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oH A Ge BD Ae a Sh ge 00d sot Fs PIE WA he Ry 028s Gat GOT 9 aN HE 9: fe 

a cones Np teddob 14k GH Wien ged ed, Crmarecer ee kote ee rst vf 
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aA 


TEROCHIDIN GS 


OF THE 


GENERAL MEETINGS FOR SCIENTIFIC BUSINESS 


OF THE 


ZOOLOGICAL 


OF LONDON; 


1922, pp. 483-1276, 


witH 36 PLATES AND 232 TEX'l-FIGURES. 


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


LONDON: 


MESSRS. LONGMANS, GREEN, AND CO, 


PATERNOSTER ROW, 


SOCLETY 


OF THE 


\ 
ales Qk | 
‘ ipa 


COUNCIL AND OFFICERS. 


OF THE 


ZOOLOGICAL SOCIETY OF LONDON. 


1922. 


Patron. 
His Masgesty THe Kina. 


COUNCIL. 
His Grace Tae Duke or Beprorp, K.G., F.R.S., Presideni. 
Tur Hon. Cecit Barine, M.A. | Cou. Sir A. Henry McManon, 


Lr.-Cou. 8. Monckron Copr- G.C.M.G., K.C.I.E., Vice- 


MAN, M.D., F.R.S. President. 
CHARLES Drummonp, SEsq., EK. G. B. Mnapr-Watpo, Esa. 
Treasurer. P. Coatmers MircHens, Esq., 


ALFRED Ezra, Esq., O.B.E., C.B.K., M.A., D.Sc., LL.D., 


Vice-President. E.R.S., Secretary. 
Huew S. Guapsronze, Hsq., | THe Ear or Onstow, O.B.E. 
M.A., F.R.S.E. Masor Aupert Pam, O.B.H. 
Tse Rr. Hon. Tue Viscount | Tur Lorp QUEENBOROUGH. 
Grey, K.G., P.C. THE Marquess oF SLIGO, 
Sir Srpney F. Harmer, K.B.B., |: F.S.A., Vice-President. 


M.A., D.Se., F.R.S., Vice- | Pror. G. Eiuiot Smiru, M.A., 
President. | M.D., F.R.S. 


Masor THe Lorp Aastarr | RicHarp 8. Tayior, Esq. 


Rosert Innes-Ker, D.8.O. AntHony H. WINGFIELD, Esq. 
Pror. Ernest W. MacBript, | A. Smita Woopwarpb, Ksq., 
D.Se., LL.D., F.R.S., Vice- | LL.D., F.R.S., Vice-Pre- 
President. sident. 


PRINCIPAL OFFICERS, 


P. Cuaumers MircHety, ©.B.i., M.A., D.Sc., LL.D., F.RB.S., 
Secretary. 

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

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

Epwarp G. BouLencEr, Curator of Reptiles. 

Miss L, E. Curesman, F.E.S., Curator of Insects. 

C. F. Sonntac, M.D., Anatomist. 

N.S. Lucas, M.B., Ch.B., Pathologist. 

R. J. Orrierp, M.A.. Hon. Parasitologist. 

Prof. G. H. Wootpriner, F.R.C.V.S., Hon. Consulting Veterinary 
Surgeon. 

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

F. W. Bonn, Accowntant. 

W. H. Cour, Chief Clerk. 


f 


: P Ve 
I N a WAAOLS ‘ 


LIST OF CONTENTS. 


1922, pp. 483-1276. 


EXHIBITIONS AND NOTICES. 


THE SEcRETARY. Remarks upon a Medal struck to com- 
memorate the 150th Anniversary of the Royal 
Nea denny Ol ele iumnal iene ce caten War nanan antec menue 


Mr. F. Martin Duncan, F.Z.S. Exhibition of cinemato- 
graph films illustrating the life-history of the Wood 
Ant (Formica rufa) and Common Wasp (Vespa 
GCPROGHOUGTA) Sip SROs Me! Satake Ren Le BL AAR aber Cra re 


Dr. Cuas. F. Sonnvac, F.Z.8. Exhibition of specimens 
and lantern-slides illustrating points in the anatomy 
andephvsiolagy) ob! Whales 222502204. econ 


The Srcrerary. Report on the Additions to the Society’s 
Menagerie during the month of April, 1922............ 


Mr. R. I. Pococr, F.R.S., F.Z.8. Exhibition of a living 
example of the rare Bush Dog, Speothos venaticus ... 


Mr. EK. G. BouLencsr, F.Z.S.,and Mr. F. Martry Duncan, 
F.Z.S. Exhibition of cmematograph record illus- 
trating the life-history of the Axolotl (Amblystoma 
LU GRITC LID Nada 8 se eles) Sere MOPS As ga eoa eS Aae dle UnUelaies ec ates 


My. D. Sers-Suira, F.Z.S. Exhibition of photographs of 
recent important additions to the Society’s collection 
OSES IEC SMa ea RMR Na AR CRS eA est ibe ae sacar ie tule 


Rev. H. N. Hurcninson, M.A., F.Z.S. Exhibition of a 
plaster-cast model reconstruction of the marine reptile, 
TPIT ORUIRTAS VOOR OOS phn SOB oe BU ORE an AHO ORE ENN ORG R ALE? 


Page 


833 


833 


833 


833 


834 


834 


834 


iv 


The Secretary. Report on the Additions to the Society’s 
Menagerie during the month of May, 1922 ............ 


The Secrerary. On the Council’s scheme to establish an’ 


ANOUGHEIONIN ,atasnpnboosdbabae aanboneengos ogobonodaonoRuaennad sos 
The Secrerary. Report on the Additions to the Society’s 
Menagerie during the months of June, July, August, 
enna Sry owenin sie WGTUPA" Soap scuneevbonsodnboccognocs sApanooacds 
The Secrerary. Exhibition of photographs of Zebras and 
Onyvactirom lem yarn chew ee nie ccin tne ei acicis) Weccarasns 
Mr. E. T. Newton, F.R.S., F.Z.S. Exhibition of tanned 
Seago peu GEO ON tN eON ie eee ae lato stant meeps ssi occa eae 
Mr. C. Tare Ruean, F.R.S., F.Z.8. Exhibition of specimens 
OL lenthermmadetrompolarics pskanisi ye ee nee tcc: 
Prof. G. Eitior Smiru, M.D., F.R.S., F.Z.S. Exhibition 
of photographs of a fossil tooth (flesperopithecus) from 
ING loa shee N Eten TE Ss a i ee elena Rie. Suna es eetee 


Dr. A. SmirH Woopwarpb, F.R.S., F.Z.S. Exhibition of 
drawings and photographs comparing teeth of a Bear, 
Chimpanzee and Hesperopithecws |)...\c assesses eee 

The Srcrerary. Report on the Additions to the Society’s 
Menagerie during the month of October, 1922......... 

Prof. EK. W. MacBrips, F.R.S., F.Z.8. Exhibition of 
lantern-slides bearing upon Dr. Kammerer’s experi- 
OVSLANAS) COVAL, LATO) AT OVER Mv tidbatababadongospadoadadsncdsuaode, 6 

Dr. A. SmrrH Woopwarp, F.R.S., F.Z.S. Exhibition of a 
skull and tusks of a Mammoth from Siberia............ 

Mr. D. Sera-Smiru, F.Z.8. Exhibition of the shed lining 
of the gizzard of a Hornbill 


eee oes cer ces esse se ere esses eee 


1276 


1276 


26. 


29. 


30. 


3l. 


32. 


33. 


34. 


30. 


36. 


At 


PAPERS. 


A Study of the Remarkable Tortoise, Testudo loveridgii 
Blgr., and the Morphogeny of the Chelonian Carapace. 
By Joan B. Procrer, F.Z.8. (Plates I.-II1.; Text- 
figures 1-21 ; and Table.) 


Ce en 


. Notes on the Anatomy of Cacopus systoma, an Indian 


Toad of the Family Hngystomatide. By D. W. 
Devanesen, M.A. (Text-figures 1-19.) 


. A Review of the Cetacea of the New Zealand Seas.—I. 


By W. R. B. Ontver, F.L.8.,F.Z.8. (Plates I.-IYV.). 


On the Dental Characters of certain Australian Rats. 
By Prof. F. Woop Jonss, M.B., B.Sc., F.Z.S. (Text- 
MUUICOS MND), asset ak eivoua Lumtisctyees ay rbonaaaseercmr Mes 

On the Structure of the Enamel in the Primates and 
some other Mammals. By J. Tuornron Carrer, 
Bee Sen n( Plates Ac V LUNG: ic nasa latd Wee oa tics koe 


Miocene Proboscidia from Baluchistan. By C. Forsrer 
Coopmr, M.A., F.Z.8. (Plates 1-1V.; Text-figures 
Tee eee cece en le ate sts aersciuteasactinle « alioa/sme sure aclaicitbe Marte 


On Commerson’s Dolphin and other Species of Cephalo- 
rhynchus. By Sir 8. ¥. Harmer, K.B.E., F.R.S., F.Z.8. 
(Lelie meicy feted ld BLA even dg Se ae RAM Mine sh rir DEG ROR eda) 


The Comparative Anatomy of the Tongues of the 
Mammalia.—VII. Cetacea, Sirenia, and Ungulata. 
By Cuartrs F. Sonnrac, M.D., F.Z.8. (Text- 
TUSURESEZO HOON) meters, (atin Me cme iaNuas Wan ietie st. Jat th Aik 


The External Characters of Scartwrws and other 
Jerboas, compared with those of Zapus and fedetes. 
By R. I. Pococx, F.R.S.,F.Z.S. (Text-figures 29-37.) 


A Revision of the Isopod Genus Ligia (Fabricius). By 
Haroxp G. Jackson, M.Sc., F.Z.8.. (Plates I. & IT.). 


Monograph of the Hymenopterous Family Stephanide. 
By Ernest A. Exuiort, F.L.8., F.Z.5. (Text-figures 
ela) sb ebueect Bon bor be tie ter ad Ncnn Pac RAC anNRG Phin Erie Anon Rea 


Page 


483 


587 
599 
609 


627 


31. 


38. 


39. 


40. 


Al. 


43. 


44, 


45 


Vv! 


Notes on Hast African Birds (chiefly nesting habits and 
stomach contents) collected 1915-1919. By Arrnur 
TOVEREDCE, WEES s0ONVIEZAS) later en ee nnel coe peer aieme 


On the Myology and Classification of the Wombat, 
Koala, and Phalangers. By CHanrtes F. Sonnrac, 
M.D., F.Z.8. (Anatomist to the Society). (Text- 
HURSADN erst) EE US Rees «KA Rae eeu Me hb 


Experimental Evidence that Commensalism may be 
beneficial to Crustacea. By Epwarp B. Povutron, 
D.Se., F.R.S., F.Z.8., Hope Professor of Zoology and 
Fellow of Jesus College in the University of Oxford. . 


Description of a New Lizard of the genus Chalcides, 
from the Gambia, living in the Society’s Gardens. 


By HE. G. Boutencer, F.Z.8. (Curator of Reptiles) ... 


On the Pavasitic Nematoda collected from Mammalian 
Hosts which died in the Gardens of the Zoological 
Society of London during the years 1919-1921 ; with 
a description of three new Genera and three new 
Species. By G. M. Vuvmrs, M.R.C.S., L.R.C.P., 
F.Z.S., Beit Memorial Research Fellow, and Assistant 
in the Department of Helminthology, London School 
of Tropical Medicine. (Text-figures 1-10.) ............ 


On the Cestode Parasites from Mammalian Hosts which 
died in the Gardens of the Zoological Society of 


London during the years1919-1921; with adeseription ~ 


of a new Species of Cyclorchida. By G. M. VEverrs, 
M.R.C.8., L.R.C.P., F.Z.S., Beit Memorial Research 
Fellow, and Assistant in the Department of Helmin- 
thology, London School of Tropical Medicine. (Text- 
BAO TERS ET VARS ASG ra ARAM IN BAUS OAR ey ec 


On a further Collection of Mammals from the Inner 
Hebrides. By Ivor G. 8. Monracu, F.Z.S. (Text- 
TOKO DT ge) ED) nea mee aR ANI AA ISL chs MN LS 


Two new Indian Species of the little-known Genus 
Aulodrilus Bretscher of the Aquatie Oligocheta 
belonging to the Family Tubificide. By H. R. 
Menara, M.Sc., Zoological Department, Benares Hindu 
University. (From the Zoological Laboratory, Cam- 
bridge.) (Plates ].—I1I.; Text-figures 1-9.)............ 


A Contribution to the Anatomy of a Hammerhead 
Shark (Zygena maiieus Shaw). By J. H. Luoyp, 
M.Sc. (Birm.), F.Z.8., and Eprrw M. SuepparD, B.Sc. 
(Wales), F.Z.8., Zoological Department, University 
College, Cardiff. (Text-figures 1-7.) .................. 


Page 


837 


863 


897 


899 


901 


929 


943 


46. 


47. 


48. 


49. 


50. 


5]. 


53. 


vil 


On an Instance of Commensalism Horweon a Hermit 
Crab and a Polyzoon. By R. Kirxparricr, F.Z.S., 
and Dr. J. Mrerzenaar. (Plates I. & IT.)............... 


New Species of Trinidad Moths. By W. J. Kays, 
HPI Sima (alenbep i yawn nasty oh Mean eas cur cssceas can sdneeadd 
The Nematode Genus Physaloptera Rud. By R. J. 
Ortiterp, M.A., Research Assistant, Dept. of 
Helminthology, London School of Tropical Medicine ; 
Hon. Parasitologist to the Society 1921-1922. (Text- 
ATU CSM AA ede eats See ape dnt eae eM uct laa ucle sutras 


The Oligocheta of the Oxford University Spitsbergen 
Expedition. By J. StepHenson, M.B., D.Sc., F.Z.S., 
Lecturer in Zoology, Edinburgh University. (Text- 
dOUIEES LG. ree oe IN RN ei ce RE i ae eae 3 


On the Anatomy of Scaphula Benson, with a description 
of a new Species. By ExENDRANATH GuosH, M.Sc., 
M.D., F.Z.8., F.R.M.S., Professor of Biology, Medical 
College, Calcutta. (Text-figures 1-7.) .................. 


The Foetal Membranes and Placentation of Chiromys 
madagascariensis. By Prof. J. P. Hitu, F.R.S., and 
R. H. Burne, M.A. (With an Appendix on the 
External Characters of the Feetus, by R. I. Pococr, 
F.R.S.) (Plates 1—-V1I.; Text- figures 1) rene 


. On the External Characters of the Beaver (Castoride) 


and of some Squirrels (Scturidw). By R. I. Pococr, 
FE.RB.S., F.Z.8. (Text-figures 38-60.) .................. 


On the Morphology of the Chondrocranium of the 
Larval Herring (Clupea harengus). By F. R. WELLs. 
(Qextoicumesel lila )ickek tusaensnee eae cat eter teased 


. On the Colours of Water-Mites. By C. S. Etron, New 


Colle sem Oraondisarm sense emai tec cess uch stlvae sores oe 


. Notes on certain Crossopterygians. By Errk A: son 


Stens1o, Upsala. (Plate I.; Text-figures 1-6.) ...... 


Page 


983 


991 


999 


1109 


1139 


1145 


1171 


ALPHABETICAL List. 
OF THE: 
CONT RI BUTO KS, 
With References to the several Articles contributed by each. 


(1922, pp. 483-1276.) 


Page 
Boutencrr, H.G., F.Z.8. (Curator of Reptiles). 
Description of a new Lizard of the genus Chalcides 
from the Gambia, living in the Society’s Gardens ......... 899 
Boutencer, E. G., F.Z.S., and Duncan, F. Marmin, F.Z.S. 
Exhibition of cinematograph record illustrating the 
life-history of the Axolotl (Amblystoma tigrinum)......... 834 
Burne, R. H. See Hiut, J. P. 
Carter, J. Toornion, F.Z.S. 
On the Structure of the Enamel in the Primates and 
some other Mammals. (Plates I.-VII.)........... Wes d99 
Cooprr, C. Forsrrer, M.A., F.Z.S. 
Miocene Proboscidia from Baluchistan. (Plates I.-IV.; 
Merete fore ss An) cak wh OM sa tae) Mt LaAUS iene lee Net el) 609 


DEvVANESEN, D. W., M.A. 


Notes on the Anatomy of Cacopus systoma, an Indian 


Toad of the Family Engystomatide. (Text-figures 1-19.) 5 


ii 


é 
1x 


Duncan, F. Martin, F.Z.S. (Librarian to the Society). 
Exhibition of cinematograph films illustrating the 
life-history of the Wood Ant (Yormica rufa) and Common 
IISA UCSMQNGEINUGII LC IVE se seh oh ar wotiste Wee tain ee ot -biclan 833 


Duncan, F. Marvin. See Bouteneer, HE. G., F.Z.8. 


