically distinct
trom each other, and in my description of this species I have
endeavoured to give a detailed justification of this procedure.
The calcisponge fauna of the Cape Verde Islands shows distinct
affinity with that of the western side of the Atlantic. In a
letter to Professor Minchin, Mr. Crossland suggested that the
distribution of these species might prove interesting because the
Islands receive currents from the Gulf Stream. This forecast has
been to some extent corroborated, for Lewcosolenia panis has been
recorded from the coast of Florida (immediately in the course of
the Gulf Stream, of course), and Leucandra typica and Leucandra
rudifera were dredged by the ‘ Challenger’ off Bermuda (also not
far from the Gulf Stream) on the same day and from the same
spot. I think, therefore, that it is not impossible that the Gulf
Stream may be a factor in the distribution of all three species;
but it should not be forgotten that the North Equatorial Current,
which takes its origin near the Cape Verde Islands and meets the
Gulf Stream as the latter issues from the Gulf of Mexico, might
produce precisely the same results by distributing the sponges in
the opposite direction, that is, by carrying them westwards instead
of eastwards. The distribution of Lewcandra typica may not be
of much value in estimating these factors of dispersal, because
having been recorded from Australia it is evidently a widely
distributed if somewhat uncommon species, but in the case of
L. panis and of L. rudifera, each of which has only been found
once before, the facts here stated are of greater significance. Leu-
candra sericata is a sponge inhabiting the Atlantic coast of South
America. Of the remaining two species which are not new,
L. canariensis has a very wide range and Sycon quadrangulatum
has been recorded from the Arctic, from the eastern Atlantic, and
from the Mediterranean. °
Where it has been necessary in the following pages to refer to
individual specimens, I have done so by the use of the Registered
Number (#..) which I have attached to each specimen in the
collection *. The numbers in square brackets—thus [1]—+refer to
the works given in the list of literature at the end. The list only
includes such memoirs as I have had to refer to in the text.
Throughout this paper I have followed the classification of the
Calcarea, set forth by Dendy in [9] and [11]. According to this
system the genera Leucosolenia and Leucandra are very com-
prehensive, the former including all the Homocela and the latter
being extended to include species such as my ZL. verdensis, which
_* The collection has been placed in the University Museum of Zoology at
Cambridge.
1908. ] CAPE VERDE ISLANDS SPONGES. 759
has no oxeote spicules. In the case of these two genera the
classification is to be regarded, I: think, as the expression of a
suspension of judgment. There can be little doubt that both
Leucosolenia and Leucandra will eventually have to be split up
into several distinct genera; but until this can be done with some
reasonable likelihood of finality, it would seem wisest not to
attempt the task.
I take this opportunity of expressing my gratitude to Professor
A. Dendy, F.R.S., who placed the collection at my disposal for
examination, and who has throughout the research given me
invaluable assistance and advice. I have also to thank my friend
Mr. F. J. Bridgman, Marshall Scholar in the Royal College of
Science, who has very kindly sketched the external form of several
of the Heteroceela (Pl. XL. figs. 4,5,7,and 9). And finally I must
acknowledge my indebtedness to the authorities of the British
Museum (Natural History), who gave me access to the National
Collection of Calcarea and have afforded me special facilities for
consulting the literature on the subject, much of which is not
easily obtainable elsewhere.
LEUCOSOLENIA PANIS (Haeckel). (Plate XL. fig. 1 & text-fig. 155.)
Synonymy :—
Ascandra pams Haeckel [14].
The collection contains three specimens of this beautiful little
sponge, all dredged from a depth of 20 fathoms near North Point,
Text-fig. 155.
Spicules from Lewcosolenia panis.
a = Trivadiates. 6 = Quadriradiates. c= Oxeotes. All X 120.
Boa Vista Island. The external form and general characters of
these agree with the description of the species given by Haeckel,
760 MR. A. G. THACKER ON [June 16,
but the largest of the three specimens is not more than one-sixth
the size of that figured by him. The sponge is composed of a
compact reticulum of Ascon-tubes, but there is no common in-
vesting skin or pseudoderm covering the whole colony and no
endogastral network.
Colour (in alcohol) pale brown.
The Skeleton, which consists of all three kinds of spicules, shows
only slight differences from Haeckel’s description (text-fig. 155).
The triradiates are regular and fairly sharply pointed; they
vary in length from 0°12 mm. to 0°18 mm. and in basal width
from 0°011 mm. to 0°016 mm. The quadriradiates are less nume-
rous than the triradiates and differ from them only by the presence
of the fourth ray, which is the same length as the facial rays but
only a little more than half as thick and is straight and sharply
pointed. The oxeote spicules are somewhat smaller than those
described by Haeckel; they vary in length from 0°35 mm. to
0:55 mm. and in maximum width from 0-025 mm. to 0-035 mm.
They are either quite straight or very slightly curved and are
sharply pointed, especially at one end. A few extremely fine
hair-like oxea occur in my specimens; their presence is not to be
regarded as of much systematic importance.
Distribution. Atlantic coast of North America; Florida (Haeckel
[14]); Cape Verde Islands (Crossland Collection).
LEUCOSOLENIA ATLANTICA, sp. n. (Plate XL. fig. 2 & text-fig.
156.)
The collection contains two specimens of this species, both
dredged from a depth of 20 fathoms off North Point, Boa Vista
Island. The sponge is composed of large Ascon-tubes which for
the most part are much separated from each other, but which
occasionally fuse together into a larger mass and then separate
from each other again (Plate XL. fig. 2). There is of course no
pseudoderm uniting the whole colony, and there is no endogastral
network. The diameter of the Ascon-tubes varies from 0°8 mm.
to 1:2 mm. and the thickness of the wall is 0°2 mm. The oscula
have a diameter of about 0°7 mm. The colour (in alcohol) is
straw-yellow.
The Skeleton consists of middle-sized triradiates, of middle-sized
quadriradiates, of large quadriradiates, and of oxeotes (text-fig.156).
The triradiates are regular and have cylindrical, rather sharply
pointed rays. The average size is: length 0°12 mm., width of
rays at base 0°01 mm. These spicules do not vary much in size
eae are distributed without order in the thickness of the body-
wall.
The middle-sized quadriradiates resemble the triradiates except
for the presence of the fourth ray. The apical rays are feebly
developed on these spicules, being thinner than the facial rays
and not more than one quarter as long.
1908. ] CAPE VERDE ISLANDS SPONGES. 761
The large quadriradiates are also regular, have conical, fairly
sharply pointed rays, and are distributed without order in the
walls of the Ascon-tubes. The average length of their facial
rays is 0°2 mm., and the thickness of the same at their bases
is 0°035 mm. The apical rays are straight or slightly curved,
are sharply pointed, are usually about half as long as the
facial rays, and have a basal thickness of 0:°025 mm. These
spicules are quite numerous and do not vary much in size. The
oxeote spicules are arranged transversely in the thickness of the
sponge-wall, and their outer ends, which are slightly bent, project
beyond the surface. They are fairly sharply pointed at both ends.
Their average length is 0°3 mm. and their maximum thickness
0009 mm. They are cylindrical, do not vary much in size, and
are quite numerous.
Text-fig. 156.
Spicules from Leucosolenia atlantica.
a = Triradiates. & = Small quadriradiates. ce = Large quadriradiates.
d = Oxeotes. All X 120.
A few very fine hair-like, but not very long oxea are to be
found in parts of the sponge. These are very probably young
forms, but even if this is not the case I do not think they are to
be considered of much systematic value.
This species appears to be more closely allied to Lewcosolenia
lawa Kirk [15], than to any other previously described Ascon. °
It is distinguished, however, from the New Zealand form (1) by
having the quadriradiates differentiated into spicules of two
distinct sizes, (2) by having oxea considerably thinner than those
of L. laaa, and (3) by its looser external form.
Distribution. Cape Verde Islands (Crossland Collection).
762 MR. A. G. THACKER ON [June 16,
LEUCOSOLENIA CANARIENSIS (Miklucho-Maclay). (Plate XL.
fig. 3 & text-figs. 157-160.)
Synonymy :—
Nardoa canariensis Miklucho-Maclay.
Nardoa sulphurea Miklucho-Maclay.
Nardoa rubra Miklucho-Maclay.
Tarroma canariense Haeckel (Prodromus).
Tarroma sulphurewm Haeckel (Prodromus).
Tarroma rubrum Haeckel (Prodromus).
Ascaltis canariensis Haeckel [14].
Ascaltis compacta Schuftner [22].
Ascaltis canariensis Lakschewitsch [16].
Ascetta coriacea, n. var. Ascaltis coriacea Fristedt [13].
Leucosolenia nanseni Breitfuss [4].
Ascetta coriacea Arnesen [1].
Leucosolenia tenuipilosa Dendy [12].
The examination of a number of Ascons in this collection has
convinced me that I have to deal with a sponge that has been
previously described under several different names and from
various widely separated localities. The sponge in question is to
be regarded as a close ally of Leucosolenia coriacea, one of the
commonest of Homocela, but it differs from the latter species in
one essential character; some of the spicules develop a fourth
ray, thus becoming quadriradiates.
In 1872, in his great monograph ‘ Die Kalkschwaimme,’ Haeckel
described a sponge from the Canary Islands which was charac-
terised by having small, completely regular triradiate and quadri-
radiate spicules with bluntly pointed rays; this sponge he called
Ascaltis canariensis. The form had been previously described
both by himself and by Miklucho-Maclay under three specific
names, a misconception which arose owing to the supposed specific
value of the different colours exhibited by different specimens of
the species. This Ascon was similar to LZ. coriacea, but differed
from the latter by the presence of quadriradiates and by the
smaller size of its spicules.
Some thirteen years later Lakschewitsch [16] recorded the same
species under the same name from Minorca.
In 1898, in describing a collection of Calcarea from Spitzbergen
(a preliminary account of which he had issued two years pre-
viously [3]), Breitfuss [4] described a sponge which he considered
new to science and which he named Leucosolenia nansent.
Having regard to the results of the present investigation, I think
this form cannot be regarded as specifically distinct from.
Haeckel’s <Ascaltis canariensis, and indeed Breitfuss himself
noticed the great similarity between the two forms, for at the
end of his description of Z. nansent he says :—‘‘ Das Skelet von
Leucosolenia nansent erinnert etwas an Leucosolenia (Ascaltis)
canarivensis (M.-Mcl.), unterscheidet sich aber von dieser durch
die Grésse der Nadeln, welche bei letzterer Species beinahe dreimal
1908. ] CAPE VERDE ISLANDS SPONGES. 763
kleiner sind und nur eine Schenkelliinge von 0:04-0:06 mm. bei
einer Dicke von 0:003—0:005 mm. erreichen. Ausserdem ist die
Innenseite der Réhren bei LZ. nanseni stets mit Papillen besetzt,
dagegen bei LZ. canariensis nur bei der Varietat papillata H.”
The measurements which he gives for the spicules of LZ. nanseni
are: length of rays 0:113 to 0:145 mm., thickness at base 0:008
to 0-014 mm. This makes the spicules, as Breitfuss points out,
nearly three times as large as in L. canariensis. I have found,
however, a series of specimens which completely links up the
two forms in this respect. There is one other slight difference
between the two forms which Breitfuss does not mention in his
comparison: in L. canariensis, Haeckel says that the apical rays
of the quadriradiates are straight and are sometimes longer,
sometimes shorter, than the facial rays; whereas in LZ. nanseni
Breitfuss states that the apical rays are slightly curved and are
usually only half the length of the other rays. In this character,
too, I have found specimens intermediate between the two forms.
Since papille were present on the inner surface of the Ascon-tubes
in some of Haeckel’s Ascaltis canariensis, the presence of these
structures in LZ. nanseni is not a specific distinction between the
two forms.
_ L. nansent appears to me to resemble a sponge described by
Schuffher [22] under the name of Ascaltis compacta, even more
than it does the original LZ. canariensis. A. compacta, which was
found off Mauritius, has regular triradiates and quadriradiates
with rays reaching a maximum length of 0:12 mm. and a maximum
thickness of 0-012 mm. The apical rays of the quadriradiates
are 0-084 mm. long, have a basal thickness of 0-009 mm., and are
sharply pointed and slightly bent at their extremities precisely as
in L. nansem. The ratio of the length to the thickness of the
rays is slightly less than in LZ. nanseni, being usually less
than 10 to 1 and sometimes as low as 7 to 1. Schuffner
separated his sponge from Haeckel’s A. canariensis because
(1) it had no papille on the inner surfaces of the Ascon-tubes,
and because (2) of the different shape of the apical rays of the
quadriradiates. With regard to the latter point, I have, as stated
above, found specimens intermediate between the two forms, and
have also found much variation within the limits of the same
specimen. And as for the papille, it is truly remarkable that
whereas one of Schuffner’s reasons for separating his sponge from
A. canariensis was that it never had these structures, Breitfuss
separated his sponges from A. canariensis partly on the ground
that they always did have the papille. This affords a very good
illustration of the kind of confusion which must necessarily arise
if attempts are made to utilise structures, which are known to be
indifferently present or absent within one species, as_ specific
distinctions between that species and other members of the genus.
It therefore appears that this dscaltis compacta is not distinct
from either Z. nanseni or A. canariensis, and I include it in the
species LZ. canariensis.
764 MR. A. G. THACKER ON [June 16,
Finally, a sponge described by Dendy [12] three years ago from
Ceylon under the name Leucosolenia tenwipilosa is to be regarded
as a variety of this species and as standing in the same relationship
to typical specimens of L. canariensis as L. coriacea ceylonensis,
Dendy, does to the typical ZL. coriacea. This variety, L. canart-
ensis tenwipilosa, has regular triradiates and quadriradiates with
rays O0‘-l mm. long by 0:012 mm. thick, the apical rays of the
quadriradiates being very variable but usually longer than the
facial rays. The distinguishing feature of the variety is the
presence of hair-like oxea. Similar hair-like oxea occur in several
of the Cape Verde Islands specimens under discussion.
The collection contains a considerable number of specimens
which I consider belong to this species. ‘They were dredged by
Mr. Crossland from various localities, Boa Vista Island, Porto
Praya, and elsewhere. The sponge forms rather massive colonies
of reticulating Ascon-tubes (Plate XL. fig. 3). The exhalent
openings are true oscula and there is no pseudoderm or endogastral
network, but in examples where the tubes anastomose very closely
more or less indefinite inter-canals are formed. Hach colony
is attached by a somewhat constricted base, and in several cases
there is a short stalk. None of the colonies in my specimens is
very large; the larger sponges have a diameter of from 1 cm. to
2cem. The diameter of the Ascon-tubes varies considerably in
different specimens and in different parts of the same specimen ;
this variation ranges from 0°15 to 0-4 mm. The walls of the
tubes are from 0:02 to 0:035 mm. thick. The colour (in alcohol)
is brown.
The Skeleton consists of regular triradiates, and of quadriradiates
which differ from the triradiates only in the presence of the
fourth ray; some of the triradiates possess a knob evidently
representing an incipient fourth ray. The rays are usually fairly
bluntly pointed (though there is some variation in this respect)
and are nearly cylindrical. The apical rays of the quadriradiates
project at right angles to the facial rays. There is not much
variation in the size of the spicules in the same individual, but
much difference between the spicules of different specimens. The
measurements of the spicules of the following series of five speci-
mens show how completely this series links up JL. canariensis
on the one hand and Z. tenwipilosa and L. nanseni on tae otaer
the sizes of the spicules of these sponges having beer given
above :—
Specimen R.N. 6.—Lengta of rays, 0:06-0:07 mm.
Thickness of rays, 0-006—-0-007 mm.
Apical rays of quadriradiates straight and equal or nearly
equal in length to facial rays.
Specmen RN. 8.— Length of rays, 0°08-0-09 mm.
Thickness of rays, 0°007—0:008 mm.
Apical rays of quadriradiates straight and from one-half to
two-thirds the length of the facial rays.
1908. ] CAPE VERDE ISLANDS SPONGES. 765
Specimen R.N. 9.—Length of rays, 0°1-0-11 mm.
Thickness of rays, 0°009-0-:01 mm.
Apical rays of quadriradiates straight but variable in form ;
sometimes half as thick and nearly as Jong as the facial
rays; sometimes equally thick but not much more than
half as long as the facial rays.
Specimen RN. 14.—Length of rays, 0°12-0°15 mm.
Thickness of rays, 0°013-0-016 mm.
Apical rays of quadriradiates either straight or bent at their
extremities, and from two-thirds to equal length of the
facial rays.
Specimen R.N. 16.—Length of rays, 0'15-0:19 mm.
Thickness of rays, 0°014—-0-016 mm.
Apical rays of the quadriradiates very variable; either
straight or bent at their extremities; either nearly as
thick and half as long as the facial rays, or else much
thinner but longer than the facial rays.
It will be noted that the rays in the first specimen are only
very slightly larger than in Haeckel’s L. canariensis, that in the
second and third specimens the size is intermediate between
Text-fig. 157. Text-fig. 158.
Spicules from four specimens of Leucosolenia canariensis, showing variation
in the size of the spicules in different specimens. All X 120.
L. canariensis and L. nanseni, that in the fourth specimen the
size is almost the same as in L. nanseni, and finally that in the
: 5 ve ‘ ;
fifth specimen the spicules are larger than in Breitfuss’s Ascons.
All the specimens contain some triradiates with knobs repre-
766 MR. A. G. THACKER ON [June 16,
senting incipient fourth rays. As has been already stated, several
specimens contain some thin hair-like oxea, and in some examples
these are quite numerous but in others very scarce; the latter
specimens obviously form connecting links between the typical
form of the species and the variety LZ. canariensis tenuipilosa.
These spicules have a thickness of 0:0015 to 0:003 mm., and vary
in length from 0-05 to 0°3 mm.; they ave sharply pointed.
I found no papille on the inner surface of the Ascon-tubes ;
as these structures were present in some and absent in other
specimens of Haeckel’s ZL. canariensis (and absent in those de-
scribed by Lakschewitsch), this character is manifestly not to be
considered as of specific value.
It remains to discuss the relationship between ZL. coriacea and
L. canariensis. As previously remarked, the only essential dif-
ference between them is that ZL. canariensis has quadriradiate
spicules; but in the Cape Verde Islands examples there appears
to be every gradation between specimens in which the quadri-
radiates are very numerous (quite 50 °/, of all the spicules) and
other specimens in which they are very scarce, and it is obvious
that the latter approach very nearly to LZ. coriacea. Nor is this
all, for Ascons having a few quadriradiate spicules have been
identified and described as L. coriacea. Breitfuss mentions [4]
that his LZ. nanseni is very like a sponge described as Ascetta
coriacea (n. var. Ascaltis coriacea).by Fristedt [13] in 1887.
Fristedt gives only an incomplete description of this sponge and
gives no measurements of the spicules, but he says that the apical
rays of the quadriradiates are more slender than the facial rays
and are slightly curved. He says that he was doubtful at first as
to whether he should identify the sponge as Ascaltis canariensis,
but did not do so because (1) of the different shape of the apical
rays, and (2) of the wide separation of the localities where the
respective specimens were found—his sponges being arctic.
Again, in the year 1900 Arnesen [1] described some sponges
from Norway which he called Ascetta coriacea, but which had some
quadriradiate spicules.
It must therefore, I think, remain more or less a matter of
opinion whether ZL. coriacea and L. canariensis should be main-
tained as separate species; but if they are to be so maintained
it is quite certain that these sponges of Fristedt and Arnesen
should be called LZ. canariensis.
Distribution. Canary Islands (Haeckel [14]); Cape Verde Is-
lands (Crossland Collection); Minorca (Lakschewitsch [16}) ;
Ceylon (Dendy [12]}); Mauritius (Schuffner [22]); N. Atlantic
and Arctic Oceans, and the White Sea (/ristedt [13], Arnesen [1],
Breitfuss [4] [5}).
SYCON QUADRANGULATUM (Schmidt).
Synonymy :—
Syconella quadrangulata O. Schmidt [21].
Sycortis quadrangulata Haeckel [14].
1908. | CAPE VERDE ISLANDS SPONGES. 767
Sycortis quadrangulata Bowerbank (Norman) [2].
Sycandra quadrangulata Lendenfeld [17].
Sycon quadrangulatum Breitfuss ['7].
The collection contains one small specimen of this well-known
species. The length of the specimen is 3°5 mm. and its width
lmm. It was dredged from a depth of 10 fathoms.
Distribution. Mediterranean, Straits of Gibraltar, coasts of Por-
tugal and Brittany (Schmidt [21)}, Haeckel [14], and Lendenfeld
[17}); Guernsey (Bowerbank [2]); the White Sea (Breitfuss ['7]) ;
the Cape Verde Islands (Crossland Collection).
SYCON CAMINATUM, sp.n. (Plate XL. fig. 4 & text-fig. 161.)
Mr. Crossland collected ten specimens of this new sponge,
several of which are quite young individuals. The general form
assumed by this Sycon is shown in Plate XL. fig. 4. It grows
erect, either singly or in groups, and its greatest diameter is about
one quarter its height. The largest specimen has a length (ex-
cluding the proboscis or peristome) of 14 mm. and its greatest
width is 4mm. The terminal osculum in this specimen has a
diameter of 18 mm. A very characteristic and well-developed
peristome or proboscis is present in all cases, and from this struc-
ture I have derived the specific name. The length of this peri-
stome is variable; in one specimen it reaches as much as 4 mm.,
the length of the body of the sponge being only 8mm. There is
no true oscular crown.
The general structure of the sponge and the character of the
body-wall are typical of the genus Sycon. There are radial
chambers stretching through the whole thickness of the body-wall,
crowned by tufts of oxea, and flanked by definite incurrent canals.
There is a well-developed gastral cortex. The body of the sponge
is nearly circular in transverse section. The thickness of the
body-wall in the middle of the sponge, and therefore the length
of the radial chambers, is 1 mm. The breadth of the radial
chambers at their distal ends is 0°16 mm., and at their proximal
ends 0'1 mm.; they are either spherical or somewhat ellipsoid in
cross-section. The inter-canals open widely on to the exterior at
their distal ends, are irregularly ellipsoid or nearly circular in
cross-section, and have about the same diameter as the radial
chambers; their length is equal to the thickness of the wall, 1 mm.
The arrangement of the radial chambers, and the manner in which
their crowning tufts of oxea project from the surface, are so
regular that under an ordinary hand-lens the surface of the sponge
gives an appearance of almost chess-board regularity. The dis-
tance which divides these crowning tufts from each other is, in
the middle portion of the sponge, about 0°16 mm.
Colour (in alcohol) light brown.
The Skeleton consists of trirvadiates, quadrirvadiates, and oxea,
and may be said to consist of four divisions: (1) that of the radial
chambers, (2) that of the tufts of spicules at the ends of the radial
chambers, (3) that of the gastral cortex, and (4) that of the
peristome.
Proc. Zoou. Soc.—1908, No. X LIX. Ag
768 MR. A. G. THACKER ON [June 16,
The tubar skeleton consists almost exclusively of triradiate
spicules. The majority of these spicules are markedly sagittal
but there is much variation in this respect, some spicules being
almost equiangular and equiradiate, whilst others have an angle
amounting nearly to 180° and the other two angles only a little
more than 90° each. In typical spicules the sagittal form is due
both to this difference of angles and to the greater length which
is attained by the basal ray. All the rays are usually straight ;
but sometimes the paired rays are slightly curved throughout
their lengths, in which case they always have their convex sides,
never the concave, towards the basal ray; and sometimes the
basal ray is curved at its extremity. In a few cases the basal ray
is shorter than the paired rays. he size of an average triradiate
of the tubar skeleton is as follows:—Length of paired rays,
0:09 mm. ; width at base of rays, 0°012 mm. Length of basal
ray, 0°11 mm.; width at base, 0°012 mm. The great majority of
the spicules are of this size or very close to it, but the rays are
occasionally considerably longer, although never much wider.
A few quadriradiate spicules, with facial rays resembling the rays
of the triradiates and with short apical rays, occur in the proximal
portions of the radial canals; they are, however, very scarce and
are probably of little systematic importance.
The crowning tufts at the ends of the radial chambers are
composed of oxea of three sizes. The largest of these (text-fig.
161, 6) reach a length of 0°8 mm. and a maximum thickness of
0°05 mm. They are usually sharply pointed at both ends, but
they taper to the point more gradually at one (the projecting) end
than at the other, the thickest part of the spicule not being in the
middle of its length; this character is very marked in some
spicules, but in others it is scarcely noticeable. These spicules
are somewhat flattened, are straight or nearly straight, and have
about one-third of their lengths projecting beyond the surface of
the sponge. The tufts also consist in part of much smaller straight
or somewhat curved, sharply pointed oxea (text-fig. 161, c), of a
diameter only about equal to that of the rays of the triradiates ;
these reach a length of 0°25 mm. and have a maximum thickness
of 0:01 mm. Jn addition to these there are a number of very fine
hair-like oxea which may perhaps be young spicules.
The gastral cortex, which is well developed and is four or five
layers of spicules thick, consists of triradiates and quadriradiates,
the apical rays of the latter projecting freely into the gastral
cavity. As in the tubar skeleton, there exists considerable varia-
tion in the spicules; the triradiates are typically somewhat longer
and narrower than those of the radial canals, but otherwise there
is little difference between the triradiates of the two regions.
The average length of the rays is 0°12 mm., and their width at
base 07008 mm. One angle is frequently greater than the other
two, thus making the spicules sagittal. The facial rays of the
quadriradiates resemble the rays of the triradiates; the apical
rays are short and stout and sharply pointed; they are either
1908. ] CAPE VERDE ISLANDS SPONGES. 769)
straight or slightly curved at their extremities; their average
length is 0°05 mm. and their width at base 0:011 mm. The
majority of the quadriradiates, like the triradiates, are sagittal.
Text-fig. 161.
Spicules from Sycon caminatum.
a = Tubar triradiates. 6 = Large oxeotes. c=Small oxeotes: d= Gastral
quadriradiates. All X 120.
The peristome consists of an irregular and complicated network
of spicules similar to those which occur in the other regions of the
sponge. There are large and small oxea like those crowning the
distal ends of the radial chambers, there are triradiates like those
of the tubar skeleton, and also triradiates and quadriradiates
resembling those of the gastral cortex. Some of the sagittal tyi-
radiates in this region have the concave curve of the paired rays
towards the basal ray. The very minute hair-like oxea which
project from the ends of the chambers do not occur in the peri-
stomal region.
_ This Sycon is most nearly allied to Sycandra (Sycon) helleri
Lendenfeld, a sponge found in the Adriatic Sea [17]. The species
differ, however, in a number of points, perhaps the most important
of which is the presence in my species of the remarkably well-
A; 49%
770 MR. A. G. THACKER ON [June 16,
developed peristome. There is also never an oscular fringe of
spicules in S, caminatum, a structure which is often present in
S. helleri. S. caminatum is further distinguished by the presence
of more than one kind of oxea at the distal ends of the radial
chambers, by the absence from the radial chambers of sagittal
triradiates with the concave curves of their paired rays turned
towards the basal rays, and lastly by the merely sporadic occur-
rence of quadriradiates in the tubar skeleton, these spicules being
numerous in that region in S. helleri.
Distribution. Cape Verde Islands (Crossland Collection).
GRANTIA INTERMEDIA, sp.n. (Plate XL. fig. 5 & text-fig. 162.)
Mr. Crossland collected a single specimen of this interesting
new species. This is an egg-shaped sponge (Plate XL. fig. 5)
6 mm. high and 5 mm. broad. The outer surface is coarsely
hispid, large oxeote spicules projecting in every direction, and
there is a conspicuous osculum at the top (diameter 1 mm.) with
a well-developed oscular fringe, 15 mm. high. The specimen
was dredged from a depth of 20 fathoms off North Point, Boa
Vista Island. Its colour (in alcohol) is pale brown. The dermal
cortex is 0°16 mm. thick, the chamber-layer is 1:4 mm. thick, and
there is a feebly developed gastral cortex, making the total thick-
ness of the body-wall about 1:6 mm. The diameter of the gastral
cavity (at its widest part) is 1°7 mm.
The specimen is not sufficiently well preserved to enable one to
make out the structure of the canal-system in any very great
detail; but the exhalent canals are well developed, and the
chambers are very much branched. The canal-system is really
intermediate between the form typical of the genus Grantia on
the one hand and that of the genus Lewcandra on the other ;
in short, it is of the “sylleibid” type. The tubar skeleton is,
however, articulate and only shows slight signs of becoming
scattered, and for this reason I place the species in Grantia, not
in Leucandra.
The Skeleton consists of all three forms of spicules.
The tubar skeleton is composed of both triradiates and quadri-
radiates. The former are very variable in shape and are usually
very irregular. Their rays are sometimes straight, sometimes
curved, and all three angles are frequently unequal (text-fig.
162, a). The rays vary in length from 0°12 mm. to 0°24 mm.
and in thickness from 0:011 mm. to 0014 mm. Some of the
triradiates in the tubar skeleton show a strong tendency to become
sagittal, and it is the basal rays of these that attain the greatest
length. There are a few, but only a very few, quadriradiates in
the tubar skeleton; the apical rays of these are the same thick-
hess as, but shorter than, the facial rays and are curved at their
extremities. The facial rays resemble the rays of the triradiates.
None of the rays of either triradiate or quadriradiate spicules is
very sharply pointed.
The dermal cortex consists of a compact mass of spicules resem-
bling those of the tubar skeleton ; the vast majority of the spicules
1908. ] CAPE VERDE ISLANDS SPONGES. til
are triradiate, but there are a few quadriradiates as well. Both
triradiates and quadriradiates are sometimes sagittal, and in this
case they have their basal rays pointing towards the base of the
sponge, as is usual in the genus.
The gastral cortex is 0°07 mm. thick, and is composed of tri-
radiate and quadriradiate spicules similar to those occurring in
the tubar and dermal cortical skeleton, but in this portion of the
sponge the quadriradiates form a much larger percentage of the
whole number of spicules than they do in the tubar skeleton or
dermal cortex. The apical rays of the quadriradiates project into
the gastral cavity. There are sagittal triradiates and sagittal
quadriradiates with basal rays pointing towards the base of the
sponge.
Text-fig. 162.
Spicules from Grantia intermedia.
a = Trivadiates X 120. 6 = Quadriradiates X 120. e¢ = Oxeotes X 24.
There are also some “sub-gastral sagittal triradiates,” viz.,
triradiates just buried within the gastral cortex and having a
basal ray pointing horizontally outwards towards the dermal
surface—in the same direction as the basal rays of the irregularly
sagittal triradiates of the tubar skeleton point; the sagittal tri-
radiates in this position closely resemble the sagittal triradiates
in other situations.
There are some enormous oxeote spicules projecting from the
surface of the sponge. These are not arranged in groups, but
emerge from the surface without definite order and either at right
angles to the surface or more or less obliquely. Their proximal
ends are buried more or less deeply in the tubar layer of the body-
wall. The spicules are spindle-shaped, are rather bluntly pointed
at both ends, reach a length of 2 mm., and have a maximum
thickness of 0°06 mm. (text-fig. 162, c).
The oscular fringe is composed of very long and very fine oxeote
spicules. These reach a length of 5 mm. and have a maximum
thickness of 0°008 mm.
Distribution. Cape Verde Islands (Crossland Collection).
1702 MR. A. G. THACKER ON [June 16,
LEUCANDRA VERDENSIS, sp.n. (Plate XL. fig. 6 & text-fig. 163.)
The collection contains only a single specimen of this species.
It was dredged from a depth of 10 fathoms and is a single Leucon
person of sac-like form, nearly cylindrical in shape (Plate XL.
fig. 6).
“The length of the specimen is 6 mm. and its maximum width
3mm. There is a terminal osculum which is naked and the
surface of the body is smooth. The thickness of the body-wall is
about 1 mm. and the diameter of the central gastral cavity is
about the same. The colour of the sponge (in spirit) is brown.
Text-fig. 163.
Spicules from Leuweandra verdensis.
«@ = Parenchymal triradiates. } = Regular triradiates of dermal cortex. c=Sagittal
triradiate of dermal cortex. d= Gastral quadriradiate. e¢ = Gastral tri-
radiate. All x 120.
The canal-system is typically leuconoid ; the flagellate chambers,
which are scattered about in the parenchyma between the dermal
and gastral cortices, are spherical or sac-shaped and have a
maximum diameter of about 0:06 mm.
The dermal cortex, which is pierced by passages leading into
large inhalent canals, is only about 0:02 mm. thick ; and the gastral
cortex, which is similarly pierced by apertures leading from the
exhalent canals, is of about the same thickness.
- The Skeleton consists of triradiates and quadriradiates. There
are no oxeote spicules,
The main mass of the skeleton, that of the chamber-layer, con-
sists of numerous large, irregularly arranged, regular or sub
regular triradiate spicules (text-fig. 163, @). They do not vary
1908. | CAPE VERDE ISLANDS SPONGES. 773
much in size; the average length of their rays is 0°3 mm. and
the thickness at base is 0°04 mm.
The skeleton of the dermal cortex consists of three kinds of
spicules :—(1) Regular or sub-regular triradiates similar to those
of the chamber-layer but smaller; average size 0°2 x 0°025 mm.
(2) Sagittal triradiates : basal ray pointing towards the base of the
sponge; these vary up to a maximum size of—paired rays 0°3 x
0:04 mm., basal ray 0°2x0°04 mm.; the basal ray is always
shorter than the other two. (3) Sagittal quadriradiates, similar
to the sagittal trivadiates except for the presence of a straight
apical ray, about one quarter the length of and three quarters the
thickness of the paired rays; these spicules do not appear to reach
such a large size as the sagittal triradiates.
The gastral cortex consists of small quadriradiates and tri-
radiates ; the former are mostly sagittal with curved paired rays,
a long straight basal ray, and a rather short curved apical ray
(text-fig. 163, d). The length of the paired rays in these
spicules is on the average 0°15 mm. and the width 0-015 mm. ;
the basal ray is rather longer and the size of the apical ray is
0:05 x0:009 mm. The quadriradiates are sometimes irregular,
sometimes nearly regular in shape, but they vary very little in size.
The triradiates (text-fig. 163, ¢) of the gastral cortex also vary
in shape, but are usually rather irregular; the average size of
their rays is 0°15 x 0-015 mm.
Distribution. Cape Verde Islands (Crossland Collection).
LEUCANDRA RUDIFERA (Poléjaeff). (Plate XL. fig. 7 & text-fig.
164.)
Synonymy :—
Leuconia rudifera Poléjaeff [19].
A few fragments of this species, which is characterised by the
possession of very peculiar minute oxeote spicules in the gastral
cortex, were collected off Bermuda by the ‘Challenger’ and were
described by Poléjaeff. Mr. Crossland dredged three specimens,
all of which are complete, so that J am fortunately able to add
the description of the external form to that of the skeleton which
was given by Poléjaeff.
The sponge (Plate XL. fig. 7) is composed of an irregularly
massive bulbous Leucon individual, and in the largest specimen
reaches a height of 2cm. and a maximum width of 1‘7 mm. The
smallest specimen is egg-shaped ; no doubt this is the young form
and the irregularly massive appearance is acquired with growth.
There is a large circular osculum at the top; there is a peristome,
and two of the specimens possess oscular crowns. The thickness
of the body-wall varies very much: in the largest specimen it is
as thick as 8 mm. in one place and as thin as 2 mm. in another.
The maximum diameter of the central gastral cavity is 6 mm.
The external surface is only slightly hispid, for the large oxea
either do not project at all or project only slightly. The internal
surface is rough owing to the projection of the apical rays of the
gastral quadriradiates.
774 MR. A. G. THACKER ON [June 16,
The canal-system is typically leuconoid ; the flagellated chambers
are spherical and have a diameter of from 0:03 to 0°05 mm.
Many of the incurrent and excurrent canals are very wide; they
attain a diameter of 1 mm. and in some cases of even more.
The Skeleton consists of the following elements :—(1) Large
triradiates of the parenchyma. (2) Triradiates of the dermal
cortex. (3) Sagittal quadriradiates and triradiates of the gastral
cortex. (4) Large oxeotes, only slightly thicker than the large
triradiates. (5) Minute oxea. (6) Minute spined grapnel-like
oxea of the gastral cortex; these latter form the main distin-
guishing character of the species. Finally, the two smaller speci-
mens have oscular fringes consisting of long fine smooth oxea.
The large triradiates of the parenchyma (text-fig. 164, a),
which make up the main mass of the skeleton, are more constant
in shape than those in Poléjaeff’s specimens. They are regular
or very nearly regular and have sharp-pointed rays. The average
size, from which there is not much variation, is—length 0-45 mm.,
basal width 0:045 mm. A small minority of the spicules depart
from the regular form and are sagittal or irregular in shape, both
rays and angles being unequal.
Text-fig. 164.
Spicules from Leucandra rudifera.
a = Parenchymal trivadiates X 24. 6 = Dermal triradiate X 24. ec = Gastral
quadriradiate X 24. d= Large oxeotes X 24. e = Small oxeotes (of dermal
conten ete.) X 120. f= Minute spined grapnel-like oxeotes of gastral cortex
x b
_ The triradiates of the dermal cortex are variable in shape, being
either sagittal or irregular; the thickness of their rays is about
half that of those of the parenchymal triradiates, being on the
average 0°022 mm.; the rays are frequently slightly curved, are
fairly sharply pointed, and reach a length of 0°35 mm. (text-
fig. 164, 5).
The gastral cortex consists almost entirely of quadriradiates
1908. |] CAPE VERDE ISLANDS SPONGES. 775
(text-fig. 164, c). These are sagittal and resemble those
described by Poléjaeff. Lateral rays 0°35 mm. long, basal ray
0:2 mm. long; apical ray varies in length from 0:05 to 0°12 mm.,
usually slightly curved. All rays of the same basal diameter,
0015 mm. There are a few triradiates in the gastral cortex ;
these only differ from the quadriradiates by the absence of the
apical ray. As in the case of Poléjaeff’s specimens, these quadri-
radiates also occur in small numbers in the walls of the exhalent
canals.
Some large oxeote spicules are scattered about in the paren-
chyma ; the greatest thickness of these is 0°055 mm. and they
reach a maximum length of 2 mm., but usually do not exceed
15mm. Many of these spicules do not project beyond the dermal
cortex at all, and over the greater part of the surface of the sponge
they never have more than one-third of their length projecting ;
near the osculum, however, they extend much further beyond the
surface. The spicules are spindle-shaped and sharply pointed at
both ends (text-fig. 164, d). They are not numerous.
There are also some very small oxea ; these are either straight
or slightly curved, and are fairly sharply pointed at both ends.
They are shorter than those described by Poléjaeff; I found none
longer than 0°25 mm. and they are usually only 0:15 mm. long.
Their thickness is 0°003 mm. They occur either singly or in
small bundles in or just beneath the dermal cortex, beyond which
they often project. They are not numerous.
The minute and very peculiar spined grapnel-like oxea (text-
fig. 164, f) of the gastral cortex form the main specific character of
this Leucon. It is these spicules which Poléjaeff describes as
“verticillate ” oxea, but this term does not appear to be quite
accurate, for the spines are not really arranged in verticils, but
project on each side of the spicule alternately down the length of
the shaft. These oxea reach a length of 0:06 mm. and their
thickness varies from 0:0007 to 0°0011 mm. They are sometimes
quite straight and sometimes curved throughout their length, but
usually the spicule is straight in its distal and bent in its proximal
part, which also seems to be slightly flattened and is slightly
wider than the distal half, but I have not found any spicules so
much expanded in this region as that figured by Poléjaeff in
plate vil. fig. 3.@, 11. [19]. Ina few of the spicules the spiked
head is much reduced, these spicules being evidently intermediate
between those with large heads and ordinary spindle-shaped oxea.
Two of the specimens possess oscular fringes composed of
spicules of the usual form—long, very fine oxea of maximum
length 1°3 mm. and width 0:0025 mm. There is of course
nothing remarkable in the presence of oscular fringes in two of
the specimens and its absence in the third, for there are many
Calcarea in which this structure is indifferently present or absent.
There is a peristome, and this structure is best developed on the
specimen without an oscular fringe. It is composed of a complex
network of spicules of all those types which occur in other regions
of the body—large triradiates, small triradiates, small sagittal
776 MR, A. G. THACKER ON [June 16,
quadriradiates, large and small oxea, and minute grapnel-like
oxea.
Colour (in alcohol) white.
Distribution. Bermuda (Poléjaef) ; Cape Verde Islands (Cross.
land Collection).
LEUCANDRA SERICATA (Ridley).
Synonymy :—
Aphroceras sericatum Ridley [20].
Leuconia sericatum Breitfuss [8] [6].
Mr. Crossland collected two specimens of this species. One of
these is a solitary pear-shaped Leucon person 1] mm. high and
with a maximum width of 5 mm.; the other specimen consists of
three Leucon persons united at a common base, the central and
largest of the three individuals being 2 cm, high and having a
maximum width of 8mm. The terminal oscula are protected by
well-developed oscular fringes; the osculum of the largest indi-
vidual is 2°5 mm. in diameter. The scattered flagellate chambers
are usually somewhat elongated, occasionally being four times as
long as broad ; their width varies very little and is about 0°045 mm.
The primary incurrent canals have a diameter of 0° ‘2 mm. and the
ultimate excurrent canals are about the same size.
The Skeleton agrees with the description given by Ridley.
Many of the spicules of the gastral cortex lack apical rays, and
are therefore of course triradiates.
Colour (in alcohol), One specimen is dirty white and the other
brown.
Distribution. 8.E. coast of Brazil (Ridley); Straits of Magellan
(Breitfuss); Cape Verde Islands (Crossland Collection).
There is no doubt that this sponge is very closely allied to the
much commoner species, Z. aspera. It is to be distinguished from
the latter however, (1) by the inferior ratio of the thickness of
the large oxea to that of the parenchymal triradiates, and (2) by
the size and shape of these triradiates themselves, which are larger
than the corresponding spicules in ZL. aspera and are constantly
sagittal. The presence in ZL. sericata of an oscular crown composed
of special oxeotes, which is given by Ridley as a difference between
the two forms, is a distinction which will not hold good, as this
structure also occurs in many specimens of L. aspera.
LEUCANDRA TYPICA (Poléjaeff).
Synonymy :—
Leuconia typica, var. massa Poléjaeff [19].
» var. tuba Poléjaeff [19].
Leucundra typica, var. tuba Lendenteld [18].
Leucandra ta ypica Dendy [10].
Leucona typica Breitfuss [6 1.
Myr. Crossland dredged five specimens of this widely distri-
buted species. One specimen consists of a mass of very wide,
very short, united Leucon persons, and therefore is of the shape
1908.] CAPE VERDE ISLANDS SPONGES. 177
typical of the var. massa; the total width of this specimen is
2°3 em. and its height is much less. Another specimen, which
consists of a single Leucon person, is elongated and has the form
typical of the var. tuba; its height is 10 mm. and its maximum
width 4mm. The other three specimens are intermediate in size
and shape between the foregoing specimens, and consequently are
of a form intermediate between the two varieties of the species
which were established by Poléjaeff.
The general anatomy and skeleton of my specimens agree with
the descriptions given by Poléjaeff and v. Lendenfeld; the only
peculiarity is in the oxeote spicules. The majority of these
resemble the corresponding spicules of the specimens hitherto
described, 2. ¢., they are small spindle-shaped or rather cylindrical,
sharply pointed, straight or slightly curved, and very variable in
size, reaching a maximum length of 0°3 mm. and a maximum
diameter of 0°005 mm. A minority of the oxea are, however,
peculiar in that they are shorter and wider in proportion to their
leneth than are the ordinary spicules, and are also markedly
flattened and more or less arrow-headed ; the arrow-head is some-
times very conspicuous.
The characters by which Poléjaeff endeavoured to divide this
species into two varieties, ¢wha and massa, do not appear to be
constant. In external form I have, as stated above, three speci-
mens which are intermediate between the varieties. The other
two characters on which Poléjaeff established his varieties are as
follows :—
(a) Apical rays of gastric quadriradiates not longer than
0:06 mm. in tuba, but reaching 0-1 mm. in massa.
(6) Oxeote spicules not longer than 0°15 mm. in massa, but
reaching 0°3 mm, in tuba.
Now with regard to (a) I find that in my specimen, which is of
the tuba external form, the apical rays in question reach a length
of 0-1 mm., and this is not exceeded in any of the other specimens.
Whilst with regard to (5) the oxea reach much the same maximum
length in all the specimens—viz. about 0°3 mm.—and are if
anything rather shorter in my specimen of twba than in the
others. For these reasons I do not think it desirable to retain
the two varietal names.
Distribution. Bermuda (Poléjaeff); East coast of Australia
(v. Lendenfeld) ; Cape Verde Islands (Crossland Collection).
LEUCANDRA CROSSLANDI; sp.n. (Plate XL. fig. 8 & text-fig. 165.)
A large number of fragments and several complete specimens
of this species were dredged by Mr. Crossland from a depth of
15 to 20 fathoms near Boa Vista Island.
The sponge (Plate XL. fig. 8) is pear-shaped, is from twice
to three times as long as it is broad, and usually several Leucon
individuals grow together attached to one another at a common
base. The individuals grow to a size of 1:5. em. by 0°8 cm. Both
external and internal surfaces of the body are rough. On large
specimens the oscula have a diameter of 15 mm.; there is never
778 MR. A. G. THACKER ON [June 16,
a true oscular crown, but an oscular rim or peristome is present ;
this varies very much in size, In some specimens being as short as
1 mm., or even less, whilst in others it reaches the enormous
length of 7mm. The thickness of the body-wall varies from
1 to 25 mm., and the maximum diameter of the central gastral
cavity is 3mm. ‘The canal-system is of the form usual in the
genus; the flagellate chambers are spherical or sac-shaped and are
rather large, having a maximum diameter of 0°09 mm. The
incurrent and excurrent canals havea maximum width of 0-25 mm.
The width of the dermal cortex is 0°1 mm. and that of the gastral
cortex 0:06 mm.
The Skeleton consists of the following elements :—(1) of middle-
sized triradiates of the parenchyma, (2) of small triradiates of the
dermal cortex, (3) of gastral quadriradiates, (4) of large oxea,
three times as wide as the parenchymal triradiates, and (5) of
minute oxea.
The main mass of the skeleton is made up by the middle-sized
triradiates of the parenchyma (text-fig. 165,a). These vary some-
what in shape and size, but are usually sub-regular with rays
0°25 by 0035 mm. Not infrequently they are slightly sagittal
with the basal ray shorter than the lateral rays. The rays are
sharply pointed.
Text-fig. 165.
Spicules from Leucandra crosslandi.
a = Parenchymal triradiates x 120. 6 = Dermal triradiates X 120. ¢ = Gastral
quadriradiates x 120. d= Large oxeotes X 24. e = Minute oxeotes X 120.
‘The dermal cortex consists of a mass of slender-rayed sagittal
triradiates ; the lateral rays spread widely at an angle of from
140° to 160°, and are from twice to three times as long as
the basal ray (text-fig. 165, 6). A typical spicule of this kind
gave the following measurements :—Length of lateral rays
0-18 mm. ; length of basal ray 0:07 mm.; thickness of all rays
at base 0°012 mm. The rays are sometimes considerably more
slender than this.
The skeleton of the gastral cortex consists of sagittal quadri-
1908. ] CAPE VERDE ISLANDS SPONGES. Cs)
radiates (text-fig. 165, c). Similar spicules line the exhalent
canals. The lateral rays are rather stout and are slightly curved ;
they are somewhat longer than the basal ray, which is straight.
The apical ray is from one-third to one-half the length of the
lateral rays. The average length of the lateral rays is 0°15 mm.;
the average thickness of the facial rays at their bases is 0°02 mm.
The apical rays are less thick at their bases and are sometimes
curved at their extremities.
The large oxeote spicules are slightly spindle-shaped, are
slightly curved, and are from twice to three times as thick as
the parenchymal triradiates (text-fig. 165, d). They are com-
pletely buried in the parenchyma or project slightly beyond
the dermal surface or project much beyond. They are sharply
pointed at both ends. In length they vary from 1 mm. to 1:5 mm.
and in width from 0:08 mm. to 0°1 mm.
Minute oxea (text-fig. 165, e) are to be found in all parts of
the sponge, but are most numerous in the dermal cortex, from
which they sometimes project beyond the surface of the sponge.
They are pointed at both ends and are quite or nearly straight.
Their average length is 0°15 mm., and their average maximum
thickness 0°0035 mm. They vary considerably in length but very
little in thickness.
The peristome consists as usual of a network of spicules of all
the types which occur in other parts of the sponge.
Colour (in alcohol) white.
This Leucon, which I have much pleasure in naming after
Mr. Crossland, appears to be more nearly allied to Leucandra
crambessa Haeckel, than to any other previously described species.
It differs from this species, however, by the presence of the
minute oxea, by the larger size of the parenchymal triradiates—
these being about twice the size of those in Z. crambessa—and in
sundry minor details of anatomy.
Distribution. Cape Verde Islands (Crossland Collection).
LEUCANDRA GEMMIPARA, sp.n. (Plate XL. fig. 9 & text-fig. 166.)
Mr. Crossland collected a considerable number of specimens of
this species. The sponge (Plate XL. fig. 9) is elongated and
somewhat flattened, and forms proliferous groups of incompletely
separated Leucon individuals. The individuals grow to a height
of 1-2 em. and a width of 4mm. Both external and internal
surfaces of the body are fairly hispid. On the larger specimens
the oscula have a diameter of 15 mm.; all the specimens have
oscular fringes spicules, but on young individuals it is only feebly
developed. The thickness of the body-wall varies very little; it
is about 1 mm. or slightly less, whilst the maximum diameter of
the central gastral cavity is 2mm. The canal-system is typically
leuconoid; the flagellated chambers are spherical or sac-shaped
and are rather large, having a maximum diameter of 0°1 mm.
The maximum diameter of the excurrent canals is only 0:15 mm.,
and of the incurrent canals even less. The thickness of the dermal
cortex is 0°12 mm., and that of the gastral cortex 0:07 mm.
780 MR. A. G. THACKER ON [June 16,
The Skeleton consists of the following elements :—(1) Rather
small triradiates of the parenchyma; (2) small dermal triradiates ;
(3) gastral quadriradiates and triradiates; (4) large oxea, four
times as thick as the parenchymal triradiates ; (5) minute oxea ;
and (6) long slender oxea of the oscular fringe.
The main mass of the skeleton is composed of parenchymal
triradiates (text-fig. 166, a). These are subregular in shape
and their rays are almost always undulated; the length of the
rays varies from 0:12 mm. to 0:22 mm., and the maximum thick-
ness from 0:015 to 0:024 mm. They are sharply pointed.. A few
of these spicules sometimes develop apical rays of variable size
and shape, thus becoming quadriradiates.
The dermal triradiates are similar to those of the parenchyma,
but are smaller; they rarely exceed 0-1 mm. in length or 0:01 mm.
in diameter.
3 Text-fig. 166.
Spicules from Leucandra gemmipara.
a = Parenchymal triradiate X 120. 6 = Gastral quadriradiate X 120.
c = Large oxeotes X 24. d = Small oxeotes X 120.
The central gastral cavity and a great part of the canal-system
are lined by sagittal quadriradiates and triradiates (text-
fig. 166, 6). The wide-spreading lateral rays are usually slightly
shorter than the basal ray, and are either straight or slightly
curved; all the rays are sharply pointed. The apical ray is
curved at its extremity. A typical quadriradiate gave the follow-
ing measurements :—Length of lateral rays 0°12 mm. ; length of
basal ray 0°15 mm.; length of apical ray 0-06 mm.; maximum
thickness of facial rays 0°012 mm.,and of the apical ray somewhat
less. There are some triradiate spicules of similar form.
Numerous large oxea project from the outer surface of the
sponge. They issue from the dermal cortex at a very acute angle
with the surface and their projecting ends all point towards the
osculum. They are spindle-shaped and are slightly curved, and
gradually but sharply pointed at both ends (text-fig. 166,c). Fully
half the length of the spicule is usually projecting beyond the
1908. ] CAPE VERDE ISLANDS SPONGES. 781
‘surface. These oxea reach a length of 2 mm., and their maximum
-thiekness when full grown is from 0° O7 mm. ‘to 0-08 mm.
In the dermal cortex there are bundles of very small oxea,
many of which project from the surface at various angles. These
spicules (text-fig. 166, d) are more or less spindle- ‘shaped and
are nearly or quite straight ; their diameter is 0-002 mm., and
they reach a maximum length of 0°25 mm. They occur almost
exclusively in the dermal cortex, but an occasional isolated spicule
is to be found in the parenchyma.
The oscular fringe is composed of very long thin oxea. They
‘reach a length of from 1 mm. to 2 mm., and their maximum
diameter is 0-004 mm.
In addition to the fringe there is, as Teele a short peristome
encircling each osculum.
Colour (in alcohol) yellowish-white.
This sponge bears a distinct resemblance to the well-known
Leucandra aspera (Schmidt). It differs from this species, how-
ever, by the presence of minute oxea in the dermal cortex.
Leucandra gemmipara is also not unlike the above described
L. crosslandi, but it is to be distinguished from the latter
(1) by the almost complete confinement of the small oxea to the
dermal cortex, (2) by having much more slender parenchymal
triradiates, and (3) by having smaller and different shaped dermal
triradiates. In addition to these and certain other minor differ-
ences, I may mention that there is an oscular fringe present on all
my specimens of L. gemmipara, but that this structure is not to
be found on any of the specimens of ZL. crosslandi. I hesitate,
however, to include this among the characters of specific value.
Distribution. Cape Verde Islands (Crossland Collection).
REFERENCE List oF LITERATURE.
f1] ArneEsen, E.—“‘ Spongien fra den norske kryst. I. Calcarea.
Systematisk katalog med bemerkninger og bestemmels-
estabel.” In Afhandlinger og Aarsberetning udgivne
Bergens Museum. 1900. No. 5.
[2] BowErpank, J. S—‘“‘ Monograph oi the British Spongiade.”
1864-1882.
[3] Brerrruss, L. L.—* Kalkschwimme der Bremer-Expedition
nach Ost-Spitzbergen im Jahre 1889.” Zool. Anz. xix.1896.
[4] “Die Kalkschwammfauna von Spitzbergen.” Zool.
Jahrbiicher, Syst. xi. 1898.
[5] ——. ‘“ Die Kalkschwimme des Weissen Meeres.” Mém.
Acad. des Sciences St. Pétersbourg, ser. 8, vol. vi. 1898.
6] “Catalog der Calcarea der Zoologische Sammlung
der koniglichen Museums fiir Naturkunde zu Berlin.”
Archiv f. Naturg. 1898.
[7] ——. ‘Die Arctische Kalkschwimme.” Archiv f. Nature.
1898.
8} ——. “Die Kalkschwiimme der Sammlung Plate.” Zool.
Jahrbiicher, Syst. 1897-1898. Supplement iv.
782 ON CAPE VERDE ISLANDS SPONGES. [June 16,
[9] Denpy, A.—“‘ A Monograph of the Victorian Sponges.
Part I. The Homocela.” Trans. R. Soc. Vict. vol. i.
part 1. 1891.
‘Synopsis of the Australian Calcarea Heteroecela.”
Proc. R. Soc. Vict., new series, vol. v. 1892.
‘Studies on the Comparative Anatomy of Sponges.
The Structure and Classification of the Heteroceela.”
Quart. Journ. Micro. Soc. vol. xxxv. p. 159. 1894.
. ‘Report on the Sponges collected by Prof. Herdman
at Ceylon in 1902.” Report of the Pearl Oyster Fishe-
vies, Gulf of Manaar. Royal Society, 1905. Part i.
pp. 59-246. 2
[13] Frisrepr, K.—‘“ Sponges from the Atlantic and Arctic
Oceans and the Behring Sea.” A. E. Nordenskiold.
‘Vega ’-Expeditionens Vetenskapliga Iakttagelser, vol. iv.
Stockholm, 1887.
(14] Harcxer, E.—** Die Kalkschwimme.” 1872.
[15] Kirx, H. B.—* New Zealand Sponges.” 3rd paper. Trans.
N. Z. Institute, vol. xxvii. 1895.
[16] Laxscuewirscu, P.—* Ueber die Kalkschwiémme Menoreas.”
Sitzungsberichte der Naturforscher-Gesellschaft bei der
Universitit Dorpat, vol. vii. p. 336. 1885.
[17] v. Lenpenretp, R.—“ Die Spongien der Adria. Die Kalk-
schwiimme.” Zeitschr. wissenschaft. Zool. vol. ii. 1891.
‘Monograph of the Australian Sponges. Preli-
minary description and classification of the Australian
Calcispongie.” Proc. Linn. Soc. N.S.W. vol. ix. p. 1083.
1884.
[19] Poutsazrr, N.—‘‘ Report on the Calcarea collected by
H.M.S. ‘Challenger.’” 1883.
[20] Riptsy, S O.—“Spongida. Zoological Collections made
during the Survey of H.M.S. ‘ Alert.’” Proc. Zool. Soc.
Lond. 1881.
[21] Scumipr, O.—‘‘ Die Spongien des Adriatischen Meeres.”
1862-1866.
[22] Scnurrner, O.—“‘ Beschreibung einiger neuer Kalk-
schwimme.” Jenaische Zeitsch. f. Naturw. v. der medi-
cinisch-naturwissenschaftlichen Gesellschaft zu Jena,
WO sale MIST
[10]
[11]
[12]
(18:
EXPLANATION OF PLATE XL.
Fig. 1. External form of Leucosolenia panis, X 3.
2. 5 ES Leucosolenia atlantica, X 2.
3h PA FA Leucosolenia canariensis, X 4.
4, 5 A Sycon caminatun, X 4.
Be a 5 Grantia intermedia, X 4.
6. 5 0 Leucandra verdensis, X 4.
Les x i Leucandra rudifera, X 2.
e 55 5 Leucandra crosslandi, X 3.
C
i 35 Leucandra gemmipara, X 4.
No. 57.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON*
April 28th, woe.
Dr, Henry Woopwarp, F.R.S., Vice-President,
in the Chair.
The Secretary read a Report on the additions that had been
made to the Society’s Menagerie during the month of March
1908.
Mr. C. Tate Ruean, M.A., F.Z.8., exhibited a specimen of an
Australian Cat-fish (Cnidoglanis megastoma Richards.), and pointed
out that the so-called second dorsal fin was in reality a procurrent
portion of the caudal, differing from the dorsal and anal fins in
the absence of basal supports.
Mr. Henry ScHERREN, F.Z.8., exhibited on behalf of Mr. Watrrr
Burton, F.Z.8., and briefly described some melanistic and black
Leopard-skins from Abyssinia.
Prof. D’Arcy W. Tuompsoy, F.Z.S., gave an explanation, illus-
trated by models, of his suggestion that the shapes of eggs were
due to physical causes, being determined by the peristaltic pressure
of the oviduct on liquid bodies contained in flexible membranes.
Mr. C. Tare Ree@an, M.A., F.Z.S., read a paper entitled ‘‘ A
Revision of the Sharks of the Family Orectolobide.” Twenty-one
species were described and were referred to eight genera. Atten-
tion was called to the great differences in form, coloration, &ec.
among the members of the family, corresponding to differences in
habits and environment.
* This Abstract is published by the Society at 3 Hanover Square, London,
W., on the Tuesday following the date of Meeting to which it refers. It will
be issued, free of extra charge, to all Fellows who subscribe to the Publications,
along with the ‘ Proceedings’ ; but it may be obtained on the day of publication
at, the price of Siapence, or, if desired, sent post-free for the sum of Six
Shillings per annum, payeble in advance.
20
Mr. F. E. Bepparp, F.B.S., F.Z.S., Prosector to the Society,
gave an account of his identification of an Oligochete Worm
obtained in considerable numbers from a well near Cambridge,
England. He described the worm as a new species of the genus
Phreatothrix, the only other species of which had been described
thirty years ago from the underground waters of Prague.
A Memoir by Mrs. E. W. Sexton, entitled “On the Amphipod
Genus Trischizostoma,’ and communicated to the Society by
Dr. W. T. Catan, F.Z.S., was based on a rich material obtained
by the steamer of the Marine Biological Association in the Bay
of Biscay, and by the steamer of the Irish Department of Agri-
culture off the west coast of Ireland.
A short paper was read by Mr. Henry Scuerren, F.Z.8., on
“Certain Errors respecting George the Fourth’s Giraffe.” The
Author adduced evidence to show that the time the animal lived
at Windsor had been much understated, and added details as to
its life in confinement, the presentation of the skin and skeleton
to the Museum of the Society, and the notes made by R. B. Davis
while painting a picture of the animal at Windsor.
Mr. C. L. Boutencer, F.Z.S., communicated observations on
the breeding-habits of a Cichlid Fish (Zilapia nilotica) which he
had been able to make in the course of a visit last year to
Lake Qurun in the Faytim province of Egypt.
A memoir entitled “A Revision of the Oriental Pelobatid
Batrachians (Genus Megalophrys)” was read by Mr. BEepparp,
F.R.S., in the absence of the Author, Mr. G. A. BouLENGER,
F.R.S., Vice-President of the Society.
The next Meeting of the Society for Scientific Business will
be held on Tuesday, the 12th May, 1908, at half-past Hight
o'clock pP.M., when the following communications will be
made :—
1. C.F. JENKIN.—The CU. Crossland Collection of Calcarea
from Zanzibar and Wasin (British Hast Africa).
Days Le 1th, TURNER, F.4.8.—Notes on the Australian Fossorial
Wasps of the Family Sphegide, with Descriptions of new Species.
3. J.T. Cunntnesam, M.A., F.Z.S.—The Heredity of Secondary
Sexual Characters in relation to Hormones: a Contribution to the
Theory of Heredity.
21
The following communications have been received :—
1. OuprieLtp Tomas, F.R.S., F.Z.S., and R. C. Wroucuron,
F.Z.S.—The Rudd Exploration of 8. Africa.—X. List of Mammals
collected by Mr. Grant near Tette, Zambesia.
2. The Rev. T. R. R. Stespine, M.A., F.B.S., F.L.S., F.Z.S8.—
Zoological Results of the Third Tanganyika Expedition, conducted
by Dr. W. A. Cunnington, 1904—1905.—-Report on the Isopoda
Terrestria.
3. F. HE. Bepparp, M.A., F.R.S.—On the Anatomy of Ané-
echinomys and some other Marsupials, with especial reference
to the Intestinal Tract and Mesenteries of these and other
Mammals.
4, Prof. H. G. Szrney, F.R.S., F.Z.8—The Armour of the
Extinct Reptiles of the Genus Pareiasaurus.
Communications intended for the Scientific Meetings of the
ZOOLOGICAL Society oF Lonpon should be addressed to
P. CHALMERS MITCHELL,
Secretary.
3 HANOVER SQuaRE, Lonpon, W.
May 5th, 1908.
No. 58.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.
May 12th, 1908.
F. DuCaneE Gopmay, Esq., D.C.L., F.R.S8., Vice-President,
in the Chair.
The Secrerary read a Report on the additions that had been
made to the Society’s Menagerie during the month of April
1908.
Mr. W. Woopianp, F.Z.8., exhibited preparations of a new
gland he had found in certain teleostean fishes, and made the
following remarks :—‘“ This new gland is diffuse in form and is
intermingled with the veins and arteries which subdivide to form
the numerous parallel capillaries of the rete mirabile found in
connection with all teleost ‘red bodies.’ It is quite distinct
from the gas-gland, and consists of rows of huge columnar cells,
which are situated in close connection with the blood-vessels,
which possess large nuclei and nucleoli and are packed with
numerous large spherical granules derived from the red-corpuscle
disintegration concerned in the generation of the oxygen con-
tained in the swim-bladder. These granules, thus abstracted by
the gland-cells from the blood, are carried away by special ducts
appertaining to the gland. The discovery of this important
gland confirms Jeger’s view as to the mode of generation of
the bladder oxygen. This gland exists in Gobius, Syngnathus,
Peristedion, Box, and some other genera.”
Mr. T. A. Cowarp, F.Z.S., exhibited a specimen of a Petrel,
Estrelata neglecta Schleg., which was the property of Mr. Arthur
Newstead, of Cheshire, and had been picked up dead, yet in a
* This Abstract is published by the Society at 3 Hanover Square, London,
W., on the Tuesday following the date of Meeting to which it refers. It will
be issued, free of extra charge, to all Fellows who subscribe to the Publications,
along with the ‘Proceedings’; but it may be obtained on the day of publication
at the price of Sixpence, or, if desired, sent post-free for the sum of Six
Shillings per annum, payable in advance.
24
quite fresh condition, at Tarporley in Cheshire, on April Ist, 1908.
This bird is a native of the Southern Pacific, and has almost
certainly never been recorded from the northern hemisphere, and
certainly never from Kurope before.
The Secretary, on behalf of Mr. R. LypEKxKeEr, F.R.S., F.Z.S.,
exhibited the tanned skin of a Wild Cat, obtained by The Hon.
Mason Mitchell, of the American Consular Service, in Sze-chuen.
Mr. Lydekker had compared it with a light-coloured skin of
Felis temmincki from Sikkim, and described it as a new local race
of that species.
Mr. C. Davies SHERBORN, F.Z.S., exhibited a specimen of chert
from the Middle Culm-measures (Carboniferous) of Christon
Down, near Doddiscombe Leigh, Devonshire, showing numerous
large and well-preserved Radiolaria,
A memoir by Mr. C. F. Jenxiy, entitled ‘The Cyril Crossland
Collection of Calcarea from Zanzibar and Wasin (British Hast
Africa),” was communicated by Prof. A. Dunpy, F.R.S., F.Z.8.
Mr. R. E. Turner, F.Z.S., read a paper entitled ‘“‘ Notes on
the Australian Fossorial Wasps of the Family Sphegide, with
Descriptions of new Species.” Highty species were described as
new; and the absence of the genera Oxybelus and Philanthus,
otherwise of world-wide range, from Australia was commented on.
Mr. J.T. Cunntinenam, M.A., F.Z.8., communicated a paper
entitled “The Heredity of Secondary Sexual Characters in
Relation to Hormones, a Contribution to the Theory of Heredity.”
The paper contained an examination and criticism of the most
important recent investigations and theories on the subject by
evolutionists of various schools, namely the theory which attributes
such characters to constitutional causes such as male katabolism,
Prof. Karl Pearson’s biometrical investigation of sexual selection
in man, Castle’s Mendelian theory of the heredity of sex, and
Geoffrey Smith’s views on dimorphism of males and parasitic
castration in Crustacea. The author maintained that all these
contributions were more or less inconsistent with the known facts
concerning the connection between the development of secondary
sexual characters and the functional activity of the primary
gonads. He drew attention to the recent discovery and experi-
mental proof on the part of physiologists that the development of
the characters was due to the stimulus of a chemical substance or
hormone produced by the testis or ovary, and passed into the
blood, and suggested that conversely hormones from parts of
the soma might affect the gametes in the gonads. In this way
the hypertrophy of a part of the body due to external stimulation
might modify the corresponding determinants in the gametes so
25
as to produce some hereditary effect in succeeding generations.
Mr. Cunningham added that his theory was an interpretation in
terms of modern physiology of Darwin’s theory of pangenesis.
The next Meeting of the Society for Scientific Business will
be held on Tuesday, the 26th May, 1908, at half-past Hight
o'clock p.m., when the following communications will be
mace :—
1. OuprietD Tuomas, F.R.S., F.Z.S., and R. C. Wroveuton,
F.Z.S.—The Rudd Exploration of 8. Africa.—X. List of Mammals
collected by Mr. Grant near Tette, Zambesia.
2. The Rev. T. R. R. Steppine, M.A., F.R:S., F.L.S., F.Z.S.—
Zoological Results of the Third Tanganyika Expedition, conducted
by Dr. W. A. Cunnington, 1904-1905.—Report on the Isopoda
Terrestria.
3. F. EK. Bepparp, M.A., F.R.S.—On the Anatomy of Ané-
echinomys and some other Marsupials, with especial reference
to the Intestinal Tract and Mesenteries of these and other
Mammals.
4. Prof. H. G. Suetey, F.R.S., F.Z.S.—The Armour of the
Extinct Reptiles of the Genus Pareiasaurus.
5. The Hon. N. Caries Rotruscuitp, M.A., F.Z.8.—New
Siphonaptera.
The following communications have been received :—
1. OtpFinLD THomas, F.R.S., F.Z.S.—The Duke of Bedford’s
Zoological Exploration of Eastern Asia.—X. List of Mammals
from the Provinces of Chili-h and Shan-si, N. China.
2. JAMES Rircuim, M.A., B.Sc., and D. C. McIntosz, M.A.,
B.Sc., F.R.S.E.—On a Case of Imperfect Development in Zchinus
esculentus.
3. Prof. E. A. Mincuin, M.A., V.P.Z.8S., and D. J. Rem,
M.B., C.M., F.Z.8S.—Observations on the Minute Structure of the
Spicules of Calcareous Sponges.
Communications intended for the Scientific Meetings of the
ZOOLOGICAL Society oF Lonpon should be addressed to
P. CHALMERS MITCHELL,
Secretary.
3 Hanover Square, London, W.
May 19th, 1908.
No. 59.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON*
May 26th, 1908.
Professor EH, A. Mincutn, M.A., Vice-President, in the Chair.
A paper was read by Messrs. OLDFIELD Tuomas, F.R.S., F.Z.S.,
and R. C. Wroucuton, F.Z.S., on Mammals collected by Mr. OC.
H. B. Grant near Tette, Zambesia, being the tenth and last of
the series of papers on Mr. C. D. Rudd’s Exploration of South
Africa.
The importance of this collection was due to the fact that
Tette was the place where Dr. Peters obtained most of the speci-
mens on which his ‘Saugethiere von Mossambique’ (1852) was
based, and the specimens now collected were therefore topotypes
of his species and in consequence of great value in working out
S. African Mammals in general.
104 specimens were referred to, belonging to 32 species.
A résumé was then given of the papers which had been pub-
lished on the collections presented to the National Museum by
Mr. Rudd, both those of the present series and others based more
or less directly upon his specimens.
The exploration had lasted five years, and its results formed
the largest and most complete collection that the Museum had
ever received from any one source. Besides duplicates, 1541
specimens had been registered in the Museum, a large number of
new species and subspecies had been discovered and described,
and many more old and inexactly described species were now
represented by good series of well-prepared skins and skulls.
The collection had, in fact, revolutionized our knowledge of
S. African Mammalogy, and it was impossible to exaggerate the
* This Abstract is published by the Society at 3 Hanover Square, London,
W., on the Tuesday following the date of Meeting to which it refers. It will
be issued, free of extra charge, to all Fellows who subscribe to the Publications,
along with the ‘ Proceedings’ ; but it may be obtained on the day of publication
at the price of Sixpence, or, if desired, sent post-free for the sum of Six
Shillings per annum, payable in advance,
28
benefit that such an exploration was to Zoology in general, and
to the National Museum in particular.
Great credit was also due to Mr. C. H. B. Grant for the
manner in which he had carried out the mission entrusted to
him by Mr. Rudd.
The Rev. T. R. R. Srespine, M.A., F.R.S., F.Z.8., reported
that the small collection of terrestrial Isopoda made by Dr. Cun-
nington on the Third Tanganyika Expedition consisted of four
species. For two of these he instituted the new genus Anchi-
philoscia, distinguished by more penicils on the mandibles and a
different cleavage of the second maxille from Philoscia as founded
by Latreille in 1804, The paper insisted on the need of some
enthusiast able and willing thoroughly to revise all the forms
which had clustered under and about the generic name Philoscia.
Mr. F. E. Bepparp, M.A., F.R.S., F.Z.8., Prosector to the
Society, gave an account of a communication on the anatomy
of Antechinomys and some other Marsupials, with special
reference to the intestinal tract and mesenteries of these and
other Mammals. With the aid of a series of diagrams, the
author described four grades or types into which he divided
the modes of suspension of the mammalian intestinal tract.
Prof. H. G. Seeney, F.R.S., F.Z.8., read a paper on the
dermal armour of the extinct reptiles of the genus Puareia-
saurus. The existence of a dermal armour in Pareisaurus had
been doubted by some authors, but Prof. Seeley was able to
exhibit some actual specimens of scutes which had been obtained
by Mr. J. Van Renen, south of Fraserberg, Cape Colony.
Prof. SEeLEy also exhibited the skull of an extinct reptile of
the genus Diademodon, on which he proposed to found a new
species, and gave an account of the further evidence which it
afforded of the structure and dentition of these South African
reptiles.
A communication from the Hon. N. Cuaries Roruscurp,
M.A., F.Z.8., contained descriptions of many new species of
Siphonaptera.
Addition to Abstract, No. 58, May 12th, 1908.
Mr. LyprexKxer described under the name Felis temmincki
mitchelli the Wild Cat which had been obtained by the Hon.
Mason Mitchell in Sze-chuen, and stated that it differed from a
light-coloured skin of /. temmincki from Sikkim by the much
lighter colour of the upper parts, which were golden tawny, with
a comparatively narrow dorsal streak of light rufous, and by its
tail being golden rufous above.
29
The next Meeting of the Society for Scientific Business (closing
the Session 1907-08) will be held on Tuesday, the 16th J une,
1908, at half-past Hight o’clock P.M., when the following com-
munications will be made :—
1. OLDFIELD Tuomas, F.RB.S., F.Z.S.—The Duke of Bedford’s
Zoological Exploration of Eastern Asia.—X. List of Mammals
from the Provinces of Chili-li and Shan-si, N. China.
2. JAMES Ritcuie, M.A., B.Sc., and D. C. McIntosu, M.A.,
Bebe) BRS. Ho "On a Caso of Imperfect Development in
Echinus esculentus.
3 Prot.’ B.A: Mincuin, M.A., VIZ. Say and Deh Rei,
M.B., C.M., F.Z.S.—Observations on the Minute Structure of
the Spicules of Calcareous Sponges.
Any A. Cuapman, M.D., F.Z.8.—Two new Genera and a new
Species of Indian Lycenide,
5. F. E. Brpparp, M.A., HRS LZ. Sw Contribution to
the Knowledge of Rhinoderma darwini.
6.) BBR Bepparp, M.A., F.R.S., F.Z.8.—Some Notes upon
the Anatomy of Chiromys madagascariensis, with references to
other Lemurs.
7. Miss Annte Porter, B.Se.— Leucocytozoin musculi, sp. n.,
a Parasitic Protozotn from the Blood of White Mice,
Communications intended for the Scientific Meetings of the
ZoeLocicaL Socrery or Lonvon should be addressed to
P. CHALMERS M ITCHELL,
Secretary.
3 HANover Square, Lonpon, W.
June 2nd, 1908.
No. 60.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON,
June 16th, 1908.
Dr. Hunry Woopwarp, F.R.S., Vice-President,
in the Chair.
The SzcrETARY read a Report on the additions that had been
made to the Society’s Menagerie during the month of May 1908.
Dr. A. SuitH Woopwarp, F.R.S., F.Z.8., exhibited photographs
and fragments of skin and bone of a Mammoth and a Rhinoceros
discovered in an ozokerite mine at Starunia, Galicia. The car-
cases of these animals appeared to have found their way into an
old marsh saturated with petroleum, which had completely
preserved them. The photographs and specimens had been
received from Dr. George von Kaufmann, who intended to present
them to the British Museum.