Ex.uiorr, Ernest A., F.L.S., F.Z.8. 
Monograph of the Hymenopterous Family Stephanide. 
(iesrehomimes l= 82) ohio re ay eee ee nN Sei 705 


Exton, C.5. 
OnitherColourszotm Weavers Vitesse nesseee eee eee 2S 


GuosH, EkenDRANATH, M.Sc., M.D., F.Z.S. 
On the Anatomy of Scaphula Benson, with a description 
Ouamew Species: (Next-ngures V7. cence eee: 1139 


Harmer, Sir 8. F., K.B.E., F.R.S., F.Z.8. 
On Commerson’s Dolphin and other Species of 
Ocpratorhynchus. (lates WI eee wane seri oben er 627 


Hi, J. P., F.R:S., Burne, R. H., and Pocock, R. I., F.R.S. 
The Fetal Membranes and Placentation of Chiromys 
madagascariensis. (Plates 1-IV.; Text-figures 1-6.) ... 1145 


Hurcuinson, Rev, H. N., M.A., F.Z.8. 
Exhibition of a plaster-cast model reconstruction of the 


marine reptile, Peloneustes philarchus ............0ccee essen 834 


Jackson, Haroup G., M.Sc., F.Z.S8. 
A Revision of the Isopod Genus Ligia. (PlatesI.-III.) 683 


Jones, F. Woop, M.B., B.Sc., F.Z.8. 
On the Dental Characters of certain Australian Rats. 
(eset trom eel ase fa ee nies siteeic\e iol daeyaale miieinin 587 


Kaye, W. J., F.E.S. 
New Species of Trinidad Moths. (Plate I.)............ oon 


x 


Kirkpatrick, R., F.Z.S., and Mrrzevaar, Dr. J. 


On an Instance of Commensalism between a Hermit 
Crab andyaybolyzoousg (elaiesi ills dy alls)\ne aaa sean 


Lioyp, J. H., M.Sc. (Birm.), F.Z.S., and Saepparp, Epirn 
M., B.Sc. (Wales), F.Z.8. 


A Contribution to the Anatomy of a Hammerhead 
Shark (Zygena malleus Shaw). (Text-figures 1-7.)...... 


Loveripes, Artuur, F.E.S., F.Z.S. 


Page 


983 


971 


Notes on East African Birds (chiefly nesting habits - 


and stomach contents) collected 1915-1919.................. 


MacBripe, Prof. KE. W., F.R.S., F.Z.S. 
Exhibition of lantern-slides bearing upon Dr. 
Kammerer’s experiments on Amphibia ..................... 


Meura, H. R., M.Sc. 

Two new Indian Species of the little-known Genus 
Aulodrilus Bretscher of the Aquatic Oligocheta belonging 
to the Family Tubificide. (Plates I—III.; Text-figures 
ISO) Sse aes er ere a err cee ae tiey geod er aM ne 


Merzeiaar, Dr. J. See Kirnxparricr, R., F.Z.S8. 


MircHei, Dr. P. CHAtmErs, C.B.E., M.A., D.Sc., LL.D., 
F.R.S. (Secretary to the Society). 

Remarks upon a Medal struck to commemorate the 

150th Anniversary of the Royal Academy of Belgium ... 


Report on the Additions to the Society’s Menagerie 
lumens telove) soavovatel mn Ont yore) NOL sos ibe ecco ndoaoncconocasusaa: 


Report on the Additions to the Society’s Menagerie 
cumin eichenmont hone aya iO 22g eaen en ne Cen ean ieee 


On the Council’s scheme to establish an Aquarium 


Report on the Additions to the Society’s Menagerie 
during the months of June, July, August, and September, 
1922 


837 


1276 


943 


Xi 


Mircuett, Dr. CHALMERS (cont.). 


Exhibition of photographs of Zebras and Oryx from 


Report on the Additions to the Society’s Menagerie 
durmentne month ot@ctober, NO22) yy ee a teases 5. 


Monragau, Ivor G. S., F.Z.S. 
On a further Collection of Mammals from the Inner 
ielebnidest) (Rext=toume Wy) eer Seite te yc eee ives sare slcsinie ae 


Newron, EH. T., F.R.S., F.Z.8. 


Exhibition of tanned skin of a F MO OU saree sale senate 


Outver, W. R. B., F.LS., F.Z.S. 
A Review of the Cetacea of the New Zealand Seas.—I. 
(TRIGNIES IWIN he bebna Soe ecso Ea eBen SenboaA HonaeuERESoUnoROscapcs 


Ortiepp, R. J., M.A. 


The Nematode Genus Physaloptera Rud. (Text-figures 
UALR) Poe yaea va aca sara Maes als Wet SCY alter Tate NEIL MAE er 


Pocock. Ryde HRS: REZiS. 
The External Characters of Scarturws and other 
Jerboas, compared with those of Zupus and Pedetes. 
(Mext=fsumes! 29 3nd saan aeener tance atest me Sh anaes 


Exhibition of a living example of the rare Bush Dog, 
DS PEOUNOS VEMOLICUS Meine Wave ANN e NEUSE Ne dart lnts| elsio)tenle elas a 


On the External Characters of the Beaver (Castorid@) 


and of some Squirrels (Sciwridew). (Text-figures 38-60.) 
Pocock, R. I. See H1u1, J. P., and Burne, R. H. 


Poutton, Epwarp B., D.Sc., F.R.S., F.Z.8. 
Experimental Evidence that Commensalism may be 


loeroelaroney! TK) (COEUISTRKGSE,  dodbdsoodatocacaceaue dodesussebooboeennous 


929 


1275 


DoT 


999 


659 


834 


1171 


897 


xll 


Page 
ProctEr, Joan B., F.Z.S. 

A Study of the Remarkable Tortoise, Testudo loveridgit 
Blgr., and the Morphogeny of the Chelonian Carapace. 
(Plates I.-III.; Text-figures 1-21, and Table.)............ 483 

Reean, C. Tarts, M.A., F.R.S., F.Z.S. 

Exhibition of specimens of leather made from Sharks’ 

SEDANS or Saar 8 Re nRepA apna AU a eS aE Mee HELE Bre 1275 
Seru-Smirh, D., ¥.Z.S. (Curator of Birds). 

Exhibition of photographs of recent important additions 
to the Seciety’s collection of Birds ........................... 834 

Exhibition of the shed lining of the gizzard of a 
Hetorpmilor li Wes eset iss niotlthe sora al att beece ths harms NECN SA OR 1276 

SHEPPARD, EpirH M., B.Sc., F.Z.S. See Luoyp, J. H., M.Sc., 
E.Z.S. 
SmitH, G. Evuior, M.D., F.R.S., F.Z.S. 

Exhibition of photographs of a fossil tooth (Hespero- 

pithecus) trom: Nebraska! sci.) ssccdsescedsersedaiie seiesee, 1275 
Sonnac, Cras. F., M.D., F.Z.8. (Anatomist to the Society). 

The Comparative Anatomy of the Tongves of the 
Mammalia.—VIT. Cetacea, Sirenia, and Ungulata. 
(ext omouTes 25 OMe WNir umes oe ners oN Aa eo mn cau el 639 

Exhibition of specimens and lantern-slides illustrating 
points in the anatomy and physiology of Whales ... ..... 833 

On the Myology and Classification of the Wombat, 
Koala, and Phalangers. (Text-figures 31-43.) ............ 863 

Srensi0, Erik A: son. 

Notes on certain Crossopterygians. (Plate I.; Text- 

(DOL ES TIES ORV GHA agree aie Ger anaD ranean Beanie a inreta | Sen SINE 6 1241 


STEPHENSON, J., M.B., D.Sc., F.Z.S. 
The Oligocheta of the Oxford University Spitsbergen 
Ese diiron. a) (dlhex ttl onices il G.)\ien mreanerat see eens eee 1109 


xiii 
: Page 
Vevers, G. M., M.R.C.S., L.R.C.P., F.Z.S. 


On the Parasitic Nematoda collected from Mammalian 
Hosts which died in the Gardens of the Zoological Society 
of London during the years 1919-1921; with a descrip- 
tion of three new Genera and three new Species. (Text- 
HU GUUICC SPM JAM cra ea a) Secs seraee pec au emeet Ne Na etme are MON Cad ves 901 
On Cestode Parasites from Mammalian Hosts which 
died in the Gardens of the Zoological Society of London 
during the years 1919-1921; with a description of a new 
Species of Cyclorchida. (Text-figures 1] & 12.)............ 921 


WE LLs, F. R. 


On the Morphology of the Chondrocranium of the 
Larval Herving (Clupea harengus). (Text-figures 1-11.) 1215 


Woopwarp, Dr. A. Smiru, F.R.S , F.Z.S8. 
Exhibition of drawings and photographs comparing 
teeth of a Bear, Chimpanzee, and Hesperopithecus ......... 1275 


Exhibition of a skull and tusks of a Mammoth from 
Siilloe ra area arash oleracea GaN UR EM IN Aaah AL GN ait a Canc 1276 


INDEX OF ILLUSTRATIONS, 


Allactaga indica, Fig. 34, p. 672. 
Amia, Fig. 11, p. 1227. 


Ancylostoma pluridentatum, Figs. 1, 2, 


p. 904. 

Apodemus sylvaticus, Fig. 32, p. 668. 

Atlantoxerus getulus, Wigs. 43, 52, 
pp. 1180, 1196. 

Aulodrilus kashi, Pls. 1., If., p. 948, 
Higs. 1-7, pp. 947, 949, 950, 955, 
959, 962. 


8, 9, pp. 963, 965. 


Australian Rats, fragments of jaws, | 


Figs. 1, 2, pp. 588, 589. 


Boriza kalodonta, P\.1., p. 991. 

Bryocodia cryptogramma, Pl. L., p. 991. 

Bryolimnia monodonia, Pl. J., p. 991. 

Bunolophodon angustidens, Pls. I.—-IV., 
p. 609. 


Cacopus systoma, Figs. 1-11, 18-19, 
pp. 528-530, 534-541, 545-547, 549- 
552. 


Callithria (Callicebus), Pl. IV., fig. 3, | 


prod: 

Callosciurus notatus, Fig. 45, p. 1184. 

prevostt, HKigs. 41, 43, 45, 59, 
pp. 1176, 1180, 1184, 1209. 

Castor canadensis, Fig. 39, p. 1174. 

fiber, Bigs. 40, 58, 59, 60, pp. 1175, 
1206, 1209, 1210. 

Centronia plorator, Pl. I., p. 991. 
Proc. Zoou. Soc.—1922. 


stephensoni, Pl. IIL., p. 943, Figs. | 


Cephalorhynchus commersonit, Pls. 1.— 
III., p. 627. 

—— eutropia, Pl. LIL., p, 627; Fig. 25, 
p. 641. 

heavisidei, Pls, 11., IL1., p. 627. 

hectort, Pl, il., p. 557. 

Characostomum asinitium, Figs. 3, 4, 
p. 905. 

Chiromys, Pl. V. fig. 2, p. 599. 

madagascariensis, Pls. I-VI, 
Figs. i-6, pp. 1145, 1150, 1154, 
1166-1168. 

Citellus beecheyi, Big. 57, p. 1204. 

(Otospermophilus) beecheyi, Figs. 
44, 54, pp. 1181, 1199. 

Clupea harengus, Bigs. 1-11, pp. 1216, 
1217, 1219-1227. 

Conopeum commensale, Pl. 1., p. 983. 

—— lacroixit, Pl. I., p. 983. 

Cyclorchida crassivesicula, Bigs. 11, 12, 
pp- 924, 925. 

Cylicospirura subequalis, Figs. 6, 7, 
p. 910. 

Cynomys ludovicianus, Kigs. 39, <1, 43, 
54, 59, pp. 1174, 1176, 1181, 1199, 
1209. 


Delphinapterus leucas, Fig. 27, 
Delphinus delphis, Fig. 26, p. 642. 
Diastephanus, Fig. 1, p. 710. 
alutaceus, Fig. 6, p. 815. 
sulcatus, Figs. 7, 8, p. 820. 
Dicentria klagest, Pl. L, p. 991. 

b 


Xvl INDEX OF ILLUSTRATIONS. 


Dicentria nondescripta, Pl. 1., p. 991. 

Dictyonosteus arcticus, Figs. 1-8, 
pp. 1242, 1248, 1250. 

Dinotherium sp., Pl. IV., p. 609. 

Diplocercides kayseri, Figs. 
pp. 1260, 1262. 


5, 6, 


Enchytreus crymodes, Fig. 6, p. 1135. 

Epitomiptera marmorata, Pl. 1., p. 991. 

—— purpurascens, Pl. I., p. 991. 

var., Pl. L., p. 991. 

Huglaucomys jimbriatus, Figs. 42, 55, 
pp. 1178, 1201. 

Husthenopteron fordii, Fig. 4, p. 1255. 


Kutamias quadrivitiatus, Figs. 41, 42, | 


51, 97, pp. 1176, 1178, 1194, 1204. 
Huxerus erythropus, Figs. 
pp- 1181, 1197. 


Fenatopus, Fig. 1, p. 710. 

rugiceps, Fig. 5, p. 79d. 

Funambulus palmarum, Figs. 42, 46, 
pp- 1178, 1186. 

Funisciurus leucostigma, Figs. 48, 57, 
pp. 1189, 1204. 


Galago, Pls. 11I., 1V., VI., p. 599. 

Geosciurus capensis, Figs. 41, 48, 58, 
59, pp. 1176, 1180, 1197, 1209. 

Glaucomys, Figs. 56, p. 1202. 

—— volans, Fig. 42, p. 1178. 

Grampus griseus, Fig. 26, p. 642. 


FTalicore indicus, Fig. 29, p. 646. 

Heliosciurus punctatus, Figs. 43, 48, 
pp. 1180, 1189. 

Hemiacodon, Pl. VI, fig. 1, p. 599. 

Hemistephanus, Fig. 1, p. 710. 

maculipennis, Fig. 4, p. 774. 

Henlea (Heniea) brucei, Big. 1, p. 1128. 

Hydrias albiochrea, Pi. 1., p. 991. 

Hydromys chrysogaster, Figs. 3-5, 
pp. 990-592. 

Hypena brodescens, Pl. 1., p. 991. 


Indris, Pls: '‘I., V., p. 599. 


44, 53, | 


Jaculus jaculus, Figs. 29, 30, 33, 36, 
pp- 661, 662, 670, 676. 


Kogia breviceps, Pl. I1., p. 557. 


Lagenorhynchus albirostris, Big. 26, 


p. 642. 


| Lamontia calibana, Pl. 1., p. 991. 


Lemur, Pl. 11, fig. 2, p. 599. 
-—— catia, Pl. II. fig. 1, p. 599. 
Leporillus jonesi, Figs. 8-10, pp. 595- 


Ligia baudiniana, Pl. II. figs. 17, 18, 
p. 683. 

cinerascens, Pl. IT. fig. 9, p. 683. 

exotica, Pl. II. fiy. 10, p. 683. 

—— glabrata, Pl. I. fig. 5; Pl. IL. 
fig. 6, p. 688. 

-— gracilipes, Pl. Il. figs. 
p. 688. 

——— lone, 1, UL, ite, I, 
p. 688. 

= walica, Pl. 
p. 685. 

occidentalis, Pl. Il. figs. 7, 8, 
p- 683. 

—— pailasii, Pl. I. fig. 4, p. 683. 

—— perkinsi, Pl. XL. fig. 1, p. 683. 

—— pigmentata, sp. n., Pl. L. figs. 2, 
3, p. 683. 

Lumbricillus egialites, 
pp. 1127, 1128: 

necrophagus, Bigs. 4, 5, pp. 1181, 

1152. 


13-15, 


1) igs) | (ee 


Figs. 2, 3, 


Macrocystis pyrifera, Pl. 1., p. 988. 
Macropus ruficollis, Pl. VIl. fig. 1, 
1s OE), 
Manatus americanus, Big. 29, p. 646. 
Marmota marmota, Figs. 59; 43, 52, 
pp. 1174, 1181, 1196. 
Membranipora tehuelcha, P\. L., p. 988. 
—— tuberculata, Pl. L., p. 983. 
Mesoplodon bidens, Fig. 28, p. 644. 
bowdoini, Pls. II1., IV., p. 557. 
Microcebus, Pl. III. fig. 2, p. 599. 
Microtus agrestis fiona, Anterior part of 
jugal, Fig. i p. 939. 


INDEX OF ILLUST 


Neobalena marginata, Pl. 1., p. 557. 
Neostephanus, Fig. 1, p. 710. 

Notomys cervinus, Figs. 11, 12, p. 598. 
Nyeticebus, Pl. VI. fig. 3, p. 599. 


Papillosetaria traguli, Fig. 10, p. 913. 
Parangitia micapennis, P\. I., p. 991. 
nmigrofulgens, Pl. L., p. 991. 
Parastephanellus, ig. 1, p. 710. 


Paraxerus cepapi, Figs. 48, 48, 57, 
pp. 1180, 1189, 1204. 

Pedetes caffer, Figs. 30, 31, 35, 36, 
pp. 662, 665, 673, 676. 

Perodicticus potto, Pi. VI. fig. 4, 
p. 599. 

Petaurista albiventer, Bigs. 53) 57, 


pp. 1201, 1204. 

Petrochirus granulimanus, Pl. 1., p. 1. 983. 

Phalanger orientalis, Figs. 31, 32, 37, 
39, 40, 43, pp. &65, 867, 872, 875, 
888, 894. 

Phascolarctos cinereus, Rigs. 38-41, 43, 
pp. 878, 875, 883, 885, 894. 