Dr. C. I. Forsyta Masor, F.R.S., F.Z.S., exhibited the lower
jaw of a young Canadian Beaver in which there was present on
each side a small conical tooth anterior to the deciduous pre-
molar. He considered the supernumerary premolar to be a case
of atavism.
Dr. ForsyrH Magor exhibited a set of drawings made from
examples of two species of Castor from the East Runton Forest-
bed, and remarked that truly Forest-bed species were found in
association with Pliocene species. He discussed incidentally the
numerous species of recent European Beavers admitted by
Professor Matschie.
* This Abstract is published by the Society at 3 Hanover Square, London,
W., on the Tuesday following the date of Meeting to which it refers. It will
be issued, free of extra charge, to all Fellows who subscribe to the Publications,
along with the ‘ Proceedings’ ; but it may be obtained on the day of publication
at the price of Sixpence, or, if desired, sent post-free for the sum of Séx
Shillings per annum, payable in advance.
32
Dr. ForsytH Magor also exhibited photographs of Pliocene
Bovine from specimens in the Florence Museum, stating that
these unpublished figures showed the great variability of the
Pliocene Bovine. He added that he endorsed Falconer’s opinion
that these Pliocene Bovine were nearly related to the primitive
Buffaloes from the Siwaliks.
Mr. Oupriznp Tuomas, F.RS., FZS., read a paper on
Mammals from the Provinces of Chih-li and Shan-si, N. China,
collected by Mr. M. P. Anderson, being the tenth of the series
of papers on the results of the Duke of Bedford’s Zoological
Exploration of Eastern Asia.
Very little material had hitherto existed from this part of
Northern China, although a certain number of specimens had
been sent to Paris by Pére David, and it was therefore of great
importance to have a series representing the species he discovered
for comparison with mammals from other regions.
The present collection consisted of about 100 specimens,
belonging to 20 species, of which several are new, the most
notable being
CAPREOLUS BEDFORDI, sp. 0.
Size rather larger than in C’. capreolus, much smaller than in
CO. pygargus. General colour in winter pelage more or less rufous,
quite unlike the grey-brown of C. capreolus. Antlers small and
slender.
Condylo-basal length of a male skull 207 mm.
Hab. Chao-Cheng-Shan, Shan-si, 8000’.
Type. Old female. Original number 1615.
The collection had, as before, been presented to the National
Museum by the Society’s President, the Duke of Bedford, K.G.
Dr. F. A. Baruer, F.Z.S., communicated a paper by Messrs.
JAMES Rirosre, M.A., B.Sc., and D. C. McInrosu, M.A., B.Sc.,
F.R.8.H., entitled “On a Case of Imperfect Development in
Lchinus esculentus.”
Professor H. A. Mincutn, V.P.Z.S., and Dr. D. J. Ret, F.Z.S.,
read a paper on the minute structure of Calcareous Sponge-
spicules. The primary object of this investigation was to demon-
strate, by means of photo-micrographs, certain structures, the
existence of which had been strenuously denied by some of the
most competent of previous investigators : namely, the presence,
after the spicules had been cautiously decalcified, of a residue in
the form of an axial filament which could be stained and rendered
evident by certain dyes, in addition to the sheath universally
acknowledged to exist. The axial filament was found to be very
distinct in the spicules of Clathrinide, but much less so in those
of Leucosolenude and Heterocela. Incidentally the study of the
33
axial filaments led to some interesting conclusions regarding the
comparative morphology of the two principal types of spicules,
monaxon and triradiate, occurring in calcareous sponges.
Dr. T. A. Cuapman, F.Z.8., F.E.S., read a paper pointing out
that Cyaniris chennellu of de Niceville was not a Cyantris (Celas-
trina Tutt), but belonged to a new genus near to Hveres; and
that a specimen in Col. Bingham’s collection placed with chennelliz
was a species almost entitled to be placed in Cyaniris, for which
he proposed new generic and specific names. Another specimen
of the latter species was in the Tring Museum. It was suggested
that de Niceville had both these species together in dealing with
chennelliz, and unfortunately selected as his type the one that was
not a Cyaniris. The two forms probably fly together and are
therefore mimetic.
Mr. F. E. Bepparp, F.R.S., F.Z.S., gave an account of two
communications, entitled respectively, ‘“A Contribution to Know-
ledge of the Batrachian Rhinoderma darwini” and ‘Some Notes
upon the Anatomy of Chiromys madagascariensis, with references
to other Lemurs.”
Miss ANNIE Porter, B.Sc., read a paper, communicated by
Mr. H. B. Fantham, F.Z.8., on “ Leucocytozoon musculi, sp. n., a
Parasitic Protozoén from the Blood of White Mice.” The para-
sites occur in mononuclear and transitional leucocytes and free in
the plasma. The free trophozoites are gregariniform vermicules,
their average size being 10:9 uw long by 5:1 u broad. Small free
forms enter leucocytes and grow partly at the expense of the
nucleus. A cytocyst is formed. The endoglobular parasites
average 8 uw long by 5 pw broad. Schizogony occurs in the bone
marrow. A schizont produces about twelve merozoites, each 4°4
by 0°8 ». Two parasites were sometimes seen in one host cell,
suggesting association, and in one case possible exchange of
chromatin occurred. Positive evidence of male and female forms
was not obtained. Vermicules were found in the gut and
Malpighian tubules of lice, ectoparasitic on the mice, but no sexual
cycle in the louse was observed. Perhaps the lice serve merely
as mechanical agents in the spread of infection.
In a memoir from Mr. E. Meyricn, B.A., F.RS., F.Z.S.,
entitled “ Descriptions of African Micro-Lepidoptera,” 108 species
and 11 genera of Tortricina and Tineina from the African Region
(especially the Transvaal) were described as new.
Prof. A. Drnpy, F.R.S., F.Z.8., communicated a paper by
Mr. A. G. THacker, A.R.C.8,(Lond.), on a collection of Calcareous
Sponges made by Mr. Cyril Crossland in the Cape Verde Islands.
ee
%
:
This Meeting closes the Session’ 1907-1908. T
(1908-1909) will begin on November 3rd next.
Communications intended for the Seientific Meet:
ZOOLOGICAL SocrETY oF LONDON should be addressed to
3 Hanover Square, Lonpon, W.
June 23rd, 1908.
CONTENTS (continwed).
June 16, 1908.
: 7 ; Page
The Secretary. Report on the Additions to the Society’s Menagerie during the month of
PMNs ON) Og staat tent saiciavnicic'c Uriel c w/t «viel eivin Fie es oye Saag deni ia ae eet Tre 629 |
Dr. A. Smith Woodward, F.R.S., F.Z.8. Exhibition of photographs and fragments of skin
and bone of a Mammoth and a Rhinoceros from Galicia ....... SRMIG Oo Cee ob 5 oa cc 630
_ Dr. C. I. Forsyth Major, ERS, F.Z.S. Exhibition of, and remarks upon, the lower jaw
Coa ee OMIM AAO Aint iE CAV OTMG/yeleneltis|c(elers\otei cove o:+18\syah ale\iover nis Gel a witle s <eaae'beale opsielalaials 630
Dr. ©. I. Forsyth Major, F.R.S., F.Z.S. Exhibition of drawings of, and remarks upon,
some remains of two species of Castor from the Hast Runton Forest-bed ............ 630
Dr. C. I, Forsyth Major, F.R.S., F.2.8. Exhibition of, and remarks upon, photographs of
Pliocene Bovine ..... Sacto SUS 6 © sce arta ire Ur ee A gOLy P SeaN eEA ee Nia a 635
1. The Duke of Bedford’s Zoological Exploration in Eastern Asia.—X. List of Mammals
from the Provinces of Chih-li and Shan-si, N. China. By Oxprieup Tuomas, F.R.S.,
Beare reeman (Calas PRENG NCL Ma iersretratatea are a. 8/ei'co «pia sicroia fous Slarcjsie #4 ete ee Gales so gine ieee 635
2. On a Case of Imperfect Development in Hehinus esculentus. By James Rrrcuts, M.A.,
B.Sc., The Royal Scottish Museum, and D. C. McInrosn, M.A., B. a F.R.S.E.
E (blake PRONONG DDE) eh ops) cea foieie oc eie shel ayeres «62 Mae elas Rea cs Nene ACs rat Sect he: Story. coat uate ancl Se 646
3. Observations on the Minute Structure of the Spicules of Calcareous Sponges. By E. A.
Mincuin, M.A., V.P.Z.S., Professor of Protozoology, University of London, and D. J.
ro Me, CAME eH ZS. \\(Plates XXXTV.=XXMVEL) ..oc. aces caer e fawicle nee ents 661
4. Two new Genera (and a new Species) of Indian Lycenids, By T. A. CuApman, M.D.,
Me re Ear CO OOO MLAS fais corey ncibhs a cainrigie foe Sietevas citi wiele’s: es a'e:0: eu anSlo 6 2g wields opelatormtame « 676
5. A Contribution to the Knowledge of the Batrachian Rhincderma darwini. By Frank
BEDE D ATID seal VIG ALE Ie ese eM A Siumetan A melard” Smead sal bine Bs 0M eo eave teiare cetera 678
«6. Some Notes upon the Anatomy of Chiromys madagascariensis, with references to other
Lemurs. By Frank EH. Bepparp, M.A., EVR.S., B.Z.8.. 250... e cee eet e ect e wei 694:
7. Leucocytozoon musculi, sp. n.,a Parasitic Protozoda from the Blood of White Mice. By
Annie Porrtsr, B.Sc. inde Zoological Research Laboratory, University College,
Tondon. (Plate XXXIX.) 2.2... i ele eee teeter eee eee eer seen ce encase 703
8. Descriptions of African Micro-Lepidoptera. By E. Meyricr, B.A., F.B.S., F.Z.S8...... 716
9. On Collections of the Cape Verde Islands Fauna made By Cyril Crossland, M.A.
(Cantab.), B.Sc. (Lond.), F.Z.S8. (late of the Gatty Marine Laboratory, St. Andrews
_ University), from July to September 1904.—The Calcareous Sponges. By A, G. THacker,
A.R.C.8. (Lond.), Research Scholar in Zoology at the Royal College of Science.
epi ratsce NA im Megas so) ox dino ta, a:4 4 ash @ afc slo miele. witalgts\s (ala apsh ore) aataeiGra a erate ween create 757
LIST OF PLATES,
1908, pp. 431-782.
Plate
XXVI. Australian Sphegide: « Oreterehe sali siaeto mae
©. ues CONVERUS (Boddeearenee
XXVIII )
‘ae x thew Siponaptena MAE epee Ok eee ete
X. | ‘ sh
pee )
XXXII. Capreolus bed fordi, on no dln sig nc ae aw
XXXIII. Abnormal Zehinus esculentus ....+ tigi Ne
XXXIV. :
Seen Spee of Caleareous Sponges -...--. 4 vase
XXXVIL.) :
XXXVIII. Ancillary appendages of: 1. “Bethea chennai, x x
2. Notarthrinus binghami, x 4B...... ae 4
XXXIX. Leucocytozoin musculi . 6 st onc asda es vik
XL. Cape Verde Islands Calcarea ...........--
NOTICE.
The ‘ Proceedings’ for the year are issued in four nee page
so that the complete reference i is now P. Z. 8. 1908, p.sst~ Yih
is as follows:— ?
nes » March and April, —
DS » May and June,
ss » November and December,
4 PROCEEDINGS
ae OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF THE
E ZOOLOGICAL SOCLETY
|| _ OF LONDON.
= —ttCéG0,
eee 183- 983.
~ Part LV. CONTAINING PAPERS READ IN
NOVEMBER ann DECEMBER.
_ APRIL 1909.
PRINTED FOR THE SOCIETY, :
SOLD AT THEIR HOUSE IN HANOVER SQUARE.
LONDON:
MESSRS. LONGMANS, GREEN, AND CO}.
? '- PATERNOSTER ROW. =~ leas eo es.
ie ei NAS el eh Ly = me Ut elt e Be aed ae LN Seth lh Pay ae =
oo SS SSS RS De ee Cees
3 a S
Ove y ——--
1) _y ha .
[Price Twelve Shillings. |
LIST SOR SC ONT EN Es:
1908, pp. 783-983.
November 3, 1908.
Page
The Secretary. Report on the Additions to the Society's Menagerie during the months of
June, July, August, and September, 1908 = ...<l.0 «0-080 +s sess ste e + s\n 783
Prof. E. A. Minchin, M.A., V.P.Z.S. Exhibition- of drawings of trypanosomes and
trypanoplasms of freshwater fishes ......... sisie bs eity ME o:6als am fayotore) winlqiepelcicrs eee 785
"The Secretary. Exhibition of a photograph of a young Malayan Tapir ..... Bese Beer sia fi's1%,
1. Description of a new Species of Toad from Sumatra. By Grorrrey Mrapx-WaAxpo, B.A.
(Plate, KEL) © aiaceewicais''s 2c locevns oa '5i, agen cin eherols att emiava ys = eterna er sees. F806
to
. On Mammals from Inkerman, North Queensland, presented to the National Museum by
Sir William Ingram, Bt., and the Hon. John Forrest, By Ouprizip Tuomas, F.R.S.,
F.Z8., and iGuy Dotuman, BLA; > (Plate XI) ces os oe ee ee ee ¢ ovine eae
3. The Sze-chuen and Bhutan Takins, By R.Lypexxur, (Plate XLIII.)....-..+...... 795
4. On an Indian Dolphin and Porpoise, By R. Lyprxker. (Plates XLIV, & XLV.) .... 802
November 17, 1908.
The Secretary. Report on the Additions to the Society’s Menagerie during the month of
October 1908 .)..5 «sys. Bee ae Son a4 Son6 Oe Wyarecctee ca aha siaraarerees ence eae 809
My. EH. E, Austen, F.Z.S. Exhibition of Hving specimens of a Fly, Hermetia illucens L.,
caught:in) Manchester’, .\.4is0.08% ¢,c ale e/niscie so ore See eee ae ee osocurs ls)
Notice of the Reports on the Ruwenzori Expedition Collections ........ Wa diet ohe) oidtaie anemia . 809
1. Contributions to the Morphology of the Group Neritacea of Aspidobranch Gastropode.—
Part I. The Neritide. By Prof. Girzerr C. Bourne, D.Se., F.Z.S. (Plates XLVI- ~
BRC V TE)! sche ewten ee sive 6 balata. ie oieceteln bre carats Bate oie ie ORs ale he) eae ea [2S aeeaeee 3810
2. An unknown Lemur from the Lushai Hills, Assam. By N. Annanpatz, D.Se., C.M.ZS.,
Superintendent; Indian Museum i. Sarey) ee ca ce Sake jae 858
Contents continued on page 3 af Wrapper.
THE ZOOLOGICAL SOCIETY OF LONDON.
Tats Society was founded in 1826 by Sir Sramrorp Rarrizs
5]
Mr. J. Sasrne, Mr. N. A. Vieaors, 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.
COUNCIL,
HIS GRACH THH DUKE OF BEDFORD, K.G., President.
Grorce A. Bovutencer, Esa,
_F.R.S., Vice-President.
Pror. Jonn Rose Braprorp,
M.D., D.8e., F.R.S., Vice-
President.
Aurrep H. Cocks, Ese., M.A.
Tue Re. Hon. Kart or Cromar,
P.C., G.C.B.
CHartes Droummonp, KEsa.,
Treasurer.
Sir Epwarp Dvranp, Br., C.B.
FReprrick Gittert, Kse., Vice-
President.
F. DuCane Gopmay, Esa,
D.C.L., F.R.S., Vice-President.
Tre Marquis or Hamitton, M.P.
E. G. B. Meavz-Watpo, Esa.
| Pror. Epwarp ALFRED MINcHIN,
M.A., Vice-President.
P. CHatuers Mircnett, Ese.,
M.-A:, D:Se., LE-D:., E.R.S.,
Secretary.
Tue Lorp Newton.
W. R. O«itviz-Grant, Esa.
ALBERT Pam, Esa.
Tue Hon. Cuarztes Rotuscutip,
M.A.
Davin Setu-Suira, Esa.
Oxprietp Tomas, Hsa., F.R.S.
Avusyn Truvor-Bartyr, Ksa.,
M.A.
Henry Woopwarp, Ksa., LL.D.
F.R.S., Vice-President.
2
The Society consists of Fellows, and Honorary, Foreign, and
Corresponding Members, elected according to the By-Laws. It
carries out the objects of its foundation by means of the collection
of living animals at Regent’s Park, by its Library at 3, Hanover
Square, W., and by its scientific publications.
The Office of the Society (3, Hanover Square), where all
communications should be sent, addressed to “The Secretary,” is
open from Ten till Five, except on Saturdays, when it closes at
Two P.M.
The Library, under the superintendence of Mr. F. H. Waterhouse,
is open daily at the above hours, except in September.
The Meetings of the Society for General Business are held at the
Office on the Thursday following the third Wednesday in every
month of the year, except in September and October, at Four p.m.
Commencing on January 20th, 1909, these Meetings will be held
on the third Wednesday of the month at 5 p.m.
The Meetings for Scientific Business are held at the Office twice
a month on Tuesdays, except in July, August, September, and
October, at half-past Hight o’cleck p.m.
The Anniversary Meeting is held on the 29th. of April, or the
nearest convenient day, at Four p.m.
The Gardens in the Regent’s Park are open daily from Nine o’clock
until Sunset. Mr. R. I. Pocock, F.LS., is the resident Superin-
tendent. The Prosectorium for Anatomical and Pathological work
at the Gardens is under the charge of Mr. Frank E. Beddard, M.A.,
F.R.S., Prosector, assisted by Mr. H. G. Plimmer, M.R.C.S.,
Pathologist to the Society.
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 £50.
No person can become a Frttow until the Admission Fee and
First Annual Subscription have been paid, or the annual payments
have been compounded for.
Frttows elected aiter the 31st. of August are not liable for the
Subscription for the year in which they are elected.
3
PRIVILEGES OF FELLOWS.
Frettows have Personal Admission to the Gardens with Two
Companions daily, upon signing their names in the book at the
entrance gate.
The Wire or Huspanp of a Fettow can exercise these privileges
in the absence of the Fellow.
Every Fsttow is entitled to receive annually 60 undated Green
Cards, and, when no specific instructions are received, the supply
will be sent in this form. If preferred, however, 20 Green Cards
may be exchanged for a book containing 2 Orders for each
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may also be obtained in lieu of 20 Green Cards. A Green Card
may also be exchanged for 2 Buff Cards for the use of Children
under 12 years of age.
It is particularly requested that Fellows will sign every Ticket
before it goes out of their possession. Unsigned Tickets are not
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Green and Buff Tickets may be used on any day and in any year,
but in no case can two Children be admitted with one Adult
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Meeting, after which the privilege lapses. Frxtows are likewise
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to 1881, if above the value of Five Pounds.
Fettows also have the privilege of subscribing to the Annual
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and Publications relating to Zoology in each year, for the sum of
* The Saturday Orders are not available if the Fellow introduces friends
personally on that day.
4
One Pound Ten Shillings. Separate divisions of volumes 39 to
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Any Fettow, having paid all fees due to the Society, is at liberty
to withdraw his or her name upon giving notice in writing to the
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Ladies or Gentlemen wishing to become Fellows of the Society
are requested to communicate with the undersigned.
P. CHALMERS MITCHELL,
Secretary.
8 Hanover Square, London, W.,
April 1st, 1909.
MEETINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON
FOR
SCIENTIFIC BUSINESS.
(AT 3 HANOVER SQUARE, W.)
1909.
Turspay, AprIn .. 6 and 27 | Turspay, NovemBer 9 and 23
He MAY tia lal amide a om DrceMBER 14
* JUNE.... 15
The Chair will be taken at half-past Hight o'clock in the Evening
precisely.
LIST OF THE PUBLICATIONS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.
THE scientific publications of the Zoological Society of London
are of two kinds—“ Proceedings,” published in an octavo
form, and “ Transactions,” in quarto.
According to the present arrangements, the “ Proceedings”’
contain not only notices of all business transacted at the scien-
tific meetings, but also all the papers read at such meetings
and recommended to be published in the “Proceedings ” by
the Committee of Publication. A large number of coloured
plates and engravings are issued in the volumes of the
“ Proceedings,” to illustrate the new or otherwise remark.
able species of animals described therein. Amongst such
illustrations, figures of the new or rare species acquired in a
living state for the Society’s Gardens are often given.
The “ Proceedings” for each year are issued in four parts,
on the first of the months of June, August, October, and
April, the part published in April completing the volume
for the last half of the preceding year. From J anuary 1901
they have been issued as two half-yearly volumes.
The “Transactions” contain such of the more important
communications made to the scientific meetings of the Society
as, on account of the nature of the plates required to illustrate
them, are better adapted for publication in the quarto form.
They are issued at irregular intervals.
Fellows and Corresponding Members, upon payment of
a Subscription of One Guinea before the day of the Anni-
versary Meeting in each year, are entitled to receive the
Society’s Publications for the year. They are likewise
entitled to purchase the Publications of the Society at 25 per
cent. less than the price charged for them to the Public. A
further reduction of 25 per cent. is made upon purchases of
Publications issued prior to 1881, if they exceed the value of
five pounds.
Fellows also have the privilege of subscribing to the
Annual Volume of the Zoological Record for a sum of 30s.
(which includes cost of delivery), payable on the Ist. of J uly
in each year; but this privilege is forfeited unless the
subscription be paid before the Ist. of December following.
The following is a complete list of the publications of the
Society already issued,
TRANSACTIONS* OF THE ZOOLOGICAL SOCIETY OF LONDON.
Ato. 16 vols. and Index. peeee Bros tite
Vol I., containing 59 Plates.... (1833-85) .... 253) 113) 6, £418 OF
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P. CHALMERS MITCHELL,
Secretary.
3 Hanover Squarr, Lonpon, W.
April 1st, 1909.
1908.] THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 783
November 3, 1908.
FREDERICK GILLETT, Esy., Vice-President,
in the Chair.
The Secretary read the following report on the additions made
to the Society's Menagerie during the months of June, July,
August, and September, 1908 :—
JUNE.
The registered additions to the Society’s Menagerie during the
month of June were 141 in number. Of these 115 were
acquired by presentation, 9 by purchase, 6 were received on
deposit, 5 by exchange, and 6 were born in the Gardens. The
total number of departures during the month, by death and
removals, was 144.
Amongst the additions special attention may be called to the
following :—
One Dent’s‘Cercopitheque (Cercopithecus denti), from the Ituri
Forest, new to the Collection, received in exchange on June 19th.
One Markhoor (Capra megaceros), 3, born in the Menagerie
on June 6th.
Two Somali Ostriches (Struthio molybdophanes), 3 2, from
Somaliland, purchased on June 15th.
The special Collection of Australian Animals reached the
Gardens on June 9th, the total number of arrivals being 603,
viz., Mammals 164, Birds 332, and Reptiles 107. Of these 200
were acquired by presentation, 193 by purchase, 6 were received
on deposit, and 204 by exchange.
_ The following were reported as being new to the Society's
‘Menagerie :—
Mammals.
Potorous platyops.
Perameles nasuta.’
Phascogale flavipes.
Hchidna setosa.
1 Broad-faced Rat Kangaroo.
1 Long-nosed Bandicoot.
2 Pouched Mice.
2 Hairy Hchidnas.
Brrps.
3 Blue Wrens.
2 Black-and-white Fantails.
1 Tree Runner.
1 Cuckoo Shrike.
1 Australian Oriole.
4 Japanese Hawfinches.
1 Harmonious Shrike-Thrush.
4 Spiny-cheeked Honey-eaters.
1 Spine-bill Honey-eater.
Proc. Zoot. Soc.—1908, No. L.
Malurus cyaneus.
Rhipidura tricolor.
Climacteris scandens 2
Graucalis mentalis.
Spectotheres australis 2
Coccothraustes robustus.
Collyriocincla harmonica.
Acanthogenys rufigularis.
Acanthorhynchus tenuirostris.
50
784
5 New-Holland Honey-eaters.
3 Lewin’s Honey-eaters.
8 White-plumed Honey-eaters.
4 White-eared Honey-eaters.
1 Yellow-faced Honey-eater.
6 Yellow-throated Honey-eaters.
1 Fulvous-fronted Honey-eater.
THE SECRETARY ON ADDITIONS TO THE MENAGERIE. | Novy. 3,
Meliphaga nove-hellandie.
Ptilotis chrysotis.
» penicillatus.
5, leweotis.
» chrysops.
Myzantha flavigula.
Glyciphila fulvifrons ?
2 Shoveller Ducks.
*
Spatula rhynchotus.
REPTILES.
Amphibolurus decresii.
Gymnodactylus miliusi.
Pseudelaps diadema.
1 Swan River Lizard.
3 Naked-toed Lizards.
2 Diademed Snakes.
JULY.
. The registered additions to the Society’s Menagerie during the
month of July were 276in number. Of these 170 were acquired
by presentation, 19 by purchase, 51 were received on deposit,
10 by exchange, and 26 were born in the Gardens. The total
number of departures during the month, by death and removals,
was 183.
Amongst the additions special attention may be called to the
following :—
A Swan-Island Capromys (Capromys thoracatus), new to the
Collection, presented by Dr. Percy R. Lowe on July 17th.
Two Arabian Hyraxes (Procavia syriaca), new to the Collection,
presented by H.H. The Sultan of Muscat on July 2nd.
A male Grévy’s Zebra (Hquus grevyt), born in the Menagerie
on July 24th.
A Great Anteater (Jyrmecophaga jubata), presented by J. 8.
da Costa, Esq., on July 25th.
A magnificent Collection of 30 Birds of Paradise belonging to
Six species, new to the Collection, obtained in co-operation with
Sir William Ingram, Bt., on July 3rd.
AUGUST.
The registered additions to the Society's Menagerie during the
month of August were 130 in number. Of these 82 were
acquired by presentation, 4 by purchase, 16 were received on
deposit, 4 by exchange, and 24 were born in the Gardens. The
total number of departures during the month, by death and
removals, was 203.
Amongst the additions special attention may be called to the
following :—
A Kiang (Zguus hemionus), $, from Tibet, deposited on
Aug. 6th.
A Barasingha Deer (Cervus dwvaucelli), $ , born in the Mena-
gerie on Aug. 8th.
1908.| ON TRYPANOSOMES FROM BRITISH FRESHWATER FISHES. 785
A Martial Hawk-Eagle (Spizaétus bellicosus), from 8. Africa,
presented by R. P. Burra, Esq., on Aug. 11th.
A Tiger- Bittern (Vatisgae 5p -asiliense), from Manaos, Brazil,
presented by Col. Don Pedro Suarez, on Aug. 13th.
SEPTEMBER.
he registered additions to the Society’s Menagerie during the
month of September were 197 in number. Of these 91 were
acquired by presentation, 10 by purchase, 34 were received on
deposit, 3 by exchange, and 59 were born in the Gardens. The
total number of departures during the month, by death and
removals, was 161.
Amongst the additions special attention may be called to the
following :—
Two Aye-Ayes (Chiromys madagascariensis), from Madagascar
purchased on Sept. 12th.
ne Tiger (felis tigris sondaica), S$, from Deli, Sumatra,
deposited on Sept. 11th.
One Jaguar (Felis onca), from Marajo, presented by G. L.
Andrews, Esq., on Sept. 4th.
One Black- vumped Duiker (Cephalophus melanorheus), new to
the Collection, from Benguela, presented by H. F. Varian, Esq.,
on Sept. 12th.
Six White-throated Pigeons (Columba albiqularis), from the
Moluccas, deposited on Sept. 24th.
Two South-American Mudfish (Lepidosiren paradoxa), from
Para, new to the Collection, presented by the Goeldi Museum on
Sept. 4th.
Professor E. A. Minchin, M.A., V.P.Z.8., exhibited some
drawings of trypanosomes and trypanoplasms of freshwater fishes
(pike, tench, bream, perch, and eel) studied by him at Sutton
Broad Laboratory. After some remarks upon methods of obtain-
ing and studying these blood-parasites, he pointed out that there
were many problems connected with them still unsolved, particu-
larly those relating to their transmission from fish to fish. It
was generally believed that the infection of fisn was effected by
the intermediary of leeches. A great obstacle to the study of
these questions was the lack of any monographs dealing with
leeches in a general way, no such work having been published
since that of Moquin-Tandon in 1846, and the hope was ex-
pressed that some zoologist would pr oduce a handbook or mono-
graph of the British leeches. Such a work would be of the
greatest assistance to those studying fish-trypanosomes, a group
of parasites which might one day become of great economic
Importance.
50*
786 MR. GEOFFREY MEADE-WALDO ON A [ Nov. 3,
The Secretary exhibited a photograph (text-fig. 167) of a young
Malayan Tapir, and remarked that he had been unable to find
accurate drawings of the young of this species. The photograph
had been given to him by the Right Hon. Sir Cecil Clementi
Smith, P.C., G.C.M.G., M.A., Honorary Member of the Society,
and had been taken from a living example which had been a pet
in his house.
Text-fig. 167.
Young Malayan Tapir, photographed from a living example.
The following papers were read :—
1. Description of a new Species of Toad from Sumatra.
By Grorrrey Mrapu-Waxpo, B.A.*
(Plate XLT.)
Buro VALHALLA, sp. 0.
Head once and a third to once anda half as broad as long:
snout as long as diameter of the orbit, short and blunt. Nostrils
nearer the eyes than the tip of the snout; the eyes equidistant
between snout and the angles of the jaws; interorbital space
flat, its width about equal to that of the upper eyelid, and slightly
greater than that between the nostrils.
* Communicated by E. G. B. MzapE-Wa.po, F.Z.S.
Lee Fea Sin WMO, olin
Green del.et Chromo lith
BUEO VALAAMIAs , Sp. nov.
I. Upper surtace of head. 2. Underside of foot.
1908. ] NEW SPECIES OF SUMATRAN TOAD 787
Tympanum two-thirds to three-fourths diameter of the eye,
very distinct: cleft of the mouth extending back to the posterior -
border of the eye.
Fingers short, blunt; 3rd longest, and the Ist rather longer
than 2nd and 4th, which are about equal in length ; subarticular
tubercles single; two moderate carpal tubercles, the inner quite
twice as large as the outer, both elliptical in shape.
Hind limb moderately elongate, tibia as long as femur ; a con-
spicuous gland on each calf; the tarso-metatarsal articulation
reaches the eye. ‘Toes moderately long, about one-half webbed ;
subarticular tubercles small and single; two small metatarsal
tubercles, the inner more prominent than the outer. No tarsal
fold.
Upper surface covered with anastomosing wrinkles, and with
pores, very conspicuous and different in size. A large prominent
elliptical or oval parotoid gland behind the eye on each side, the
length of this gland contained once and a half in the length of
the head. The lower surface granular, granules of uniform size
and evenly distributed.
Olive-brown above, with a few slight traces of darker markings,
in one specimen a distinct black line along the inner margin of
the parotoid glands. Iris bright yellow, towards the corners
thickly vermiculated with black.
Length 82 mm. from snout to vent.
Two females from Pulo Weh Island, off N. Sumatra.
I propose that the name Bufo valhalle be given to the species,
as I was travelling on Lord Crawford’s yacht ‘ Valhalla’ when
the specimens were obtained.
There seem to be several species very nearly allied to this toad.
I have compared the living examples with specimens of ufo
olivaceus Blanford, Bufo stomaticus Liitken, and Bufo ander-
sonii Bler., in the collection of the British Museum (Natural
History), with the kind help of Mr. G. A. Boulenger, F.R.S.
The extreme prominence of the parotoid glands seems to be
the most marked characteristic, and in this it differs considerably
from B. olivacews, in which these glands are depressed. Another
difference between these two species is to be found in the skin of
the upper surface; in Bufo olivaceus it is nearly smooth, in
B. valhalle, on the contrary, it is wrinkle-covered and of a porous
consistency.
B. stomaticus closely resembles it, but may be distinguished
from it by the toes being three-fourths webbed, whereas 2. val-
halle has them only one-half webbed. The parotoid glands in
B. stomaticus ave only nearly as long as their distance from the
end of the snout.
B. andersonii can be distinguished by the presence of a tarsal
fold.
No specimen of Bufo sumatranus Peters, was available for com-
parison, but in that species the tympanum is only one-fourth the
width of the eye, and it is also the possessor of a tarsal fold.
788 MESSRS. 0. THOMAS AND G. DOLLMAN ON [ Nov. 3,
Another characteristic of B. valhalle is the presence of a con-
siderable glandular swelling on the calf of the hind limb; in none
of the above-mentioned species does this occur; it 1s, however,
conspicuous in the British species B. calamita.
B. valhalle is capable of jumping along at a good pace, and is
very active in catching any insect, however fast; I have frequently
seen them jump quite 6 inches from the ground to catch a moth
running up the side of their cage.
On Mammals from Inkerman, North Queensland, pre-
sented to the National Museum by Sir William Ingram,
Bt., andthe Hon. John Forrest. By OLprreLp THomas,
F.R.S., F.Z.S., and Guy Dotiuman, B.A.
[Received August 7, 1908. |
(Plate XLII.)
After he had made the interesting collection of mammals from
Alexandria, Northern Territory, of which an account was given
two years ago*, Mr. W. Stalker was sent by the same generous
donors to Inkerman, their station on the Burdekin River, near
Townsville, in the southern part of North Queensland, and he
has there formed the very fine collection of which we now give a
list.
This region was hitherto almost entirely unrepresented in the
National Collection, such of the few Queensland specimens as we
possessed being either from the far north, on the Cape York
peninsula, or from near Brisbane; aud the present collection
therefore, including as it does admirable series of all the local
mammals, 1s of exceeding value to the Museum, while its general
scientific interest also proves to be very great.
A study of the collection shows ‘clearly that this part of
Queensland belongs to the northern fauna, ‘the species being all
either those of North Australia, or nearly related to them, while
such southern forms as are represented are generally subspeci-
fically separable from their allies of New South Wales and South
Queensland. The new Wallaby (Macropus ualabatus ingrami)
and the Water Rat (Hydromys chrysogaster reginw) may he
quoted as instances of such subspecific differences; while the
presence of /soodon torosus instead of obesulus, of Dasyurus hal-
lucatus, Macropus agilis and many others, are evidence of the
essentially northern relations of the Townsville region.
Altogether this is one of the most important ‘Australian col-
lections that the Museum has ev er received, and we are therefore
very deeply indebted to Sir William Ingram and the Hon. John
Forrest, the donors of this most colivetnle addition to our National
Gollecnone:
* P. Z. 8, 1906, p. 536.
INVEONT SNIVEVIVA SNdOoUoVvW
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1908. | MAMMALS FROM NORTH QUEENSLAND. 789
1. PrERoPpuUS GOULDI Peters.
3.458. ©. 466.
6. 509. Mt. Elliot, Townsville.
2. PrEROPUS SCAPULATUS Peters.
3. 457, 462, 463, 489, 499, 500, 501, 502
©. 320, 464, 490, 491.
3. CHALINOLOBUS GOULDI Gray.
@. 496.
4, SCOTEINUS GREYI Gray.
Q. 492, 493, 497.
5. MINIOPTERUS SCHREIBERSI Kuhl.
©. 354, 356.
Forearm 47 mm.
6. MINIOPTERUS AUSTRALIS Tomes.
Gs 40) OLaaay
Forearm 38 mm.
7. NYCTINOMUS PLANICEPS Peters.
3. 495.
The skull of this specimen is not so flattened as those of other
examples in the Museum, being rather intermediate in this
respect between V. planiceps and VV. norfolcensis.
8. CANIS Dinco Blumenb.
One specimen.
9. HyDROMYS CHRYSOGASTER REGINA, subsp. nov.
3. 391, 394, 396, 419, 420, 423, 487,506. 2. 508, 393, 398,
400, 422, 505.
In general appearance and size like true 17. chrysogaster, only
very much greyer in colour, especially on the upper surface.
General colour of back dark greyish, between ‘‘olive” and
“sepia” of Ridgway, slightly darker in the middle line. Under
surface white, tinged with buff, though not nearly so rich a buff
as in the other Eastern forms. ‘Tail blackish brown proximally,
the terminal three or four inches white.
Dimensions of type (measured in the flesh) :—
Head and body 336 mm.; tail 320; hind foot 66; ear 19.
Skull—basal length 54:7 mm.; basilar length 52:2; zygomatic
breadth 31:7; length of nasals 20°5; palatilar length 28:5;
palatal foramina 6°7; upper molar series 9.
Type. Adult male. B.M. No. 8.8.8.253. Original number 396.
Collected 11th July, 1907.
From all the described East Australian Hydromys, whether
790 MESSRS. 0. THOMAS AND G. DOLLMAN ON [ Nov. 3,
the New South Wales “lutr “lla ”* is or is not distinct from
the Tasmanian “ chrysogaster,” this Queensland Water-Rat is
separable by its darker colour, less suffused with yellowish or buffy.
Mr. Stalker’s series is remarkably uniform in this respect.
Geoffroy’s Hydromys leucogaster was also from Tasmania, and
probably represents a partially albinistic phase of 1. chrysogaster.
Gould’s H. fulvolavatus is more buffy throughout. Its name
may prove to be tenable for the South Australian representative
of the genus.
Jourdan’s H. fulvo-venter +, a name hitherto entirely over-
looked, was based on a specimen believed to be from Swan River,
but the description is wholly inapplicable to any W. Australian
specimen, and there can be little doubt that the type was an
example of H. chrysogaster, which, coming to M. Jourdan with
his specimen of Macropus irma, was er oneously supposed to have
come from the same place.
10. Mus CULMORUM, sp. n.
3. 296, 297, 298, 299, 300, 321, 324, 327, 329, 445, 481.
o. 278, 295, 301, 303, 304, 305, 306, 322, 323, 325, 326, 330,
361, 381, 434, 445, 447, 449, 450.
One ° in spirit.
Heath eel Burdekin R.; Beach Mount; Mt. Abbot.
A coarse or spiny-haired fulvous Rat with a whitish belly,
allied to MW. tunneyi. Size about as in Mus rattus, or rather
smaller. Fur sparse and coarse, more or less mixed with flattened
spines. General colour above brownish fulvous, varying con-
siderably according to the degree of spimousness. Sides more
buffy. Under surface whitish, often with a tinge of yellow, the
hairs pale slaty basally on the belly, whitish throughout on the
throat and sometimes on the inguinal region. Ears rather short,
practically naked. Upper surface of hands and feet white. Tail
of medium length, longer than in J. tunneyi, its rings (at base)
averaging about 10 to the centimetre, thinly haired, dull brownish,
little hghter below. Mamme, as usual in this group, 2—3=10.
Skull of somewhat the peculiar short broad shape of that of
AM. tunneyi, to which there is no doubt the species is most nearly
allied, but its characters are not so extreme. Its bulle, although
much jarger than those of MW. assimilis, greyi, and terre-regine,
are not so large as those of JZ. twnneyi, and the molars not quite
so broad. The interorbital region is narrower, and not so heavily
ridged.
Dimensions of the type (qhenstmed in the flesh) :—
Head and body 150 mm. ; tail 135; hind foot 29; ear 17.
Skull—greatest length 35: 5 mm. > hrsilen length 30; greatest
breadth 19; ; length of nasals 12:2 : Saysewon bie breadth 4°8 ;
palatilar leneth 16: 7; diastema 9°7 ; palatal foramina 7°5 ; oreatest
dumeter of bulle 9; length of upper molar series 7:1.
“ Hydromys lutrilla, ‘Macleay ” Gould, Mamm. Austr. 1. Introd. p. xxxvi, 1863.
+ Ann. Sci. Nat. (2) viii. p. 372, 1837.
1908. ] MAMMALS FROM NORTH QUEENSLAND. 791
Type. Adult female from Beach Mount.. B.M. No. 7.9.15.21.
Original number 330. Collected 5th May, 1907.
This Rat would appear to be the commonest species of Central
Queensland, and it is probable that some of the specimens that:
have been referred to MW. greyi, M. assimilis, M. terre-regine, and
other species really belong to it. The types of all the latter are
in the British Museum, and we have been able to assure ourselves
of its distinctness from them.
Its only near relative as yet described is W/. tunneyi of the
Northern Territory, and from this it is distinguishable by its
longer tail, rather smaller size, and smaller bulle.
Among the series obtained by Mr. Stalker, there is a good deal
of variation in the breadth of the teeth, the extremes being so far
apart as to suggest specific distinctness. No external characters
corresponding to the tooth differences can be found, however,
and we prefer for the present to put the whole series under one
heading.
11. Mus sp.
Q. 437.
An intermediate species not at present determinable.
12. Mus PATRIUS, sp. n.
3. 406,411. @. 407, 408, 409, 410.
So & @ in spirit.
A Mouse of the forresti-delicatulus group, of about the size of
hermannsburgensis.
Size medium within the group, decidedly larger than JZ. deli-
catulus. General colour above pale wood-brown, becoming slightly
more buffy on the sides. Belly greyish white, fairly sharply
defined, the bases of the hairs slaty, their tips white. Hars of
medium size, practically naked, their few fine hairs buffy. Hands
and feet white; sole-pads 6, rather larger and less sharply defined
than in J. delicatulus, the sole between the pads naked. Tail
slightly longer than head and body, finely haired, brown above,
whitish on sides and below. Mamme 0—2=4.
Skull in size and shape agreeing with that of WZ. hermanns-
burgensis, but the bulle of the normal small size, those of
Prof. Spencer’s species being unusually large.
Molars with the laminee obliquely twisted, as already described
in Mus forresti, although not so much as in that species. A well-
marked anterior supplementary cusp present on m’.
Dimensions of the type (measured in the flesh) :—
Head and body 65 mm. (range up to 70); tail 66 (range to 71) ;
hind foot 18; ear 12.
Skull—greatest length 22:7 mm.; basilar length 17:2; nasals
8-2; interorbital breadth 3:2; breadth of brain-case 10°5; pala-
tilar length 10:1; diastema 6; palatal foramina 4°2; length of
upper molar series 4°2.
7192 MESSRS. 0. THOMAS AND G. DOLLMAN ON [ Noy. 3
Type. Aduit female. B.M. No. 7.8.9.44. Original number 408.
Collected 27th July, 1907.
This species may be distinguished from J/. delicatulus by its
larger and from J/. fares by its smaller size, and from JZ.
hermannsbur gensis by its comparatively small bulle. MW. nove-
hollandice Waterh., which is somewhat similar externally, is not
a member of this ‘eroup, having normal molars without supple-
mentary anterior cusps.
My. Stalker states that he dug these mice out of holes, in each
of which he found one male and two females.
13. Mus muscuuus LL.
A number of the introduced House-Mouse.
14. Macropus GIGANTEUS Zimm.
3. 349, 353, 362, 376, 415, 471, 472. 2. 279, 428.
15. MACROPUS ROBUSTUS ERUBESCENS Sclat.
3. 346, 369, 379, 403, 431. 2. 338, 368, 390, 474, 488.
16. MAcRopUSs UALABATUS INGRAMI, subsp. noy. (Plate XLII.)
@. 425, 465, 468, 469. Inkerman.
Most nearly moe to MW. uw. apicalis Giinth., but smaller and
much greyer in colour.
General colour of back light buffy grey, the buff colour
becoming more dominant towards the posterior part of the body.
Under surface of body grey, washed over with a rich tawny-buff
tint. Head similar in colour to J/. uv. apicalis, but lighter in the
light parts. Prominent light buff-colouved areas round the bases
of the ears, continuous with the lateral face-stripes and practically
meeting on the crown. Backs of ears black, with hght margins.
Forehead with an indistinct median black stripe. Fore limbs
light buffy grey, the light-coloured shoulders contrasting strongly
with the dark str ipes behind them. Hands and feet black. Tail,
for the greater part of its length black, basal part greyish and
tip white.
Skull very much smaller than that of JZ. w. apicalis, and with a
less elongated nasal region. Palate narrower and teeth smaller.
Dimensions of the type (Gens ed in the flesh) :—
Head and body 630 mm.: tail 640; hind foot 195: ear 78.
Skull—hasal length 105 mm. ; condylo-basal length 111;
zygomatic breadth 60; nasals, length 48, greatest breadth 198;
central breadth 12, consti iction it 3 : palate length 66; diastema 23 ;
tooth-row from p' to m’® 36:5; length of secator (prego
length of three anterior molar form teeth 21.
The skull of an old female with worn teeth has a basal leneth
of 107 mm.
Type. Sub-adult female. B.M. No. 8.8.8.65. Original number
468. Collected 17th October. 1907.
This Wallaby would appear to be intermediate between the
1908. | MAMMALS FROM NORTH QUEENSLAND. 793
northern J/. uw. apicalis Giinth. from Cape Grafton, and the
southern JZ. walabatus of New South Wales and Victoria, but is
paler and smalier than either.
We have named this handsome animal in honour of Sir
William Ingram, to whose initiative and generosity the acquisi-
tion of the collection is due.
17. Macropus parryi Benn.
g. 285, 334, 341, 388, 389, 395, 470, 478.
Q. 370, 399, 477, 480.
18. Macropus aqiiis Gould.
3S. 270, 271, 274, 290, 292, 293, 294, 392, 421, 429, 508, 511.
©. 280, 291, 346, 372, 380, 418, 424.
©. 512. Mt. Hlliot, near Townsville.
19. PETROGALE ASSIMILIS Ramsay.
6. 275, 232, 286, 335, 373, 430, 473, 482.
Q. 272, 281, 287, 307, 310, 333, 336, 385, 455, 456, 466, 483
484.
This series of skins indicates that P. assimilis of Ramsay *
should be recognised as distinct from P. penicillata Gray, with
which it was doubtfully united by Thomas yt, who had at that
date no specimens of it for examination.
20, LAGORCHESTES CONSPICILLATUS PALLIDIOR, Subsp. nov.
3.416. @. 413, 432.
Most nearly allied to Z. c. leichardti Gould, but differing in
being much lighter in colour. The general tawny colour of the
upper surface of L. ¢. leichardti is represented by a light fawn-
colour, the difference beg very evident on the sides and
posterior half of the back.
Collett has already pointed out £ the existence of these light-
coloured Queensland specimens, and it would appear, taking into
consideration the type locality of Z. ¢. leichardti§, that this
Eastern form is deserving of subspecific rank.
Dimensions of the type (measured in the flesh) :—
Head and body 470 mm. ; tail 450; hind foot 156; ear 50.
Skull—hasal length 70 mm.; zygomatic breadth 47°55; nasals,
length 28, greatest breadth 17, least breadth 13, constriction
breadth 13°4; palate, length 44°7, breadth outside m* 24-2,
breadth inside m' 15; palatal foramen 5:2; diastema 9; length
of upper cheek-teeth 28.
Type. Adult female. B.M. No. 8.8.8.104. Original number
A413. Collected 30th July, 1907.
* Ramsay, Proc. Linn. Soc. N.S.W. i. 1876, p. 360; u1. 1877, p. 11.
+ Cat. Mars. B. M. p. 67, 1888.
+ Zool. Jahrb. ii. p. 889, 1887.
§ The type locality of D. c. leichardti is the “country between Port Essington
and the Gulf of Carpentaria.”
794 ON MAMMALS FROM NORIH QUEENSLAND. iN ov. 3,.
21. AXPYPRYMNUS RUFESCENS Gray.
¢. 510. Elhot R., Townsville.
22. 'TRICHOSURUS VULPECULA Kerr.
6. 253, 256, 258, 264, 350, 358, 375, 387. 2. 268, 277,
401, 436.
3. 476. Mt. Abbot.
Some of these specimens exhibit a distinct tawny coloration,
such as is found to a greater degree in the more northern form
described by Ramsay from the Bellender Ker Mts., N. Queens-
land, under the name of Phalangista johnstonii *.
23. PHASCOLARCTUS CINEREUS Goldf.
Skull (2).
My. Stalker informs us that this is the most northern point at.
which the Koala occurs.
24. Iscopon rorosus Rams.
3. 280, 318, 397, 404, 405, 412, 437. 9. 331,345, 382, 439.
One female in spirit.
These specimens show that Ramsay’s Perameles torosus +,
described from Rockingham Bay, N. Queensland, should stand as
a species distinct from /. macrourus Gould, with which it was
united in the Catalogue of Marsupials.
All of them are much larger and more heavily built than the
type of 7. macrourus, a character that is very evident in the size
of the skulls. The following are the average skull dimensions of
6 adult males, compared with the type skull, which is also that of
an adult male.
I. torosus. I. macrourus.
Greeaibestalemeblt aeesrenen cr. 88 mm. 7o°5 mm.
Zygomatic breadth............ 40 37
Length of upper tooth-row | 47 43
TROON ates WO)” » sa eawee aways J
25. DasyurRUSs HALLUCATUS Gould.
Gr. 828) 348, ol 384. | Oe aii, 332,.and mos oaraue
6. 485. Mt. Abbot.
In No. 7.9.15.36 the posterior half of the body is wholly devoid
of the characteristic white spots.
26. TACHYGLOSSUS ACULEATUS Shaw.
So Ds: Oe Al, SIAL
Represents Collett’s Echidna acanthion, described from west of
Rockhampton.
* Proc, Linn. Soc. N.S.W. 2nd ser. iii. p. 1297, 1888.
+ Proc. Linn. Soc. N.S.W. ii. p. 12, 1877.
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1908. ] ON SZE-CHUEN AND BHUTAN TAKINS. 795
3. The Sze-chuen and Bhutan Takins.
By R. Lyprexxer.
(Received September 2, 1908. ]
(Plate XLIII.* and Text-figures 168-171.)
The first recognition of a Takin distinct from the typical
-Budorcas taxicolor of the Mishmi Hills is due to the late
Professor A. Milne-Edwards, who in 1874 (‘ Recherches pour
sservir 4 Histoire naturelle des Mammiferes,’ p. 367, pls. Ixxiv. &
Ixxix.f) described and figured a representative of the species from
Moupin under the name of Budorcas taxicolor, var. tibetanus.
Here I may take the opportunity of mentioning that Moupin is
stated in all zoological works that have come under my notice to
be in Eastern Tibet. As a matter of fact, it is, as pointed out
to me by my friend Mr. Thomas, situated in Sze-chuen; a
circumstance which clears up a number of difficulties and mis-
conceptions with regard to the range of the animals of this part
of Central Asia. The name Budorcas sinensis has been applied
to the Takin of Kansu, which, as shown by a specimen in the
Tring Museum, is inseparable from the Sze-chuen animal; the
authority for the name I am, however, unable to find.
Milne-Edwards described the male of the Sze-chuen Takin as a
_yellowish-red animal; and also stated that the female is paler
and greyer. Neither his description nor his plate of the male is,
however, satisfactory ; and as mounted specimens of both male
-and female are now exhibited in the public galleries of the British
Museum (Natural History), I consider that they should be figured
before their colouring is deteriorated by exposure.
The male specimen (Plate XLII. fig. 1), which was stated to
-come from Sze-chuen, was purchased by the Trustees of the
British Museum from Rowland Ward Ltd. in 1905. The female
(Plate X LITT. fig. 2), on the other hand, was given by Mr. Mason
Mitchell, of the American Consular Service in Sze-chuen, to
Mr. Rowland Ward in 1908, by whom it was, in turn, presented
to the British Museum. in noticing the presentation of the latter
“Specimen in the ‘ Field’ newspaper (vol. exi. p. 790, 1908), I stated
that the presumption was that it represented the cow of the race
described by Milne-Edwards. The original sender of the specimen
-stated, however, in a letter to Mr. Ward that there are two
distinct kinds of Takin in Sze-chuen, differing not only in colour,
but also in siz2 and in habits; the smaller red kind—known to
the Chinese as yea-nw (wild ox)—associating in small herds, while
the larger grey one—the twnu-yea (big wild ox) of the Chinese—-
-goes about singly, or at most in pairs. I accordingly suggested
that if this statement were borne out by the facts, the larger grey
race might be named Ludorcas taxicolor mitchelli.
When, however, the grey female was mounted and_ placed
* For explanation of the Plate see p. 802.
+ The work was published in parts from 1868 to 1874; as I do not know the dates
of publication of the separate parts, I have given the date of completion of the work.
796 MR. R. LYDEKKER ON [Nov. 3,
beside the yellow male (with which it is approximately equal in
size, although its horns are considerably smaller), I could not
entertain any doubt as to the two representing the different sexes
of one and the same animal. Whatever, therefore, may be the
truth with regard to the Chinese story, I cannot but regard
the two specimens in the Museum as severally representing the
male and female of the Ludorcas taxicolor var. tibetanus of
Milne-Edwards. Both specimens are subadult animals.
Text-fig. 168.
Imperfect skull and horns of adult male of the Mishmi Takin.
The distinctive characteristics of the Mishmi and the Sze-chuen
Takins (which I now consider worthy of specific separation) will per-
haps be made most readily apparent by the following comparison :—
1. Buporcas TAxtcoLor.—Size large, although precise shoulder-
1908. } SZE-CHUEN AND BHUTAN TAKINS. CMe
height not ascertainable, owing to the bad mounting of the
British Museum specimen. General colour (apparently in both
sexes) of upper-parts cigar-brown, with an elongated tawny
“saddle” on the back, becoming much darker on the under-
parts, and passing into deep blackish brown on the limbs; the
ears and the whole of the head in advance of the same, together
with the entire under surface of the lower jaw, and a dorsal
stripe extending from the occiput to the root of the tail, black.
No distinct beard in male; and tail apparently not distinctly
tufted, and brown in colour.
Horns (text-fig. 168) stout, elevated into a strong, oblique,
prominent, longitudinal ridge at the base, with the long smooth
tips situated in a plane different from that of the basal portion,
and generally directed (when fully adult) almost straight upwards,
but inclining somewhat backwards at the extreme tips.
2. BuDORCAS TIBETANUS.—Size probably equal to that of the
last; height at shoulder of subadult male 404 inches. General
colour of upper-parts of subadult animals in winter coat orange
or grey, strikingly different in the two sexes; dorsal stripe not
extending further forwards than the withers ; under-parts lighter
than back ; black on head confined to the backs of the ears, a ring
round each eye, the front of the face in advance of the eyes, and
the extreme tip of the inferior surface of the lower jaw, thus.
forming a striking contrast to the light area. <A distinct beard
on the throat of the male; tail strongly tufted, and blackish in
colour.
Horns (text-fig. 169, p. 798) more slender than in tawicolor,
with much less development of the oblique basal prominence, and
the long tips in the same plane as the basal portion; these
tips inclining somewhat inwards and also decidedly backwards
throughout their length.
In the male, the whole of the fore-quarters, exclusive of the
black areas, bright golden-yellow, gradually becoming more and
more grey posteriorly till it passes on the hind-quarters into
grizzled grey, which is continued on to the limbs, and also forms
a vertically elongated patch on the lower part of the shoulder.
Dorsal stripe extending as far forwards as the withers.
In the female (which, as in the type species, has much smaller
horns) the yellow on the fore-quarters of the male replaced by
dirty white; and the limbs a darker grey, becoming nearly black
on the knees and hocks. Dorsal stripe not extending further
forwards than the middle of the back.
These differences are, in my opinion, amply sufficient to justify
the recognition of the Sze-chuen Takin as a species distinct from
the typical Mishmi animal and its smaller Bhutan representative.
With its bright golden-yellow or white fore-quarters, contrasting
strongly with its black face, ears, and eye-rings, the former is
indeed a much more strikingly coloured animal than the latter,—
a feature in which it agrees with several of the other Sze-chuen
mammals.
798 MR. R, LYDEKKER ON [ Nov. 3,
My only doubt is whether the name tibetanus, as being some-
what misleading, ought not to give place to sinensis. On the
other hand, it is possible that the species may cross the border
dividing Sze-chuen from Tibet.
Text-fig. 169.
Skull and horns of old male of the Sze-chuen Takin, collected by Mr. J. W. Brooke
Coming to the Bhutan Takin, which I described in the ‘ Field’
for 1907 (vol. ex. p. 887) as a small local race of the typical
species under the name of Budorcas taxicolor whitei, giving also a
preliminary notice in the Society’s ‘ Proceedings’ for the same
1908. | SZE-CHUEN AND BHUTAN TAKINS. 799
year (p. 749), I regret to say that the two skins there referred to
as being in the possession of the Hon. Walter Rothschild have
Text-fig. 170.
Skull and horns of old male of the Bhutan Takin.
gone elsewhere than to the Tring Museum, and are therefore
unavailable for fuller description and figuring. I must therefore
Proc. Zoot. Soc.—1908, No. LI. Dt
800 MR. R. LYDEKKER ON [ Nov. 3
rely mainly on the two pairs of horns (those of an old bull and of
a subadult female) presented to the British Museum by Mr. J.
Claude White, C.M.Z.S., British Commissioner in Sikhim, which
form the type of this race.
Before proceeding further, it may, however, be well to mention
that the Bhutan Takin, which lives high up on the mountains on
the open zone between the upper limit of forest and the snow-
line, is completely cut off by deep river-gorges from the habitat
of its Mishmi representatative. On this ground alone it,
therefore, undoubtedly belongs to a separate race.
As to the height attained by this race, I cannot, in the absence
of a mounted skin, give even an approximate estimate; but I
believe it to be somewhat smaller than the typical Ludorcas
taxicolor, although its chief claim to distinction rests on the
relatively small size of its horns.
The type horns of the old bull are somewhat worn at their
tips, as they also are at the base, where the rugosities have in
consequence disappeared. When entire, they would probably
have measured about 15 or 16 inches in length along the front
curve, as against from 20 to 24 inches in adult bulls of tke
Mishmi Takin. The horns of a younger bull, associated with
one of the skins referred to above, measured 13 inches, against
18 inches in horns of a Mishmi bull of about the same age.
Horns of cows are much smaller. Similar characters are shown
by the horns of the skull here figured (text-fig. 170, p. 799), which
was sent to the Museum by Mr. White.
I have been hitherto unable to institute an exact comparison
between the skins of the Mishmi and the Bhutan Takin, but I
think the latter has a somewhat smaller lighter dorsal saddle than
the former. Both agree in their general dark colour, and in the
under-parts being darker than the back, as well as in the large
extent of black on the head, and the extension of the dorsal stripe
to the occiput.
In this place it may be mentioned that the skull of a large
ruminant from the Phocene of the Siwalik Hills, N.E. India,
described by the late Professor L. Riitimeyer (by whom it was
regarded as probably indicating a hornless species) as Bucapra
daviest, appears to represent an animal nearly related to the
Takin. Certain details in the form of the skull, as well as of
the teeth, seem, however, to indicate the generic distinctness of
Bucapra from Budoreas. I have little doubt that, when entire,
the Siwalik skull carried horn-cores of a type approximating
more or less closely to those of the Takin.
Since the above was written the Museum has received from
Mr. J. W. Brooke, skins, skeletons, and skulls of Takin of both
sexes and of several ages from Sze-chuen, where the animals were
killed in rhododendron and bamboo jungle at an elevation of
about 10,000 feet. The skulls and skins include those of a fully
1908. | SZE-CHUEN AND BHUTAN TAKINS, 801
adult bull and cow; the bull is larger and darker than the figured
specimen. The skull represented in text-fig. 169 (p. 798) belongs
to the bull obtained by Mr. Brooke.
Special interest attaches to the skull of a very young Takin
(text-fig. 171), with the three pairs of milk-molars just about to
cut the gum, and no traces of horns whatever. Unfortunately no
‘skulls of Serow or Musk-Ox of corresponding age are available
for comparison, so that it is impossible at present to get much
further in the matter of the Takin’s relationships.
Text-fig. 171.
Skull of very young Sze-chuen Takin, collected by Mr. Brooke.
Compared with that of a somewhat older Serow, in which horns
are developed and the first true molars in use, the skull of the
baby Takin is mainly distinguished by the great elevation of
the frontal region, and the shortness and breadth of the nasal
bones,—characters which become greatly exaggerated in the
adult. The premaxillee are still more widely sundered from the
nasals than is the case in Serows; and above the supraoccipital,
wedged in between the hind portions of the parietals, is a large
undivided interparietal, which I believe to be represented in the
Serow skull. Allowing for the greater breadth of that of the
Takin, the two skulls seem in other respects to present a great
general similarity.
[ dddendwm.—Since the paper was read I have received a letter
pile
802 MR. R. LYDEKKER ON AN [ Nov. 3,
from Mr. Brooke, in which it is stated that old males of the:
Sze-chuen Takin grow to a very large size, as, indeed, is indicated
by the skin and skull sent to the Museum. Also, that in summer
the long and rough orange or reddish coat is replaced by one of
short greyish hair. |
EXPLANATION OF PLATE XLII.
Subadult male (1) and female (2) of the Sze-chuen Takin (Budoreas tibetanus);
in winter coat, from the mounted specimens in the British Museum.
4. On an Indian Dolphin and Porpoise.
By R. LypEKKER.
[ Received September 2, 1908.)
(Plates XLIV. & XLV.*)
Once more I am indebted to the Director—Lieut.-Colonel F. W.
Dawson—of the Trevandrum Museum for sketches and measure-
ments of certain Cetaceans recently captured on the Travancore
coast. These are represented by three specimens, referable to two
species; they differ to a greater or less degree from the typical
forms of all the Dolphins and Porpoises hitherto described from
Indian waters, and from the world generally. The most remark-
able fact about the new specimens is that two of them are Bottle-
nosed Dolphins, referable to Z'ursiops, of which genus, in addition
to the typical 7’. twsio, I have already recognised (Proce. Zool. Soc.
1905, vol. i. pp. 125-128) three, if not indeed four, Indian
species.
Col. Dawson informs me that the, two examples of this.
apparently new Bottle-nose were caught by fishermen about six
miles to the north of Trevandrum, in the spring of the present.
year (1908). The skeletons of both were preserved ; and one of
these has, at my request, been presented by Col. Dawson to the
British Museum, as it seemed desirable that an apparently new
form should be represented in the chief English collection.
The following particulars concerning these two specimens.
(A and £B) have been supplied to me from Trevandrum :—
Ale B.
ittgen abi ft2, ame
Length from tip of snout to cleft in flukes .... 9 0 9 4
Dox tosorigamyot dorsal fim 25... 0:5 acs eeeaee eee 4 0 4 2.
‘yf UU PPORR ys ies snaBt ete. serwesichpines < seem 2 0 Leal
yc Pemba SrOOVE ..5. «aw adsaue. ts eee dD 6 Dia
5. 6) AA SLOOVE: oiisch.canistemsioys sade - eee eee 6 6 6 7
Leneth ofjdorsal fim 9 .0.j.-4-s0-cee- + eee 16 Lig
le ee lt vil alee
Expanse of flukes: "200. 3.0..0..- 0: - see eee Ze Ih 2 tty
* For explanation of the Plates see p. 808.
‘dS VNWOIOHAGOAN 2 ‘INOSMVG SdOISHAL
Tyr
ATIX Id @06T S Za
PAS (OVINUAS) ACME INTL WET WAL CONS)
Ta eS
SR
INISE Wal SONS Sp 74 val
1908. } INDIAN DOLPHIN AND PORPOISE. 803
A, Be
Length of beak from groove which seid itied foe ft. in.
MERON MEA astailcstist cist, tae atetieeh. abtse Aye oe | Ai 5
Hength of genital groove s..4essdesceaeecee oo if 8
i small SrOOVE tobe aa: dt esterases neh oe 2 3
Cre ho mma bne iene eae. oc Mo tal dk sc cde eee fail Lad
Greatest breadth of body ................000eeee ly lly).6 1,8
Pe geehero bro DOG s 24! sety. cep. cee roen os re eens Lis
» eircumference in front of dorsal fin .| 3 10 4 0
Smallest circumference at root of tail ......... 11 12
Pere nt Oi Wonca fi .5...2sr0.i2. «calc cts Dede oe sox oe ee net [reas 23
Length from snout to blow-hole ............... is Teac) Lys
As regards shape, the body is rather elongate, with a promi-
nent ridge extending from the back of the dorsal fin to the
middle of the flukes. Both the flippers and the dorsal fin are
distinctly faleate. The eyelids are well developed and somewhat
mobile ; while the blow-hole is, as usual, placed somewhat to the
left of the middle line. In front of the blow-hole is a fatty
cushion, marked off from the moderately tapering beak by an
ill-defined Y-shaped groove. The lower jaw projects somewhat
in advance of its fellow.
The colour of the two specimens is described as follows :—
A.—Upper-parts deep glistening black, becoming somewhat
lighter below, with a pinkish tinge round the anal and genital
apertures ; under side of lower jaw and muzzle dull white.
£.—Above deep glistening plumbeous black, abruptly passing
into pale slaty on the sides; genital and anal regions lighter ;
lips dull white.
The teeth, which are relatively large, with rugose crowns,
number :
ua) 26 :
i and 5 = 49 and 51 in 4, and
on 55
= and 2 = 47 in B.
The vertebre, of which the first two are in each case fused
together, number :
Co Dale ie 20: Can 25 —- 04. imi a and
ORC De 2 igh, Cae 28) —) Al imteb.
The first four pairs of ribs are two-headed.
The pterygoids are in contact, and the mandibular symphysis
1s short.
The phalanges number :
ee Oe 7 PVE ese Vee, indy, ance
es, EE OF 7, HV So Vik oe Ee
Both specimens evidently belong to the same species, and from
the general contour of the head, body, fin, and flippers, coupled
804 MR. R. LYDEKKER ON AN ' Nov. 3,.
with the number of vertebre and teeth, the large size of the
latter, the presence of four pairs of double-headed ribs, and the
approximation of the pterygoids, there can be little hesitation in
referring that species to the genus 7ursiops.
As regards comparison, I think it will simplify matters to.
reproduce, with some slight modification, the synopsis of the
species of the genus given in my above-mentioned paper in the
Society’s ‘ Proceedings’ for 1905 :—
1. Tursiops tursio.
a
Type specimen: Teeth 5, = 44.
Vertobre: C.% Dales llr On. 27 of
Pterygoids in contact.
Tursiops abusalan.
Type specimen : Teeth = as
Wertebree > (©. (2) 12> ilo e222 on — no
Pterygodids (8) 3 in contact.
~ [ES
9)
Indian specimen: Teeth 5= andi Fe = 54 and 53.
WVertebre 2 Cl 7, Delain tin Ca Jor — 60)
Pterygoids divergent.
3. Tursiops catalania.
Type specimen: Teeth OF
Vertebre : C. 7, D. 12, ii los Ca, 24 —= 98.
Pterygoids diver, gent (2).
fet . 25
Indian specimen *: Teeth 55 and 5 = = 50 and 51.
Womans Cy G4. 1D), Ia I, Li, Ca. 24 = 61.
Pterygoids divergent.
4. Tursiops parvimanus.
25
teeth >, — 49.
a4
Vertebree = 62.
5. Tursiops gilli.
F 22 23
Type specimen: Teeth oo and 55 = 44 and 45.
Vertebree (?).
Indian specimen : ee me nd ; = = 55 and 53, or (in
D2
young) 5 = and 5 22 =e land 50.
Vertebre : c os D. 12, L. 16, Ca. 23 = 58.
Pterygoids divergent.
If the foregoing identifications be correct, we shall have the
* Tursiops fergusoni, Lydekker.
1908. ] INDIAN DOLPHIN AND PORPOISE. 805
following external characters of the four definable species of
Tursiops included in the above list :—
1. Tursiops tursio. Kuropean Seas.
Size large: 9 ft. 6 in.
Upper surface blackish.
Under-parts white and unspotted.
bo
. Tursiops abusalam. Red Sea and Indian Ocean.
Size smaller: 7 ft. 24 in. (type), 6 ft. 11 in. (India).
Upper surface dark greenish.
Under-parts whitish and spotted with green in adult;
whitish in young.
(Sv)
. Tursiops catalania. N, Australia to Indian Ocean.
Syn. (?) 7’. ferguson.
Size about the same as last: 7 ft. 8 in. (type), 7 ft. 43 in.
(India).
Upper surface dark slate.
Under-parts yellowish *, flecked with lead-colour.
Tursiops gilli. N. Pacific to Indian Ocean.
Size, Indian specimen, 6 ft. 8 in.
Whole surface blackish, tending to lighten slightly on the
under-parts, with a tinge of reddish in Indian specimens.
In addition to the above, Mr. F. Lahillet has described
(without reference to my paper) a Bottle-nosed Dolphin from the
La Plata estuary under the name of Tursiops gephyreus, of which
the leading characteristics are as follows :—
Teeth 55 = 45.
Vertebre: OC. 7, D. 13, L. 17, Ca. 24 = 61.
Pterygoids divergent.
Phalanses: 0 15 Ul. 7; 110.6, UV. 2) V.1.
Size, large, about 7 ft. 2 inches (276 cm.).
General colour leaden grey, becoming somewhat lighter on the
under-parts ; three or four reddish circles on the sides in advance
of the vent.
Mr. Lahille considers his Bottle-nose as nearly allied to
T. catalania, of which it may indeed be only a large race. In
addition to its size and colouring, and slight differences in the
number of the teeth and vertebre, it is distinguished by its nar-
rower beak and premavxille and much broader temporal region.
That the new Indian Bottle-nose (Pl. XLIV. fig. 1) is quite
distinct from 7’. tursio, 7. catalania, and T. abusalam, in all of
which the under-parts are light-coloured, is certain. In general
colour it agrees much more closely with the Travancore specimen
>
* In the type the under-parts are described as whitish ; if the orange tint of the
Trevandrum specimens is a specific character, then the name 7. fergusoni will be
available for the Indian form.
+ An. Mus. Nac. Buenos Aires, ser. 3, vol. ix. p. 347, 1908.
806 MR. R. LYDEKKER ON AN
[ Nov. 3,
provisionally referred to 7’. gilli (Proc. Zool. Soc. 1905, vol. 1
pl. xii. fig. 1), but the under-parts are hghter, and there appear
to be slight differences in the form of the beak and dorsal fin.
These might be considered individual variations; but the differ-
ence in the number of the vertebre is so great, while the relations
of the pterygoids are also different, that I cannot refer the two
specimens to the same species. From 7’. yephyreus the Travancore
Bottle-nose differs, among other features, conspicuously in regard
to the number of joints in the flippers. ‘There accordingly seems
no other course but to regard the new Trevandrum Bottle-nose as
an undescribed species
a view in which I am supported by the
taxidermist at the Trevandrum Museum, who has had under his
hands all the Travancore Cetaceans described by myself.
I there-
fore propose the name Z'ursiops dawsoni for the new specimens,
taking the skeleton in the British Museum as the type.
The second species to which I have to refer on the present
occasion 1s a representative of the Finless Porpoises, Veophocena
(Neomeris), taken by fishermen off Trevandrum in June last and
purchased by the local museum. While agreeing in all general
respects with the typical Neophocena phocenoides, this specimen
(PI. XLIV. fig. 2) differs by the circumstance that the purplish-red
patches on the lips and throat are replaced by pale grey areas of
corresponding shape; while there are likewise numerous irregu-
larly disposed, narrow, lead-coloured streaks on the under surface
of the lower jaw, not noticed in descriptions of the ordinary form.
The general colour is uniform dark plumbeous, becoming gradually
paler on the flanks and under-parts. The teeth are
= 39 and 41. The vertebre number ©. 7, D. 14, L. 12
20 12 ae
ig *24 5H
Ca. 26
= 59; and of the fourteen pairs of ribs, seven are double-headed.
Bee as phocenoides is generally described as having
about | aa 5 (=3 5) teeth, while its vertebral formula is given as
Cane D. 12, 13, L.4+Ca. 38-43 = 57, 58, or (maximum) 63,
nothing of any decisive importance can be inferred in these
respects with regard to the new specimen, which, for the present,
at any rate, I prefer to leave unnamed.
It may be useful to publish the following dimensions of this
specimen, as supplied from Trevandrum :—
Length from tip of snout to notch in the tail .
Do. to origin of flipper, right side ............
es 3 w left ‘side™-.4; ars.
oo Benitalls See A sn A eNO Eee ae
SF) oe ZEN © OSAMU se scrooganas025 9. EE, 2
Length of flippers, right side ..................
+ left*sider sie. ee aes
Expanse On tall rare ce ee oscil aie caesar otra a
Greatest circumference of body ...............
Smallest Us a at root of tail...
Genital onooverse ss ere ener SH a ae A. 5
bo bo
in.
)
ih
102
1908. ] INDIAN DOLPHIN AND PORPOISE. 807
Mammary groove half-an-inch in length; situated on the
posterior half on each side of the genital groove, which is placed
on a ridge 6 inches long, marked off from the abdominal region
and gradually widening towards the anal opening; ear- -hole
minute.
Since the foregoing was written [ have received from the
Director of the Trevandrum Museum a sketch and description of
another Dolphin purchased from the local fishermen on the
eleventh of August 1908.
Of this specimen: (Pl. XLV.) the following particulars have
been sent me :—
is tis
Extreme length from tip of snout to notch on the tail-
flukes
From tip of snout to the basal angle of the prenarial
PNGLTDOR Ss CLO TENUKOIINI See ee geese me Sn ar OREN ne
Prominipron snout boanele Of mou .....s.2e5.2 ee acess
Wowio, anterior amele Of Lhe CYC). ar. i. ns- a ctecec saucy “ee
MMM ONIN OL Mase yates be laieiaty- «Sei ceescyinn cist Ra RER Seta ee
“ flippers RP ee ee ee ae tas oy asi cfagnncl tee eg
J OUTG BON Oil 6KCTGSP TN Tae AN ole See ee oer a Leni iatent
e anterior commissure of genital grove ............
NiGIOUG: PNG ASRS aoe ee Re ee Aan a un rapa ee
Length of front margin of flipper along the curve ......
- Bs ‘ dovsal fin HRA STEEN THROES
Hero lnba@ he, CONS ATM cos. saa ca sn ccasisseecncreveesceeestr ures a:
Greatest girth of body in front of dorsal fin ...............
=a mavelhoclisaves @lorsseulistBay Ba soct otdtion sir
Smallest Circumuerencerat LOOU Ok ball) Qija, yee neces
AB aA SEs Op NUE SE 02 ncteials 5 sig 9 -Beip elie vioyresaid02 morefoajesyontelys
yj
bo
WIR
fk TE RS) [NO Tp
i
De
KFHONWONNACWOORWwWwMO
Doe oS
Hye 11 in. longest diameter ; ear vertically ovate, length 3 mm. ;
genital groove 8 in.; vent 21 in. in diameter; between genital
and anal openings 8 in.
A prenarial adipose elevation marked off from the tapering
snout by a U-shaped, ill-defined groove. Body fusiform, much
compressed towards the tail, the prominent backward extension
of the dorsal fin gradually shelving off into the caudal ridge, which
is continued along the corresponding line of the under surface.
Flippers feebly falcate.
The colour is uniform pale plumbeous, washed with pale brown
and becoming lighter towards the under side; the body being
profusely flecked with long pear-shaped pale pinkish and dark
plumbeous markings of varying sizes. Angle of mouth, margin
of upper lip, and top of snout mottled with whitish ; lower lip
creamy white mottled with brown ; belly mottled with white, less
so on the pectoral region. There are also groups of milk-white
parallel striations in different parts of the body and on the
flippers and flukes.
3 36
meni 2 =e
36 X 36 = 144; moderate, conical, the tips curving
808 ON AN INDIAN DOLPHIN AND PORPOISE. [ Nov. 3,.
inwards and enamelled, and the sides compressed with the base-
expanded. Pterygoids narrow, separated from each other in the
middle line, with their inner border divergent posteriorly.