Phascolomys mitchelli, Figs. 31, 32, 35, 
36, 39, 41-43, pp. 865, 866, 869, 
871, 878, 885, 889, 894. 

Phastia maricolor, Pl. 1., p. 991. 

Phocena communis, Fig. 27, p. 648. 

Phostria duplicata, Pl. I., p. 991. 

Physaloptera sp., Big. 44, p. 1081. 

acuticauda, Hig. 26, p. 1051. 

— alata, Fig. 10, p. 1026. 

— anomala, Fig. 8, p. 1022 

antarctica, Figs. 36, 3/7, pp. 1068, 

1069. 

bonnet, 

1036. 

capensis, Figs. 42, 43, pp. 1078, 

1079. 

clausa, Figs. 2-4, pp. 1013-1015. 

colubri, Fig. 38, p. 1071. 

— dilatata, Fig. 41, p. 1076. 

—— gracilis, Fig, 19, p. 1040. 

longissima, Wig. 20, p. 1041. 

magnipapiula, Big. 51, p. 1059. 

—— malayensis, Pig. 23, p. 1047. 

manxillaris, Fig. 7, p. 1020. 


Figs. 16, 17, pp. 1035, 


RATIONS. XVil 


Physaloptera monodens, Bigs. 13, 14, 

pp. 1031, 1032. 

mordens, igs. 27, § 
1054. 

—— muris-brasiliensis, Fig. 9, p. 1024. 

obtusissima, Fig. 15, p. 1038. 

-—- papillotruncata, Fig. 5, p. 1017. 

—— paradoaa, Figs. 33, 34, pp. 1062, 
1064. 

phrynosoma, Fig. 18, p. 1038. 


Jaa ZN 


28, pp. 1053, 


—— preputialis, 
pp. 1043, 1045. 

quadrovaria, Fig. 32, p. 1061. 

retusa, Bigs. li, 12, pp. 1028, 
1029. 

— semilanceolata, Fig. 6, p. 1018. 

simplicidens, Hig. 39, p. 1073. 

terdentata, Figs. 24, 25, pp. 1048, 
1049. 

—— tumefaciens, 
pp. 1056, 1057. 

-— turgida, Hig. 40, p. 1075. 

varant, Wig. 3d, p. 1066. 

Fig. 25, 


Figs. 29, 30, 


Physeter macrocephalus, 
p. 641. 
Plusiodonta cupristria, Pl. I., p. 991. 


Pococera brunnapex, Pl. 1., p. 991. 


Polymastodon, Pl. VIL. figs. 2-5, 
p99! 

Propithecus, Pl. 1., Fig. 3, p. 599. 

Protoxerus stangeri, Fig. 47, p. 1187. 


Pseudochirus peregrinus, 


pp. 865, 868. 


Figs. 31, 34, 


Rana breviceps, Fig. 12, p. 548. 

heaadactyla, Bigs. 4, 12, pp. 584, 
543. 

— tigrina, Fig. 12, p. 543. 

Rattus greyi, Wigs. 6, 7, pp. 598, 594. 

Ratufa indica, Figs. 39, 42, 46, 57, 
pp. 1174, 1178, 1186, 1204. 

Rheithroseiurus macrotis, 
p. 1187. 

Rifargia brunnipennis, Pl. I., p. 991. 


Fig. 47, 


Salmo, Fig. 11, p. 1227. 
Samea delicata, Pl. I., p. 991. 
Sanys implicata, Pl. I., p. 991. 


XVIL1 


Seaphula celox, Figs. 
1143. 

minutia, Hig. 6, 7, pp. 1148, 
1144. 

Scarturus tetradactylus, Figs. 29, 31- 
34, 36, 37, pp. 661, 665, 668, 670, 
672, 676, 679. 

Scirtopsda orientalis, Pigs. 29-31, 34, 
36, pp. 661, 662, 665, 672, 676. 

Sciurus carolinensis, Fig. 49, p. 1191. 

hudsonius, Fig. 51, p. 1194. 

—— hypopyrrhus, Fig. 49, p. 1191. 

niger rufiventer, Fig. 50, p. 1192. 

saltuensis, Fig. 38, p. 1172. 

—— vulgaris, Figs. 38, 42, 50, 57, 
pp. 1172, 1178, 1192, 1204. 

Samoans 12k, Woy des By (ds SEE 

Setaria javensis, Fig. 9, p. 912. 

Sicista subtilis, Fig. 32, p. 668. 
Sisyrosea guaica, Pl. 1., p. 991. 

Syngnathus, Wig. 11, p. 1227. 

Syngria delicata, Pl. 1., p. 991. 

Stephanus, Big. 1, p. 710. 

coronator, Figs, 2, 3, p. 739. 

Streptopharagus armatus, Fig. 8, p. 911. 


1-5, pp. 1139- 


INDEX TO ILLUSTRATIONS. 


Tarsius, Pls, I11., 1V., p. 599. 

Testudo horsfieldii, Big. 21, p. 517. 

-——— ibera, Figs. 1, 11, pp. 487, 501. 

——- loveridgii, Pls. I-II1., p. 483; 
Figs, 1-21, pp. 487, 490-495, 497, 
499-501, 503-506, 508-510, 512, 513, 
517. 

Thairopora armata, Pl. 1., p. 988. 


Tomeutes fasciatus, Figs. 43, 45, 
pp. 1180, 1184. 
Troglostrongylus  troglostrongylus, 


Fig. 5, p. 908. 
Tursiops tursio, Fig. 27, p. 645. 


Ingulata, Tongues of, Fig. 30, p. 6d1. 
Aerus rutilus, Wig. 57, p. 1204. 


Zapus hudsonianus, Figs. 29-382, 
pp. 661, 662, 665, 668. 

Zygena malleus, Kigs. 1-7, pp. 
9738-975, 978, 980. 


ale 


INDEX. 
1922.—Pages 483-1276. 


[New names in clarendon type. 


Systematic references in italics. 


(z.8.t.) indicates additions to the Society’s Menagerie. | 


Abdimia abdimii, 861. 
Acercus lutescens, 1238. 
Atluredus viridis (2. 8. t.), 834. 
Agapornis personatus, 8d4. 
Alactagulus, 682. 
Alcedo semitorquata, 850. 
Allactaga, 667, 669, 672, 674, 675, 678, 
680, 682. 
Allotheria, 604. 
Amblyospiza unicolor, 842. 
Amblyostoma tigrinum, 834. 
Amydrus morio riippelli, 843. 
Ancistrodon contortria (z. 8. u.), 1275. 
Ancylostoma pluridentatum, 908. 
Andropadus insularis, 846. 
Anthreptes collaris, 838. 
Antilope, 654. 
Apodemus hebridensis 
subsp. n., 935. 
larus, subsp. n., 936, 
——- —— tire, subsp. n., 934. 
tural, subsp. n., 935. 
—— sylvaticus, 660, 670. 
Aquila rapax, 855, 
Arca americana, 1141. 
pexata, 1140. 
Ardea cinerea, 859. 
Proc. Zoou. Soc,—1922, 


ghia, 


Ardea melanocephala, 859. 

Armadillo, 683. 

Astur sphenurus, 856. 

tachiro (? nyanz@), 856. 
Asturinula monogrammica, 858. 
Atlantoxerus, 1181, 1188, 1195, 1197. 
getulus, 1181, 1195. 

Auchenia, 654. 

Aulodrilus kashi, sp. n., 246, 963. 
limnobius, 943. 

—— pleuriseta, 944., 

—— remex, 943, 948. 
stephensoni, sp. n., 963. 
Aulophorus furcatus, 943. 


Balena australis, 558. 

mysticetus, 640, 

Balenoptera acutorostrata, 563. 

boops, 640. 

-—— borealis, 562, 640. 

musculus, 561, 640. 

physalus, 562. 

Barbatia, 114.1. 

Belomys pearsonii, 1203. 

Berardius arnuxii, 567. 

Bison, 654. 

americanus (%Z.8.L.), 1276. 
Cc 


xx 


Boriza kalodonta, sp. n., 992. 

Bos, 654. 

Bothriocerus americanus, 736. 

europeus, 730. 

Bothrioneuron iris, 943. 

Bracon serrator, 718. 

Branchiodrilus hortensis, 948. 

Branchiura sowerbyi, 948. 

Breviceps verrucosus, 538. 

Brotogerys jugularis apurensis (4. 8. L.), 
1278. 

Bryocodia cryptogramma, sp. n., 
993. 

Bryolimnia monodonta, sp.n.,993. 

Buarremon brunneinucha (2.8. u.), 1274. 

Bubo africanus, 854. 

Budorcas, 654. 

Bunolophodon angustidens, 609. 

longirostre, 610. 

Butastur rufipennis, 857. 

Byssarea, 1142. 


Cacatua leadbeatert (2. s. u.), 835. 
Cacopus systoma, 527. 

Calidris arenaria, 862. 
Callithrix, 602. 

Callosciurus, 1172, 1188. 

—— finlaysoni, 1185. 

notatus, 1178, 1185. 
prevosti, 1178, 1184. 
Camelus bactrianus, 649. 

——— dromedarius, 649. 
Caprimulgus europeus, 849. 
fossei mossambicus, 849. 
Carapus fasciatus (2.8. L.), 1273. 
Cardiocranius, 675. 

Cassicus wropygialis (z. 8. L.), 835. 
Castor, 1178, 1175, 1182. 

— fiber, 835, 1207. 

Cavia, 677. 

Centronia plorator, sp.n., 991. 
Centropus monachus, 852. 
Cephalophus, 654. 

spadia (z. 8. L.), 833. 
Cephalerhynchus, 643. 
commersoni, 627, 629. 

—— eutropia, 631, 633, 641. 
— heavisidei, 631, 633. 


INDEX. 


Cephalorhynchus hectori, 580. ik 

Ceratophrys cultripes (z. 8. u.), 835. 

Cerchneis tinnunculus, 854. 

Cercopithecus patas (4. 8. u.), 1275. 

Chalcides, sp. n. (z. s. u.), 1274. 

—— armitagii, sp. n., 899. 

Chalcomitra amethystina (4. 8. u.), 1278, 
1274. 

senegalensis inestimata, 837. 

verreauxt, 838. 

Chaleopelia chalcospilos, 853. 

Chameleon gracilis (4. 8. u.), 1274, 
1275. 

Characostomum asmilium, 905. 

Charadrius alexandrinus, 862. 

hiaticula, 862. 

Chiromys, 601, 602. 

madagascariensis, 1145. 

Chlorophis irregularis (2. 8. u.), 1274. 

Cichladusa guttata, 846. 

Ciconia ciconia, 861. 

nigra (4%. 8. u.), 1276. 

Cireaétus pectoralis, 857. 

Circus macrurus, 858. 

Cisticola erythrops, 846. 

—— lugubris, 846. 

Citellus, 1174, 1182, 1200. 

—— (Otospermophilus) beechyi, 1179, 
1198. 

citellus, 1198. 

Civettictis civetta (z. 8. L.), 1278. 

Clupea harengus, 1218. 

Coccystes cafer, 853. 

glandarius, 852. 

Celogenys, 677. 

Colius indicus pallidus, 851. 

striatus affinis, 851, 

Coliuspasser ardens tropicus, 839. 

—— eques, 839. 

-— yacksoni, 839. 

laticauda, 839. 

Colobus polycomus (Z. 8. L.), 833. 

Connochetes, 654. 

Coracias caudatus, 849. 

-——— garrulus, 849. 

Cornopeum commensale, sp. n., 
985. 

——— facroirtt, 985. 


INDEX. 


Corvus scapulatus, 844. 

Corythornis cyanostigina, 851. 
Coturnix delegorguei, 862. 
Crateropus kirki, 845. 

platycercus (z. S.L.), 1274. 
Crax alector (Z. 8. u.), 835. 

sclatert (az. 8. L.), 835. 

Crex crex, 862. 

Cricetomys gambianus (2. 9. u.), 833. 
Crotalus confluentus (z. s. u.), 1275. 
horridus (az. s. u.), 1275. 
Cuculus canorus, 853. 

Cursorius gallicus (z. s.t.), 1276. 
Cyclanorbis senegalensis (a. 8. L.), 1275. 


Cyciorchida crassivesicula,sp.n., | 


924. 
-~—— omalancistrota, 923. 
Cylicospirura, gen. n., 909. 
—— subequalis, sp. n., 910. 
Cynelurus jubatus (4. 8. L.), 1275. 


XX1 


_ Diastephanus frontilinea, 813. 


Cynomys, 1174, 1180, 1182, 1198, 1200. | 


—- ludovicianus(z. s. u.), 1274. 
Cypselus affinis, 848. 


Dacelo gigantea (2.8. u.), 834. 
Delphinapterus, 643. 

leucas, 640, 641, 643. 
Delphinus, 643. 

commersoni, 627. 

—— delphis, 585. 
Dendraspis angusticeps (4. 8. %.), 1274. 
Dero limosa, 948. 
Diastephanus alutaceus, 814. 
bilineatus, 805. 

—— biréi, 804. 
brevipetiolatus, 81. 
carinifrons, 811. 
celebensis, 822. 
chinensis, 814. 

dohrni, 817. 

elegans, 810. 

fasciatus, 826. 
Hlaviceps, 824. 
flavidentatus, 809. 
flavifrons, 825. 
flavomaculatus, 807. 
—— flavonotatus, 826. 


JSuscidens, 804. 

Fuscinervis, 824. 

—— gracilis, 805. 

leucodon, 802. 

lewcodontus, 810. 

maculifemur, 819. 

—— nigripes, 812. 

nova-quineensis, 807. 

——- pallescens, 803. 

—— parviceps, 812. 

quadridens, 818. 

rothkirchi, 808. 

—— ruficollis, 819. 

salomonis, 821. 

-—— similis, 824. 

—— simillimus, 823. 

sulcatus, 820. 

—— szépligetit, 828. 

-— terebellus, 818. 

— tertianus, 806. 

togoensis, 797. 

trilineatus, 815. 

trilobatus, 816. 

Dicentria nondescripta, sp. n., 
991. 

Dicotyles, 653. 

Dicrurus afer lugubris, 844. 

Dictyonosteus arcticus, 1241. 

Dinotherium, 609, 620. 

Diplocercides kaysert, 1268. 

Diplodontus despiciens, 1252. 

Dipus mauritanicus, 679, 680, 682. 

Dissodectes dickinsont, 854. 


Echinococcus granulosus, 926. 

Hianus ceruleus, 855. 

Hlephas africanus, 654. 

—— indicus, 654. 

Koglaucomys, 1177. 

jfinbriatus, 1208, 1205. 

Enchytreus sp., 1135. 

—— albidus, 1110, 1112, 1133. 

crymodes, sp. n., 11383. 

EKohippus, 606. 

Ephippiorhynchus senegalensis (2. 8. L.), 
1275. 


XX1i 


Epitomiptera marmorata, sp.n.. 
995. 

purpurascens, sp. n., 994. 

Keuus, 653. 

asinus, 648. 

caballus, 648. 

-—— chapmanni, 648. 

kiang (2. 8. u.), 1273, 

EHrinaceus, 603. 

KHubelum, 683. 

Euchoreutes, 669, 671, 675, 680. 

Hupagurus bernhardus, 984. 

Hupsychortyx leucopogon (4. 8. u,), 835. 

Eurocephalus riippelli, 845. 

Husthenopteron fordi, 1249. 

Eutamias, 1177. 

quadrivittatus, 1193. 

Huxerus, 1197. 

erythropus, 1180, 1183, 1195, 1197. 

Hylais eatendens, 1238. 

hamata, 1232. | 


Falco biarmicus, 854. 

-—— cuvieri, 854. 

Felis tigris (a. 8. u.), 1273, 1275. 
Fenatopus albomaculatus, 781. 
annulitarsis, 787. 
aratifrons, 799. 

arcuatus, 793. 

birdi, 804. 

— brevicollis, 789. 

—— celebensis, 822. 
claripennis, 798. 

-— conradti, 789. 

fasciatus, 826. 
Fernandopoensis, 788. | 
flavicollis, 799. 
formosanus, 786. 

—— fuscinervis, 824. 

indicus, 784. 

var. swmbana, 785, 
-—— yar. sulcaticollis, 787. 
var. P, 785. 
iridipennis, 796. 
lactetpennis, 790. | 
longicauda, 794. 


longicollis, 790. 
natalicus, 793. 


INDEX. 


Kenatopus nigripes, 812. 
—— nova-guineensis, 807. 
-—— ocellatus, 784. 

—— piceicornis, 796. 
—— punctatus, 797. 
ruficeps, 734. 
rugiceps, 794. 

—— schlettereri, 788. 
similis, 824. 
simpsoni, 792. 
sulcaticollis, 787. 
sumbanus, 785. 
—— togoensis, 797. 

—— turcomanorum, 791. 
variidens, 793. 
Fenus diadema, 827. 
Formica rufa, 833. 
Fridericia leydigt, 1111. 
Fringillarie tahapizi (a. 8. u.), 1275. 
Funambulus, 1178, 1188. 
—— palmarum, 1188. 
étristriatus, 11'79. 
Funiscturus, 1179, 1183. 
——- cepapt, 1179, 1190. 
isabella, 1189. 
leucostigma, 1179, 1189. 