Palatines somewhat W-shaped, with the median suture extend-
ing nearly halfway between the widely separated pterygoids.
Ribs 12 pairs; six pairs double-headed.
Vertebrie= C7, 1). 12 9 Ca. 25) — Ol he torch moe
cervical vertebre are united, but the epiphyses are not fused with
the centra of the vertebree. Symphysis of the mandible one-third
the length of the ramus.
Bhalanges 1) bl) (4 Lo Vers
There can be no doubt from this description and the figure that.
this Dolphin is referable to the genus Sodalia, as redefined by
myself in the ‘ Journal of the Bombay Natural History Society,’
vol. xv. pp. 412 & 413, 1903*. It is equally evident that it 1s
identical with the Speckled Dolphin (Sotalia lentiginosa ~); and
this being so, it is apparent that the young Trevandrum Dolphin.
named and figured by myself in the paper just cited as Sotalia
Jergusont (pl. D) cannot (as I suggested might prove to be the
case) be separated from S. lentiginosa, the absence of spotting
being a feature of immaturity.
The present specimen renders necessary the following slightly’
amended definition of Sotalia :—
Teeth medium, smooth, and numerous ees D).
Pterygoids separate.
Palatines W-shaped, with a long symphysis below the.
pterygoids.,
Vertebrze 49-55.
In SN. lentigiwosa the number of the teeth in the adult may now
—— = 68 or 72; and that of the vertebre as
C7, Ded) or) 12, 10 or79; Car 21) or 23 = 49) orb tara
least this is, I think, the best way of hens bares the vertebral
formula of the present specimen with that of the type of
S. fergusont.
be given as
EXPLANATION OF THE PLATES.
PratE XLIV.
Wig. 1. Tursiops dawsoni. +5 nat. size.
Fig. 2. Neophocena sp. About + nat. size.
Both from the neighbourhood of Trevyandrum.
Prate XLV.
Sotalia lentiginosa.
From the neighbourhood of Trevandrum.
* In the figure on p. 414 the lettering pal. and pt. should be transposed..
t= Steno | lentiginosus Blanford.
1908. ] ON TROPICAL FLIES CAUGHT IN MANCHESTER. 809
November 17, 1908.
Prof. EK. A. Mincuty, M.A., Vice-President, in the Chair.
The Secretary read the following report on the additions:
made to the Society's Menagerie during the month of October
1908 :—
The number of registered additions to the Society’s Menagerie:
during the month of October was 131. Of these 38 were acquired
by presentation, 5 by purchase, 83 were received on deposit, and
5 were born in the Gardens. The number of departures during
the same period, by death and removals, was 186.
Among the additions special attention may be directed to :—
One Vicuna (Lama vicugna), from South America, presented
by T. Rome, Esq., on October 16th.
A Collection of 26 Birds of Paradise, including 5 Count Raggi’s.
Birds of Paradise (Puradisea raggiana), 7 Lawes’ Birds of Para-
dise (Parotia lawesi), 10 Hunstein’s Birds of Paradise (Diphyllodes
hunsteini), 1 New Guinea Rifle-bird (Péilorhis intercedens), 2 Violet
Manucodes (Phonygama purpureo-violacea), and 1 Prince Rudolph’s
Bird of Paradise (Paradisornis rudolphi), never previously im-
ported, from S.E. New Guinea, deposited by Sir William Ingram,.
Bt., #.Z.5., on October 5th.
A Collection of Mammals, Reptiles, and Birds, including a
Black-and-White Sparrow-Hawk (Accipiter melanolewcus), from
S. Nigeria, presented by Dr. W. F. Macfarlane, F.Z.8., on
October 50th.
Mr. E. E. Austen, F.Z.S8., exhibited living specimens of a Fly,
Hermetia illucens L., caught in Manchester and received from
Dr. W. E. Hoyle. Since larvee of this fly (a native of Tropical
South America, Central America, and the West Indies, breeding’
in decaying vegetable matter) had previously been found in
Liverpool in cargoes of raw rubber from the Amazons, it was:
thought that these specimens might possibly have been introduced
into Manchester in a similar way.
The Reports on the Ruwenzori Expedition Collections were:
communicated to the meeting, aud will be published entire in.
the ‘ Transactions.’
810 PROF. G. C. BOURNE ON THE | Nov. 17,
The following papers were read :—
1 Contributions to the Morphology of the Group Neritacea
of Aspidobranch Gastropods.—Part I. The Weriicde.
By Prof. Gitzert C. Bourne, D.Sce., F.Z8.
[Received October 27, 1908.
(Plates XLVI.-LXVI.* and Text-figure 172.)
While the Haliotide, Fissurellide, Pleurotomaride, Trochide,
Patellide, and other members of the Aspidobranch Gastropoda
have received a large amount of attention from morphologists,
the Neritacea have, until recent years, attracted little interest.
The anatomy of so familiar a species as the European Weritina
Auviatilis was imperfectly known till the appearance of Lenssen’s
memoirs in 1899 (25) and 1903 (26). ‘Thiele’s (39) short but
accurate descriptions of various organs of tropical species of
Neritidee have added largely to our knowledge of the group, but
even when these are taken into consideration it can hardly be
said that a sufficiently comprehensive comparative account of the
Neritide exists in a form available for students of molluscan
anatomy.
It has been too readily assumed that the Neritacea, forming, as
they do, an extremely specialized section of the Rhipidoglossa,
are unlikely to retain any considerable traces of primitive
organization, or to yield evidence bearing on the ancestry of the
Gastropoda. Thus Pelseneer (30) writes: ‘ D’autre part les
Neéritacés sont plus spécialisés que tous les autres Rhipidoglosses
(Haliotis ete.) ou la commissure viscérale est déja croisée, par :
1°. Lexistence d’une seule branchie et d'un seul osphradium.
2°. Liexistence d’un seul rein. 3°. Liexistence d’un orifice
genital propre. 4°. L’existence d’yeux a cavité fermée. 5°. La
séparation plus compléte des ganglions pleuraux et pedieux.”
There is no doubt that the Neritacea are specialized in these
respects, but this is no reason for regarding them as probably
uninteresting subjects for anatomical study, for animals highly
specialized in some respects may, nevertheless, retain many
primitive features, and there are so many points in which the
Neritacea seem to approximate to the Pectinibranchs, e.g. the
existence of a single kidney with a slit-shaped opening into the
mantle-cavity, the complex genital ducts with accessory glands,
&e., that it has long been a matter of interest to determine
whether they are intermediate between the less specialized
Aspidobranchia and the Pectinibranchia, or whether their
apparent resemblances to the latter group are due to convergence.
Finally, the Helicinide, interesting because they are terrestrial
and pulmonate, have not been the subject of any comprehensive
* Kor explanation of the Plates see p. 884.
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1908. | ASPIDOBRANCH GASTROPOD MOLLUSCS. 811:
anatomical memoir since Isenkrahe published an incomplete
account of their structure in 1867, and it is high time that this
gap in our knowledge of Gastropodan anatomy should be filled
up. If comparative anatomists have given but small heed to the
Neritacea, the conchologists and systematists have done their full
share of work on the group, and the works of Martens (27) and
Pilsbry and Tryon (40) give a most complete account from a
taxonomical point of view. But, as is often the case when classi-
fication is founded on external characters only, such as the shell and
the operculum, a more complete study shows that it rests on
insecure foundations ; and while I am unable to do more than
criticize the genera Verita and Neritina as usually defined, I shall
bring forward evidence which will, I hope, induce authors more.
conveniently situated for the study of these forms than I am to
undertake a revision of the family Neritide, based upon anatomical
characters.
The name Neritacea—the group has not been raised to the.
rank of a suborder or even of a tribe or section—was used by
Lamarck as a collective designation for the recent families of
Neritide, Neritinide, and Helicinide. To these have been added
the Neritopside, Titiscaniidee, Scutellinidee, Hydrocenide, and
Proserpinide, and the fossil families Maclureide and Naticopside.
It is not part of my present intention to criticize the recent
and extinct families and genera that have been founded by
conchologists, nor to discuss the probable relationships of the
paleeozoic forms which, like the genus Deshayesia, have been held
to occupy a position intermediate between the Neritide and
Naticidie (see Pethd (33), who refers it to the latter family).
But as it will appear in the latter part of this paper that the
result of my anatomical investigations is to show that the
Neritacea retain some primitive characters, and in so far as they
are specialized do not show any approximation to the Pectini-
branchia, but are contrariwise modified in a special direction,
which culminates in the terrestrial Helicinide, kec., it is of
interest to consider how far the group may be regarded as of
undoubted geological antiquity. The family Neritide is of
respectable antiquity. The genus Werita is represented by the
subgenus Lissochilus Petho in the Triassic and Jurassic, and by
the subgenus Otostoma d’Archiac in the Cretaceous of Europe,,
Algiers, and Asia Minor. The genus Deshayesia, which is con-
sidered by some conchologists ‘to “present a very remarkable.
combination of the characters of Verita and Natica and appears
to establish a passage between those genera” (Pilsbry and
Tryon, vol. x. p. 5), is from the Eocene and Miocene of the Paris
and Bordeaux basins, and if it is really a Neritid, its Naticid
characters must be due to convergence and must not be taken as
indicating a passage between the Aspidobranchiate and Pectini-
branchiate Gastropods, for such a passage must have been effected
long before the Tertiary period.
The genus Veritina, if indeed it is as distinct from Nerita as.
‘812 PROF. G. C, BOURNE ON THE [Nov. 17,
conchologists assume, is found fossil from the Liassic onward,
and is most numerous in species in the Miocene and Pliocene.
The subgenus Veritodonta Brusina, from the Tertiary of Dalmatia,
is of special interest, as beimg possibly a forerunner of the pul-
monate Hydrocena, now living in the same region. The genus
Neritoma Morris is found in the Jurassic of Europe, and the
subgenera Veridomus Morris and Lycett and Onchochilus Petho
in the Oolite and the Triassic and Jurassic respectively. The
genus Deianira Stoliczka is from the lacustrine deposits of
the Cretaceous of Europe, and Velates Montfort from the Ter-
tiaries of Europe, India, and Madagascar. ‘The limpet-like
Pileolus Sowerby, resembling the modern Septaria, dates from
the Jurassic and Cretaceous. The Neritopside, differing from
the Neritide in the characters of radula and operculum, are
yepresented by a single recent species, Veritopsis radula, from
the E. Indies and Polynesia, but are fossil from the Secondaries
and Tertiaries. Since, with the exception of Deshayesia, there is
no doubt as to the relationship of these extinct genera, it is clear
that even in earlier Secondary times the Neritacea were dif-
ferentiated into marine, estuarine, and freshwater forms more or
less resembling those of the present day, and must have been
derived from an earlier stock, which we may look for in primary
‘formations. But the remains of Neritacea from paleeozoic deposits
-ave at the best doubtful. As for some species of Verita which
have been described from this period, it is only necessary to
quote von Martens (27): ‘‘ Hinige angebliche Arten von Nerita
aus den paleozoischen Formationen sind betreffs der Gattung
héchst zweifelhaft, wie es tiberhaupt meist eime unsichere und
choffnungslose Sache ist, paleeozoische Gastropoden auf Gattungen
‘der Gegenwart zu beziehen.”
The family Maclureide, of which Maclurea Lesueur, from the
‘Cambrian and Silurian of N. America and Scotland, is the sole
genus, is placed near the Neritide because of its opercular
-apophyses, but its affinities are very doubtful, and it has at
various times been placed in the Solariide, Atlantide, Pleuro-
tomariide, or between the Bellerophontide and Haliotidee.
Naticopsis M‘Coy, ranging from the Devonian to the Trias, and
‘the subgenus Zrachydomia Meek and Worthen, from the Car-
boniferous, are placed in the Neritopside because of the characters
of the operculum, but the shell is more like that of the Naticide.
If these paleozoic genera are really allied to the Neritacea, the
latter group is of great antiquity. On the other hand, the most
specialized of all the Neritacea, the pulmonate Helicinide,
Hydrocenidee, and Proserpinide, are only found in late tertiary
deposits, and have clearly been evolved in comparatively recent
‘time from Neritiform ancestors.
The sole exception to this statement is furnished by the genus
PDawsoniella from the Carboniferous of Illinois. It is found in
association with shells of the genus Pupa, and there can be no
doubt that it was of terrestrial habit. Formerly placed in the
1908.] ASPIDOBRANCH GASTROPOD MOLLUSCS. 813
genus Helix, it is now regarded as a member of the Helicinide,
but differs from the latter in possessing a large basal columellar
eallosity covering the umbilical region. The operculum is
apparently unknown. ‘The affinities of Dawsoniella must be
considered doubtful, but in any case its resemblance to the
Helicinide is to be regarded as due to convergence, and not to
natural affinity. In the first place, it is highly improbable that
if Helicinide had existed from the Carboniferous onwards, no
trace of their remains should have been discovered in secondar
and earlier tertiary deposits. In the second place, the Helicinide
are unquestionably derived from the Neritide, and, as we have
seen, the Neritide were not established in Carboniferous times.
It is interesting, however, to note that the genus Naticopsis
(Trachydomia) occurs in the same formation as Dawsoniella, and
it seems probable that we have here an interesting case of parallel
development. If Vaticopsis,a marine form, was the forerunner
of the later Neritide, it would seem to have given rise in
Carboniferous times to the terrestrial Dawsoniella, just as the
marine Neritide have given rise in later time to the terrestrial
Helicinide ; and the ancestry and conditions of life being similar,
the two terrestrial forms acquired such a similitude that their
shells have been classified together in the same family.
Though the several genera of recent Neritacea have been
studied with minute care from a systematic point of view, we
have no very satisfying account of their habits. The genus
Nerita is confined to tropical or subtropical seas, and if we accept
for the moment the limitations of the genus as defined by con-
chologists, all the species are marine and are found for the most
part between tide-marks, clinging like limpets to the rocks.
Some of the more brightly coloured species live on coral-banks.
It has been remarked by several travellers that they are capable
of enduring a considerable amount of exposure to the air. Thus
‘Quoy and Gaimard (36) were surprised to see Verite attached to
black rocks under the full glare of a tropical sun, without
apparent injury, and they observed that these animals always
retained a few drops of water in their shells which they ejected
when forcibly torn from their attachment. C.B. Adams observed
a West-Indian species living in crevices in the rocks between the
tide-marks at the height of three-quarter ebb-tide, and the young
forms were even higher up, attached to rocks and stones which
were only wetted by spray. Practically nothing is known of the
breeding-habits of Nerita, and in view of the complexity of
the accessory genital organs, especially in the female, observations
on this point are very much to be desired.
As to the extent to which different species of WVerita are
tolerant of brackish or even of fresh water, very little information
of a satisfactory character is forthcoming. Many species are
recorded from bays at the mouths of rivers or from estuaries,
where there must be a considerable admixture of fresh and salt
water. Verita lineata Chemnitz is recorded as ascending the
814 PROF. G. C. BOURNE ON THE [ Nov. 17
Saigon River in Cochin China, as far as 20 or 25 miles from its.
mouth, and must therefore be capable of living in fresh or at
least slightly brackish water. This species 1s a ty pical member of
the genus Verita, having a thick shell with spiral coste, a den-
ticulate outer lip, and a ‘granular operculum, and is (omen ale as.
being one of the few species with these characters which is
tolerant of both salt and fresh water, though there are several
species of Weritina, usually a freshwater genus, which live in
brackish water or are even marine. It seems probable, however,
that several species of Verita are capable of existing in brackish
water, since many of them are recorded from bays at the mouths.
of ei ge rivers, and an observation made by Mrs. Longstaff—to
whom I am indebted for some well- preserved specimens of Verita
plicata Linn.—shows that fresh water is not fatal evento
exclusively marine species. Mrs. Longstaff attempted to Kull
some individuals of this species by immersing them in fresh
water: they were apparently uninjured, but did not lke the new
conditions and crawled up the side of the vessel in which they
were placed, fixing themselves round its rim, apparently ready to
withstand a considerable sojourn in the air.
This question of habitat, in fresh, brackish, or salt water, is of
some importance in considering the generic distinction between
Nerita and Neritina. ‘The species of the latter genus are mostly
inhabitants of fresh water, and some are found only at the
sources of streams, far away from the sea. The numerous
European species, of which JV. fluviatilis is the most familiar
example, are freshwater forms, but occasionally occur in brackish
or even salt water. nies many of the tropical species are partly
or wholly marine, e. g. V. waluensis Lesson of the Indian Ocean
and Polynesia. Ao and Gaimard (35) found WV. auriculata Lam..
in the sea. Dr. Ed. von Martens (27) describes the following
S. American forms as ‘‘species submarine.” WV. virginea is common
in brackish water, but var. listeri, of E. Nicaragua, is found
throughout the river San Juan, and also in Greytown harbour
in localities where the water is alternately brackish and fresh.
NV. picta Sowerby from 8. Panama was observed in abundance-
on a mudbank covered at times with fresh water, and has been
described as strictly marine by C. B. Adams. AV. viridis, not rare
in the Mediterranean and in the Caribbean Sea, also in the
Bermudas, is truly marine and lives on Zostera. From all of
which it follows that whereas WVerita 1s very rarely found in fresh
water, Veritina is much more easily accommodated to different
conditions of life.
In what precedes, I have accepted the usual distinctions.
between the two genera founded upon the characters of shell and
operculum, but it is a question, as will be more clearly shown in
the sequel, whether these characters are of sufficient importance:
to afford generic distinctions between forms, the internal anatomy
of which is, in nearly all respects, so similar as to be practically
indistinguishable; and the further question will be raised as to.
1908. | ASPIDOBRANCH GASTROPOD MOLLUSCS. 815
whether anatomical differences of an important character do not
indicate the distinction of Neritina fluviatilis—and possibly of
other nearly allied European species from the tropical forms
usually classed in the same genus. It may be pointed out here
that the conchological characters relied upon in the deter-
mination of the two geneva are confessedly somewhat obscure.
In general, Verita has a thicker and more solid shell, usually
ornamented with spiral ribs, but these may be absent, as in
polita, morio, picea, &e.; when ribs are present they usually project
slightly beyond the outer lip, which is then dentate, but it may
be smooth. The inner surface of the outer lip is generally
dentate, but this character may be absent. The operculum is
usually solid, with a granular outer surface, or with a marginal
zone, and the shape of the apophyses springing from its inner
side is claimed to be characteristic.
Dr. Ed. von Martens (27), the leading authority on the
classification of the Neritacea, maintains that the denticulation
on the inner side of the outer lip and the characteristic sculpture
of the operculum are the most constant differences between
Nerita and Neritina, “in den meisten Fallen auch der allgemeine
‘die Meerbewohnerin verkiindende Habitus, der aber bei den
kleineren schwarzen Arten weniger hervortritt.”. In Neritina
the shell is usually thinner and less solid, not ornamented with
spiral ribs, though these are present in .V. cornea Linn. and
especially in its variety swbsulcata, and spirally arranged rows of
spines or nodules are not uncommon. ‘The outer lip is smooth
and not dentate (it is distinctly crenulate in V. granosa Sowerb.
and iV. aculeata Gmelin) and the imner denticulations of the
outer lip are wanting. The operculum is not granular or sculp-
tured, but minute granulations may be discerned with the aid of
a lens in several species; I found them specially well marked in a
specimen of V. recluziana Guillou. As for the apophyses, after
making a careful comparison of these structures in all the species
of both genera that I possess, I have concluded that they offer so
many examples of convergence that they are quite unreliable for
the purpose of generic distinction, but, as I make no pretension to
skill as a systematist, my judgment in this matter must be taken
for what it is worth.
A certain number of Neritine have the last whorl of the shell
broadly expanded, the aperture enlarged, and the spire reduced,
so that they acquire a secondary symmetry ; such, for instance,
are .V. dilatata Brod. and WV. crepidularia Lam. It is interesting
to note that those forms which tend towards a bilateral symmetry,
such as the two species quoted, and also VV. auriculata Lam.,
NV. tahitensis Lesson, NV. bicanaliculata Récluz, are all from the
Indian or Pacific Ocean, and are clearly intermediate between the
more common spirally coiled Neritine and the genus Septaria
(= Navicella Lamarck), which is confined to the same regions.
On the other hand, the expanded American species of Veritina,
e. g. latissima Brod. and intermedia Sowerby, retain the distincs
Proc. Zoou. Soc.—1908, No. LIT. 52
816 PROF. G. C. BOURNE ON THE [Nov. 17,
dextral coil of the spire and their shells are asymmetrical, in
general form somewhat like Haliotis. The same is the case with
NV. oweniana Gray, from the W. Coast of Africa, Fernando Po,
and Cape Palmas ; and thus it appears that the Neritine of rivers
running into the Indian and Pacific Oceans (but not of American
rivers on the Pacific slopes) have given rise to the lmpet-like
fluviatile genus Septaria, whilst those of the Atlantic shores have
followed a different line of evolution and have in no case given rise
to Septaria forms.
The genus Septaria Férussac is commonly referred to in
zoological works as Vavicella Lamarck, but Ferussac’s name has
the priority and, according to all rules of zoological nomenclature,
ought to be adopted. It appears to be a characteristically fresh-
water genus, living on the roots of Nipa palms and other trees on
the banks of rivers in India, Mauritius, Bourbon, N. Australia,
and the Pacific Islands. The Septarie of the Mascarene Islands
and Madagascar affect the vicinity of waterfalls, and are found
adhering to stones out of the water but constantly wetted by
spray. Due allowance being made for the secondary symmetry
which they have acquired, the internal anatomy of individuals
of this genus closely resembles that of Verita and the tropical
Neritine. Little or nothing is recorded of the breeding-habits
and development of Septaria, but some specimens of S. Lougain-
villei Récluz, a Fijian species In my possession, have egg-cases,
resembling those of Veritina fluviatilis, attached to the shell, each
case containing a number of veliger larve.
No more than a passing mention can be made here of the
Titiscaniide, of which 7%tiscania Bergh is the sole genus. It is
a slug-like marine form, probably widely distributed in the Indo-
Pacific seas, but hitherto recorded only from the Philippines and
Mauritius. It isa highly specialized member of the Neritacea,
with something of the form and habits of a Nudibranch. The
shell is lost, but the ctenidium and mantle-cavity, though reduced,
remain. Bergh (2) has given an account of the anatomy of this
remarkable genus, from which it is evident that it is rightly
placed among the Neritacea, and probably near to Veritopsis,
because of the absence of the median plate in the radula. There
are, however, many imperfections in Bergh’s account of the
anatomy. Believing that the supra-intestinal nerve was absent
in Verita, he failed to find it in Jitiscania, and we are left in
doubt as to whether it is really present in the latter genus cr not.
For similar reasons doubt must be thrown on his description of
the heart with a single auricle, and his account of the generative
organs is so wanting in precision and detail that one can only
gather from it that the accessory glands and ducts are complicated,
and may very possibly resemble those of Verita.
The remaining families of the Neritacea, the Helicinide, Pro-
serpinide, and Hydrocenidee, are, as is well known, terrestrial
pulmonate forms, with the habits of snails. Their distribution
coincides on the whole closely with that of the tropical Merite
1908.] ASPIDOBRANCH GASTROPOD MOLLUSCS. 817
and Neritinw. The Helicinide occur chiefly in islands in the
Indo-Pacific region, and in the Antilles, reaching their maximum
in the last-named locality. As has been already remarked, they
are only found fossil in later Tertiary deposits, with the exception
of the remarkable Dawsoniella, which occurs in the Carboniferous
of Illinois, and is apparently related to Trachydomia, a subgenus
of Vaticopsis, from the same formation. The present distribution
and the geological history of these Pulmonate Neritacea suggest
problems which will be dealt with in the second part of this paper.
With the exception of Dawsoniella, they are unquestionably
descended from Neritid or Neritinid ancestors, and it is difficult
to account for their dispersal in islands so widely separated from
one another without making assumptions which differ widely from
accepted doctrines of animal evolution.
It was my original intention to undertake a monographic
account of the anatomy of the Neritacea, but as the work proceeded
it became evident that, owing to the difficulty of obtaining suit-
able material, and the great number of species which must
necessarily be subjected to microscopical examination before com-
pleteness could be attained, this would be a task of many years’
duration. The results already attained seem of sufficient import-
ance to merit publication, and may induce zoologists travelling or
living in tropical regions to give attention te a group deserving
more attention than it has received. I therefore purposely omit
a detailed account of certain organs, such as the buccal bulb, the
radula, and the histology of the alimentary tract, the mantle, &c.
My chief attention has been directed to the nervous system, the:
kidney, the pericardial’ cavity and its connection with the rest of
the celom, and the generative organs.
The difference between the aquatic Neritide and the terrestrial
pulmonate Helicinide is so considerable that they must be dealt
with in separate sections of this memoir, and in each case, in
order to avoid a confusion of my own observations with a criticism
of the work of other authors, I will preface my statements with a
short account of the literature of the subject.
Family NERITIDZ.
The first important contributions to the anatomy of WVerita are.
those of Quoy and Gaimard (85 & 36), whose figures and descrip-
tions of the nervous system and alimentary tract leave much to be
desired, and it is not necessary to enter into an examination of
their errors and omissions. But they made some observations on
the generative organs which, though far from complete, have not
been followed up, and have scarcely been noticed by any subse-
quent author except Bergh. It will be best to quote their
description in full:—‘‘Dans le sexe femelle est un groupe
Vorganes qui mérite quelque attention. On y voit l’extrémité
du rectum, puis un corps pyriforme trés-allongé entouré en partie
dune sorte de glande striée en travers, qui s’ouvre vers le bas.
52*
818 PROF. G. C. BOURNE ON THE [Nov. 17,
Cet organe est creux et contient dans sa cavite, accollés les uns aux
autres, plusieurs corps en massue allongée, finissant en filaments.
Ils sont résistants, comme fibreux, et paraissent grenus a la
loupe. Nous ne pouvons deviner usage de ce petit appareil, qui
remplit sans doute quelques fonctions relatives a la géneration,
puisqu’on ne le trouve que chez les femelles. | Plus en dehors est
Vutérus, composé dune poche pyriforme et dun renflement qui
lui est accollé, lequel contenait une grande quantité-d’ceufs, ronds
blanes et crétacés. L’oviducte, gros long et tortueux, fait com-
muniquer cet organe avec lovaire, placé au bord droit du foie.”
Though this description and the figure accompanying it are
inexact, it is evident that the “corps pyriforme” is the sperma-
tophore-sac, the ‘corps en massue allongée” are the sperma-
tophores, the “ glande striée en travers” is the ootype, with its
glandular walls, the “uterus,” as described and figured by these
authors, has no separate existence, but the ‘“renflement qui lui
est accollé” is the crystal-sac, which does, in fact, open into the
distal end of the ootype. It also seems probable that MM. Quoy
and Gaimard mistook the spherical crystalline concretions in the
erystal-sac for ova.
Of the male, Quoy and Gaimard give a very insufticient account
of the accessory generative organs, but observed the excessively
long coiled region of the sperm-duct to which I have given the
name of epididymis.
In an earlier memoir Quoy and Gaimard (35) gave a super-
ficial account of the structure of Werita, which only merits
attention because it contains two figures showing the modification
of the cephalic integument at the base and to the inside of the
right tentacle of the male, which has been referred to, but
seldom correctly figured or described, by subsequent authors as
a “cephalic penis.” In the figures referred to this structure
is represented in the correct position, but simply as a conical
eminence, without any detail.
From the time of Quoy and Gaimard there is no work dealing
with the anatomy of Verita till that of Bouvier in 1886. Von
Jhering (22), in his well-known work on the nervous system of
mollusca, abruptly removes the whole of the Neritacea from their
position alongside of the other Rhipidoglossa and places them in
a class Orthoneura, which has long since been broken up, its
contents being restored to their proper places by subsequent and
more exact observers. But his investigations were confined to
the nervous system of WVeritina fluviatilis, and had he carefully
studied the anatomy of some of the larger species of Verita or
of a Septaria he would probably have paused before promulgating
the opinions set forth in his lengthy memoir.
Bouvier (8), in a preliminary note published in 1886, gave a
short account of the principal external features of the anatomy
of Verita and some details of the nervous system, but the reader
should turn for a more complete account to his great work (9)
on the nervous system of prosobranchiate Gastropods. As he
1908.] ASPIDOBRANCH GASTROPOD MOLLUSCS. 819
himself subsequently corrected his errors about the nervous
system, it is not necessary to dwell on what is now a matter of
history. Failing to recognize the extremely fine supra-intestinal
nerve in any of the Neritide he dissected, he wrote with charac-
teristic emphasis, “Il n’y a pas de commissure viscerale croisée,”
and classed the Neritacea as ‘“‘ Rhipidoglosses orthoneuroides,”
Following de Lacaze-Duthiers, he identified the swollen origin
of the subintestinal nerve with its sheath of ganglion-cells as the
subintestinal ganglion, but, curiously enough, did not observe the
large ganglion on the visceral commissure, afterwards discovered
and called the subintestinal by Béla Haller (20) and Boutan (6).
But his description of the nervous system is much in advance of
anything that preceded it. He was the first to discover the
course of the subintestinal nerve and of the left pallio-branchial
nerve. He discovered and described correctly the labial com-
missure, characteristic of the more primitive prosobranchs. This
commissure, as he says, is “ trés facile a préparer,” and it is curious
that Bela Haller, who succeeded in the much more difficult task
of tracing the supra-intestinal nerve, should have emphatically
denied the existence of this very obvious labial commissure.
There is a further point of difference between these two authors,
in which Bouvier appears to me to be correct. Béla Haller
describes no less than fourteen transverse commissures behind the
anterior commissure of the pedal nerve cords, whereas Bouvier
found, as I find, nothing more than fine nerves passing from the
inner sides of the cords to the muscles of the foot. In Bouvier’s
brief account of some of the more important anatomical features
of Verita peloronta there is a curious misprint, which has created
some confusion among some subsequent authors. On p. 47 he
writes: ‘“ Au fond de la cavité branchiale, a droite, se trouve
la branchie bipectinée, libre en avant, en arriére rattachée au
manteau a droite et & gauche par un expansion de la lame
médiane, de sorte que le fond de la cavité branchiale est divisé
en deux étages superposés. A gauche de la branchie se trouve
le rein; il souvre dans la cavité branchiale par un orifice en
boutonniere situé dans la paroi antérieure du nucléus.” The
words italicised ought to be transposed: the ctenidium, of course,
is on the left side of the mantle-cavity, and the kidney is
to the right of the ctenidium. Bouvier further describes the
. so-called cephalic penis “toujours assez réduit dans les Nérites,”
and gives a figure of a remarkable development of this organ in
Neritina cariosa. Though his description and figure are not very
clear on this point, Bouvier appears to have determined the true
position of the osphradium in Verita, but as he did not examine
the structure of this organ and did not recognize the ganglia con-
nected with it, his determination is rather of the nature of a
conjecture than of proof.
The next contribution to the anatomy of WVerita is that of
Rémy Perrier (34), whose researches were confined to the
kidneys and associated organs in WNerita peloronta, Neritina
820 PROF. G. C. BOURNE ON THE [Nov. 17,
oweni, and Septaria (Navicella) janelli. The position and general
anatomy of the kidney of Veritina flurratilis had been previously
described by Landsberg (24); and Perrier adds some details
relating to the trabecular structure of the excretory portion of
the kidney and the reno-pericardial canal. He did not, however,
fully elucidate the relations of the glandular and non-glandular
parts of the kidney, and described the latter as a closed sac inter-
vening between the kidney and the pericardium. This error was
afterwards corrected by Lenssen. The most important part of
Perrier’s work, in so far as it relates to the Neritide, is his
account of the heart. He discovered and gave an accurate figure
of the left auricle and showed that, contrary to Landsberg’s state-
ments, the ventricle is in fact traversed by the rectum.
Bergh (2) in 1890, as an addendum to his paper on 7%tiscania,
gives an account of the anatomy of Verita peloronta and Neritella
(Neritina) pulligera. This is the first attempt, since Quoy and
Gaimard, to give a complete account of the anatomy of Verita,
but it is unfortunately very incomplete and contains some
serious errors and omissions. For example, Bergh denies the
existence of a second auricle, and lays considerable stress on its
absence. He describes the eyes as open, whereas they are in fact
closed. His description of the nervous system, correct enough as
far as it goes, is no advance on the original description of Bouvier.
He gives a more or less detailed and tolerably correct account of
the buccal bulb, odontophore and radula, and notes the presence
of salivary glands, but mistakes an cesophageal dilatation for the
stomach, and describes the true stomach as enlargements of
the hepatic ducts. The position of pericardium and kidney are
correctly described without adding anything to previous know-
ledge of the subject; but the reno-pericardial duct was not
recognized. All of Bergh’s specimens appear to have been
females, and he makes an attempt to describe the complicated
accessory generative ducts and glands, but, as he says, ‘‘ bei den
vorlegenden Materiale konnten die ganz unklaren Verhiltnisse
dieser Theile nicht genauer eruirt werden.” He recognized, how-
ever, the spermatophore-sac, and gives a good outline figure of a
spermatophore of V. pulligera.
In 1892 two short papers by Boutan (6) and Bouvier (10), the
atter published very shortly after the former, established the
existence of a supra-intestinal nerve in Verita and Septaria, thus
restoring the Neritacea to their proper place among the Strepto-
neurous Rbipideslecs. Shortly afterwards Boutan (7) published
a further account of the nervous system of WVerita polita and
Septaria (Navicella) porcellana, in which the course of the supra-
intestinal nerve is correctly figured, but he failed to recognize the
supra-intestinal ganglion which Bouvier had signalized in the
previous year. Boutan appears to have been in error as to the
position of the osphradium, which he says “s’étend le long du
septum branchial qui réunit la branchie au plancher de la cavité
et est a peine distinct & l’eil nu.’ The osphradium, as I shall
1908. | ASPIDOBRANCH GASTROPOD MOLLUSCS. 821
show, is in fact in front of the suspensory membrane of the
ctenidium, close behind the thickened margin of the left side of
the mantle and in front of the anterior end of the left columellar
muscle. Close below the osphradium is a complex of ill-defined
ganglionic enlargements, and as Boutan failed to find the true
osphradium he missed the ganglia lying beneath it. In this
same paper Boutan argues, erroneously as I now think, that the
so-called subintestinal ganglion of Bouvier is not a member of
the group of visceral ganglia and gives figures of the pleuro-pedal
ganglia of Verita and Septaria correcting the older figures of the
latter author.
In the following year Béla Haller (20), in the course of his
studies on docoglossate and rhipidoglossate Prosobranchs, gave a
tolerably full account, not only of the nervous system, but also
of the alimentary tract, kidneys, and genital organs of Werita
ornata. This work contains a curious mixture of acute and
accurate observations and incomprehensible errors and omissions.
His elaborate figure of the nervous system is in some respects
the best that has been published, but in other respects is most
misleading. As has already been mentioned, he flatly denies the
existence of a labial commissure, which is not only certainly
present, but much easier to dissect than in any other Rhipido-
glossate. I can positively assert that the numerous pedal
commissures figured by Béla Haller are not present: Bouvier
was perfectly correct on this point. In a simple dissection, one
may easily make mistakes in attempting to trace delicate nerves
through the mass of muscle in which they are embedded, but a
study of microscopical sections leaves no room for error. A
careful examination of a series of sections of several species fails
to reveal any trace of transversal commissures posterior to the
main pedal commissure. B. Haller discovered the supra-
intestinal nerve, independently it seems of Boutan and Bouvier,
and gives a fairly correct figure of the crossed visceral commissure.
Like Boutan he identifies the elongated ganglion on the right of
the crossed visceral commissure as the subintestinal, but he did
not see the stout nerve given off from it, almost immediately
swelling up to form the genital ganglion lying on the oviduct or
sperm-duct. In respect of the supra-intestinal and branchial or
osphradial ganglia, Haller gives a complicated figure which, as
far as I am able to reconstruct these ganglia from serial sections,
may be correct, but after many attempts I have been unable by
simple dissection to verify his account. These ganglia are covered
by the thickened and folded epithelium of the osphradium, which
in all the species at my disposal is too opaque to allow the
ganglia to be seen by transparency.
Haller’s description of the alimentary tract is much more
accurate than that of his predecessors. He gives a good account
of the position and general relations of the stomach, cesophagus,
and course of the intestine, but his observations on the buccal
bulb, salivary glands, &c. seem to me defective. He describes
822 PROF. G. C, BOURNE ON THE [Nov. 17,
and figures a posterior diverticulum of the buccal bulb which I
have failed to discover either in sections or by dissection, and
his drawing of the salivary glands is incorrect in detail. In
describing the heart he has, curiously enough, fallen into the same
error as some of his predecessors, since he categorically denies the
presence of a rudimentary right auricle: ‘‘da von einem rechten
Vorhof nicht einmal ein Rudiment mehr erhalten ist.”
After criticizing Perrier’s account Béla Haller gives a somewhat.
detailed description and a figure of the kidney of Verita ornata,
but neither description nor figure is correct. According to him
the kidney is an acinous gland, not differentiated into anterior
and posterior lobes differing in histological structure. The ducts
of the acini unite and open by large apertures directly into the
bladder (Urinkammer). The reno-pericardial canal opens into
the bladder and is dilated into a large sac which runs back
posteriorly between the pericardium and the ureter, and is
identified with the cavity described by Perrier as lying between
pericardium and kidneys and incapable of being injected from the
general body-cavity. I shall prove, in due course, that the
glandular part of the kidney is not acinous, that there is a histo-
logical differentiation between the anterior and posterior moieties,
that there are not several ducts leading from the glandular part.
to the ureter, and that the reno-pericardial canal opens not into
the bladder but into the glandular portion.
As for Haller’s description of the male and female generative
organs, I need only say that his work is scarcely an advance on
that of Claparéde, and he failed to discover the remarkable com-
plexity of these organs, which, indeed, could hardly have been
discovered without careful and laborious reconstruction of
sections.
It could not be guessed from the title ‘‘ Die systematische
Stellung der Solenogastren und die Phylogenie der Mollusken ”
that Thiele’s (89) memoir, published in 1902, contains a number
of new and acute observations on the morphology of the Neritacea.
Interpolated as they are in a lengthy discussion of the phylo-
genetic history of the Gastropoda, Thiele’s results are somewhat
difficult to summarize, and it is to be regretted that he did not
see fit to embody them in a separate memoir. He studied
sections of Verita pica, Septaria parva and suborbicularis, Scu-
tellina cinnamomea, and Helicina japonica. It should be noticed
in the first place that he places Scutellina without comment among
the Neritidze. Scwtellina was classified by Fischer (15) among the
Docoglossa, by Pilsbry and Tryon (40) near the Haliotide, and I
have been unable to discover what author detected their relation-—
ship to the Neritide. It is clear, however, from Thiele’s account.
of the female generative organs that it belongs to the last-named
family. After touching on various points of the anatomy of
the Neritide, such as the ctenidium, which he compares with
that of the Acmeide rather than the Trochide; the subpallial
sense-organ, which he describes and figures correctly but is
1908.1} ASPIDOBRANCH GASTROPOD MOLLUSCS. 823
inclined to identify with the subpallial sensory tracts of Patellide,
&e., rather than with a true osphradium; the left columellar
muscle, which he considers to be derived from the subdivision of
the primitive right muscle; the salivary glands, in respect of
which he corrects the statements of Haller and Amaudrut, Thiele
proceeds to give a more detailed account of the accessory genital
organs. Though his diagrams are too schematic, his drawings of
sections too few in number, and his description too condensed
to convey a clear impression to anyone unfamiliar with these
complex structures, his account of the female organs of Verita pica
and Septaria parva is very exact, both as regards the general
anatomy and the histology. I shall have occasion to refer
frequently to it in the descriptive part of this paper. It need
only be mentioned here that he does not appear to have found
spermatophores in the spermatophore-sac, and therefore is obscure
as to the function of this organ. Though he found and has
figured the peculiar crystalline concretions in the crystal-sac, he
names this structure the uterus—for insufficient reasons, as it
appears tome. He did not discover the oviduco-ccelomic funnel,
and does not mention the presence of the third duct, which I have
called the ductus enigmaticus in Septaria parva. It is of course
possible that it is not present in this species. The description of
the female organs of Scutellina cinnamomea leaves no doubt that
this form is a member of the Neritide. The description of the
male organs of Helicina japonica will be dealt with in the second
part of this paper, and | can supplement it by an account of the
female organs of <dlcadia. Thiele regards the ‘“receptaculum
seminis,” 7. e. the spermatophore-sac, as the representative of the
right kidney in female Neritide, and though I do not agree with
this conclusion it is not far from the truth.
Further on Thiele gives a description of the kidney in Verita
pica and in Septaria, and here also makes more accurate obser-
vations than any of his predecessors. He also notices the
extension of the pericardial cavity to the right side of the animal
in Septaria, and makes a just comparison between the conditions
obtaining in this animal and the Cephalopoda. In conclusion,
Thiele suggests that the Neritide may have been derived from
the Trochide, but points out features in which they show a
- resemblance to the Docoglossa. The latter, however, as he says,
are more probably analogies than homologies, as the radula and
the structure of the generative organs preclude any idea of close
relationship between these groups.
The genus WVeritina, owing to the abundance of the common
NV. fluviatilis in Kuropean rivers, has been more often and more
thoroughly studied than the genus Werita. It is not necessary
to do more than refer to the works of Moquin-Tandon (28),
Claparede (12), and Landsberg (24), or to the paper on the develop-
ment of Weritina by Blochmann (4), because the results obta*ned
by these authors have already been discussed and entirely super-
seded by the admirable papers of Lenssen (25 & 26). In the first
824 PROF. G. C. BOURNE ON THE [Nov. 17,
of these two papers Lenssen deals with the digestive and genital
systems, giving a detailed account of the bucco- pharyngeal
cavity, the odontophore, the cesophagus with its glandular
appendages, and the stomach. He and Gilson (18) are the only
authors besides Thiele who have published an accurate account
of this system of organs in the Neritacea. Gilson and Lenssen
discovered the remarkable fact that in WVerttina, a dicecious
-Gastropod, the female ducts are diaulic, whereas the male ducts
are monaulic; and the latter author gives a thorough and accurate
description of the very complex arrangements of both male and
female organs. I have only to say that I have carefully verified
Lenssen’s statements and find nothing to correct and very little to
add to them as regards the species examined, Veritina fluviatilis,
but I find considerable and important differences in some of the
tropical Neritine.
In his second paper Lenssen deals with the nervous, circulatory,
respiratory, and excretory systems of WVeritina fluviatilis. Here
he has not been in some respects as accurate as in his first paper.
For instance, in the description of the nervous system (p. 297)
he confuses the labial with the buecal commissure. It is clear
both from his text and figure that the commissure that he dis-
covered is the buecal commissure, but he calls it the labial.
It is practically impossible to dissect out the true labial
commissure in so small an animal as WV. fluviatilis, and it is
exceedingly difficult to trace it in sections; but I have satisfied
myself that it exists. Im other respects Lenssen’s account and
figure of the nerve-centres appear to be correct, and I can confirm
his statement that there are no transverse commissures behind
the single large commissure uniting the anterior ends of the pedal
cords.
As regards the visceral and pallial nerves Lenssen makes a
considerable advance on his predecessors and he accepts Bouvier’s
identification of the subintestinal ganglion. He discovered,
apparently without being aware that Boutan and Béla Haller had
anticipated him in this matter, the ganglion on the subintestinal
nerve at the point where the latter turns rather sharply from
right to left to course close below the surface on the dorsal side
of the pedicle attaching the anterior’ part of the body to the
visceral mass. His account of the relations of this ganglion and
of the nerves given off from it is for the most part very exact, but
he does not appear to have observed that the genital nerve (doc.
cit. pl. i. fig. 1, ms.) almost immediately enlarges to form a
ganglion ‘of considerable size, closely attached to the oviduct (or
sperm-duct). He further describes a structure which he hesitates
to identify as the rudiment of the right ctenidium. ‘A cet
endroit,” he says, “il existe un organe creux, l’homologue, peut-
&tre des mamelons découverts chez les patelles et dautres proso-
branches. Cet organe renferme un grand nombre de globules
sanguins et semble, par conséquent, dépendre soit de Vappareil
“cinelisisariee soit de Yappareil respiratoire. I] fait saillie dans la
1908. | ASPIDOBRANCH GASTROPOD MOLLUSCS. 825
cavité branchiale et s’ouvre 4 sa base dans le sinus sanguin que
nous venons de signaler.” Further on (p. 312) he discusses the
homology of this organ and suggests that it may represent the
right ctenidium or the right osphradium, but gives no decided
opinion on this point. A description and discussion of the
significance of this organ will be found on p. 864.
Though he made, as he tells us, a careful search for it, Lenssen
failed to discover the supra-intestinal nerve; but being cautious
he does not venture to affirm that it does not exist. Nor am I
ready to deny its existence, but after searching most carefully
through several series of sections I am unable to discover a trace
of it; and it is very possible that this nerve, extremely small in
Nerita and the tropical species of Weritina, has actually dis-
appeared in VV. fluviatilis. In the descriptive part of this paper
I shall have something to add to Lenssen’s account of the left
branchial ganglion and the osphradium. It is not necessary for.
me to refer at length to Lenssen’s account of the circulatory,
respiratory, and excretory systems. Though somewhat short, his
descriptions of these systems are accurate so far as they go, and
he is the first author to give a true and intelligible account of the
kidney and reno-pericardial duct.
I make no separate reference to the literature bearing on the
anatomy of Septaria (Navicella). This genus has not been studied
in detail by any author, but Bouvier, Boutan, and others have
described the nervous system in the works already quoted. As
my interest in the Neritacea dates from some dissectious of
Septaria which I made for the purposes of my class, and as the
secondary symmetry acquired by this genus makes it a very
favourable object for describing and figuring the somewhat
complex relations of the ccelom and genital ducts in the Neritacea,
I will begin the account of my own work with a description of its
anatomy.
Genus SeprariA Férussac.
The species of this genus available for my researches were
WS. borbonica Bory, S. depressa Lesson, both forming part of the
collections of the Oxford Museum, and S. bougainvillei Reécluz,
from the British Museum. The number of specimens at my
disposal was small, and I unfortunately dissected the only two
specimens of Septaria borbonica that I possessed before I had
made myself thoroughly familiar with the problems of Neritacean
anatomy. A specimen of S. depressa was cut in horizontal and
one of S. bougainvillei in transverse sections. Both these
specimens proved to be females. There are some minor points of
difference in the anatomy of the two species which will be referred
to in due course.
A dorsal view of S. borbonica is given in fig. 1*. The roof
* The figure-numbers 1-69 in this Memoir refer to the figures on Plates XLVI.—
LXVL., which are described on pp. 885-837. There is only one text-figure (text-
fig. 172, p. 855).
826 PROF. G. C. BOURNE ON THE [ Nov. 17,
of the mantle-cavity has been cut through and largely removed
to show the principal organs of the pallial complex. The head.
is relatively large; the tentacles short and swollen at their bases ;,
the eyes, as in all Neritacea, borne on prominences at the outer
sides of the bases of the tentacles. Owing to the abortion of
the visceral spire the animal has acquired a secondary symmetry,
which does not, however, extend to the more important systems
of organs. The right and left columellar muscles, em.l and em.r,
are subequal in size and symmetrically disposed right and left of
the body. The visceral spire is reduced to a triangular mass at
the posterior end of the body. To the right side of the mass is.
the ovary, ov.; the left side is occupied by the stomach covered
over by the liver.
The mouth, situated on the ventral side of the head, is at the-
end of a very short snout, which can be scarcely retractile. The.
foot is large and oval, occupying nearly the whole of the ventral
surface behind the snout: it is surrounded by a rudimentary
epipodial ridge. The operculum is wedged in between the viscera
and the upper surface of the foot, extending as far forward as the.
posterior end of the buccal bulb. It is functionless, at any rate
as regards the closing of the aperture of the shell, but it seems to.
give support to the muscles of the foot, and retains a rudiment of
the apophyses characteristic of the opercula of the Neritide.
On the dorsal side, after the removal of the roof of the mantle-
cavity, the single bipectinate ctenidium, the post-torsional left,
is seen lying obliquely across the mantle-cavity, its base attached
to the left side and its free end pointing forward and to the right.
As in most ctenidiate Neritacea, the proximal moiety of the
ctenidum is attached by a suspensory membrane to the right and
left walls of the mantle-cavity, in consequence of which arrange-
ment the posterior half of the cavity 1s divided into an upper and
a lower chamber.
The heart, enclosed in a spacious pericardial cavity, lies on the
left side, just behind and below the posterior end of the left
columellar muscle. It cannot be seen in a dorsal view, but its.
position is indicated by pc. The rectum, after traversing the
ventricle, crosses obliquely from right to left just in front of the
visceral mass, 1s partly embraced by the complex mass of accessory
genital glands and ducts, g.d, and opens by the anus near the
anterior end of the right coluinellar muscle. The kidney, £, lies
between the rectum and the basal half of the ctenidium : it opens.
by a slit-shaped pore into the lower chamber of the mantle-cavity
close to the right side of the base of the ectenidium, but the
opening cannot be seen in the drawing. It is perhaps necessary
to state here that the kidney is the post-torsional left, as has.
been fully recognized by recent authors on Molluscan anatomy.
It is therefore the homologue, not of the large functional kidney
of other Aspidobranchia, but of the so-called papillary sae of
Trochide and Haliotide, and of the rudimentary left kidney of
the Docoglossa.
1908. ] ASPIDOBRANCH GASTROPOD MOLLUSCS. 827
The Alimentary Tract.
Fig. 2 is an illustration exhibiting the macroscopic characters
‘of the buccal bulb, cesophagus, stomach, and intestine. It would
be possible to write at considerable length on the structure and
histology of these various regions, but I purposely refrain from
“«loing so, although my preparations have enabled me to study
them with considerable accuracy. Allowance being made for
small and unimportant differences in proportion and detail, the
‘structure of the alimentary tract of Septaria is so closely similar
to that of Veritina fluviatilis as described by Lenssen (25), that it
‘is superfluous to give a description which would be little more
than a repetition of his accurate observations. I need only call
attention to one or two minor points. I find that in Septaria, as
in WV, fluviatilis, there are seven buccal cartilages, three pairs and
one median and azygos. The smallest pair, discovered for the
‘first time by Lenssen, does not strictly belong to the odontophore,
but lies in the antero-inferior walls of the buccal bulb and serves
as supports for a pair of pads, covered by a horny cuticle, against
which the right and left halves of the anterior end of the radula
work. In Septaria there is a small pair of glandular sacs, one
‘on each side, opening into the lateral extensions of the subradular
diverticulum of the buccal cavity. These have been noted by
‘Thiele in Nerita pica, but are not recorded by Lenssen in
Neritina fluviatilis. The csophagus in Septaria passes to the
left on leaving the buccal bulb and shows clear traces of the
larval torsion so carefully described by Amaudrut (1). Just
before its junction with the stomach it expands considerably and
receives three large ducts from the liver. The cesophagus may
be said to join the stomach tangentially; hence its aperture is
prolonged backwards as a wide groove, bounded by thickened
‘epithelial lips, which, while they differ slightly in detail, have
the same relations as are described by Lenssen in Weritina
fluviatilis, The stomach of Septaria consists, as is the case in all
‘the Neritide I have examined, of a dilated cesophageal and a
narrower pyloric moiety. In the former there is a large and
prominent epithelial ridge, described by Lenssen as the “ créte
stomacale,” conspicuous for its triangular appearance in section,
Its extremely long columnar epithelial cells are always covered
‘by a thick apparently cuticular product, which in appearance and
composition seems to be similar to the cuticular ning continuous
with the crystalline style found in somany Molluscs. In Septaria
‘there is a small digitiform diverticulum of the cesophageal moiety
of the stomach, situated between the lower end of the “ eréte
stomacale” and the upper border of the cesophageal groove.
‘This diverticulum, which is probably homologous with the spiral
diverticulum of the stomach of Haliotis, appears to be absent in
Neritina fluviatilis. The intestine and rectum do not call for
any special mention. The histology of the different regions
‘varies, and the variations have been sufficiently described by
828 _ PROF. G. C. BOURNE ON THE [ Nov. 17,
Lenssen. The disposition of the coils of the intestine and their
relations to the stomach, cesophagus, and radula-sac are indicated
in fig. 2, as is also the position of the heart and the fact that the
ventricle laps completely round the rectum. The radula-sac is
large and usually of considerable length, but varies considerably
in different specimens. When long it is involved in the coils of
the intestine and its posterior part always passes ventrad of the
cesophagus but dorsad of the stomach.
The Nervous System.
The main features of the nervous system have already been
described by Bouvier (9 & 10) and Boutan (7). The latter
author, correcting and amplifying the earlier account of Bouvier,
describes a supra-intestinal nerve completing the streptoneurous
condition of the visceral nerve, and gives an amended figure of
Bouvier’s drawing of the pleuro-pedal nerve-centres. In my
earlier dissections I failed to identify the supra-intestinal nerve,
but have been able to follow its course more or less completely in
my serial sections, and am able to verify Boutan’s statements as
far as they go. In one particular I can add to them. Boutan
traced the supra-intestinal nerve from its origin from the right
pleural ganglion along the right side of the body, whence it turns
over the gut towards the left side and courses, as he says, “‘ dans
la cavité branchiale, au niveau du tiers inférieur de la branchie.”
It is hardly correct to say that the nerve passes into the branchial
cavity. After a considerable amount of trouble I have been able
to trace the nerve as far as the osphradium, the precise character
and position of the latter organ having been overlooked by
Boutan. The supra-intestinal nerve on arriving at the left side
of the body passes obliquely forward in the connective tissue
underlying the integument on the dorsal side of the left
columellar muscle. Near the anterior end of this muscle the
nerve passes upward, and without any ganglionic enlargement on
its course, it joins the elongated ganglion underlying the osphra-
dium in the left anterior corner of the mantle-cavity. The
osphradial ganglion is also supplied, as is the case in Verita and
Neritina, by the symmetrical left branchio-pallial nerve, emana-
ting from the left pleural ganglion. This large nerve traverses
the columellar muscle and passes almost direct to the osphradium,
where it enlarges to form the above-mentioned ganglion. From
the ganglion a branch passes along the anterior border of the left
suspensory fold of the ctenidium and may be traced without
difficulty nearly to the tip of the latter organ. Another branch
passes backwards, nearly parallel to the columellar muscle. I
have not been able to trace this nerve in its entirety, but have no
doubt’ that it is the continuation of the supra-intestinal nerve,
and joins the visceral ganglion in the vicinity of the uropore,
thus completing the visceral loop. If this is the case the strepto-
neury is complete, as it is in Vertta and the larger tropical species
of Neritina.
1908. | ASPIDOBRANCH GASTROPOD MOLLUSCS. 829:
In order to avoid repetition of details I will pass briefly over
the rest of the nervous system of Septaria. In all essential
features it resembles the nervous systems of Verita and the
tropical Neritinz, which I shall describe in greater detail in the
subsequent part of this paper. I need only say here, because
Béla Haller has thrown doubt upon these points in his description
of the nervous system of Nerita ornata, that there is a well-
defined labial commissure in Septaria, and that I can find no
trace of transverse commissures, posterior to the main anterior
commissure, between the pedal cords in this genus.
The position and structure of the subpallial sense-organ or
osphradium in the Neritide has been correctly described by
Bernard (3) and Thiele, but the latter author throws doubt on
its homology with the true osphradium of other Mollusca, and
other authors give doubtful or incorrect descriptions of it. In
Septaria this sense-organ is easily distinguished in transverse
sections as a prominent ridge of epithelium running forward from
the anterior end of the left suspensory fold of the ctenidium
along the roof of the mantle-cavity and ending only a short.
distance behind the thickened anterior edge of the mantle. It.
lies almost in the angle formed by the union of the mantle with
the left columellar muscle, and its position at the inhalant side of
the mantle-opening is consistent with the function usually
attributed to an osphradium, that of a sense-organ for testin g the
quality of the water before it passes over the ctenidium. The
cells covering this ridge are higher, their nuclei are more closely
crowded together and stain more deeply than those of the
adjacent mantle epithelium. Under a high power of the micro-
scope the epithelial ridge can be resolved into three parallel
strips. The two outer strips (@-> and 6-ce in fig. 17) consist
solely of columnar epithelial cells with granular cytoplasm and
rather large nuclei. The free ends of these cells bear cilia which
in the groove shown on the lower side of fig. 17 are longer than
elsewhere. The central strip (6-0 in fig. 17) is largely composed
of the same elements, but its character of a sensory epithelium is
well shown by the presence of a number of attenuated sense-cells,
interspersed between the larger columnar cells. The nuclei of
the sense-cells are smaller and more elongated than those of the
columnar cells; their cell-bodies stain deeply in carmine, and in
many cases it can be seen that their inner ends are prolonged
into fine fibrils which traverse the thin layer of muscle-fibres and
connective tissue underlying the osphradium and pass inte the
osphradial ganglion. The size and position of this ganglion at.
the place where it is joined by the large branchio-pallial nerve
are shown in fig. 17.
The respiratory and circulatory systems have been worked out
in some detail by Lenssen in Weritina fluviatilis, and their
arrangement is similar in Septaria, but allusion must be made
to one or two points in which Lenssen’s account is defective.
The ctenidium in Septariu is an elongated triangular organ, its
free pointed extremity directed forward and to the right; its
830 PROF. G. C. BOURNE ON THE [Nov. 17,
base attached to the body-wall on the left side. It consists of a
flattened axial plate containing blood-spaces, with numerous
gill-lamelle running transversely across its dorsal and ventral
surfaces. The ctenidium is therefore typically bipectinate.
Unlike that of V. fluviatilis its posterior half is attached, on the
left side to the mantle close to its union with the columellar
muscle, on the right side to the lower surface of the kidney, by
a membranous suspensory fold. Thus the posterior part of the
mantle-cavity is divided into an upper and a lower chamber. A
large blood-vessel runs along each edge of the axial plate, and in
the: upper and lower wall of each vessel there is a stout band of
longitudinal muscle-fibres, which must serve as retractors of the
gill rand also assist in the circulation of blood through the gill-
lamelle. On the right-hand is the afferent and on the left the
efferent branchial venous sinus. The two do not communicate
with one another at the apex of the gill but only by the lacunar
passages in the gill-lamelle and the axial plate. The general
course of circulation in the gill is as follows :—Blood is brought
to the gills from the large venous sinus underlying the kidney
by the “afferent branchial sinus. This sinus does not com-
municate, as may be seen by inspection of fig, 20, with the
cavities of the axial plate, except at very rare intervals, but it is
in free communication above and below with the cavities of the
gill-lamelle, and its blood passes into these latter and circulates
through them. The cavities of the gill-lamellz and also that of
the axial plate are broken up by numerous trabeculee or partitions
passing from wall to wall and are bounded by a thin layer of
connective tissue containing a few muscular fibres. The margin
of each gill-lamella is somewhat swollen and contains a cavity or
vessel which, as far as I can ascertain, is continuous from one
end of the lamella to the other, and also is in free communication
with the irregular spaces below. At the opposite side of the gill
these marginal vessels open into the efferent branchial sinus.
The innermost cavities of the gill-lamelle open from place to
place into the system of lacune in the axial plate, and these,
uniting to form a large lacuna near the left edge of the axial
plate, also open at frequent intervals by large apertures into the
efferent branchial sinus. It 1s obvious that the blood on entering
the gill-lamelle from the afferent sinus may either take a direct
course to the efferent sinus by way of the marginal vessels of the
lamelle, or may traverse the lacune in the adaxial part of
the lamelle, pass into the system of lacune in the axial plate,
and thence be discharged into the efferent sinus.
The distribution of ciliated epithelium on the faces of the gill-
lamelle is of some interest. Each gill-lamella is an extremely
delicate plate of semilunar form, its straight inner margin
attached to the axial plate, its curved margin free and, as
described above, somewhat thickened. When one attempts to
separate the lamelle by the aid of needles one recognizes that
their central portions adhere very closely together, whereas thei:
1908. ] ASPIDOBRANCH GASTROPOD MOLLUSCS. 831
lateral portions are readily parted from one another. Whena
single lamella is separated out, stained and examined under the
microscope, it has the appearance shown in fig. 23, the dark
central tract with diverging horns being the expression of
numerous deeply stained and closely packed nuclei in this region.
In short, the epithelium covering the lamelle is not uniform.
‘The following arrangement can be determined in section :—The
lateral tracts of each face of each gill-lamella are clothed by a
cubical epithelium containing isolated or grouped gland-cells of
oval shape with clear contents. The epithelial cells of these
tracts (if they are ciliated at all, which I am inclined to doubt)
bear exceedingly short and fine cilia. The thickened margin of
the lamella always bears three or four “frontal” cells at its
extreme edge; these cells, as in the gills of Lamellibranchia,
carry a vide of short rather stiff cilia. External to them are a
few an devoid of cilia, and at the extreme ends of the lamellee
large gland-cells alternate with the epithelial cells of this region.
Sections through the dark median band with its two horns show
that this is a tract of more columnar cells, closely packed together,
with deeply staining nuclei, each carrying a tuft of very long
cilia which interlock with those of the adjacent lamella and are
the cause of the adherence noted above.
There are no supporting rods or skeletal bars, such as those
described by M. F. Woodward in Pleurotomaria, but, as shown in
fig. 21, the connective tissue underlying the epithelial cells is
thickened near the attachment of each gill-lamella to the axial
plate. There is some resemblance between the arrangement of
the lateral cilia in Septaria and other Neritidze and in Pleuro-
tomaria, and by parity of reasoning the ciliated tracts of the
former genus must differ from those of Lamellibranchia in the
same manner that Woodward has shown them to differ in the
case of the latter genus. It is interesting to note the structural
analogies of gastropod and lamellibranch gills. Im the case of
the Neritidee the ciliated tracts fulfil the same functions as the
ciliated discs of the Filbranchia. But their arrangement is
different. The cell-mechanism is the same, but it cannot be
doubted that it has been independently evolved in the two
groups, affording a good instance of the evolution of similar but
not identical structure in similar organs subject to similar
-conditious.
Lenssen, describing the gills of Veritina fluviatilis, has given
an incorrect account of the epithelium. He figures an almost
uniform covering of ciliated cells, and among them a few gland-
cells. I have found the same arrangement in WV. fluviatilis as in
Septaria, and Lenssen would appear either to have altogether
overlooked the ciliated tracts, or to have confused in a single
drawing and description the ciliated cells of the one tract and
the glandular cells of the other. It is curiously difficult to obtain
good preparations of the gills of V. fluviatilis, and if my attention
had not been called to the subject by the much more obvious
Proc. Zoo. Soc.—1908, No. LITI. 53
832 PROF. G. C. BOURNE ON THE [ Nov. 17,
arrangement of the cells in Septaria, 1 should have overlooked it.
As it is I have identified a simuar gill-structure in Verita plicata,
NV. variegata, Neritina gagates, and NV. longispinosa. The de—
scription given above is therefore characteristic for the ctenidium
of the Neritide, and will not be repeated in dealing with other
species.
The branchial nerve, derived from the ganglion underlying the
osphradium, runs through the suspensory membrane and along
the outside of the efferent branchial vessel in the free moiety of the
gill. This nerve has been noted by nearly all previous authors, but.
only Bernard has described a nerve running down the other side of
the gill, m connection with the afferent branchial vessel. Such
a nerve, embedded in the dorsal longitudinal muscle, is present in
Septaria and is shown in fig. 20,b67.n. As far as I can determine
from my sections it communicates with the nerve on the efferent
side by a slender connection at the tip of the gill. It is thickest
in the posterior part of the course, and passes into the right.
suspensory membrane in the direction of the visceral ganglion ;
but try as I might, I could not trace it through the suspensory
membrane to the ganglion.
The Circulatory System.
Lenssen’s account of the heart and blood-vessels in WV. fluviatilis
is in all essential features applicabie to Septaria. I need lay stress.
on a few points only. The ventricle in Septaria is thick and
muscular and is completely wrapped round the rectum. I wish
to emphasize this point, because some authors have stated that the.
ventricle is only par tially attached to the rectum in Neritide, and
have contrasted this with the conditions found in Haliotide and.
Trochide. Practically the supposed difference does not exist.
The blood from the efferent venous sinus passes directly into the-
left auricle, and at the point where the efferent sinus joins the
auricle a large pallial sinus, bringing back blood from the numerous
lacunze of the roof of the mantle-cavity, opens into it. This
pallial sinus has been recorded by Lenssen in JV. fluviatilis, but I
think he underrates its importance. In Septaria, at any rate, its
diameter is nearly equal to that of the efferent branchial sinus,
and it is connected with so extensive a system of blood-lacunz in
the mantle that there can be no doubt that the latter is a very
important auxiliary organ of respiration. The blood, therefore,
which reaches the ventricle by way of the left auricle has been
oxygenated either during its passage through the ctenidium or
through the mantle, but none of the pallial blood passes through
the ctenidium. The importance of the pallial circulation in
Septaria is further indicated by the relatively considerable size of
the right auricle. This organ, the relations of which may easily be.
traced in sections or by simple dissection, passes from that part of
the ventricle lying posterior to the rectum, across the pericardial
cavity, and is attached to the left body-wall just at the level of the.
1908. j ASPIDOBRANCH GASTROPOD MOLLUSCS. 833:
left posterior corner of the foot. Here it receives blood from two
venous sinuses: the one bringing back blood from the left posterior
region of the body-wall, the other from a considerable system of
lacune in the posterior part of the foot. The former of these two
sinuses runs in close connection with the posterior lobe of the
kidney, and it would appear that we have here an arrangement
whereby oxygenated blood from the mantle is also enabled to get
rid of its waste nitrogenous products before it is returned to the
heart. This is the reverse to what occurs in the case of the blood
passed through the ctenidium, which is first purified of its nitro-
genous waste matter during its passage through the sinuses of the
anterior lobes of the kidney and is afterwards carried to the
ctenidium by the afferent branchial sinus. The right auricle, like
the left, is covered by the glandular tissue of the pericardial gland.
As Septari ia has undergone a considerable amount of detorsion, it
is evident that the position of the right auricle, and its connection
with the left and posterior part of the body-wall and foot, is a
secondary phenomenon, due to its movement from right to left,
in the direction of the hands of a clock, during the. process of
detorsion. In fact, one can only use the term “right” in a
morphological sense, to indicate that this auricle w ould be on the
right if the typical gastropod torsion had been maintained. In
the genera Verita and Neriéina, as will be seen, the rudimentary
aut ieled is more distinctly on the Tight side, but even in these forms
posterior to the ventricle. The size of the right auricle varies
much in the Neritide. It is always present, but in some of the
typically marine forms such as WVerita peloronta it is so small and
unimportant that it might easily be overlooked, and indeed its
existence has been denied by Béla Halier. In the tropical
Neritine, many of which are semi-aquatic in habit, spending no
inconsiderable part of their lives on the roots of trees above low-
water mark, or even above high-water mark of neap-tides, the:
right auricle is larger and receives the same blood-supply as in
Septaria. The last-named genus is one that, according to all
accounts, has progressed further than any other Neritid in the
direction of a terrestrial life, living as it often does on stones in
the vicinity of waterfalls where it is only wetted by spray. One
might expect, therefore, that it would exhibit a more marked
tendency to the replacement of a branchial by a pallial respiration,
and such has been shown to be the case. It is of special interest
to observe the connection between a more highly developed pallial
respiration and the increased size and importance of the right
auricle, for, as I shall show in the second part of this memoir
dealing with the Helicinide, there is every reason to believe that,
in the last-mentioned family, in which the ctenidium is lost and
the respiration is entirely pallial (or, as it is called, pulmonary), the
single auricle that persists is the right and not, as has generally
been supposed, the left.
The courses of the main efferent ov arterial vessels proceeding
from the ventricle, and the venous sinuses in the foot and visceral
53*
834 PROF, G. C. BOURNE ON THE [Nov. 17,
cavities, are so similar in Septaria to those described by Lenssen in
Neritina fluviatilis, that it is not necessary to enter into any
further description of them.
The Hemocele and Colom.
The cavities in which lie the buccal bulb, the cesophagus,
radula-sac, and the coils of the intestine are, as is usually the case
in Molluscs, a vast blood-space or hemocele. In most Neritide
and in the Helicinide this space is largely filled up by a parenchy-
matous tissue, which from its structure and position appears to
be metabolic in function and probably serves for the storage of
reserve material. Where present it is aggregated along the courses
of the chief arterial vessels, and a certain amount of it is to be
found surrounding the anterior aortain Septaria. This metabolic
tissue, however, is never abundant in Septaria, and I postpone
a description of it to the section dealing with the WVerite and
Neritine. In Septaria, and indeed in all Neritide, the large
heemoceelic cavity does not extend as a wide space into the visceral
mass of the spire. In this region nearly the whole of the space
contained within the body-walls is occupied by the ovary, the
stomach, and the mass of the liver lying above and to the right
side of the stomach. It is only on the left side that some distinct
posterior prolongations of the hemoccele are visible surrounding
the intestine, the left and lower sides of the stomach, and the
radula-sac. The great anterior heemoccelic space surrounding the
pharyngeal bulb and the greater part of the coils of the intestine
is often referred to by writers on molluscan anatomy as the
anterior body-cavity, and allusion is sometimes made to a sort of
diaphragm or partition shutting off this anterior cavity from the
visceral cavity behind. In fact, there is no definite diaphragm or
dissepiment, and the appearance of one is due to a somewhat
complex union of muscular membranes connected with the kidneys
and genital ducts, but particularly to a large venous sinus with
muscular walls passing from the right towards the base of the
ctenidium. In this space lies the elongated visceral ganglion.
Theoretically the visceral heemoceele is to be regarded as a
continuous space, the hinder part of which is filled up by the
stomach and liver. This can be understood by reference to fig. 41,
illustrating a horizontal section through the ventral part of the
body of WVeritina gagates.
With regard to the coelom, Lenssen observes with much truth
that in Neritina fluviatilis it is the most extensive space in the
whole body. ‘‘ Kn evant,” he says, “elle (la cavité péricardique)
se prolonge jusqu’a la base de la branchie, et de la, s’élargissant
de plus en plus en arriére, elle s’étend sur toute la largeur du
corps et divise l’animal en deux portions bien nettes. Sa paroi
supérieure se confond avec la base du rein et se prolonge sous
Yutérus; elle sépare le rein dufoie. Sa partie inférieure, chargée
de pigment, enveloppe le massif formé par les circonvolutions de
1908. | ASPIDOBRANCH GASTROPOD MOLLUSCS. 835:
Vintestin et du sac radulaire dans la région antérieure du corps
et se prolonge en arriére jusqu’a I origine du rein.” This
description is tolerably exact, but the ramifications of the ccelomic
cavity are extremely difficult to make out in Veritina, and a much
clearer picture of the extent and relations of the coelom can be
obtained from the more symmetrical Septaria. In this genus, as
may be seen in fig. 4, the ecelom is, relatively to the size of the
body, a vast space extending from vight to left across the entire
width of the visceral mass.
Two divisions can be recognised, the per ‘icardial and the gonadial
celom. The former lies to the left : ; 1t is traversed by the bend
of the rectum, contains the heart, and communicates by the reno-
pericardial duct with the kidney. The gonadial division of the
ccelom lies on the right, and may be described in general terms
as intervening between the liver and ovary and the dorsal body-
eal in the region of the visceral mass (fig. 4, g.co.). Near the
anterior border of the visceral mass it Pane) a spacious cavity
extending downward on the right side of the body to the level of
the floor of the visceral hemocele, and here it enters into
remarkable relations with the oviduct. As shown in figs. 3, 4,
and 5, the oviduct, which pursues a nearly straight course forward
from the ovary to the accessory genital apparatus, crosses the
gonadial celom, and at this point opens into it by a distinct
but short and narrow oviduco-cceelomic duct, the details of
which are indicated in fig. 5. The duct is lined by a cubical
epithelium bearing fine cilia, and this, near its opening into the
gonadial division of the ccelom, is replaced by an epithelrum
contaiing closely crowded, deeply stained nuclei and bearing
longer cilia. The similarity between this oviduco-ecelomic funnel
and duct and the reno-pericardial canal of the left side is obvious,
and affords evidence of the former existence of a right kidney,
into which, as in other Rhipidoglossa, the ova were discharged to
find their way to the exterior by the right uropore. The full
significance of these relations will be dealt with further on: it
need only be said here that the existence of an oviduco-ccelomic
funnel, opening into a special division of the cclom, is not
parallelled in any other adult Gastropod, except the allied genera
Nerita and Neritina.
The pericardial division of the ccelom is of largesize : it extends
forward to the base of the ctenidium and is continued for some
little distance along the left edge of the latter, parallel to the
efferent branchial vessel, as a narrow diverticulum. Posteriorly
the pericardial cavity extends along the left side of the stomach
nearly to the pointed extremity of the visceral mass. The
gonadial division of the ccelom is a more irregular cavity. In the
anterior part of the visceral mass, immediately behind the
accessory genital organs, it is of considerable vertical depth,
reaching from roof to floor of the visceral sac in front of the
liver and ovary. A little further back it extends over the liver
and ovary and to the right of the latter as far as the floor of the
836 PROF. G. C. BOURNE ON THE [ Nov. 17,
visceral sac, but on the left it appears in sections only as a narrow
cleft reaching as far as the rectum. Posteriorly the gonadial
coelom gradually becomes smaller and smaller, as the liver and
ovary project further into it, and eventually it is reduced to a
comparatively narrow space between the rectum above and the
vadula-sac and liver below. In the region of and behind the
ventricle the two divisions of the cceelom communicate freely with
one another by a large slit-like passage which, as shown in fig. 4,
c.ap., lies dorsad of the stomach. The pericardial floor leading to
it is deeply pigmented.
The EKxcretory System.
The kidney in Septaria is more easily studied than in any other
member of the Neritide, and my investigations confirm the
correctness of Lenssen’s somewhat brief account of this organ in
Neritina fluviatilis. The kidney of Septaria is an elongated organ
lying transversely across the posterior third of the body, closely
attached to the left and anterior side of the rectum where the
latter passes across from the pericardium to join the complex mass
formed by the accessory genital organs (fig. 1, A.). The left and
posterior end of the kidney lies close below the dorsal body-wall
(fig. 4, K.); its right and anterior moiety passes into the dorsal
wall of the mantle-cavity. Throughout the whole of its extent it
is in close relation to the pericardial division of the ccelom.
The kidney may be described in general terms as a tubular organ
bent upon itself in such a way that its two ends are anterior and
open respectively into the mantle by the uropore and into the
pericardial division of the ccelom by the reno-pericardial duct.
Only the dorsal limb of the tube is glandular, its cavity being
traversed by a number of deep infoldings of its lateral walls, which
in turn give off secondary folds (fig. 18, A’.). There is no question
of an acinous structure such as has been described in Verita ornata
by Béla Haller (20). As Lenssen describes for Veritina fluviatilis,
the partitions which cross the cavity of the glandular part of the
kidney ot Septaria, though they may sometimes appear in sections
to cross from wall to wall, are really infoldings of the lateral walls
terminating in free edges within the cavity of the kidney. The
whole of the glandular part is surrounded by a system of blood-
sinuses (fig. 19, 6.s¢.), connected with smaller sinuses running in
the partitions, and a distinct sinus or vessel may often be recog-
nized along the free edges of the latter. The epithelium clothing
the walls and partitions of the glandular part of the kidney is
non-ciliated, but is not of uniform character throughout. In the
right and anterior moiety of the kidney the cells are dilated by
the presence of a large transparent vacuole at their free ends ; the
nucleus lies in the basal part of the cell, stains faintly, and is
surrounded by a small amount of cytoplasm. In the left and
posterior moiety of the kidney the cells are more prismatic, are
less vacuolated, their nuclei stain more deeply and are situated
1908. | ASPIDOBRANCH GASTROPOD MOLLUSCS. 837
nearer to the middle of the cell-body. The different appearance
“of these two regions of the glandular part of the kidney, as seen
under a low power of the microscope, is represented in fig. 18.