Galago, 602, 603. 
Galoncus tridentatus, 901. 
Geosciurus capensis, 1180, 1183, 1195, 
1197. 7 
Giraffa, 647, 649. 
Giaucomys, 1177, 1205. 
stramineus, 1203. 
-—— volans, 1203. 


Globicephalta melena, 578. 
Grampus griseus, 579, 641. 
Gymnura, 603. 


Hematopus ostralequs, 862. 
Haliaétus vocifer (% .8. u.), 834. 
Halcyon albiventris orientalis, 851. 
—— chelicuti, 851. 

—— leucocephalus, 851. 

Haticore, 647. 

Harpolestes senegala ortentalis, 844 


INDEX. 


Heliosciurus punctatus, 1179, 1188,1190. | 
| Hypochera orientalis, 838. 

| Hyrax, 654. 

| Hystrix africe-australis (z. s. u.), 1278. 


Hemiacodon, 604. 

Hemimastodon, 609. 

Hemistephanus angulicollis, 766. 

—— collarifer, 763. 

cylindricus, 776. 

damelicus, 753. 

erythrocephalus, 767. 

—— glabricoxis, 764. 

granulatus, 771." 

intermedius, 771. 

—~ limpidipennis, 769. 

—— macrurus, 770. 

maculipennis, 773. 

marginalis, 764. 

pehlkei, 772. 

—— peruanus, 765. 

—— rujiceps, 774. 

suwbmaculatus, 773. 

tener, 767. 

——— vadosus, 175. 

—— wiistneii, 768. 

Hemisus, 585. 

Henlea (Henlea) brucei, sp. n., 
1121. 

—— (Henleanelia), sp., 1125. 

heleotrophus, 


Sp. Me, 


1119. 
(Hepatogaster), sp., 1126. 
Hesperopithecus, 1275. 
Himantopus himantopus, 862. 
Hirundo emint, 847. 
puella wnitatis, 847. 
smithi, 847. 
Hyena hyena (2.8. u.), 12738. 
Hydractinia echinata, 989. 
Hydryphantes ruber, 1232. 
Hydrarachna distincta, 1252. 
Fuscata, 1232. 
—— globosa, 1232. 
paludosa, 1234. 
—— schneideri, 1232. 
Hydrias albiochrea, sp. n., 996. 
Hydromys chrysogaster, 589. 
Hygrobates longipalpis, 1236. 
Hylopetes alboniger, 1205, 
Hyomoschus, 649, 654. 


XKili 


Elypena brodescens, sp. n., 995. 


Ichneumon coronatus, 718. 
—- serrator, 718. 


Indris, 602. 


| Jaculus jaculus, 659, 664, 667, 669, 


674, 675, 678, 682, 


Kogia breviceps, 566. 


| Lagenorhynchus, 643. 


albirostris, 641. 

eruciger, 628. 

flowert, 628. 

obscurus, 581. 

Lagonosticta senegala, subsp. n., 
842. 

Lamontia calibana, sp. n., 997. 

Laniarius sublacteus, 844. 

Lanius collaris humeralis, 844. 

Leiperenia galebi, 901. 

Leporitius jonest, 595. 

Lepralia bifurcata, 988. 

—— edar, 983 


| Ligia, 683. 


baudiniana, 698. 
—— cinerascens, 693. 
dilatata, 701. 
exotica, 693. 
— glabrata, 692. 
-— gracilipes, 695, 
hawaiensis, 696. 
italica, 695. 
natalensis, 700. 

- nove-zealandie, 697. 
—— occidentalis, 692. 
- oceanica, 691. 

- olfersii, 694. 


-—— pallasiz, 691. 


perkinsi, 699. 


| —— pigmentata, sp. n., 699. 


—— richardsone, 701. 
simon, TOL. 


c3 


XXIV 


Limnesia fulgida, 1232. 

—— undulata, 1236. 

Limnodrilus claparédeianus, 955. 

—— socialis, 943. 

Lissodelphis peronti, 582. 

Lophoaétus occipitalis, 855. 

Lophoceros deckeni, 850. 

-— melanoleucus suahelicus, 850. 

neumanni, 850. 

Loxodon cyclotis (2.8. u.), 1276. 

Lumbricillus egialites, sp. n 
1126, 1133. ; 

Sranciseanus, 1129. 

henkingi, 1129. 

—— nicrophagus, sp. n., 1130 
1133. 

—— pagenstecheri, 1110, 1129. 

Lybius zombe, 853. 

Lyngria delicata, sp. n., 996. 


? 


Macronyx croceus, 838. 

Malaconotus poliocephalus blanchoti, 
845. 

Manatus, 646. 

Marionina ebudensis, 1110. 

Marmota, 1178, 1182, 1200. 

bobak, 1200. 

—. caudata, 1200. 

—- himalayana, 1200. 

—— hodgsoni, 1200. 

— marmotta, 1179. 

monax, 1200. 

Marmotops, n., 1200. 

—— monax, 1200. 

Megaladapis grandidiert, 603. 

Megaptera nodosa, 564. 

Megischus acutus, 742. 

—— americanus, 736. 

—— annulator, 736. 

antinori, 719. 

— borneensis, 735. 

brasilianus, 736. 


brunneus, 725. 
— canadensis, 827. 
claripennis, 758. 
coronator, 738. 


—— Cylindricus, 776. 


08) 


INDEX. 


Megischus ducalis, 740. 
erythrocephalus, 767. 
europeus, 730, 
Hloridanus, 828. 

— froggattii, 734. 
Jurcatus, 827. 
indicus, 784. 
insidiator, 724. 
insularis,738. 


longicaudatus, 738. 
longicollis, 790. 
maculifrons, 751. 
maculipennis, 773. 
—— niger, 733. 
nigricauda, 772. 
— palliditarsis, 756. 
ruficeps, 737, 774. 
rujicollis, 'T56. 
spoliator, 828. 
submaculatus, 773. 
— tarsalis, 829. 
tarsatus, 729. 


—— —— yar. nigricans, 733. 

— texanus, 730. 

—— viduus, 738. 

—— violaceipennis, 746. 

Megiseleius, read Megischus. 

Melanophoyx ardestaca, 858. 

Melierax gabar, 856. 

Melittophagus bulleckoides, 849. 

Membranipora denticulata, 984. 

—— membranacea, 986. 

——- savartti, 987. 

——— tehuelcha, 989. 

tuberculata, 986. 

Merops apiaster, 850. 

Mesenchytreus sp., 1136. 

gelidus, 1110. 

solifugus, var, rainierensis, 1110. 

Mesocestoides lineatus, 922. 

litteratus, 922. 

Mesoplodon bidens, 640, 648. 

bowdoini, 569. 

—— grayi, Did. 

layardt, 574. 

Microtus agrestis fiona, subsp n, 
940. 


? 


Monodon monoceros, 640. 
Monopylephorus africanus, 967. 
parvus, 943. 

Motacilla flava campestris, 838. 
vidua, 838. 

Mus musculus musculus, 933. 


Naia bungarus (4.8. u.), 1273. 
Naidium pleuriscta, 944. 

Nais josine, 1119. 

Nectarinia famosa (2.8. u-), 1278. 
reichenowi, 837. 
Neohalena marginata, 559. 
Neostephanus albomaculatus, 781. 
alluaudi, 777. 
camerunus, 779. 
crassipes, 780. 

—— globiceps, T79. 

insignis, 778. 

natalicus, 793. 

‘penthert, 781. 

Nestor notabilis (z. 8. u.), 1276. 
Notharctus, 604. 

Nothodectes, 604. 

Notomys cervinus, 597. 
Numenius arquatus, 862. 
pheopus, 862. 
Nycticebus, 603. 


Orcella brevirostris, 640. 
fulminalis, 642. 

Orcinus orca, 577. 

Oriolus brachyrhynchus letior, 843. 


Ortholophus leucolophus (2. 8. t), 1273. 


Otaria californica (4. 8. L.), 839. 
Ourebia nigricaudata (4.8. u.), 1275. 
Ovibos, 654. 

moschatus (Z. 8. L.), 1274. 

Ovis, 654. 


Pagurus bernhardus, 897. 
—— cuanensis, 897. 
Paikea, gen. n., 574. 
hectort, 575. 
Paleopropithecus, 603. 
Pandion haliaetus, 854. 


INDEX. xxXV 


Papillosetaria, gen. n., 913. 

-—— traguli, sp. n., 913. 

Papio papio (4. 8. u.), 1273. 

Parangitia micapennis, sp. n., 
994, 

— nigrofulgens, sp. n., 99+. 

Parastephanus albiceps, 751. 

brevistigma, 756. 

claripennis, 758. 

damellicus, 753. 

eburneus, TH2. 

--— glaber, 749. 

—— levicollis, 750. 

—— maculifrons, 751. 

—— malayanus, 760. 

—— martini, 748. 

—— orbitalis, 748. 

palliditarsis, 756. 

pictipes, T54. 

—— pygmeus, 750. 

— rubripictus, 759. 

rufidornatus, 758. 

—— rufo-ornatus, 758. 

seitus, T57. 

Passer griseus suahelicus, 838. 

Pedetes caffer, 659, 664, 667, 669, 672, 
674, 675, 678. 

Peloneustes philarchus, 334. 

Pelycodus, 604. 

Penelope purpurascens (4. 8. b.), 830. 

Peradectes, 604, 606. 

Pernis apivorus, 85d. 

Perodicticus potto, 602. 

Petaurista, 1177. 

albiventer (magnificus), 1200, 1205. 

caniceps, 1203. 

philippensis, 1203. 

yunnanensis, 1208. 

Petinomys fuscocapillus, 1205. 

Petrochirus granulimanus, 983. 

Petrodromus, 603. 

Phacocherus, 653. 

africanus (Z. 8. L.), 1274. 

Phalanger orientalis, 863. 

Phascolarctos cinereus, 863. 


| Phascolomys, 608. 


—— mitchelli, 863. 


XXvVi INDEX. 
Phastia maricolor, sp. n., 992. 
Phenacolemur, 604. 

Phocena, 648. 

Pholidauges leucogaster (2. s.u.), 1274. 
verreauxt (z,8.u.), 1274. 
Phostria duplicata, sp. n., 995. 


Physaloptera gracilis, sp. n., 1039. 
—— guiartt, 1092. 

ancurva, 1093. 

imermis, 1094. 

| —— inflata, 1096. 

| — leptosoma, 1085. 


Physaloptera, 1004, 1011, 1080. 
abbreviata, 1087. 
abjecta, 1100. 
—— acuticauda, 1050. 
affinis, 1088. 
——- alata, 1025. 
var. chevreuxt, 1083. 
var. nowvellt, 1083. 
alba, 1099. 
--— aloisii-sabaudie, 1098. 
— amphibia, 1099. 
anomala, 1021. 
—— antarctica, 1067, 1099. 
-— bilabiata, 1096. 

—— bonnei, sp. n., 1034. 
brevicauda, 1100. 
brevispiculum, 1090. 
brevivaginata, 1083. 
britanica, 1097. 

—— bulbosa, 1102. 
—— capensis, sp. n., 1078. 
caucasica, 1089. 
—— cesticillata, 1091. 
chame@leontis, 1086. 
~— circularis, 1093. 
citilh, 1095. 

clausa, 1012. 
coelebs, 1092. 
~—— colubri, 1070. 

— constricta, 1102. 
contorta, 1102. 
—+ crassa, 1096. 
crost, 1085. 
— — dentata, 1097. 
—— digitata, 1090. 
—— dilatata, 1076. 

dispar, 1092. 
elegantissima, 1091. 
Susiformis, 1097. 
galiniert, 1085. 
—— gemina, 1090. 
getula, 1082, 


— limbata, 1095. 
-— longissima, sp. n., 1041. 


| -—— magnipapilla, 1058. 


—— malayensis, sp. n., 1046. 
malleus, 1100. 

— masxillaris, 1019, 

-—— megalostoma, 1096. 
—— mépivites, 1091. 
monodens, 1030. 

—— mordens, 1052, 

—— mucronata, 1103. 

—-~ muris-hrasiliensis, 1023. 
—--— nasilionis, 1096. 

— numidica, 1086. 

—— obtusissima, 1082. 
ovata, 1103. 

pallaryt, 1087. 
papilloradiata, 1101. 
—— papillotruncata, 1016. 
—— paradoxra, 1062. 

—— phrynosoma, sp. n., i037. 
—— prepitialis, 1043. 

—— pyramidalis, 1095. 
—— quadrovaria, 1060. 

— rara, 1081. 

retusa, 1027. 
rotundata, 1101. 

—— ruwenzorri, 1093. 

—-—- saginata, 1103. 

sciuri, 1094. 

-—— semilanceolata, 1017. 
—— simplicidens, sp. n., 1072. 
—— sonsinot, 1098. 

—— spiralis, 1098. 

—— spirula, 1101. 

striata, 1103. 


| xii strongylina, 1104. 


—— subalata, 1084. 


| —-— tacapensis, 1082. 


—— tenutcollis, 1104. 
—— terdentata, 1048. 
—— torquata, 1091. 


INDEX. 


Physaloptera torresi, 1088. 

——- truncata, 1097. 

—— tumefaciens, 1055. 

—— turgida, 1074. 

—— varani, 1065. 

Physeter, 648. 

—— macrocephalus, 565, 641. 

Pimpla coronator, 738. 

Piona longipalpis, 1232. 

Platanista gangetica, 640, 643. 

Plecostomus conmersoni (z. 8. u.), 1274. 

Ploceus nigriceps, 842. 

—— reichenowi, 842. 

—— speker, 845. 

Plusiodonta cupristria, sp. n., 
992. 

Pococera brunnapex, sp. n., 996. 

Podargus strigoides (z. 8. L-), 834. 

Poicephatus fuscicapillus, 853. 

Polumastodon, 605. 

Porolepis ? sp., 1259. 

Potamocherus, 653. 

Prinia mystacea tenella, 846. 

Propithecus, 601, 603. 

Prosymna meleagris (a. 8. L.), 1275. 

Protoxerus, 1190. 

—— stangert, 1190. 

Psammophis sibilans (a. 8. u.), 1275. 

Pseudochirus peregrinus, 863. 

Pseudorca crassidens, 577. 

Péteropus medius (%. 8. u.), 1276. 

Ptilodus, 605. 

Pulsatri« perspicillata (z. 8. u.), 1275. 

Pycnonotus barbatus micrus, 845. 

-—— tricolor pallidus, 845. 

Pygeretmus, 675. 

Pyromelana flammiceps, 840. 

— — nigriventris, 840. 

—— xanthomelana, 840. 

Pytelia melba kirki, 842. 


Quelea cardinalis, 840. 
—— intermedia, 840. 


ana breviceps, 542. 
——- hexadactyla, 542. 
—— tigrina, 542. 
Rattus greyi, 593. 


XXXVI 


Ratufa, 1178, 1188. 
bicolor, 1185. 
—— indica, 1185. 
macrourus, 1185. 


| Redunca nagor (4. 8. u.), 1275. 


Rhacophorus buergeri (z. 8. u.), 1274. 
Lhamphocalus dimidiatus (a. 8. t.), 839. 
Rheithrosciurus, 1183, 1188. 
Rhinoceros, 647, 658. 

Rhinochetus gubatus (2. 8. u.), 834. 
Lhodospiza obsoleta (z. 8. u.), 1273. 
Rhyncocyon, 603. 

Rifargia brunnipennis, sp. n.. 993. 
Riparia rufigula, 847. 


Sagartia parasitica, 897. 
Saltator artricollis (z. s. u.), 1276. 
Samea delicata, sp. n., 997. 


| Sanys impilacata, sp. n., 994. 


Scaphula celox, 1159. 

—-— minuta, sp. n., 1143. 

Scarturus tetradactylus, 659, 662, 664, 
667, 669, 674, 675, 678, 682. 

Schizorhis africana leucogastra, 852. 

Schlettererius cinetipes, 714. 

rufipes, 713. 

Seirtopoda orientalis, 659, 664, 667, 
669, 674, 675, 678, 682. 

Sciwropterus, 1203, 1205. 

—— (Hylopeies) alboniger, 1203. . 

(Petinomys) fuscocapillus, 1203. 

Seiurus, 1172. 

carolinensis, 1177. 

—— (Neosciurus) carolinensis, 1191. 

—— griseus, 1177. 

—- (Tamiasciurus) hudsonius, 1193. 

— hudsonius, 1177. 

—— (Kchinosciwrus) hypopyrrhus, 1190. 

hypopyrrhus, 1177. 

—— niger (rufiventer), 1177, 1192. 

saltuensis, 1173, 1177. 

vulgaris, 1173, V177. 

Scopus wmbretta (z. 8. L.), 834. 

— — —— bannermanni, 861. 

Seleucides chrysocephalus (2. 8, L.), 834. 

Serinus striolata affinis, 858. 

Setaria, 911. 

-—— javensis, sp. u., 911. 