It may be thought that this apparent distinction is due to a
difference in the secretory activity of the cells, but I have found
it to be so constant a feature in the different species of Verita
and Neritina that I have studied that I think there must be
some functional differentiation between the two regions. The
glandular part of the kidney opens posteriorly into the non-
glandular part or bladder, the latter being a large flattened sac
of irregular form running forward between the glandular part
and the pericardium. It is this sac which Perrier described as a
closed cavity interposed between the kidney and the pericardium,
and Béla Haller was scarcely more correct in identifying it as the
reno-pericardial duct. Anteriorly the bladder expands towards
the right (fig. 19) to form a diverticulum lying below and behind
the venous sinus in which the ascending portion of the sub-
intestinal nerve is contained. The tough membranous walls of
this diverticulum of the kidney-sac and of the venous sinus form
together with the anterior wall of the gonadial ccelom the so-called
diaphragm referred to on p. 834. The left anterior corner of the
bladder passing below the glandular part turns upward as a
narrow passage lying between the latter and the base of the
‘etenidium, and twisting over to the right opens into the lower
chamber of the mantle-cavity by the uropore below and to the
jeft of the afferent branchial sinus. By far the greater part of
the bladder is lined by a non-ciliated flat epithelium, but in the
duct leading to the uropore this is replaced by a ciliated epithelium
composed of very long attenuated and transparent ciliated cells. The
reno-pericardial canal, as is the case in all Neritide, is very large.
It opens, as shown in fig. 18, rp.c., into the anterior end of the
glandular part of the kidney; thence running to the left and
posteriorly, it becomes closely attached to the wall of the duct of
the bladder in the vicinity of the uropore, and twisting downwards
and inwards it describes an @-shaped curve and opens by a wide
ciliated aperture into the base of the anterior diverticulum of the
pericardial cavity referred to aboveas extending forward alongside
the efferent branchial sinus. The epithelial cells of the reno-pert-
cardial duct are very large and each bears a tuft of long stiff cilia.
Lenssen has given a figure of this characteristic epithelium in
Neritina fluviatilis, and describes it as a good example of a
discontinuous epithelium. His figure is a good representation of
the appearance usually seen in sections, but from what I have
seen in some well-preserved specimens I think that the apparent
discontinuity is due to contraction produced by reagents.
It follows from the above description that the kidney of Septaria
(the kidney of other members of the Neritid is similar) is not a
simple glandular sac, but is composed of a glandular and non-
glandular part the relations of which are very similar to those ob-
served in the lamellibranchiate kidney. In Septaria bougainviller
838 PROF. G. C. BOURNE ON THE [ Nov. 17,.
there is no communication between the anterior ends of the
glandular and non-glandular parts. The exception is shown in
fig. 19, representing part of a horizontal section through Septaria
depressa. In this the terminal portion of the non-glandular part
is seen to communicate with the glandular part by a small but
perfectly distinct orifice in the vicinity of the renal aperture of
the reno-pericardial duct. I can find no trace of such an orifice
in S. bougainville?, but it occurs in several other Neritide. It is
not an accidental rent in the wall, the epithelium passing in
unbroken continuity round the lips of the orifice.
It is now so generally understood that the kidney of the
Neritide is the post-torsional left, that it is not necessary for me
to insist upon the further evidence in support of this opinion
derivable from the facts just mentioned. But it is perhaps.
necessary to allude to the question as Fleure (17) in a compara-
tively recent paper has attempted to prove that the single kidney
of the Neritide and Pectinibranchia is the left post-torsional
and corresponds with the large functional left kidney of most
Rhipidoglossa. Unfortunately for his argument his paper 1s
followed by that of Miss Drummond (13), in which the fact
previously insisted on by von Erlanger (14) is placed beyond all
doubt, namely that the existing kidney of Paludina (and
presumably of other Pectinibranchs) is the post-torsional left.
The rudiment of the post-torsional right kidney becomes converted
into the genital duct in the course of embryonic development
It is clear, from the presence of the oviduco-ccelomic funnel, that.
the same thing has happened in the Neritide.
The Generative Organs.
Gilson (18) was the first to give an intelligible account of the
female generative organs of Veritina fluviatilis, and their structure
was subsequently worked out in great detail by his pupil Lenssen.
More lately Thiele has described these organs in several species of
Neritide, including Septaria (Navicella) parva and suborbicularis.
A further description would therefore seem superfluous, were it
not that Septaria differs in some not unimportant particulars
from WVeritina, and Thiele’s account of the former genus is little
more than a brief note, omitting histological detail, and, moreover
defective in at least one very important particular. Moreover, as.
a result of a comparison of Lenssen’s and Thiele’s work with my
own and of an attempt to homologize the different cavities, ducts,
and glands in the different forms that I have examined, | have
come to conclusions somewhat different from those of the two
authors named, and have to suggest a new nomenclature for the
different parts. Gilson and Lenssen have shown that the female
ducts of Neritina fluviatilis ave diaulic; Thiele has made the same
statement for Verita and Septaria. A veference to the diagram
(fig. 3) will show that the female ducts of Septaria are triaulic
There is the large ovipository aperture (Ov.ap.), through which the.
1908. ] ASPIDOBRANCH GASTROPOD MOLLUSCS. 839)
eges enclosed in their egg-shells are extruded ; the copulatory or
vaginal aperture (vag.ap.), through which the sperm of the male
is received ; and a third minute aperture (ap.de.) situated further
back, whose function I cannot determine: I shaJl refer to it as
the aperture of the ductus enigmaticus. This third aperture is
not present in Veritina fluviatilis nor in the marine species of
Nerita, but is present in all the fresh- or brackish- water tropical
species hitherto classed as Veritina. To begin with a description
of the diagram, fig. 3. The ovipository aperture leads into a vast
pouch with thick glandular walls. This has been called the
“uterus” by Gilson and Lenssen, the ‘‘shell-gland” by Thiele. As.
there is no doubt that the egg-shell is formed from the secretion
of the glands opening into this cavity the latter name is appro-
priate, but it will avoid confusion if I borrow a name from an
analogous structure in the Platyhelmia and call it the ‘“‘ ootype.”
A flattened saccular diverticulum (cry.s.), lying on the right
side of the terminal part of the ootype, opens into the latter not
far from the ovipository aperture. In the species | examined it.
is of small size and does not contain concretions or foreign
bodies, but it is clearly homologous with the “ poche a cristaux ~
of Lenssen, and I shall refer to it as the erystal-sac. In Nerita
and Weritina it is relatively large and filled with spherical
crystalline bodies. Thiele calls this sac the uterus, but the name
is inappropriate, for there is no evidence whatever that the ova
are passed into it.
The relations of the ootype may be studied in the series of
transverse sections (figs. 6-12). Anteriorly it lies above and
partly embraces the rectum. In this region its walls are very
thick and glandular, its cavity large, crescentic, and simple.
Further back it lies to the right of and apart from the rectum,
and its cavity becomes more complicated in form. The whole
organ is spirally twisted, so that the concavity of the crescent,
which was at first directed downwards, is in fig. 9 directed
upwards. In fig. 10 the left-hand corner of the cavity is seen to.
be prolonged into a diverticulum, which in fig. 11 turns sharply
to the right and runs back parallel to its former course. From
this, which may be described as the terminal part or fundus of the:
ootype, two passages are given off in different directions. The
one, which I shall call the egg-duct, leads to the oviduct and
through this to the ovary. ‘The other establishes a connection
with the vagina and ductus enigmaticus and serves for the admis-
sion of spermatozoa to the fundus of the ootype.
The egg-duct (eg.d. in figs. 10-13) is characterized by the
different forms of glandular tissue constituting its walls. Its.
opening into the cavity of the ootype is embraced by a glandular
thickening. distinguishable from the fact that its cell-contents do
not stain in any of the ordinary dyes: this I call the clear gland.
As shown in figs. 11, 12, 13, it is largely situated in a tongue-
shaped projection of the right-hand wall of the fundus, which
appears to function as a valve guarding the passage from the
‘840 PROF. G. C. BOURNE ON THE [ Nov. 17,
fundus into the egg-duct. The dorsal side of the egg-duct is
capped by a considerable glandular mass (m.gl.), whose contents
stain deeply in hematoxylin : it may therefore be identified as a
mucous gland. Beyond this is a narrow ring of glandular tissue
whose cell-contents stain bright rose-red colour in eosin and
carmine dyes: for this reason I shall refer to it as the rose-
coloured gland (figs. 12-14, re.gl.). Beyond the rose-coloured
gland the egg-duct dilates to form a distinct chamber with thick
glandular walls. This chamber corresponds to the “ premiére
ampoule ” of Lenssen ; I shall call it the thalamus (figs. 14, 15, 16,
th.). It occupies the right-hand posterior corner of the genital
complex, and the gland surrounding it may from its position and
staining-properties be identified as the vitelline gland. The
oviduct, a narrow tube lined by a columnar ciliated epithelium,
emerges from the lower left-hand corner of the thalamus, runs
towards the left, is thrown into a few convolutions, then turns
sharply backward and downward and enters the connective-tissue
layer forming the floor of the large right extension of the gonadial
celom. In its passage below this ccelomic space it gives off a
short branch to the right, which immediately opens into the
celom by a ciliated oviduco-ccelomic funnel (figs. 3 & 4, od.cf.).
Beyond the oviduco-cceelomic funnel the oviduct branches
repeatedly ; the branches subdivide and end in a number of
claviform acini lined by a germinal epithelium and containing ova
in all stages of development. The ovary, thus constituted, is a
fairly extensive organ lying to the right of the liver and spreading
for some distance over its dorsal surface.
The second passage leading out of the fundus of the ootype isa
narrow thin-walled tube which passes to the right and immediately
dilates to form a sae of considerable size lying between the
posterior end of the ootype and the rectum. As this sac always
contains free spermatozoa in greater or less abundance, it may be
called the sperm-sac. Its walls are thin, usually much folded, and
lined by a simple cubical ciliated epithelium without any trace of
glandular structure. In longitudinal section it is seen to be
U-shaped.
From the bend of the U a short duct is given off posteriorly.
It has thick muscular walls, and ends in a thick-walled dilatation,
filled with spermatozoa. This is the receptaculum seminis
(=spermatheca of Gilson). The limbs of the U are prolonged
forward as two narrow ducts, which acquire thick muscular walls
and open separately into the mantle-cavity. The outer or right-
hand duct corresponds to the ‘ connecting duct” of Gilson and
Lenssen. I shallcallit the vaginal canal. ‘The inner or left-hand
duct is not represented in WNerita or in WNeritina fluviatilis,
and as its function is obscure I have named it the ductus
enigmaticus. Thetwo ductsrun forward close to one another and
to the rectum. The ductus enigmaticus is relatively short and
straight, and eventually opens into the mantle-cavity by a minute
pore situated on the ventral side of the genital complex and some
1908. | ASPIDOBRANCH GASTROPOD MOLLUSCS. 841
distance posterior to the anal and ovipository apertures (fig. 6,
ap.de.). The vaginal canal runs parallel to the ductus enigmaticus
as far as the opening of the latter, and then bends abruptly back-
ward. Its lumen becomes very narrow, and its muscular wall
relatively thick; after a short course backward it opens by a
minute pore into a sac (sp.s.), which corresponds to the copulatory
vesicle of Gilson and Lenssen, the receptaculum seminis of Thiele.
As may be seen in the diagram, it is the dilated posterior end of
the vagina (vag.), and is of small size in Septaria borbonica and
bougainvillet. But in the tropical Weritinw, in Nerita, and,
according to Thiele, in Septaria parva it is relatively of enormous
size and invariably contains a number of spermatophores of
complex structure. Hence I shall call it the spermatophore-sac,
although I have not found a trace of spermatophores in any of the
specimens of Septaria that I have examined. 'The lumen of the
spermatophore-sac gradually diminishes anteriorly and passes
without sensible alteration of structure into the vagina, which
opens into the mantle-cavity by the vaginal aperture situated on a
prominent papilla some little way in front of the anus.
Gilson has given a clear account of the process of fertilization
in Neritina fluviatilis. Owing to the small size of the spermato-
phore-sac it must be somewhat different in Septaria borbonica
and bougainvillei. The spermatozoa must be deposited by the
male in the vagina and must travel by way of the vaginal canal to
the receptaculum seminis, where they are disposed, just as Gilson
describes, in a very regular manner, all heads turned towards the
centre of the vesicle and all tails directed outwards. At the time
of impregnation the spermatozoa must be passed, by contraction
of the muscular walls of the receptaculum and its duct, into the
fundus of the ootype and thence into the egg-duct, where the ova
are fertilized. The function of the ductus enigmaticus can only be
guessed at. It may serve to admit water into the cavity of the
sperm-sac, or contrariwise may serve for the expulsion of fluids
accumulating in the sperm-sac.
Histology of the Genital Ducts.
From what precedes it will have been gathered that there are
five different kinds of glands on the course of the ootype and
ege-duct, viz., the vitelline gland, the rose-coloured gland, the
mucous gland, the clear gland, and the ootype or shell-gland.
Similar glands occur in the same positions in all the Neritide I
have studied, including Veritina fluviatilis. The histology of the
shell-gland of the last-named species bas been correctly described
by Lenssen, but he gives a very summary, and, in the case of the
vitelline gland at least, a somewhat incorrect account of the
remainder. The important thing is that all the glands are of
the same fundamental structural plan, and only differ from one
another in the proportions and staining-properties of the cells and
the secretions produced by them. It may therefore be inferred that
842 PROF. G. C. BOURNE ON THE [ Nov. 17,
the community of structure is due to community of origin ; and if
it can be shown that there is a transition between the secretory
epithelia of the glands and the epithelia of other regions, there is
a presumption in favour of the view that the glands are derived
from those other epithelia. A transition can be demonstrated
between the epithelium of the mantle-cayity and that of the
shell-gland. The epithelium lining the mantle-cavity differs con-
siderably in different parts, but im the neighbourhood of the
ovipository aperture it consists of fairly high columnar ciliated
cells, among which are numerous gland- cells with granular contents,
which in Septar ia stain bright. green in picro-indigo-carmine.
This epithelium is continued over the lip of the “ovipository
aperture into the terminal part of the lumen of the ootype, and,
extending further down on the right side than elsewhere, forms
the lining of the erystal-sac. Elsewhere it quickly undergoes a
change: the gland-cells disappear, and the ciliated cells increase
in length, become attenuated, and are separated by considerable
intercellular spaces (fig. 24). This simple ciliated epithelium is
continued downward for some little distance, and only gradually
becomes complicated by the appearance at first of a few club-
shaped gland-cells lying between the ciliated cells. The gland-
cells contain large granules, highly eosinophilous, or staining
bright green in picro-indigo-carmine. The gland-cells soon
become more abundant, and some of them, while retaining their
connection with the surface, tend to take up a deeper position.
Their swollen inner ends, containing the nucleus and most of the
cytoplasm, pass through the thin layer of muscle-fibres under-
lying the ciliated epithelium and embed themselves in the
surrounding connective tissue, their distal ends being drawn out
into fine tubes which pass between the ciliated cells and open into
the cavity of the ootype. In good preparations the walls of these
tubes are quite distinct, and each tube contains a single row of
eosinophilous granules, but swells out in the intercellular space
between the ciliated cells, so that it appears to terminate in a
claviform vesicle distended with granules (fig. 25). A little
further down the gland-cells increase in number and form groups,
and these groups passing into the surrounding connective tissue
form at first shorter, but in the greater part of the ootype
relatively long, club-shaped masses surrounding its cavity. ‘The
club-shaped masses have the appearance of and have been
described as crypts, but are not to be regarded as such, for what.
appears to be the cavity of the crypt is occupied by the ‘tubes, and
there is no lumen into which the secretion is discharged, but each
cell has its own duct opening on the surface. This may readily
be seen in transverse sections of the so-called crypts in good
preparations. It would be more correct to describe each group of
cells as a bunch, the deeper cells having very long hollow stalks.
and the more superficial cells shorter stalks; all the stalks pass.
between the ciliated epithelial cells, become slightly swollen, and
open on the surface. The manner in which this somewhat.
1908. j ASPIDOBRANCH GASTROPOD MOLLUSCS. 843
elaborate arrangement is derived from a comparatively simple
mixed ciliated and glandular epithelium is very well shown in the
terminal chamber of the male ducts of Verita (fig. 52), Precisely
the same fundamental structure is found in all the glands on the
course of the female ducts. In the ootype gland the cell-bodies are
coarsely granular and stain deeply in hematoxylin, the granules
of secretum are highly eosinophilous, and the ciliated cells are
moderately long. In the “ clear gland” the cytoplasm of the
gland-cells is scarcely granular, is not stained appreciably by any
of the ordinary dyes, the “stalks” or ducts of the cells are
relatively large, their contents clear and unstained by reagents.
The ciliated cells are somewhat widely spaced, and conspicuous
because they are not hidden by granules in the secreting ducts
(fig. 27).
In the “mucous gland” the bunches of glandular cells are
large ; the gland-cells are dark and granular and their cytoplasm
seus deeply with hematoxylin and picro- indigo-carmine. ‘The
secretory granules are not so large as in the uterine gland, are
of unequal size, and as they are not eosinphilous, but stain blue
with picro-indigo-carmine and deeply with hematoxylin, they are
probably mucinogenous. The ciliated epithelial cells betw een
which the unicellular ducts run are very much elongated.
In the “ rose-coloured gland” the bunches of gland-cells are
rather small; their evtoplasm stains rose-pink with picro-indigo-
carmine, carmine, or eosin; they are not granular, and their
ducts contain a non-granular coagulum which is highly eosino-
philous. The ciliated cells are short.
In the “ vitelline gland ” (fig. 28) the cell-bunches are of
moderate size ; the gland-cells have a reticular cytoplasm staining
faintly with picro-indigo-carmine or hematoxylin; the secretory
granules ave small, of equal size, and faintly stained by the
reagents mentioned ; the ciliated celis forming the boundary
epithelium are clearly defined and of moderate length.
It is evident that, although it isnot possible to attribute precise
functions to the different glands, they are to be regarded morpho-
logically as differentiations of a tract of mixed glandular and ciliated
epithelium, the histological characters of which are such that it is in
the highest degree probable that it has been formed as an invagi-
nation of the mantle-epithelium—that is to say, of the ectoderm.
The histology of the vagina, vaginal duct, sperm-sac, and ductus
enigmaticus is quite different. “These organs are non-glandular,
and are all lined by a very similar ciliated cubical epithelium,
which at first sight does seem to be very similar to the epithelium
of the oviduct, and different from that of the mantle- -cavity. But
a careful examination with high powers of the microscope shows
that they are different. In the oviduct the cytoplasm of the cells
is differentiated to form a distinct refractive external border, the
cilia ave longer and stouter, the nuclei are more elongated and
stain more intensely than is the case in the bursa copulatrix and
the ducts leading from it. Moreover, when the vaginal duct is
844 PROF. G. C. BOURNE ON THE | Nov.47,
traced to its aperture, which in Septaria lies some way in front of
the anal and ovipository apertures, the epithelium of the duct.
is seen to pass without any distinction of histological character
into the epithelium of the adjacent part of the “mantle- -cavity,
which latter is not in this place glandular, as it is close by the
anal and uterine orifices, but is a simple ciliated columnar
epithelium resembling the epithelium of the terminal portion of
the vaginal duct in the minutest particulars. The ductus enig-
maticus opens into a region of the mantle in which the epithelium
has been greatly moditied by the abundant development of long
beaker- shaped mucous cells, and in this case the transition fen
the epithelium of the duct to that of the mantle is abrupt (fig. 6).
From a consideration of these facts I am inclined to the opinion
that the whole of the sperm-sac and its two ducts is also formed
as an invagination of the mantle-epithelium, and that therefore:
the whole of the accessory organs contained in the genital complex,
with the exception of a short length of the oviduct, are ectodermal
structures secondarily attached to the oviduct ; and in this I differ:
from Thiele (39), who regards the whole of the vagina and sperm-
sac (which he calls the receptaculum seminis) as the representative
of the right kidney of the Neritidee. There can be no doubt that
the distal portion of the oviduct represents a part, probably the
terminal part or duct, of the right kidney. The existence of the
oviduco-ceelomic funnel is sufficient evidence of this homology..
But in my opinion the limit between kidney derivative and
ectodermal derivative is indicated in Septaria by the opening of
the oviduct into the thalamus. Here there is an abrupt change
in the histological character of the epithelium, and a reference to
figs. 8, 13, & 16 shows that the thalamus and the egg-duct.
intervene between the oviduct and the sperm-sac. I have given
strong reasons for believing that all the glands of the thaiamus and
ege-duct are derived from the ectoderm; and if Iam right this.
cir ‘cumstance militates against Thiele’s view that the spermato-
phore-sac represents the right kidney, for it can hardly be.
maintained that ectodermal structures have pushed their way into-
the primitive kidney and divided it into two widely separated
parts, one opening to the exterior, the other communicating with:
the ccelom by a ciliated funnel.
I am unable to give a description of the male organs of Septaria,
as all the specimens that I have examined by means of sections.
were females.
From an inspection of fig. 4 it might be inferred that the-
extensive tubular gland lying in the dorsal body-wall to the right.
of the rectum, and therefore occupying on the right side of the
body a position similar to that occupied by the kidney on the left,
is a representative of the glandular part of the right kidney.
Such an inference, however, cannot besustained. The histological’
characters of the gland in question are indistinguishable from
those of the hypobranchial mucous gland of other Rhipidogloss
(e. g. Fissurella). The anterior lobe of the gland lies immediately-
1908. | ASPIDOBRANCH GASTROPOD MOLLUSCS. 845.
behind the genital complex—-some of its tubules are seen in
figs. 15 & 16,—and its duct issues from the anterior lobe, passes.
ventrad of the receptaculum seminis, and opens into the right-
hand side of the mantle-cavity just below and to the right of the
oviduco-ccelomic funnel. It will be observed that the gland in
Septaria lies on the right side of the rectum, and cannot therefore
be the exact homologue of the hypobranchial gland of the
Trochide, which hes to the left of the rectum, between the latter
and the left ctenidium to which it is related. It must rather be
regarded as the homologue of the right hypobranchial gland of
the dibranchiate Rhipidoglossa, and it seems probable that it
represents the additional gland of the right side described by
M. F. Woodward (41) in Plewrotomaria. The right hypobranchial
(or additional hypobranchial) gland should be related to the right
ctenidium, but this has apparently disappeared in the Neritide..
I cannot find any vestige of it in Sepéaria, but in various species
of Verita and Neritina there is a small vascular organ projecting
into the mantle-cavity close to the aperture of the hypobranchial
mucous gland. This has been described in Neritina fluviatilis
by Lenssen under the name of the “organe creux,” and I shall
show in the latter half of this paper that partly on account of its
relation to the hyprobranchial gland there are good reasons for
regarding it as the vestige of the right ctenidium.
Genera NeritA Adanson and Neririna Lamarck.
It would be possible to give a detailed account of numerous
minute differences in the alimentary tract, minor branches of the
nervous system, excretory organs, &c. in the various species of
Nerita and Neritina that I have studied; but the enumeration
of these details would be not only tedious but unprofitable. . I
have satisfied myself that in all essential features of the-
anatomy of the alimentary tract, nervous system, respir atory and
circulatory systems, and excretory organs, the various species.
enumerated below are so similar to one ‘another that they may be
included in a single description. Moreover, their main anatomical
features are so like those of Septaria that I may spare the reader
the trouble of perusing a mass of detail which would differ only
in unessential matters from what has already been described in.
the last-named genus. It is otherwise with the generative ducts,
especially the female ducts. I shall have to point out that the
specimens I have had the opportunity of examining fall into
three groups, defined by the constitution of the genital ducts.
The first group comprises the marine species unquestionably
belonging to the genus WVerita, as defined in conchological works.
.The second group comprises the tropical species hitherto classed
in the genus Veritina and the genus Septaria. ‘The third group
comprises the European WVeritina fluviatilis, and to this must
probably be added the various European species of Veritina ; but
I cannot say anything definitely on this subject, for I have not.
846 PROF. G. C. BOURNE ON 'THE [Nov. 17,
yet had the opportunity of studying the anatomy of the European
species other than fluviatilis.
I was at first disposed to arrange these three groups as different
genera, but have hesitated to do so, because I have perceived
that it is impossible to make a reconstruction of the family
Neritidz on the basis of the slender amount of material at my
disposal. I can only indicate the conclusions I have come to and
express an opinion upon the probable relationships within the
family, leaving to future workers in this field the task of testing
the validity of my suggestions.
T am inclined to think, with Moquin-Tandon (28), that the
species hitherto classed in the genera Werita, Neritina, and
Septaria, agreeing as they do in all fundamental anatomical
features, should be grouped together in a single genus JVerita.
In this genus I recognize four sections—as a result of further
investigations more will probably have to be added,—Werita, s.s.,
Paranerita, Septaria, and Neritina. The section Verita com-
prises (so far as the present state of our knowledge permits a
definite statement) all those truly marie species hitherto classed
in the genus Verita. The section Paranerita includes the tro-
pical brackish- and fresh-water forms hitherto classed in the
genus Veritina, but which differ from Neritina fluviatilis in the
arrangement of the female ducts, in the presence of a distinct
“supra-intestinal nerve, and other minor features. The section
Neritina includes Weritina fluviatilis and probably ali the
European, Mediterranean, and Western Asiatic species of
Neritina included in the section Zheodowus of Montfort. The
subgenus Seplaria is equivalent to the genus Septaria ot
Férussac. Though I do not regard this as more than a pro-
visional arrangement, I shall make use of it, for clearness’ sake, in
the following part of this paper. It will be found, I believe,
that as our knowledge of the Neritidee advances, the various
species will fall into a number of geographical groups and sub-
groups, each group including marine, estuarine, and fluviatile
forms, but the consideration ‘of this sug ggestion must be left to
the latter part of this paper.
Adopting the provisional nomenclature suggested in the
preceding paragraph, the material at my disposal has been as
follows :-—
NVerita, 8.8.
AV. peloronta Linn. This, the well-known “ Bleeding-tooth,” is a
West Indian species.
iV. plexa Chemnitz. This species is widely distributed in the
Indian Ocean.
NV. lineata Chemnitz. From the Malaysian Islands and N.-
Australia.
(The above formed part of the spirit-collections of the British
Museum of Natural History, and I am indebted to Mr. E,
A. Smith for the opportunity of studying their anatomy.)
4908.] | ASPIDOBRANCH GASTROPOD MOLLUSCS. 847
LV. plicata Linn. This species is widely distributed in the Indian
and Pacific Oceans. My specimens were obtained, through
the kindness of Mrs. G. B. Longstaff, from Ceylon.
NV. melanotraga KE. A. Smith. Specimens of this Australian
species were kindly collected for me by Mr. Geottzey W.
Smith, of New Sales, Oxford,
Paranerita.
NV. variegata Lesson. From the East Indies and Polynesia. I
am indebted to Mr. K. A. Smith for specimens of this species,
which is synonymous with V. gagates Morch.
NV. gagates Lam. From Mauritius.
NV. longispina Récluz. From Mauritius. My specimens of the
last-named two species formed part of the collections of
the Oxford Museum, and in the same collections I found a
number of fairly well-preserved spirit-specimens of an
unknown species from Fiji, which I could not determine
because the animals had been extracted from their shells and
were not accompanied by any note of identification.
Neritina.
NV. fluviatilis Miller is common in the Isis and Cherwell and in
the smaller streams near Oxford.
In dealing with the general anatomy of these species I shall
-chiefly occupy myself with a description of those features in
which they differ from one another, but it will also be necessary
to enter into some details about organs in which they resemble
-one another, but in respect of which there is disagreement among
previous authors.
EHaternal Features.
Septaria, as has been shown, has undergone so much reduction
of the visceral spire that it has acquired a secondary external
symmetry, but the three sections Verita, Paranerita, and Nert-
tina vetain to a much greater extent the primitive gastropod
asymmetry. It is obvious, however, that they are tending
towards a secondary external symmetry, the visceral spire being
relatively small and making scarcely more than half a turn.
The shell-muscle or columeliar muscle is a striking feature in the
Neritide. It is always paired and subsymmetrical, Both muscles
are coarsely fasciculated; that of the right side is somewhat the
stouter, that of the left side somewhat longer in an antero-
posterior direction, The two muscles are attached to the right
and left inner surfaces of the shell within the area, and their
impressions are not easy to see in those shells in which the area is
strongly toothed or highly developed. The muscle-fibres of each
side converge downwards and inwards from their surfaces of
attachment “to the shell and pass into the powerful muscular
mass forming what is really the anterior end of the opercular
Proc, Zoot. Soc,—1908, No. LIV. 5A
848 PROF. G. C. BOURNE ON THE [ Noy. 17,.
lobe, but in contracted specimens this end always appears to be
posterior in position. The subequal development of the right.
and left columellar muscle is without doubt related to the
peculiar development and functional importance of the semilunar:
operculum, the straight and morphologically anterior edge of which
works against the anterior edge of the so-called area in a hinge-
like manner. When the animal is fully extended the operculum
is turned back and pressed close to the flat surface of the area ;
and an examination of a relaxed specimen or of a living Weritina.
fuviatilis shows that it is a mechanical necessity that there
should be an equal pull at both ends of the semilunar operculum
in order to bring about an effective closure of the aperture when
the animal retreats into its shell. There can be no doubt that.
the operculum is a very important factor in the economy of the
Neritide, and that, in the course of evolution, its growing
importance has had much to do with the tendency to reduction
of the visceral spire exhibited by members of this family. It.
can hardly be doubted that the physiological effect of a well-
developed left shell-muscle, exerting a pull upon the organs of
the left side every time that the animal withdraws itself into its.
shell, will be to draw those organs over from right to left and
thus to uncoil the typical gastropod spiral in opposition to the.
action of the right muscle. When both muscles are equally
developed and inserted symmetrically on the interior of the shell,
as is the case in existing Neritide, their joint action must tend to
the establishment and maintenance of a symmetrical arrangement.
of such organs as are affected by their action, and we find, in
fact, that the disposition of the two muscles produces an
appearance of symmetry in the region of the head and foot,.
which to a certain extent affects the organs of the pallial complex
and of the proximal part of the visceral mass. It is known that.
in Fissurella the secondary symmetry of the adult is acquired in
the course of development by the relatively great growth of the-
foot, the formation of a shell-muscle on the left as well as on
the right side of the body, and the subsequent rapid growth
of the right side, causing a shifting back of organs from right.
to left. But in this case the increased functional importance of
the sole of the foot and the loss of the operculum and opercular-
lobe have been the main factors in determining the final relations
of foot, shell, and visceral mass. In the Neritide different.
conditions have prevailed. The sole of the foot remains relatively
small; the operculum and opercular lobe become large and
important, and by their presence prevent any posterior union of
the right and left shell-muscles; the visceral spire remains:
connected with the cephalo-podial regions by a relatively narrow
pedicle. he different lines of evolution are well illustrated by
Septaria, in which the sole of the foot has enlarged so much that
it rivals that of a Fisswrella, a Patella, or a Capulus in size and
importance, and the shell has become almost patelliform, but the
operculum, though functionless, retains its place between the-
1908. } ASPIDOBRANCH GASTROPOD MOLLUSCS. 849
posterior part of the foot and the visceral mass, and the last+
named is interposed between the hinder ends of the elongate:
and perfectly symmetrical right and left shell-muscles. Of course,
the analogy between the Fissurellide and Neritide is incomplete:
in many respects. The ancestor of the latter family must have.
suffered great reduction in the organs of the post-torsional right.
side, such as the ctenidium, before any tendency to detorsion
manifested itself. All that I wish to point out here is that, in
any discussion of the causes of torsion and detorsion in gastropod
molluses, the action of the muscles should be considered more.
carefully than has hitherto been the case. I do not propose to-
pursue the subject further in this place, as it would involve a
long and detailed comparison of the regional anatomy of many
different gastropods, but before leaving it I must guard myself
against the imputation of giving a Lamarckian explanation of a
particular course of molluscan evolution. I have suggested a.
physiological explanation, and this is necessarily Lamarckian in
form. ‘The reader can easily translate my suggestion into terms.
of Natural Selection, and it would be no compliment to his.
intelligence if I were to occupy an additional page of print in
doing so.
Of other external features, I need only allude to the following.
The snout is so short that it can hardly be said to exist; there is
neither pretentacular nor postentacular elongation in the cephalic
. or postcephalic regions in the sense indicated by Amaudrut (1).
The opercular lobe is relatively very large and muscular; its
outline is semilunar, the left corner deeply indented to receive
the apophyses of the operculum (fig. 29). The epipodium is.
represented by a low ridge, devoid of any tentacles or outgrowths,
extending from behind the tentacles to the posterior end of the
operculum on each side. It is somewhat enlarged in the opercular
region, but is inconspicuous in contracted specimens, because it is.
compressed between the hind end of the foot and the opercular
lobe.
The anterior border of the mantle is muscular, and in marine
species (Verita) more or less frilled. The remainder of the
mantle is extremely thin.
The tentacles are conical, commonly decorated with alternate
stripes of black and white, and are short in all the fresh-
water species (Paranerita and Neritina) that I have examined,
but long and slender in the marine species (Verita).
Various accounts have been given of the so-called cephalic penis
in the Neritide. It was figured by Quoy and Gaimard (35),
referred to in somewhat doubtful terms by Moquin-Tandon (28),
overlooked by Claparéde (12). Bouvier (9) gives a remarkable
figure of this organ in Veritina cariosa, and Thiele makes a very
short reference to it and gives a small figure of its appearance
in transverse section. The organ in question is without doubt a
modification of the inner part of the base of the right tentacle
and is a good external indication of the male sex. Its position
*%
54
850 PROF, G. C, BOURNE ON THE - [Nov. 17
and appearance are illustrated in fig. 30 (Weritina fluviatilis),
fig. 31 (Werita melanotraga), and fig. 32 (Paranerita gagates). It
differs somewhat in size and shape in the other species I have
examined, but I have never seen it bifurcated as described by
Bouvier, nor of the shape figured by him for WV. cariosa.
In all the species I have examined it is a solid muscular out-
growth of the base of the right tentacle, innervated, so far as I am
able to determine from sections, by a slender branch of the right
tentacular nerve. Its external side, that is the side nearest the
tentacle, is deeply grooved ; the lower lip of the groove is swollen
and prominent and clothed with a iayer of long cylindrical
epithelial cells, among which are numerous sense-cells. Elsewhere
the surface of the organ is covered by a moderately high ciliated
cylindrical epithelium, the cilia being specially well developed
along the lower surface, but the groove itself is lined by a simple
non-ciliated cylindrical epithelium. The hinder end of the
groove opens almost behind and above the base of the right
tentacle, and there is no sign of any furrow or ciliated tract
leading from the groove to the aperture of the male duct. The
male opening lies close to the base of the right eye-stalk, but
rather in advance of it, and it is not easy to understand how any
connection can be established between it and the penial groove.
Communication on the ventral side of the tentacle seems to be
hindered by the presence of the epipodial ridge. It is, however,
difficult to judge from the examination of contracted specimens,
The organ is probably very extensile, and its shape seems adapted ~
for grasping and conveying the complicated spermatophores of
Nerita and Paranerita to the vaginal opening of the female;
but I must confess that I have been unable to find any proof of
this. Ihave found no trace of spermatophores either in the male
ducts or in the groove of the penis in any. species, and I have
failed to observe the act of copulation in the numerous WVeritina
fluviatilis that U have kept in aquaria. ; '
The relative positions of heart, rectum, kidney, and ctenidium,
as seen in an external view of the entire animal, are shown in
fig. 29 and need no further comment.
In the marine species (Nerita) there is a well-developed
opercular gland opening by a median aperture just in front of
what is morphologically the anterior edge of the operculum,
between it and the posterior edge of the mantle-flap. (In a
contracted specimen, such as that drawn in fig. 29, the anterior
edge of the opercular lobe appears to be posterior.) This gland is
very large in Nerita plicata, melanotraga, and peloronta; smaller
but still conspicuous in lineata and plexa. In freshwater species
(Paranertta and Neritina) it can scarcely be said to exist, being
‘represented only by a glandular area, sometimes produced
internally into a few crypts, in the angle between the mantle
and the opercular lobe. The gland in Werita is clearly an
invagination of this glandular area, and there is a gradual and
interesting transition from the external epithelium to the highly
1908. | ASPIDOBRANCH GASTROPOD MOLLUSCS. 851
modified epithelium forming the bulk of the gland. The duct is
short and wide, and as it enters the hemoccele is accompanied by
a stout strand of muscle, from which flat muscular partitions
radiate in all directions. The lumen of the duct enlarges greatly,
and forms a sort of sac the cavity of which is broken up into a laby
rinth, formed by the ingrowth of the muscular partitions carrying
before them the highly modified glandular epithelium of the sae.
The whole forms a large and compact organ lying below the
viscera, and very conspicuous when the animal is disserted. The
structure of the epithelium is peculiar (fig. 34) and sections of
it are not easy to interpret. Judging from the transitional
epithelium lining the duct, it appears that the ordinary cylindrical
epithelial cells become very slender and are compressed by the
excessive development of the gland-cells lying between them:
they lose their cilia; their nuclei, reduced to a small size, stain
very deeply and are shifted to the external ends of the cells, the
cytoplasm radiating outwards from the nuclei to form a cover
over the distal ends of the adjacent gland-cells. They have in
fact undergone a modification similar to that of the cover-cells of
the testis of Hydra. The gland-cells appear to be of two kinds:
clear cylindrical cells with finely granular contents staining
faintly in hematoxylin, and cells of similar shape but filled with
irregular and highly eosinophilous globules. J have satisfied
myself, however, that these are only different phases in the
secretory activity of the same kind of cell. The pale, finely
granular cells represent the resting condition. Their nuclei are
very small and deeply staining and are either close to the base
and on one side of the cell or somewhat higher up and close to
one side. Some of these cells are vacuolated, their contents stain
lightly in eosin, and their granular contents begin to aggregate
into globules, this process generally commencing at the basal end
of the cell. In later stages the whole cell-body is filled with
eosinophilous globules of irregular size, and these are finally
discharged, by rupture of the cover-cells, into the lumen of the
gland. The space between the hinder flap of the mantle and the
opercular lobe is commonly full of the sticky coagulated secretion
of the gland, but what its function is I am unable to guess.
From its large size, the opercular gland must be of considerable
physiological importance, but it is remarkable that it only occurs
as a highly developed and differentiated structare in marine
forms. Possibly its secretion is useful in assisting the animals to
retain their hold on rocks washed by the waves.
The Alimentary Tract.
This may be treated very briefly, as the structure of the buccal
bulb, cesophagus, stomach, and intestine is very constant through-
out the family, and the details in which the various species differ
from one another are so small and unimportant that they may be
passed over. A sketch of the alimentary canal of Paranerita
852 PROF, G. C. BOURNE ON THE | Nov. 17,
longispina, with the coils of the gut unravelled, is given in fig. 35,
and the relative positions of buccal bulb, intestine, and stomach,
and the division of the last-named into esophageal and pyloric
moieties, are indicated in fig. 41. The characters of the
epithelial lining of different portions of the gut, and the ridges
and folds into which the lining of the stomach is thrown, are
remarkably similar in all the species I have examined, and Lenssen’s
description of the digestive system of Neritina fluviatilis may be
taken as typical for all the Neritide. Such small differences as
occur chiefly affect the relative size and length of the radular sae,
and the length and consequently the complexity of the coils of
the intestine. Marine species pass a considerable quantity of
sand through their bodies, and their intestine is longer and
thicker-walled than is the case in freshwater species. The
radular sac is also longer, and the radular teeth coarser and more
powerful, in marine than in freshwater forms. The relative size
of the pharyngeal bulb also differs. It is, for example, very large
in WVerita peloronta and plexa, relatively small in Paranerita
varigata and gagates, but in every case the essential structure
and relations of the cartilages of the odontophore, the salivary
glands, and diverticula of the pharyngeal cavity are the same.
There are seven buccal cartilages in all Neritide, as described by
Lenssen, and the differences in size observed in the pharyngeal
bulb are due to the greater or less size of these cartilages and
the more or less powerful development of the intrinsic muscles
attached to them. In all cases I have found the pair of lateral
glandular sacs opening into the lateral extension: of the sub-
radular diverticulum as described by Thiele. In some species
the walls of these sacs are pigmented and their lumina contain
dark-coloured concretions. J must add that I have in no case
been able to find salivary glands with long ducts such as are
described by Béla Haller in Werita ornata. (‘In diesen munden
dorsalwarts an der gewohnlichen Stelle mit sehr langen, wohl
differenzirten Ausfuhrungsgiingen, zwei compacte acinése Buccal-
driisen,” loc. cit. p. 131, Taf. xi. fig. 123.) These salivary or
buccal glands always have the structure and relations described
by Lenssen. They are closely applied to the hinder end of the
pharyngeal bulb and pass round to its lower surface, embracing
the origin of the radular sac. They have no ducts, properly
speaking, but are large saccular diverticula of the anterior end of
the cesophagus, and are comparable with the ceesophageal pouches
of other Rhipidoglossa.
The Nervous System.
I have already given on pp. 819-821 an analysis of the different
accounts of the nervous system given by different authors, from
which it may be gathered that while there is agreement on the
main points, there are still questions about which there is
considerable difference of opinion, Having nothing to add on
1908. } ASPIDOBRANCH GASTROPOD MOLLUSCS. 853
the subject of the cerebral ganglia, the labial commissure, the
buccal commissure, and the pedal centres to the excellent figures
and descriptions of Bouvier, I will pass on at once to the
‘disputable points.
It has been clearly demonstrated by Bouvier (10), Boutan (7),
and Haller (20) that the marine Nerite (Nerita) and Septaria
are typically streptoneurous, the supra-intestinal branch of the
visceral commissure existing as an extremely fine nerve whose
relations are normal, Lenssen (26) has shown that with one
exception the general character of the nervous system is the same
in Neritina fluviatilis, but this exception is important, He could
not find any trace of the supra-intestinal nerve, nor of any
connection between the visceral and branchial ganglia, I have
taken great pains to discover this nerve in the same species, but
have net succeeded in discovering it either in sections or by
dissection.
Though it would seem on ¢ priori grounds improbable that the
nervous system of Weritina fluviatilis, so similar to that of
Nerita in all other details, should differ from it in the
deficiency of so characteristic a nerve, I can only add my
testimony to that of Lenssen, and state that I believe that it
does not exist. If we are right in this conclusion, it affords
an additional reason for separating, as I have done, Neritina
fluviatilis from the tropical freshwater forms which I have classed
together under the name Paranerita. In the latter there is no
question of the completeness of the crossed visceral commissure.
I have found it in all my specimens, and its course is very
eorrectly described by Bouvier. In this, as in other respects,
I find that the nervous systems of Merita and Paranerita are
practically identical.
But in spite of all the labour that has been bestowed upon the
subject there is still an uncertainty or deficiency in the descriptions
and figures hitherto given of the ganglia on the visceral com-
missure. I have devoted a considerable amount of time to this
‘question, working both by means of sections and dissections and
hope to elucidate some points that have hitherto been obscure.
Not being satisfied with existing figures of the pleuropedal
centres, I have given in fig. 36 a drawing founded on the
combined results of dissections and reconstruction of serial
sections. As the drawing is fully lettered, 1 may spare the
reader the trouble of a detailed description of the nerves issuing
from the pedal and pleural ganglia. J have made use of the same
lettering as Bouvier to designate the different nerves, so that my
drawing can be more easily compared with his. I will only call
attention here to the branch of the cerebro-pleural connective
labelled s, and to the nerves labelled op. The former diverge
gradually from the connective, pass to the outside of the anterior
extrinsic muscles of the pharyngeal bulb, and are distributed to
the walls of the cephalic region behind the tentacles. They are
‘quite distinct nerves, but do not appear to have been noticed
854 PROF. G. C. BOURNE ON THE { Nov. 17,.
before. The nerves op. pass under the pleural ganglia and take-
their origin from the swollen anterior ends of the pedal cords..
They are slightly asymmetrical, but are distributed on both sides.
of the body to the opercular lobe.
As I have already pointed out, authors differ in their identi-
fication of the subintestinal ganglion. The conical enlargement
of the origin of the subintestinal nerve—it has in all species the-
shape represented in fig. 36, and is not a rounded ganglion as
figured by Bouvier—was identified by de Lacaze-Duthiers as.
the subintestinal ganglion; and Bouvier and Lenssen have
accepted this view. Boutan and Haller, on the contrary, claim
that the subintestinal is represented by an elongated ganglionic
swelling on the right of the visceral commissure just where the
latter turns towards the left to run through the large blood-sinus.
leading in the direction of the uropore. Concerning this ganglion.
I have something to say. As isshown in the accompanying text-
figure (172), it is an elongated and rather diffuse ganglion from
which three main nerves are given off. The most anterior (2")
passes inwards, forwards, and ventralwards and supplies (as far
as I am able to determine) the coils of the intestine lying
below and to the side of the anterior part of the radular sac.
Lenssen has described a similar nerve in WV. fluviatilis, but im
this species it is given off some distance in front of the ganglion,
and, curiously enough, he says that some of its branches are
distributed to the stomach ; as a matter of fact, they pass in an
opposite direction. The second nerve (n°) is rather stout and
passes outwards into the tissue overlying the posterior part of
the right columellar muscle. It does not penetrate the muscle,
but turns sharply backwards, and I was able to trace it in sections.
as far as the pedicle of the visceral mass, and thence alongside
the oviduct to the ovarian follicles. Lenssen has described a
similar nerve in JV. fluviatilis arising like the first well in front of
the ganglion ; this he calls a columellar nerve, but I am of the:
opinion that it has the same distribution as that just described,
though I could not follow it very well in my sections. The
third nerve has been quite incorrectly described by previous.
authors. As shown in the text-figure, it is very short and stout,
and passing downwards enters at right angles a stout and
relatively long nervous cord (g.g.), which is thickly coated with
nerve ganglion-cells. This cord may properly be described as a:
genital ganglion. It is closely attached to the gonaduct, and its.
lower end (gn*) turns sharply backward and passes to the spermato-
phore-sac (or epididymis in the male), on the surface of which it.
breaks up into a number of fine fibres. Its upper end (gn’)
accompanies the gonaduct in its course towards the complex of:
accessory genital glands, and on its arrival into the complex
breaks up into a number of fine branches whose further course:
I was unable to follow, but there is no doubt that they are
distributed to the different genital glands. This nerve corresponds,
without doubt, to the recto-genital nerve of other Streptoneura,
1908. ] ASPIDOBRANCH GASTROPOD MOLLUSCS. 855:
and the large ganglionic enlargement on its course is associated
with the great importance and complexity of the accessory genital
organs in the Neritide.
Beyond this recto-genital nerve the visceral commissure in its
course from right to left is somewhat thinner than it is in the
region of the ganglionic swelling just mentioned, but it remains
invested with a coating of nerve-cells, and is only gradually and
slightly enlarged in front of the uropore to form the visceral
ganglion described and figured by all previous authors. I must
Text-fig. 172.
Liane Ge:
WIS — 9.9
gn:
An.
A dissection of Nerita plexa, showing the course and distribution of the visceral
nerves and the visceral, branchial, and osphradial ganglia: semidiagrammatic.
An, anus. 67.n, branchial nerve. cm.r, right columellar muscle. Cry.s, crystal
sac. Ct, ctenidium. gd, complex ot genital ductsand glands. 4g.g, genital ganglion.
gm, nerve passing to the spermatophore-sac. gn”, nerve passing to the organs of
the genital complex. m}, anterior (pallial) branch of the branchio-pallial nerve. m2,
posterior (branchial) branch of the branchio-pallial nerve. os, osphradium. o0s.g,
osphradial ganglion. Ph, pharyngeal bulb. sb.n, subintestinal nerve. sp.g, supra-
intestinal ganglion (identified as a separate ganglion only in Nerita plicata and
Paranerita gagates). sp.n,supra-intestinal nerve. Urp,uropore. V1,right-hand
enlargement of the visceral ganglion. J), left-hand enlargement of the visceral
ganglion. These two enlargements are much closer together than represented in the
figure; the tissues lying between them have been stretched by turning back the
genital complex, and the middle part of the visceral ganglion is represented as if
stretched to a corresponding amount. It is almost invariably broken in actual
dissection.
856 PROF. G. C. BOURNE ON THE ; [ Nov. 17,
add that, contrary to Lenssen’s statement, the genital nerve in
NV. fluviatilis is given off from and not in front of the first
ganglionic swelling, and that the characters of the genital
ganglion &e. are practically identical with those described above.
The nerves given off from the enlargement of the visceral
commissure in the vicinity of the uropore are distributed, as all
previous authors have described, to the kidney and pericardium.
If I have described the origin and distribution of these nerves
at some length, it is because they are of importance in determining
the homologies of the first ganglion on the visceral commissure
which Boutan and Haller have identified with the subintestinal
ganglion. Now this ganglion, whatever its size and position,
never gives rise to nerves supplying the viscera and gonads. It
is essentially the ganglion of the right side of the mantle, and in
the primitive dibranchiate Aspidobranchia supplies the nerve
going to the post-torsional right ctenidium and osphradium.
When these disappear, as in the Trochide and Pectinibranchia,
either there is no definite subintestinal ganglion or it tends to
approximate itself to the left pleural ganglion as in the Cerithiide,
or, aS is more often the case, it may enter into close relations with
the right symmetrical pallial nerve and innervate the right side
of the mantle. On the other hand, the genital and visceral nerves
always issue from the visceral ganglion (or ganglia if more than
one is present) at the hinder end of the visceral commissure, and
this original connection is maintained with great persistence even
in the short-looped euthyneurous forms. Hence it would be
contrary to what is observed in other gastropods if the genital,
intestinal, and stomach nerves issued from the subintestinal
ganglion ; and the conclusion is that Boutan and Haller were
wrong in their identification, and that the ganglion in question
isa member of the visceral series. This is the more likely when we
consider that in other Rhipidoglossa—in 7’rochus, for instance—
the abdominal ganglion is an elongated and ill-defined enlarge-
ment occupying a considerable section of the posterior part of
the visceral commissure. As I have stated, there is a continuous
and thick cortex of nerve ganglion-cells investing all that part of
the visceral commissure of the Neritide lying between the
ganglion from which the genital nerve proceeds and the ganglion
adjacent to the uropore. The whole of this thickened section is
to be compared with the elongated visceral ganglion of Z'rochus,
and the swellings at its two ends, which do not in fact form such
distinctly separate ganglia as might be inferred from figures and
descriptions, may be regarded as concentrations of nerve-cells—in
other words, incipient but not yet separate ganglia at the two
ends of a long and ill-defined tract of ganglion-cells. This view
is strengthened by the fact that the swelling at the right end
from which the genital and other nerves proceed bears the same
relations to the oviduco-celomic funnel (which is evidently a
relic of the right kidney) as the swelling from which the renal
and pericardial nerves proceed bears to the reno-pericardial funnel
1908. ] ASPIDOBRANCH GASTROPOD MOLLUSCS. 857
of the left kidney. The great elongation of the ganglionated
posterior tract of the visceral commissure is readily explicable
when it is borne in mind that, as I have already described for
Septaria and shall describe further on for Verita, the embryonic
condition of the celom has been retained in this family and its
cavity stretches across the body from left to right, the two
primitive kidneys being separated by a considerable space, and
opening at widely separate points into the celom. In short,
this region of the body is broader from right to left than is usual
in Gastropoda, and the visceral ganglionic swelling is elongated
accordingly.
I must admit that it may be urged against this view that if the
“organe creux” described by Lenssen in iV. fluviatilis, and
found by me in a corresponding position in the various species
examined, is really a vestige of the right ctenidium, the swollen
right end of the ganglionic enlargement bears the same topo-
graphical relation to it that the subintestinal ganglion bears to
the right ctenidium in dibranchiate Rhipidoglossa. I confess
that I was at first inclined to agree with Boutan and Haller in
identifying what I now regard as a specialized part of the visceral
ganglion as the subintestinal, and was inclined to argue that the
outgrowth must represent the right ctenidium because of its
relation to the ganglion, and that the ganglion must be the
subintestinal because of its relation to the relic of the ctenidium.
This was so flagrant an example of the circulus in definiendo that
I was led to reconsider the question, and I am satisfied that the
origin of the genital nerve and the relationship to the oviduco-
‘celomic funnel and to the right side of the ccelomic cavity
‘outweighs all other evidence.
If, then, Boutan and Haller were wrong, de Lacaze-Duthiers
and Bonvier were right in their identification of the subintestinal
ganglion. To test this point, ] have made a careful study of
sections through the pleuro-pedal centres. These goto show that
the enlargement from which the subintestinal nerve proceeds is
really a nerve-centre, and is therefore the representative of the
‘subintestinal ganglion. To make this point clear, the question
must be asked, what do we mean when we speak of a ganglion ?
It is a swelling caused by the presence of the nerve ganglion-
cells investing a central core of nerve-fibrils. It is something
more than this, it is a nervous relay, in which some of the fibres
enter into nerve ganglion-cells, and in which the dendrites of
the ganglion-cells are intermingled and in contact with one
another. As faras I am aware, nobody has yet attempted to
work out the courses of the nerve-fibres and to trace them to
their connection with groups of nerve-cells in the Mollusca. I
am unable to do more than touch the fringe of a subject that
offers a large field for future research, as my preparations were
not made with this purpose and are inadequate to disentangle the
complex of nerve-fibres and cells. I have, however, ascertained
the following facts :—Each nerve-centre consists of a core of nerve-
858 PROF, G. C. BOURNE ON THE [ Nov. 17,
fibres and dendrites surrounded by a cortex of nerve ganglion-:
cells. There are three kinds of ganglion-cells: (1) Small bipolar
cells, very numerous, closely crowded together, and with deeply
staining nuclei. They occur in great numbers in all the nerve-
centres and are the only cells extending from the centres along
the nerve-trunks. (2) Large apparently unipolar cells, with
larger and less deeply staining nuclei than in 1. These are
searce and locally aggregated in the nerve-centres. (3) Large
multipolar cells more numerous than the second kind and with
similar nuclei. They are localized chiefly at the sides of the
centres, and in the pedal cords their axial processes can often be:
distinctly traced into nerve-fibres.
The presence of the larger cells of the second and third kind is
characteristic of a separate nerve-centre or ganglion. These facts.
are illustrated, but on a very small scale, in figs. 37-40.
Fig. 37 is a section, taken somewhat obliquely, through the
pleural and pedal centres, just behind the union of the former
with the latter, and including the connection between the two
pleural ganglia. The details of the pedal centres need not con-
cern us; it is sufficient to allude to the fact that they give
evidence of a considerable degree of complication. In the pleural
centres it 1s evident that there are two principal groups of nerve-
cells in each ganglion : a dorso-lateral and a ventral. A bundle
of nerve-fibres, originating from the dorso-lateral group of the
left ganglion, runs across to the right and turning downwards
traverses, but does not enter into connection with, the ventral
group of cells of the right ganglion and is joined by another band
of fibres originating from a distinct group in the lateral region of
the right ganglion. These two bundles unite to form the root of
the subintestinal nerve. Fig. 38, representing a section some-
what posterior, shows the subintestinal root still small and
traversing the ventral cell-group of the right ganglion. Fig. 39
passes through the base of the conical subintestinal ganglion and
shows the same bundle of fibres as in the last section, now
considerably larger; and above it a new bundle surrounded by a
very thick mass of ganglion-cells, among which is a relatively
large number of the larger kinds. The intervening sections
would show that the fibres of the upper bundle pass into or take
their origin from the thick dorsal and lateral mass of nerve-cells.
Fig. 40 is a section taken some way further back through the
subintestinal nerve and shows the two bundles of fibres still
distinct and surrounded by a thin layer of small nerve-cells.
Eventually when the layer of nerve-cells dies out the two bundles
of fibres can no longer be distinguished from one another. There
can be no doubt that the bundle / in fig. 37 is the origin of the
subintestinal nerve from the left pleural ganglion, and that the
bundle z represents the much-abbreviated zygoneurous connection
of the subintestinal with the right pleural ganglion. The mass
of cells sb in fig. 39 is the subintestinal ganglionic centre, and
the uppermost of the two bundles of fibres marked / is formed by.
1908. ] ASPIDOBRANCH GASTROPOD MOLLUSCS. 859
fibres of the subintestinal nerve originating in that centre. It is
possible that the two bundles represent afferent and efferent
fibres, but there is no proof of this. The facts enumerated,
however, are sufficient evidence that Bouvier was right in his
identification of the subintestinal glanglion. The sections show
further that the commissure between the two pleural ganglia
signifies much more than a zygoneurous connection between the
right pleural and the subintestinal. The bulk of the commissure
is made up of two stout bundles of fibres (1 & 2 in figs. 37 and
38), running transversely and connecting respectively the dorso-
lateral and the ventral cell-groups of the right and left pleural
ganglia. The upper bundle is related to the origins of the
cerebro-pleural connectives, the lower bundle to the origins of the
right and left pallial and columellar nerves. It is evident that
the shortening of the anterior part of the visceral loop and the
approximation of the subintestinal ganglion to the pleural centres
is connected with a crossing over of nerve-tracts belonging to the
symmetrical pallial centres, and that this transference is quite
independent of the zygoneurous or dialyneurous connections of the
visceral commissure. It is suggestive that this intimate union
between the right and left pallial centres occurs in a group of
Rhipidoglossa in which there are two subequal and sub-
symmetrical shell-muscles, innervated from the pleural ganglia.
I have given reasons for believing that the development of the
left muscle is correlated with the development and increased
functional importance of the operculum, and that when once it is
established its action must produce a tendency towards the
secondary symmetrical disposition of the organs connected with
or influenced by it. This tendency is manifest in the nerve-
centres. Theasymmetrical centres become of less, the symmetrical
centres of greater, functional importance. The two shell-muscles,
acting together as a pair, must be subject to nervous co-ordination,
This has been effected by the development, in conformity with
the physiological needs of the organism, but through the operation
of natural selection, of transverse connections between the pleural
eentres, and these, be it noted, are of two kinds. There is a con-
nection by way of the cerebro-pleural connectives, providing, as
one may legitimately infer, for the co-ordination of sensory
impulses arriving from the cephalic sense-organs. And there is a
connection by way of the symmetrical pallial and columellar
nerves providing for the co-ordination of sensory inypulges
arriving from the anterior borders of the mantle and of motor
impulses travelling outwards to the two shell-muscles.
These considerations suggest a fruitful field of enquiry into the
‘causes which have led, firstly, to the separation of pleural from
pedal centres ; secondly, to the various degrees of approximation
or separation of pleural and cerebral centres ; and, thirdly, to the
approximation of the asymmetrical to the symmetrical pallial
centres with the concomitant shortening or partial suppression of
the visceral commissure. J have many indications that an
860 PROF, G. C. BOURNE ON THE [ Nov. 17,,
enquiry on these lines would help to elucidate a number of
doubtful problems in gastropod morphology ; but the subject is a
large one, requiring much comparative study of different forms of
nervous systems and must be left for a future occasion.
Respecting the supra-intestinal ganglion and its connections,
my observations serve to bring together and harmonise the:
scattered descriptions of previous authors. The text-figure on
p. 855 shows the relations of the supra-intestinal ganglion, the
symmetrical left branchial and pallial nerves, the osphradium,
and the osphradial ganglion. The supra-intestinal ganglion is.
always very small and hard to discover, but I have determined
its presence in WVerita plicata and Paranerita gagates. It is con-
nected by a very short branch with a long cord of nerve-fibres
ensheathed by nerve ganglion-cells which underlies the osphradium..
This cord must be regarded as a diffuse osphradial ganglion. It.
receives the bulk of its nerve-supply from the left symmetrical
branchial nerve, and this connection has been noted by several
authors ; Thiele (39) in particular gives a good figure of it as seen
in section. The posterior end of the osphradial ganglion is con-
tinued into the clearly defined nerve which passes along the edge-
of the right or anterior suspensory membrane of the ctenidium
and is continued along the afferent side of the gill. I can con-
firm Bernard’s statement that this nerve is continued round the:
apex of the gill and down its efferent side. The position of the.
osphradium has been correctly described by Bernard (3). It lies,
as indicated in the text-figure, in front of the attachment of the
suspensory membrane of the ctenidium to the roof of the mantle-
cavity, and may be distinguished by the naked eye as a small
furrow bounded by two ridges of thickened epithelium. — Its.
minute structure does not differ in any important particular from
that of Septaria.
The Respiratory and Circulatory Systems.
The ctenidium, heart, and blood-vessels require little or no
description. They are constructed on the same plan throughout.
the Neritide, and as Lenssen has treated the subject very fully
in his memoir on Neritina fluviatilis and I have already written
all that I have to add to his account in dealing with Septaria,
there is no more to say. The right auricle is always present, its.
position in Paranerita gagates being indicated im fig. 42. It
is somewhat less well-developed in marine forms (e. g. WVerita
peloronta) than in the freshwater forms, but always carries.
blood back to the ventricle from lacune in the posterior and left
side of the body-wall. It is separated by the whole width of the-
visceral pedicle from, and has no connection with, the little hollow
organ lying in the right-hand posterior corner of the mantle-
cavity which I have referred to as possibly being a relic of the-
right ctenidium, and this fact must be admitted as evidence
against the suggested homology. The gill-lamelle of the ctenidium_
1908. | ASPIDOBRANCH GASTROPOD MOLLUSCS. 861.
of all species much resemble those of Septaria, but the extent
and shape of the ciliated tracts vary slightly in different species.
—not to such an extent, however, as to make it worth while
to write a separate description for each.
The Hemocele and Celom.
The only noteworthy feature about the hemoceele is that it:
tends to be filled up by an abundant development of vesicular
connective tissue, which penetrates into all the spaces between
the viscera, except the ccelomic spaces, and is specially accumu-
lated round the blood-vessels. It is naturally much altered by
the action of reagents in tropical specimens long preserved in
spirit, but may conveniently be studied in Weritina fluviatilis.
As JI have not yet been able to obtain Paravicini’s memoir on
the connective tissue of Gastropods, nor have I had time to.
make a comparative study of this tissue in freshly killed
specimens, I will only shortly mention the appearance presented
by this tissue in Weritina. In starved specimens which have.
been kept for a long time in aquaria, the connective tissue
consists chiefly of a number of stellate cells united by their:
processes, or, if one prefers to express it so, of a reticulum of
protoplasm with nuclei at the nodes. These nuclei (fig. 59,
ret.) are small, oval, and deeply staining. In well-nourished
specimens recently taken from the river the meshes of the-
reticulum are filled with vesicular cells (‘ Langer’s cells”) with
larger faintly staining nuclei. These cells are filled with
granules which stain bright yellowish green in picro-indigo-
carmine. They accumulate round the blood-vessels, and are often
so abundant as to obscure the network of connective tissue..
Apparently they are derived from small ameeboid cells which
contain similar large nuclei such as that marked am. in fig. 59..
There can be little doubt that these cells are metabolic in function
and serve to store up reserve material, though I have not been
able to demonstrate the presence of glycogen in them as Blund-
stone (5) has in other molluscs. Masses of fatty-looking tissue of
this nature surround the intestines, liver, nerve-cords, and even
the accessory genital organs of tropical species of Werita and
Paranerita and their presence is a great hindrance to dissection..
I have found that this tissue blackens slightly with osmic acid in
Paranerita, indicating the presence of fat; and this is worth
noticing, for fat-cells are said to be absent from the connective.
tissue of Gastropods. The vesicular cells of Veritina are not
blackened by osmie acid. :
The ccelomic cavity is fully as extensive, and has much the
same relations as in Septaria, but owing to the retention of the
spiral coil of the visceral mass, and the complications arising from
the excessive development of the spermatophore-sac of the female
or the epididymis of the male, it is difficult to give an intelligible
account of it; but I hope that with the assistance of the diagram,,.
862 PROF. G. C. BOURNE ON THE | [ Nov. 17,
fig. 58, the reader will be able to understand the following
account of it. For descriptive purposes and without prejudice to
any theoretical conclusions, the coelom may be regarded as con-
sisting of a left or pericardial division (figs. 29, 42, 58, pe.co.)
and right, or gonadial division (figs. 41, 43, & 58, g.co.). The
pericardial division lies on the left side of the body and is of
considerable vertical depth. Anteriorly it extends to the base of
the ctenidium, posteriorly to the union of the pyloric and
esophageal divisions of the stomach (fig. 42). Its outer wall is
very thin; its inner wall tough and muscular. Its posterior end
is traversed by the rectum, and in it lie the ventricle and the two
auricles of the heart. On the inner or right side the pericardial
ccelom is continued into a wide passage (c.ap.) which runs under
the kidney, above and in front of the anterior lobes of the liver.
From this passage a diverticulum is given off on the left side,
which passes under the non-glandular part of the kidney and
stretches forward towards the uropore (fig. 43), below which it
ends in a dilatation and receives the celomic opening of the reno-
pericardial canal. To the right the passage widens out to form
the gonadial ceelom, a flattened sac of irregular form lying between
the base of the genital complex and the liver. Its right-hand
corner is produced into a diverticulum extending as far as the
posterior end of the right columellar muscle, and here it comes
into cloge relation with the gonad, but it does not surround any
portion of the latter organ, as is the case in Septaria. The
oviduco-ccelomice funnel opens into the right-hand corner of the
gonadial ccelom (fig. 41, od.c.f.) at the base of the diverticulum
just mentioned and not far from the columellar muscle. In the
male the sperm-duct runs very close to the wall of the same part
cof the ccelom, but after an assiduous search through many series
of sections I have been unable to find any trace of a spermiduco-
celomic funnel. The cavity of the gonadial division of the
ceelom is largely obliterated by the spermatophore-sac of the
female or the epididymis of the male, which projects into it like a
hernia from the posterior end of the genital complex, and, carrying
the posterior wall of the celomic cavity in front of it, les
enclosed in a ccelomic pocket in the position shown in figs. 44, 49,
and 58. It is clear that these relations are similar to those
described for Septaria, the difference being that in the latter
genus the straightening out of the visceral spire has allowed the
gonadial ccelom to extend much further along the right side of
the body, so that its relation to the gonad is obvious. There can
pe no doubt, however, that in the more spirally coiled WVerita,
Paranerita, and Neritina the gonad has been derived from the -
‘wall of the right corner of the gonadial celom. The oviduco-
coelomic funnel of the female is evidence of this primitive
connection ; and as the cavity of the gonad is morphologically a
part cf the ccelom, the latter must be regarded as co-extensive
with the gonad, and therefore as occupying the whole of the
-apex of the visceral spire. If, as a result of the large development
1908. | ASPIDOBRANCH GASTROPOD MOLLUSCS, 863
of the liver and intestines, the visceral spire were elongated, the
gonad would remain at its apex, and the gonaduct would be
correspondingly elongated. This, in fact, is the typical position
of the gonad in multispiral gastropods, and the primitive connec-
tion of gonad with ccelom is well illustrated by the Neritide.
The Hxcretory System.
The kidney in Verita and *Paranerita has essentially the same
structure and relations as in Septaria and Neritina fluviatilis.
It consists of a glandular and a non-glandular part or bladder
communicating posteriorly behind the rectum. The bladder is a
very wide sac (fig. 43, A’) lying between the glandular part and
the pericardium. Anteriorly it twists under the glandular part
and leads into a small thick-walled chamber (fig. 47) which opens
to the exterior by the uropore (fig. 45, Urp.). As in Septaria
the posterior moiety of the glandular part seems always to differ
somewhat in histological character from the anterior moiety; this
difference is indicated in figs. 43, 44, & 46. The only direct
connection between the glandular and the non-glandular parts
of the kidney is at the posterior end, but there is an indirect
anterior connection, similar to that described in Septaria depressa.
The reno-pericardial funnel opens, as explained above, into a
diverticulum of the ccelom which passes below the non-glandular
part of the kidney (fig. 46). Its further course is that of an @,
and for some distance it projects into the bladder, and eventually
becomes attached to the wall of the chamber which opens to the
exterior by the uropore (fig. 45). Passing up the wall of this
chamber it opens into it by a small but distinct ciliated passage
(fig. 47, cil.p.), and continuing its course up the wall of the
uropore-sac dilates to form a canal of considerably wider diameter
which divides into two branches. The main branch passes to the
left over the uropore-sac, and opens into the glandular part of the
kidney ; the smaller branch passes to the right and is connected
with some small detached renal lobes lying in the blood-sinus
leading to the afferent branchial vessel. The characteristic
epithelial lining of the reno-pericardial canal, described in detail
by Lenssen and Thiele, is confined to the section lying between
the ccelomic opening and the ciliated connection with the uropore-
sac. This section is very long; I have calculated that it is at
least ‘9 mm. long in a specimen of Paranerita gagates measuring
13 mm. in length. The transition from the characteristic epithe-
lium of the duct to an ordinary columnar ciliated epithelium, and
from the latter to the glandular epithelium of the kidney, is
shown in fig. 47.
The wide thin-walled sac which I have described as the bladder
or non-glandular part of the kidney is generally named the
ureter. J have not used this term because in those Gastropods in
which a long ureter is present running alongside of the rectum
(e.g. Paludina) there is evidence that it is formed from the
Proc. Zoot. Soc.—1908, No. LV. peel 5)
864 PROF. G. C. BOURNE ON THE [Nov. 17,
mantle and therefore lined by an ectodermic epithelium. IL
believe that the homologue of such an ectodermal ureter is to be
found in the structure which I have referred to as the uropore-
sac. As shown in fig. 45, the mantle-epithelium surrounding the
uropore is modified, consisting of high columnar ciliated ‘cells.
This epithelium passes round the lips of the ur opore, and appears
to line the uropore-sac, which, however, should be described as a.
recess rather than a sac, for it is widely open below into the non-
glandular part of the kidney, and only has the appearance shown
in fig. 47 in sections passing through its upper part. For some
reason the epithelium lining this recess is very much macerated
in all my specimens and I cannot give a clear account of it; but
the thick wall appears to be made up of a number of glandular
crypts, or, rather, of bunches of gland-cells, which have passed
through the basement-membrane into the subjacent connective
HSene. as is the case with the various accessory genital glands.
The ciliated epithelial cells retain their position on the outside of
the basement-membrane. Lower down, in the vicinity of the
uropore, the glandular structure eradually disappears, but the
columnar ciliated cells are continued as a broad band running
back for some distance along the anterior and inner wall of the
bladder (fig. 46, ci.ep.). The histological characters of this
ciliated band leave little doubt that it is an ingrowth of the
ectoderm, and its function is obviously to create a powerful
ciliary current in the direction of the uropore.
The position of the little folded organ which has been referred
to as possibly representing the right ctenidium is indicated im
fig. 43, R.ct. In some specimens it is only represented by a
thickening and modification of the mantle-epithelium exactly like
that round the uropore. In other specimens the epithelium is
thrown into folds, but this may be due to contraction in spirit.
The mantle-wall is thickened by a special development of muscular
strands below the patch of modified epithelium, and in connection
with these is a blood-space, which is nothing more than a diverti-
culum of the large transverse blood-sinus enclosing the visceral
ganglion. The organ in question has, therefore, the same relations
on the right that the functional ctenidium has on the left to the
transverse blood-sinus. Furthermore, its position with regard to
the hypobranchial mucous gland is precisely that of the right
ctenidium to the right hypobranchial gland in dibranchiate
Rhipidoglossa. Hence, in spite of its distance from the rudi-
mentary right auricle and the fact that, so far as I am able to
determine, 1t recelves no special nerve-supply, I regard it as
a vestige of the right ctenidium. The hypobranchial gland
yarieS considerably in size in the different species of Verita and
Paranerita, but its position is always the same. The mass of
secreting tubules forming the body of the gland lie in the roof
of the right-hand side of the mantle-cavity, and chiefly in the
posterior part of the swelling formed by the genital complex.
Their position in the male of Paranerita gagates is shown in
1908. ] ASPIDOBRANCH GASTROPOD MOLLUSCS. 865
fig. 48, Hy.g., and in the female in figs. 62 & 63, Hyg.
The duct of the gland opens into the right-hand posterior
corner of the mantle-cavity by a fairly large pore situated
below the posterior end of the swelling formed by the genital
complex, and at the base of the vestigial right ctenidium (fig. 49,
Hy.g.a.). As a consequence of the great size of the sper-
matophore-sac and epididymis in Paranerita, the hypobranchial
gland is carried back with it and some of its tubules may lie
at the posterior end of the body, apparently among the viscera,
but really in the tissue surrounding the spermatophore-sac
(figs. 62 & 63) or its equivalent in the male. The tubules are
lined by a simple columnar glandular epithelium, the cells of
which have small nuclei and clear contents. The hypobranchial
gland attains relatively enormous dimensions in the Helicinide,
penetrates among the viscera, and forms a considerable part of
the mass of the visceral spire.
The Generative Organs.
The position of the gonad is the same in both sexes, and the
oviduct or sperm-duct formed by the union of the ducts of the
numerous ovarian or spermatic follicles passes up on the right-
hand of the spire towards the posterior end of the right columellar
muscle and then turns inward to pass into the base of the genital
complex. It is not necessary to add anything to the account
given by Lenssen of the structure and histology of the ovary and
testis in Veritina fluviatilis. It holds good for all the Neritidee
that I have studied.
Lenssen gives a rather summary—but, on the whole, an
accurate—description of the male accessory organs of WV. fluviatilis.
It is open to criticism in matters of detail, but these are not of
sufficient importance to justify a discussion of them in this place.
There is very little difference between the male organs of the
different species of Veritide. Of Nerita I have examined lineata
and melanotraga; of Paranerita, gagates, variegata, and longi-
sping; and I have verified Lenssen’s description of Weritina
fluviatilis. Fig. 50 is a drawing, founded upon sections and
dissections, of the male organs of Paranerita gagates, and those of
variegata, longispina, and of the unknown species from Fiji are
indistinguishable. The sperm-duct is seen passing forward and
upward with a slightly sinuous course close to the external body-
wall on the right side. Just behind the posterior end of the
right columellar muscle it comes into close relations with the
gonadial extension of the cceelom and forms a few coils close against
its wall. At this pomt the sperm-duct turns abruptly to the
left and is immediately thrown into most complicated convolutions
which are closely applied to one another and form a large ovoid
mass—the epididymis. This, as has been explained above, is
enclosed in a fine membranous bag, and is so large that it
projects backwards into the ccelomic cavity and ae across to
866 PROF. G. C, BOURNE ON THE [ Noy. 17,
the left side of the body. Its relations are clearly seen in fig. 44.
It can be seen that the sperm-duct narrows somewhat on entering
the epididymis, forms a great number of very fine convolutions
on its left side, and passing to its hinder end increases appreciably
in diameter. The ultimate coils lie on the right side, are greatly
swollen and filled with spermatozoa. The last coil finally emerges
from the epididymis-sac close to where the sperm-duct entered it,
and immediately contracts to form a rather thick-walled tube,
which I shall call the middle tube. The sperm-duct before it
enters the epididymis is lined by small cubical ciliated cells. On
entering the epididymis the epithelium changes its character,
The cilia are lost; the cells become relatively large—in the
narrower coils three or four suffice to surround the lumen of the
duct,—their cytoplasm becomes dark and granular and their
nuclei clear with a sparse chromatic reticulum. In the thick
terminal convolutions (fig. 55) the epithelial cells are very large
and their cytoplasm is full of deeply-staining granules. So far
as I am able to judge, the spermatozoa mature during their
passage through the coils of the epididymis. In the follicles of
the testis and in the initial part of the sperm-duct they have
globular or ovoid heads, which do not stain deeply, and short
filiform tails. In the large terminal coils of the epididymis the
heads are elongate, stain mtensely with hematoxylin, and the
tails are longer and apparently thicker. They are, however, so
matted together that it is impossible to make out details. But it
is clear that the chromatin has increased in quantity and that the
spermatozoon, as a whole, has increased in bulk. The characters
of the epithelial cells are suggestive of the function of elaborating
material for the nutrition of the spermatozoa. In the middle
tube the epithelium again becomes cubical and richly ciated. The
middle tube passes into the genital complex, runs some way
forward in its wall, makes an abrupt turn backwards, and at once
enters a small thick-walled chamber which I shail call the
thalamus, as 16 evidently corresponds to the chamber into which
the oviduct opens in the female.
There is very little pigment on the course of the sperm-duct in
the species of Paranerita that I have studied: if any is present
it is on the walls of the middle duct, but I have only observed
this in P. gagates.
The thalamus is a narrow and elongated chamber situated in
front of the posterior third and near the inner or left side of the
genital complex (figs. 44 & 48, th.). It is lined by a very long
ciliated columnar epithelium sharply marked off from the ciliated
lining of the middle tube of the sperm-duct. This abrupt passage
from one kind of epithelium to the other indicates, I believe, the
boundary between the mesodermic and ectodermic structures.
Anteriorly and posteriorly the thalamus receives the ducts of a
compact acinous gland which I shall call the prostate (figs. 48 &
AY, prs.). It is clearly formed by evaginations from the wall of
the thalamus and has a distinct lumen, Its epithelial lining is
1908. | ASPIDOBRANCH GASTROPOD MOLLUSCS. 867
shown in fig. 54. It is interesting as exhibiting in its simplest
form a mixed epithelium consisting of ciliated and glandular
cells. The latter are very large with basal nuclei surrounded by
dense granular protoplasm, the outer ends of the cells containing
eosinophilous granules in a protoplasmic reticulum. The ciliated
cells are very small and wedged in between the glandular cells in
a very regular manner. ‘The prostate corresponds to what
Lenssen calls the “glande annexe” in VW. fluviatilis, in which
species it is relatively of enormous size. Lenssen did not observe
that the epithelium is mixed, as just described, but though it is
not so easy to distinguish its characters in the former, there is no
difference between WV. fluviatilis and Paranerita in this respect.
The thalamus, then, is a split-like cavity extending some
distance in front of and behind the entrance of the sperm-duct.
Below and at the sides it communicates widely with the vast
cavity which I shall call the terminal chamber. Lenssen has
called it the ‘poche semilunaire” from its appearance in section.
This is in reality a large pyriform sac, of which the outer
wall remains thin and but slightly glandular, while the inner
wall is greatly thickened by glandular differentiation of its
epithelium and projects like a demi-column into the cavity of the
sac, reducing its lumen to a crescentic slit (figs. 44, 48, & 50).
This projecting glandular column is attached along a slightly
spiral line, is much thicker posteriorly than anteriorly, and its
hinder end loses its attachment to the inner wall and projects
backwards into the cavity of the sac. Hence sections through
this end show not a semi-lunar but a circular cavity surrounding
a central pillar (fig. 49). The thalamus opens into the recess
where the column becomes free from the inner wall. Anteriorly
the terminal chamber diminishes in diameter; its walls become
less glandular and more muscular, but the glandular thickening
on the inner side is continued for some distance forward and
eventually ends in a free projecting process, not far from the
external aperture.
Posteriorly a gland of some size (figs. 48 & 50, 6.g/.) opens into
the terminal chamber. This lke the prostate has a distinct
lumen, and is a saccular outgrowth of the hinder part of the
terminal chamber with much-folded walls. Its epithelium is
wholly glandular, without any admixture of ciliated cells, and the
elements composing it are loaded with globules which stain
deeply in hematoxylin and are therefore probably mucinogenous.
Each globule contains a minute spot, staining brightly in carmine,
and the whole epithelium has a very characteristic appearance,
which I have represented in fig. 56.
That the terminal chamber corresponds to the ootype of the
female there can be no doubt. The structure of the glandular
walls is identical. The outer wall is only feebly glandular compared
with the inner, but its structure, shown in fig. 52, is interesting as
illustrating the steps by which a mixed ciliated and glandular
epithelium, such as that of fig. 54, has become modified into the
368 PROF. G. C. BOURNE ON THE [ Nov. 17,
complex glandular organ depicted in fig. 53. The illustrations
explain themselves and need no comment.
Comparing the above with Lenssen’s account of the male
organs in WV. fluviatilis, it must be observed that in the latter
species there is no posterior gland; the prostate is relatively of
great size; the thalamus (not described by Lenssen, but distinctly
represented) is relatively small and unimportant; the epididymis
is small, deeply pigmented, and does not project into the ccelom,
but remains compacted against the base of the terminal chamber,
and like it slung up to the roof of the mantle-cavity.
The male organs of Verita lineata are depicted in fig. 51.
The epididymis has been partly unravelled to show that it
consists of a long greatly convoluted narrow tube, deeply
pigmented throughout its extent, and a comparatively short, less
deeply pigmented, wide tube. In its natural position the epidi-
dymis is packed closely against the hind end of the terminal sac
and lies in the mantle-roof without projecting into the celom. In
other respects the male organs of this species (and of WV. melano-
traga and plicata, which are indistinguishable) closely resemble
those of Paranerita. It will be observed that Werita resembles
N. fluviatilis in the size, pigmentation, and position of the
epididymis, but differs from it in possessing a posterior gland and
in the relatively small size of the prostate.
Thus there is a close resemblance between the male organs of
the Neritide. They are monaulic; there is no gonaduco-ccelomic
funnel; and Jerita stands midway between Paranerita and
Neritina.
The same relations as regards the different sections are shown
in the female organs. A diagram of the female organs of
Paranerita gagates is given in fig. 60 and sections in figs. 61,
62, & 63. Their structure is very similar in P. longispina and
variegata. The essential features are the same as in Septaria.
The ducts are triaulic. The ovipository aperture les close
alongside of the anus and leads into a large ootype whose cavity
and glandular walls are so closely similar to those of Sepiaria that
they need no further description. At a short distance from its
aperture the ootype gives off dorsally and rather to the right a
diverticulum, which soon enlarges to form a thin-walled sac of
some considerable size filled with spherical calcareous concretions ;
this is, of course, the crystal sac. The hinder end or fundus of
the ootype divides into two passages whose course is somewhat
simpler than in Septaria. That on the right, the egg-duct, after
passing through a “clear gland,” a mucous gland, and a ‘“ rose-
coloured gland,” leads into the thalamus (th.), which is surrounded
by a well-developed vitelline gland. The thalamus receives the
oviduct, and there is a distinct oviduco-ceelomic funnel (fig. 41,
od.c.f.) opening into the gonadial ccelom a short distance
behind the posterior end of the right columellar muscle. The left-
hand passage leads, as in Septaria, into a bilobed sperm-sac, and
this receives the muscular duct of the receptaculum seminis
1908. | ASPIDOBRANCH GASTROPOD MOLLUSCS. 869
(fig. 63, R.s.). The left horn of the sperm-sac is produced into
the ductus enigmaticus, of which the minute opening into the
mantle-cavity is shown in fig. 61, ap.de. Thus tar the identity
with Septaria is nearly complete, but there are differences in
the remaining structures. The right horn of the sperm-sac is
continued into the vaginal canal, which is very long and thrown
into a number of convolutions occupying a considerable part of
the middle third of the genital complex (fig. 62, vag.c.). The walls
of the vaginal canal are very thick and muscular; their thickness
increases at the anterior end, where the canal makes an abrupt
turn backward and opens into the vagina. The form and
relations of the last-named are clearly shown in fig. 60, this part
of the drawing being a careful drawing from a dissection. The
vagina is a thick-walled muscular tube, opening anteriorly into
the mantle-cavity by a slit-shaped aperture rather behind and to
the right side of the ovipository aperture in P. gagates (fig. 61,
vag.ap.). Posteriorly the vagina increases gradually in diameter,
its walls gradually become thinner and less muscular, and its
hinder end expands suddenly to form the thin-walled spermato-
phore-sac, which projects into the ccelomic cavity and bears the
‘same relations to it and the adjacent organs as does the epididymis-
sac in the male. There are some minor differences in the shape
of these organs in the different species of Paranerita. In P. longi-
spina, for instance, the vagina projects some way beyond the
ovipository and anal apertures and its free extremity is coiled in
the shape of $ (fig. 64.). In the same species the vagina is much
longer than in P. gagates, is of the same diameter throughout, and
its posterior end makes a half-turn round the spermatophore-sac
before opening into it.
In P. gagates and variegata the walls of the vagina and vaginal
canal are longitudinally ridged internally. The vaginal canal is
lined by a columnar, ciliated, and non-glandular epithelium. The
vaginal epithelium is made up of larger cells, still ciliated but
more glandular in appearance and filled with chromophilous
granules. The epithelium lining the spermatophore-sac consists
of elongated columnar cells filled with chromophilous granules,
but not, as far as I could determine, ciliated. I cannot, however,
be certain on this point, for the epithelium and contents of the
spermatophore-sac were macerated in all my specimens. The
spermatophore-sac always contains a number of fusiform spermato-
hores embedded in an abundant coagulum. Their structure will
be described later.
The female organs of WVerita differ to a considerable extent
from those of Paranerita and Septaria. They have been con-
cisely described in WV. pica by Thiele (39). Working with
sections he has correctly observed the main features and_ his
drawings of sections are perfectly accurate. But it would take
a much larger number of drawings than he was able to give
to present a clear idea of the complicated ducts and cavities
present, and his diagram gives a very poor idea of the actual
870 PROF. G. C. BOURNE ON THE [Nov. 17,
structure. J must therefore repeat and add to his observations,
but take the opportunity of remarking that he shares with Gilson
and Lenssen the credit of having first elucidated the structure of
these remarkable organs in the Neritide.
I have studied the female organs of Werita plicata, melano-
traga, lineata, and plera. They are all very similar to one another
and to V. pica, as described by Thiele. The following account
founded on JV. plicata may therefore be regarded as applicable to
the subgenus. As shown in the diagram fig. 65, there is a great
similarity as regards the ootype, crystal sac, the various glands
surrounding the egg-duct, and in the thalamus between WVerita
and Paranerita and Septaria. I have already given so sufficient
an account of these structures that [need not recapitulate im this
place, but will confine myself to the differences.
The oviduct takes the usual course close to the right side of the
pedicle of the visceral sac, and opens in the usual place imto the
coelomic cavity by an oviduco-celomic funnel, particularly large
and distinct in iV. melanotraga. Turning into the base of the
genital complex the oviduct passes in front of the receptaculum
seminis and opens into a relatively large thalamus. The position
of the thalamus is indicated in figs. 65, 67, 68, th. Its cavity
is relatively rather large, and the epithelium lining it has the
characters described for Septaria, but is unusually long, and
between the ciliated epithelial cells open the ducts of the uni-
cellular vitelline glands. The oviduct enters about the middle of
the thalamus and its characteristic ciliated, cubical, non-glandular
epithelium is continued downwards and forms the lining of the
inner—that is the left-hand—side of a duct, or rather a fissure, for
although very narrow in transverse section it is of considerable
vertical depth, leading from the thalamus towards the base of the
uterus. This fissure is the equivalent of theegg-duct. Its right-
hand wall is formed by the elongated epithelium bordering a large
glandular mass, which corresponds to the mucous gland in Sep-
iaria. As in Nerita it stains intensely blue with hematoxylin,
T have no doubt that it isa mucous gland. The lowest edge of
this gland is shown in fig. 68, m.gl. At the lower level represented
in fig. 67 the mucous gland is replaced by another which corre-
sponds to the “rose-coloured gland” of Septaria, but the left side
of the egg-duct is still lined by the simple, cubical, ciliated epi-
thelium. Beyond the rose-coloured gland, at a higher level than
that shown in fig. 67, the egg-duct receives through a distinct
duct the secretion of the mucous gland of the opposite side; a-
portion only of this is shown in the figure at m.gl.’. At this
point the egg-duct enlarges, bends very sharply round, and becomes ~
continuous with the lower portion or fundus of the ootype cavity
(Oo.t). The simple ciliated epithelium dies out at the bend and is
no longer seen in this region. Passing upwards again, we find
the ootype extending far forward alongside of the rectum, and in
the middle of it on either side the two lobes of the “ clear gland,”
as indicated in outline in fig. 68. The clear gland soon disappears .
1908. | ASPIDOBRANCH GASTROPOD MOLLUSCS. 87k
and the two sides of the ootype are formed by a thick mass of
the characteristic glandular tissue, but the two ends of its fissure-
like cavity remain very thin.
To return again to the lower level depicted in fig. 67. Some-
what further down than this the right-hand por tion of the wall
of the ootype disappears, and the bottom of its cavity ends in a
large thin-walled sac, which is really the lowest portion of the sac
shown on the right- hand of figs. 65, 67, & 68. This is clearly
comparable to the sperm-sac of ‘Paranerita and Septaria, but it 1s
not bilobed and its structure and further relations in Ni erita are
different, and can best be explained by reference to the diagram
fig. 65. The sac turns upwards and is closely applied to the left
face of the rectum. Its outer or abrectal wall is deeply pigmented,
rendering it a very conspicuous object in dissections, and is
thrown Seo a number of folds r unning on the whole longitudinally.
The pigment is deposited in the bodies of the very distinct band of
columnar ciliated cells which forms the outer wall of the sac, and
may be described as a plicated ciliated band rather than a groove.
This band of ciliated cells, at a lower level than is shown in fig. 67,
passes round the left and lower side of the sac and is continued
into the ciliated epithelium of the egg-duct, which again, as
described above, is continuous with the Giliated ‘epithelium of the
oviduct. The rest of the walls of this pigmented sac are extremely
thin and non-ciliated. It is aremarkable feature that the adrectal
part of the cavity of the sac and also its lower part adjoining the
base of the uterine gland is broken up by a number of trabecule,.
which in arrangement greatly resemble those of the glandular
part of the kidney, but they are not in this case glandular. Some
of these trabecule are shown in fig. 67. They are much more
developed in VV. melanotraga than in NV. plicata, and they have
been noted by Thiele in V. pica. At the top of the pigmented
sac the abrectal ciliated band becomes a distinct groove, and this
separates off as a duct, at first thin-walled, and lined by the same
pigmented ciliated epithelium as the groove. The duct soon
diminishes notably in diameter ; its pigmented ciliated epithelium
gives place to a non-pigmented epithelium composed of much
smaller cells but still ciliated, and at the same time it acquires a.
thick external muscular coat. This duct (fig. 65, vag.c.) pursues a
very slightly sinuous course posteriorly (it is somewhat longer and
more convoluted in V. melanotraga), and while its lumen continues.
to diminish, its muscular coat increases greatly m_ thickness.
(fig. 68, vag.c.). It is evidently, I think, the homologue of the
vaginal canal of Paranerita and Septaria. It opens into a dila-
tation, with less thick but muscular walls, lined by an epithelium
which is continuous throughout all the structures I have yet to
describe and whose characters I will specify presently. From the
lower end of the dilatation a diverticulum is given off, sometimes,
as shown in the diagram fig. 65, in the form of a narrow stalk
with a bulbous termination, sometimes a simple blind tube of
subequal diameter throughout. It contains spermatozoa and is
872 PROF. G. C. BOURNE ON THE [Nov. 17,
evidently the receptaculum seminis. Anteriorly the dilatation
narrows and is continued forward alongside and to the left of the
rectum as the vagina, which opens by a slit-shaped aperture into
the mantle-cavity on the posterior and inner side of the swelling
formed by the terminal portions of the ootype and rectum. At
about two-thirds of its length from its anterior end the vagina
gives off from its posterior side a diverticulum, which at once
expands to form a capacious spermatophore-sac containing several
spermatophores. This does not, as in Paranerita, project into
the celom, but remains closely attached to the dorsal body-wall,
projecting, indeed, very little backwards beyond the receptaculum
seminis. In 1. plicata, in which the kidney (as also in Septaria)
extends far over to the right side of the body, the posterior end
of the spermatophore-sac is partly imbedded in this organ.
The inner walls of vagina, receptaculum seminis, and sperma-
tophore-sac are thrown into longitudinal folds and clothed by a
similar epithelium which has been accurately described by Thiele.
The cells are all of one kind, non-ciliated, with basal nuclei and
clear vacuolated outer ends. They appear to be glandular, but do
not contain the chromophilous granules characterizing the epi-
thelial cells lining the similar structures in Paranerita. In any
case they differ from the ciliated cells of the vaginal canal.
It will be observed that there is no ductus enigmaticus, and the
female organs of WVerita are therefore diaulic, agreeing in this
respect with WNeritina fluviatilis, but differing from Paranerita
and Septaria.
Lenssen has given so sufficient an account of the female organs
of Neritina fluviatilis that I need do no more than say that I
have carefully verified his statements and find them correct. It
is perhaps necessary to repeat that the large thin-walled sae at
the base of the ootype, which he cails the ‘poche de confluence,” is
the equivalent of the sperm-sae of other forms; that what he calls
the “ connecting canal” is the vaginal canal; and that what he
calls the “ bursa copulatrix” is the equivalent of the spermatophore-
sac. J may further point out that the position of the receptaculum
seminis on the course of the vaginal canal is another feature in
which .V. fluviatilis more nearly resembles Werita than Para-
nerita and Septaria.
There is apparently no oviduco-celomic funnel in V. fluviatilis.
I have carefully examined several series of sections in the ex-
pectation of finding it, but have failed to discover a trace. The
oviduct as it passes from the ovary to the thalamus runs in the
wall of the gonadial ccelom, but makes no communication with
this cavity, and I have made use of sufficiently high powers of the
microscope to be able to say that I have not overlooked this
structure because of its minute size. Iam satisfied that it does
not exist.
It is not, perhaps, very profitable to discuss the homologies of
such complicated organs as the genital ducts of the Neritide in
the absence of any exact knowledge of their development, and I
1908. | ASPIDOBRANCH GASTROPOD MOLLUSCS. 873
am not yet ina position to supply this defect in our knowledge
of the group. But as Thiele has homologized the spermatophore-
sac and vagina with the right kidney of other Aspidobranchs, a
criticism of his conclusions will not be out of place.
In considering this question, we must start from the fact
established by von Erlanger (14) and Miss Drummond (18) that
in Paludina the post-torsional right kidney makes its appearance
in the course of embryonic development, but is eventually arrested
and becomes the gonaduct. Miss Drummond has given a very
instructive figure (oc. cit. pl. vii. fig. 6) showing the reno-peri-
cardial opening of the right kidney still open, after the latter has
acquired a connection with the gonad. This is the permanent
condition in female Neritide (except Veritina fluviatilis), and it
cannot be doubted that in this family the gonaduct is, as in Palu-
dina, the representative of the right kidney. This being the case,
the gonopore—that is to say, the opening of the gonad into the
kidney—must be looked for on the course of the oviduct, some-
where behind the reno-pericardial (oviduco-ceelomic) opening.
All that lies in front of the last-named may be kidney, or part of
it may be derived from the mantle-epithelium, either by invagi-
nation or by the closing in of a primitively open groove.
The facts do not warrant our expecting that the kidney and the
gonaduct should have acquired separate openings into the mantle-
cavity as Thiele supposes. Such an expectation, indeed, would be
nonsensical, for the kidney and gonaduct are one and the same
thing. But it is possible—and this, I think, is what Thiele means
—that of the two mantle-openings in the diaulic wVerita one is
the primitive aperture of the right kidney, the other secondarily
acquired, whether by invagination of the mantle-epithelium or
by a secondary outgrowth from the kidney. Basing his opinion
on the histological characters of the epithelium, which in Werita
has, but in Paranerita and Septaria has not, a resemblance to the
epithelium of the left functional kidney, Thiele decides that the
vagina is the true renal aperture and the spermatophore-sac the
representative of the right kidney. He does not push this homo-
logy to its logical conclusion and assert, what must be true if his
view were correct, that the vaginal canal, sperm-sac, receptaculum
seminis, fundus of the ootype, egg-duct, and thalamus as well as
the oviduct itself, are all representative of the right kidney. He
further supposes that in the monaulic male the right kidney has
disappeared. But the sperm-duct no less than oviduct must be
formed from the arrested post-torsional right (pretorsional left)
kidney ; and as the male pore obviously corresponds to the ovi-
pository aperture of the female it would appear more probable
that the latter, and not the vaginal aperture, is the representative
of the primitive uropore. And this, I believe, is the more correct
view of the case.
I have pointed out that the true generative opening into the
kidney must be situated behind the oviduco-ccelomic funnel in
the Neritide. If, now, we make a comparison with the more
874 PROF. G. C. EOURNE ON THE [Nov. 17,.
primitive Aspidobranchia, with Plewrotomaria, Haliotis, Trochus,
Cemoria, we find that the gonaduct enters the kidney at no great.
distance from its external aperture, and close to the reno-pericardial
canal, when this structure is present. The glandular part of the
kidney lies behind the entrance of the gonaduct. It would be
contrary to what we see in all other forms if we were to find, as
we should if Thiele’s view were correct, the glandular part of the
kidney situated in front of the opening of the gonad and the
veno- pericardial canal, between these and the renal pore. The
presence of an anterior lobe of the right kidney in Pleurotomaria
and Haliotis does not invalidate this reasoning, as may readily be
seen on consideration of its relation to the ureter or non-glandular
part of the kidney. But, it may be asked, if the complex of
glandular tissue and ducts lying in front of the oviduco-celomic
funnel in the Neritide do not represent the glandular part of the
kidney, what do they represent? I have no doubt that they are,
in large part, analogous to the modified glandular terminal part.
of the ureter described by M. F. Woodward (41) in Pleurotomaria,
or, to seek a nearer homology, to the glandular sac forming the
ureter in the left functional kidney of the Neritide themselves.
I have already instituted a comparison between this glandular
ureter and the various glands found on the course of the ootype
and egg-duct, and have given reasons for believing that both are
derived from an invagination of the mantle-epithelium. If these
comparisons are correct, the conclusion follows that the ovipository
aperture in the female and the single pore of the male are the
representatives of the ureter of the right side. The vaginal
aperture of the female has therefore nothing to do with the
primitive right renal opening. As to how it has been established
I will not, in the absence of embryclogical evidence, hazard an
opinion. I will merely point out that the formation of accessory
sexual ducts is a common phenomenon. The ductus enigmaticus
of Septariaand Paranerita is an example. So alsoare the vaginal
ducts of the triaulic Doridide and Elysiide. In the Platyhelmia.
multiplication of the female orifices, e. g. in Zrigonoporus, 1s
common; and I do not think it altogether fanciful to say that.
there is some analogy between the Laurer-Stieda canal of Tre-
matodes and the ductus enigmaticus of the Neritide.
The spermatophores of Werita and Paranerita require some
description. They are very similar in general appearance in all
species I have studied. As shown in fig. 69, a spermatophore
consists of a cylindrical body, rounded at one end and produced
at the other end into a long hollow filament. In several cases I
have seen this filament engaged in the aperture of the vaginal
canal, as represented in fig. 64, and extending for a long
distance into its lumen. It is therefore evident that the contents
of the spermatophore—the spermatozoa—are voided through the
filament into the lower end of the vaginal canal, possibly into the
receptaculum seminis, and do not pass into the lumen either of
the vagina or of the upper part of the vaginal canal. Usually
1908. ] ASPIDOBRANCH GASTROPOD MOLLUSCS. 875
there are half a dozen or more spermatophores in the sac, but
sometimes only one. In no case have I found an empty sperma-
tophore-sac. Though I have tried in many different ways, I have
not been able to make any preparations giving a satisfactory
demonstration of the structure of the spermatophores. They are
brittle, and are always contracted and distorted by the action of
reagents ; probably the study of fresh specimens is necessary for
the elucidation of their mechanism. As shown in figs. 66 and 69,
the cylindrical body contains a central cavity filled by a mass of
spermatozoa. The wall of this cavity is formed of a thin layer
of a hard brittle substance which must be of the nature ef chitin.
Around it is a protoplasmic layer (fig. 66, III.), from which a
number of fine filaments radiate to an external wall composed of
an elastic homogeneous substance. The radiating fibres pass
from the inner to the outer walls at regular intervals, so that the
body of the spermatophore appears in a side view to be made up
of a number of segments. The layer of protoplasm surrounding
the central capsule is filled with chromophilous granules, but
there is no trace of nuclei. Both the inner capsule and the outer
wall are continued into the filament, which is therefore a double
tube. There is some evidence that the filament is coiled up
within the capsule and afterwards shot out much as is the filament
of a nematocyst, but of this I cannot be certain. It is a curious
thing, of which I can offer no explanation, that neither I nor any
‘of my predecessors have seen any trace of a spermatophore in the
male organs. It seems certain that they must be formed in the
terminal sac, but there is no positive evidence as to their origin.
It is interesting to note that in the freshwater forms, Septaria
and WVeritina, there are no spermatophores. Gilson has followed
out the process of fertilization in WV. fluviatilis, and it is clear
from his account that spermatophores do not exist in this species.
Lenssen, it is true, alludes to the probability of their occurrence,
but he did not discover them, and my experience is the same
‘as his. J have found in one or two specimens a number of
‘spermatozoa agglutinated together in a mass of coagulum in the
vagina, but I could not detect any structure resembling that of
the spermatophores of Verita and Paranerita. The reduced size
of the spermatophore-sac in Septaria borbonica and S. bougain-
villet affords evidence that spermatophores are not formed in these
‘species. Against this must be set the fact that Thiele describes
a large spermatophore-sac in S. parva, and in fig. 128 gives
the outline of a large irregular mass in its interior, but he
makes no mention of spermatophores.
In fig. 57, a, 6, c, d, e, | have given drawings of the different
‘forms of concretions found in the crystal sac of Nerita melano-
traga. They dissolve readily in dilute acids with evolution of
bubbles, leaving an organic residue in which I could not find any
‘trace of a nucleus. They are composed of a number of crystalline
prisms radiating from the centre of the concretion and projecting
‘on the surface as shown in @and 6. Sometimes the crystals are
876 PROF. G. C. BOURNE ON THE [ Nov. 17,
arranged in several concentric layers as in d, and in other cases,
as in c, a concretion is made up of an aggregate of several smaller
concretions. As the crystal sac is always full of these concretions
in Nerita and Paranerita they must be of some importance,
and I think that they are connected with the formation of an
external calcareous layer of the .egg-shell. The egg-cases of
Nerita and Paranerita are not known, but those of Septaria
bougainvillet have an external calcareous envelope which is readily
dissolved in dilute acids leaving a horny layer beneath. The
crystal sac is very small in Septaria, and this may account for
the calcareous layer being very thin. If this surmise be correct,
the “ crystal sac” is a calcigenous gland, as hinted in a footnote
by Thiele, though for some unexplained reason he prefers to call
it the uterus.
Before bringing this part of my work on the Neritacea to a
close, 1 may conveniently discuss the various questions arising
out of the facts enumerated. In the first place, there is the
question of the inter-relationship of the existing members of the
Neritide. Leaving Scutellina out of the question, because I
have not been able to obtain specimens of this genus, I have to
justify my subdivision of the members of the family into the
groups NVerita, Paranerita, Septaria, and Neritina. The fact
that the female Veritina (sensu restricto) is diaulic and Para-
nerita triaulic is in my opinion sufficient to separate these forms
from one another. Further than this Weritina resembles Verita
more closely than Paranerita, not only in being diaulic, but also
in the characters of the epididymis, in having the epididymis
and spermatophore-sac restricted to the mantle-region instead of
projecting backwards into the ccelom, and in the position of
the receptaculum seminis on the vaginal canal. Veritina, again,
is more specialized than any other of the Neritide in that it has
lost the supra-intestinal nerve and the oviduco-ccelomic funnel.
The evidence of comparative anatomy therefore points to its
having been evolved independently of Paranerita from a marine
Nerita stock, and this conclusion is strengthened by a consider-
ation of the evidence afforded by distribution in space and time.
Assuming, as we are amply justified in doing, that all estuarine
and freshwater forms are descended from marine Neritide,
the various species of Veritina inhabiting rivers debouching into
the Mediterranean, Caspian, and Northern European seas must
have been derived from a marine form inhabiting those seas. At
the present time no member of the genus WVerita (s. stricto) is
found in any of them. Veritina viridis, it is true, is found in
the Mediterranean, but this is probably a fluviatile form which
has found its way back to the sea, for even WV. fluviatilis occurs
in brackish and sometimes in salt water. The ancestral marine
forms must therefore be looked for in geological strata, and it is
significant that, whereas Veritia is common in Tertiary deposits
and extends back as far as the Lias, the most recent fossils
1908.] ASPIDOBRANCH GASTROPOD MOLLUSCS. 877
recognized as members of the genus Verita occur in the Upper
Cretaceous—the subgenus Otostoma, for instance, in the Upper
Cretaceous of Europe, Algeria, and Asia Minor. Making due
allowance for the fact that the distinction between NVerita and
Neritina is not very obvious, especially in fossil shells, it is
clear that the numerous examples of the latter genus found in
freshwater tertiary deposits must have been derived from marine
forms that have long since disappeared from European seas, and
the existence of Weritinw in European secondary strata pushes
back its origin to a remote period. It is probable on the paleon-
tological evidence that the European species form a distinct
geographical group, and the coincidence of anatomical evidence
makes the probability well nigh a certainty.
That Septaria is derived from Paranerita—the females of both
are triaulic—and that the latter is descended from the marine
Nera still abundant in tropical seas, is beyond all doubt. But
the geographical distribution of these forms presents problems
which become more difficult the more one reflects upon them.
The species of Paranerita that I have studied come from localities
as far apart as Fiji and Mauritius, but belong to the Indo-Pacific
region, the marine life of which is tolerably uniform in character.
The anatomy of these species is so similar that they are prac-
tically indistinguishable from one another, though I have noted
small differences between them. Paranerita, however, is only
exceptionally a marine form. Most of the species are fluviatile,
some (e.g. P. cornea from the Philippines) are amphibious or
almost terrestrial in habit, and it is a singular thing that,
although largely continental, they abound in oceanic islands.
Similarly Septaria, an exclusively freshwater genus, is charac-
teristically insular, and species scarcely distinguishable from one
another are found in the Mascarene Islands, in Fiji, and other
Pacific islands. How have these freshwater forms reached their
present habitats? Surely not by the ordinary means of dispersal,
for the animals, adapted as they are to existence in fresh water,
cannot have migrated over the whole Polynesian area, across
great extents of deep ocean. Nor could the egg-cases of Septaria,
which are attached to the shell of the parent, have been wafted
uninjured by any conceivable agency across the Indian Ocean.
If we fall back on the stereotyped explanation that the species
now isolated are representatives of a genus which is still widely
distributed and has been throughout long periods of geological
time, it is still insufficient, for it assumes what will not readily
be granted, the existence of former land-connections between
distant oceanic islands, between the Mauritius Islands, Samoa,
and Fiji. It is a tempting supposition that, as the marine
Nerita is universally distributed in tropical seas and as Para-
nerita is abundant in rivers running into seas where Verita is
abundant, and as the anatomical characters of the two forms
are singularly alike, and as the conchological characters sepa-
rating Paranerita from Nerita are just those which are
878 PROF. G. C. BOURNE ON THE [Nov.17,
characteristic of freshwater shells—viz. relative lightness and
smoothness of shell and operculum due to the greater proportion
of organic over calcareous material, and might therefore be
attributed to the direct influence of external conditions,—then,
wherever circumstances were favourable, marine forms ascended
rivers and as a result of changed conditions of life assumed the
characters which in our artificial systems of classification are
attributed to Paranerita (the tropical Veritina of previous authors).
T can see no @ priori objection to this supposition, for if evolution is
still going on within a group of animals as it has gone on in past
times, marine Neritids must still be passing into estuaries, and
from estuaries into rivers, and as they change their conditions of
life so they must react to their surroundings and undergo modifi-
cations of structure. And as the organization of marine Neritids
is extremely similar in all parts of the world, and as the con-
ditions obtaining in rivers are also very similar, a similar
environment acting upon a similar organization must produce
‘similar results. Let no one object that the environment does
not have a direct influence on the organism. It can be proved
that it has in certain Mollusea. Take a sample of oysters that
have been reared for two years, say, in the Scheide and another
sample reared for a similar period, say, in the Bay of Arcachon.
Their shells will have such distinct and easily recognizable
characters that an experienced eye will have no difficulty in
identifying them. Take both samples and lay them down, say,
at Whitstable, and leave them there for another two years. At
the end of that time the two samples will still be distinguishable
because of the characters of the first two years’ growth. But in
the last two years’ growth they will exactly resemble each other,
and this new growth will have neither Schelde characteristics
nor Arcachon characteristics, but Whitstable characteristics
differing from both the former. These facts are well known to
oyster-merchants, and I have personally verified them. They
are proof of the direct action of the environment on the grow-
ing shell, and if only conchological evidence were forthcoming,
T should be inclined to accept the supposition put forward
above. Indeed, before I made a detailed study of the genital
organs, I thought that it was the best explanation of the problem,
but when I found that the females of Verita were diaulic, and
those of Paranerita triaulic, and that there were parallel dif-
ferences in the male organs, the explanation no longer satisfied
me. It is inconceivable that such a structure as the ductus
enigmaticus could have been independently evolved several
times over. At the same time I think it probable that many of
the fluviatile Neritidee have been independently derived from
marine Neritide. and I have entered upon this discussion in the
hope that others will make an anatomical examination of species
from different localities and determine how far they differ from
one another. An anatomical study of freshwater Neritide from
the Atlantic seaboard 1s very much to be desired.
1908. | ASPIDOBRANCH GASTROPOD MOLLUSCS. 879
As to the larger questions of the relationship of the Neritide
to other groups of the Rhipidoglossa, and whether they can be
regarded as representatives of the stock from which the Pectini-
branchia were derived, I think that some evidence can be offered
from the foregoing pages. The Neritide are commonly held to
be a highly specialized and at the same time an annectant group
leading to the Architenioglossa. Such a view, though it may
contain an element of truth, cannot, in my opinion, be held
without considerable modification.
To deal first with the position of the Neritide among the
Rhipidoglossa. Thiele admits that he cannot assign them a
satisfactory position. He is inclined to derive them from the
Trochidz, but also detects resemblances to the Acmzide; as he
rightly observes, they cannot be derived from the latter family,
for it is docoglossate.
If we compare the Neritide with the rest of the Rhipidoglossa
we see that they possess a number of primitive characters common
to the whole group. Such are, the short snout, without pre-
tentacular or post-tentacular elongation ; in the nervous system,
the presence of a labial commissure, of elongated buccal ganglia,
of a long cerebral commissure, of elongate pedal nerve-cords
sheathed in ganglion-cells. The pharyngeal bulb is large, situated
far forward in the head and embraced by the cerebral com-
missure. The ventricle of the heart is traversed by the rectum,
and although that of the right side is much reduced there are two
auricles. The ctenidium is typically bipectinate and is less
modified than in the Trochide, for the lower gill-lamelle are
equal in size to the upper.
The characters indicating a higher degree of specialization than
in other Rhipidoglossa are as follows:—The eyes are closed,
the supra-intestinal nerve is reduced or even absent, and the
left symmetrical pallial nerve takes a principal share in the
innervation of the ctenidium. The subintestinal ganglion is
closely approximated to the right pleural and is united by a very
short zygoneurous branch with the latter. A direct commissural
connection is established between the right and left pleural
ganglia. Only a single functional kidney—the left post-torsional
is present, its fellow of the right side having been converted
into the gonaduct. The accessory genital ducts are extremely
complicated and in the female are diaulic or triaulic. The
visceral spire is reduced and the animal has acquired a secondary
symmetry emphasized by the presence of a left as well as a right
columellar muscle. In possessing a single (left) ctenidium the
Neritide stand on the same level of organization as the Trochidze
or Turbonide.
It might be concluded from all this that the Neritide are
nothing more than Rhipidoglossa which have been specialized in
certain directions while retaining many of the primitive features
characteristic of all the Aspidobranchia, and that they so far
resemble the monobranchiate forms (Trochide and Turbonide)
Proc. Zoo. Soc.—1908, No. LVI. 56
880 PROF. G. C. BOURNE ON THE [Nov. 17,
that they have probably descended from them. But this con-
clusion cannot be sustained. In some important respects the
Neritide are more primitive than any of the Rhipidoglossa.
The celom, in particular, extending as it does across the whole
width of the body, retains features which may be called embryonic
when we compare it with von Erlanger’s and Miss Drummond's
account of the development of Paludina, but must surely be
regarded as primitive when we consider the probable phylogeny
of the Gastropoda. No such extensive cceelomic space has been
described in any other gastropod, and to find a parallel to it we
must refer, as Thiele has done, to the Cephalopoda. When a
feature is shared by representatives of two orders now widely
separate, and is also shown by embryological evidence and by
a priori reasoning to be primitive, there are very good grounds
for regarding it as ancestral. In other Rhipidoglossa the ccelom
is reduced to a pericardial sac surrounding the heart. This
pericardial sac, as embryology teaches us, is the reduced repre-
sentative of a primitively much more extensive space. This more
extensive space is preserved in the Neritide, and the conclusion
is that they have inherited it from ancestors more generalized in
this respect than the remainder of the existing Rhipidoglossa.
This ancestor must have been older even than Pleurotomaria, for
the ccelom is reduced to a pericardial sac in this genus.
The same conclusion is reached by a consideration of the
excretory organs. In Pleurotomariide, Haliotidz, Trochide,
Turbonide, and Fissurellide the post-torsional left kidney is
reduced to a small sac, and in the first four families this
‘papillary sac,” as it is called, has undergone modification. It
no longer serves for the elimination of waste matters from the
blood, but is phagocytic. In the Neritidz the left post-torsional
kidney is large and persists as the functional excretory organ ; it
is the right kidney that has changed its function and undergone
reduction. It cannot be doubted that the ancestral Gastropod
possessed paired functional kidneys as do the Chitonide and
among the Fissurellide Cemoria (fide Haller). The obvious
inference is that the families in which the left kidney is modified
to form a papillary sac are to that extent modified, and that the
Neritidee are descended from an ancestor in which this modifica-
tion had not yet taken place. The persistence of the left kidney
in Neritide, therefore, is to be regarded as an ancestral rather
than as a specialized character, and as evidence that this family
cannot have been descended from Trochide or Turbonide, in which
specialization has taken a different direction.
These arguments indicate that the Neritide are descendants of
a very primitive stock (a conclusion sufficiently supported by
Paleontology), from which the remainder of the Rhipidoglossa
and probably other groups of Gastropoda were also derived.
If these conclusions are accepted, the question of the relation-
ship of the Neritide to the Tzenioglossa is simplified. In this
case we have to consider whether the special characters of the
1908. | ASPIDOBRANCH GASTROPOD MOLLUSCS. 881
Neritide appear, and if so, to what extent, in the more primitive
Tenioglossa. If they do not reappear, it cannot be maintained
that the latter group is descended from the former. Secondly,
whether such resemblances as there may be between the two
groups may be attributed to inheritance from a common ancestor
or to convergence, or whether possibly both these factors have
taken a share in producing these resemblances, and, if so, what
share.
Taken by itself, the persistence of the left post-torsional kidney
as the functional excretory organ in the Neritide and Pectini-
branchia would be strong evidence of their relationship, but ,
relationship does not imply that one group is descended from the
other. It may be remote and may only indicate that both
groups are descended from a common ancestor, and this is clearly
the true conclusion in the matter. Among the special characters
of the Neritide those of the nervous system are the most
important. If the Pectinibranchs were descended from a Neritid
stock, we should expect to find in the more primitive members of
the suborder traces of the special features of the Neritid nervous
system. But we find nothing of the sort. The generalized
Pectinibranchs such as Paludina, Cyclophorus, Littorina, or
Cyclostoma are typically dialyneurous. In none of them is the
subintestinal ganglion approximated to the left pleural. There is
not a zygoneurous connection between the right pleural and the
subintestinal ganglia. There is no trace of a direct commissural
connection between the right and left pleural ganglia. The
supra-intestinal nerve shows no sign of reduction or disappearance.
Paludina, as Bouvier has shown, is quite rhipidoglossan in
respect’ of its nervous system. The evidence is clear that the
archaic Teenioglossa cannot have descended from the Neritide.
On the other hand, the persistence of the left kidney as the
functional excretory organ, and the fact that the permanent
relations of the reduced right kidney (gonaduct) of female
Neritide to the ceelom almost exactly represent an embryonic
phase in Paludina, are coincidences which must almost certainly
be attributed to inheritance from a comyon ancestor. At some
remote age the Prorhipidoglossan stem must have divided into
two branches. In one the left kidney underwent reduction, and
this branch gave rise to the Pleurotomariid, Haliotide, Trochide,
Fissurellide, and probably also to the Docoglossa. In the other
branch the left kidney retained its size, and with the reduction
of the right kidney became the only excretory organ. From
this branch all the Gastropoda which retain the left kidney only
—the Neritide, the Pectinibranchia, and the Kuthyneura—are
descended. The last two must have branched off at a very early
period, while the ancestral form still retained all the primitive
characters of the nervous system, as these are preserved in
Paludina and in Acteon. The Neritacea remain as the much
modified representatives of the primitive stock. Their special
characters are peculiar to themselves and are not to be explained
56*
882 PROF. G. C. BOURNE ON THE [ Nov. 17,
by reference to any other existing group. Thus the resemblance
of the shell of some members of the group to the Naticide is due
to convergence. So also the possession of a single functional
etenidium on the left side, a character shared by the Trochidee
and Pectinibranchs, does not indicate that the Neritide stand
midway between these forms, but only that the right ctenidium
has been suppressed independently in each. I have brought
forward evidence showing that the loss of the right ctenidium is
comparatively recent in the Neritide, probably as a consequence
of the great development of the accessory genital organs which
oceupy all the space on the right side of the spacious mantle-
cavity.
A consideration of the Helicinide, the most specialized of the
Neritaceze, must be postponed to the second part of this memoir.
List OF THE PRINCIPAL WORKS REFERRED TO.
1. Amauprour, A.—La partie antérieure du tube digestif et la
torsion chez les Mollusques gastéropodes. Ann. des Sci. Nat.
(8) vii. 1898, p. lL.
2. Bercu, R.—Die Titiscanien, eine Familie der rhipidoglossen
Gasteropoden. Morph. Jahrb. xvi. 1890, p. 1.
3. Bernarpb, F.—Recherches sur les organes palléaux des Gas-
_téropodes prosobranches. Ann. des Sci. Nat. (7) 1x. 1890,
. 88.
4. ee ieee F.— Ueber die Entwicklung der Neritina fluvi-
atilis Mill. Zeits. f. wiss. Zool, xxxvi. 1882, p. 125.
5. Buunpsrong, BE. R.—Glycogen in Vesicular Cells of Molluses.
Proc. Roy. Soc. Lond. xxxvili. 1885, p. 442.
6. Bouran, L.—Sur le systeme nerveux de la Werita polita.
Comptes Rendus Acad. Sci. Paris, exiv. 1892, p. 1133.
7. Ibid Mémoire sur le systeme nerveux de la Verita polita et
de la Navicella porcellana. Arch. de Zool. expér. et gén. (3)
i. 1893, p. 221.
8. Bouvier, E. L—Le systéme nerveux et certains traits
WVorganisation des Neritide et des Helicinide. Bull. Soe.
Philomath. de Paris, (7) x. 1886, p. 93.
9. Ibid.— Systeme nerveux, Morphologie générale et Classification
des Gastéropodes prosobranches. Ann. des Sci. Nat. (7) iii.
1887.
10. Ibid.—Le systeme nerveux des Néritidés. Comptes Rendus
Acad. Sei. Paris, cxiv. 1892, p. 1281.
11. Brock, J.—Zur Neurologie der Prosobranchier. Zeits. f.
wiss. Zool. xlvii. 1889, p. 67.
12. CuararreDE, E.—Anatomie und Entwicklungsgeschichte der
Neritina fluviatilis. Miiller’s Arch. f. Anat. und Physiol.
1857, p. 109.
13. Drummonp, 1. M.—The Development of Paludina vivipara.
Quart. Journ. Micr. Sci. xlvi. 1902, p. 97.
14. Eruancer, R.von.—On the paired Nephridia of Prosobranchs.
Quart. Journ. Mier. Sci. xxxii. 1892, p. 587.
1908. | ASPIDOBRANCH GASTROPOD MOLLUSCS. 883
15.
16.
We
18.
19;
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.
34.
35.
36.
37.
Fiscuer, P.—Manuel de Conchyliologie. Paris, 1887.
Fiemmine, W.—Ueber Bindesubstanz und Gefisswandung im
Schwellgewebe der Muscheln. Arch. f. mikr. Anat. xiii.
1877, p. 818.
Frievre, H. J.—Notes on the Relations of the Kidneys in
FHaliotis tuberculata ete. Quart. Journ. Mier. Sci. xlvi. 1902,
Dende
Gitson, G.—The Female Organs of Veritina fluviatilis. Proc,
Malac. Soc. Lond. 11. 1896, p. 81.
Hauer, B.—Untersuchungen iiber marine Rhipidoglossen.
Morph. Jahrb. ix. 1884, p. 1.
Ibid.—Studien iiber docoglosse und rhipidoglosse Proso-
branchier. Leipzig, 1894.
Tbid.— Betrachtungen itiber die Phylogenese der Gonade und
deren Miindungsverhiiltnisse bei niederen Prosobranchiern.
Zool. Anzeiger, xxiii. 1900, p. 61.
JHERING, H. von.—Vergleichende Anatomie des Nerven-
systems und Phylogenie der Mollusken. Leipzig, 1877.
Ibid.—‘‘ Gibt es Orthoneuren?” Zeitsch. f. wiss. Zool. xlv.
1887, p. 499.
LanpsBEeRG, B.—Ueber das Herz und die Niere von NVeritina
fluviatilis. Zool. Anzeiger, v. 1882, p. 661.
Lenssrn, J.—Systeme digestif et systeme génital de la Ver-
tina fluviatilis. La Cellule, xvi. 1899, p. 177.
Ibid.—Systéme nerveux, systéme circulatoire, syst¢me respi-
ratoire et systéme excréteur de la Veritina fluviatilis. La
Cellule, xx. 1902, p. 289.
Martens, F. von.—Die Gattungen WNerita und Neritopsis.
Systematisches Conchylien-Cabinet, Bd. 11. Heft 27, 1889.
Moguin-Tanpon, A.— Histoire naturelle des Mollusques
terrestres et fluviatiles de France. Paris, 1855.
PELSENEER, P.—‘‘ Gibt es Orthoneuren?” Bull. Sei. de la
France et de la Belgique, (3) xix. 1888, p. 46.
Ibid.— Classification générale des Mollusques. Bull. Sci. de
la France et de la Belgique, (4) xxiv. 1892, p. 347.
Tbid.—Les reins, les glandes génitales et leurs conduits dans
les Mollusques. Zool. Anzeiger, xix. 1896, p. 140.
Tbid.—The Mollusca, in a Treatise on Zoology, edited by
E. Ray Lankester.—Part V. London, 1906.
Perué, J.—Die Kreide- (Hypersenon) Fauna des Peter-
wardeiner Gebirges. Paleeontographica, li. 1906, p. 109.
Perrter, R.—Recherches sur l’anatomie et Vhistologie du
rein des Gastéropodes prosobranches. Ann. des Sci. Nat.
(7) vii. 1889, p. 61.
Qvoy et Gaimarp.— Voyage autour du Monde: Zoologie.
Paris, 1824, p. 460.
Ibid.— Voyage de l’Astrolabe: Zoologie, ii. 1834.
RAnDLEs, W. B.—Some Observations on the Anatomy and
Affinities of the Trochide. Quart. Journ. Micr. Sei. xlvii.
1904, p. 33.
884 PROF. G. C. BOURNE ON THE [ Nov. 17,
38. Srmmrorn, H.—Gastropoda prosobranchia. Bronn’s Tierreichs:
Mollusca, 1896-1907.
39. Turee, J.—Die systematische Stellung der Solenogastren und
die Phylogenie der Mollusken. Zeitschr. f. wiss. Zoologie,
Ixxii. 1902, p. 249.
40. Tryon, G. W.—Manual of Conchology, vol. x. Philadelphia,
1888.
41. Woopwarp, M. F.—The Anatomy of Plewrotomaria beyrichit.
Quart. Journ. Micr. Sci. xliv. 1901, p. 215.
42. Woopwarp, 8. P.—A Manual of the Mollusca. London,
1875.
EXPLANATION OF THE PLATES.
Prates XLVI-LXVI.
Lettering in all the figures, except figs. 36-39.
af.br. Afferent branchial vessel. mf. Muscle-fibres.
An. Anus. | m.gl. Mucous gland.
ap.de. Aperture of the ductus m.t. Middle tube of gonaduct.
enigmaticus. O. Otocyst.
ax. Axial plate of ctenidium. | od. Oviduct.
b.gl. Basal gland of the male | od.ef. Oviduco-ceelomic funnel.
genital organs. | Oe. (Esophagus.
bpm. Branchio-pallial nerve. | Oot. Ootype.
br.x. Branchial nerve. Oot.ep. Kpithelium of ootype.
b.si. Blood-sinuses of kidney. | Op. Opercular lobe.
c.ap. Aperture between pericardial Os.g. Osphradial ganglion.
and gonadial ccelom. | Ov. Ovary.
cil.e. Ciliated cells. | Ov.ap. Ovipository aperture.
cil.ep. Ciliated epithelium. P. Pedal nerve-centres.
cil.p. Ciliated passage into ureter. | pe.co. Pericardial celom.
ceil.tr. Ciliated tract of gill-lamelle. | Ph. Pharyngeal bulb.
el.gl. Clear gland. | prs. Prostate.
em.l. Lett columellar muscle. | R. Rectum.
em.r. Right columellar muscle. | R.auw. Right auricle.
Cp. Cephalic penis. R.ct. Right ctenidium.
Cry.s. Crystal-sac. re.gl. Rose-coloured gland.
Cé. Ctenidium. Rd. Radula.
D.en. Ductus enigmaticus. | R.p.c. Reno-pericardial canal.
H. Kye. R.s. Receptaculum seminis.
Hd. Ege-duct. | £&.s.d. Duct of receptaculum seminis.
ef.br. Kifferent branchial vessel. S.g. Salivary gland.
Epd. Hpididymis. Sp. Spermatophores.
fr.cil. Frontal cilia. sp.d. Sperm-duct.
g-co. Gonadial coelom. sp.ep. Epithelium of sperm-duct.
Gd. Genital complex. Sp.s. Sperm-sac.
gl.c. Gland-cells. Spz. Spermatozoa. |
Hy.g. Hypobranchial gland. St. Chsophageal division of stomach.
Hy.g.a. Aperture of hypobranchial St’. Pyloric division of stomach.
gland. T. Tentacle.
Int. Intestine. tch. Terminal chamber.
int.c. Interstitial cells. th. Thalamus
K. Glandular portion of the Ts. Testis.
kidney. Ur. Ureter.
Kk’, Bladder or non-glandular por- Ur.p. Uropore.
tion of the kidney. V. Ventricle of heart.
L.au. Left auricle. Vag. Vagina.
Ti. Liver. Vag.ap. Vaginal aperture.
Li.d. Hepatic ducts. | Vag.c. Vaginal canal.
“msc. Longitudinal muscles of gill. | —_—-vis.gn. Visceral ganglion.
M. Mantle. | vis.n. Visceral nerve.
M.c.. Mantle-cavity. vt.g. Vitelline gland.
M.ep. Mantle-epithelium.
1908. | ASPIDOBRANCH GASTROPOD MOLLUSCS. 885
Fig.
Fig.
Pratt XLVI.
1. Dorsal view of Septaria borbonica, 2. The mantle has been cut through
and partly removed to show the organs of the pallial complex (x 5).
2. A semi-diagrammatic drawing to show the course of the gut in Septaria.
3. A diagram of the female genital organs of Seplaria.
Pratt XLVII.
4. A transverse section through the anterior part of the viscera mass of
Septaria bougainvillei, showing the connection between the pericardial
and gonadial divisions of the ccelom aud the oviduco-ccelomic funnel,
od.c.f.
5. The oviduct and oviduco-ccelomic funnel as shown in the last figure, but
magnified 300. This figure is reversed.
Prares XLVIII.-L.
Figs. 6-16. Transverse sections taken at different levels through the genital complex
Fg.
Fig.
Fig.
a
Fig.
of Septaria bougainvillei, 2. For full description of these figures, see
text, pp. 839-841.
17. Septaria bougainville?, section through the osphradium and osphradial
ganglion. XX 300.
Prate LI.
18. Portion of a horizontal section through Septaria depressa, showing the
uropore (U7.p.), the opening of the reno-pericardial canal (Rp.c.) into the
glandular part of the kidney (K.), the pericardium (pe.co.), &c. :
19. A section more ventral than that shown in fig. 18, showing the opening of
the reno-pericardial canal ito the pericardium, and the large extent of
the non-glandular part of the kidney.
Prats LILI.
. 20. An oblique section through the left or efferent edge of the ctenidium of
Septaria bougainvillei.
21. The more median part of the same section, showing the ciliated junctions
of the gill-lamelle.
22. A section through the right or afferent edge of the same ctenidium.
23. A surface view of a single gill-lamella of Septaria horbonica, showing the
relative extent of the ciliated and non-ciliated tracts.
24. The ciliated epithelia of the terminal portions of the ovipository and vaginal
ducts of S. bougainvilleit. Highly magnified.
25. The ootype epithelium somewhat further down the duct, showing gland-
cells containing eosinophilous granules pushing through the basemeut-
membrane into the subjacent connective tissue.
Puatre LIII.
26. A portion of the ootype gland and ootype epithelium of S. bougainvillei.
X about 200.
27. A portion of the “clear gland” from the genital ducts of S. bougainvillet,
2 , showing the elongated ciliated epithelial cells lining the lumen of the
duct, and the bunches of clear secretory cells the long ducts of which pass
between the epithelial cells. Magnified about 750.
28. A portion of the epithelial living of the thalamus and the vitelline gland
of S. bougainvillei, 2, showing the bunches of granular secretory cells
and their ducts filled with granules. Magnified about 500.
Puate LIV.
29. Left side view of N. (Paranerita) gagates, removed from its shell.~ X 5.
The pericardium has been opened to show the relative positions of heart,
ctenidium, rectum, and kidney.
30. The right tentacle and cephalic penis of Neritina fluviatilis. X 10.
31. The right tentacle and cephalic penis of Nerita melanotraga. X95.
32. The right tentacle and cephalic penis of NV. (Paranerita) gagates. X 5.
33. A horizontal section through the cephalic penis of NW. (Paranerita)
variegata.
34, A portion of the opercular gland of Nerita melanotraga, showing the
eland-cells in different phases of activity and the interstitial or covering
cells (int.c.).
886
Fig. 35.
36.
37.
Fic. 42,
43.
Fig. 47.
48.
49.
PROF. G. C. BOURNE ON THE [Nov. 17,
Prate LV.
View of the pharyngeal bulb and alimentary tract of NV. (Paranerita)
longispina unravelled. :
A dorsal view of the pleuro-pedal nerve-centres and principal nerves
issuing from them in NV. (Paranerita) gagates. ‘The lettering in this
and the succeeding figures is the same as that adopted by Bouvier (9).
Cd. Right pleural ganglion. Cg. Left pleural ganglion. O. Otocyst.
P. Pedal centres. 61. Left symmetrical branchial nerve. d!, d?. Anterior
pallial (parietal) nerves of the right side. e!, e?. Anterior pallial (parietal)
nerves of the left side. f#. Supra-intestinal nerve. /!. Subintestinal
nerve. kl, Cerebro-pedal connective. k?. Cerebro-pleural connective.
ld. Right columellar nerve. 7g. Left columellar nerve. m. Pallial
branch of the left branchio-pallial nerve. m!. Right pallial nerve.
op. Opercular nerves, issuing from the upper surface of the pedal centres.
s. Branches of the cerebro-pleural connectives passing to the walls of the
head. Sd. Subintestinal ganglion.
A section taken somewhat obliquely through the pleural and pedal centres.
2. Origin of the lower of the two direct commissural connections between
the pleural ganglia. Z. Zygoneurous connection between the right
pleural ganglion and the subintestinal nerve. Other letterimg as in
fig. 36.
Pratt LVI.
. A section through the pleural centres posterior to that shown in the
preceding figure. 1&2. Upper and lower direct commissural connections
between the pleural ganglia.
9. A still more posterior section through the pleural centres.
. A transverse section through the proximal end of the subintestinal nerve,
showing two bundles of nerve-fibres.
. Part of a longitudinal section through WN. (Paranerita) gagates, 9 , showing
the relation of the spermatophore-sac (Sp.s.) to the gonadial division of
the ccelom (g.co.) and the opening of the oviduco-ccelomic funnel (od.c.f.)
into the latter.
Prate LVII.
A horizontal section through N. (Paranerita) gagates, 8, showing the
two divisions of the stomach, the extent of the lower part of the peri-
cardial ccelom, and the relations of the right auricle.
A similar section taken at a somewhat higher level, showing the extension
of the pericardial ccelom towards the right side and the opening of the
reno-pericardial funnel into it.
Prate LVILII.
. A similar section, more dorsal than fig. 43, showing the relations of the
epididymis to the viscera.
. A section through the uropore, more highly magnified, showing the relations
of the visceral ganglion and reno-pericardial canal.
. A section showing the opening of the reno-pericardial canal into the peri-
cardial division of the celom in WN. (Paranerita) variegata and the
extension of the ciliated epithelium of the ureter (ci.ep.) into the
bladder,
Prats LIX.
A section through the ureter and adjacent part of the kidney in W. (Para-
nerita) variegata, showing the opening of the reno-pericardial canal into
the glandular part of the kidney and the ciliated passage (cil.p.) leading
directly from the latter into the ureter.
A eee section through the genital complex of WN. (Paranerita)
gagates, 8.
Part of a horizontal section through WN. (Paranerita) variegata, showing
the opening of the hypobranchial gland (Hy.g.a.) into the mantle-cavity.
1908.]_ - ASPIDOBRANCH GASTROPOD MOLLUSCS. 887
Fig. 50.
51.
~
54,
57.
58.
59.
Fig. 60.
61.
Vig. 62.
63.
Fig. 65.
52.
Pratt LX.
The male organs of NV. (Paranerita) gagates, semi-diagrammatic.
A similar representation of the male organs of Nerita lineata.
Part of a section through the outer wall of the terminal chamber of the
male ducts of WV. (Paranerita) variegata, illustrating the formation of
bunches of unicellular glands in the connective tissue underlying the
epithelium of the chamber. X about 1000.
Prate LXI.
. Part of a section through the inner wall of the terminal chamber of the
same species, showing a bunch of unicellular glands the ducts of which pass
between the ciliated epithelial cells lining the chamber. XX about 1000.
Part of a section through the prostate of the same species, showing the
gland-cells and ciliated interstitial cells (cil.c.). > about 1000.
. Part of a section through the terminal coils of the epididymis of the same
species, showing the tube filled with spermatozoa and the granular non-
ciliated cells (ep.) lining the tube.
Prate LXII.
. Part of a section through the basal gland of the male organs of the same
species, showing groups of gland-cells contaiming vesicular bodies, each
with a brightly staiming spot.
a, b,c, d, e. Different forms of concretions from the crystal sac of Nerita
melanotraga.
A diagram illustrating extent and relations of the coelom in Paranerita.
Reticular connective tissue and metabolic cells from Neritina fluviatilis,
X about 1000. vet. Protoplasmic reticulum with nuclei. set. Metabolic
cells with granules.
PratE LXIII.
The female organs of N. (Paranerita) gagates, semi-diagrammatic. This
figure, as also figures 64 and 65, represent the organs as they appear when
dissected from the ventral surface. The ootype and rectum, which in their
natural position lie dorsad of and partly to the left of the vagina, are
thrown over to the right side of the figure.
A longitudinal section through the anterior part of the genital complex of
N. (Paranerita) variegata, showing the vaginal aperture and the aperture
of the ductus enigmaticus.
‘Pirate LXIV.
A section from the same series through the middle of the genital complex.
A section from the same series through the lower part of the genital
complex.
. A drawing of the vagina and spermatophore-sae of NV. (Paranerita) longi-
spina: the spermatophore-sac has been laid open.
Prate LXV.
A semi-diagrammatic representation of the female organs of Nerita
plicata.
66. A section of a spermatophore of Nerita plicata. I. Central capsule filled
67.
with spermatozoa. II. Chitimous wall of the central capsule. III.
Layer of protoplasm with chromophilous granules. 1V. Radiating fibres.
V. External wall.
A transverse section passing through the bottom of the fundus of the
ootype of Nerita plicata, illustrating the relations and structure of the
sperm-sac. For further description see pp. 870 & 871.
Puate LXVI.
Fig. 68. A section from the same series as fig. 67, taken at a higher level and
showing the connection of the thalamus with the fundus of the ootype.
69. A spermatophore of Nerita plicata.
888 DR. N. ANNANDALE ON AN [| Nov. 17,
2. An unknown Lemur from the Lushai Hills, Assam.
By N. Aynanpaz, D.Sc., C.M.Z.S., Superintendent,
Indian Museum.
[ Received November 3, 1908. |
(Text-figure 173.)
My. T. D. La Touche, of the Geological Survey of India, has
recently shown me a remarkable photograph taken by himself
during the Lushai Expedition of 1889-90. It represents two
individuals of a small mammal evidently allied to the Slow Lemurs
(Vycticebus), but differing from all known Asiatic species of the
order in possessing a thick, bushy tail. The photograph is not
clear as regards the tail, but Mr. La Touche assures me that
it was present.
Text-fig. 173.
An unknown Lemur from Assam.
The new Lemur is white in colour, with a narrow black
mid-dorsal stripe extending from the occiput to the base of the
1908. ] UNKNOWN LEMUR FROM ASSAM. 889
tail, a dark triangular patch round each eye, and the anterior
surface of the ears dark. The tail is apparently very thick and
cylindrical, shorter than the head and body, and without definite
markings. The lmbs are comparatively short and stout. The
head is large and round, the face flat, the muzzle small, the ears
short and rounded; the eyes are perhaps a little smaller than
those of Nycticebus tardigradus, but are separated by less than
their own diameter. The fur is apparently close and woolly.
Hab. “Caught near Fort Lungleh, Dec. 1889” (La Touche).
Evidently an inhabitant of dense jungle on the outer ranges of
the Lushai Hills, Assam.
Mr. La Touche tells me that the individuals he photographed
were caught in the jungle and escaped from captivity after a
short confinement. They were habitually so slow in their move-
ments that no precautions were taken to. prevent their escape ;
but when once they had got out of their cage they vanished
rapidly. They were fond of hanging upside down, as the upper
animal of the photograph (text-fig. 173) isdoing. It will be noticed
that in this position the tail does not hang down but is supported
against the side of the box. Possibly it is prehensile, but this
is not clear. The lower animal in the photograph is evidently
asleep. It sits with its head tucked in under its chest, much as
WVycticebus does; the tail is also tucked in under the body.
Possibly on examination the skull of this interesting Lemur
would show further differences from the known Indian genera,
WVycticebus and Loris. Nycticebus and Loris have a rudimentary
tail or no tail at all; Tarsius, the only other Asiatic genus, which
is the type of a separate family and only occurs in the Malay
Archipelago, has a long, thin tail with a tuft at the end. The
closest extra-Asiatic allies of Vycticebus and Loris are the Pottos
(Perodicticus) from W. Africa, which have short or rudimentary
tails. Even the type of coloration of La Touche’s Lemur, however,
so closely resembles that of the Indian forms that it is impossible
that the new genus is widely separated from them. Among the
Madagascar genera, moreover, /ndris—like Tarsius, the type of a
separate family—has a rudimentary tail; and even in the genus
Perodicticus, as it is now defined, there is a considerable difference
in the length of this organ in different species.
[Since the above was written I learn from Col. E. W. Loch
that the tailed Lemur of the Lushai Hills is well known to him.
I defer the publication of a technical description and the naming
of the genus until it has been possible to examine specimens.—
January 5, 1909. |
890 MR. R. I. POCOCK ON [ Dee. 15,
December 15, 1908.
Dr. Henry Woopwarp, F.R.S., Vice-President,
in the Chair.
The Secretary read the following report on the additions
made to the Society's Menagerie during the month of November
1908 :—
The number of registered additions to the Society's Menagerie
during the month of November was 91. Of these 50 were acquired
by presentation, 18 by purchase, 12 were received on deposit, 10 by
exchange, and one was born in the Gardens.
The number of departures during the same period, by death
and removals, was 190.
Amongst the additions special attention may be directed
to :— ;
Two Walruses (Odobenus rosmarus) from Franz Josef Land ;
purchased on Nov. 23rd.
One Sumatran Civet (Viverra tangalunga) from Sumatra; pre-
sented by A. R. Heath, Esq., on Nov. 24th.
Two Pardine Genets (Genetta pardina) from Warri, Southern
Nigeria ; presented by E. G. Stevens, Esq., on Dec. 13th.
Mr. Frederick Gillett, V.P.Z.S., gave an account of his recent
Hunting Trip to the Thian Shan, illustrated by lantern-slides.
Mr. R. I. Pocock, F.L.S., F.Z.8., Superintendent of the Society’s
Gardens, exhibited photographs of a Sumatran Tiger, recently
purchased by the Society, and made remarks upon this animal and
upon the other Tigers at present living in the Gardens. He
said :—‘ This Tiger, a male, was one of a litter obtained by
Mr. Pinckney at Deli in Sumatra. Its ground-colour is noticeably
darker and duller in hue than in the Indian and Siberian Tigers
in the adjoining cages. The stripes are numerous, closely placed,
and broad, nearly all of those on the sides of the body, behind the
shoulders, and on the hind-quarters, being looped or reduplicated.
The shoulder is scantily striped, and the outside of the fore leg
nearly unstriped, except for one or two narrow stripes across the
wrist and a few abbreviated stripes along the back of the leg below
the elbow, which are continuous with those on the inner side of
this limb. The inner sides of both fore and hind limbs are fully
striped to the feet. The pale areas over the eyes, on the cheeks,
chest, belly, and inside of the limbs are only dirty white and not
sharply defined from the yellow-brown hue of the rest of the body.
The yellow-brown hue of the muzzle extends over the whisker-
area down to the black patch round the corner of the mouth
1908.] THE SUMATRAN TIGER. 891
and separates the white patch on the front of the upper lip from
the white of the cheek.
Except for the multiplication and duplication of the stripes,
this specimen seems to agree with other Sumatran Tigers that
have beendescribed. In his Monograph of the Felide, for ¢ example,
Elliot remarks that Sumatran Tigers are smaller than Indian
examples and do not exhibit any white about the face and throat,
those parts being buff, while the general colour is dark red, but
with the stripes distributed in the typical style.
Text-fig. 174,
Sumatran Tiger (from a specimen now living in the Society’s Gardens).
Our Sumatran Tiger is also small. His age is uncertain,
however, and he may be no more than about three years old. His
weight is probably only about half that of our large Indian Tiger.
He stands about 29 inches at the shoulders.
The Sumatran Tiger was originally named Lelis tigris nigra by
Lesson (Nouv. Tabl. R. Anim., Mamm. p. 50, 1842.) But since no
description was subjoined, nigra must be regarded as a nomen
nudum. Fitzinger subsequently described it as Tigris sondaica
(SB. kais. Akad. Wien, lvili. pt. i. p. 454, 1868), and this name
has been universally and correctly adopted,
Our Sumatran specimen resembles in the nature of its stripes
the Persian Tiger described and figured by Dr. Heck (Lebende
892 MR. R. I. POCOCK ON [ Dee. 15,
Bilder ete. p. 157), but may be at once distinguished by the
indistinctness and small extent of the white areas of the head and
body, by the absence of the fringe of hair on the belly, and the
shorter hair of the cheeks and throat. The Persian race has
been named Ff’. tigris virgata by Matschie, in allusion to the
completeness of the pattern of stripes.
Of Indian Tigers the Society possesses at the present time three
examples: one large male from Mysore, presented by A. Forbes,
Esq., C.S.1., and two females from Nepal, presented by H.R.H. the
Prince of Wales. The latter are remarkable for the reduction
both in number and length of their stripes, of which scarcely any
Text-fig. 175.
Nepalese Tiger (from a specimen now living in the Society’s Gardens).
show a sign of looping. The greater part of the shoulder, the
outside of the foreleg, and a large portion of the costal area of the
thorax are without stripes; while on the inner side of the fore leg
the only stripe that persists is the brachial stripe, a constant
feature in many species of Felis. Onthe hinder part of the body
and on the hind-quarters the stripes show a strong tendency to
abbreviation, in addition to being comparatively thin and widely
separated. From their general appearance I am convinced that
these two specimens came from the same litter, a conclusion which
lessens the systematic value one might be inclined to attach to
1908. ] THE SUMATRAN TIGER. 893
the features they have in common. Be it noted, moreover, that
another Tigress which came at the same time from the same
country was as fully striped as our other Indian Tiger, though much
less so than the individual from Sumatra; and that a thick-
coated Siberian specimen in the British Museum is as poorly
striped as the two Nepal specimens here described. These Nepal
Tigers do not develop a thick winter coat, although they are kept in
the open all through the cold weather. Indian Tigers are regarded
systematically as typical representatives of Felis tigris.
Of Mantchurian Tigers the Society has a fine pair presented by
the Duke of Bedford. From their facial similarity I should say
that they undoubtedly came from the same litter. Beyond the
fact that they were shipped from Vladivostock, their exact locality
is unknown. They seem to be typical members of the Man-
tchurian race, and differ from our Indian Tigers in having a con-
siderably greater extent of white and a correspondingly lesser
extent of yellow on the body, head, and limbs. They also develop
a thick coat in the winter. The male stands about 38 inches at
the shoulder.
The race to which these Tigers belong was named /. tigris
mongolica by Lesson (Nouy. Tabl. R. Anim., Mamm. p. 50, 1842);
but since the name was unaccompanied by a diagnosis, it cannot
stand, although it has been adopted by Matschie, Trouessart, and
Lydekker, who at the same time reject the name nigra given by
Lesson tothe Sumatran race. I adopt, therefore, the name longi-
pilis proposed by Fitzinger (SB. kais. Akad. Wien, lviii. pt. i.
p. 455, 1868).
The four desertbed races of Tigers may be briefly characterized
and contrasted as follows :—
a. Pale areas of the head, body, and limbs dirty to buff white and
small in extent; size small or medium ............ sondaica.
a’, Pale areas of the head, body, and limbs clean white, sharply
defined and greater in extent.
6. A copious mane on the cheeks and throat and along the
belilvgan sive mae drum ys. Matar foee dees eset virgata.
6’. Mane on cheeks shorter; practically none on throat and
belly.
c. White on belly, face, and inside of legs considerably more
extensive ; winter coat thick and woolly ... longipilis.
c'. White on belly, face, and inside of legs much less extensive;
winter coat short and not markedly longer and thicker
than that of the summer S220). 220s ees tigris.
894 MR. F. E, BEDDARD ON THE ANATOMY | Dee. 15,
The following papers were read :—
1. Some Notes on the Muscular and Visceral Anatomy of
the Batrachian Genus /emisus, with Notes on the
Lymph-Hearts of this and other Genera. By Franx E.
Bepparp, M.A., F.R.S., F.Z.8.
[ Received October 23, 1908. |
(Text-figures 176-190.)
The existing knowledge of this genus of Frogs is limited, so far
as I am aware, to the external characters*, the osteology, and
certain points in theanatomy of the tadpole t. I therefore take the
opportunity afforded by the death of the only example of a species
ot Hemisus (H. guttatwm) ever possessed by the Society to lay
before the Meeting a few notes upon the structure of the “ soft
parts” of the adult, as a further contribution to the anatomy
of the Engystomatide.
As I have had only one individual for examination, my account
of the anatomy of Hemisuws cannot aim at being comprehensive.
I have, however, been able to get together a considerable number
of facts upon the anatomy of many organs and systems of organs
in this Frog, which I treat of in the following order :—
Dorsal Musculature, p. 894.
Ventral Musculature, p. 898.
Hyoid and its Musculature, p. 907.
Musculature of the Thigh, p. 912. .
Abdominal Viscera, p. 913.
Thymus Gland, p. 915.
Posterior Lymph-Hearts and Sacs, p. 916.
Posterior Lymph-Hearts of Yenopus, p. 924.
Lymph-Hearts of Rana guppyi, p. 930.
Résumé of Characters of Hemisus, p. 932.
Résumé of principal new Facts, p. 933.
§ Muscles of the Dorsal Surface.
Contrary to what is found in Breviceps and Rhinoderma, the
depressor mandibule of Hemisus is quite large and well developed.
It arises in the ordinary way from the fascia dorsalis over-
lying the latissimus dorsi and crosses the scapula on its way to
its insertion on to the lower jaw. Of this muscle the outer
margin is thicker than the rest, though there is no abrupt break —
dividing the muscle into two sections.
* Boulenger’s Cat. Batr. Sal. B. M. 1882, and literature therein cited ; Gadow,
in Cambridge Natural History. vol. viii. 1901, relating to Reptiles and Amphibians.
+ Bles, “ Notes on Anuran Development, &c.,” Budgett Mem. Vol., Cambridge.
£ “On the Anatomy of Breviceps,” P.Z.S. 1908, p.11. “On the Anatomy of
Rhinoderma,”’ P. Z. 8. 1908, p. 678. ;
1908.] OF THE BATRACHIAN GENUS HEMISUS. 895
The latissimus dorsi is a large muscle, the origin of which
commences some way behind the scapula and extends forward to
a point about on a level with its posterior border. It arises from
the middle line of the back and underlies the fascia dorsalis which
is closely adherent to it. It should be mentioned in considering
this muscle that the humerus is not free from the body. It is
closely connected with the fascia covering the body and a strongish
band connects the fascia dorsalis with the very elbow. . This state
of affairs must necessarily, one would suppose, have influenced
the adjacent musculature. In any case, the latissimus dorsi
blends early with the infraspinatus, and indeed it is dificult to
distinguish between the two muscles anywhere. The conjoined
muscles narrow rapidly to form a thick muscle a little way from
the insertion on to the humerus.
The cucullaris is a very massive muscle and is attached up to
the very tip of the suprascapula, along its anterior border.