XXVIil 


Sicista loriger, 661, 664, 667, 669, 674, 
675, 682. 
Sisyrosea guaica, sp. n., 996. 
Sorex araneus, 930. 
—— —— araneus, 931. 
grantt, 932. 
minutus minutus, 938, 
Speothos venaticus, 834. 
Spermestes, sp., 841. 
Spreo superbus, 843. 
Stenella euphrosyne, 582. 
pseudodelphis, 583. 
Stephanus acutus, 742. 
—— andinus, 727. 
annulator, 736. 
anomalipes, 731. 
antinori, 719. 
bicolor, 732. 
—- borneensis, 735. 
brasiliensis, 736. 
— brevicollis, 789. 
-— brevipetiolatus, 816. 
—— brunneus, 725, 
capitatus, 774. 
ceylonicus, 735. 
cinctipes, 714. 
collarifer, 763. 
comma, G44. 
conradti, 789. 
coronator, 738. 
coronatus, 718. 


erassicauda, (28. 
-—— cylindricus, 776. 
damellicus, 753, 759. 
diadema, 8277. 
diversus, 827. 
ducalis, 740. 
erythrocephalus, 767. 
europeus, 730. 

—— flavomaculatus, 807. 
— froggattti, 734. 
Frontalis, 828. 

——- furcatus, 736. 

—— gigas, 720. 

—— globiceps, 779. 
hematipoda, 741. 
—— hornianus, 726. 

—- indicus, 784. 


INDEX. 


Stephanus insignis, 778. 
intermedius, (71. 
lacteipennis, 790. 
lanceolatus, 741. 
leucodontus, 810. 
—— limpidipennis, 769. 
lucidus, 740. 

—— macrurus, 770. 
maculipennis, 773. 
malayanus, 760. 
marginalis, 764. 
martini, T48. 
natalicus, 798. 
—— niger, 733. 

—— nigricans, 733. 
—— nigricauda, 722. 
—— pachylomerus, 723. 
pallescens, 803. 
pulosus, TAA. 
pygmeus, 750. 
rubripes, 734. 
ruficeps, T37. 
rufipes, 716. 
rufofemoratus, 743. 
rufo-ornatus, 758. 
rugosus, 745, 
salomonis, 821. 
saussuret, "737. 
schlettereri, 788. 
serrator, 718. 
sickmannt, 727. 
—— spoliator, 828. 
submaculatus, 773. 
sulcifrons, 724. 
—— tarsatus, 729. 

-—— tener, 767. 

—— terebellus, 818. 
—— texanus, 730. 

—— tibiator, 721. 

—— togoensis, 797. 
-— tortus, 731. 
turcomanorum, 731. 
—— unicolor, 725. 

-—— vadosus, 775. 

—— villosus, 743. 
violaceipennis, 746. 
—— wustneti, 768. 

—— wanthocephalus, 724. 


Strepsiceros, 654. 
Streptopharagus armatus, 910. 
Struthio camelus (4.8. u.), 1273. 
Suberites domuncula, 897. 
Jicus, 989. 

Sus, 647, 653. 

—— babirussa, 648. 

scrofa, 648. 

Sylvieltta whytei, 846. 


Tachornis parvus, 848. 

Tamias, 1174, 1183. 

Tapirus, 653. 

americanus, 648, 

— bairdi, 648. 

— indicus, 648. 

terrestris (zZ. 8. L.), 1274. 
Tarsius, 602, 6038. 

Tchitrea cristata suahelica, 846. 
Testudo calcarata (z. 8. u.), 1274. 
tbera, 493, 502. 

loveridgu, 484. 
Thairopora armata, 987. 


Thomonys bulbivorus (z. 8. u.), 1274. 


Tomeutes, 1188. 

— hippurus, 1185. 

vittatus, 1185. 

Totanus hypoleucus, 862. 
Trachyphonus emini, 853. 
Tragelaphus scriptus (z. 8. .), 1273. 


INDEX. 


Tragulus, 648, 649, 654. 
Tricholema lachrymosa, 853. 
Tringa minuta, 862. 

subarquata, 862. 
Troglostrongylus, gen. n., 906. 


Tubifex (Ilyodrilus) bedoti, 953. 
tubifex, 943, 955. 
velutinus, 944. 

Tupaia, 603. 

Tursiops, 643. 

truncatus, 584. 

Turtur senegalensis, 853. 
Tympanistra tympanistra, 853. 


Ureginthus bengalus angolensis, 841. 
niassensis, 841, 


Varanus niloticus (z.8.t.), 12175. 
Vespa germanica, 833. 


Xerus, 1180, 1183. 
—-— rutilus, 1181, 1195. 
Xorides coronatus, 718. 


678, 680, 682. 
hudsonianus, 659. 
Ziphius cavirostris, 576. 
Zygena malleus, 971. 


PRINTHD BY TAYLOR AND FRANCIS, RED LION COURT, FLEBT STREET, 


xxix 


troglostrongylus, sp. n., 907. 


Zapus, 660, 664, 667, 669, 674, 675, 


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PROCEEDINGS 


OF THE 


GENERAL MEETINGS FOR SCIENTIFIC BUSINESS 


OF THE 


ZQOLOGICAL SOCIETY 
OF LONDON, 


1922. 


PART III. 


CONTAINING Paces 483 To 835, witH 23 Plates" 


87 'lex?-FIGURES. 


SEPTEMBER 1922. 


PRINTED FOR THE SOCIETY, 
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LIST OF CONTENTS. 


1922, Parr III. (pp. 483-835). 


EXHIBITIONS AND NOTICES. 


nu Sucretary. Remarks upon a Medal struck to commemorate the 150th Anniversary 
of the Royal Academy of Belgium...... 


Mr. F. Martin Dunean, F.Z.S. Exhibition of cinematograph films illustrating the life- 
history of the Wood-Ant (formica rufa) and Common Wasp (Vespa germanica) . 


Dr. Cnas. F, Sonnrac, F.Z.S. Exhibition of specimens and lantern-slides illustrating 
points in the Anatomy and Physiclogy of Whales 


ee ee ee ee wee ee Oe ee we PH Oe te ee oe 


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


of April, 1922 


Mr. R. I. Pococr, F.B.S., F.Z.S. Exhibition of a living example of the rare Bush- pees 
Speothos venaticus nao nda 

Mr. B. G. Bounznenr, F.Z.S8., and Mr. F. Martin Duncan, F.Z.8. Exhibition of cinema- 
tograph record illustrating the life-history of the Axolotl (Amblystoma tigrinuwm).. . 


Mr. D. Suru-Smirn, F.Z.S. Exhibition of photographs of recent important additions 
to the Society’s collection of Birds . Beas cei 

Rey. H. N. Hurontnson, M.A., F.Z.S. Exhibition of a plaster-cast model reconstruction 
othe maximexeptile Peloneustes pRtanCHiUs) wines |« mie elst-)e) ct) sla) cies eis) e eres sie 
Tne Stcretary. Report on the Additions to the Society’s Menagerie during the month 
OIF MeN NOMA eGo Gun EW ve odo subloce 


Tne Secretary. On the Council’s scheme to establish an Aquarium 


Page 


833 


833 


833 


833 


834 


834 


834 


834 


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(2 natural size) 


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VITTY & SEABORNE, LTD., 
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P.Z.S. 1922. Procter PI. Il. 


TESTUDO LOVERIDGII BLGR. @ 


(? natural size) 


VITTY & SEABORNE, LTD., 
LONDON. 


P.Z.S. 1922. Procter PI. Ill. 


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PAPERS. 


26. A Study of the Remareable Tortoise, Testudo loveridgit 


Blgr., and the Morphogeny of the Chelonian Carapace. 
ite JOAN B. Procter, F.Z.8. 


[Received May 22, 1922: Read June 18, 1922.] 


(Plates I—III.*; Text-figures 1-21; and Table.) 


INDEX. Page 
introductrony mere te cea enn tien a eee Coen 9 
General Account .. Ui ae Sie 1484, 


Material and Meter 3 Cimactes fess UMiaR Won, 
Habitat and habits. 


Rixbermalt Charactersiueeseensne od ckee eects ee ee 487 
Shape and proportions; Coloration; Head; Limbs; 
Tail; Epidermal shields. 
Skeleton ......... 496 


Bony carapace ae adult, ‘Boar miscre on Nee adult; (Sails 
Vertebral column; Ribs; ; Pectoral girdle; Pelvic 


girdle. 

Development ......... BKeb ope Cbobao AHS ahs Haw eoadaceepeoeae! | MOOT 
Ribs ; Carapace; Plaetron. 

Discussion ....... 514 


Morphogeny of the oan apace eand Plastroy on; fou igin 2 the 
costal and neural plates. 


RSIRUTERSTO AE, nl cunnh aomar nee a Ree Hd CoecHBEe acai acpaiace eerae ae lite il 532) E 

INIOGESW Aas cea eee eM oe. oan reese Amel naan, SANNA! Abas Meee. Oe 

Bibliography 524, 
INTRODUCTION. 


In the preparation of the present paper, I am indebted to 
many persons for help and various courtesies. My especial 
thanks are due to Mr. A. Loveridge, for his generosity in allowing 


* For explanation of the Plates, see p/ 526, 
Proc. Zoou. Soc.—1922, No. XXXIV. 34 


484 MISS JOAN B. PROCTER ON THE 


me to use the whole of his valuable collection for dissection and 
general study. Also to Sir John Bland-Sutton, for his gift of 
X-ray plates of the types and young, and facility to study the 
tortoises themselves beneath the fluorescent screen at Middlesex 
Hospital at a time when no specimens could be spared for dissec- 
tion; andalso to Mr. R. H. Burne, of the Royal College of Surgeons, 
for investigating the structure of the denticulated jaw, and the 
ribs, and for making slides of them. 

The purport of the present work is to furnish a detailed 
description of this extraordinary Soft Tortoise, based upon the 
large series of specimens which have passed through my hands; 
especially of the peculiarly interesting structure of the bony cara- 
pace and its development, compared with cases of fenestration in 
Testudo and other groups. The recently discovered young is also 
described in comparison with the adult, and various notes on the 
habits of living specimens by Mr. Loveridge are appended. 


GENERAL ACCOUNT. 
Material and history. 


In 1920, when Mr. G. A. Boulenger first examined this tortoise, 
then new to science, there were no specimens available for the 
making of askeleton. The carapace, which feels soft and springy, 
was examined by cutting a flap in the abdominal shields, removing 
the viscera, and holding the animal against the light. When 
this is done, no bony matter of any kind can be detected; the 
carapace is seemingly translucent without ribs or costal plates: 
it was therefore described as boneless. The view obtained in this 
way is deceptive, as it is limited to the region of the second and 
third costal and vertebral shields on account of the restrictions 
of the fenestra in the plastron through which it is viewed; the 
shadows of the reduced costal plates, which are there, are entirely 
obscured by the black markings in the epidermal shields. 

The young specimen described in his note*, although agreeing 
in almost every particular with the type, excepting its convexity, 
was an anomalous specimen, not the young of loveridgii. In 1921, 
Mr. Loveridge sent a Kaffir on an expedition to hunt for further 
material, and the true young was found in the type-locality | 
This has a depressed carapace, but does not resemble the adult in 
general appearance. Since then further expeditions have suc- 
ceeded in capturing more specimens, both adult, young, and half- 
grown, so that at one time and another I have been able to 
examine twenty-three spirit specimens, besides two specimens 
at present living in the Zoological Gardens. It has also been 
possible to prepare skeletons both of adult and young, and to 
make dissections. 

After examining this new material, Mr. Boulenger agrees 


* Boulenger, C.R. Acad. Sci., Paris, t. 170, p. 264 (1920). 


REMARKABLE TORTOISE, TESTUDO LOVERIDGII. A485 


with me in my three principal amendments to his original 
description :— 
1. That the type-specimens are fully adult. 


2. That the adults have bony neural and costal plates present 
to a certain degree. 


3. That the young has a more or less depressed carapace. 


Characteristics. 


Testudo loveridgti is already wel! known as the Soft-shelled 
Tortoise with the flat carapace, both these characteristics being 
entirely foreign to Zestudo as previously defined. 

In general appearance it looks as if it had been crushed in youth 
and had only survived by a miracle. When taken in the hand 
alive it has a boneless feeling which is uncanny ; both carapace 
and plastron react to pressure on the abdominal region with a 
springy motion, and the animal is able to inflate itself to a slight 
degree. 

Mr. Boulenger pointed out in his note that in the case of the 
plastron this softness is due to an enormous diamond-shaped 
fenestra, usually met with in the newly-hatched or very young 
of other species. The viscera thus exposed are protected by 
extremely thick, soft, dermis of a very tough nature. 

The true structure of the carapace was revealed by X-rays quite 
accidentally, my original object being to compare the supposedly 
boneless adult with the young, in which the ribs show normal 
development. When viewed through the fluorescent screen, the 
adults presented an extraordinary appearance, the ribless vertebree 
and more or less normal limb-girdles being overlaid by a bony 
network, intimately correlated with the net of sutures between 
the epidermal shields, and formed by the partial development of 
the neural and costal plates which had been apparently wanting, 
and which in other species compose the solid bony carapace (vide 
Pls. II. & II.). The significance of this bony network only 
became clear after considerable study of the X-ray plate taken 
at the time. The skeleton subsequently prepared corroborated 
every detail revealed in this plate, and also showed that the 
vertebre are very remarkable in form. The animal is further 
remarkable in possessing what appear to be teeth, acrodont in 
type and similar to those met with in Agamoid lizards. Their 
regularity renders them quite unlike the serrations met with in 
some tortoises. The horny sheaths of the alveolar ridges are 
similarly denticulated. 

In the detailed description which follows, these structures will 
be dealt with at greater length. 

The species 1s still in a state of great instability, for apart from 
the immense range of variation to which it is subject, many 
abnormalities occur, such as the presence of horny shields proper 


to the more primitive Chelonians. 
34* 


486 MISS JOAN B. PROCTER ON THE 


Affinities. 

Testudo loveridgnt is undoubtedly closely related to 7’. tornieri 
Sieb. Even before the skeletal characters were known, when the 
former was thought to be without a bony carapace, it was recog- 
nized as being nearer to the latter species than to any other; 
in fact, Mr. Loveridge took his first specimens to be torniert. 
Comparison of their skeletons show that they are similar in 
structure. 

Briefly, they are probably both derived from the geometrica 
group, loveridgii being one step further removed than tornieri. 

T. tornieri differs from 7. loveridgu in that the carapace is not 
quite so depressed. The bony plates beneath are rather more 
developed, and slightly different in arrangement. It has a ninth 
pair of well-developed costal plates, and the whole animal is 
relatively much longer, being about half as broad as long instead 
of two-thirds. 

The supracaudal shield is also entire, instead of divided as is 
usually the case in loveridgw. It possesses, however, all the . 
special peculiarities of Joveridgii, including the deficient ribs, and 
fenestrated carapace and plastron, which produces a soft-shelled 
condition. It also appears, from its photographs, to have both 
jaws regularly denticulated. 

Siebenrock’s two specimens from Bussisi and Lindi are 
described as having soft shells, but he considered the condition 
of the carapace which he dissected away in one individual to be 
pathological (1904). The finding of a third specimen by Lonn- 
berg at Njoro (1911) and the recent discovery of loveridgii, 
however, prove that it is merely physiological. 


Habitat and habits. 


7. loveridgti is found in the arid rocky country round Dodomo 
and Tabora in Tanganyika Territory. Its distribution seems 
to be restricted to rocky kopjes where great boulders of grey 
granite are scattered; the characteristic vegetation is of the thorn- 
bush type; capers are also numerous. 

The tortoise is good at climbing up the face of the rock, and 
turns over easily after falling on its back. It can only be 
dislodged from crevices with difficulty, as it is able to inflate itself 
to a certain degree, and thus wedge itself firmly, using its legs 
as struts. It is also able to squeeze itself beneath stones, on 
account of the springy structure of its carapace. 

It is probable that it lays a single oval egg (about 35 mm. long 
and 23mm. in diameter), as such a one was recently dug up by 
Mr. Loveridge in the enclosure in which several females were 
penned. The embryo extracted from this egg is almost at the 
point of hatching, and [ have little doubt that it is loveridgii; 
but it is not possible to be certain, as females of 7. pardalis and 
Cinixys belliana were also in the pen, and tortoise-embryos 
are very similar to one another, and at this stage the more 
striking characteristics of loveridgu are not manifest. 


REMARKABLE TORTOISE, TESTUDO LOVERIDGII. A87 


Further particulars as to habits will be found in the notes 
by Mr. Loveridge appended to this paper. 


EXTERNAL CHARACTERS. 
Shape and proportions. 

The carapace of the adult is from three-quarters to four-fifths 
broad as long (actually 71 to 83 per cent.), excessively depressed, 
either flat or concave in the vertebral region; depth usually 
about one-fifth of the length. In text-fig. 1 (q¢. v.) the depressed 
carapace of loveridgii is compared with that of the normal zbera. 
Marginal region turning downwards and outwards abruptly, 
excepting the anterior border, which remains flat; marginals 
four to eight sometimes perpendicular. Sides straight, either 


Text-figure 1. 


Carapace of a normal Tortoise, T. ibera (a), compared with that 
ot T. loveridgii (b). 


parallel or oblique ; anterior edge of shell straight or forming 
a slightly inverted curve, feebly serrated; posterior boider 
rounded or obtusely pointed, also slightly serrated. Some- 
times the greatest width is in the pectoral and sometimes 
in the pelvic region, but in the majority it is equal at and 
between these two positions. This depends upon the oblique 
or parallel trend of the sides of the carapace. In the table of 
measurements the breadth is always taken at mid-carapace. 


488 MISS JOAN B. PROCTER ON THE 


The young are as broad as long and about one-third deep as 
long, and the carapace is uniformly convex and cordiform in 
outline. With growth these proportions change, the tortoise 
becoming narrower and more depressed. In order to show 
the precise nature of this change in form, the specimens in 
the appended table are graded according to age; the regular fall 
of the percentage values for breadth and depth are thus obvious, 
and could be well expressed by graphs. 

In both adults and young the dimension of depth does not 
properly apply to the carapace s. st7., but to the thickness of the 
entire animal. 

The plastron is large and projects beyond the carapace in front 
and falls short of it behind; fore-lobe truncate, hind-lobe deeply 
cleft, the two about equal in length and nicely shaped around 
the limbs; bridge very variable in width, usually about once and 
a half the length of either lobe. In the young the breadth of the 
entire plastron is greater in proportion. 


Coloration. 


The system of markings is similar to that met with in the 
geometrica group; the pattern, however, does not form regular 
stars, on account of the excentric positions of the areolars. These 
surfaces are reddish brown or rust-colour, black and pale horn- 
coloured bands radiating from them to the shield-edge; the rays 
are sometimes very strongly marked and regular, sometimes one 
colour predominating and sometimes the other, according to 
whether the black rays are broad or narrow, strongly marked 
or indistinct (vide Pl. I.). In the young the dark pigment is 
irregularly massed around each areolar. Each shield has its 
characteristic design, according to its contours; in well marked 
specimens that of the fifth vertebral resembles a rising sun, as 
depicted in decorative art; those on the marginals, the aster- 
pattern of old blue china. 

On the plastron the system is the same, but the yellow or horn- 
colour greatly predominates, partly owing to the larger areolars 
and partly to the fact that the black rays do not always reach to 
the shield-edge. 

The head and limbs are brownish or yellowish, sometimes 
indistinctly mottled. 

Owing to the burrowing proclivities of these tortoises, the live 
specimens usually have their beautiful markings entirely obscured 
by dirt, which lodges in the deep concentric striations of the 
shields. 

Mr. Loveridge notes that the specimens from Tabora are 
lighter in colour than those from Dodoma. 


Head. 
Head moderate; tympanum as large as or larger than eye: 
frontal usually broken up into four shields; prefrental large, 
subcordiform, divided in one specimen; a pair of large supra- 


489 


REMARKABLE TORTOISE, TESTUDO LOVERIDGII. 


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490 MISS JOAN B. PROCTER ON THE 


nasals meeting each other by a mesial suture of variable length, 
usually between one-fourth and two-thirds length of prefrontal 
(vide text-fig. 2a). 

Horny coverings of sheaths of alveolar ridges of maxillee and 
mandibles with regular, clearly-defined denticulations, 15 to 20 
(usually about 20) on each outer, and 6 to 9 (usually about 7) on 
each inner maxilla; 12 to 15 on each outer and 5 to 9 or 10 on 


Text-figure 2. 


- b. 
. a. Upper surface of head. (Adult; nat.size.) 1b. Side view of head. 
: ce. Horny teeth enlarged. 


each inner mandible. Beak tricuspid, with one or two of these 
denticulations between each cusp. Sometimes the points of these 
odontoids look precisely like the regular teeth of an Agamoid 
lizard ; but in other individuals they are not so well developed. 
They may be more so in the upper than the under jaw, or on the 
outer or the inner ridges, but whenever distinct are extremely 
regular (vide text-fig. 2b,¢). Im all cases they are more developed 
in the horn than in the underlying bone. 


Limbs. 


Lower part of fore-limbs covered with large claw- or scale-like 
shields, three very much enlarged on the anterior surface above 
wrist; normally five well-developed claws. 

Tibial portion of hind-limb often longer than radial portion of 
fore-limb, covered with irregular juxtaposed horny shields the 
size of those on the humeral portion of the fore-limb; scales on 
sole enlarged and claw-like in adults; normally four strong 
claws. In an abnormal specimen these claws are so overgrown 
that the outer one measures 13 mm. A group of enlarged 
tubercles on hinder side of thigh in some individuals. 

The limbs are not nearly so club-shaped as in most species of 
Testudo; both wrists and ankles are supple and defined by a well- 
marked crease in the skin; the foot projects considerably (vide 
Pl. I.). This is especially marked in the young. 


Tail. 


Tail normal, much longer and stouter in the male than in the 
female. 


REMARKABLE TORTOISE, TESTUDO LOVERIDGII. 491 


Epidermal shields. 


The epidermal shields are highly characteristic in form, but 
subject to great variation individually. In several specimens 
supernumerary shields and other abnormalities occur; for 


Text-figure 3. 


Epidermal shields of the carapace. (Adult; {nat. size.) 


instance, one adult has twelve marginals on the left side, one has 
twelve on both sides, and in one the posterior marginals form a 
deeply serrated border; the male type has only four vertebrals, 
one female has the third vertebral divided longitudinally, and 


492, MISS JOAN B. PROCTER ON THE 


two have extraordinary embossed vertebrals; three individuals 
possess an intergular. Mr. Loveridge mentions two specimens, 
which unfortunately escaped, in which small azygous shields 
occur, also several in which the supracaudal is single or semi- 
divided. 

Each shield is also subject to considerable individual variation 
of size and shape. 

Text-figure 4. 


aa 


' 


Sketch of carapace of male type-specimen, showing abnormalities 
in epidermal shields ($ nat. size). 


The shield surfaces in the young are excessively rugose or 
coarsely granulated, each one bordered narrowly by a raised band 
of smooth horn; with age these rugosities wear smooth, but the 
ensuing growth-rings, also ef smooth horn, are extremely regular, 


REMARKABLE TORTOIS£, TESTUDO LOVERIDGII. 493 


and the concentric striations between are deep and strongly 
marked. This applies to the plastron as well as the carapace. 

Nuchal shield very variable; much broader than long and cleft 
anteriorly in young specimens; about as long as broad to three 
times as long as broad, usually a little more than twice, and not, 
or but slightly, notched in adults. 

First vertebral subtetragonal, usually slightly longer than 
broad, or as Jong as broad; the anterior border longer than the 
posterior, both convexly curved ; lateral borders straight, slightly 
oblique. On the male type this shield is greatly elongated (vide 
text-fig. 4). 

Second and third vertebrals hexagonal, broader than long, the 
suture between them usually a little shorter than the length of 
either shield, a little longer than the suture between first and 
second or third and fourth. 

Fourth vertebral with very short posterior border, not more 


Text-figure 5. 


Young specimen, showing relatively broader shields (nat. size). 
a. Carapace. b. Plastron. 


than half length of suture between third and fourth. In the 
male type this shield is subtriangular or cordiform, and widely 
separated from the last vertebral by the fourth costals, 

Fifth vertebral obtusely pointed and flat anteriorly, in the 
same plane with the preceding vertebrals ; posteriorly its border 
is curved, and bent downwards sometimes very abruptly; the 
lower edge in some cases curves in so as to form an angle with 
the more obliquely set supracaudals. 

Costals highly characteristic in form, In the ordinary Testudo 
the sutures between these shields form straight lines more or less 
at right angles to the long axis of the carapace; in 7’. loveridgii 
they form curves, particularly pronounced in the case of thase 
between the fourth pair and fifth vertebral. The areolars are 
also excentrically placed. In 7. ibera, for instance, the border 


494 MISS JOAN B. PROCTER ON THE 


around the areolar is about twice as broad in front as behind, and 
situated high up in the shield, the distance between it and the 
vertebrals being less than the distance between it and the costal 
posterior to it, and a quarter or less the distance between it and 
the marginals; in fact, the relative widths of the shield-borders 


Text-figure 6. 


Epidermal shields of the plastron. (Adult; $ nat. size.) 


(top : back : bottom : front) are roughly expressed by the geo- 
metrical progression—2:4:8:16. In Z. loveridgii this ratio is 
better expressed—4:2:4:5 or 4:1:4:6, the areolar being 
mid way between vertebrals and marginals, but situated very far 
back in the shield. The areolar of the fourth costal is often 


REMARKABLE TORTOISE, TESTUDO LOVERIDGII. 495 


actually in contact with the anterior margin of the fifth verte- 
bral. This condition is not marked in the young, but is accen- 
tuated by each growth-ring. The fourth pair, in the male type, 
as already remarked, are broadly in contact mesially, in front of 
the last vertebral; in no other instance does this occur although 
they are sometimes in close proximity. The abnormal form of 
these shields on the male type is shown in text-fig. 4. 

The second costal is usually about equal in width to the third 
vertebral. 

The marginals are also extremely characteristic and very greatly 
reduced indepth. The first pair are normal in outline but a little 
longer than deep; the second and third are about one and a half 
times as long as deep; fourth to eighth greatly reduced, once and 
a half to twice as long as deep, as against about two-thirds long 
as deep in 7’. cbera; ninth and (or) tenth deepest, a little deeper 
than long; tenth and eleventh similar but slightly smaller. In 
the young the marginals are more or less uniform, as deep as 
broad or a little deeper; the sutures dividing them are somewhat 
oblique, making them rhombic in shape (vide text-fig. 5 a). 


Text-figure 7. 


Intergular shield (nat. size). 


The supracaudal is paired in all the specimens examined with 
the exception of a female living in the Zoological Society’s 
Gardens, and three young specimens, in which it is single. 

The epidermal shields of the plastron are not peculiar in any 
way (vide text-fig. 6). 

Gulars moderate, truncate and rounded anteriorly, forming 
together a bow-shaped edge, projecting beyond the carapace and 
forming a support for the animal’s chin; lateral edges shorter 
than the median suture; gulo-humeral sutures directed obliquely 
backwards and inwards, meeting each other at an angle of 
60° to 140°. 

An intergular is present in three out of twenty-three speci- 
mens. In the smallest specimen it is small but distinct (vide 
text-fig. 5 b); in one female it is somewhat smaller than a gular 
(vide text-fig. 7), and in a male it is well developed and as large 
as a gular. In all three it is protuberant and kite-shaped, the 
short sides in front. 


496 MISS JOAN B. PROCTER ON THE 


Humerals about twice as broad at humero-pectoral suture as 
from apex to apex, or their median suture. 

Pectorals narrow, median suture two-fifths to three-fifths 
length of median humeral suture. 

Abdominals more or less square, median suture two to four 
times length of median pectoral suture. 

Femorals very variable, median suture usually longer than 
median pectoral suture ; postero-lateral corners projecting beyond 
lateral edges of anals. 

Anals.—Ano-femoral sutures slightly oblique, directed for- 
wards; lateral borders at right angles to them; median sutures 
short, each anal cut away posteriorly and ending in a pronounced 
point of variable shape, the resulting cleft between them forming 
an angle of 90° to 120°. The width and depth of this cleft is 
apparently not dependent on sex, as one would suppose from the 
great difference in the relative sizes of their tails. 

The inferior surfaces of the marginals which cover the 
bridges are extremely narrow, from three to four times as long 
as deep. 

SKELETON. 

Besides the complete skeleton of one adult, and the radiographs 
of the types, a considerable amount of accessory material has 
been available. An excessively interesting individual, in which 
development had been arrested at a stiil earlier stage, decomposed 
in transit, thus enabling me to use most of the skeletal parts for 
study. In this specimen the bony plates were as thin as tissue- 
paper and extraordinarily flexible. Parts of other tortoises 
also disintegrated and formed interesting checks on the first 
skeleton. 

Of the young, an X-ray plate shows that the limb-girdles are 
normal, as in the adult. The structure of the carapace has been 
studied in a series of six, ranging from 42 to 82 mm. in length; 
in these it was dissected off and the inside aspect cleaned. <A 
complete preparation of both carapace and plastron was made 
from a specimen 60 mm. in length, and the plastron of the 
42 mm. individual studied, without its removal, from the inside. 

This material was amply sufficient to show the normal structure 
and development, besides some interesting variations of the 
skeleton of 7’. loveridgw, and to form highly interesting com- 
parisons with other species. 

The word “plate” is repeated after the name of each dermal 
bone, in order to distinguish them from the epidermal shields of 
the same names. 


The bony carapace of the adult. 

In general appearance the bony carapace is extraordinary. 
Such portions of the neural and costal plates as are present, are 
formed of translucent bone, and when damp are springy and 
flexible, making it possible to depress or elevate the carapace as 
if it were supported upon springs. 


REMARKABLE TORTOISE, TESTUDO LOVERIDGII. AQT 

Where the epl idermal ae oe for tures ae each cae the 

bone a eath is dee see ved, som mene s forming per ions, 

but ee ey tea ven Sain Gale Pls. “ae © IIL. ond 
te as is. 8). 

eath ae ertebr. al ae Pe er a e fenestree shaped 


like TR ae ri S aa eee and fourth 


ee a | 


Os a 


/ —————— 
f ———3 
Anmepl Se uma 


[ om WWAg z = 
(ya UD a ee 


marginal plates, mp.; nuchal, nw. ; 
divided mesially each by a bony bridge for oe OLY udimentary 
eural plates; the first and fourth ane sma aie mpat a wi ith ft ne 
second an ird (wide text-fig. 8). 
Beneath the costal shields there are also lar ae milar 
in shape to the shields under which they are situ ie a "Tha t 


498 MISS JOAN B. PROCTER ON THE 


under the first is entire; those beneath the second and third 
divided mesially by a narrow transverse bridge of bone formed by 
the third and fifth costal plates. There is no large fenestra 
beneath the fourth in the normal adult. 

Nuchal plate moderate, posterior corner free, forming anterior 
edge of first median fenestra, which exposes the ninth vertebra. 

First neural plate (text-fig. 8, np.1) very much reduced in 
comparison with that of a normal Testudo; anterior edge pointed, 
bordering first fenestra by which it is separated from the nuchal 
plate ; narrow lateral limbs, uniting by short sutures with first 
pair of costal plates (cp. 1); narrow posterior limb, entering into 
second median fenestra, where it forms part of the dividing 
bridge and unites by short suture with the second neural. 

Second neural plate (np. 2) vestigial, barely wider than the 
vertebre beneath, with straight lateral edges, uniting by short 
sutures with preceding and succeeding neurals, and forming the 
main part of the fenestra! bridge. 

Third neural plate (np. 3) octagonal, very much reduced, oblique 
edges free, curved invertedly, bordering the inner corners of the 
second and third fenestre, which this plate divides one from the 
other by uniting with the second costal plates (cp. 2) by narrow 
sutures, thus forming a narrow bridge between them; also form- 
ing similar connections with second and fourth neurals. 

Fourth neural plate (np. 4) similar to second but somewhat 
broader in proportion, forming the longitudinal dividing bridge 
across the third median fenestra. 

Fifth neural plate (2p. 5) precisely similar to the third. 

Sixth neural plate (wp. 6) similar to second and fourth but 
broader still, dividing fourth median fenestra, which is, however, 
much smaller than the two preceding. 

Seventh neural plate (np. 7) normal, hexagonal, without free 
edges, uniting by six sutures with the sixth and eighth neural, 
and seventh and eighth costal plates respectively; those formed 
with the eighth costals slightly longer than the others. 

Highth neural plate (mp. 8) normal, sub-tetragonal, without free 
edges, uniting by oblique sutures with the eighth costal, and by 
transverse ones with the seventh neural and first pygal plates, 
the latter suture the longer and curved. 

First pygal plate broadly cordiform or septagonal, set almost 
at right angles to the horizontal plane of the neurals, curving in 
where the marginal epidermal shields meet the fifth vertebral, 
and curving out more obliquely to form a point beneath the supra- 
caudal shields. It forms sutures with the eighth neural and 
costal, eleventh marginal (mp. 11), and second pygal (p. 2) plates, 
that with the eighth neural about twice as long as any of the 
others and curved invertedly. 

Second pygal normal, equal in size to the eleventh marginal 
plate. 

First costal plates (cp. 1) shaped like wings, uniting by short 
sutures with the lateral edges of first neural (which forms a 


REMARKABLE TORTOISE, TESTUDO LOVERIDGII. 499 


bridge between them) and second costal plates, thus separating 
the first lateral fenestra from the first and second median 
fenestra on each side; the broadly curved anterior edges form 
an uninterrupted suture with part of nuchal, whole of first and 
second, and inner corner of third marginal plates in most cases, 
but sometimes the distal portion is undeveloped, and therefore 
less wing-like, or cleft, or perforated beneath the shield-sutures 
(vide text-fig. 9). 


Text-figure 9. 


Left first costal plate from beneath, showing perforation beneath vertebro-costal 
shield-suture ; distal end incompletely developed, and cleft. (Nat. size.) 


Fenestre between first and second costal plates are from frout 
to back as long as one marginal, from side to side two or three 
times this length. 