When the latissimus dorsi is cut and reflected I can find no
muscle comparable exactly to the transversely running rhom-
boideus (or retrahens scapule) of Rhinoderma. The position of
that muscle is occupied by fascia binding the suprascapula to the
middle line of the back, in which no muscular fibres can be de-
tected on dissection. The cutting and reflection of the latissimus
dorsi, and the fact that the suprascapula thus exposed is a narrow
plate of cartilage with a concave posterior boundary-line, brings
unto view certain muscles connecting the transverse processes of
the third and fourth vertebre with the scapula and suprascapula,
which have received various names in Lana.
Inasmuch as these muscles have not been described in the
large female Rana guppyi, where they are naturally peculiarly
clear, and in which Frog they appear to differ slightly from the
corresponding set of muscles in Rana esculenta, it will not be
useless to describe these muscles before proceeding to deal with
those of Hemisus.
In Rana guppyi the muscles in question, which obviously
vesemble, as has been pointed out, the serratus group of muscles of
higher animals, can be divided into two groups :—those which are
inserted on to the under surface of the suprascapula and those which
are inserted on to the under surface of the scapula. The direction
of the two sets of muscles is totally different, and their course
indeed lies nearly at right angles. The broad cartilaginous edges
of the suprascapula nearly completely cover this system of
muscles. The group which are inserted on to the suprascapula
consist of four muscles, of which one, the rhomboideus, has been
-already described by myself in this species *.
The second is a large flat muscle arising from the free end of
the transverse process of the fourth vertebra, which I take to be
the retrahens scapule of Ecker (with which therefore I was wrong
in identifying the rhomboideus in my description of Pipa quoted
* Cf. memoir on Pipa, P. Z.S. 1895, p. 836.
Proc. Zoou. Soc.—1908, No. LVII. 57
896 MR, F. E, BEDDARD ON THE ANATOMY [ Dec. 15,
below) and which is perfectly distinct from the rhomboideus of
Rana guppy to which I have just referred. The muscle is inserted
on to a considerable area of the suprascapula and is in contact in
front with the insertion of the rhomboideus, which is in its turn
in contact with the msertion of the cucullaris. At its origin the
muscle is connected with the tendinous intersection of the
longissimus dorsi as well as with the transverse process of the
fourth vertebra. Between this muscle and the next to be men-
tioned lies the anterior lymph heart which is described on another
page *. This next muscle arises from the end of the transverse
process of the third vertebra and is distinctly composed of two
parts. Hach of these is a flat sheet of muscular fibres.
The two sheets are perfectly distinct at their origin. For the
outer of the two does not extend so far along the surface of the
cartilage posteriorly as does the inner muscle. At their insertion,
however, close to that of the retrahens scapule, there is no dis-
tinction between the several layers of this muscle, which must
therefore be regarded as simply double-headed. It differs there-
fore from the transverso-scapularis tertius or serratus (Kicker) of
Rana esculenta, with which I believe it to be homologous. The
insertion of this muscle is in contact with that of the retrahens
scapule. The fourth muscle is not a flat muscle like the last:
two, but is narrow and more or less oval in transverse section.
It arises independently of the last muscle from the anterior
edge of the free end of the transverse process of the third
vertebra, and is inserted quite away from the serratus near the
anterior border of the suprascapula outside of the insertion of
the levator anguli scapule. This muscle is not mentioned by
Kcker, unless, indeed, it is this which is the transverso-scapularis
tertius.
The remaining muscle of the “ serratus ” series is obviously the
homologue of the transverso-scapularis major (Ecker) of Rana
esculenta, and is the only muscle of the series which is inserted
upon the scapula. As in &. esculenta, it arises by two heads,
one from each transverse process of vert. 3 & 4. That arising
trom the transverse process of the fourth vertebra is very much
the larger and both heads are entirely fleshy. The insertion of
this muscle on to the scapula lies between the insertions of the
sterno-cleidomastoid and the protrahens scapule, which are the
two head muscles of the scapula corresponding to the single head
muscle of the suprascapula referred to above *.
When in Hemisus the latissimus dorsi has been cut through
and reflected, two muscles belonging to the serratus series aie
exposed throughout their entire length and a third very nearly
so. The two which are fully exposed belong to the suprascapula,
and the third is very obviously the equivalent of the transverso-
scapularis major which is inserted on to the scapula.
* Vide p. 930.
+ The sternocleidomastoideus of. R. guppyi veally consists of two parts, a much
larger part and a smaller which is inserted separately by a longish tendon.
1908. ] OF THE BATRACHIAN GENUS HEMISUS. 897
The two muscles belonging to the suprascapular series of the
serratus group arise respectively from the transverse processes of
the third and fourth vertebre. That arising from the fourth
vertebra, and which therefore represents the retrahens scapule of
Rana, is much the more slender of the two. It is a flat strap-
Shaped muscle of much the same diameter throughout. It is
inserted on to the end of the suprascapula by a flat tendon. The
shorter and wider muscle arises from the transverse process of
the third vertebra and is inserted on to the suprascapula along a
wider line of insertion than that of the last-described muscle,
but in contact with it at its extremity. It corresponds, as I
imagine, to that double-layered muscle in Rana guppyt which I
have identified provisionally with the transverso-scapularis tertius
of Rana esculenta. The chief reason which leads me to this
inference is that the anterior lymph-heart lies between it and the
retrahens scapule just described. Moreover, the muscle is dis-
tinctly composed of two layers, or rather it may be better to
speak of the lower layer as a distinct muscle, since it is more oval
in section than the superjacent layer. In this case the deeper
layer may be really the equivalent of the third “ serratus” muscle
described above in Rana guppyi. The two muscles (or three)
which have been just described run in a direction which is not
far from parallel to the longitudinal axis of the body, being
directed obliquely inwards from behind forwards to that axis, and
they constitute those muscles belonging to the serratus system
which are inserted upon the suprascapula. There is also besides
the cucullaris, which has been already referred to, another head
muscle, the levator anguli scapule, which is also attached to and
beneath the suprascapula.
There now remain certain muscles of the serratus complex which
are inserted upon the scapula. Of these there is first of all the
obvious homologue of the transverso-scapularis major of Rana.
This consists, as in Rana, of two heads arising respectively from
the transverse processes of the third and fourth vertebre. The two
heads are entirely fleshy and more equal in size than in Rana; they
combine to form a single muscle which is inserted low down on the
scapula. The direction of this muscle is quite at right angles to
that of the suprascapular series of the serratus complex. A second
large muscle lies in front of that which has just been described,
and its fibres run about parallel with those of the transverso-
scapularis major, and are inserted on to the junction of the scapula
and the suprascapula if the junction is fixed by the change of
direction of the bony scapula from the chiefly osseous, partly cav-
tilaginous supra-scapula; indeed, perhaps the bulk of the fibres
are really attached to the suprascapula. This muscle arises well in
front of the third vertebra, but its origin is not covered by the
suprascapula. I suppose that it may be compared with the trans-
verso-scapularis minor (Ecker) of Rana esculenta; but the origin
is different and the muscle actually and relatively much larger.
When the abdominal viscera are removed or pushed aside, the
57*
898 MR. F. E. BEDDARD ON THE ANATOMY [ Dec. 15,
internal surface of the ilia and their muscles are exposed, as I
have recently figured * in the genera Megalophrys, Rana, Pelobates,
and Ceratophrys. The conditions observable in Hemisus when
a dissection of this kind is made are more like those of Ceratophrys
than those of any of the other genera to which I have just referred.
The ilium is exposed for the greater part of its length and devoid
of muscular covering, for the ilio-coccygeal origin does not extend
at all over the ventral surface of the bone.
The ilio-lumbaris avises towards the anterior end of the ilium,
exactly in the way in which I have figured it in Ceratophrys.
It is, however, a rather more solid muscle and passes up to the
origin of the cesophageal muscle + without a break except for
tendinous intersections which correspond to the transverse
processes of the successive vertebrae. Moreover, it abuts closely
upon the centra of the vertebrie, at any rate anteriorly. There
is no long lateral slip of this muscle as in the Pelobatide f.
S$ Ventral Musculature.
The two pectorales abdominis differ from those of many Frogs
in that they meet in the middle line ventrally. The rectus
abdominis absolutely ceases to be visible with their origin, and is,
in facet, anteriorly to this line covered by them, a peculiarly
strong inscriptio tendinea forming the boundary line between
the two muscles. Another peculiarity of this inscriptio tendinea
besides its strength and toughness, which is doubtless in relation to
the importance of the pectoralis attached to it, is the fact that this
tendinous seam is firmly attached to the skin. So firm and so
direct (i.e. not through a special septum such as those which
divide the other subcutaneous lymph-spaces) is this connection
that some fibres of the muscles concerned have the appearance
of arising from the skin. The two pectorales abdominis are
not only continuous at their origin from this tendinous seam and
septum, their fibres are nearly in contact for some little space
in front of this; for there is a prolongation forwards of the seam
at right angles to the rest, from which the innermost fibres of each
pectoralis abdominis arise. This is not, however, continued far
towards the sternal region. A triangular or, indeed, almost
V-shaped space is left between these two pectorales abdominis
and the pars sternalis anteriorly, as is shown in the figure
(text-fig. 176), which is uncovered by any muscular layer and
where the posterior region of the pectoralis sternalis is exposed.
In the middle ventral line of the body the septum between
the pectoralis abdominis and the rectus abdominis is pretty
well at right angles to the longitudinal axis of the body.
Laterally the line of origin of the pectoralis curves more and
more anteriorly, so that at the sides of the body the origin of the
* P.Z.S. 1907, p. 332, text-fig. 93, p. 333, text-fig. 94, &e.
+ For which see p. 915.
ft P.Z.S. 1907, p. 905.
HSS) <7 OF THE BATRACHIAN GENUS HEMISUS. 899
pectoralis abdominis is not far from the armpit. Furthermore,
in this region the fibres of the muscle very distinctly arise from
Text-fig. 176. - a
rn gia
Ventral musculature of Hemisus guttatum; the skin has been largely removed,
but no muscles have been cut and reflected.
f. “Thymus gland.” m. Submentalis muscle. R. Rectus abdominis mascle; the
letter points to the first inseriptio tendinea. p.abd. Pectoralis abdominis
S.m. & Sm.’ Two portions of submaxillaris muscle.
900 MR. F. E, BEDDARD ON THE ANATOMY [ Dec. 15,
the skin itself and entirely from the skin, not merely by a few
fibres here and there as may be the case towards the middle line
of the body.
The pectoralis cutaneus is completely absent. The septum which
divides the thoracic from the ventral lymph-sac runs across the
pectoralis sternalis at about its middle; but I observed no trace of
the muscles in or about this septum which are so obvious in Rana.
These muscles would appear to be not unfrequently unrepresented
among the Batrachians. In the present species they can hardly
be represented by the cutaneous fibres of the outer part of each
pectoralis abdominis described above.
The sterno-radialis, as in Rana, arises from the omosternum,
and its origin is limited to the omosternum. Instead of being
overlapped by, it overlaps the anterior part of the pectoralis
sternalis. It is a broadish strap-shaped muscle, but not relatively
so large as in Rana.
The pectoralis sternalis is divided, as in Rana, into a portio
anterior and a portio posterior. The first of these two halves of
the muscle is not visible superficially for the whole of its extent.
Only a small part appears before any dissection is made, as may
be seen in the text-figure accompanying this description (text-
fig. 176, p. 899). The origin of most of it underlies the origin of
the sterno-radialis, and some underlies the origin of the pectoralis
posterior. The latter is a very large and deep fleshy muscle,
much larger than the portio anterior. It is triangular in form
and overlaps, as already stated, a part of the portio anterior.
There is a tendinous seam running along it for about half of its
course before reaching the humerus, on to which a part of the
pectoralis abdominis is inserted. The muscles of the two sides of
the body meet at a tendinous seam from which they chiefly
arise, but they take origin also from the very obliquely set
coracoids.
Coraco-humeralis and pectoralis minor.—There are in Hemisus
three strong and fleshy muscles which correspond, as I presume, to
the two muscles thus named in Rana (2. guppyi*) to their origin
from the coracoid; they lie one behind the other. The most
posterior of the series (text-fig. 177, p.) is quite visible superficially
before any dissection of the ventral musculature is made. It is in
contact with, but obviously separate from, the pectoralis sternalis
posterior. In section the muscle is at first crescent-shaped, since it
partly underlies, as well as being parallel to, the part of the
pectoralis already referred to. Further on the muscle becomes
flatter, and is inserted upon the humerus by a flat strap-shaped
short tendon immediately ventral of the insertion of the pectoralis
abdominis. It may be that this muscle is really referable to the
pectoralis sternalis rather than to the coraco-humeralis ; but in
any case it is perfectly distinct from the pectoralis from origin to
insertion.
* See Beddard, “ Anatomy of Pipa,” P. Z.S. 1895, p. 837.
1908. ] OF THE BATRACHIAN GENUS HEMISUS. 901
Immediately underlying the last-deseribed muscle (when the
animal is viewed in the ordinary position of dissection) is a much
broader muscle which I take to represent that muscle which I
Text-fig. 177.
Ventral musculature of Hemisus guttatwm, with the skin removed and some of
the superficial muscles cut and partly removed.
f. Fat-mass, lying within a lymph-sac covered by pectoralis abdominis and floored
by a delicate layer of muscle (r) partly belonging to the rectus abdominis and
partly to the obliquus internus. d. Fibrous wall of femoral lymph-sac cut
irregularly near to its origin from the reflected border of the rectus abdominis.
p. Muscle (cut across) which is perhaps part of the coraco-humeralis.
‘s. Coraco-humeralis. s'. Pectoralis minor.
902 MR. F. E. BEDDARD ON THE ANATOMY Dee. 15,
7)
have just referred to in Rana guppyi and Pipa* as the pecioralis
minor. It arises (text-fig. 177, s.’, p. 901) from the more internal
part of the coracoid not only below the pectoralis sternalis, but
from the opposite (7. e. the dorsal) side of the coracoid bone. It
arises by several partly separate strands, is fan-shaped, and rapidly
narrows to a cylindrical muscle, which is inserted on to the opposite
side of the humerus to the pectoralis, and is doubtless a muscle of
antagonistic action.
In front of this hes the third muscle of the series which [am now
considering (text-fig. 177, s.). It isa short rather broad muscle
arising from the humeral half of the coracoid. It is attached to
the humerus just below the insertion of the first-described of the
three muscles belonging to the present series. I think that there
can be little doubt that it really corresponds to the coraco-humeralis
of Rana.
The obliquus 1s quite extensive on the dorsal surface of the
body, the fibres having precisely the same direction as those of
the obliquus externus in Lana, i.e. obliquely from before back-
wards and outwards. Whenthe animal is pinned in a dissecting-
dish with the ventral side downwards, the whole of the flanks
are seen to be occupied by this muscle up to the large vacuity
posteriorly occupied by the saccus iliacus. Dorsally the fibres
originate laterally of the ilia and expanded sacral transverse
processes from the tough aponeurosis which covers the dorsal
musculature loosely, and is attached by a downward band to the
ilia and sacral transverse processes before it becomes confluent
with the obliquus externus. Posteriorly the muscle appears to
end in a slightly thickened concave margin at the saccus iliacus.
This ending, however, is only apparent; there is a folding over
exactly such as will be described in the case of the rectus in the
pubic and femoral region, but less in extent, and caused in exactly
the same way, or, at any rate, correlated with an anatomical fact
of the same nature. For in the muscle now being described there
is a firm insertion along the bend of the muscle-layer of the
dorsal wall of the iliac lymph-sac. The fold in this dorsal region
of the obliquus is by no means so deep, however, as is that of the
rectus abdominis ventrally. It is plain all the same from following
them out that the fold in question is perfectly continuous from
the ventral region to the dorsal, and it follows therefore that
there is no strict demarcation between the rectus abdominis and
the obliquus externus in this Frog. That is to say, there is no
line of demarcation between the deeper flap of the rectus and the
obliquus externus. The superficial flap of the rectus, as already
said, ends upon the skin. The two parts of the muscle are thus
nearly at right angles here, and the posterior sheet runs
almost dorso-ventrally, forming the anterior boundary of the.
iliac lymph-sac and exposed by cutting open one of the septa
ns A redissection of the muscles in question in Rana guppyi shows the presence of
the “pectoralis minor” in that Frog, as I have asserted.
1908. | OF THE BATRACHIAN GENUS HEMISUS. 903:
of the lymph-sac. When the dorsal part of the muscle now
under description is cut through by an. incision running
parallel with the long axis of the body, it can be plainly seen
to be a single though fairly thick layer of muscle. There is
no layer underneath it. There is, in fact, in this region, that is
along the entire back, but one obliquus muscle. There is,
however, a strong fascia covering the muscle dorsally. This latter
may really represent the obliquus externus as well as a portion of
the rectus abdominis already described as being inserted upon the:
ékin. For the muscle which I am now describing has, in spite of
the direction of its fibres, more in common with an obliquus.
internus. When the fibres are traced ventralwards they are seen
to end ina digitate fashion on the sides of the body in a delicate
membrane. Anteriorly the muscle extends to within a very
short distance of thé scapula, but not in the least touching it.
It is bounded, in fact, anteriorly by the origin of the depressor
mandibulx. It is interesting to notice how thoroughly this Frog
Hemisus differs from its ally Breviceps in the oblique muscles.
In the latter they are both well developed and fleshy throughout.
In Hemisus the muscles are largely defective as muscular tissue ;
and on the ventral side there is only the delicate membranous
continuation of the obliquus.
This sheet of the body-wall is partly muscular and partly forms
a delicate membrane of connective tissue, in which no muscular
but only wavy connective-tissue fibres can be detected by the
microscope. As to the latter tract, I shall presently mention it
in describing the rectus abdominis muscle. When the rectus is
cut across, reflected in the middle region of the body, a delicate
membrane comes into view which underlies the rectus and is the
membranous part of the obliquus internus veferred to. It is even
suggestive of an omentum, such is its freedom from the rectus.
It is not, however, attached to the viscera which it covers, save
here and there by an emergent blood-vessel. It extends all over
the body-cavity right back to the neighbourhood of the bladder—
in fact, to the posterior boundary of the abdominal cavity. It is
quite thin and transparent. It appears to me that this membrane
must be referred to the obliquus internus, since it is absolutely
continuous with a sheet of muscle laterally which can be nothing
else than the obliquus internus, as well as the muscular sheet
anteriorly which bounds the thoraco-abdominal cavity.
The rectus abdominis in this Frog is much more like that of
Breviceps than of Rhincderma. For it has only one inscriptio
tendinea between its origin at the pubes and the inscriptio ten-
dinea to which the pectoralis abdominalis is attached. The fibres
too are arranged in a fan-shaped way like those of Breviceps, and do-
not run only in an anterior direction parallel with the long axis
of the body as in Rhinoderma and many Frogs. In the middle
line of the body the fibres run postero-anteriorly ; laterally they
are quite oblique in direction. Furthermore, it will be noticed
‘904 MR. F. E. BEDDARD ON THE ANATOMY [ Dec. 15,
from the drawings (text-figs. 176 & 177, pp. 899, 901) that, as in
Breviceps *, the boundary-line between the thigh ventrally and the
trunk ventrally is entirely occupied by these muscles, a separate
obliquus externus not being visible on this view of the animal.
Whereas in Rana, when the skin is reflected from the abdomen and
thigh, the obliquus externus as well as the rectus are seen to form
the boundary-line between limb and trunk. There is another
important difference which this muscle shows and in which is re-
sembles Breviceps. The rectus abdominis overlaps a considerable
portion of the thigh, to the extent indeed of 6 mm. or so. Under
the free edge of the muscle laterally a seeker can be pushed.
There is, however, a plain distinction laterally between the rectus
abdominis and the obliquus muscle (for the moment I leave it
undecided whether it is to be regarded as externus or internus),
which is not merely the lateral and dorsal extension of the rectus.
It will be noticed that the one inscriptio tendinea (see text-fig.
177) which exists behind the origin of the pectoralis abdominis,
and along the course therefore of the rectus abdominis, does not
reach the edge of the muscle which overlaps the thigh musculature
and towards which it tends. In this region then it is impossible
to discriminate between rectus and obliquus, on the assumption,
that is, that we have here reached the border-line of the two.
I am disposed, however, to think that this lateral extension of the
rectus is wholly rectus; for a careful dissection shows that it ends
by being inserted upon the skin and its fibres are not continuous
with those of what is obviously the obliquus muscle described
above as originating from the dorsal aponeurosis.
When the pectoralis abdominis is cut through and reflected
the anterior portion of the rectus abdominis is brought into view.
This les at a much lower plane than the posterior region of the
muscle. For there is a deep cavity between it and the covering
pectoralis abdominis. This cavity is not merely a lymph-space.
It contains an elongated body which I describe later in connection
with the thymus. This cavity then is floored (examined in the
ordinary position of these muscles when dissected from the ventral
surface) by a delicate layer of muscles (text-fig. 177, 7.) which is by
far thinner than the rectus abdominis of which it is the forward —
continuation from the anterior inscriptio tendinea. The muscular
fibres, however, do not extend over the whole of the cavity thus
exposed. ‘Towards the middle line the muscular fibres form an area
which is not only thicker in its muscular tissue than more laterally,
but definitely arises from the inscriptio tendinea. Laterally there
is no such origin from the inseriptio tendinea where the rectus
abdominis and the pectoralis abdominis meet, and this sheet has
been described as a part of the obliquus internus.
The lateral portion of the rectws abdominis, under which, when
it covers the thigh, a probe can be passed, as already mentioned,
demands a more detailed consideration. It is to be noted, in the
* P, Z. 8. 1908, p. 16, text-fig. 3. + See p. 915.
1908. ] OF THE BATRACHIAN GENUS HEMISUS. 905
first place, that there is here not simply the matter of a muscle
extending loosely over the proximal region of the thigh during
its relaxed condition. The edge of the rectus which lies upon the
thigh is bound down to the skin of the leg by a septum of con-
nective tissue (text-fig.177, 2.) which forms the wall of a lymph-sac
belonging to the system of femorallymph-sacs. When this septum
-and the flap of muscle is cut through transversely by a pair of
scissors, the section is seen to be V-shaped, the edge of the V being,
-of course, the line along which the wall of the lymph-space already
referred to is inscribed. The ventralmost flap of the V is natu-
rally the muscle exposed on a dissection from the ventral surface,
-and is what has been described as the rectus abdominis. The more
dorsal flap is folded under this up to the very line (the mid-
ventral line of the body) where the muscles of the two sides of
‘the body meet, and is inserted on to the edge of the pubis.
Although here the fibres of the superficial flap of muscle are
‘accurately antero-posterior in direction, while those of the sub-
jacent flap are exactly at right angles to them—running, that is to
say, in a lateral direction—the directions of the fibres become
-coincident at the apex of the V which the two flaps of muscle form.
Tt should now be mentioned that the deep-lying flap of muscle
-of which the fibres are consistently lateral in direction throughout
is not a continuation of the obliquus internus. The membranous
‘sheet which represents the latter muscle in this region of the
body underlies and is free from the layers of muscle which have
just been described. The lower flap of the muscle of one side of
the body is quite distinct from that of the other, since they are
divided by the line of the pubis from each edge of which they
arise. The superficial flaps are, however, quite united in the
middle line, and posteriorly, at any rate, no linea alba is to
be seen.
The submentalis (text-figs. 176, 178, m., pp. 899, 906) has
‘a shape which is evidently influenced by the shape of the
jaw and is also a considerably larger muscle than in Rana.
The anterior extremity of the lower jaw, instead of forming a
uniform curve as in Rana, has a perfectly straight or square
xegion anteriorly, which is shown in the accompanying text-figure
(text-fig. 178). The breadth of the jaw here is fully 5 mm., and
‘the length of the submentalis is therefore only a little less and it
has not in so marked a degree as in other Frogs a lenticular shape.
It has the form of a cylinder tapering to both extremities. Its
‘fibres can be seen to run straight across from one side of the jaw
to the other in the middle region of the muscle. At both ends
they curve upwards and are inserted into the angle of each man-
dibular ramus where the straight anterior portion of each, which
is at right angles to the longitudinal axis of the body, passes into
the side of each ramus. I have described in 2hinoderma darwini*
a pair of triangular muscles lying behind the submentalis which
I compared to the genioglossus. It might perhaps—though at
* P, Z. 8. 1908, p. 686, text-fig. 1464.
906 MR. F, E. BEDDARD ON THE ANATOMY [ Dec. 15,
present any suggestion as to the homology of the muscles lacks a
firm base upon comparative anatomy—be more reasonable to
regard the muscles in question as a part of the submentalis. In
any case, I do not find the least trace of this muscle in Henvisus.
Text-fig. 178.
gh.
SEN.
Ventral musculature of neck-region in Hemisus guttatum.
m. Submentalis muscle. g.h. Geniohyoid ; the white lines dividing the two lateral
parts of the muscles from the median practically unpaired portion represent
the hypoglossal nerve. Sé.i. Sternohyoid muscles ; the three separate muscles
are shown. ‘'o the left of these are seen the petrohyoidei.
Submaxillaris— Although Hemisus differs from Rhinoderma in
the matter just referred to, the two agree in the specialization of
the submaxillaris proper (not including the subhyoideus, which
was formerly regarded as being a part of this muscle) into two
regions. The conditions observable in Hemisus are shown in the
figure referred to above (text-fig. 176, p. 899). The main mass
of the muscle, which is all that exists in Rana and many other
Batrachians, is indistinguishable posteriorly from the subhyoideus.
Fach muscle is divided from its fellow in the middle line of the
1908. ] OF THE BATRACHIAN GENUS. HEMISUS. 907
throat by a considerable tendinous interval anteriorly. This
non-muscular interval diminishes in breadth posteriorly until it
practically disappears in the region of the subhyoideus. In
addition to this the swhmawillaris consists of an anterior layer of
fibres on each side which are comparable to an almost similarly
placed layer of fibres in Rhinoderma. A thin layer of fibres runs
on each side from the fascia covering the submentalis to the ramus
of the jaw in an oblique direction, and overlies almost at right
angles the section of the submaxillaris which is contiguous. This is
clearly shown in text-figure 176, s.m.',and needs no more elaborate
description.
The subhyoideus is of about the same proportions as in Rana,
and passes behind the ramus of the lower jaw on its way to the
cornua of the hyoid. That it is attached to the cornua of the
hyoid and not to the wall of the skull is quite apparent. An
examination of text-figure 176 would seem to show an additional
muscle belonging to the series which form the floor of the mouth,
and arising on either side from the anterior extremity of the
sternum. I am unable, however, to give any further details
about this muscle than are displayed in that figure. It may of
course be merely an anterior slip of the pectoral series (including
the sterno-radialis).
§ Hyoid and its Musculature.
The hyoid cartilages of Hemisus are peculiar in several respects.
The main features of this part of the skeleton can be understood
by a reference to the accompanying text-figures (text-figs. 179, 180,
pp- 908, 909). The body of the hyoid is rather long and narrow.
The anterior hyoid processes of the body of the hyoid join the
anterlor cornua much in the way that is to be seen in the hyoid
ot Breviceps*. Furthermore, the two anterior cornua or cerato-
hyals themselves join ventrally of the median body of the hyoid
and project in the shape of a rather broad plate for some way
backwards over the latter. The hyoglossus muscle therefore passes
through an actual foramen in the hyoid, which it completely fills.
Tt is evident, however, that this hyoglossal foramen isnot absolutely
homologous with that of Xenopus 7, but is more comparable to the
nearly completeforamen seen anteriorly in the hyoid of Pelodytes <.
For the foramen in Xenopus is an actual perforation of the body of
the hyoid, whereas in Hemisus the foramen is produced beyond
the end of the body of the hyoid and by the approximation of the
origins of the anterior cornua of the hyoid. Were there a com-
plete foramen in the hyoid of Pelodytes punctatus it would be
more comparable to that of Henisus in that the anterior cornua
enter into its formation. It would not, however, be strictly
homologous; for in Pelodytes and Pelobates there are a pair of
* P. Z.S. 1908, p. 12, text-fig. 2.
+ See Ridewood, Journ. Linn. Soc., Zool. vol. xxvi. pl. 8. fig. 1, h.
t Ridewood, P. Z. 8. 1897, pl. xxxv. fig. 10.
908 MR. F, E. BEDDARD ON THE ANATOMY [ Dec. 15,.
lateral foramina as well as the median notch nearly converted into.
a foramen in these genera (and, it may be added, in Megalophrys*).
I take it that in Hemisus the single median foramen embraces.
Text-fig. 179.
A. Ventral surface of hyoid of Hemisus guttatum partly cleared of muscle (the-
geniohyoids are removed and the hyoglossus cut through twice and the middle
part removed).
c. The anterior cornu of one side. ec’. The posterior cornu in which the absence of
dotting indicates bone. h. Thin portion of anterior cornua, which meet m the
middle line to form a ventral and backwardly projecting sheet of cartilage, with
a rounded posterior margin which partly covers the hyoglossus (divided just
behind the edge of this cartilage). h.g. Posterior region of hyoglossus.
ph. Sternobyoideus posterior ; the anterior petrohyoideus is seen anteriorly to
be inserted on to the body ot the hyoid, where it nearly meets its fellow in the -
middle line ; above this is seen the insertion of the anterior of the sterno- .
hyoids.
B. Section through body of hyoid in a longitudinal direction.
these lateral foramina as well as the median notch, since it is-
bounded laterally, not only by the roots of the anterior cornua,
* Beddard, “On Pelobatide,” P. Z. 8S. 1907, p. 895, text-fig. 237.
1908. | OF THE BATRACHIAN GENUS HEMISUS. 909:
but also by the anterior lateral processes of the body of the hyoid..
It is perhaps possible to compare the lateral foramina in the
hyoid of Breviceps with the lateral foramina of Pelodytes and
Pelobates.
Text-fig. 180.
Hyoid of Hemisus guttatum and its musculature. Ventral aspect.
h. Anterior border of hyoid: the letter points to the plate formed by the union ot
the thinner portion of the anterior cornua which underlie the hyeglossus
muscle, which muscle passes above them through the foramen thus formed.
Sé.h. Sternohyoid muscles (3), over which the hypoglossal nerve is seen to pass.
and to supply, by one branch, the hyoglossus muscle. ‘lo the left of the figure
are seen the petrohyoidei.
The anterior cornua near to the wall of the skull are bars of
translucent cartilage of approximately equal diameter throughout.
Towards its attachment to the body of the hyoid each bar gets much
wider, as is shown in text-figure 179. The wider region of each
cornu is due to the existence there of a semilunar tract of cartilage
reinforcing the outer edge of the bar and becoming excessively
thin along its anterior convex border. This cartilage is perfectly
continuous with each cornu, but has the appearance of an
‘910 MR. F.'E. BEDDARD ON THE ANATOMY | Dec. 15,
extrinsic addition to it, because of its lack of translucency. This
art of the hyoid cartilages is, in fact, whiter and more opaque
than the bluish translucent cornu. This is not so evident where
it is so very thin (7. ¢. at and near to its free edge) as it is where
the two tracts of cartilage fuse to form the hood which has
already been spoken of.
The body of the hyoid is, as already stated, long and rather
narrow ; it is also slightly oval in contour, is very thick, opaquely
white coloured, and has a swollen appearance with a convex
surface. It is obviously very thick without further proof by section
with a scalpel (text-fig. 179, B., p. 908). It therefore contrasts
greatly with this cartilage in, at any rate, many other Frogs, where
it is thin and flat and even translucent. When the body of the
hyoid of Hemisus is divided up by a longitudinal incision it is seen
to present a rather complicated structure which accounts for its
external appearance when uninjured. Anteriorly the cartilage is
not particularly thick; it underlies and is closely adpressed to
but is not continuous with a plate of bone which immediately
underlies the wall of the pharynx. This is not, 1t will be observed,
precisely the same thing as the ossifications which sometimes
occur in the body of the hyoid among Batrachians *, nor obviously
can it be compared ‘to the splint of bone found underlying the
body of the hyoid in Pelodytes and figured by Ridewood 7. We
have in Hemisus a plate of bone overlying the cartilaginous body
of the hyoid, from which it is completely separate and non-
atherent.
This plate of bone in its turn is closely connected with the
ventral wall of the pharynx. The cartilaginous plate which under-
lies this bone when divided longitudinally presents a remarkable
appearance, which is also shown in text-figure 179, B. Anteriorly
the cartilaginous plate is relatively thin and flat. It is behind
the insertion upon it of the petrohyoid that the cartilage has the
convex outline and swollen appearance already commented upon.
This turgid region is seen to be formed by a division into two
plates of the cartilaginous hyoid which do not absolutely meet but
come into the closest contact possible short of fusion posteriorly.
Imbedded in thespace between the two layers of the dichotomously
divided sheet of cartilage is a lenticular mass of a softish con-
sistency and a spongy appearance. Iam unable to suggest what
this is, unless it is developing bone. Though the above descrip-
tion is incomplete in this, it is obvious from the facts which I
have been able to ascertain definitely that the hyoid complex in
Hemisus is very remarkable.
There are other Frogs than the Engystomatid in which the
hyoid apparatus is in certain respects like that of Hemisus. I
have already directed attention to the likenesses shown in the
hyoid of Breviceps. The thin layer of rather different-looking
* K.g., Rappia sp., Cyclorhamphus marmoratus, &c., Parker, Phil. Trans. 1881.
+ Loe. cit. pl. xxxv. fig. 10.
1908. ] OF THE BATRACHIAN GENUS HEMISUS. 911
cartilage fixed on to the ceratohyals and extending backwards in
Hemisus to form a ventral hood over the hyoglossal muscle is
represented in other Engystomatid Frogs though to a less degree;
for I identify this cartilage with that termed “ extra-hyal” by
the late Prof. W. K. Parker*. This anatomist has figured a
small piece of cartilage so named in Hngystoma carolinense where
it caps the forward convexity of the ceratohyal. The same
structure is depicted in the same place exactly in Zomopterna
breviceps (¢ = Rana breviceps). In another Frog, Pywicephalus
rufescens (= Rana rufescens), the extra-hyal cartilage is shown to
be much larger, extending for a long way down the ceratohyal ;
but there is nothing like the hood of Hemisus. In Callula, how-
ever, nothing of the sort is figured. But the Engystomatid
Diplopelma ornatum differs from its congener D. berdmorei and
agrees with Hngystoma in possessing this cartilaginous cap.
Phryniscus also seems to want this structure.
The Ayoglossus is a very thick muscle, as is usual; it forms a
single muscle for the most part where it traverses the lower
surface of the body of the hyoid. But a distinct slip on either
side is quite distinguishable from the main body of the muscle,
from the very origin of the muscle from the posterior cornua of
the hyoid. The fibres of origin of the hyoglossus can be stripped
away from the shaft of the posterior cornu, with which bone they
have no relation except at the very tip, where they arise con-
tiguous with the insertion of the petrohyoideus posterior (see text-
fig. 178, p. 906). The hyoglossus, in fact, merely covers ventrally
the shaft of the posterior cornu ; it is not at all wrapped round it.
The petrohyoideus is shown in text-figs. 178, 179, & 180. The
most remarkable feature about this series of hyoid muscles is
the insertion of the anterior part of the muscle upon the ventral
surface near the middle line of the basihyoid. This muscle divides,
as will be seen, the insertion of the first from those of the second
and third portions of the sternohyoid (see text-fig. 179). The
last petrohyoid is attached to the tip of the thyrohyal and does
not extend beyond it on to the larynx.
The origin of the sternohyoideus I am unable to describe fully.
But the greater part at least seems to be derived from the under
surface of the conjoined coracoids. Whether any of it is formed
as a direct continuation of the rectus abdominis I do not know.
In any case the muscle is divisible from at least very near to its
origin into three distinct slips, which run forwards in close contact
and as one muscle. Whether these three separate s] ips correspond
or not to the three muscles in Breviceps, of which I have referred
two to the sternohyoid and one to a derivative of the obliquus
which I have termed in that Frog “ hyoabdominal” +, I am
uncertain; but their insertion one after the other in both Frogs
is in favour of this comparison. Moreover, the origin of the
hyoabdominal in Breviceps, a little way behind and to the outside
* Phil. Trans. 1881.
+ P. Z.S. 1908, p. 12, text-fig. 2, Hy.abd., and p. 23, text-fig. 5, hy.abd.
Proc. Zoou. Soc.—1908, No. LVIII. 58
912 MR. F. E. BEDDARD ON THE ANATOMY [ Dec. 15,
of the sternum, may be covered over in Hemisus by the greatly
modified coracoids, and thus cut off from its relations with the
obliquus. The insertions of the sternohyoid are shown in text-
fig. 179, p. 908. They lie one behind the other to the outside of
the hyoglossus. It is clear that in the disposition of this muscle
Hemisus is nearer to Breviceps than to Rana.
The geniohyoideus (see text-fig. 178) has the two usual insertions
posteriorly which exist in other Batrachia Salientia. They are
divided from each other for a long way up the muscle (towards
the mouth) by the main anteriorly running branch of the hypo-
glossal nerve. The lateral insertion on to the body of the hyoid
of the geniohyoideus is preceded by a gathering together of the
fibres of the muscle into a thick strap-shaped band which curves
round the insertion of the sternohyoid (the most anteriorly
inserted slip of that muscle) andis inserted below it—out of sight
in the ordinary position of dissection from below. The inner
portion of the muscle is a very delicate layer of fibres which with
its fellow of the opposite side completely covers the hyoglossus
and extends to the very end of the hyoid apparatus. In this the
muscle resembles that of Greviceps, but, as pointed out in my
description of the latter genus *, the peculiarity is not important.
The omohyoid appears to be completely absent, as in Breviceps.
I could discover no trace of that muscle in Hemisus.
§ Musculature of the Thigh.
When the skin is removed from the inside of the thigh the
following muscles are brought into view, which are named in
order from the anterior border of the thigh, viz., the vastus
internus, adductor longus, sartorius, adductor magnus, rectus
internus major, and rectus internus minor.
There is therefore nothing particularly striking so far about
these muscles, which resemble those of Breviceps and even Rana.
It is to be noted, however, that the adductor longus is hardly at
all exposed and is almost completely covered by the sartorius.
In this feature Hemisus differs from Lreviceps t and is nearer to
Rhinoderma = and Rana.
Tn describing the anatomy of Lreviceps $ I have directed special
attention to the partial origin of the rectus internus minor from
the skin. The same characteristic feature occurring also in
Rhinoderma is a bond of union between these two, in some other
respects, not very closely allied genera. This peculiar attachment
of the muscle in question to the inner surface of the skin of
the leg is also noticeable in Hemisus, but not nearly to so
marked an extent as in Lreviceps. The insertion upon, or
origin from, the skin of the thigh of a few fibres of the muscle
was, however, plain enough.
The semitendinosus is formed by the union of two fleshy heads.
* P. Z.S. 1908, p. 14. + P. Z. S. 1908, p. 25, text-fig. 6.
+ P.Z.S. 1908, p. 691. § P.Z. S, 1908, p. 26.
1908. | OF THE BATRACHIAN GENUS HEMISUS. 915
When the thigh-muscles are viewed from the outside more
muscles are visible without dissection than in the corresponding
view of Rana. Several of these are shown in text-fig. 183, p. 919.
The rectus anticus femoris is a thin and slender muscle as in Rhino-
derma *, and broadening out it ends on the surface of the vastus
internus, the head of origin of which is also visible on the present
view. On the same side of the gluteus the dio-psoas is also
visible without any further dissection. The vastus externus is
not at all of unusual size as it isin Brevicepst; and the biceps,
which comes next, is perhaps rather large. The pyriformis,
-distinctly broad, is as usual inserted dipping down between this
muscle and the last of those belonging to the dorsal series, which
‘is, of course, the semamembranosus.
§$ Abdominal Viscera.
The liver consists of three lobes and is rather different from
that of Breviceps. In the first place, it does not in the very least
-conceal the heart. Secondly, it does not only possess three lobes
as in Rana, but the right half of the liver is much smaller than
the two lobes which together constitute the left half of the gland.
“These two lobes are not completely separated. The gall-bladder
is large and spherical and almost hidden by the right lobe.
The gastro-hepatic ligament shows an interesting structural
feature which may or may not be common to other Frogs. The
ligament in question is, of course, part of the ventral mesentery of
the alimentary canal. The ligament does not extend far forward
along the stomach ; it is limited to about the last one-third of an
inch of the stomach before it bends abruptly to join the duodenum.
In this course four gastrohepatic vessels, which later join to form
two, and which are separated by a nearly if not quite anangious
-section of the membrane from that close to the duodenum which
-earries the main portal vein.
The stomach (text-fig. 181, st., p. 914) when cut open is seen to
be marked internally by thick rug, of which I counted eleven near
to the end of that organ. It is noteworthy that in this region at
-any rate five or six of these ruge are to some extent connected by
transversely running folds, a coarse network being thus formed.
‘The remaining plice of the internal surface of the stomach are
completely detached folds without any connecting folds. These
simple separate folds occupy the ventral surface of the organ ; the
others are dorsal, and the internal surface of the stomach is fairly
evenly divided between them. As in Lhinoderma, the stomach
does not end where it bends sharply to the right and narrows.
Tt is thus V-shaped and its lining ends very abruptly where the
-duodenum begins.
The duodenum (text-fig. 181, 7.) is marked by very fine and
transverse delicate folds or valvule conniventes. ‘There is no
* P, Z, S. 1908, p. 691. + P. Z. S. 1908, p. 26.
58*
914 MR. F. E. BEDDARD ON THE ANATOMY [Dee. 15,
intermediate region between these and the strong longitudinal
folds of the stomach. Moreover, the thickness of the walls of
Text-fig. 181.
Alimentary tract of Hemisus guttatum opened to show the varying characters
of the lining membrane in different regions.
c. Colorectum. 7. Small intestine. S¢. Stomach.
the Btomeien is at once diminished at the commencement of the
1908. | OF THE BATRACHIAN GENUS HEMISUS. 915
duodenum. Thus the pyloric region almost projects in a valve-
like fashion into the duodenum. The small intestine measures
when stretched out 55 mm. Throughout its whole course the
small intestine is occupied by rather closely set valvule conni-
ventes, and nowhere can I detect any distinct reticulate arrange-
ment of the folds such as I have figured in Rhinoderma. Here
and there faint indications of such are present, but nothing that
can be seriously compared to the reticulations found in 2hino-
derma or at the commencement of the duodenum as figured in
Rana esculenta. The colorectwm (text-fig. 181, ¢.), into which
the ileum opens, suddenly projecting indeed into its interior, is
marked by longitudinally running folds. There is a remarkable
resemblance to the stomach in that on one side of the gut the folds
are simple, while on the other side there are transverse folds
connecting them, and thus a network is formed.
The kidneys ave of considerable length; the right kidney
measures 19 mm., which may be compared with the total body-
length of 60 mm.
The oviducts are thick and very much coiled. The straight
piece which intervenes between the funnel and the coiled region
is very short. ‘The oviducal funnel lies behind the root of the
lung. It is unusually long, and that of the right side at any rate
measured 7mm. It is an open groove and is attached all along
to the cervical aponeurosis, the layer of the obliquus internus
which closes the abdominal] cavity anteriorly.
The lungs hang quite freely into the abdominal cavity ; they
are only attached to the liver quite at the root as in Rana guppyi*.
The diaphragm or esophageal muscle is not an extensive muscle
in Hemisus. In this it agrees with its allies. It appeared to me
also that, as in Breviceps t, the muscle is entirely inserted upon
the cesophagus.
§ The Thymus Glands.
In describing the anatomy of Breviceps I have pointed out the
large size, compared to those of Rana, of the thymus glands.
In the Batrachian with which I deal in the present communi-
cation, a pair of bodies which I take to be the thymus glands
are much larger still, proportionately as well as actually. In
Rana esculenta they ave described and figured as being about
3mm. in length. In the specimen of Hemzsws upon the dissection
of which the present paper is based, and which measured 60 mm.
(24 inches about), each thymus consisted of two discrete portions ;
the larger on each side was quite 7 mm. across at its greatest
diameter. The position and size of the organs are shown in
text-figure 176, f., p. 899. They lie on each side behind
the subhyoideus muscle and in front of the sterno-radialis and
upon the deltoid, to which especially they are attached by flat
bands of connective tissue. Each gland is flat and plate-shaped,
of rounded contour; from the middle of the ventral surface a
* P, Z.S. 1907, p. 349, text-fiz. 99, m. ~ P. Z. 8. 1908, p. 27.
916 MR, F, E. BEDDARD ON THE ANATOMY | Dec. 15,.
process arises which runs forward as far as the angle of the lower
jaw, which has almost the appearance of a duct, but which is, of
course, no such thing. The glands are white and appear to be-
chiefly composed of fat.
In addition to these structures there is on each side of the body
another “gland” of the same general appearance, but smaller,
which I refer to the same category. These are flat circular bodies.
2-5-3 mm. in diameter. These lie on either side well behind the
head and just behind the scapula, covered over by the fascia which
is continued into the depressor mandibule, and have a special!
relation to the thickened lateral edge of that muscle, as is described
elsewhere *. Their position is considerably more posterior than
the thymus glands of Rana and than the additional glands of
Hemisus already described. It is well known + that among the.
Amphibia there are in the adult various remains of bodies derived
from cells belonging to the branchial system of the tadpole, which
have been termed ‘ postbranchial bodies,’ ‘‘ Epithelk6rperchen,”
&e. I take it that in Hemisws these various structures which
have just been described are also to be referred to the same.
category. They are, however, unquestionably much larger rela-
tively as well as actually than in at least some other Amphibia
Salientia.
§ Posterior Lymph-Hearts and Associated Sacs.
The enormous size, actual as well as relative, of the posterior
lymph-hearts of Breviceps = is at present an unique fact in the.
anatomy of the Batrachia Saiientia. On grounds of aftinity the
existence of equally or nearly as large posterior lymph-hearts.
might have been expected in Rhinoderma darwini; but a careful
search convinced me that Rhinoderma$ was unlike Breviceps:
in this important and remarkable peculiarity. In Hemisus,.
however, I find an equivalent of this structure, which is very
different in its character, though retaining certain features which
lead to the inference that we have in this genus a modified
and degenerate homologue of the enormous posterior lymph-heart.
of Breviceps verrucosus.
As is the case with Breviceps, though not to so great an extent.
as in that Frog, the thigh of Hemisus is fairly enclosed within
the contour of the body. The body does not, however, extend
beyond the tip of the coccyx as it does in Breviceps.
There is thus in Hemisus, as in Breviceps, a space left dorsally
on each side of the posterior region of the vertebral column,
behind the oblique muscles, which is floored by the muscles of the-
proximal region of the thigh. This space is cut off from the:
lymph-spaces covering the rest of the thigh by a transverse and
* Cf. p. 894.
+ Maurer, “Schildriise, Thymus und Kiemenreste der Amphibien,” Morph. Jahrb.
xi. 1888, p. 298.
i Beddard, “ Anatomy of Breviceps,’ P. Z.S. 1908, p. 33.
§ Id., “On Rhinoderma,” P.Z.S. t. c. p. 678.
1908. ] OF THE BATRACHIAN GENUS HEMISUS. O17
oblique septum starting in the neighbourhood of the coccyx.
The length of this space is some 11 mm., the total length of the
body being, from snout to tip of coccyx, 58 mm. It is therefore
proportionately smaller than the corresponding space in Greviceps,
which contains the lymph-heart ; the greater size in Breviceps 1s,
however, effected by the extension of the body behind the tip of
Text-fig. 182.
Dorsal view of Hemisus guttatum, with the skin of the back partly removed.
f. Fat-body lying in saccus jliacus and covered by a membranous wall.
c. Cutaneous muscle.
the coceyx. This cavity is exposed by the removal of the skin,
which is particularly thick and glandular on the dorsal surface of
the body generally in this Frog. To the wall of the greater part
of the cavity the skin is not adherent and can be, therefore,
918. MR. F. E. BEDDARD ON THE ANATOMY [ Dee. 15,
removed withont tearing the structures which it covers.. When
this has been done (see text-fig. 182, 7, p. 917), a transparent and
toughish membrane is revealed, which completely covers the cavity
and is continuous with the septum bounding it on the side of the
leg, to which reference has been already made. Anteriorly and
laterally, however, the membrane described is adherent to the skin.
Below this membrane, and of course perfectly visible through it, is
a large fat-body quite similar in appearance to the structure which
I have termed the thymus gland and to the abdominal fat-bodies
at the anterior end of the gonads. These fat-bodies are lobulated
and extend some little way down the side of the body in the
direction of the abdomen. The transverse diameter is thus the
greatest and they measure in this direction about 15mm. In
the opposite direction the diameter is 8 mm., showing that these
bodies nearly fully occupy the suprafemoral cavity, which is now
under discussion. These fat-masses are also of considerable
thickness, and lobulated upon the lower surface as well as the
upper. They are by no means to be confused with the mass of
apparently coagulated lymph which I have described in a corre-
sponding position in Breviceps verrucosus. The general aspect
of this region of the body will be understood after a reference
to text-figure 182.
So far there is nothing exactly corresponding to the lymph-
hearts of Breviceps. There is merely a correspondence in the
existence of a space lying above the thigh and to the side of the
coceygeal region of the vertebral column in the two Frogs.
There is, moreover, the important difference that in Hemisus
this space is largely occupied by the bulky fat-body already
described. Anteriorly to the fat-body on each side, as is Shown in
the accompanying drawing (text-fig. 182, c.), a broad strap-shaped
muscle is to be seen; this is not to be confused with the obliquus
externus which lies in front of it, and, indeed, until a dissection is
made, almost, if not quite, in contact with it. Whether this muscle
is morphologically a portion of the obliquus which has become
detached from the rest of that muscle and diverted to a separate
function is another matter; it is not in any case in anatomical
continuity with it. This muscle arises from the ileum below,
passing upwards to the dorsal surface of the body and at the
same time outwards. Its oblique course ends chiefly upon the
skin, but also upon the membrane, uniting the fat-body to the
skin in this region.
When the membrane covering the fat-body, and connecting it
with the mass of bone and muscle constituting the caudal and
pelvic regions of the body, is cut and the fat-body pushed aside.
towards the side of the body (away from the middle line) it is seen
(text-fig. 183) to fill but loosely the space in which it lies and from
which it is, indeed, partly cut off by membranes which cover it
below. This membrane is largely fenestrated, so that the space
which contains the fat-body is not shut off completely from the
space lying below it. It is also attached by flat strands of membrane
1908. ] OF THE BATRACHIAN GENUS HEMISUS. 919
to another membrane lying beneath and rather in front of the
fat-body and partly covered over by the strap-shaped cutaneous
muscle already in part described.
Text-fig. 183.
Saccus iliacus of Hemisus guttatum.
The fat-body (f ) has been pushed over to the left; strands of connective tissue
uniting it with the walls of the sac are shown.
s. Anterior septum of lymph-sac.
This membrane bounds the cavity below, and when it is cut
through (as is shown in text-fig. 184, ¢, p. 920) the obliquus muscle
is exposed ; but as the membrane is quite free of the muscle it would
seem to be really the dorsal wall of a subsidiary sac lying on the
obliquus. It seems to me to be clear that this large sac, almost
filled up by the fat-body already described, and in which also
lies—as will be described presently—the lymph-heart, corresponds
to the saccus iliacus of the Frog, Rana esculenta. Furthermore,
the muscular slip which I have referred to as traversing the sac
in Hemisus near to its anterior wall appears to me to be probably
the musculus cutaneus iliacus of Rana. This sac and the muscle
920 MR. F. E. BEDDARD ON THE ANATOMY [ Dec. 15,
is figured in the edition of Kcker’s ‘Frog’ which has been
translated by Dr. Haslam*. There is a fuller account of the
lymph-saes of the Common Frog in the more recent edition of
Gaupp. I figure here (text-fig. 185) for purposes of comparison
with Hemisus the saccus iliacus in the large female of Hana
guppy. It is a much more elongated sac in proportion to its
breadth than is the case with that of 2. esculenta, as displayed
in the figure cited from Dr. Haslam. Its walls also are
Text-fig. 184.
Saccus iliacus of Hemisus guttatwn.
The fat-body is pushed to the right; the strands of connective tissue connecting it
with the walls of the sac are shown.
e. Obliquus muscle revealed by cutting open the floor of the sac.
attached by branched threads of stout connective tissue, which
present the exact appearance, as will be seen in the figure,
of the chorde tendinee of the valves of the mammalian heart.
There are two of these branched stays between the dorsal
and ventral walls of the sac. Gaupp mentions these structures.
as occurring in the lymph-saes of Rana, but does not parti-
cularize their existence in the saccus iliacus. They are not
represented in the figure, to which I have referred, of the sac
in Rana esculenta. This sac communicates by a wide orifice
* Fig. 173, p. 259.
1908. ; OF THE BATRACHIAN GENUS HEMISUS, 921
with the abdominal cavity at a point corresponding to that of
the ostium in Rana esculenta, and also at its posterior extremity
where it dips underneath the sac lodging the posterior lymph-
heart. A seeker pushed through each ostium eventually appears:
at the same place in the abdominal cavity. In Haslam’s edition
of Ecker’s ‘ Frog’ it is mentioned that the hinder portion of the
posterior lymph-heart lies in the saccus iliacus ; but in Gaupp’s
edition it is stated that the lymph-heart lies in a special sac ot
its own.
Text-fig. 185.
Saceus iliacus of Rana guppyi opened.
e. Cutaneous muscle. gl. Gluteus. O. Ostium, leading into abdominal cavity.
tr. Trabeculie connecting walls of the lymph-sac. °
The latter statement is clearly correct for Rana guppyt, where
the lymph-heart lies just in front of the pyriformis muscle in a
special sac, which is at least not in open communication with
the saccus iliacus, and which indeed overlies it. There is thus
an obvious difference between Hemisws and Rana in that the
posterior lymph-heart of the latter lies in a special sac of its own,
whereas that of Hemisus is contained in the saccus iliacus, as is
plainly shown in text-fig. 186 (p. 923), where the posterior boundary
of that sac touches the posterior wall of the heart. There is a
further difference in that the interior of this lymph-sac is largely
divided by trabeculz, in the interstices of which is lodged the
922 MR.-F, E, BEDDARD ON THE ANATOMY [ Dec. 15,
large lobate fat-body already described and figured (text-figs. 182,
183, pp. 917, 919). At the same time it will be noted that the
“* chorde tendinex ” of the lymph-sac of Rana would seem to fore-
shadow (or to be the remains of) this trabecular system. The
mass of trabecule and contained fat-mass is carried to an extreme
in the opposite direction in the case of Xenopus, where (see
p. 924) a tough mass of connective-tissue fibres and interspersed
fat surrounds the lymph-hearts.
It is said, and im a sense correctly, that there are no lymphatic
glands in the Frog. The absolute truth of this generalization
(which of course applies to other lower Vertebrates as well as the
Frog) depends upon what is meant by the term ‘lymphatic gland.”
For if we regard a lymphatic gland as an enlargement on the
course of a lymphatic vessel the lumen of which is subdivided by
trabecule, then the structures described here in Hemisus are at
least not very unlike lymph-glands. One cannot but think, in
view of the masses of fatty tissue with which they are partly
plugged, that they must play some important part in the function
of the lymphatic system; and the existence of fatty masses in
Xenopus, described below*, strengthens this supposition in that
it shows that the structure is not unique.
The lymph-heart itself is of considerable size, though not
so colossal as in Greviceps. It measures fully 4 mm. in length
and is rather elongate in form. It is displayed in text-fig. 186, .2.,
from which it will be seen that the heart occupies quite a normal
position. It les, however, quite definitely within the large
saccus iliacus which has just been described. In Rana esculenta
the lymph-heart of the same pair is said by Gaupp7 to lie in
a special sac of its own. This is certainly not the case with
Hemisus. The posterior lymph-heart lies at the inner angle of
the saccus ilacus. The heart is in contact with the vastus
externus, upon which it lies ventrally; with the ilio-coccygeal
muscle to the inside. On the opposite side its wall lies freely
within the lymph-sac already mentioned and described at length.
It is in front of the pyriformis muscle. It is not very closely
related to the glutzeus muscle. The septum bounding the saccus
iliacus posteriorly is attached along the vastus externus and ends
on the wall of the heart, or at least is firmly attached to the
posterior wall of the heart at this point. Just in the angle
where the septum in question and the heart meet is a smallish
circular ostium (text-fig. 186, O.), which I take to be a communi-
cation between the saccus iliacus and the heart. A small blood-
vessel, as is shown in the figure (m), runs along the outer side of the
heart forwards, dipping down to the gluteal muscle; it is possible
that this is the vein into which the heart opens. The heart is
rather pear-shaped and lies with its long axis parallel to the long
axis of the body. The narrow end isanterior. The heart is easily
to be distinguished from the posterior septum of the saceus iliacus
* See p. 924. + See his edition of Ecker’s ‘ Frog.’
1908.] OF THE BATRACHIAN GENUS HEMISUS. 923
which is attached to it, and from such septa generally, by its
yellowish colour, due of course to the fact that its walls are
muscular and not formed of connective tissue. The outer wall
of the heart is quite smooth and has a bronzy appearance, due to
its muscular walls; the internal surface is sculptured into raised
Text-fig. 186.
Lymph-heart (posterior) of Hemisus guttatum and associated structures.
Uh. Lymph-heart cut open to show its interior. x, Vein referred to in the text.
O. Ostium referred to in the text.
muscle-masses here and there with, of course, depressions between.
The fibres in the cuter wall of the heart run largely, as I have
convinced myself by a microscopic examination, across its long
diameter.
924 MR. F, E. BEDDARD ON THE ANATOMY [Dec. 15,
§ The Posterior Lymph-Hearts of Xenopus.
The posterior lymph-hearts of this genus are not, so far as I
am aware, known. As I have dissected them in two individuals
of this African Frog, and as they present certain remarkable
differences from those of other Frogs, | think it worth while to
append to my account of these structures in Hemisiws some notes
upon the lymph-hearts of Yenopus. When the animal is dissected,
a mass of yellow fat is seen to lie upon the thighs and to spread
Text-fig. 187.
Hinder part of body of one side of Xenopus levis.
1h. Mass of fat in which lie lymph-hearts—distinguished as light circles.
s. Sense-organs of lateral line.
upwards for some way on to the back. The mass of fat upon the -
proximal region of each thigh is, as is to be seen in the annexed
figures (text-figs. 187, U.A., & 188, U.h.), of roughly circular out-
line. It is seen on excavation by the scalpel to be of some
depth, and I regard the space in which it hes as representing the
saccus iliacus of the lymph-system of Hemisus and other Frogs.
There is, however, no empty sac here. The fat entirely fills the
1908. | ‘OF THE BATRACHIAN GENUS HEMISUS. 925
cavity. Theactual fat itself is contained in a very close meshwork
of fibres of connective tissue. When this is cut into and pressed
the actual fat readily escapes and floats up to the surface of the
water in the dissecting-dish.
The figure to which reference has been made (text-fig. 188)
shows the saccus iliacus and adjacent structures intact on the right
side save for the removal of the skin. The thin membrane covering
the saccus iliacus is left intact. The cut edge of the membrane
lying further to the right and continuous with this is the inner
lateral wall of the femoral lymph-sac. On the left side the
wall of the saccus iliacus is not shown, having been removed by
tearing it away, and thus exposing the spongy fat-laden interior
Text-fig. 188.
LA.
Hinder part of body of another individual of Xenopus levis.
A. Single lymph-heart seen through delicate membrane which covers iliac fat-body
1.h. Two lymph-hearts lying in fat-body and cut open.
of the lymph-sac in question. This wall does not fit loosely, but
is intimately connected with the fat-holding reticulum below, so
that it has to be removed in fragments. The border-line between
the two sacs is shown by a depression running along the mem-
brane; it is along this line that the boundary between the sacci
jliacus and femoralis is fixed. At the inner upper corner of the
saccus iliacus, where it abuts upon the middle line of the body, a
triangular flap of the covering of the saccus iliacus is cut and
reflected. This shows a portion of a muscle which is presumably
926 MR. F. E, BEDDARD ON THE ANATOMY [ Dec. 15,
to be regarded as the cutaneus iliacus, and which has already
been described in Hemisus and Rana guppyt.
Tt will be noted that, as in Hemisus, this muscle lies within the
saccus iliacus, and that it is entirely attached to the wall of that
sac at its insertion, and does not reach the skin at all. In
Hemisus some of the fibres of the corresponding muscle are thus
intercepted, but the rest reach the skin. In Rana all of the
fibres of the muscle reach the skin. It is noteworthy that the
wall of the saccus iliacus, as is clearly shown in the figure, is
distinct from the overlying skin and not fused with it as are the
dorsal walls of other lymph-spaces such as the adjoining femoral
to the lateral septa, to which reference has already been made.
On the left side the attachments of the septa bounding the femoral
lymph-sac are shown in their attachments to the skin. On the
right side a circular elevation is visible (text-fig. 188, A, p. 925),
lying pretty well in the centre of the area occupied by what I here
compare to the saccus iliacus. This, when cut open, proved to be
the single lymph-heart of that side of the body ; 1t contained a large
orange-coloured clot (presumably of lymph), the darker colour of
which,as compared with the surrounding tissues, was obvious before
cutting into the lymph-heart. The clot was roughly spherical in
outline, slightly flattened from above downwards. Its greatest
diameter was 4mm. Thus the heart itself may be considered to
be a little larger. On the opposite side of the body there was no
single lymph-heart corresponding to that of the right side. There
were most distinctly visible, when the surrounding spongy tissue
was carefully cut away, two perfectly detached and separate posterior
lymph-hearts. Between them was some of the spongy tissue
which fills up the lymph-sac, and there was no open communication
between the two hearts, whatever may be the facts with regard
to a communication by means of finer tubes. These two lymph-
hearts lay one behind the other in a perfectly straight line,
parallel with the longitudinal axis of the body. A careful com-
parison of the relative positions of the two lymph-hearts of the
left side of the body with the single lymph-heart of the right
side of the body, showed very clearly that the latter occupied a
place midway between the anterior wall of the anterior and the
posterior wall of the posterior left lymph-hearts. It would thus
appear to correspond to the two of them—that is, the single right
lymph-heart has been produced from a conerescence of two
originally separate lymph-hearts, or it has given rise by division
to two lymph-hearts on the left side. Of these two lymph-hearts
of the left side the anterior was distinctly the larger, which is not
very well shown in the figure referred to (text-fig. 188). This ~
larger left lymph-heart, moreover, contained a blood-clot which
was not to be seen in the interior of the hinder and smaller lymph-
heart of the left side. This suggests, of course, that the systole and
diastole of the two consecutive hearts are not synchronous,
but that one is in systole, while the other is in diastole. The
remains of the lateral line which are so prominent in Xenopus as
1908. ] OF THE BATRACHIAN GENUS HEMISUS. 927
stitch-like along the sides of the body did not appear to me to have
any exact relation to the series of lymph-hearts of the left side,
that is, they did not accurately overlie them. The hearts lay to
the inside of the lateral line, as is also shown in the figure.
The soft and yet toughish and even sticky tissues which form
the fat-holding plug which nearly, or quite, fills the saccus
iliacus, and in which the hearts are imbedded, are not altogether
easy of dissection, and the exposure of the lymph-hearts with
much neatness is very difficult. It is possible that in the
specimen which has just been described I have overlooked a
lymph-heart; for in two others which I have dissected there were
undoubtedly three pairs of posterior ltymph-hearts. In one of
these, which happened to be rather a small individual, the hearts,
at any rate of one side of the body, were distinctly visible directly
the skin was removed and the fatty mass exposed (see text-fig. 187,
p. 924). This latter was, as usual, very yellow. Conspicuous—
this time by their paler and browner colour—-were the three
lymph-hearts, of which the most anterior was not only the most
conspicuous but the largest. The other two lying in a row behind
it were, however, quite evident, though probably they would
escape the attention of anyone not aware of their existence.
Text-fig. 189.
Mliac fat-body of Xenopus levis dissected to show three lymph-hearts.
Indeed, it was after the discovery by dissection of three
lymph-hearts on each side of the body of a third specimen, that
IT noted the external appearance of these lymph-hearts in the
small specimen to which reference has just been made. In this
latter specimen the three lymph-hearts of the right side are repre-
sented, as seen by dissection, in text-figure 189, the upper wall
of each heart having been removed in order to display the interior.
It happened that this particular specimen was especially favourable
Proc. Zoou. Soc.—1908, No. LIX. 59
928 MR. F. E. BEDDARD ON THE ANATOMY [ Dec. 15,
for displaying the structures in question. The preservative
alcohol had hardened the hearts and the surrounding tissue in
such a way as to render them very tough and had decolorized the
hearts, while rather deepening the yellow of the fat-mass. It
was thus found convenient to cut through with a pair of scissors
the fat-mass of the left side of the body along a line presumed to
pass through the hearts. Text-figure 190 shows that this was
successfully accomplished, and represents the two sides of the cut
whereby the mass of fat and the contained lymph-hearts were
divided longitudinally.
Text-fig. 190.
Vliac fat-body of Xenopus levis cut longitudinally into two halves to show
cavities of lymph-hearts.
This longitudinal section passed, I believe, very nearly through
the middle of each lymph-heart. It will be seen that they are not
all of them of exactly the same size, though the differences between
them are not very great. In the dissection (text-fig. 189, p. 927)
of the hearts on the right side of the same Frog it will be noted
that the last of the three hearts is considerably the largest of the
three, perhaps twice as large as either of the others. This did
not appear to be the case on the left side of the body. What is _
particularly striking about these hearts when seen evenly divided,
and in section, is the great thickness of their walls. This is not
exaggerated in the drawing to which reference has been made.
The dissection of the corresponding hearts on the opposite side
of the body (text-fig. 189) of this specimen shows the apertures in
the walls of the hearts which presumably admit lymph into their
1908. ] OF THE BATRACHIAN GENUS HEMISUS. 929
interior. The thickness of the walls is also shown in this
drawing. It is doubtless partly owing to the strong contraction
-of the hearts at the moment of death.
IT am not aware that the posterior hearts of Xenopus have
been described. The pectoral lymph-hearts, however, of this
Frog are described and figured by Dr. Bles in his beautiful
memoir * upon the larval development of Xenopus. There is but
-a single pair, and each is enveloped in a lymph-sac of its own.
Dr. Bles remarks that the early development of these structures
in Xenopus is remarkable, since in Rana temporaria and Bufo
calamita the author found that the pelvic lymph-hearts do not
appear until after the metamorphosis. This, however, does not
argue that these structures are new formations and not com-
parable with the lymph-hearts in Urodeles. It is hardly likely
that the anterior and posterior lymph-hearts are not parts of the
same series, and therefore the early development of the pectoral
hearts in Yenopus is to be set off against the late appearance of
the pelvic hearts in certain Frogs, including, as it is to be supposed,
Xenopus itself.
Inasmuch as the lymph-hearts of Rana are connected with
veins supplying the fore and hind limbs respectively, a suggestion
may be made as to the retention (or, if my opinion be not accepted,
the multiplication of these hearts) of three pairs of posterior
lymph-hearts in _Yenopus. While in most Frogs, indeed in all, the
hind limbs are the powerful swimming-organs of the animal, and
exceed in size the relatively feeble fore limbs, the disproportion
reaches its maximum in Xenopus. Of this Frog Dr. Bles justly
writes :—‘“ The size of the arm is altogether out of proportion to
the size of the leg, which is an extremely powerful swimming-
organ. The limbs of Xenopus as a Frog are paralleled by the
limbs of Macropus asa Marsupial” +. The excessive size of the
hind limbs in Yenopus bears some relation to the triple lymph-
hearts. It is true that I have not succeeded in finding the veins
into which the hearts open. But it can hardly be doubted, from
the position which they occupy, that their orifices are into veins
connected with the legs.
Attention may be drawn to the variability of these posterior
lymph-hearts in Xenopus. This fact, as it appears to me, is of
itself evidence, though naturally not of a positive character, that
the structures in question are not a new formation, but are
derivatives of the chain of lymph-hearts in certain Urodeles.
The variability affects, as I think, the number of the hearts,
which does not only differ in individuals, but from side to side of
the same individual. But even if I am mistaken in this and
have simply failed to find the supposed missing hearts in those
specimens where only one or two appear to exist, there still
remains the variability in point of size. There is, I hope, no
* “The Life-history of Xenopus levis Daud.,” Trans. Roy. Soc. Edinb. vol. xi.
pt. ili. 1905, p. 789.
+ Trans. Roy. Soc. Edinb. fom. cit. p. 819.
59*
930 MR. F. E. BEDDARD ON THE ANATOMY [ Dee, a,
doubt whatever about this. An inspection of my figures, which
were drawn quite independently of any directions from myself,
will settle this matter. It is well known that variation is apt:
to affect organs or series of organs undergoing reduction. And, —
therefore, I dwell upon this fact as bearing upon the view upheld
here, that in this Frog we havea persistent multiplicity of lymph-
hearts, such as that which characterizes certain Urodeles, ,but
which is considerably reduced.
The interest attaching to these facts concerning the posterior
lymph-hearts of Xenopus is not only that they are—so far as I
am aware—a contribution to the anatomy of that Frog. Their
chief interest centres in the possibility of comparing them with
the multiple lymph-hearts of certain Urodeles. With regard to
this matter Dr. Wiedersheim sums up as follows in his ‘ Vergleich-
ende Anatomie der Wirbelthiere’* :—‘“ Bei Salamandra macu-
losa und Stredon pisciformis sitzen zahlreiche Lympherzen langs
des Sulcus lateralis unter der Haut und zwar entfallen bei dem
erstgenannten Thier auf dem Schwanz (beide Seiten zusammen-
gerechnet) 10-12, auf den Rumpf mindestens ebensoviel. Bei
Siredon pisciformis finden sich jederseits 8 rhythmisch pulsirende
Liympherzen die wie bei Salamandra maculosa aus ovalen, von
quergestreifter Muskulatur umwickelten Bliischen bestehen.”
These facts were first discovered by Weliky ft.
§ The Lymph-Hearis of Rana guppyi.
Although it is usually stated in books and memoirs dealing
with the anatomy of ana that the posterior lymph-heart is a
single structure on each side, this is not the universal view.
Hoyer has given an account, with figures in the text, of the
lymph-heart of Rana esculenta. A longitudinal section of one of
the hearts given by him shows three distinct cavities, of which
the hindermost is the larger. It is to be noted, however, that
this figure is hardly convincing of itself as to the existence of
three separate hearts, such as we haye seen exist in Xenopus.
For the muscular wall shown in Hoyer’s figure extends without a
break over the three cavities; there isno such complete separation
as is indicated in text-figure 189 (p. 927) of the present communi-
cation. And so far, therefore, Dr. Hoyer’s figure bears out the
statement of Oehl (which is quoted by him), viz. “ Das Herz gelappt
ist,” and that of Ranvier (also quoted by him), who observed
that each apparently single posterior ‘lymph-heart was divided
by partitions into several divisions. Weliky, however, according
to Hoyer, found that each heart “aus 3 gesonderten Abtheilungen
besteht: dass also 3 hintere Lympherzen jederseits vorhanden
sind.” This, of course, agrees precisely with the facts which I
have detailed in the present paper with regard to Xenopus. At
the same time it cannot be disputed that the conditions obtaining
* Zweite Auflage, Jeni, 1886. + Zool. Anz. 1884.
{ Bull. Acad. Cracovie, 1904, p. 228.
1908. | OF THE BATRACHIAN GENUS HEMISUS. 931
in Xenopus are more primitive than those found by the above
writers to occur in Kana. The hearts are larger and, extending
as they do over a larger area of the body, are more widely
separated, and thus more completely distinct than can be the
case with Rana, when so small a cleft contains the three hearts—
or the trifid heart,—which are not indeed difficult to overlook
altogether. Whatever may be the case with Rana esculenta, I
found in a specimen of Rana guppyi a single lymph-heart on each
side posteriorly, and measuring about 6 mm. in length; it lay in
quite the usual position in front of the pyriformis muscle, but
well behind the gluteus, separated from it, in fact, by the posterior
end of the ium.
The direction of the heart is rather obliquely outwards, very
nearly parallel to the pyriformis muscle. Although each heart may
be accurately described as a single heart, the cavity is completely
divided across its major length (7. e. transversely to the long axis
of the pyriformis muscle) by a septum of the same appearance
and texture as the general parietes of this lymph-heart into
two quite separate chambers. This division, however, is merely
a septum; there 1s no question of any separation of the obviously
single heart into two hearts. Nor is there any external constric-
tion of a marked character which could fairly justify a statement
that there were two consecutive lymph-hearts present on each side.
I may say that the lymph-hearts on both sides of the body of
this Frog were identical. This, it may be observed, is a very
different condition of the posterior lymph-heart to that which
has just been described in Xenopus. In the latter genus, I repeat,
there are three distinct lymph-hearts on each side, in Rana guppyi
only a single heart the cavity of which is divided *.
I may take this opportunity of calling attention to the anterior
lymph-heart in Rana guppyi, where it 1s very conspicuous on
account of its large size. It is quite 9 mm. iong (when slightly
stretched perhaps) and lies in a lymph-sac completely covered by
the suprascapula, which has to be lifted up in order to display it.
The sac and the contained heart are bounded by the longissimus
dorsi and the retrahens scapule on the inside, and by the trans-
verso-scapularis tertius on the outside+. The greater part of
the lymph-heart consists of a single chamber, in which there is
no trace of any septa. At the inner side, however, an incomplete
septum partly separates off a very small chamber, which is about
1-5 mm. in breadth. This contains, I believe, the orifice into the
vein. It would be better perhaps to describe the incomplete
septum as a valvular flap which regulates the flow of blood and
lymph. In any case, there is no complete division of this anterior
lymph-heart into two chambers, such as has been described in
the posterior lymph-hearts of the same Frog.
* Mr. Burne kindly allows me to quote a letter in which he informs me that
the posterior lymph-hearts in a specimen of this Frog in the College of Surgeons
Museum are quite similar.
+ For these muscles, v. supra.
932 MR. F. E, BEDDARD ON THE ANATOMY [ Dec. 15,.
§ Résumé of principal Muscular and other Characters
of Hemisus.
The principal characteristics of the genus Hemisws which f
have been able to elucidate in the foregoing pages, are as.
follows :—
(1) The rectus abdominis has but one inscriptio tendinea..
It cannot be distinguished laterally from the obliquus.
externus and is folded over itself at edge of the abdomen,
where it joins the thigh, in a remarkable way.
(2) The obliquus internus consists of a thick layer of muscles,
arising mainly from the dorsal aponeurosis to the outside
of the ilia, which end upon a thin membrane which repre-
sents the ventral portion of this muscle. It is uncovered
for the greater part by the obliquus externus.
(3) The sternohyoid consists of three distinct muscles inserted
separately and behind each other on to the hyoid body.
(4) There is no omohyoid.
(5) The submaxillaris appears to be specialized into three:
tracts on each side, of which the fibres run at different.
angles to each other.
(6) The dorsal portion of the depressor mandibulz is present.
(7) The latissimus dorsi has an origin extending considerably
beyond (behind) the scapula and joins early with the
infraspinatus.
(8) There is no rhomboideus muscle.
(9) There are altogether four “serratus” muscles on each side:
of the body; 1 and 2arise respectively from the transverse
processes of vertebra 3 and 4; 3 arises from the transverse:
processes of both those vertebre; 4 arises from the neck
laterally in front of the third vertebra.
(10) In the thigh the rectus internus minor partly arises from
the skin.