Second costal plates (cp. 2) slender, feebly developed, widely 
separated from the neural; anterior and postericr heads forming 
narrow sutures with first and third costal plates beneath costo- 
vertebral shield-sutures, main arms supporting transverse costal 
shield-suture, and uniting distally with the fourth marginal plate. 
Length of each (measured from end to end) three times breadth 
(measured from front to back); median head forming a short- 
pointed process directed inwards, but widely separated from the 
vestigial second neural plate. The X-ray plates of the types 
show that these processes are sometimes connected with the 
vestigial rib-heads by the ligament-like periosteal tissue of the 
abserbed rib. 

Third costal plates (cp. 3) extremely slender, rib-like, and feebly 
developed. Main arms half or less than half breadth of those of 
second, from which they are separated by a fenestra about half 
as broad as those preceding. Hach has three heads, each one of 
which is twice as broad as the main arm, the median uniting by 
suture with the third neural plate, thus forming a bridge between 
second and third median fenestrz on each side; anterior and 
posterior heads directed obliquely outwards, similarly united with 
the corresponding heads of second and third costal plates, which 
are directed obliquely inwards beneath the costo-vertebral shield- 
sutures. 

In the specimen in which development has been arrested at a 
Proc. Zoou. Soc.—1922, No. XXXV. 35 


500 MISS JOAN B. PROCTER ON THE 


very early stage, only the merest traces of bone-deposit are 
present in place of the rib-like arms of these plates (vide text- 
fig. 10). 

Fourth costal plates (cp. 4) in form precisely like second, twice or 
more than twice width of third, sometimes with very ragged edges ; 
widely separated from neural by third median fenestre, forming 
a suture with sixth marginal plates, sometimes with corner of 
seventh also. Fenestra between third and fourth similar in size 
and shape to that between first and second, about twice width of 
that between second and third. 3 

Fifth costal plates (cp. 5) precisely like third in every way, 
connecting by suture with seventh marginal plates. Fenestra 


Text-figure 10. 


Carapace of adult arrested at an earlier stage in development. 
(Left side, from within ; 4 nat. size.) 


between fourth and fifth similar to that between second and 
third, about half width of that between third and fourth. In 
the poorly developed specimen already referred to (text-fig. 10) the 
rib-like limb is represented by a few bone-granules in isolated 
patches. 

Sixth costal plates (cp. 6) similar to second and fourth, but 
rather more developed. Anteriorly narrowly in contact with fifth 
beneath the vertebro-costal shield-sutures, otherwise separated 
by fenestre slightly narrower than any of the preceding. Central 
head in closer proximity to the neurals than those of second 
and fourth; the dividing fenestra about half as large as in 
the former instances, Posterior edge quite straight, forming 


REMARKABLE TORTOISE, TESTUDO LOVERIDGII. 501 


a suture with seventh costal plate throughout its length from 
fenestra to ninth marginal, excepting in the abnormal specimen, 
in which it is partially separated from the seventh by an 
elongated fenestra. 

Seventh costal plates (cp. 7) straight, with angular head and 
pointed distal end which is projected some distance into middle 
of ninth marginal. Anterior facet of head free, bordering fourth 
vertebral fenestra, posterior forming suture with seventh neural 
plate; main anterior border forming suture with sixth costal 
from end to end, posterior border forming a similar suture 
with eighth. On tke upper third of the suture between the 
seventh and eighth costal plates there is a small oval fenestra, 


Text-figure 11. 


Diagram of marginals 5, 6,and7. a. 7. loveridgii. b. T. ibera. 


smaller on one side than on the other in the individual figured, 
but of variable size in others. It is always very ragged in 
outline. 

Eighth costal plates (cp. 8) pentagonal, broad, with the exception 
of the very small fenestra described above, completely in contact 
with the surrounding plates—7.e., seventh costal, seventh and 
eighth neural, first pygal, ninth, tenth, and eleventh marginal 
plates. A small triangular process is projected into the anterior 
portion of tenth marginal. A small kidney-shaped fenestra with 
jagged edges pierced above the ilio-sacral regicn, anteriorly 
reaches just beyond the groove made by the fourth costal and fifth 

30* 


502 MISS JOAN B. PROCTER ON THE 
vertebral shields and proceeding backwards almost to the eighth 
neural. 

Presumably this fenestra and the small oval one between the 
seventh and eighth costals are all that remain of the large kidney- 
shaped one, similar to those between the preceding costal and 
neurals, which is present in youth. 

The marginal bones are somewhat wider than the corresponding 
shields; first, second, and third markedly so; sixth to tenth deeply 
notched in the middle for the reception of spurs from the corre- 
sponding costal plates. Fifth, sixth, and seventh enter imto the 
composition of the plastron beneath, and will be dealt with as 
part of the plastron. 

When one considers the great depth of the lateral marginais 
in ordinary tortoises in comparison with their abnormal shallow- 
ness in 7’. loveridgii, the shallow carapace of the latter is to 
some extent explained. In text-fig.11 the fifth, six,and seventh 
marginal plates, drawn diagrammatically, of 7. loveridgu and 
T. ibera ave compared. 


The bony plastron of the adult. 


The bony plastron is also composed of bone thin to trans- 
lucency. In the centre there is an enormous diamond-shaped 
fenestra reaching from ento- to Xiphiplastron, and side to side 
from marginals to marginals. It is very ragged in outline, 
variable in size, and lies beneath the pectoral and abdominal 
shields, so that the greater part of the abdomen is unprotected 
by bone (vide text-fig. 12). 

Epiplastrons (ep.) shapely, oblique, moderate in size; posterior 
borders serpentine. 

Entoplastron (ené.) in the shape of a diamond, the anterior and 
posterior corners of which have been truncated ; posterior margin 
free, corresponding with the humero-pectoral shield-suture above, 
entering into the central fenestra, into which a sharp bony spur 
projects, representing the tip of the sternum, which is fused to the 
inner surface of this plate in the normal way. 

Hyoplastrons (hp.) very small, rectangular, obliquely set; 
separated from each other by the anterior angle of central 
fenestra; widely separated from the hypoplastrons by marginal 
plates. 

Marginal plates (mp.) 5, 6, and 7 enter into the composition 
of the plastron on each side of the fenestra, spreading obliquely 
inwards to a most unusual degree, number 9 running beneath the 
pectoral shields to the abdominal, and terminating with a free 
oblique edge in the central fenestra; n umber 6 borders the lateral 
corner of the central fenestra on each side. 

Hypoplastrons (Ayp.) completely separated from each other by 
the posterior angle of the central fenestra; produced to form 
part of the posterior plastral lobe, 


REMARKABLE TORTOISE, TESTUDO LOVERIDGII. 503 


Xiphiplastrons (zp.) form slightly oblique sutures with the 
hypoplastrons; their anterior median corners enter into the 


Text-figure 12. 


Bony plastron of an adult, viewed from witheut (4 nat size). 
Fenestra |{ |; deep grooves beneath epidermal shield-suture -2-2-2===> * 


epiplastrons, ep.; entoplastron, enp.; hyoplastrons, hp. ; Inaperalbositnaray, in 
xiphiplastrons, zp.: marginal plates, mp. 


posterior corner of the central fenestra, which separates them 
from each other anteriorly; a broad open cleft posteriorly ; 
lateral edges bi-lobate. 


* The groove beneath the median longitudinal shield-sutures is equally deep, but 
is not marked, as it would obscure the bony ones beneath. 


504 MISS JOAN B. PROCTER ON THE 
Skull. 


Differing from the common TZ. ibera in the following 
particulars :— 

The prefrontals are larger than the frontals, and widely 
separated from the post-frontal bridge; temporal and post-orbital 
arches stouter in proportion, especially the latter; annulus tym- 
panicus larger, stapes as slender as a hair. 

Maxillary with three alveolar ridges, the inner one of which is 
comparatively weak, the outer one, or maxillary edge, denticulated 
with what appear to be minute pointed teeth of the acrodont 
type, but which have not the composition of true teeth; the number 
is about 20 on each outer, and about 7 on each inner, maxillary 


Text-figure 13. 


€, 


Skull of adult (nat. size). a. Palatal view (horny sheaths not blackened). Ib. Side 
view (horny sheaths blackened). Horny sheath of maxillary partially dissected 
away to show denticulations in the bone itself. @. Mandible (horny sheaths 
blackened). 


ridge. One or two similar denticles between the cusps of the 
premaxillary. These denticles are so fragile that if the adherent 
horny sheath is removed in the usual way, they are completely 
destroyed, but they can be clearly seen without a lens if the horn 
is partially removed. In text-fig. 13 this has been done, and the 
remaining horn blackened in the profile view, in order to dis- 
tinguish it from the maxilla. 

All the cranial foramina seem to be proportionately small. 

The mandible has both its alveolar ridges denticulated in the 
same manner as the maxilla. In the figure the horny sheaths, 
in this case entire, are blackened. 


Vertebral column. 

First eight (cervical) vertebre as in 7’. ibera. 

The dorsal vertebre (ninth to eighteenth in the complete 
series) differ profoundly from the ordinary pattern; they are 
entirely without spinous processes and the centra are greatly 
depressed instead of compressed, and hollowed to a thin shell 
for the reception of the spinal cord, the vertebre being thus a 
simple depressed tube, adherent to the bony neural plates of the 
carapace. In all other species the vertebral column is separated 


REMARKABLE TORTOISE, TESTUDO LOVERIDGII. 505 


from the carapace by wide arches, the contact with it being 
formed by long thin neural processes; In no case is the main 
column, even narrowly, in contact (excepting 7”. torniert). 

It is in some cases impossible to detect what remains of the 
neural arch in a section, for it has degenerated to such an 
extent that in some individuals it appears to be wanting 
altogether. Possibly it is absorbed to a variable extent in the 
same manner as the ribs. In the vertebral sections figured (vide 
text-fig. 14), it seems to be represented by an extremely thin 
layer beneath the neural plate, forming a meagre roof to the 
neural canal. ‘The specimen to which it belongs, however, is the 
most degenerate of any that have been examined. More normal 
individuals, however, cannot be spared for the necessary dissec- 
tion. Figs. ¢ and d show diagrams of sections through the 
middle and towards the anterior end of the eleventh vertebra ; 


Text-figure 14. 


Diagrams of mid-dorsal vertebra. a. Anterior end. b. Posterior end. @. Section 
towards one end. d. Section through middle. e. Side view of -vertebre with 
lateral expansions of neural plates cut off. f£. Two vertebree bent apart, to 
show thinness and flexibility of the combined neural plate and neural arch. 
Epidermal shield, es; dermis, d@; neural plate, np; neural arch, xa; centrum, c. 


figs. a and b, the vertebra viewed from each end; fig. e, the 
twelfth and thirteenth with part of eleventh, side view, with 
the lateral portion of the third neural plate cut through; fig. f, 
the twelfth and thirteenth, bent apart to show the thin flexible 
covering of the neural canal. The relative positions of the 
articulation of the vertebre and the neural plate sutures vary, 
usually alternating with each other. The points of juncture 
between the 5th and 6th neural plates and 14th and 15th 
vertebre, however, coincide, one above the other. 

The ninth vertebra is, of course, highly specialised for the articu- 
lation of the neck. The facets for the articulation of the first 
(non-functional) ribs are situated at the extreme anterior end. 
Vertebree ten (2nd dorsal) to thirteen are progressively more 
elongate, and have the tubercles for the articulation of the rib- 
heads at the extreme anterior end on each side as in the ninth: 
in some of the young, articulation takes place at the juncture of 
each consecutive pair of vertebre. Fourteenth to seventeenth 
become progressively shorter, and the facets for the corre- 
sponding rib move further backwards in a lateral ridge as the 
ribs become directed obliquely in this direction. Highteenth 


506 MISS JOAN B. PROCTER ON THE 


nineteenth, twentieth, and twenty-first vertebre progressively 
shorter, their diapophyses elongate, slender, distally in contact 
aud supporting the iliac crests, those of the nineteenth the longest. 
Diapophyses of anterior caudal vertebra also elongate. Caudals 
normal, twenty-four in number, stouter in the male than in the 
female, according to the radiographs. 


libs. 
The ribs may be considered as absent in the adult, the part 


Text-figure 15. 


NU 


oS oe 
aD - | “Gp 
oe 2S 
lp Sea 
eee 
ee 


Bony carapace of an adult, from within (# nat. size), showing vertebre, and 
vestigial ribs. (Ribs not entirely absorbed in this specimen.) * 


normally adherent to the carapace having become completely 
absorbed. The free or capitular portion is sometimes present as 


* Sutures between costal plates 2, 3 and marginal plates 4, 5, accidentally omitted. 


REMARKABLE TORTOISE, TESTUDO LOVERIDGII. »: 507 


a vestigial thorn-like structure, but is more often absent 
altogether, or represented by a thin ligamentous simulacrum 
(vide Pls. IT. & ITI.). 

The first pair, which in all tortoises are rudimentary, are 
normal. The second (first functional) have the free capitular 
portion sometimes present, although reduced, adherent terminally 
to the first costal plate, and in contact with the first pair in the 
usual manner (vide text-fig. 15). Third pair absent or vestigial. 

Fourth pair, when present, similar to the second, but usually as 
slender as a hair; in contact with third pair of costal plates. 
Fifth pair absent or vestigial. Sixth pair like the fourth, but still 
more vestigial and directed slightly backwards. Seventh pair 
absent or in some specimens present, and bridging the fenestre 
and meeting the corresponding process of the sixth costal plates, 
directed slightly backwards. Eighth pair rather depressed, 
directed still more backwards, in contact with the seventh pair 
of costal plates just in front of the seventh neural plate. 
Ninth pair slender, directed strongly backwards and flattening 
out beneath the head of the eighth costal plates. 

In the young specimen (60 mm. long) deseribed in the next 
chapter, a tenth pair of ribs are present. 


Pectoral girdle. 
The bones of the pectoral girdle and fore-limbs, as compared 
with those of 7’. ibera, ave slender, and the coracoids are not so 


strongly dilated ; in other respects they are similar (vide Pls. LI. 
& ITT.). 


Pelvic girdle. ; 

The bones of the pelvic girdle and hind-limbs, also compared 
with those of 7’. ibera, are somewhat more slender and those of 
the pelvis less dilated (vide Pls. I1.&11I.). The ilia are not fixed 
above as in the ordinary tortoises, as they terminate just beneath 
the last median fenestra in the carapace, whilst in tortoises with 
a, solid carapace they articulate with a bony prominence beneath 
the eighth costa] plate. 


DEVELOPMENT. 

Until April of this year, only adult or extremely young speci- 
mens were available, and it was, therefore, only possible to guess 
at the intermediate skeletal forms. Mr. Loveridge has, however, 
now sent over a series of young and half-grown, which illustrate 
the processes which are taking place and corroborate my earlier 
speculations. 


Ribs. 

In the young the ribs are perfectly normal (vide text-figs. 17, 
18) but step by step they disappear. The portion adherent to 
the carapace becomes flattened, and disappears in a manner 
similar to that met with in other members of the genus. The first 
portion to become completely absorbed is that which underlies 


508 Miss JOAN B. PROCTER ON THE 


the oblique connecting bridges between the costal plates, which 
in turn underlie the vertebro-costal epidermal shield-sutures: 
that is to say, the point where the rib becomes adherent. 

The capitular free portion of the rib absorbs in a peculiar way. 
At 71 mm. these free ribs still appear to be present when the 
carapace of a spirit specimen is raised; when explored, however, 
they prove to be chiefly composed of soft pliant material, like 
ligament. At 81mm. there is stilla splinter-like rib within this 
soft material, but this becomes more and more wasted by 
absorption, until it is lost absolutely in the adult, or represented 
by a tiny thorn-like splinter one or two mm. in length—the 
extreme capitulum of the original rib. Mr. Burne has very 
kindly investigated the structure of this soft portion, excised 
from a young specimen. Longitudinal sections show that with 


Text-figure 16. 


Microscopie section through capitulum of absorbing rib (X 100), in which almost 
nothing but periosteum is left. (After a sketch by Mr. Burne.) Periosteai 
fibres, p; cartilage, c; areas of calcification, b. 


the exception of two small areas of calcification in the capitulum 
itself, no bone is left (vide text-fig. 16). These areas are situated 
in a part of the original cartilage; the rest of the section shows 
nothing but parallel bundles of what appears to be connective 
tissue. Near the edge of the sections there are, however, a row 
of multi-nucleated cells which Mr. Burne considers to be osteo- 
clasts. He says: ‘‘i presume what happens is that the bone 
after being formed in the rib is absorbed by these osteoclasts, 
and the periosteum—possibly thickened—is left, forming a simu- 
lacrum of the rib.” There is no doubt in my mind that this is the 
right interpretation ; also that this process of absorption, carried 
only to an early stage, is responsible for the excessive slender- 
ness of this portion of the ribs in other species. 


The carapace. 


In a speeimen 42 mm. in length, probably newly hatched or 
only a few months old, the plates are commencing their develop- 
ment, and areata particularly interesting stage (vide text-fig. 17). 

The nuchal is more or less fully formed within the marginal 
ring, but does not extend beyond. The neurals, excepting the 
third and fifth, consist of minute oval areas of yellowish bone- 
granules towards the anterior of each underlying vertebra. The 
third and fifth are laterally dilated to some extent, the fifth being 
almost in contact with the head of the fifth costal plate. These 
two neurals underlie the two middle vertebral shield-sutures. 
The thin layer of coarse bone-granules which composes each 


REMARKABLE TORTOISE, TESTUDO LOVERIDGII. 509 


one of them is distinguishable without a lens. In pulling up the 
carapace, quite half of these plates separated easily from the 
vertebre and adhered to the carapace, or rather to the dermis, 
which shows that they are not completely fused with the vertebree 
at this stage. 