(11) There are two pairs of large fat-masses lying in lymph
spaces and another pair corresponding in position to the
thymus lying beneath the skin.
(12) The stomach is elongate, with a narrower portion bent to:
form a U with the major portion.
(13) The small intestine is traversed throughout by closely set.
transverse folds.
(14) The fat-bodies are very large.
(15) The oviducts are much coiled and with a long drawn out.
gutter-hke funnel.
(16) The lymph-sacs are not particularly large and in the-
usual position.
(17) The hyoid has large extra-hyals anteriorly which fuse
ventrally below the hyoglossus muscle, and the body of the-
hyoid is very thick, with a bony plate distinct from and
overlying the convex and swollen cartilaginous layer.
—
The above 17 characters appear to me to be the chief anatomical
1908. | OF THE BATRACHIAN GENUS HEMISUS. 933
distinguishing features of Hemisus, besides certain osteological
and external peculiarities which I do not deal with here. Of the
former there are more that are peculiar to Hemisus than of those
which ally it to its allies. Hemisus is peculiar, so far as is at
present known, in Nos. (1), (2), (8), (6), (7), (41), (13), (17)—that
is, in nearly one-half of those which I have selected.
Hemisus agrees with both of its allies, Breviceps and Rhino-
derma, in (5)—at any rate, as to the fact that there are various
additional muscles, not present in Rana, upon the floor of the
mouth ; there is, however, no detailed agreement between the
three genera in the disposition of these muscles. In (10), (12)
Hemisus agrees with Rhinoderma, and differs from Sreviceps
in (16). Hemisus agrees with Breviceps and differs from Rhino-
derma in (4), (8), (14). I am not quite certain as to the re-
maining features of the anatomy, which I have made use of as
indications of closer or more remote affinity. These facts, and
indeed others which will be found in the foregoing pages, do
not, as it seems to me, permit of a very decisive placing of
Hemisus with regard to the two remaining genera of Engysto-
matid Frogs whose anatomy is known. The particular likenesses
which Hemisus shows to Breviceps, as opposed to Rhinoderma,
may be increased by the addition of the fact of the partial
inclusion of the limbs within the area of the trunk, and by the
division of the rectus abdominis muscle by only a single inscriptio
tendinea. But Breviceps remains, after all, an extremely specialized
type in many ways. General reflections upon the arrangement
of these Frogs will, in fact, be better deferred until more
anatomical facts have been collected.
§ Résumé of principal new Facts.
It may be convenient to extract from the foregoing account of
Hemisus, and of Xencpus and Rana, the following principal new
facts which I have been able to ascertain :—
(1) Hemisus is characterized by the existence of three pairs of
large-lobed fat-bodies, of which one pair correspond in
position to the thymus in other Frogs, the second lie in a
cavity (?a lymph-sac) behind the shoulder-girdle, and the
third pair are contained in a sac partly overlapping the
thigh, which is to be compared with the saccus iliacus of
Rana. The prerenal fat-bodies are also very large.
(2) Xenopus has a similar pair of fat-bodies in the repre-
sentative of the saccus iliacus, the fatty tissue, however,
straying further forward on to the back.
(3) The saccus iliacus in Hemisus and Xenopus is divided by
trabecule in the interstices of which lies the fat-body ;
the commencement of such a division of the saccus iliacus
is seen in Rana guppyi. It is possible to compare these
structures with lymph-glands.
934
(4)
(5)
(6)
MR. G. A. BOULENGER ON A [ Dec. 15,
Hemisus has a single pear-shaped’ posterior lymph-heart,
which, unlike that oe Rana, lies in the saccus iliacus.
Xenopus has a chain of three perfectly distinct posterior
lymph-hearts on either side of the body, which lie in the
saccus iliacus.
In Rana guppyt the single posterior lymph-heart of each
side is completely divided into two consecutive chambers.
(7) Neither in Hemisus nor in Rana guppyi is there any
(8)
division of the anterior lymph-heart.
Hemisus possesses a hyoid which is remarkable in several
ways and unlike that of other Batrachians. The extra-
hyals are large and meet in the middle line below and not
in contact with the body of the hyoid; the latter consists
of a cartilaginous plate continuous with the corona, which
is greatly thickened posteriorly by a nucleus of laxer
tissue, and above which lies a plate of bone—not imbedded
in it, but distinct from it.
(9) A comparison of Hemisus with Breviceps and Rhinoderma
(10)
allows of the extraction of certain characters apparently
distinguishing the Engystomatide, 7. e. specialization of
muscles of floor of mouth, division of sternohyoid, con-
nection of rectus internus minor with skin.
Hemisus, though a burrowing and ant-eating genus like
Breviceps, shows comparatively few special structural like-
nesses to it. The principal resemblances are: partial
inclusion of limbs within the trunk; (?) absence of omo-
hyoid and rhomboideus ; great strength of muscles
attached to the shoulder and fore limb, which, however,
are not entirely the same muscles in the two types; the
modifications of the muscles of the hyoid and the floor of
the mouth, which are to some extent similar in the two |
types. But the many differences in the abdominal and
dorsal musculature obscure and outweigh the special
likenesses, which might be referred to similarity in habits
and mode of life.
Description of a new Species of Lacerta from Persia.
By G. A. Boutencer, F.R.S., V.P.Z.8.
[ Received October 13, 1908. j
(Plate LX VII.*)
LACERTA CHLOROGASTER.
Head moderate, once and three-fifths to once and three-fourths —
as long
2 as broad; snout moderately long, obtuse. Rostral not
touching the nostril ; one postnasal; a single anterior loreal;
four (rarely five) upper labials anterior to the subocular ; a
complete series of granules between the supraocular and the
* For explanation of the Plate see page 936.
aELIUS WD) OREO) WsLO) Walkral alow "1
UML SulonyD Je Tsp uaeg’p
UA te SOGL S 42d
1908. ] NEW SPECIES OF PERSIAN LIZARD. 935
supraciliaries: occipital usually shorter and broader than the
interparietal ; temple covered with smaiil smooth scales, with a
large masseteric disk and a curved tympanic shield; a large
anterior supratemporal, usually in contact with the fourth supra-
ocular. A feeble or very indistinct gular fold; 20 to 27 gular
scales on a line between the collar and the third pair of chin-
shields ; collar with feebly serrated edge, composed of 7 to 9
plates. Dorsal scales hexagonal, longer than broad, strongly
keeled, juxtaposed or faintly imbricate ; lateral scales more feebly
keeled, smooth towards the ventrals, a little smaller than dorsals,
3 or 4 corresponding to the length of a ventral plate; 44 to 50
scales across the middle of the body. Ventrals in 6 longitudinal
series, the second series on each side from the median line the
largest ; 25 to 30 transverse series. Preanal plate large, bordered
by a single series of scales. The hind limb does not reach beyond
the shoulder. 27 to 30 lamellar scales under the fourth toe.
Femoral pores 14 to 18. ‘Tail twice, or nearly twice, as long as
head and body; upper caudal scales strongly keeled, pointed
posteriorly. Head and back greyish-olive in the male, the sides
and limbs yellowish-green with a black network, or black with
small yellowish-green spots; a few turquoise-blue spots may be
present behind the shoulder. Pale golden-brown above in the
female, with small blackish spots and a dark brown lateral band
with wavy outlines. Lower parts yellowish-green to bright
grass-green, the males with a series of turquoise-blue spots on
the outer ventral plates and with the throat often blue or bluish-
green ; anal region and lower surface of hind limbs often lemon-
yellow. Iris brownish.
ic Or
MotalMleme thes Ve) see see caer <b akan s ae 182 mm. 137 mm.
TELEE VCS Shani orc nes depmes seca oon ene i 13
"INV ie eh OF Yon Pe Vet |. ed ye Gas On Cee 10 8
From end of snout to fore limb ... 2A 18
- - vent ...... 61 57
Tikogeylhtonl ose Oe ote Sy ses ae anneee Ree 24. 19
le liiayol lbiaml oy yeh, oe dane se sabe noe ROS BORE 34 29
Bete ca tl Pons SORES Peele Biancs sal ae PAL 80 *
This species, which is intermediate between JL. praticola
Eversm. and Z. taurica Pall., was first discovered in May, 1907,
by Mr. R. B. Woosnam at Enzeli, on the south coast of the
Caspian Sea. Thanks to the courtesy of Mr. H. N. Rabino,
British Consul at Resht, I have since received further specimens
from the same locality, some of which reached me alive or recently
dead, thus enabling me to describe the natural coloration.
Mr. Woosnam has furnished me with the following note
respecting the occurrence of L. chlorogaster :—
“These lizards were caught on the narrow dry sandy peninsula
between the Caspian Sea and the large salt-lagoon at Enzeli,
* Tail reproduced.
936 DR. E. LONNBERG ON WART-HOG | Dec. 15,
where they were quite common and frequented chiefly the sandy
banks and dry reed-fences around the gardens. They are pro-
bably to be found all along the south coast of the Caspian, for
although none was obtained during the journey along the coast.
from Resht to Asterabad Bay, this may be accounted for by the
fact that it was then early in the year, February and March, and
too cold. But I once or twice caught a glimpse of a lizard, among
the scrub on the dry sand-dunes near the shore, which I feel sure
now must have been this species. None was obtained on the
smaller western peninsula, but I should not like to say they are
not to be found there, for they probably are, and I expect the
species exists all along the south coast of the Caspian Sea where it.
is dry and sandy.”
EXPLANATION OF PLATE LXVII.
Lacerta chlorogaster, male, natural size, and enlarged views of upper
surface of head and anal region.
3. Remarks on some Wart-Hog Skulls in the
British Museum. By Dr. Eryar Lonnpere, C.M.Z.8. Ke.
[ Received October 19, 1908. |
When recently describing the mammals collected in German
East Africa by Prof. Dr. Y. Sjéstedt * I made some remarks
about different races of Wart-Hogs, and, with some hesitation, I
expressed the opinion that “for the present at least” five such races.
must be ‘discerned and distinguished by names.” Since then
I have had the opportunity, thanks to the kindness of Mr. Oldfield
Thomas, of studying the material of Wart-Hogs in the British
Museum (Nat. Hist.), and, thanks to the kindness of Dr. 8.
F. Harmer, that in the University Museum of Cambridge. It was
quite easy to recognize among this material the five races men-
tioned in the paper quoted above, in such cases where they were
represented by skulls of adult specimens, especially boars.
A few remarks about these skulls may be of some value for
future study of these animals, as I did not have access to speci-
mens of all five races when writing the first paper.
Phacocherus africanus (Gmelin) appears to be the largest or one
of the largest of these races. A skull of an adult boar of this kind
in the British Museum from the typical locality, Cape Verde,
measures 440 mm. in length, but the extreme tips of the nasals are.
not complete, so that this measurement should be a little longer.
The postorbital portion of this skull is very long, measuring 59mm.,,
but it is at the same time very broad, viz. 58 mm. across the flat
area. By this characteristic Ph. africanus is very easily distin-
guished from Ph, eliani, which also has along but at the same time
* Lonnberg: Mammals, in Wiss. Ergebn. d. schwed. zool. Exp. nach dem Kili-
mandjaro, dem Meru etc., 1905-1906, unter Leitung von Prof. Dr. Yngve Sjéstedt.
1908. ] . SKULLS IN THE BRITISH MUSEUM. 937
very narrow postorbital portion of the skull. Thisis proved by the
following measurements obtained from specimen No. 69.10.24.47
in the British Museum, ¢ ad. from Zorilla, Abyssinia. The length
of this skull from the tip of the nasals to the occipital crest is
388mm. The postorbital portion has a length of 60 mm., but the
breadth of its flat area is only 244 mm. These differences in the
dimensions of the postorbital portion of the skull become still
more striking if they are expressed in percentages of the length
of the skull. In such a case the length of the postorbital portion
of the skull of Ph. africanus will be found to be 13-4 °/, and the
width of its flat area almost the same, or 13:1 °/, of the length of
the skull. The same percentages for Ph. eliani are respectively
15:4 °/, and 6:3 °/,. Ph. africanus has a comparatively narrow
interorbital space, which corresponds to only 30 °/,* of the length
of the skull. In Ph. eliani the interorbital region is a little
broader, so that it corresponds to 31°7 °/, of the length of the skull.
The combined characteristics of the postorbital and interorbital
portions of the skulls of these two races give the impression
that they are comparatively longer than those of other races.
The skulls of these races are therefore at once distinguished from
others. Ph. massaicus has a comparatively long postorbital
region, viz. about 14 °/, of the length of the skull, but as it is
very broad at the same time, its flat area measuring about 14°5 °/,,
and the interorbital region as well is very broad, being 38°8 °/, of
the length of the skull, no confusion with other races is possible.
Ph. sundevallii has a somewhat shorter postorbital region, viz.
13:7 °/,, and the flat area of the same is considerably narrower,
viz. 11°/,: at the same time its interorbital region is much
narrower than in Ph. massaicus, so that the percentage expressing
its relation to the length of the skull, 32°38, resembles that of
Ph. cliani. In such a way these four races may be easily
distinguished from each other, if the material is derived from
adult males.
Two Wart-Hog skulls in the British Museum from Angoni-
land (No. 8.2.14,1, ¢ ad., and 8.2.14,2, ¢ jun.) show some
affinity to swndevallii. Their interorbital width is respectively
33-4 °/, and 32:1°/, of the length of the skull (thus rather
similar in this respect to Ph. sundevallii), but the postorbital
portion is smaller than in Ph. sundevallii, its length beg about
11:7 °/, and the width of its flat area 10-1 °/, of the length of the
skull in the adult. In the younger specimen the last-mentioned
dimension is still smaller (8°1 °/,), as always is the case with the
young ones, and cannot be considered. More material is needed
before anything can be decided about this Wart-Hog. It may,
however, be added that its choane are rather wider, 36 mm. in
the adult, than in the typical Ph. sundevallii, 29 mm.
Two other skulls of Wart-Hogs in the British Museum
one from Ukanga, Lake Nyassa (No. 91.5.9,3, ¢ ad.) and
* The interorbital measurements are always counted at the middle of the orbit.
938 DR. E. LONNBERG ON WART-HOG - || Decgglta;
another from Lake Mweru (No. 94.3.8,17, ¢ ad.), are quite
similar inter se. Their interorbital width is respectively 28°8 °/,
and 29°3 °/,, the length of their postorbital portion 12 °/, and
12:3 °/,, and the length of the postorbital flat area 11‘1 °/,
and 11:4 °/, of the length of the skull. A third skull from Lake
Mweru (No. 94.3.8,18, ¢ jun.) is also similar with regard to the
first two dimensions, viz. 30°/, and 12 °/., but the postorbital flat
area is narrow, 8°7 °/,, im consequence of its youth. It appears
from this as if the Wart-Hogs of the country between Lake
Nyassa and Lake Mweru agreed in having a comparatively very
narrow interorbital region, narrower than in Ph. sundevallii.
The width of the flat postorbital portion is similar to that of
Ph. sundevallii, but the length of the same is somewhat shorter.
With regard to other measurements, it seems as if the skulls
from Angoniland, Nyassa, and Lake Mweru had a somewhat
longer preorbital portion (distance from tip of nasals to anterior
orbital margin) than both Ph. swndevallii and Ph. massaicus.
Comparative studies of more material of fully adult animals
may thus in the future prove that the Wart-Hogs inhabiting the
countries adjoming Lake Nyassa and Lake Mweru are racially
different both from swndevallii, inhabiting Natal and probably
Transvaal] and the southern parts of Portuguese East Africa, and
from Ph. massaicus, inhabiting the Masai country in German
Hast Africa.
How widely Ph. massaicus is distributed cannot be stated for
the present. The skull of a young Wart-Hog of male sex from
Uganda in the British Museum (No. 95.4.3.42) agrees so far with
Ph. massaicus 11 having a very broad interorbital region, which
corresponds to 37 °/, of the length of the skull. But, on the other
hand, the postorbital portion is rather small, its length being only
11°/, and the width of its flat area only 10°5 °/, of the length of
the skull. If this smallness be not due to the youth of the
specimen, there must exist a separate race of Wart-Hog in
Uganda which should be easily recognized by the two combined
characteristics: great interorbital width and shortness of the
postorbital region.
Jn the specimen examined the length of the postorbital region
exceeds the width of its flat area by 2 millimetres only, but,
as experience proves that the latter dimension increases more
with age than the former, it is probable that in adult Wart-Hog
boars from Uganda the width of the postorbital flat area is
greater than the length of the same portion of the skull (as also
is the case in Ph. massaicus).
In the collections of the British Museum are two Wart-
Hog skulls, numbered 0.3.27.16 and 0.3.27.17, which aroused
the interest of the present writer more than all the others.
Both these, which were presented by Lord Delamere, have no
traces of upper incisors, and 0.3.27.16 has no incisors in the
lower jaw either, with the exception of two small pea-shaped
rudiments lying in corresponding grooves of the jaw-bone ; these
1908. | SKULLS IN THE BRITISH MUSEUM. 939
rudiments appeared to represent the median pair. Specimen
0.3.25.17, on the other hand, is provided with four well-developed
incisors in the lower jaw. The lower incisors are thus subject to
great variation, but the upper ones appear to be constantly
missing, as the premaxillary is too thin to carry any teeth, justas
in Ph, ethiopicus. The two skulls in question resemble the Cape
Wart-Hog (Ph. ethiopicus) in other respects, too, both in general
shape and with regard to particular features, as will be seen from
the following comparison. The postorbital portion of the skull is
very short in Ph. ethiopicus, about 10°3 °/, of the length of the
skull in a specimen in the Royal Natural History Museum of
Stockholm (brought home by Sparrman), 10°5 °/, in specimen
0.3.27.17 and 10-9 °/, in specimen 0.3.27.16 in the British Museum.
The width of the postorbital flat areais greater than the length of
this portion, viz. 13°3 °/, in Sparrman’s specimen, which is the
oldest, and respectively 11:9 °/, and 11:5 °/, in Lord Delamere’s
two specimens. The interorbital width is rather greater in
Sparrman’s specimens, viz. 36°5 °/., than in the two others, respec-
tively 33:0 °/, and 34-7 °/,. There was no locality indicated on the
labels of Lord Delamere’s two Wart-Hog skulls, and I believed,
therefore, judging from their resemblance to Ph. ethiopicus, that
they originated from the Cape. Mr. Oldfield Thomas, however,
kindly informed me that this was not the case, as Lord Delamere
had travelled in North-eastern Africa, Somaliland, and British
East Africa, and the skulls were most probably from Somaliland.
This made the matter more complicated, but at the same time
more interesting, as it was not probable that the same race of
Wart-Hog inhabited two countries so far apart when the inter-
vening countries were occupied by widely different races. A
renewed examination revealed also that Lord Delamere’s Wart-
Hogs differed in some respects from Ph. ethiopicus, although the
general shape of the skull (especially the postorbital portion) was
similar. The nasals of Ph. ethiopicus are ‘anteriorly rather
evenly convex, but form in their posterior portion behind the
foramina infraorbitalia a voof-like ridge or elevation”*. In
Lord Delamere’s Wart-Hogs the nasals are rather flat along their
whole extent, without forming any ridge posteriorly. The choane
are broad, much widened posteriorly in Lord Delamere’s Wart-
Hogs, but in Ph. ethiopicus they are not wider behind than in front
and the margins are parallel. The sphenoidal pits are completely
open, not covered by any bony roof, in Lord Delamere’s Wart- Hogs,
but the lateral walls formed by the pterygoids are high so that adeep
canal is formed. The distance from the hind margin of foramina
palatina to the hind margin of the palate measured in a straight
line is respectively 50 and 51 mm. in the two specimens of
Lord Delamere’s Wart-Hogs, but only 35 mm. in Ph. ethiopicus,
although the latter specimen is older and a little larger. As these
differences'are quite recognizable and more material, no doubt,
on direct comparison, will add other characteristics osteological
* Lonnberg, JU. c. p. 55.
940 MR. R. LYDEKKER ON [ Dec. 15,
as well as external, I think it will be correct to distinguish this
Wart-Hog, presumably from Somaliland, by a separate name,
and I venture to propose to call it Phacocherus delamerei.
T hope that sportsmen who visit Somaliland may have their
attention drawn to this animal and bring home satisfactory
material to fully elucidate this question.
As the Eurasian members of Suid are less specialized than
the African genera of the same family, it must be assumed that
the hogs originated on the Eurasian continent, the more so as
the oldest known fossil remains belonging to this family have
been found in Europe. ‘The genera of Suidze less specialized than
the Wart-Hog, even Potamocherus and Hylocherus, have the
postorbital portion of the skull comparatively much longer and the
interorbital region much narrower than the corresponding parts
of Phacocherus. Thus a comparatively long postorbital and a
narrow interorbital region of the skull in a Wart-Hog indicate
a less specialized race. It agrees well both with the geographical
distribution , and origin that the most northern races of
Phacocherus, viz. Ph. eliani in Abyssinia and Ph. africanus
Senegambia, are the least specialized. Ph. massaicus further
south has retained a rather Jong postorbital region of the skull,
but acquired a great interorbital breadth. The Wart-Hogs from
Lake Mweru and Nyassa, on the other hand, have still a narrow
forehead, but their postorbital portion is somewhat shortened,
while Ph. sundevallit has the forehead broader but the postorbital
portion not so much shortened. Finally, in the Cape region, the
most specialized of all Wart-Hogs, Ph. ethiopicus, with very short
postorbital portion and a comparatively broad forehead, is found.
It has also completely lost the upper incisors, while the
lower ones ave absent or rudimentary. Ph. delamerei, which at
present must be regarded as inhabiting Somaliland, has in
that country independently reached a similar stage of specializa-
tion as Ph. wethiopicus at the Cape. It is probable that this
analogy between the Wart-Hogs of the Cape and Somaliland
depends upon similar natural conditions of the two countries, and
if that be so there might no doubt be found other instances of
parallel development within the saine geographical areas.
4, On Two Chinese Serow-Skulls.
By R. LypEexKerr.
[ Received October 3, 1908. |
(Text-figures 191-192.)
When I described the immature specimen of the White-maned
Serow (Wemorhedus argyrochetes) of Sze-chuen in the Society's
‘Proceedings’ for 1905, vol. 11. p. 329, pl. viii., some doubt was
expressed at the meeting as to whether the animal was anything
more than a local race of the widespread Vemorhedus sumatrensis ;
1908. } TWO CHINESE SEROW-SKULLS. 941
and it has recently been relegated to that grade*. Iam now in
@ position to demonstrate its right to specific rank.
Text-fig. 191.
A B
Female skulls of Nemorheedus sumatrensis milne-edwardsi (A) and
N. argyrochetes (B), from Sze-chuen.
* See Proc. Zool. Soc. 1908, p. 185. I am afraid I cannot follow my friend
Mr. Pocock in transferring the name Nemorhodus to the Gorals; it has been too
long in use for the Serows.
949 MR. R. LYDEKKER ON [ Dec. 15,,
In September last Mr. J. W. Brooke* presented to the British
Museum (Nat. Hist.) the skins and skulls of an adult male and
female of the White-maned Serow obtained by himself and Mr. C.
H. Mears at Towquan, 60 miles north of Kanshieu, N.W. China.
That they belong to this species is evident from the fact that
the greater part of the fore-legs and the whole of the hind-
legs, inclusive of the lower portion of the thighs, are bright rusty
red. Both skins, as indicated by the teeth and horns of the
associated skulls, belong to fully adult animals; the horns being
much larger than those of the immature mounted specimen in the
British Museum. From that specimen, and also from one recently
figured by Mr. M. W. Lyon, the new skins differ, however, by
the absence of such a distinctly white mane as occurs in the
immature female.
Text-fig. 192.
A female of the Sze-chuen race of the Serow, Nemorhcedus sumatrensis milne-
edwardsi, from a photograph by Mr. J. W. Brooke.
Tn all the races of the ordinary Serow (WV. swmatrensis), as 1s
well shown in the figures illustrating the paper by Mr. Pocock
already mentioned, the skull is relatively broad and _ short,
with the nasal bones likewise proportionately broad and short,
although there is a considerable degree of local variation in this
respect. These characters are exemplified in a third skull (text-
fig. 191 A, p. 941) sent home by Mr. Brooke. I believe this to
belong to the same species as the Serow shown in the annexed
reproduction from a photograph (text-fig. 192), which is evidently
* [We much regret to note that since the reading of this communication news
of the murder of Mr. Brooke in China has reached England. |
1908. | TWO CHINESE SEROW-SKULLS. 943
one of the dark-coloured races of V. swmatrensis allied to the
Malay representatives of that species, the scalp-skin accompanying
the skull being of a dark type.
The skull and photograph indicate a Chinese representative of
the ordinary Serow, which is doubtless the one to which Pére
David gave the name WV. milne-edwardsi. The following
dimensions are taken from the skull :—
iixGreme basal leneth® 229,22 i. eee Daa oeikees 11 inches.
as zygomatic WCE Le" LH SUR EAS OE oe
err et irons sive: Fe Shs, VN, Bie seein
Width Olmeasallsne ie 2eaMey) AS Oe ee a aeE
This skull and scalp belong, it should be added, to a female.
Of a very different type are the male and female skulls from
‘Towquan, these (text-fig. 191, B, p. 941) being narrower and
longer with longer and narrower nasals. The dimensions of these
two skulls are as follows :—
bo — Ot
CO} OD|H RIA
Male. Female.
ectreme basalllemoth \..7....--....-.- 12 114 inches,
si zygomatic padioh See ee ) 43 es
Wemethvol masalser. sks: es scare - ae. 43 42,
An Coram AS Oa 13 DS ale
From this it will be evident that while in the White-maned
Serow the basal length and the nasal length exceed the corre-
sponding dimensions in the ordinary species, the width of the
nasals is less. A further important difference is to be found in
the basisphenoidal region, which is much narrower in the White-
maned than in the ordinary species; while the premaxille are
also longer and narrower in the former thay in the latter.
The following measurements of the two animals taken im-
mediately after death have been supplied by Mr. Brooke. In
taking the measurements the tape is stated to have been laid
flat on the body without pulling.
Male; shot March 30th, 1908.
Height at shoulder from back of foot 44”.
Nose to root of tail 68”.
Lip (upper) to horn 113".
Kar 82" (length), ear 7 (widest part).
Cir cHaten ence of hind-leg level with the point where the
loose skin from the belly joins the leg 23”.
Widest part of body (2. e. just behind withers and over
breast-bone under body) 453”.
Neck where it joins the body 37",
Narrowest part of body in front of hind-iegs 382”.
Female; shot April Ist, 1908.
Height at shoulder from back of foot 433”.
Nose to root of tail 613".
Bottom of upper lip to root of horn 13”.
Proc. Zoot. Soc.—1908, No. LX. 60
944 MR. R. I. POCOCK ON WARNING COLORATION __[ Dec. 15,
Length of ear 9”; widest part 73”.
Hind-leg level with the point where the loose skin joins the
belly (as before) 18”.
Widest part of body (as before) 46”.
Narrowest part of body in front of hind-legs 383".
These dimensions indicate animals rather larger than Sumatran
Serows; and in some degree justify Mr. Henry’s statement as to
the Chinese animal beingas large as a cow.
In my opinion, Mr. Brooke’s specimens fully justify the re-
cognition ot Vemorhedus argyrochetes as a valid species, especially
as it appears to inhabit the same district as WV. milne-edwardsi.
The ears of the White-maned species appear to be rather larger
than those of the other.
Mr. Brooke mentions that the White-maned Serow is known to
the natives as ‘‘ Nikka” and the dark species as ‘¢ Nik-lu.”
The Sze-chuen race of the true Serow has the back black mingled
with white; the front of the fore-legs is black to the knees,
velow which the limb is grey with patches of rusty; the hips and
posterior surfaces of the hind-limbs are rufous, the black on the
front surface extending some distance short of the hocks. These
particulars are taken from a mounted skin presented by Mr. Brooke
to the Museum.
I may add that I have given a preliminary notice of the
specimens forming the subject of this paper in the ‘ Field’ for
October 8th, 1908.
5. Warning Coloration in the Musteline Carnivora*. By
R. I. Pococx, F.L.8., F.Z.S., Superintendent of the
Zoological Society's Gardens.
[Received December 15, 1908. |
(Text-figures 193-198.)
As long ago as 1846, Hamilton Smith wrote: “The Ratels offer
one more instance of the colours of the fur being light on the
upper surface of the body and dark beneath, producing a kind of
family livery, alike m this and the Grisons, Zaxidea and Meles,
and not obliterated in Hira [Galera] and Arctonyx.”+ The
circumstance in fact is so well known that it would be profitless
to search literature for earlier and even later records. Some
later authors indeed have drawn attention to the style of colora-
tion above described as being uncommon and as a “ divergence
* Under the term Musteline I include in this paper both Weasel-like and
Badger-like Carnivora, which are generally referred to two distinct subfamilies
Mustelmae and Meline. I have not aimedat making the list of nauseous species
complete; but have based my conclusions in the main upon those which have come, as
living animals, directly under my own observation. ;
y Jardine’s Nat. Library, xv. Mammalia, pp. 205-206.
1908. } IN THE MUSTELINE CARNIVORA. 945
from the usual rule” *; and Mr. Lydekker observes: “ It is also
noteworthy that in the parti-coloured examples [of the Mustelidee]
there is a great tendency for the underparts of the body to be
darker than the upper; whereas, it is scarcely necessary to
observe, the reverse is the case in the great majority of
mammals.” +
‘Before the publication of Mr. Thayer’s paper explaining the
celative or procryptic significance of the usual style of coloration
whereby reflected lights are toned down and shadows obliterated,
it was hardly to be expected that any special inquiry would be
made as to the meaning of the peculiar livery of the Mustelines
in question; but, so far as [am aware, no suggestion has been
made on this head since the publication of that luminous idea ¢.
Yet the inference seems obvious enough that, since the colours
are reversed, their functions must also be reversed; that is to
say, if animals which are light below and dark above are concealed
on this account under a top light in their normal surroundings,
those which are light above and dark below should be made con-
spicuous under the same conditions. White on the upper side
should have the effect of enhancing reflected light, and dark on
the under side the effect of emphasising shadows.
A simple experiment demonstrates this to be a fact. If a cork
be pinned with a long pin against a sheet of brown paper of its
own colour under a top light, it may be made practically invisible,
as Thayer has shown, by painting its upper side dark and its
under side white. But if the cork be then turned over so that
its white side be uppermost and its dark side undermost, its
maximum of conspicuousness is achieved. The effect of turning
it over is much the same as that produced by immensely increas-
ing the intensity of the top light over the uncoloured cork.
In the case of Mammalia, it is exceptional for the coloration to
be of a kind that makes for conspicuousness. In the majority of
instances it is procryptic for the purpose of enabling the indi-
vidual either to escape enemies or to secure prey. Hence, if it
be claimed that the livery of these Mustelines belongs, as I think,
to the former category, it is necessary to produce in favour of the
claim evidence drawn from the bionomics of the species in addi-
tion to that deducible from the above mentioned fact that the
coloration is the very opposite of that exhibited by a very large
number of procryptically coloured forms. Sufficient evidence to
justify the adoption of this view as a useful working hypothesis,
is, in my opinion, supplied by what is known of the habits of the
species discussed in the following pages.
_ With the exception of mimetic species, animals which are
coloured so as to be conspicuous in their natural surroundings
are very often protected from enemies by distastefulness arising
from a nauseating flavour or odour, or by the possession of poison-
* J. G. Wood, ‘ Illustrated Nat. History,’ Mammalia, p. 372, 1861.
+ Royal Natural History, 11. p. 47, 1894.
t ‘The Auk,’ xiii. 1896, pp. 124 & 318.
60*
946 MR. R. I. POCOCK ON WARNING COLORATION __[ Dec. 15,
glands and stings which make them dangerous to meddle with.
They also as a very general rule have no need of procryptic color-
ation to enable them to capture wary or keen-sensed prey. Their
movements are usually slow and deliberate, and instead of avoiding
they seem rather to court observation, some indeed attracting
attention by the emission of characteristic sounds. Very com-
monly also they are hard, tough, and difficult to kill.
Poreupines of the genus Hystriz furnish a good instance of
this. Protected by their spine-armature, they are quite con-
spicuous in the dusk by reason of the predominance of white on
the dorsal surface, and they make themselves heard by shaking
their caudal rattles and uttering hoarse grunts. This I pointed
out last year (see P. Z.8. 1906, p. 902, pubd. April 1907). Sub-
sequently I noticed that the Canadian Tree- Porcupine (Hrithizon),
which which has no rattle, but is conspicuously coloured when
its spines are erected, possesses a strong and unpleasant odour
recalling that of concentrated human perspiration. This is also
very possibly one of the aposematic attributes of the species; and
I have recently come across a passage showing that exactly the
same discovery was made about forty years ago by that keen
naturalist Charles Kingsley in connection with the Brazilian Tree-
Porcupine or Coendoo (Coendu). He wrote: “ More than once we
became aware of a keen and dreadful scent, as of a concentrated
essence of unwashed tropic humanity, which proceeded from that
strange animal, the Porcupine with a prehensile tail, who prowls
in the tree-tops all night, and sleeps in them all day, spending
his idle hours in making this hideous smell. Probably he
or his ancestors have found. it pay as a protection; for no
Jaguar or Tiger-cat, it is to be presumed, would care to meddle
with any thing so exquisitely nasty, especially when it is all over
sharp prickles.” * It is interesting that the same comparison
should have been independently employed both by Kingsley and
myself in attempting to describe the scent of these Porcupines ;
and that he should have anticipated me by so many years in
assigning a protective value to it.
Up to the present time the only Mammals, apart from Porcu-
pines, claimed to be warningly coloured, so far as I am aware, are
the Skunks of America (Mephitis, Conepatus, Spilogale) and the
Zorillas of Africa belonging to the genus Jetonya and known in
Cape Colony as Cape Polecats. These are black Mustelines orna-
mented dorsally, as a rule, with broad clear white longitudinal
stripes, which sometimes coalesce or almost coalesce to form a
continuous white field. When attacked they increase their
apparent size and enhance their conspicuousness by erecting the
long hairs of their bodies and by brandishing their bushy white
tails. At the same time they eject from their anal glands a
volatile fluid, with a most repulsive, acrid and persistent odour,
Skunks in captivity are frequently quiet undemonstrative animals;
* “At Last,’ p. 248, ed. 3, 1805; quoted also by J. G. Wocd in Waterton’s
Wanderings in South America,’ p. 458, MacMillan & Co., 1879.
1908. | IN THE MUSTELINE CARNIVORA. 947
but I have seen a Cape Polecat behave in the way described
above at the sight of a small dog, uttering the while shrill squeals
of anger. Both Skunks and Cape Poiecats are said to feed natur-
ally upon any small terrestrial vertebrates they can catch, and
also upon insects. Merriam indeed describes the North-American
Skunk as preeminently an insect-eater, adding that it ‘ destroys
more beetles, grasshoppers, and the like, than all our other
mammals together,” also “‘ he devours vast numbers of mice.”
But those that have come under my observation in the Zoological
Gardens will eat fruits like bananas and dates. Hence they are
in all probability omnivorous in their native haunts; and are,
therefore, not dependent for food upon the live things they
capture.
Text-fig. 193.
Yo ‘ “troennastey oo
Cape Zorilla (Ietonya: capensis), left-hand figure, and Cape Weasel (Pecilogale
albinucha), right-hand figure.
I have been able to demonstrate experimentally and to my
complete satisfaction, that the white on Skunks and Zorillas
makes them conspicuous at night, whether it be cloudy or star-
lit, against the dark background of the soil or of low herbage.
I made the experiment with three stuffed skins, one of the
Canadian Skunk (Mephitis mephitica), one of the Cape Zorilla
(Ictonyx capensis), and one of the Libyan Zorilla (Jctonyx
libyca). The Skunk was not a good specimen, having died in bad
coat with short hair, yet I could see it at a distance of 15 feet.
The others were easily visible at twice that distance * the
Libyan Zorilla being more visible than the Cape specimen on
account of the greater amount of white on its dorsal area. The
conspicuousness of all three was enhanced when they were
made to move. On the other hand, when put upon the snow
* They would clearly be visible at a much greater distance to, Carnivora with
nocturnal vision.
948 MR. R. I. POCOCK ON WARNING COLORATION __ [ Dec. 15,
all are visible, the Skunk being the most conspicuous of the
three on account of the greater amount of black in its pelage ;
and the Libyan Zorilla the least conspicuous for the opposite
reason. The interest of this fact lies in the circumstance that
the species of Skunk mentioned above frequently has to traverse
snow-covered ground; whereas the Libyan Zorila probably
never has to do so. By twilight and daylight the specimens
were conspicuous both in vegetation, on the grass, and on
snow-covered ground.
Another Weasel which presents a very unusual type of colora-
tion is the genus Pecilogale of tropical and Southern Africa, the
typical form of which is P. albinucha of Cape Colony. The livery
is of the same style as that of /ctonyx, the body being black with
four snow-white stripes along the back. On the shoulders these
coalesce into two stripes which fuse with a large white patch
covering the fore part of the nape of the neck and the top of the
head. The tail is white. The resemblance between this animal
and Jctonyx may be an instance of true (Batesian) mimicry as
Mr, Lydekker has suggested *. On the other hand, if Pecilogale
is itself protected by an exaggerated development of the sub-
caudal stink-glands such as are found in the common weasel,
stoat, and polecat,it may be that the similarity in question is an
illustration of Miillerian resemblance. Very little seems to be
known of the habits of this rare animal, but its long and lithe
form which is typically weasel-like, suggests that it resembles in
mode of life the weasel and the stoat, to which it is more nearly
related than to the zorillas. If this be so, it would seem that
the peculiar style of coloration, so unlike the protective coloration
of weasels and stoats, must have either a true warning (apose-
matic) or a false warning (pseudaposematic) significance.
Another member of this family which I have no doubt is
warningly coloured is the Teledu (J/ydaus) of Indo-Malaya.
The general colour is blackish brown, but a white band
commonly extends from the top of the head down the spine to
the tail, the tip of which is also white. On the back of the head
and neck the area of white is increased by the hairs forming a
decided erectile crest. This animal, like Skunks and Cape
Polecats, is nocturnal. It is slow in its movements and feeds to
a great extent upon insects and worms. It also possesses stink-
glands, which exude a fetid volatile liquid. According to Hors-
field, ‘the entire neighbourhood of a village is infected by the
odour of an irritated Teledu, and in the immediate vicinity of the
discharge it is so violent as in some persons to produce syncope,”
as has been stated to be the case with the discharge of the Skunk. -
Mr, Shortridge, who has collected these animals in Java, tells me
that he believes they feed upon roots; and he has noticed the
night air tainted by their smell. Ialso owe to Mr. Shortridge
the suggestion that in Java at all events the Teledu is mimicked
* Royal Natural History, ii. p. 70, 1894.
1908. ] IN THE MUSTELINE CARNIVORA. 949
by the Ferret-Badger (Helictis orientalis). There is unquestion-
ably a close superficial resemblance between them in size, form,
and colour, although the white on the nape and shoulders of
Helictis is less extensive and there is more white on the face than
in Mydaus. Helictis, however, may be itself a protected form,
and in this case the resemblance between it and Mydaus is
probably an instance of common warning coloration usually called
Miillerian mimicry.
Text-fig. 194,
Teledu (Mydaus javanensis), upper figure, and Ferret-Badger (Helictis
orientalis), lower figure.
In the genera of Mustelide above described the coloration is
very specialised, consisting in the Skunks and Zorillas of alternat-
ing black and white bands, and in the Teledus and Ferret-Badgers
of a single white band running down the back. There are some
Skunks, however, in which the entire dorsal area is white, as if
the white stripes had extended towards the middle lne and
coalesced. It is quite possible, however, that the uniform white-
ness of the back is the more primitive livery of the two, and
that the ancestral Skunk was grey-backed, like a Ratel, later
forms becoming white-backed by the gradual whitening of the
whole dorsal area, or striped by the sorting of the hairs into black
and white bands.
One of the best-known examples of the style of coloration
mentioned above, in which the upper side is markedly lighter
than the under, is the Ratel (Jellivora), which is represented by
species or subspecies in India, Arabia, and Africa. The back and
head, sometimes white, as a rule are iron-grey, the muzzle, legs,
and under side being jet-black. Where the grey or white of the
dorsal surface meets the black of the under surface, the contrast
between the two is emphasised by a whiter line which is very
950 MR. R. I. POCOCK ON WARNING CoLORATION __[ Dec. 15,
conspicuous both on the forehead and the sides of the body.
One form only is black above as well as below.
There is abundant testimony to the unpleasantness of the odour
emitted by Ratels. Writing of the Cape Ratel, Mr. W. L. Sclater
says: “It further defends itself by emitting an offensive odour
from its anal glands.” * In his account of the habits of the
Indian species, Blanford does not mention this attribute; but I
have been told by Indian sportsmen that the characteristic is
well known, and two of the Society’s keepers, Dixon and Hoare,
who have looked after a male specimen that is still living in the
Gardens, tell me that formerly this animal, when threatened or
disturbed, would emit an odour, described as suffocating, which
could be perceived ata distance, varying according to the estimate
of the raconteur, from fifty to one hundred yards.
Text-fig. 195.
White-backed Ratel (Mellivora ratel).
Ratels are omnivorous, and can be kept in confinement in
health and strength without meat. Their liking for honey is
notorious, and is the attribute from which their generic name has.
been derived. They are known to be desperate fighters and extra-
ordinarily tenacious of life. The skin is not only very thick, but
also very loose, so that if seized by almost any part of it the animal
can reach and biteitsassailant. Ofthe African species Mr. Sclater
says: “It is very difficult to kill, only, it is said, by actually
crushing its skull or by stabbing to the heart can this be
effected.” The very small size of the ears in the Ratel is another |
noticeable feature bearing on the question of his immunity from
attack. Animals which require sharp hearing either to escape
enemies or capture prey usually at all events have large ears;
and the fact that the animals forming the subject matter of this.
* “The Mammals of South Africa,’ i. p. 112, 1900.
1908. ] IN THE MUSTELINE CARNIVORA. 951
paper have small external ears is in keeping with the theory that,
they have no enemies to fear.
Caged Ratels do not always make use of their scent-glands.
For example, two specimens of J/. capensis in the Society’s
Gardens never did so; and the keeper in charge, noticing this
difference in behaviour between them and the specimen of
A. indica above mentioned, asked me if it was a specific feature.
The explanation no doubt is that when once tamed these animals
soon learn that they are safe from enemies, and therefore do not
resort to this special mode of defence. ;
The coloration of the Grison (Grison furax=CGalictis vittata),
a South-American musteline, is very similar to that of the
Ratel (Mellivora). The whole of the upper side of the head and
body is greyish, the under side of the body and head and the
limbs being black. Across the forehead and along each side of
the head towards the shoulder at the junction of the grey and
black, there runs a whitish band which is very conspicuous as the
animal advances.
I have the independent testimony of two of the Society’s
keepers, Dixon and Heffer, that when Grisons fight or are
disturbed they stink like Skunks and Cape Polecats (Ictonyx) ;
and J. G. Wood*, writing apparently of his own knowledge,
says: “‘ The odour which proceeds from the scent-glands of the
Grison is peculiarly disgusting, and offends human nostrils even
more than that of the stoat and polecat.” The Grison also has
the reputation of being extremely savage and a most dangerous
foe to any animal it ventures to attack. It was of this species,
and probably the next, that Mr. W. H. Hudson wrote “... .-
there are[on the pampas of La Plata] two quaint-looking weasels,
intensely black in colour, and grey on the back and flat crown.
One, the Grison furax, is a large bold animal that hunts in com-
panies ; and when these long-bodied creatures sit up erect, glaring
with beady eyes, grinning and chattering at the passer by, they
look like little friars in black robes and grey cowls; but the
expression on their round faces is malignant and bloodthirsty
beyond anything in nature, and it would perhaps be more decent to
liken them to devils rather than to humans”; and again: ‘ After
watching the weasels dance for some minutes, I stepped up to the
mound, whereupon the animals became alarmed and rushed pell-
mell into the burrows, but only to reappear in a few seconds,
thrusting up their long ebony-black necks and flat grey-capped
heads, snarling and chattering at me, glaring with fierce beady
eyes.” The same author bears testimony to the absence of the
hiding instinct in the young of this species. He says: “I once
surprised a weasel [Grison furax] in the act of removing her
young, or conducting them, rather ; and when she was forced to
quit them, although still keeping close by, and uttering the most
piercing cries of anger and solicitude, the young continued
* Tilustrated Nat. History, i. p. 372.
952 MR. R. I, POCOCK ON WARNING COLORATION ___[ Dec. 15,
piteously crying out in their shrill voices and moving about in
circles, without making the slightest attempt to escape, or to
conceal themselves, as young birds do.”* These passages attest
the savage aspect and fearless behaviour of the Grison and the
suppression of the instinct to hide in young individuals. These
are precisely the attributes one would expect, if the species is
specially protected and warningly coloured.
Text-fig. 196.
ov
Wrras raat: ag
Grison (Grison furax), upper figure, and Patagonian Weasel (Lyncodon
patagonicus), lower figure.
As is the case with the Ratel and some other mustelines, the
* <The Naturalist in La Plata, pp. 15-16, 104, and 385-386, 4th ed. 1903. In
the paragraphs above quoted I have substituted the name Grison furax for
Galictis barbara. My. Hudson’s description of the larger animal, apart from his
remark that it is “ about the size of a cat,” coupled with my own knowledge of the
geographical distribution of Galera barbara and of Grison furax, convinced me
that he had applied the wrong specific name to the larger La Plata musteline. I
therefore wrote to him on the matter, and he kindly confirmed this, adding that he
was misled by a wrong label in the Buenos Ayres Museum and had discovered the
mistake subsequently. It is important that the error should be corrected, because
although obvious enough to those who know the two species under discussion, it has
already made its way into the literature of natural history. In the ‘ Royal Natural
History,’ for example, the larger of the two weasels mentioned by Mr. Hudson is
cited as the Tayra (Galera barbara), and the smaller as the Grison (Grison furax) :
whereas the larger is, as stated, the Grison, and the smaller, I suspect, the Pata-
gonian Weasel (Lyncodon patagonicus).
1908. ] IN THE MUSTELINE CARNIVORA. 953
diet of Grisons is mixed. Two now living in the Zoological
Gardens feed upon fowls’ heads, dried dates, and bread and milk.
The so-called Patagonian Weasel (Lyncodon patagonicus),
although smaller than the Grison, presents much the same style
of coloration. The hair of the body and tail is long and grey.
On the nape of the neck there is a large black patch which
emphasises a large white patch covering the top and sides of the
head and extending laterally along the neck. This is set off both
in front and below by the black colour of the muzzle, cheeks,
sides of the neck, and lower shoulder. The legs are black. Very
little seems te be knowa of the habits of this animal. It may or
may not be offensive like the Grison. If it is, its coloration is, I
think, probably genuinely aposematic. If it is not, the resem-
blance in colour between the two may be an instance of true or
Batesian mimicry. The evidence, however, that the two species
are found together is not conclusive. In favour of this view it
may be added that ZLyncodon has been recorded in Central
Argentina from Mendoza and Azul southwards to the Rio Colorado
and Rio Negro, and Grison also as far south as the Rio Colorado * ;
and according to Trouessart’s Catalogue both occur in Northern
Patagonia.
Bat exact coincidence in distribution is not essential to the
belief that the resemblance between the two animals is an instance
of Batesian mimicry or of Miillerian resemblance. It is merely
essential to show that enemies that might prey upon small
carnivora of this kind are dispersed over the areas inhabited by
the two forms. Wide ranging raptorial birds, for example, that
knew the Grison by sight in the northern parts of La Plata,
might easily mistake the Patagonian Weasel for the young of it
in the southern parts of that country.
An interesting parallel is traceable between Lyncodon and
Grison in South America, on the one hand, and Pecilogale and
Ictonyx in Africa, on the other. In both cases, we have a large
musteline which is known to have offensive stink-glands, and a
smaller one in which this attribute has not yet been recorded.
There are reasons for thinking that in both the larger species
the coloration, though widely different, is aposematic; and the
smaller form in each case resembles the coloration of its com-
patriot. The smaller forms also appear to be much scarcer than
the larger, a fact which is in favour of their coloration being
mimetic.
Singularly enough, too, Lyncodon and Pecilogale resemble each
other and differ from typical mustelines, including their hypo-
thetical models, in the reduction of the number of cheek-teeth
to three on each side in both the upper and the lower jaws, the
total number of teeth being 28. In both J/ctonyx and Grison,
on the contrary, there are four cheek-teeth in the upper jaw and
five in the lower, making a total of 34.
* Matschie, SB. Ges. nat. Fr. Berlin, 1895, p. 190.
954 MR. R. I. POCOCK ON WARNING COLORATION [ Dec. 15,
Allied to the Grison is another South American musteline, the
Tayra (Galera barbara). Itisa larger animal than the Grison
and approaches an otter in size. I cannot find in literature any
convincing evidence that this animal stinks to the same nauseating
extent as the Grison; but considering the close relationship.
between the two species, which until lately were referred to the
same genus, this must be regarded as probable. Hamilton Smith,
however, remarks that it ‘‘has a strong musky smell”*. One
that lived a few years ago in the Zoological Gardens was extra-
ordinarily tame, and Dixon, the keeper in charge of it, tells me
that it never smelt like the Grisons or Ratels, but only “like a
Badger.” This negative evidence as to its potential offensiveness
Text-fig. 197.
Bush-Dog (Speothos venaticus), upper figure, and Tayra (Galera
to} Pp ) o ) ‘J
barbara), lower figure.
must not, however, be overvalued ; for, as has been explained,
two tame specimens of the African Ratel that have recently lived
in the Gardens were never known to make use of their stink-
glands. The same is true of some Canadian Skunks (Jephitis
mephitica) we now possess, although their glands are entire; and
it is well known that Cape Polecats (Icton yx capensis) can. be
tamed and kept in houses as pets for destroying rats.
The colour of the Tayra varies, some specimens, perhaps always
* Jardine’s Nat. Library, xv. Mammalia, p. 202, 1868.
1908. ] IN THE MUSTELINE CARNIVORA. 955
young ones, being white, and others black. As a rule, however,
they are foe brown or blackish with the head and neck grey,
and there seems to be nearly always a conspicuous large yellow
patch upon the chest. They eat a variety of food, and one that
lived some years in the Zoological Gardens was fed upon dates,
bananas, figs, and a little cooked meat. That the Tayra has in
a wild state the same savage disposition when attacking or attacked
as other mustelines, hardly admits of a doubt; but very little
appears to have been recorded of its habits. i
Some years ago Mr. O. Thomas pointed out to me the
obvious resemblance between Galera barbara and the South-
American Bush-dog, Speothos venaticus ; and it occurred to me
that it might be mimetic. But I do not at present know enough
of the bionomics of the two species, to feel justified in doing more
than put forward this view as a theory for future confirmation or
refutation. In its favour it may be urged that there are no
reasons for supposing that the dog is protected in any way from
larger carnivora, and it is apparently much rarer than Galera.
A well-known European carnivore with much the same style
of body-coloration as the Grison is the Badger (Ieles meles),
which is hoary grey above and black below and on the legs. The
coloration of ihe head, however, 1s very different from that of the
Ratel, Grison, or Tayra, for it is white with a broad black band
extending on each side from the muzzle across the eye to the ear,
which is itself white-rimmed ; and the chin and throaé are black.
Badgers are slow and leisurely in their movements, and have
earned a reputation for stupidity by the fearlessness and indiffer-
ence of their manner towards things in general. Their diet is
mixed, but they subsist to a very great esteae upon vegetable food.
In no senseare they dependent: for a livelihood, so far as is known,
upon the capture of wary mammals or birds. When attacked,
they are notoriously most savage and formidable antagonists,
being gifted with exceptionally strong jaws, a thick, highly flexible
and loose skin, and wonderful tenacity of life. They also possess
stink-glands which exude a powerful and unpleasant odour.
The scent of the secretion has given rise to the epithet ‘stinking
brock,’ and forms the basis of the well-known simile ‘smells
like a badger.’
At dusk, when badgers emerge to feed, they are rendered con-
spicuous by the whiteness of the head; and looking into our
badger’s cage in the Gardens in the evening, I have pecan been
eck by the ease with which the whereabouts of the animal
could be detected, especially when on the move, by the whiteness
of this region.
Other species of J/eles and the Indian Sand- Badger (Arctonya
collaris) seem to agree with the European Badger in all respects
essential to the present argument in the matter of coloration and
mode of life; and the same I suspect is true of the American
form, Taxidea americana.
A yery unusual style of coloration is also presented by the
956 MR. R. I. POCOCK ON WARNING COLORATION —__[ Dec. 15,
Sarmatian, mottled or marbled Polecat (Putorius sarmaticus).
The whole of the upper side of the body is brown variegated with
yellowish-white spots and patches, which on the sides of the neck,
belly and thighs tend to run into longitudinal stripes, offering
« sharp contrast with the jet-black hue of the throat, legs, and
the rest of the under side. The tail is long, bushy, al largely
white. The head is mostly black, but the lips and chin are
white ; a broad white band crosses from beneath the ears over the
forehead, and the distal half of the ears is white. Blanford
remarks of this species, which is found in Eastern Europe and
Western Asia, that it has “the same disagreeable foetid odour that
Text-fig. 198.
Sreorenar <=
English Badger (Meles meles).
y
is characteristic of the common [ European] Polecat,. . . . which
is particularly distinguished amongst the weasel tribe for the evil
odour generated by the secretion a its anal glands, whence its
name of foumart or foul martin.” He also says that it feeds
on birds, rats, mice, lizards, senle® and snails. The coloration
of this animal is so different from ‘that of the ordinary weasels,
and conforms in a general way so closely with that of some other
fetid members of that tribe, the pattern of the head being
especially like that of the Libyan Zorilla (/etonyx libycus), that I
cannot help thinking it has a warning significance. Very little,
however, seems to be known of this species in its native haunts,
* Fauna of Brit. India, Mammalia, pp. 163 & 165, 1888.
1908. | IN THE MUSTELINE CARNIVORA, 957
the information quoted above from Blanford having been taken
from Hutton’s account based upon observations of living examples
in captivity.
Reference was made above to a black form of Ratel. This
occurs in the Ituri Forest and was described by Mr. Lydekker as
Mellivora cottoni*. This species, or race, has a special interest
in connection with the views advocated in this essay, from its
bearing upon the theory I have already published 7, that where
concealment is unneeded animals tend to assume a uniformly
dark coloration unrelieved by spots or stripes. As illustrations of
this were cited elephants, rhinoceroses, hippopotamusest, buffalos,
bisons, many bears, moles, ravens, rooks, and others, which, either
by their strength and size, their gregarious habits or mode of
life, are protected from carnivorous enemies and have no need of
procryptic coloration to help them in the capture of prey.
In thei habits, many Bears are very similar to badgers and
ratels. They are slow and leisurely and bold in their movements,
and feed chiefly upon roots, fruits, honey, and other vegetable
products, although they will kill and eat living prey. They are
not, however, dependent upon it §, as are the Cats, to which they
offer the greatest possible contrast both in coloration and mode
of life. They have no stink-glands like the Mustelide described
above, but are well known to be terrible antagonists when fighting.
Major Rodon, F.Z.8., has told me that the Himalayan Black
Bear (Ursus torquatus) is more than a match for leopards, and
that he has known one drive a leopard from its kill. Now this
species of Bear has a very distinct, somewhat V-shaped white or
yellow patch across the chest, which is displayed to full view
when the animal stands erect. The Malayan Bear (Ursus malay-
anus) has a similar, usually yellowish, horseshoe-shaped mark ; and
the Sloth Bear (Welursus ursinus) carries the same badge. It is
significant that this white mark is shown to an antagonist when
the bear assumes its attitude of defence, and it reminds one
forcibly of the patch described above possessed by the Tayra
(Galera barbara); and I venture to suggest that, as in that animal,
it acts as a recognition mark and danger signal.
Since the preceding pages were written, I have had the oppor-
tunity of discussing with Mr. Abbott H. Thayer the theory of
warning coloration in general and its application to Mammalia
in particular. Mr. Thayer has already published || his disbelief
* P. Z. 8. 1906, p. 112.
+ Pall Mall Magazine, Feb. 1904, pp. 179-180. |
{ It is interesting to record that three young hippopotamuses, one from Nigeria
and two from German Hast Africa, when brought to the Gardens, were pink below
aud protectively countershaded on Thayer’s principle. They were believed to be
about two years old at the time. During the two subsequent years, as they grew in
size and capability, the under side gradually became pigmented.
§ With exception of the Polar Bears, all the bears in our Gardens thrive on a diet
of ship’s biscuits and upon the bread and buns given to them by visitors.
ij Trans. Ent. Soc. London, 1903, p. 556.
958 MR. R.-I. POCOCK ON WARNING COLORATION [Dec. 15,
in warning coloration and his belief that the patterns of nauseous
species, to the conspicuousness of which in their natural sur-
roundings several observers have testified, are procryptic. Prof.
Poulton has briefly replied to this view so far as butterflies are
concerned *. I will here endeavour to do the same as regards
the Mammalia.
As stated in a demonstration given at the Gardens and sub-
sequently in conversation with me, Mr. Thayer holds that the
white markings of the Skunk, Badger, Ratel, Teledu, and Grison
gerve to conceal these animals from the ground-prey upon which
they feed. The head of a Badger or Ratel, for instance, would
lose its shape when looked at from below, because the white
tracts would be cut out against the sky; and this obliteration of
identity would be beneficial to the carnivore by enabling him to
capture field-mice and other ground-living species. Justification
for this hypothesis is found in the demonstrable fact that white
spots and patches appear as sky-holes, especially in foliage, when
viewed from a lower level; and it may be granted that the mark-
ings on the mustelines mentioned above may have the significance
claimed for them by Mr. Thayer when they are visible from
beneath. But I cannot bring myself to believe that his expla-
nation supplies the key to the guiding factor in their evolution.
Take, for example, the Teledu, the food of which is said to consist
of insects, larve, and worms. It cannot be seriously claimed
that the Teledu is helped in getting food of this nature by the
whiteness of the top of the head and neck, because worms are
blind, whilst nocturnal ground-insects at best have feeble powers
of vision. Even if the Teledu feeds also upon mice and other
vertebrates with vision something lke our own, which must be
admitted as a possibility, it is not very obvious how the narrow
median white spinal stripe can be of any procryptic use in the
way claimed. It would for the most part be invisible to the little
animals. It would on the contrary be in full view to an enemy
of larger dimensions than the Teledu, especially to one lurking in
a tree and looking down upon the musteline passing beneath.
So, too, with the Badger. This animal, as has been stated, lives
for the most part upon vegetable food, and itis difficult to believe
that the catching of mice can have had a survival value in the
history of any individuals of sufficient importance to the species
to guide the evolution of its facial coloration. On the other
hand, it is a demonstrable fact that Badgers grubbing or trotting
slowly about in the dusk, as is their wont, are quite conspicuous
to human eyes at a distance that could be covered by a wolf’s or
lynx’s spring, simply in virtue of the black and white bandings
on the head. It is quite easy to believe, however, that this same
pattern must be procryptic against a suitable background of
white rocks with black interstices or of foliage with light shafts
breaking through, especially if the animal be still; and it is quite
* “Assays in Evolution,’ 1908, p. 321.
1908. ] IN THE MUSTELINE CARNIVORA. 959
evident that the two explanations are not mutually destructive ;
but if we have to make a choice between the two, that is to say,
between the theory that the coloration of the mustelines mentioned
in this paper is aposematic or that it is procryptic, I think the
balance of evidence is in favour of the former. We have no
experimental proof of either so far as these particular species are
concerned. We do not know that any Badgers, or Ratels, or
Teledus have escaped destruction by their peculiar coloration ; but
we are justified in inferring its usefulness to the survival of the
species if it has had that effect. So, too, we do not know thet
any Badgers or Ratels or Teledus have succeeded in capturing
living prey in virtue of their peculiar coloration; but if they
have done so we are justified in inferring the comparative use-
lessness of the occurrences to the survival of the species, because
these mustelines feed mostly upon food which is insensible to the
patterns in question. The fearlessness, fierceness and tenacity
of life of these carnivora must also be reckoned with in this
connection.
The theory of warning coloration is intimately connected with
that of warning sounds. And it isa fact that many poisonous
animals like snakes, scorpions, and very large spiders stridulate
or rattle, or produce other sounds, when on the defensive or when
frightened. It is believed that they advertise themselves by this
means and warn their enemies to let them alone. If this be so,.
and no other explanation has been offered of the fact, we are
justified in inferring that advertisement that appeals to the ear is
useful to these specially protected species. The bearing of this
argument on the likelihood of the occurrence of advertisement
that appeals to the eye, is quite obvious,
I will only mention one more fact completely in accord with
the aposematic as opposed to the procryptic significance of the
coloration of the mammals discussed in this paper. A frightened
Skunk or Zorilla with long black and white hairs on end presents
exactly the same style of coloration as a common terrestrial
Porcupine with black and white quills erected. No one can
maintain that the coloration of Porcupines is procryptic for
the purpose of capturing prey, because they do not feed upon
living animals. And no one who has seen them in the dusk, can
dispute that the whiteness of the quills makes them conspicuous.
In addition to this they rattle and grunt and stamp, and appear
to advertise themselves in all ways at their disposal, and are
extremely unpleasant animals to deal with. There seems to me
to be no escape from the conclusion that their coloration is apo-
sematic. If so, that of Skunks and Zorillas, with the same style
of coloration, and equal though different unpleasantness, is also
probably aposematic; and from Skunks and Zorillas we pass to:
Badgers, Teledus, Ratels, and Grisons, all of which are more
or less patterned with white and all gifted with an unpleasant
odour.
Proc. Zoou. Soc.—1908, No. LXI. 61
960 DR. W. T. CALMAN ON A NEW [ Dec. 15,
6. On a new River-Crab of the Genus Gecarcinucus from
New Guinea. By W.T. Catan, D.Sc., F.Z.8., British
Museum (Natural History).
[Received November 8, 1908. ]
(Plate LX VIIL*)
The genus Gecarcinucus was established in 1844 by H. Milne-
Edwards t for a species, G. jacguemontii, which occurs in various
localities in the neighbourhood of Bombay. Milne-Edwards
referred the genus to the family Gecarcinide, and in this he was
followed by Dana, Wood-Mason §, and Ortmann |. Alcock 4,
however, has pointed out that, in spite of its great resemblance
to the Land-Crabs, the proper place of the genus is with the
River-Crabs (Potamonide) and it is included by Miss Rathbun **
in her monograph of the latter family.
Sir William Ingram, B.A., F.Z.S., has recently presented to the
Natural History Museum several specimens of a new Crab from
New Guinea which I refer to the same genus and in which the
Geearcinoid facies is even more strongly marked than in the
Indian species.
GECARCINUCUS INGRAMI, sp. 0.
Carapace about three-fourths as long as broad, very deep, very
convex antero-posteriorly, less so from side to side. Cervical
groove strongly marked, its lateral limbs nearly longitudinal,
becoming obscure just before reaching the antero-lateral margin.
Cardiac region partly defined posteriorly. A transverse groove
behind each branchial region. Mesogastric groove short, deep,
not forked. Branchial regions strongly convex, the gastric and
cardiac less so. Epigastric lobes prominent and rounded, separated
from front by a transverse groove which runs behind the orbits;
surface smooth, very faintly rugose near lateral margins.
Front generally a little less than one-fourth of width of carapace,
nearly vertically deflexed, its margin smooth. Seen from above its
outline is concave; from in front, its lower edge is slightly convex
and its lateral margins convergent; in the middle it is strongly bent
inwards, touching the epistome between the antennular fosse.
Orbits not entirely raised above lateral margin of carapace, roughly
quadrilateral, wider than high. Seen from in front, they are
inclined downwards and outwards, but are not produced at the
outer. corner into a gutter-like sinus as in G. jacquemontit.
Outer orbital angle hardly dentiform as seen from above; a
* For explanation of the Plate see p. 963.
+. Voyage dans l’Inde, par V. Jacquemont, Zool. Crustacés, p. 4, pl. 1., 1844.
t U.S. Expl. Exp., Crust. 1. p. 375, 1852.
§ Jour. Asiat. Soc. Bengal, xl. pt. 2, p. 190, 1871.
i Zool. Jahrb., Abth. Syst. vil. p. 732, 1894; Bronn’s Thierreich, Crust, p. 1178,
1899.
*| Jour. Asiat. Soc. Bengal, Ixix. pt. 2, p. 279, 1900.
** Nouv. Arch. Mus. Paris, (4) viii. p. 66, 1906.
ID AGS dCs INE IGOOVANIT.
G.M.Woodward del.et lith. 5 West, Newman imp.
GECARCINUCUS INGRAM.
Ea
1908. ] RIVER-CRAB FROM NEW GUINEA. 961
sight ridge runs backwards from it along the antero-lateral
margin, interrupted by the cervical groove and then forming a
small epibranchial tooth; almost immediately behind this the
ridge runs on to the dorsal surface of the branchial region and
dies away at about the anterior third of the length of the carapace.
The lower margin of the orbit is concave as seen from helow;
it has no tooth at its inner angle, which is indistinctly separated
from the inner suborbital lobe. On the lower surface of the
carapace the pterygostomial groove is well-marked and a faint
continuation of the cervical groove separates the sub-hepatic and
‘sub-branchial regions.
The epistome has the middle lobe more rounded than in G@. jac-
quemonti and the lateral portions less concave but more pro-
minent anteriorly. ‘The lobe of the first maxillipeds forming the
floor of the branchial channel is considerably more exposed than
in that species. The third maxillipeds have the ischium strongly
grooved along its whole length, the merus broader than long, “ts
anterior edge oblique and slightly concave; the exopodite extends
a little way beyond the ischium.
Chelipeds very unequal, the larger one very massive in the
male. Merus long, nearly the whole of it visible from above
beyond the carapace, each of its three edges carrying a row of
sharp spiniform teeth. Carpus with a strong spiniform tooth on
the inner side, behind which the inner edge has, at most, one or
two minute granules; the lower double tooth, present in G. jac-
quemonti, is here wanting. Chele smooth, with some faint
rugosities on the outer surface; the upper and lower margins are
rounded and there is a single blunt tooth at the proximal end of
the lower edge. The palm of the larger chela is but little longer
than high ; between the bases of the fingers on the outer side is
a large rounded or subconical tubercle. The fingersare str ongly
compressed and deep, equal to or shorter than che palm ; there is
one large serrated tooth on the immovable finger and two, smaller,
on the dactylus. The palm of the smaller chela is much longer
than, high, the tubercle on the outer side is small, the fingers are
more slender and their teeth are less prominent.
The walking-legs are very long and slender, the second pair the
longest. The merus has a small subterminal tooth above; the
propodus has a few spines on its lower, and some, more minute, on
its upper edge; the dactylus has four rows of spines.
The Abdesien of the male differs greatly from that of G. jac-
quemontu; it is strongly constricted a little way from the base,
the narrowest part being at the junction of the fifth and sixth
somites; the sixth somite is widened distally and the telson is
linguiform. The anterior part of the thoracic sternal surface .of
the male is not setose as it is in G. jacquemontit.
Locality. “‘Madeu, St. Joseph River, British New Guinea,
2000-3000 feet, W. Stalker coll.” Mr. Stalker informs me that
‘the species probably burrows in swampy ground, although he did
not actually see specimens taken from the burrows,
Lo) fs
962 ON A NEW RIVER-CRAB FROM NEW GUINEA. [ Dec. 15,
Measurements 11 millimetres :—
CO deers ea ae Se
iene thy ofenrsipaceta. 54.5. saseeseere 29) (26. 5. 29). 28
Breadth of carapace ; 24... a:2e+enscde Do ado, alae oD
dot) %fwonttich oat 3: Sat feeble yeceey 9 8 stm)
do. fronto-orbital margin... 18° 18 18 19
Length of larger cheliped ............ 762% 460.2) OBA eae
do. second walking-leg ...... LOS rn spetangrks ale
do. third walking-leg ......... GO Bde a F2- sol SS
For comparison I give the corresponding measurements of three:
specimens of G. jacquemontu * in the Natural History Museum :-—
Ge NG HM ON
Wength oimearapace. 2203. 22.56. Bo (30 fast
Breadth of carapace .............2.-+. 48 44 45
do. PROG AS, SLE SD) oe 8
do. fronto-orbital margin... 21 9 See2al
Length of larger cheliped ............ 74 68 ~~ 68
do. seed walking-leg ...... Gis) (0 67
do. third walking- alge HR ck 64 58 61
Cc
Remarks.—G@. ingrami agrees with G. Jawequemontit in the:
strongly inflated branchial regions and in the form of the efferent
pranchial orifices. The latter are bounded above by a prominent
lip formed by the everted edge of the epistome and below by the
enlarged terminal lobe of the first maxillipeds, which is left lar gely
exposed when the third maxillipeds are closed. Whether these.
characters are adequate to justify the alliance in one genus of
two species so widely separated geographically may perhaps be
disputed when the systematic arrangement of the Potamonide
comes to be more closely examined than it has hitherto been. For-
the present, the genus appears to be as well defined as are most
of those forming the family. It seems to me doubtful, however,
whether the subfamily Gecarcinucine, formed for its reception by
Miss Rathbun, can be sustained. The proportionate width of the
front, selected by that author as the most important distinction
in her key to the subfamilies and genera (Nouv. Arch. Mus.
Paris, (4) vi. p. 247), does not hold good, as the measurements.
given above show, even for G. jacquemontii, and it is still less
distinctive in the new species. On the other hand, it seems
probable that there are still to be discovered among the Pota-
monide characters more trustworthy as indications of affinity
than those hitherto employed. For example, the remarkable
form of the mandibular palp, as figured by Milne-Edwards for
G. jacquemontii, recurs not only in the species here described
* These specimens, presented by Mr. R. C. Wroughton, are trom Kaman River,
Bombay. As the measurements show, they are smaller than the type-specimens of
Milne-Edwards redescribed by Miss Rathbun, and have the carapace relatively
narrower, but in other respects they agree so closely with Milne-Edwards’s figures as.
to leave no doubt that they belong to : the same species.
1908.] ON MAMMALS FROM NORTHERN CHINA. 963
but also in several other Oriental Potamonide—I find it in
Potamon (Potamonautes) cunicularis and in P. (Parathelphusa)
tridentatwm, but not in African species referred to these two
subgenera,—and the clue thus afforded might, if followed up,
lead to results important for the classification of the family.
EXPLANATION OF PLATE LXVIII.
Vig. 1. Gecarcinucus ingrami, male, from above, natural size.
2
s is >» anterior part ot body, seen from in
front.
Bib oy 45 ,», larger chela, from the outer side.
4. 4 os » third mavyilliped.
5. 2 % . abdomen.
7, The Duke of Bedford’s Zoological Exploration in Eastern
Asia. —XI. On Mammals from the Provinces of Shan-si
and Shen-si, Northern China. By Oxprizetp THomas,
Johor theese ©
[ Received December 15, 1908. |
The collection dealt with in the present paper is a continuation
of that described in the last part (swpra, p. 635), which con-
tained a certain number of Shan-si Mammals, collected up to
February 1908. After despatching that series Mr. Anderson,
now accompanied by Mr. A. de C. Sowerby, an English resident
in Shan-si, commenced work again immediately after starting
from Tai-Yuen-Fu, whence he moved westwards and southwards,
across the Hoang-ho, down to Yen-an-fu, in Shen-si. He then
turned northwards again, crossed the Great Wall at the south-
eastern corner of the Ordos Desert, which I had asked him to
visit, and from there worked back again to the centre of Shan-si,
where, except for the interesting topotypical series of Hutamias
asiaticus senescens, picked up on the way to Peking, the collecting
of this set came to an end.
The collection as a whole is of extreme interest and value as
being the first we have received from the far eastern edge of the
inland desert area of Central Asia. It therefore gives us the
exact Eastern limit on this latitude of certain of the Central Asian
desert forms, such as Meriones, Dipus, and Ochotona, while in
other cases it shows the effect that this raised desert area has on
such forms as have penetrated to it from the lower country still
further east to the coast. This effect 1s mainly in a general
paling of colour, without alteration in structure, only one species 7,
* (The complete account of the new species described in this communication
appears here; but the names and preliminary diagnoses of those underlined were
published in the ‘ Abstract,’ No. 63 (Dec. 15, 1908).—KEprror. |
+ The Hedghog.
964 : MR. OLDFIELD THOMAS ON [ Dec. 15,
in the sense that I use that term, representing here a different
one from further east, all the others being what I should term
locally modified subspecies.
Besides these modified subspecies of the coast forms, there are
of course several interesting new species in the collection, notably
the beautiful little desert Hamster which I have named after the
Duchess of Bedford, and the curious Vole Mierotus inez. Jn all
twelve new forms are described in the present paper (see footnote
p- 963).
So far Mr. Anderson has made magnificent use of the Duke of
Bedford’s generosity, the amount and completeness of his collections.
already almost equalling those made during the Rudd exploration
of 8. Africa, and the results obtained being equally epoch-making
both for our knowledge of Hastern Asian Mammalia, and for the
improvement of our National Museum. Japan, from Saghalien to
Yaku-shima, T’su-shima, Korea, and Quelpart, and the chief areas
of North-eastern China, have now all been visited in turn, and
such collections made as to multiply a hundredfold our know-
ledge of the mammals of the region and to form a really sound
basis for further work on the subject.
No such complete and systematic survey has ever been made in
Eastern Asia before, and for this fine service to Science zoologists.
have to thank the generosity of our President.
Mr. Anderson gives me the following note on the characteristics
of Shan-si and Shen-si, the two provinces dealt with in the
present paper :—
“The provinces of Shan-si and Shen-si ave quite different in
character. The former may be briefly described as a mountainous
country with occasional large upland plains. Some peaks in
Shan-si rise above 10,000 ft., and are massive rocky mountains:
with only a comparatively thin coating of loess soil. Where the
loess figures mostly is in the plains, of which that of Tai-Yuen-Fu,
that of Ta-Tung-fu, and that of Hsiu-clou are the best examples.
The streams of Shan-si flow only in the rainy season, with the ex-
ception of the larger rivers. Northern Shen-si, on the other hand,
is a region of loess hills of almost uniform height ; the skyline of
Shen-si, seen from the mountains of its eastern neighbour, is a
straight line declining very gradually as it passes from north to:
south. The portion of Shen-si visited appears indeed like an
extension of the plateau of which Ordos is part, only this extension
has been cut into by a great many perennial streams, a process
which 1s now taking place in southern Ordos.”
1. Myorts sp.
3. 1875. Pao-teh-chow, Shan-si. 3500’.
A very old specimen with worn teeth. Related to J/. mystacinis.
6e “ > Tey ney hh 17 7 1
Bats were very rare throughout the area visited, this being
the only one seen.”—J/. P. A,
1908. ] MAMMALS FROM NORTHERN CHINA. 965
2. ERINACEUS MIODON.
Thos. Abstr. P. Z. 8. 1908, p. 44 (Dee. 15).
¢. 1846, 1853, 1855, 1856, 1857, 1862, 1869, 1871, 1873. Yu-
lin-fu, Shen-si. 4000’.
A pale-brown species allied to #, dealbatus Swinhoe; no wholly
white spines intermixed with the brown-ringed ones. P’ much
smaller than in #. dealbatus.
Size about as in #. dealbatus. Spines of back about 22-24 mm.
in length, white for two-thirds their length, then br oat ringed
with blackish brown, the ring about 4 mm. in breadth, the
terminal 3—4 mm. white (or the very extreme point darker again,
but not enough to affect the general tone). No wholly white spines
present. The resulting general colour * of the whole animal is
near ‘‘ drab.” Ears well haired. Head, sides, limbs, and tail
varying from dull whitish, or brownish white, to distinctly brown
(“ broecoli-brown”), the colour of the hairy part being evidently a
character of little value. Belly always lighter, sometimes quite
white.
Skull of about the same general proportions as in #. dealbatus.
Zygomata rather more abruptly expanded anteriorly. Premawxillee
extending backwards in a narrow point, which in most cases
reaches a forwardly projecting point of the frontals, thus completely
cutting off the maxille from the nasals; in aro cases, however,
the premaxillary and frontal points do not reach each other, so
that there is a short naso-maxillary suture, and it 1s evident that
the details of these sutures should only be used with very great
caution as distinguishing characters. In the four specimens that
I refer to #. dealbatus there is a long naso-maxillary suture.
Teeth at once distinguishable fc om those of H. dealbatus by
the conspicuously smaller size of p’, and in a lesser degree of p,
and the upper and lower canines. P’ forms a nearly equal- sided
triangle, its greatest diameter rarely exceeding 2 mm., while in
£. dad atus it is much broader than long, its transverse ’ diameter
being 3-4 mm. (in the type) and upwards.
Dimensions of the type, measured in the flesh :—
Head and body 215 mm.; tail 42; hind foot 40 (range from
35); ear 34°5,
Skull—condylo-basal length 53 mm. ; greatest breadth 36 ;
nasals 14°5 (diagonally) x 3; inter orbital breadth 18 ; palatal
length 29; front of i‘ to back of m’® 27.
Hab. as above. .
Type. Adult male. B.M. No. 9.1.1.9. Original number 1871.
Collected 11 May, 1908.
This Hedghog looks externally very like Swinhoe’s #. dealbatus,
* Even so speckled an animal as a Hedghog, when seen far off, may be said to
have a “general colour” resulting from the “intermingling of all the colours on the
surface, and it is in this sense that J always use the term ‘when describing mammals.
Some writers speak of “general colour ” for what I should term “ gr ound- colour,” a
very different thing. "y
966 MR. OLDFIELD THOMAS ON [ Dee. 15,
but is readily distinguishable by the much smaller size of p®. In
this respect Swinhoe’s Chefoo example, and the two obtained in
the same place by Mr. Anderson, closely agree with the type of
EL. dealbatus; and so far as our material goes I should not consider
Prof. Matschie’s Z. tschifuensis distinct from Swinhoe’s species,
especially as the present series shows how variable the characters
of the nasal sutures may be. The Chefoo specimens have from
10 to 20 per cent. of their spines wholly white, such spines being
in £, miodon conspicuous by their absence.
Dr. Satunin * has described two Hedghogs from Chingan and
Ussuri respectively, but both have many white spines mixed with
the dark ones. He makes no reference to the earlier described and
evidently closely related /. orientalis Allen, from Vladivostok,
the describer of which in turn ignores #. dealbatus Swinhoe.
“There appear to be large areas in North China where the
Hedghog is not found at all, and some places, of which the neigh-
bourhood of Yu-lin-fu is one, where they are remarkably common.
At the time we were at Yu-lin (April to May) the neighbouring
desert was alive with several species of beetle upon which the
Hedghog fed.
“The Hedghog seemed to be unknown in the vicinity of Pau-
teh-chow, Shan-si, but at Ning-wu-fu we heard reports of them,
though we saw none ourselves.
“Chinese name, ‘ Tsi-wei’ £ (¢si- a thorn or spine).”—J/. P. A.
The British Museum also contains another Hedghog from Shen-si
Province, collected by Father Hugh, and this again seems distinct
from any hitherto described. It may be called
ERINACEUS HUGHT.
Thos. Abstr. P. Z. S. 1908, p. 44 (Dee. 15).
A very dark-coloured, finely speckled species, quite unlike any
of the other Chinese Hedghogs. Spines light basally as in
E. miodon, but the dark ring is much broader, and is followed by
quite a narrow light ring, only about 0-5 to 0°8 mm. in length,
the point for about the same length being again dark. As a
result the whole animal is very dark with a fine whitish ticking,
and has quite a different appearance to the broadly washed whitish
of the other species. Head, limbs, and belly brown.
Hind foot of type 38 mm.
fab.. Paochi, Shen-si.
Type. Adult female. B.M. No. 0.6.27.2. Presented and collected
by Father Hugh.
The only species which this Hedghog might have been referred
to is H. hanensis Matsch., but I owe to the courtesy of
Prof. Matschie some spines from the type of that animal, and
these show quite a marked difference in the general coloration,
* Ann. Mus, Pétersb. xi. pp. 170-173, 1907
+ Bull. Amer. Mus. xix. p. 179, 1903.
f For the Chinese names given in this paper we are indebted to Mr. A. de
C. Sowerby.
1908. | MAMMALS FROM NORTHERN CHINA, 967
the bases of the spines being dark where they are white in
EF. hughi, while the terminal 3 mm. of the spines are light horn-
colour without a dark tip. The coat is also liberally mixed with
wholly white spines, which are practically absent in #. hughi.
3. Feuis carus L. (domestica auct.*).
1866 (skin without skull). Ordos near Yu-lin-fu, Shen-si.
A remarkably fine long-haired Cat, with a considerable
yvesemblance to a Kuropean Wild Cat. These cats are said to be
common in Northern China.
4, CANIS LUPUS TSCHILIENSIS Matsch.
¢. 1699. Yen-an-fu, Shen-si. 3000’.
This Wolf would appear to represent Prof. Matschie’s Lupus
tschiliensis.
“Uncommon. Much feared by the Chinese goat and sheep
herders.
‘¢ Chinese name, ‘ Lang ’.”—J/. P. A.
5. VULPES VULPES subsp.
2. 1666. 30 miles W. of Fen-chou-fu, Shan-si. 4500’.
“Common. Much hunted by the Chinese for the sake of its
skin.
“Chinese name, ‘ Hu-li’ or ‘ Hu-tai’.”—J/. P. A.
6. MARTES FLAVIGULA BOREALIS Radde.
g. 1704, 2. 1703 (mative skins without skulls). ¢. 172%
(skull only). Yen-an-fu, Shen-si. 3000’.
The fine skull no. 1727 agrees closely in dimensions with
Radde’s specimens, and is very considerably larger than an old
male skull of I. f. kuatwnensis Bonh.
“* Rather common, its tracks often seen.
‘Chinese name, ‘ Hwang-yao’.”—J/. P. A.
7. MuLEs LEProRHyNcHus M.-Edw.
@. 1870. Yu-lin-fu, Shen-si. 4000,
Although Yu-lin-fu is in the direction of the region where
Prof. Matschie’s WMeles hanensis and siningensis were procured, 1
fail to see any reason why this Badger should not be referred to
Milne-Edwards’s species, which was described from Peking.
“While difficult to capture the Badger is nota particularly
uncommon animal, as I have seen its Bade numbers of times
both in Shan-si and Shen-si.
“‘ Chinese name, ‘ Huan-tzi’.”—J/. P. A.
8. ScIUROTAMIAS DAVIDIANUS M.-Edw.
6. 1654. ¢&. 1655, 1656. 15 miles N.W. of Fen-chou-fu,
Shan-si.
* Of. Pocock, P. Z. 8. 1907, p. 149.
968 MR. OLDFIELD THOMAS ON [Dee. 15,
3g. 1720, 1824. @. 1716, 1722. Near Yen-an-fu, Shen-si.
3800".
©. 1981. 20 miles S.W. of Ning-wu-fu, Shan-si, 6600’.
‘“‘So far always found living among the rocky precipitous sides
of canyons where bushes are plentiful and some trees exist.
Nowhere common. One of thei foods is the kernel of the wild
peach. ;
“This squirrel has cheek-pouches like those of Chipmunks.”—
ie? Ae
9, KUTAMIAS ASIATICUS SENESCENS Mill.
3. 2004, 2005, 2006, 2007. ©. 2008. Mon-tou-ko, 15
miles W. of Peking. 500’. Zopotypes.
This interesting series of topotypes, which Mr. Anderson
obtained at my special request, and those next following, form a
most valuable addition to our collection of Asiatic Chipmunks
and have enabled me to gather a general idea of their local
characteristics.
In the first place, it appears evident that none of the forms
other than the original #. asiaticus can be properly called species,
as all grade into one another, each series varying to a certain
extent, and overlapping the members of the next. All seem to
be in fact members of one widely distributed species, modified by
local conditions, and one sees no sign of what is often found in
North America, where representatives of quite distinct species
may be found taking each other’s place in neighbouring localities.
Of course in N. America a much greater richness in different
types is present to be drawn upon, while here all are modifi-
cations of 1. asiaticus.
These main modifications appear to be four in number, so far
as we yet know :—(1) The sharply defined black and white five-
lined asiaticus of Russia and Siberia; (2) a more rufous form
inhabiting Saghalien, Hokkaido, the Amur region and Korea, to
which the names of wthensis, lineatus, and orientalis are assign-
able ; (3) the grey-mantled senescens of the Peking region, which
passes through an intermediate link into (4) the desert form, found
on the western edge of Shan-si and in northern Shen-si where it
borders on the Ordos Desert.
10. EUTAMIAS ASIATICUS ORDINALIS.
Thos. Abstr. P. Z. 8. 1908, p. 44 (Dec. 15).
6 - 1798, 1099, 11803, 1830; 1867, S72 t ens Ls0 Aisa
Yu-lin-fu, Shan-si. 4000’. 2
3. 1888, 1941. 9. 1887. Mts. 12 miles N.W. of Ko-lan-
chow, Shan-si, 7000’.
A pallid, semi-desert race allied to H. a. senescens.
Colour much paler throughout than in senescens. . Crown paler
and more approaching pinkish buff; shoulders and nape with
almost no grey in them, scarcely more grey-grizzled than the
1908.] MAMMALS FROM NORTHERN CHINA. 969
cream-buff cheeks and sides; rump more or less ochraceous-buff,
markedly brighter than in senescens, where it is “‘ raw umber” ;
dark dorsal stripes lightened in intensity by their hairs being
largely tipped with ochraceous; their porportionate lengths as in
senescens.
Dimensions of the type, measured in the flesh :—-
Head and body 139 mm.; tail 125; hind foot 39; ear 19°5.
Skull—ereatest length 41 mm.
Hab. of type. Yu-lin-fu, Shen-si. 4000’.
Type. Adult female. B.M. No. 9.1.1.36. Original number
1804. Collected 1 May, 1908.
“ Live in low bushes, not climbing trees.
“ Not until the latter part of April, about the time we reached
Yu-lin-fu, did the weather become warm enough to attract the
Chipmunks from their holes. They frequent the sides of the
loess gullies mostly, and are usually fairly common where found
at all. Besides the places where specimens were.collected, two
were seen at Pao-teh-chow, and, I believe, from the people’s
accounts, that they occur at Yen-an-fu. Mr. Sowerby states that
Chipmunks are common near T'ai- Yuen-Fu.
«‘ A Jarge proportion of Chipmunks, wherever we found them,
had their tails broken short.”—W/, P. A.
Tt is only in accord with the usual order of things that the
Chipmunks of the country edging the Ordos Desert should reflect
their surroundings by being markedly paler than their allies near
“Peking. So great is the difference indeed that an intermediate
link in the series might suitably have a special subspecific name,
as follows :—
11. HtrAMIAS ASIATICUS INTERCESSOR.
Thos. Abstr. P. Z. 8. 1908, p. 44 (Dee. 15).
©. 1971, 1972, 1998, 2002. Ning-wu-fu, Shan-si. 6000.
Sd. 1984, 1987, 1992,1993. @. 1982, 1985, 1986, 1989, 1990,
1991. 20 miles S.W. of Ning-wu-fu. 6000’.
Intermediate in intensity of colour between the dark grey-
mantled senescens and the pallid ordinalis. Ground-colour of
shoulders light greyish, more grizzled than in ordinalis, less than
in senescens; cheeks and sides buffy. Rump darker in tone than
in ordinalis, but in this respect nearer that form than senescens,
the general colour near clay-colour. Proportions and intensity of
dorsal stripes about as in senescens, though the dark ones are
rather more numerously grizzled with ochraceous.
Dimensions of the type, measured in the flesh :—
Head and body 145 mm.; tail 133; hind foot 30; ear 19.
Skull—greatest length 42 mm.
Hab. of type. Ning-wu-fu, Shen-si. 6000’.
Type. Adult female. B.M.No.9.1.1.42. Original number 1972.
Collected 14 June, 1908.
With 7 authentic examples of senescens before me, with 14 of
970 MR. OLDFIELD THOMAS ON | Dee. 15,
this intermediate form, and 11 of the desert ordinalis, | have
felt justified in indicating their respective degrees of differences
by subspecifie names. All are of course members of the widely
spread species #. asiaticus, with the eastern uthensis type of
which the Imperial Tombs specimens mentioned in a previous
paper tend to connect the true senescens.
The country in which this Chipmunk is found is mountainous
and broken, while #. a. crdinalis inhabits the flat region
bordering the sandy Ordos.
12. CrrELLus Moncoxticus M.-Edw.
3. 1731. Ching-pien, N.W. Shen-si. 5100’.
3. 1738. @. 1734, 1746. Ordos Desert, N.W. of Ching-
pien. 4900".
3. 1747, 1851, 1863, 1864, 1868. 92. 1748, 1858, 1865.
Yu-lin-fu, Shen-si. 4000’.
These specimens agree closely in their general sandy coloration
with Milne-Edwards’s figure of mongolicus, and with the example
obtained by Swinhoe near Suen-hwa-fu in 1863,* which may be
accepted as a topotype, for David collected a number of his
“Mongolian” specimens at this latter place, which is below, not
on, the true Mongolian plateau.
On the other hand, our Mongolian plateau specimens, both
those collected by Mr. C. W. Campbell at Hara Ussu in 1898, and
by Mr. Anderson at Taboul in 1907 7, ave so markedly darker in
colour that they might be recognized as a special plateau sub-
species as follows :—
CITELLUS MONGOLICUS UMBRATUS.
Thos. Abstr. P. Z.S. 1908, p. 44 (Dee. 15).
Size and proportions as in true mongolicus. Colour much
darker and greyer, speckled with blackish and buffy, so as to
result in a tone rather darker than Ridgway’s “isabella.” Crown
near broccoli-brown, markedly darker and less fawn than in
mongolicus. Under surface broadly washed with buffy, lips and
chin white. Sides of neck, front of forearms, and back of lower
legs more strongly suffused with tawny or tawny ochraceous than
in mongolicus, 11 which the colour is sandy or buffy. Tail-hairs
much shorter than in mongolicus, though this is probably a
seasonal character, cream-buff at their bases and tips, their
middles black, none of the strong ochreous-buff colour showing on
the upper side; below the middle line is ochraceous buff, but far
narrower and less conspicuous than in mongolicus.
Dimensions of the type, measured in flesh :-—
Head and body 197 mm.; tail 62; hind foot 37.
‘Skull—ereatest length 46°3 mm.; basilar length 37; zygo-
matic breadth 28; length of upper tooth-series 10.
* See P. Z. S. 1870, p. 445.
+ See P. Z.S. 1908, p. 105.
1908. | MAMMALS FROM NORTHERN CHINA, 971.
Hab. Mongolian Plateau. Type from Taboul, about 100 miles
N.W. of Kalgan. Alt. 5000’.
Type. Young adult male. B.M. No. 8.3.5.5. Original number
1499. Collected by M. P. Anderson, 1 Augnst 1907, and presented
by the Duke of Bedford, K.G.
The difference in general colour between wmbratus and
mongolicus might have been thought to be seasonal in its nature,
as both Mr. Campbell’s and Mr. Andergon’s specimens of the
former were collected in July and August, and the present series
in Apriland May. But some of the latter have already got their:
summer pelage on the crown, while Mr. Swinhoe’s topotypica
example of mongolicus was killed in September.
From the region inhabited by Biichner’s two species, C. alas-
chanicus and C. obscwrus, the Mongolian plateau is separated by
the western parts of the range of C. mongolicus.
“Fairly common. Usually living in valley-bottoms, or in
stretches of plain where more or less grass exists.
« Chinese name, ‘ Sa-hsu’ = Sand-rat.”—W/. P. A,
13. Merrones AucEPS Thos.
3. 1670, 1675, 1676, 1677, 1678, 1684, 1685, 1693, 1695.
©. 1679, 1683, 1694, 1701, 1702. Yen-an-fu, Shen-si. 3000’.
3S. 1736, 1737. Ordos Desert, N. of Ching-pien, Shan-si,
4900’.
¢- 1878, 1880. @. 1877, 1881, 1882. Pao-teh-chow, Shan-si.
3500".
@. 1917, 1958. Mts. 12 miles N.W. of Ko-lan-chow, Shan-si..
7000".
3. 1960. Ning-wu-fu. 6000’,
This handsome species, one of the discoveries of the present.
exploration, was described in my previous paper from a single
specimen, so that this good series is very welcome. Theskins are
on the whole very uniform, with the exception that the tail is
sometimes white below, prominently bicolor, and sometimes wholly
ochraceous, all intermediate stages between the two being present.
It is possible that Milne-Kdwards may have mixed up some
specimens of J/. auceps in his account of IZ. psammophilus, as the
two species are so similar; but I have taken as representing his
species the example unquestionably belonging to the smaller form,
which he sent to the British Museum in 1867.
‘¢ One of the commonest mammals of North China. At Yen-an-
fu they were abundant, burrowing in the farm fields; in the
grassland north of Ching-pien they were also plentiful, but here
they lived in the grassy plains, or in the bushy areas along the
edges of the plaims. This rodent was not found at Yu-lin-fu,,
and was comparatively rare near Ko-lan-chow, while at Ning-wu-
fu I did not see them.
“‘ Chinese name, ‘ Hwang-hsu ’= Yellow rat.” —J/, P. A,
972 MR. OLDFIELD THOMAS ON [ Dec. 15
14. MerronEs psAMMOPHILUS M.-Edw.
g. 1652. ©. 1653. Tai-Yuen-Fu, Shan-si. 2700".
Distinguishable from MW. auceps mainly by its smaller size,
shorter tail, and smaller bulle.
15. Meriones uneurcuLatus M.-Edw.
go. 1739, 1740. 9. 1741, Lian)” Ordos Desert NE Wenn
Ching-pien, Shen-si. 4900’.
This Gerbil is readily distinguishable from the other two
Chinese species by its black claws, grey-based belly-hairs, and
buffy instead of ochraceous tail. My. Anderson had previously
obtained a good series of it at Taboul on the Mongolian plateau.
Its skull is very like that of JZ. psammophilus, but the bulle
are smaller, and do not abut on or overlap the hinder corners of
the zygomata.
16. Mts CONFUCIANUS LUTICOLOR.
Thos. Abstr. P. Z.S. 1908, p. 45 (Dec. 15).
g. 1689, 1698, 1710. Yen-an-fu, Shen-si. 3000’.
_ Apale race of M. confucianus, smaller and more delicately
built than the Shantung J/. c. sacer.
Size rather less than in sacer. Fur soft and fine, without
spines. General colour above pale buffy (between cream-buff and
buff of Ridgway) lined with brown along the dorsal area. | Sides
clearer buffy. Under surface and inner side of limbs pure, sharply
defined, buffy white, whiter on the chin, more creamy on the
belly. Ears large, pale greyish brown with white edges. Upper
surface of hands and feet white. Tail well-haired, the scales
being practically hidden, pencilled terminally, the hairs at the
tip 6-7 mm. in length ; brown above proximally, white terminally
and below, the amount that is white varying from one-third to
two-thirds the length of the tail.
Skull smaller and more delicately built throughout than that
of I. c. sacer, the brain-case smoother and more rounded and
the ridges less developed ; palatal foramina shorter; bulle rather
Jarger ; molars smaller.
Dimensions of the type, measured in the flesh :—
Head and body 130 mm. ; tail 167; hind foot 27; ear 23.
Skull— greatest length 35°3 mm.; basilar length 27-4; greatest
breadth 17; nasals 12°7; interorbital breadth 5:3; palatilar
length 14:7; palatal foramina 6-5; upper molar series 5:6.
Hab. as above.
Type. Adult male. B.M. No. 9.1.1.92. Original number 1689.
‘Collected 13 March, 1908.
This is a more or less desert form of the widely spread
M. confucianus, smaller than J. c. sacer, paler coloured than the
true confucianus.
‘A yare animal in Shen-si, where we found it living in some
rocky barren gullies. This rat is apparently not dependent or
1908. } MAMMALS FROM NORTHERN CHINA. 973
cultivation; wherever [ have found it, namely in Shantung,
Chih-li, Shan-si, and Shen-si, it has lived in rocky places among
trees and bushes.”—J/. P. A.
17. Mts wAGNERI MoNGOoLIUM Thos.
@. 1663. 30 miles W. of Fen-chou-fu, Shan-si. 4500’.
6.1777. 2.1778. Yu-lin-fu, Shen-si. 4000'.
“Found in hillside fields near Fen-chou-fu; at Yu-lin-fu
common in fields upon the river-bank.”—J/, P. A.
18. APODEMUS SPECIOSUS subsp.
6. 1705, 1706, 1707, 1718. 2. 1709, 1719. Near Yen-an-
fu, Shen-si. 3800’.
6- 1893, 1894. 1895, 1900, 1912, 1926, 1927, 1937, 1938,
NO4AS OA TIO l902 1910, Weve eh) U/L 1957, 1959.
Mountains 12 es IN W. ot 1 lan- hon Shen -Si. 7000’.
@. 1983, 1995. 20 milesS.W. of Ning-wu-fu, Shan-si. 6600’,
“This mouse is another form which seems not to depend on
cultivation at all. It is found only in those rare spots in North
China where trees and bushes are numerous. At Yen-an-fu it
was only fairly common, but in the mountains 12 miles north-
west of Ko-lan-chow we found this animal abundant.”—J/, P. A.
19. APODEMUS AGRARIUS PALLIDIOR Thos.
6. 1657. 30 miles W. of Fen-chou-fu, Shan-si. 4500’.
G. 1713. ©. 1717. Near Yen-an-fu, Shen-si. 3800’.
6. 1936. Mountains 12 miles N.W. of Ko-lan-chow, Shan-si.
7000’.
Quite similar to the typical series from the Shantung Peninsula.
““TIn cultivated fields. Very rare in this part of China.”—
Ji yal el
20. CRICETULUS TRITON INCANUS.
Thos. Abstr. P. T. S. 1908, p. 45 (Dee. 15).
6. 1708, 1714. 2. 1715, 1725. Yen-an-fu, Shen-si. 3000
G. 1898. 2.1945. Mountains 12 miles N.W. of Ko-lan-
chow, Shan-si. 7000’.
A paler, clearer grey race of the Shantung C. triton De Wint.
External characters as in true C’. triton except that the colour
is distinctly paler and more drabby (drab-grey) as compared to
the darker “ smoke-grey” triton; the head and fore back pale
clear grey, “ grey No. 3,” markedly different from the compara-
tively dark grey of triton.
Skull essentially as in triton, but rather more delicately built
the nasal region, interorbital SHIGE, and brain-case all lightly
narrower.
Dimensions of two specimens, measured in the flesh :—
¢. Head and body 155 mm. ; tail 85; hind foot 25; ear 21.
9g (type) 29'P 168 ”m OO ” 98 ; + 2A oS oe
974 MR. OLDFIELD THOMAS ON [ Dec. 15,
Skull of type—condylo-basal length 41 mm.; basilar length 36 ;
zygomatic breadth 22°8; nasals 15:7; interorbital breadth 5-4 ;
palatilar length 17:5; palatal foramina 7°2; length of upper molar
series 5:1.
Another fully adult skull only measures 37°3 mm. in condylo-
basal length, while an example of true triton reaches 39 mm.
The type skull of C. triton, which has worn teeth, is only 33 mm.
in the same measurement, so that members of this group evidently
vary very much in the size that their skulls may attain to.
Hab, of type. 12 miles N.W. of Ko-lan-chow, Shan-si. 7000'.
Type. Old female. B.M. No. 9.1.1.123. Original number
1945. Collected 3 June, 1908.
This is evidently a pale inland dry-country form of the coast
C. triton.
“This large Hamster israre. They usually reside under bushes
at the edge of some farmfield from which they take their food.
I have sometimes found green leaves in their pouches, but more
often they carry some grain. Their clean-cut burrows usually
descend vertically into the earth.
‘Chinese name, ‘ Pan-Tsang-er.’”—JV/. P. A.
21. CrICETULUS ANDERSONI Thos.
3. 1660, 1664. 9. 1658, 1661, 1665. 30 miles W. of Fen-
chou-fu, Shan-si. 4500’.
6 .. 1667, 1668, 1671; 2672;.11682, W69U 1723. (2. Teak
1690, 1692, 1700, 1711, 1712, 1721. Yen-an-fu, Shen-si. 3000-
3800!.
3S. 1883. 9.1876, 1879, 1884. Pao-teh-chow, Shan-si. 3500’.
d. 1889, 1903, 1904, 1905, 1906, 1914, 1922, 1923, 1924, 1925.
@. 1885, 1886, 1890, 1899. Mountains 12 miles N.W. of
Ko-lan-chow, Shan-si. 7000.
9.1961. Ning-wu-fu, Shan-si. 6000/.
There is surprisingly little difference either in colour or length
of fur between these summer specimens and those obtained by
Mr. Anderson the previous winter, when he first discovered this
well-marked little species.
‘“T consider the grey dwarf Hamster the most abundant mammal
of Shan-si and Shen-si. It was common at all our collecting
erounds within these provinces, with the single exception of Yu-
lin-fu, where it seems not to exist. We failed to find it in Ordos.
also.” —M. P. A.
Lo
2. CricEtuLUS GRisEUS M.-Edw.
@. 1815. Yu-lin-fu, Shen-si. 4000!.
23. CRICETULUS BEDFORDIZ.
Thos. Abstr. P. Z.S. 1908, p. 45 (Dee. 15).
g. 1757, 1758, 1778, 1774, 1775, 1801, 1805, 1806, 1808,
1823, 1831, 1842, 1843, 1844, 1854, 1860, 1861. 2. 1731 (in
1908. | MAMMALS FROM NORTHERN CHINA. 975
spirit), 1756, 1802, 1807, 1809, 1810, 1814, 1818, 1832, 1833,
1834, 1835, 1836, 1845, 1859. Yu-lin-fu, Shen-si. 4000",
@. 2003. Wu-chai, 23 miles W. of Ning-wu-fu, Shan-si.
6100".
A small, very short-tailed species with completely hairy soles
and pure white belly.
Size very small. Fur soft and fine, hairs of back about 9 mm...
in length. General colour above drab-grey, becoming on the
flanks and posterior back more distinctly drab (‘* eeru-drab,” in
some cases approaching “ pinkish buff”). Whole of under surface,
lower part of sides, all four limbs, and tail pure snowy-white, the
line of demarcation, which runs from just below the eyes to the
top of the base of the tail, well defined, more or less serpentine,
convex upwards at the shoulders and hips, downwards on the
flanks. A prominent white patch over each eye. Hars of
medium size, their proectote dark brown, their metentote white ;
a whitish patch behind their posterior bases. Palms and soles
completely covered with white hairs, except for a naked patch at
the base of the pollex. Tail short and stumpy, about the length
-of the hind-foot, well-haired, completely white.
Skull smaller in all dimensions than that of C. roborovskii,
apparently the nearest ally of the present species.
Dimensions of four specimens, taken in the flesh :—
6 1773. Head and body 81 mm.; taill4; hindfoot 12 ; ear13.
3 1861 (type). ,, lg coe ee le os 12) | aie
g 1756. si ae ee eye rs TOY) Salas
© 1807; a smote hee “etl = A 1 1ottt ae
Skull of type—greatest length 23:1 mm.; basilar length 18;
greatest breadth 13-2; length of nasals 7-7; interorbital breadth
3°6; brain-case breadth 10°8; palatilar length 9-6; diastema 6-2 ;
palatal foramina 3°9; upper molar series 3:1.
Type locality. Yu-lin-fu, Shen-si. 4000’.
Type. Old male. B.M. No. 9.1.1.165. Original number 1861.
‘Collected 8 May, 1908.
This very beautiful little Hamster, which I have named in
honour of the Duchess of Bedford, is most nearly allied to the
Nan-Shan (. roborovskii Satunin,* with which it appears to
agree in colour and structure. But it is very markedly smaller,
the type of C. roborovskii (stated to be young) having a head and
body length of 90 mm., and a larger skull throughout, the molar
series being 3°8 mm. in length.
“The Desert Hamster is a common animal in the region of
Yu-lin-fu, where it is found in the sandhills of the desert. I was
unable to find the burrows of these animals, and it seems probable
that the shifting sand closes their entrances as soon as the animal
has passed through.
“One specimen, No. 2003, came from Wu-chai, Shan-si, a town
* Ann. Mus. Zool. St. Pétersb. vil. p. 571, 1902.
Proc. Zoou. Soc.—1908, No. LXII. 62
976 MR. OLDFIELD THOMAS ON [ Dee. 15,,
some 23 miles west of Ning-wu-fu, and situated upon a portion
of the Mongolian Plateau which extends into Shan-si.
“The Desert Hamster eats millet very greedily.
‘Chinese name, ‘ Mi-tsang-er’ (Mi=millet).”—J/, P. A.
24, Microrus MAnpDaRiInus M.-Edw.
o: 1907, 1918, 1929, 19500) Se 9081951. li Mountainsal2
miles N.W. of Ko-lan-chow, Shan-si. 7000!.
These specimens agree very fairly with Milne-Edwards’s de-
scription, and with his type, which I have examined in Paris. The
species was described from Chinese Mongolia, and its occurrence
here in Shan-si is therefore quite natural.
These are the first examples of the true J. mandarinus that
the Museum has received, those from Afghanistan previously
referred to the species being undoubtedly different. The latter
are members of the subgenus Phaiomys, and have the connection
between the second and third enamel-space of the first lower
molar which is characteristic of Pitymys and of true Phaiomys,
and may in fact be taken as equally diagnostic of the latter.
But if this be done, Wicrotus brandti and mandarinus, both of
which have five ciosed triangles in m,, cannot be considered
members of Phaiomys, and would either be referable to Wierotus,
in spite of thei long fore-claws, or Lataste’s subgenus Lasio-
podomys, founded on J. brandti, should be revived to contain
them.
The following are flesh measurements of Jf, mandarinus :—
3g. Head and body 95 mm.; tail 23; hind foot 17; ear 7.
2 . ” 90 38 20 ; ” 16; ” 8.
“A vare thing: found only upon the mountain tops (7000 feet).
among bushes and grass, or in open fields. Their burrows much
resemble those of the ‘gray dwarf hamster’ (C. andersoni); we
trapped dozens of hamsters in our endeavour to secure more of
these voles.”—W/. P. A.
25. Microrus (KorHENomys) INEZ.
Thos. Abstr. P. Z. S. 1908, p. 45 (Dec. 15).
3. 1896, 1919, 1921, 1930, 1949, 1952, 1954, 1956. 9. 1891,
1892, 1897, 1909, 1920, 1946, 1947, 1948, 1953, 1955. Moun-
tains 12 miles N.W. of Ko-lan-chow, Shan-si. 7000'.
A small pale-brown Vele with a certain skull-resemblance to-
M. (Hothenomys) melanogaster.
Fur soft and fine, hairs of back 8-9 mm. in length. General
colour above a peculiar pinkish brown, rather browner than
Ridgway’s ‘“ fawn-colour,” lined with darker brown on the head,
and often rather paler on the fore-back. Under surface pale
‘* wood-brown,” the slaty bases of the hairs showing through, Kars.
about the length of the fur, not distinguishable by colour from
the general tone. Upper surface of hands and feet dull whitish ;
fore claws slightly longer than hind; soles with 6 pads. Tail of
1908. | MAMMALS FROM NORTHERN CHINA. 977
average length and hairiness, brown above, lighter below, not.
sharply contrasted. Mamme 0—-2=4.
Skull with a marked general resemblance in form to that
of MW. melanogaster, having the same unusually broad interorbital
region, and smooth unridged surface with the angles and crests.
scarcely developed, not even the oldest specimen showing any
trace of frontal crests. Palatal foramina medium. Posterior
palate more complete than in that species, the lateral grooves
reduced to small or minute disconnected foramina, the posterior
border squarely transverse, running completely across to the
molars; ridges bounding mesoptery ‘ygoid fossa running above
(or to) this posterior border, and curving down to join it on
ts dorsal aspect. Bullee of average size.
Teeth with the triangles tending throughout to be completely
closed, thus contrasting with those of J. melanogaster in which
they are mostly open. M' with the usual 5 spaces, 3 external and 3
internal salient angles, and m* with 4, 3 and 2 respectively, and
each tooth with a tendency to the development of a small extra
postero-internal angle, very different to the large extra angle of
the allied form. M’* narrow, rather elongate, with five separated
spaces, three external and three internal salient angles, and a
long posterior lobe. First lower molar normally with four closed
triangles, and the partly open one in front of them joining the
anterior trefoil, but this is sometimes also closed, making a fifth
closed triangle. M, either without closed trian cles, or the middle
Space separ. ated into two.
Dimensions of four specimens, measured in the flesh :—
Head and body 88 mm.; tail 31; hind foot 16 ; ear 10°5
3. st SOR Fad Gin moor - 16 fe5y geMnaict alls
(type) ,, GO ree a ang i153 re LOS Oe salle
s re OO. ee tes aaiaie ~ LG p45 Wit elite.
kull of Hype eientes length 23°5 mm.; basilar length 20;
zygomatic breadth 15:5 - length of nasals 6°7 ; interor pital breadth
4-2; height of crown fr om front of alveolus of m? 7-2: ; palatilar
length 10-1; palatal foramina 4:1; length of upper molar series
(crowns) D7.
Hab. as above.
Type. Adult female. B.M. No. 9.1.1.188. Original number
1892. Collected 28 May, 1908.
This interesting little Vole would appear to be most nearly
allied to J/. (Fothenomys) melanogaster, but differs strikingly in
the closure of many of the dental triangles open in that animal.
Its palate is also much more complete posteriorly, and its fore
instead of its hind claws are slightly the longer.
In colour WM. inez is also unique, its peculiar pimkish-brown
colour being quite unlike that of any other Vole in the Museum
collection ; “indeed, owing to its colour, it was mistaken for a
ee Ricdback ” by Me. Anderson.
“ Inthe mountains N.W. of Ko-lan-chow, Shan-si, this Redback
62*
978 MR. OLDFIELD THOMAS ON [ Dec. 15,
wag found rather common in the bottoms of certain narrow,
wooded and bushy gullies. They, in company with dpodemws
speciosus, burrow in the soft loose soil beneath the bush.” —W/. P.A.
26. CRASEOMYS SHANSEIUS Thos.
go. 1931. @. 1932, 1933, 1934, 1935, 1942. Mountains
12 miles N.W. of Ko-lan-chow, Shan-si. 7000’.
Practically topotypes of this striking species, described from
Mr. Anderson’s previous collection.
The summer fur is hardly shorter than that of winter, and is
quite similar in colour and texture.
‘‘¢ Much less common than the last, but like it found in the bush-
overgrown gullies and canyons.” —WM. P. A.
97. MyYosPALAX FONTANIERI M.-Edw.
3. 1733. Ordos Desert, 8 miles N. of Ching-pien, Shen-si.
A900".
3g. 1916. Mts. 12 miles N.W. of Ko-lan-chow, Shan-si. 7000’,
g. 1966, 1973, 1997. 2. 1965, 1996, 2001. Ning-wu-fu,
Shan-si. 6000’.
3. 1988. @. 1980, 1994. 20 miles 8.W. of Ning-wu-fu,
Shan-si. 6600’.
These specimens all agree in possessing some evidence of the
white face-markings figured by Milne-Edwards in his type, but
believed by him to be inconstant. He may have mixed up with
M. fontanieri examples of the next species, in which the white
crown-streak is rarely present.
Specimens of the remarkable Mole-rats of this genus are
exceedingly rare in Museums, so that this fine series of a species
new to the Museum, and those next referred to form a most
valuable accession to the Museum collection. In addition we have
lately received from Mr. E. B. Howell some examples of
M. psilurus M.-Edw., so that the Museum now possesses examples
of every described species of the genus, including really good
series of the two now obtained.
28. MyospaLax cansus Lyon.
Myotalpa cansus Lyon, Smiths. Mise. Coll. 1. p. 184, 1907.
@. 1729, 1730. Yen-an-fu, Shen-si. 3000’.
6 1794, 1795, 1800, 1811), 18165 USi ey USloe eon alse,
1838, 1849, 1850.. 2. 1751,1791,,1796, 1797, N8l2; e207 Nee
1827, 1837, 1839, 1840, 1841, 1847, 1852. Yu-lin-fu, Shen-si,
4000’.
This species differs from the last mainly in size, the largest —
skull, which agrees closely with the figure given by Mr. Lyon,
being 45:3 mm. in condylo-basal length. Face-markings are less
developed, only five out of 28 specimens having small frontal
streaks, while the buffy nose-patch is small and often almost
obsolete.
1908. | MAMMALS FROM NORTHERN CHINA. 979
In this series there is a marked difference between the sexes,
the male skulls being markedly larger, more rugged and heavily
ridged than the female. Mr. Lyon had stated that his specimen
was a female, but on my sending him a pair of the present series
for comparison, he has been good enough to inform me that the
male agreed precisely with the type, and that the latter was
evidently wrongly sexed.
“ This small rodent mole is rather common, for traces of it, old or
new, may be found in‘nearly every field in those parts of Shan-si
and Shen-si visited. It is, however, difticult to trap ; we secured
the present series chiefly by hiring peasants to dig the animals
out and bring them to us. In walking on the surface of the
eround the rodent mole turns the long claws of the fore-feet under
the soles and walks upon the backs of the claws; they make but
slow progress. When frightened or angered this animal utters a
peculiar little squeal. We had many live ones in the course of
our work at Yu-lin-fu and elsewhere, but did not find them
inclined to be savage.
“* Chinese name, ‘ Ha-whei’ or ‘ Ha-lao.’ ”—WM, P. A.
29. Dipus sowERBYI Thos.
g. 1743, 1744. @. 1745. Ordos Desert, 30 miles W. of Yu-
lin-fu, Shen-si. 4700’.
Se oO daa ios, 1iob; NGO 1761, Li62, 1163, liGe
1765, 1766, 1768, 1770, 1771, 1772, 1776, 1782, 1783, 1784, 1785,
1787, 1788, 1789, 1792, 1824,1825. ©. 1749, 1754, 1767, 1786,
1793, 1826. Yu-lin-fu, Shen-si. 4000’.
Dipus sowerbyi was obtained by Mr. Sowerby at Yu-lin-fu and
described in a special paper *. It is the first three-toed Jerboa
discovered in the Far East, the five-toed Allactaga mongolica
having been hitherto the only known Chinese Jerboa. Reasons
for the separation of the genus Dipus from the earlier Jaculus are
given in the paper quoted.
The species is probably characteristic of the Ordos Desert and
its vicinity.
‘“‘ Inhabits the sand-dunes.”—I/. P. A.
30. ALLACTAGA MONGOLICA Radde.
3. 1978. Ning-wu-fu, Shan-si. 6000’.
“ TInhabits the loess plain near Ning-wu-fu, where its tracks are
fairly plentiful. On the plateau west of Ning-wu we saw tracks
which were probably made by this animai, but we were unable to
identify its burrow, and could not secure a second specimen.’ —
M. P.A.
31. LEPUS SWINHOEL SUBLUTEUS.
‘Thos. Abstr. P. Z. 8. 1908, p. 45 (Dee. 15).
_¢. 1674, 1681, 1696. @. 1686. Yen-an-fu, Shen-si. 3000’.
* Ann. Mag. N. H. (8) it. p. 307, 1908.
980 MR. CLDFIELD THOMAS ON (Dec. 15,
gd. 1735. Ordos Desert, N. of Ching-pien, Shen-si. 4900’.
3. 1759 (young), Pe oe Shen-si. 4000’.
A pale race of LZ. swinhoet.
General colour throughout, of head, body, and chest-band, much
paler than in true sw vee near ‘ pinkish-buff” of Ri d gway,
instead of the richer colour of swinhoei, which approaches
“ ochraceous buff.” Sides of rump with a greater tendency to the
development of grey patches. Other characters as in true
swinhoer.
Dimensions of the type, measured in the flesh :—
Head and body 466 mm.; tail 91; hind foot 111; ear (from
notch) 95.
Skull—greatest length 86°5 mm. ; basilar length 70.
Type locality. Southern Ordos Desert.
Type. Adult male; B.M. No. 9.1.1.261. Original number
1735. Collected 14 April, 1908.
This Hare is a pale Ordos Desert race of LZ. swinhoei, which
yanges in its normal form from Chefoo and Nanking westwards
to Southern Shen-si. It unfortunately happened that the type
specimen was darker than has since proved to be usual with
Chefoo specimens, and deceived therefore by the description I
gave of it in 1894, before any of Mr. Anderson’s specimens bad
arrived, Prof. Matschie has recently distinguished a Hare from
Hing-an-fu, Southern Shen-si, as Lepus filehneri. is description,
however, quite fits Mr. Anderson’s topotypical series from Chefoo,
and I think there is no doubt LZ. filchneri should be referred to
L. swinhoet.
I regret that Prof. Matschie should have been led into error by
my description of what has since proved to be the abnormal colour
of the typical specimens.
With regard to his Z. stegmanni from Kiau-chow, | would note
that a certain proportion of the specimens of ZL. swinhoei, other-
wise indistinguishable, show the peculiar light speckling of the
upper tail-surface which Prof. Matschie uses as a primary
character. One example from Chefoo shows this very clearly, as
does another from Tung-chow, E. of Peking.
Dr. Satunin has recently * described a number of Hares from
Central Asia, but I cannot find any among them similar to this
Ordos Hare. I note, however, that his LZ. kozlovi, from Kam, can
hardly escape being the same as ZL. sechuenensis de Wint.7, almost
from the same district, of which he makes no mention.
“This is the common Hare of China. Although specimens were
taken at only two localities on our long journey, they weve seen at
all stages. They live generally wherever there is cover. In the
Ordos, north-west of Ching-pien, and in the mountains near Ko-
lan-chow they were exceedingly abundant.
““Chinese name, ‘ Tu-tzi.’”— VU. P. A.
* Ann. Mus. St. Pétersb. xi. p. 162, 1906.
$ P.Z.S. 1899, p. 576.
ee MAMMALS FROM NORTHERN CHINA. 981
. OcHoron A BEDFORDI.
Thos. Abstr. P. Z. 8. 1908, p. 45 (Dec. 15).
3. 1673, 1680. ©. 1687, 1688, 1697. Yen-an-fu, Shen-si.
3000".
6. 1915. @. 1944. Mountains 12 miles N.W. of Ko-lan-
chow, Shan-si. 7000’.
3. 1967, 1969, 1974, 1975, 1976,1999. ©. 1962, 1963, 1968,
1977, 2000. Ning-wu-fu, Shan-si. 6000’.
A Pika allied to O. dawurica, but with larger bulle. Size
rather larger than in O. dawurica. General colour above of
summer specimens pale wood-brown, tending to ochraceous-bufty
on the sides of the neck, the under surface approaching pinkish-
buff. Winter specimens rather paler and greyer. Patches behind
ears inconspicuous, ochraceous-buff; ears with their proectote
black proximally paling to dull bufty ter minally ; metentote buffy ;
lips and chin white, without darker markings. Hands and feet
creamy or buffy above, the long hairs of the palms and soles
whitish brown.
Skull with the general characters of O. dawwrica, the type
of the subgenus Ochotona (see below), but larger, less strongly
convex on the forehead, with the brain-case larger, broader, and
running out to more definite postero-external angles, and the
bulla very markedly larger, perhaps the largest in the genus.
Molars broader.
Dimensions of four specimens, measured in the flesh :—
i Head and body 192 mm.; hind foot 30; ear 19.
¢ a7.
” 192 ” 9 ” OL, 9 19.
I 6 DI »
° (type). ” hee ee) ” pe > 9 ae
” 175 oD ”? Y45 55 21.
Skull ui type—greatest length 44°2 mm.; basilar length 36°6 ;
oy soul breadth 21; length of nasals 15; interorbital breadth
3°5; breadth of brain -case 17°4; palatal foramina 12:3; oblique
diameter of bulle in plane of basioccipital 13°5; length of upper
tooth-series (alveoli) 8°5.
Type locality. Ning-wu-fu, Shan-si. 6000’.
Type. Adult female. B.M. No. 9.1.1.278. Original number
2000. Collected 23 June, 1908
This Pika is very nearly allied to the Siberian and Mongolian
O. dawurica, which it closely resembles in colour, but is dis-
tinguished by its rather larger size, much larger bulls, and broader
molars.
With regard to Ochotona huangensis Matsch., from Western
Kan-su, Prof. Matschie has been good enough to send me the
dimensions of its bulle, and these appear closely to agree with
those of O. dawurica, and are therefore markedly smaller than the
unusually large bulle of O. bedfordi.
T have named this fine and distinct species in honour of His
Grace the Duke of Bedford, K.G., the importance of whose
assistance in the exploration of Eastern Asia is indicated by the
982 MR. OLDFIELD THOMAS ON [ Dec. 15,
fact that the type-specimen is No. 2000 of the Mammals collected
during the Exploration.
“¢ At Yen-an-fu we found these animals about a few bush-grown
burying-grounds in the main valley. They proved to be very shy,
for after: trapping a few we suddenly found we could catch no
more. My belief is that the remainder of the colony deserted the
place. North-west of Ching-pien Mr. Sowerby found Pika-
burrows again, but we secured no additional specimens till we
reached Ko-lan-chow, Shan-si. In this locality we did not find
them common, but in the vicinity of Ning-wu-fu they are fairly
abundant. No. 2000, taken 23 June, contained four young with
hair, and evidently about to be born.
“Chinese names: ‘ Ti-tu’ = Ground- Hare (at Yen-an-fu); ‘Hao-
tu ’= Rat-hare (at Ko-lan-chow).”—W. P. A.
In a previous paper* attention was called to the confusion
which had followed from the giving of two names, O. ogotona and
O. dauwurica, to the same animal by Pallas ; and it now appears that
this confusion has further resulted in the naming of Mr. Lyon’s
subgenera + being inaccurate. The latter had assumed that the
animal described by Waterhouse as Lagomys ogotona was really
Pallas’s species of that name, and consequently called the sub-
genus to which it belonged the typical Ochotona Link, based on
Pallas’s animal. As already noted, however, Pallas’s ogotona
was the same as his earlier dawurica, and this is one of the species
essigme alle to Mr. Lyon’s Conothoa, so that the latter name becomes
a synonym of Ochotona.
The nomenclature of the subgenera would therefore appear te
be as follows :—
1. Ocnorona Link. Type, O. dawurica Pall.
Conothoa Lyon. » QO. roylei Og.
(Incisive and palatal foramina united, open.)
2. Prxa Lac. 1799. Type, O. alpina Pall.
Lagomys G. Cuv. 1800 (nec Storr, 1780).
Type, O. alpina Pall.
(Incisive and palatal foramina separate. Frontal outline
not abnormally bowed.)
3. Ocotoma Gray, 1867. Type, O. pallasi Gray.
(Incisive and palatal foramina separate. Frontal outline
abnormally bowed.)
The last-named also includes O. ladacensis Giinth.
333), OcHOTONA SORELLA.
Thos. Abstr. P. Z. 8. 1908, p. 45 (Dee. 15).
©. 1979. 20 milesS. of Ning-wu-fu, Shan-si. 6600’. 10 June,
1908. B.M. No. 9.1.1.279. Type.
A very small species allied to O. cansa Lyon.
* P.Z.S. 1908, p. 109. + Smiths. Mise. Coll. xlv. p. 438, 1904.
1908. ] MAMMALS FROM NORTHERN CHINA. 983:
Size even smaller than in O. cansa. General colour brown,,
rather darker than Ridgway’s ‘“ broccoli-brown,” a lighter patch
across the nape. Under surface rather lighter, soiled cream-buff,
a more ocnraceous-bufty area down the centre of the belly; the
slaty bases to the hairs showing through; sides of neck more
tawny. Hars blackish grey with white edges. Upper surface of
hands and teet cream-buff, their thickly furred palms and soles.
slaty brownish.
Skull most like that of O. cansa, as figured by Lyon *, but the
upper outline is more convex, the nasals are longer and narrower,
the palatal foramina are more widely open, and the bulle are
markedly smaller.
Dimensions of the type, an adult female, measured in the
flesh :—
Head and body 140 mm.; hind foot 27; ear 18.
Skull—greatest length 36:4 mm.; basilar length 29 ; greatest
breadth 17; nasals 11°8x4; interorbital breadth 4; breadth of
brain-case 14; height of crown from alveolus of m° 10°7 ; diastema
8; palatal foramina 88x43; diagonal length of bulle 9:2;
length of upper tooth-series (alveoli) 6-7.
Hab. and Type as above.
This little Pika belongs to the O. pusilla group, and 1s no doubt.
most nearly allied to Mr. Lyon’s O. cansa, from Kan-su, of which
the Museum possesses a specimen from the Province of Sze-chuen.
It differs, however, in the skull-characters above detailed, and
clearly cannot be referred to it.
“The single specimen was taken by Mr. Sowerby in a wood upon
an abrupt hillside, where this, and probably another, had its.
burrow. The burrows, which were long and intricate, were
subsequently dug up without another specimen being found.
Examination showed No. 1979 to be an adult female, with the
uterus indicating that young had recently been born, and the
mammary glands secreting.”—J/. P. A.
* Smiths. Mise. Coll. 1. pl. xv., 1907.
No. 61.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.*
November 3rd, 1908.
Freperick Giuuett, Hsq., Vice-President, in the Chair,
The SECRETARY read a report on the additions that had been
made to the Society’s Menagerie during the months of June,
July, August, and September, 1908.
Prof. EK. A. Mincuin, M.A., V.P.Z.S., exhibited a series of draw-
ings of Trypanosomes obtained from British Freshwater Fishes.
Mr. Guorrrey Meapz-Watpo, B.A., exhibited a living speci-
men of a Toad that he had obtained in Sumatra, and described
it as belonging to a species new to science.
The Secrerary exhibited a photograph of a young Malayan
Tapir, and remarked that he had been unable to find accurate
drawings of the young of this species. The photograph had been
given to him by the Right Hon. Sir Cecil Clementi Smith, P.C.,
G.C.M.G., M.A., Honorary Member of the Society, and had been
taken from a living example which had been a pet in his house.
A paper was read by Messrs. OLpDFizLD Tuomas, F.R.S., F.Z.S.,
and Guy Dotiman, B.A., on Mammals from Inkerman, near
Townsville, North Queensland, collected by Mr. W. Stalker and
presented to the National Museum by Sir William Ingram, Bart.
and the Hon. John Forrest.
Almost no material from this part of Australia had previously
been received by the British Museum, so that this collection,
which consisted of over 200 specimens, belonging to 26 species,
was of particular value.
* This Abstract is published by the Society at 3 Hanover Square, London,
W., on the Tuesday following the date of Meeting to which it refers. It will
be issued, free of extra charge, to all Fellows who subscribe to the Publications,
along with the ‘ Proceedings’ ; but it may be obtained on the day of publication
at the price of Szxpence, or, if desired, sent post-free fer the sum of Siz
Shillings per annum, payable in advance.
36
The collection clearly showed that the Townsville region
belonged faunistically to North Australia, the species being nearly
identical with those of New South Wales and Victoria. Several
species and subspecies were described as new.
Mr. R. Lyprexnsr, F.R.S., F.Z.S., communicated papers on
Takins from Sze-chuen and Bhutan, and on an Indian Dolphin
and Porpoise.
The next Meeting of the Society for Scientific Business will
be held on Tuesday, the 17th November, 1908, at half-past Hight
o'clock p.m., when the following communications will be made :—
1. N. Annanpats, D.Se., C.M.Z.S.—A new Genus and Species
of Slow-Lemurs from the Lushai Hills, Assam.
2. Prof. G. C. Bourng, D.Sc., F.L.S., F.Z.S8.—Contributions to
the Morphology of the Group Neritacea of Aspidobranch Gastro-
pods.—Part I. The Neritide.
3. RUWENZORI EXPEDITION :——
Report on Mammalia by OLpFTeLtp Tuomas, F.RB.S., F.Z.8.,
and R. C. Wroucuton, F.Z.S.
Report on Fishes. Batrachians, and Reptiles by G. A.
Bou .encer, F.R.S., V.P.Z.S8.
Report on Mollusca by E. A. Surru, I.S.0., F.Z.8.
Report on Lepidoptera Heterocera by Sir Grorce F.
Hampson, Bt., F.Z.8.
Report on Coleoptera: Part I. Lamellicornia, by GILBERT
J. Arrow, F.E.S.; Part II. Buprestide, by C. O.
Warternovss, P.E.S.; Part IIT. Curculionide, by Guy
A. K. Marswatt, F.Z.S.
The following communications have been received :—
1. F. E. Bepparp, M.A., F.R.S., F.Z.8.—Some Notes on the
Muscular and Visceral Anatomy of the Batrachian Genus
Hemisus, with Notes on the Lymph Hearts of this and other
Genera.
2. G. A. Boutuncsr, F.R.S., V.P.Z.8.— Description of a new
Species of Lacerta from Persia. :
3. Dr. Eryarn Lonnpere, C.M.Z.S.— Remarks on some Wart-
Hog Skulls in the British Museum.
4, R. Lypexxer, F.R.S., F.Z.S.— On two Chinese Serow Skulls.
37
5. Prof. ALEXANDER Merrxr, F.Z.8.—The Development of the
Lesser Black-backed Gull, Larus fuscus, L.
6. R. I. Pocock, F.L.8., F.Z.8.—Warning Coloration in the
Musteline Carnivora.
7. Prof. G. O. Sars, C.M.Z.S.—Zoological Results of the
Third Tanganyika Expedition, conducted by Dr. W. A.
Cunnington, 1904—-1905.—Report on the Copepoda.
8. W. T. Catman, D.Sc., F.Z.S.—On a new River-Crab of the
Genus Gecarcinucus, from New Guinea.
Communications intended for the Scientific Meetings of the
ZooOLOGICAL Society or Lonpon should be addressed to
P. CHALMERS MITCHELL,
Secretary.
3 Hanover Square, Lonpon, W.
November 10th, 1908.
ee
(step fie ae
POURS Rega eas
Pie ae
LL Oe
No. 62.
ABSTRACT OF THE PROCEEDINGS
ZOOLOGICAL SOCIETY OF LONDON.*
November 17th, 1908.
Prof. K. A. Mincuin, M.A., Vice-President,
in the Chair.
The Secrerary read a Report on the additions that had been
made to the Society’s Menagerie during the month of October
1908.
Mr. H. E. Austen, F.Z.8., exhibited some living tropical flies,
captured in Manchester, and remarked on the agency of man in
extending the distribution of insects.
Dr. N. Annanpatze, C.M.Z.8., communicated a paper on “ A
new Genus and Species of Slow-Lemurs from the Lushai Hills,
Assam.” The animal is known only from a photograph of two
individuals taken in 1889 by Mr. T. D. La Touche, and resembles
Nycticebus in appearance, but has a long bushy tail, which
distinguishes it from all known Oriental Lemurs.
Mr. G. C. Bourns, D.Sc., F.LS., F.Z.S., Linacre Professor of
Comparative Anatomy in the University of Oxford, presented a
Memoir entitled ‘ Contributions to the Morphology of the Group
Neritacea of Aspidobranch Gastropods.—Part I. The Neritide.”
Asaresult of the comparative study of the anatomy of several
species of marine, estuarine, and freshwater Neritide, the Author
found that the forms hitherto classed in the genera Jeriia,
Neritina, and Septaria fall into three well-marked groups of
subgeneric value, the chief distinctive characters relating to the
* This Abstract is published by the Society at 3 Hanover Square, London,
W., on the Tuesday following the date of Meeting to which it refers. It will
be issued, free of extra charge, to all Fellows who subscribe to the Publications,
along with the ‘ Proceedings’ ; but it may be obtained on the day of publication
at the price of Sixpence, or, if desired, sent post-free for the sum of Si
Shillings per annum, payable in advance,
40
accessory genital organs. The investigation had been pursued by
the study of sections as well as by dissections, and a number of
important anatomical results were set forth. The ganglion on
the visceral loop identified by Lacaze-Duthiers and Bouvier as
sub-intestinal was shown to be the true sub intestinal ganglion,
whilst the ganglion described under that name by Boutan and
B. Haller was shown to be the enlarged right end of an elongate
and diffuse visceral ganglion. The celomic cavity was shown to
be larger in extent in the Neritide than in any other adult
Gastropods, and to be divisible into a left pericardial and a right
garadial division. The macroscopical and microscopical characters
of the accessory genitalia were described in full, and in conclusion
the relationships of the Neritidee inter se were discussed.
RUWENZORI EXPEDITION.
Mr. W. R. Ocitvie-Grant, F.Z.S., gave an account of the
expedition which he had organized to collect in the Ruwenzori
range of mountains in Equatorial Africa. He had obtained
sufficient funds through the generosity of H.G. The Duke of
Bedford, President of the Society, the Earl of Dartmouth,
Viscount Iveagh, the Hon. N.C. Rothschild, Sir Alexander Baird,
Sir Ludwig Mond, Mr. W. A. Bell, Mr. C. Ozarnikow, and
Mr. W. H. St. Quintin, Fellows of the Society, Lord Strathcona
and Mount Royal, the Trustees of the Percy Sladen Fund, and
the Worshipful Company of Fishmongers. It had been agreed
that the first set of specimens collected should be presented to the
British Museum of Natural History. The expedition had been
led by Mr. R. B. Woosnam and had consisted of that gentleman,
Mr. R. E. Dent, the Hon. Gerald Legge, and Mr. Douglas
Carruthers, with Mr. A. F. R. Wollaston as Medical Officer and
botanical collector. The results had been extremely successful,
amongst the specimens obtained having been 404 Mammalia,
2470 Aves, 135 Reptiles and Batrachia, 31 Pisces, and a very
large number of Invertebrates. (
Mr. R. B. Woosnam gave an account of the Expedition
illustrated by lantern-slides.
The following Memoirs on the Zoological Results of the
Ruwenzori Expedition were presented to the Meeting and will
be published in the Scientific Transactions of the Society :—
O. Tuomas, F.R.S., F.Z.S., and R. C. Wroucuton, F.Z.S.—
Report on Mammalia.
G. A. Boutencrr, F.R.S., V.P.Z.S.—Report on Fishes,
Batrachians, and Reptiles.
HK. A. Suirs, I.8.0., F.Z.8.— Report on Mollusca.
A. 8. Hirer, F.Z.5.—Report on Arachnida,
4]
G. J. Arrow, F.E.S.—Report on Coleoptera: Pt. I.
C. O. WaterHouss, Pres.E.8.— Report on Coleoptera: Pt. II.
C. J. GAHAN.— Report on Coleoptera: Pt. IIT.
G. A. K. MarsHatt, F.Z.8S.—Report on Coleoptera: Pt. IV.
The late Col. C. T. Brnenam, F.Z.S.—Report on Hymenoptera.
Sir Grorce Hampson, Bt., F.Z.S.—Report on Lepidoptera
Heterocera.
F. A. Heron.—Report on Lepidoptera Rhopalocera.
E. E. Austen, F.Z.8.—Report on Diptera.
W. L. Distant, F.E.S.—Report on Rhynchota.
W. F. Kirsy, F.L.8., F.E.S.—Report on Neuroptera.
W. F. Kirsy, F.L.S., F.E.S.—Report on Orthoptera.
The next Meeting of the Society for Scientific Business will
be held on Tuesday, the 15th December, 1908, at half-past Hight
o'clock p.m., when the following communications will be
made ;—
1, FREDERICK GILLETT, VERA So ™ NX Hunting-trip to Thian-
Shan, illustrated by lantern-slides.
2. F. E. Bepparp, M.A., F.R.S., F.Z.8.—Some Notes on the
Muscular and Visceral Anatomy of the Batrachian Genus
Hemisus, with Notes on the Lymph Hearts of this and other
Genera.
3. G. A. Boutencer, F.R.8., V.P.Z.8.—Description of a new
Species of Lacerta from Persia.
4. Dr. Ernar Linnsere, C.M.Z.8.—Remarks on some Wart-
Hog Skulls in the British Museum.
5. R. Lypexxer, F.R S., F.Z.8.—On two Chinese Serow Skulls.
6. R. I. Pococn, F.L.S., F.Z.8.—Warning Coloration in the
Musteline Carnivera.
7. W. T. Cauman, D.Sc., F.Z.8.—On a new River-Crab of the
Genus Gecarcinucus, from New Guinea.
8. OupFiELD Tuomas, F.R.S., F.Z.8.—The Duke of Bedford’s
Exploration of Hastern Asia.— XI. On Mammals from the
Provinces of Shan-si and Shen-si, Northern China,
42
The following communications have been received :—
1. Prof. G. O. Sars, C.M.Z.8.—Zoological Results of the
Third Tanganyika Expedition, conducted by Dr. W. A.
Cunnington, 1904-1905.—Report on the Copepoda.
2. Prof. E. A. Mincurn, M.A., V.P.Z.S.—Studies on the
Flagellate Blood Parasites of Freshwater Fishes.
3. T. Goopry, B.Sc.—A further Note on the Gonadial Grooves
of a Medusa, Aurelia aurita.
Communications intended for the Scientific Meetings of the
TZooLOGICAL Socrery oF LonpDON should be addressed to
P. CHALMERS MITCHELL,
Secretary.
3 Hanover Square, Lonpon, W.
November 24th, 1908.
No. 68.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.*
December 15th, 1908.
Dr. Henry Woopwarp, F.R.S., Vice-President,
in the Chair.
The Sscrerary read a Report on the additions that had been
made to the Society’s Menagerie during the month of November
1908.
Mr. Freperick Gituert, V.P.Z.S., gave an account of his
recent Hunting Trip to the Thian Shan, illustrated by lantern-
slides,
Mr. R. I. Pococs, F.L.8., F.Z.8., Superintendent of the Society’s
Gardens, exhibited photographs of a male Sumatran Tiger recently
purchased by the Society, remarking that the Society has at
present, living in the Gardens, examples of three out of four
known races of Tigers.
Mr. F. EH. Bepparp, M.A., F.R.S., F.Z.S., Prosector to the
Society, communicated a paper entitled ‘“‘Some Notes on the
Muscular and Visceral Anatomy of the Batrachian Genus
Hemisus, with Notes on the Lymph Hearts of this and other
Genera.”
Mr. G. A. Boutenesr, F.Z.S., V.P.Z.8., described a “New
Species of Lacerta from Persia.”
A communication was received from Dr. Einar LONNBERG,
C.M.Z.S., “On some Wart-Hog Skulls in the British Museum.”
* This Abstract is published by the Society at 3 Hanover Square, London,
W., on the Tuesday following the date of Meeting to wnich it refers. It will
be issued, free of extra charge, to all Kellows who subscribe to the Publications,
along with the ‘ Proceedings’ ; but it may be obtained on the day of publication
at the price of Stxpence, or, if desired, sent post-free for the sum of Six
Shillings per annum, payable in adyance,
44
Mr. R. Lypexker, F.R.8., F.Z.8., communicated a paper “‘ On
two Chinese Serow Skulls.”
Mr. R. 1. Pocock, F.L.S., F.Z.8., Superintendent of the Society’s
Gardens, read a paper entitled “‘ Warning Coloration in the
Musteline Carnivora,” and exhibited skins of Skunk, Badger, &c.,
to illustrate his argument.
Dr. W. T. Cauman, F.Z.8., communicated a paper ‘On a new
River-Crab of the Genus Gecarcinucus, from New Guinea.”
Mr. Ouprretp THomas, F.R.S., F.Z.8., read a paper on Mam-
mals collected in the Provinces of Shan-si and Shen-si, Northern
China, by Mr. M. P. Anderson, for the Duke of Bedford’s
Zoological Exploration of Eastern Asia. 33 species were included,
represented by 335 specimens, presented, as before, to the
National Museum by His Grace.
The following were described as new :—
ERINACEUS MIODON, Sp. n.
Near H#. dealbatus, but penultimate upper premolar much
smaller, 2 mm. in diameter.
Hab. Yu-lin-fu, Shen-si.
Type. Male. Original number 1871.
ERINACEUS HUGHI, sp. n.
Very dark-coloured, the usual light ends to the spines reduced
to a minute subterminal light band.
Hab. Paochi, Shen-si.
Type. Female. B.M. No. 0.6.27.2.
KUUTAMIAS ASIATICUS ORDINALIS, Subsp. n.
Allied to Z. a. senescens, but much paler throughout.
Hab. Yu-lin-fu, Shen-si; at edge of Ordos Desert.
Type. Female. Original number 1804.
HUTAMIAS ASIATICUS INTERCESSOR, Subsp. n.
Intermediate in general tone between subspp. senescens and
ordinalis.
Hab. Ning-wu-fu, Shan-si.
Type. Female. No. 1972.
CITELLUS MONGOLICUS UMBRATUS, subsp. n.
General colour much darker than in true mongolicus, near
isabella” of Ridgway.
Hab. Taboul, Mongolian Plateau.
Type. Male. B.M. No. 8.3.5.5.
45
Mus CONFUCIANUS LUTICOLOR, subsp. n.
Very pale. Size smaller than in the Shantung JZ. c. sacer.
Hab. Yen-an-fu, Shen-si.
Type. Male. No. 1689.
CRICETULUS TRITON INCANUS, subsp. n.
Paler and clearer grey than in true C. triéon. Skull narrower.
Hab. Near Ko-lan-chow, Shan-si.
Type. Female. No. 1945.
CRICETULUS BEDFORDI, Sp. n.
Clear drab-grey, with pure white uncer surface. Tail very
short. Feet hairy. Head and body 77 mm.; tail 12; hind
foot 12.
Hab. Yu-lin-fu, Shen-si.
Type. Male. No. 1861.
MICROTUS INEZ, sp. 1
Pinkish brown above, wood-brown below. Head and body
90 mm.; tail 35; hind foot 15-5. Molars with the triangles
mostly closed.
Hab. Near Ko-lan-chow, Shan-si.
Type. Female. No. 1892,
LEPUS SWINHOEI SUBLUTEUS, subsp. n.
General colour pinkish buff, much paler than in true swinhoev.
Hab. Southern Ordos Desert.
Type. No. 1735.
OcHOTONA BEDFORDI, Sp. n.
Allied to O. dawurica, but larger and with much larger bulle.
Skull, greatest length 44-2 mm.
Hab. Ning-wu-fu, Shan-si.
Type. No. 2000.
OcHOTONA SORELLA, sp. Nn.
Near O. causa, but even smaller. Skull length 36-4 mm.
Hab. Near Ning-wu-fu, Shan-si.
Type. No. 1979.
The next Meeting of the Society for Scientific Business will
be held on Tuesday, the 12th January, 1909, at half-past Hight
o'clock p.M., when the following communications will be
made :—
1. Prof. G. O. Sars, C.M.Z.S.—Zoological Results of the Third
Tanganyika Expedition, conducted by Dr. W. A. Cunnington,
1904-1905.—Report on the Copepoda.
AG
2. Prof. E. A. Mincuin, M.A., V.P.Z.8.—Studies on the
Flagellate Blood Parasites of Freshwater Fishes.
3. T. Goopry, B.Sc.—A further Note on the Gonadial Grooves
of a Medusa, Aurelia aurita.
4. A. E. Brown, D.Sc., C.M.Z.S.—The Tuberculin Test in
Monkeys; with Notes on the Temperature of Mammals.
5. Prof. R. Cotterr, F.M.Z.S.—A few Notes on Balena
glacialis and its Capture in recent Years in the North Atlantic by
Norwegian Whalers.
The following communications have been received :—
1. F. Woop-Jonss, B.Sc., F.Z.8.—The Fauna of the Cocos-
Keeling Atoll.
2. Grousse-DiseAse Commission Reports:
(a) A. E. Suiprny, M.A., D.Sc. F.R.S., F.Z.8.—The
Ectoparasites of the Grouse.
(6) A. E. Ssipuey, M.A., D.Sc. F.RS., F.Z.5.—The
Thread-Worms (Nematoda) of the Red Grouse (Tetrao
scoticus).
_ (c) A. E. Surprey, M.A., D.Sc., F.R.S., F.Z.S.—The Tape-
Worms (Cestoda) of the Grouse. Appendix: Parasites of
Birds allied to the Grouse.
Communications intended for the Scientific Meetings of the
ZOOLOGICAL Society or Lonpon should be addressed to
P. CHALMERS MITCHELL,
Secretary.
3 Hanover Squareg, Lonpon, W.
December 22nd, 1908.
ConTENTS (continwed).
December 15, 1908.
Page
The Secretary, Report on the Additions to the Society's Menagerie during the month of
PionpRtaer BOOS Pal ae site n awe 8 i vs ¥ scraa eee wees sales tare mingle ee share <P teh om ale eiaage =. 890
‘Mr. Frederick Gillett, V.P.Z.S. Note on a Hunting Trip to the Thian Shan ..........., 890
“Mr. R. I. Pocock, F.L.S., F.Z.S. Exhibition of photographs of a Sumatran Tiger, and
remarks upon this animal and upon other Tigers living in the Socviety’s Gardens ..... . 890
1. Some Notes on the. Muscular and Visceral Anatomy of the Batrachian Genus Hemisus,
with Notes on the Lymph-Hearts of this and other Genera. By Frank E. Bepparp,
OEE Sey EA wee aieje we hon kote ovine cn eye sialece seen cen bums otulwsie es atadine 80+
2, Description of a new Species of Lacerta from Persia. By G. A. Bou.encsr, F.R.S.,
ems mA ace MEE ee wi cod sane ea ee tise 28 Wisie a e's a aw lely nesta, obgi oul sin sls eins 93-4
3. Remarks on some Wart-Hog Skulls in the British Museum. By Dr. Eivar Loyypure,
NSN cea Cesta ole basic aie o’o\e'aeve) va eid aihveia sas Atohaiwie: hai shared rim proleaier ate 936
4, On Two Chinese Serow-Skulls. By R. LypEKKER ........-+..- Ber avi onaicis'e fi aisienen ata 940
” ;
5. Warning Coloration in the Musteline Carnivora. By ‘R. I. Pococs, F.L.S., F.Z.S.,
Superintendent of the Zoological Society’s Gardens ..........-- bon damearie ofevensrenete 4+ ©
6. On a new River-Crab of the Genus Gecareinucus from New Guinea. By W. T. Cauman,
D.Sc., F.Z.S., British Museum (Natural History). (Plate LXVIII.)..........+.-..- 960
7. The Duke of Bedford’s Zoological Exploration in Eastern Asia—XI, On Mammals from
the Provinces of Shan-si and Shen-si, Northern China. By Ouprienp Tuomas, J*.R.5.,
Ath fa icts eave akccta wis ees wee e cece ete e ete en eens cece tteeseaneeens Seas MS se DOS
MHGe pase veces cee recs cc cece enter ste ere tere meen mseren ese re see h teva ere ce derend Le
List of Cosine AUELCLa VERT OE nS emetic. scare. coin peehave e-alefcle iin let 9 olovavsnere fava Noli nid SA 5 te es ae il
List of MBER TRE CERES AR Sea aes) Oe Me ae a at Sm ee dats ees et ee Salo! ai oadeler oan eteeey shea nian ci eTsl ca chess wove el jit
Alphabetical TDS RCN EA Olay MMW LST THe 1: RPP heir eeiaay EAS SC ceresr COS MIrOn oom ec mm ucir unt) Si On aCe ix
Theta? Jen SESS Nee Ae Ree DBE n OOo imCon hn Ob.cp mre rc aia oS Oca Oce XVil
List of Text-figures -. 6.2... secre eee ae : Sh Sere atans a Tae Tome, lot Oana wee user at xix
HIRO NENV: GeenleriG LENISi cicis! uiess's' 2 dbus mls cittaielersin/ ple. sueelelennte; s slisin alvync/s's sla, Sede anaes re RADE
fyyn-t2 tls
LIST: OF PLATES. |
1508, pp. 783-983.
Pate > nl ; Pare ao
XLI. Bufo valhaile, sp. nov. 1. Upper surface of head. 2. Un-
derside of foot <2. s< <4 isenicee Genes Oe pte ae ea
XLII. Macropus ualabatus ingramt........ pyc Ste evahate eyoy eat te 183°
XLII. The Chinese Takin (Budorcas tibetanus), 1 $,2Q....-.-. 195
XLIV. 1. Tursiops dawsont, 2. Neophocena sp, ++-.++ .++++++++} ong
XLV. Sotalia lentiginosa ...-cceeceee wees seer ees boeiak oie f 802
XLVI. Septaria borbonicd .. 000. oe cece sees ets pete en eee )
XLVIT. Ccelom and oviduco-celomic funnel of Septaria ....... «. |
Sv |
XLV Le | Genital complex of Septaria bougainvillei Q ...... «++. |
KEEXS
L. Female ducts and csphradium of Septaria ....--+....+5-
GL... Kidney ‘af Septarta depressa: <i xian’. «\ne vin none eiodiete hex |
LIT, Gill-lamellee ete. of Seplaria «0... eee eee ee eee ee eee
LIIf. Histology of genital ducts of Seplavia 225.2241 ee |
LIV. Anatomy of Nerita, Paranerita, and Neritinad .,..+-..+..-
LY. Alimentary tract and nervous system of Parancrita........ |
LVI, Anatomy and Histology of Neritide ........... aren eae r 810
LVIL. Paranerita gagates ...++.....002..serere-s MOWIAe Seek Soo
ee Paranerita variegata and GOGOLES, xu 3.x) sia) +2) 5d ene ee ®
LX. Male ducts of Nerita and Paraneriia ....+-+-......+++.0
LXI. Male ducts of Paranerita ...... Sate ials SLOete + .ofen a omit ee re
LXII. Anatomy and Histology of Neritide ..........0.s.c0e:
LXIII. Reproductive organs of ParaneritaQ .. So. eae
LXIV. Female organs of Paranerita ...0ve.. esse nee ee eee eens
= | Reproductive organs of Nerita plicata Q ..... eet oc )
LXVII. Lacerta chlorogaster ...+..+ ho ee a eee 934
LX VIL; © GQecarcinueams an Gr aiid ic op cw siapasernsejn\e)o ss) aintelole etateeieteele eLeeanae 960°
NOTICE.
The ‘Proceedings’ for the year are issued in four parts, paged consecutively,
so that the complete reference is row P. Z. 8. 1908, p... . The Distribution
is as follows:—
Papers read in January and February, issued in June.
5 » March and April, oy just.
ee » May and June, » 95 October,
% » November and December,,, ,, April.
‘ Proceedings, 1908, pp. 431-782, were published on Oct. 30th, 1208,
The Abstracts of the papers read at the Scientific Meetings in
November and December are contained in this Part.
Pp 970"
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