The costals are slightly more developed, and are disposed as 
follows :—The vertebral and costal epidermal shields meet each 
other in a zigzag suture which has seven angles. At each of 
these, three shields meet corner to corner. It is beneath these 
points that the centres of the incipient costal plates are located. 
They are therefore alternately nearer to, and further from, the 
vertebre. ‘Those nearer to them tend to develop proximally, and 
are more or less rhomboidal in shape, and those further away 
develop distally down the rib, and are more triangular, the apex 
pointing towards the marginal region. 

The first, second, and third are not quite in contact with each 
other, the third, fourth, and fifth are nearly in corner to corner 


Text-figure 17. 


Carapace of newly-hatched young, from without (nat. size). 
Epidermal shield-sutures «---------------------~ ; vertebrae and ribs :::%:::. 


contact, the fifth and sixth are joined by oblique suture as in the 
adult, forming an oblique bony bridge beneath the shield-suture. 
The seventh and eighth are but commencing their formation. 

The pygals are absent altogether. 

The marginal plates are not recognizable as such at this stage, 
but minute groups of bone-granules are detectable beneath the 
shield-sutures between marginals three to eight. Posteriorly 
they are not visible, although their primary particles may be 
present in the dermis. 

At about 60 mm. the eight pairs of ribs (2nd to 9th) are still 
distinct and of normal length, but are modified, the proximal 
5 mm. having become very slender, and the remainder flattened 
and adherent to the dermal costal plates where these are present. 
In the particular specimen figured a ninth pair of ribs are fully 
developed, and reach the marginals on both sides. 

At this stage (vide text-fig. 18) the development of the 


510 MISS JOAN B. PROCTER ON THE 


dermal bones of the carapace is well advanced, showing most 
clearly how the carapace of the adult is formed. The anterior 
and median portion is developed almost as much as in the adult, 
but posteriorly large fenestre occur beneath the fourth costal 
and fifth vertebral shields, which (fenestra) are almost entirely 
roofed in with bone in the adults. 

The nuchal and marginal plates are fully formed. The neurals 
are of the adult pattern, but they are narrow, thin, and feebly 
developed anteriorly, becoming progressively broader posteriorly, 
the seventh and eighth are the broadest, followed by a much 
narrower first pygal. The first pygal is not well developed, and 
is separated from the last costal plate on each side by a very wide, 


Text-figure 18. 


SN 


/ 


oe 
Ih i (| 
A 


Carapace of young specimen 60 mm. in length, from within (% nat. size). 


subtriangular fenestra, which is represented in the adult by a 
small one of ragged outline in the last costal plate itself. 

From the study of this one stage it is easy to reconstruct the 
development of the bony carapace of the adult, and, by logical 
conclusion, to suggest a manner in which a solid carapace can be 
formed. 

Costal plates one to six are similar in form and arrangement 
to those of the adult, excepting that they do not reach the mar- 
ginal rib by about 3mm. In the case of the first, the shortness 
of the plate causes a small fenestra to be discovered between 
the underlying rib and the anterior marginal plates. 

At this stage one sees the relative positions of true ribs and 


REMARKABLE TORTOISE, TESTUDO LOVERIDGILI. 511 


costal plates, which are not clear in the ribless adults. The 
main limbs of the second, fourth, sixth, and eighth costal plates 
ave very broad, narrowly bordering the underlying rib anteriorly 
but projecting widely beyond its posterior edge. The alter- 
nating third, fifth, and seventh are but feebly developed, and 
are very little broader than the underlying rib, which they cover, 
and border narrowly and equally. This alternating arrangement 
of the breadth development, which takes place posterior to the 
companion rib, accounts for the relative widths of the lateral 
fenestre in the adult, in which the slender costal plates are 
preceded by a narrow fenestra, and succeeded by one at least 
twice as broad, the general effect being a single fenestra the 
ishape of the epidermal shield, bridged by a slender rib-like 
costal. The same principle applies to the formation of the plates 
beneath the vertebral shields. In this case the first, third, fifth, 
and seventh neurals are well developed (progressively as already 
noted), and form lateral sutures with the corresponding well- 
developed heads of the corresponding costal plates. The broad 
bridge of dermal bone thus formed is over and posterior to the 
free underlying rib-head. This makes the segment of the median 
fenestra immediately following, half the width of the remaining 
segment on each side of the vertebre, a similar arrangement to 
that met with in the case of the lateral fenestrae. The alter- 
nating second, fourth, and sixth neural plates are feebly developed, 
barely wider than the underlying adherent vertebrie, and are 
widely separated from their corresponding costals, which, though 
they are broadly developed distally, are not continued proximally. 
The large kidney-shaped fenestra in the dermal bone is, however, 
bridged by the true rib beneath; a slight deposit of dermal bone 
on the distal end of this rib-bridge forms the feeble, pointed apical 
head of the costal plate which projects into the fenestre ata 
later stage when the rib itself has absorbed. 

Thus the broad neuro-costal bridges are continuous with the 
slender main-limbed costals, and the undeveloped neurals are con- 
nected by the free portion of the rib with broad costal main limbs. 
The bory bridges formed by the oblique heads of the costal plates, 
and which separate the median from the lateral fenestre (beneath 
the vertebro-costal epidermal shield-sutures), are about as wide as 
their neighbouring broad costal or neural plates. In the same 
way narrow anterior sections of the median fenestree are opposite 
the wide and posterior sections of the lateral fenestrae, and vice 
versc. 

In the young of this size there is one great difference in the 
stage of dermal bone development, namely that of the seventh and 
eighth costal plates. In the adult, the seventh, although narrow, 
normally forms an uninterrupted suture with the sixth and eighth 
from the last median fenestra to the marginal region. At the 
present stage, the seventh is similar to the slender third and fifth 
costals, and the eighth is similar to the broad second, fourth, and 
sixth costals, the seventh being separated from the eighth by a wide 


boy) MISS JOAN B. PROCTER ON THE 


fenestra, represented in the adult bya small fontanelle, and from 
the sixth by a narrow one, absent in the adult ; the latter fenestra, 
however, is divided into two by the contact of the posterior corner 
of the end of the dilated sixth with the seventh costal, about 3 mm. 
from the marginal plates. This is caused by the relative positions 
of the implicated ribs, which become closer together posteriorly 
as the vertebrae become progressively shorter. For this reason, the 
eighth costal plate in this specimen completely roofs over the ninth 
and tenth (abnormal) ribs, the intervening space between them 
being no broader than the entire second costal plate (compare 
text-figs. 18 and 15). 

Therefore, in the first instance (42 mm.) it is the central portion 
of the carapace which is the most developed ; later (60 mm.) the 
plates are more or less equally developed all over, with the exception 
of the neurals, which are progressively enlarged posteriorly. Con- 
tinuing up the series (71, 81,and 89 mm.), the first, second, fourth. 
sixth, and eighth costal plates expand in length until they form 
sutures with the marginals; the third and fifth become the slender 
rib-like plates of the adult form; and the posterior portion of the 
carapace from the seventh costal to the pygal plates becomes 
roofed in with bone by the expansion of the seventh and eighth 
costal plates. 

The anterior part of the carapace in the adult, therefore, is 
arrested at an earlier stage in its development than the posterior. 


The plastron. 


At 42 mm. the plastron is in the initial stage of development 
(vide text-fig. 19); all nine bones are present, but very small, 
thin, and for the most part widely separated from each other. 
The epiplastrons are merely small strips of bony deposit beneath 


Text-figure 19. 


Plastron of newly-hatched young, from without (nat. size). 


the anterior portion of the pectorals. The entoplastron is better 
developed, and subtriangular in shape. The hyoplastrons are 
ragged in outline, and not in contact with epiplastrons or 


REMARKABLE TORTOISE, TESTUDO LOVERIDGII. 513 


entoplastron. Hypo- and xiphiplastrons narrow and in sutural 
contact, forming a narrow, bony band running obliquely across 
the hind lobe. The xiphiplastrons, however, do not meet each 
other. 

In the 60 mm. specimen (vide text-fig. 20) the median fenestra 
is much broader in proportion than in the adult, reaching almost 
from edge to edge of the plastron. Marginal plates five, six, and 
seven, which in the adult are produced inwards for a considerable 
distance, only overlap the line of their corresponding epidermal 
shields to a small extent, and are about equal, not graduated 
in depth as in the adult (wide text-figs. 12 & 20). The epi- 
plastrons are more or less pentagonal and not well developed 
posteriorly at their outer corners, but are in contaet with the 


Text-figure 20. 


iW: 
ans 


. 
| 


z 
ee Uta 


IN ~, i | 
ww iy 


Plastron of young specimen, 60 mm. in length, from within (nat. size). 


hyoplastrons and the entoplastron. . Hyoplastrons narrow and 
not in contact with the entoplastron, or with the fifth marginal 
plates, with which in adults they form long oblique sutures. 
Hypoplastrons slender, widely separated from each other, and 
only narrowly in contact with the sixth marginal plate; they 
form short sutures with the equally slender xiphiplastrons. 

Further development takes place principally on the inner 
borders of the plastral plates until the central fenestra has 
assumed the regular diamond-shaped form of the adult in this 
species (compare text-figs. 19, 20, and 12). 


514 MISS JOAN B. PROCTER ON THE 


Discussion. 


The morphogeny of the carapace and plastron. 

The primitive plastron, therefore, and the delicate network of 
dermal bone which forms the carapace of the adult Yestudo 
loveridgii, are brought about by arrested development. The 
fenestrze have not been formed in the completed structures by 
absorption, and do not increase in this manner with age as 
supposed. ‘The relative thickness or breadth of the plates depends 
on individual variation, specimens of abnormal fragility having 
had their development in this respect cut short at an earlier 
stage than the average individual. This principle also applies 
to such species as Testudo polyphemus, in which the carapace is 
said to become thin and form fenestrae with age. A specimen 
in the British Museum Collection of 210 mm. (76. 1.36.6) and a 
half-grown specimen of 135 mm. (73.8.13.25) have both 
carapaces and plastrons fenestrated in a manner similar to those 
of 7. loveridgii, whilst one of 225 mm. has them thin but entire. 

The ribs of the latter on the other hand, and seemingly the 
neural arch of the vertebrae, do absorb with growth; the ribs 
completely in most cases. Thus the bony plates are developing. 
whilst the ribs are degenerating by osteoclastic absorption—a 
process which is, of course, physiological, and in fact, as far as 
the ribs are concerned, carries to a logical conclusion a tendency 
which is well marked throughout the genus. 

The question which now arises is: what has led to the extra- 
ordinary degree of depression of the carapace ? Some individuals 
are positively concave above, and all have a markedly deformed 
and rickety appearance. 

Flattening is evidently not brought about, at any rate to this 
extent, by a fenestrated or thin condition of the carapace, for the 
giant tortoises which pass through these stages in the course of 
development + and have very thin carapaces, are remarkably convex 
in form. The only visible result here is the crumpled appearance 
of the shell of 7’. abingdonti. T. polyphemus, on the other hand, 
is distinctly flattened. 

Tt can be argued on the one hand that the flattened carapace is 
brought about by the habit of living beneath stones and squeezing 
into rock-crevices. This habit, induced by environment, would be 
bound to have a modifying e Breen: for, during youth, ale develop- 
ment of a domed and solid carapace would be interfered with by 
the constant application of pressure, and in a sufficient number 
of generations the ability to form a normal carapace might be lost 
altogether. The fact that the Burrowing Tortoise, 7. polyphemus, 
has a thin or fenestrated and somewhat fancy carapace 
supports this view. Could this be proved experimentally, it 
would furnish a convincing argument in favour of the heritance 
of acquired characters. 

On the other hand, it can be equally well maintained 
that an inherited tendency to the arrest in development is 


REMARKABLE TORTOISE, TESTUDO LOVERIDGILI. 515 


orthogenetic, brought about either gradually or as a mutation, and 
that the furtive habit of hiding beneath stones was the natural 
result, since the tortoise no longer possessed adequate protection 
from enemies. 

Possibly both principles come into play, the reduced armour 
and loss of ribs being orthogenetic, and the depression and relative 
condition of the vertebre being subsequently induced by the rock- 
dwelling habit. 

Again, the metabolism of bone formation may be to some 
extent influenced by climatic conditions, lack of lime in the 
soil or water etc., or by their food. In this connection it is 
interesting to note that Mr. Loveridge has recently found two 
specimens of Cinixys belliana in the 7’ loveridgit locality which 
have remarkably depressed carapaces (31 °/, of total length as 
against over 40°/,); and, further, Tornier describes one from 
Bussisi, the 7’. torniert locality, which has its bony plates 
excessively thin, and actually fenestrated. Possibly this indi- 
vidual, if let alone, might have founded a race of soft-shelled 
tortoises in this genus also. 

The actual process of development of the bony plates of both 
carapace and plastron in 7’. loveridgii is really quite normal ; in 
spite of the astounding appearance which they present in the 
adult, they are similar in general form to those of the young of 
other species. In some, of course, one set of elements may 
develop more quickly in proportion, altering the balance of the 
fenestre or their outlines, but the same process is gone through 
by them all. 

On dissecting a series of the young of Yestudo horsfieldit, I find 
that the very young (50 mm.) presents a bony network of 
costal and neural plates, modified ribs, and undeveloped plastron, 
similar to the young 7’. loveridgii (60 mm.). The two are 
compared in text-fig. 21. Series in other species, chosen at 
random, show the same thing, with, of course, variations. In 
some the dermal plates are more equalized, and in the early 
stages present a less net-like form, each costal being kite- 
shaped, rather as in the Turtles. In the Giant Tortoise 
(I. nigrita) from the Galapagos, a very young specimen (95 mm.) 
has the bony carapace still extensively fenestrated; the costal 
plates are more or less equally developed, each being dilated 
beneath the vertebro-costal sutures, and from thence narrowing 
to the marginal region, the fenestre between are therefore 
equal. The neural plates are broadly developed ; second, fourth, 
and sixth are not in contact with the corresponding costals. 
The general structure is therefore in essentials the same as in 
T. loveridgii. In Freshwater Tortoises,such as Hmys, development 
is proportionately quicker beneath the vertebral shields, a speci- 
men of 50 mm. having this part of the carapace entire, whilst 
the costal region is only half completed. 

In the following section the development of the éostal and neural 
plates will be dealt with further, in connection with their origin. 


Proc. Zoou. Soc.— 1922, No. OOO 36 


516 MISS JOAN B. PROCTER ON THE 


Origin of the costal and neural plates. 

Much has been written on the origin of these plates, and many 
ingenious theories have been propounded*. 

Most authors agree that the marginals, nuchal, and pygals are 
of dermal origin, and that the plastron is derived from the 
sternum and gastralia which have become greatly modified, but 
the question of the neurals and costals is still unsettled. 

The earliest opinion worth citing is that of Cuvier (1799), 
who considered that the costal plates were formed “ par les dilata- 
tions de huit cdtes ou batons osseux qui prennent naissance sur 
les unions des vertebres, et se terminent a un rebord qui entourent 
toute la carapace.” The idea that the plates are dilatations of the 
vertebral spinous processes and true ribs is held by many sub- 
sequent authors—Geoffroy St. Hilaire (1809), Bojanus (1819-21), 
Rathke (1848); or, further, that these plates originate in periosteal 
or differentiated tissue—Goette (1899), Haycraft (1899), Newman 
(1906). Haycraft (1891) formulates a theory that costal plates 
are rib-expansions formed where there is no differentiated 
periosteal membrane confining the rib. In the case of turtles, 
where the expansion does not reach the marginal, he states 
that the distal portions which preserve their rib-like form are 
unable to expand because “they are invested by a restraining 
periosteum.” The fact that the costal plates in most Chelonians 
develop gradually towards the marginals, passing through the 
stage normal to adult turtles, disproves this theory. 

Carus (1827) and Gegenbaur (1889) thought that the costal 
plates were greatly developed vertebral transverse processes—a 
view which, of course, is immediately disproved by the fact that 
their points of origin are distinct and widely separated from the 
vertebral column. 

O. P. Hay (1901) accepts none of the foregoing theories. He 
distinguishes three layers of bone—dermal, fascial, and cartila- 
ginous. The first is present in the armour of Dermochelys, the 
most primitive living Chelonian. . The modern costal and neural 
plates are formed not by dermal but by the fascial bone, 
which since it is completely united to the cartilage elements, 
appears to arise directly from the perichondrium, as observed by 
Goette (1899). 

That these plates are dermal in origin and similar in every way 
to the nuchal, pygal, and marginal plates was first suggested by 
Carus (1834), and was followed up by Peters (1838), Owen (1849), 
Baur (1887), Gadow (1899, 1905, 1909), and Versluys (1914). 
Many points brought to light in the present study corroborate 
this view. For instance, the fact that the true ribs are 
degenerating throughout their length by absorption within the 
periosteum whilst the costal plates are in process of actual 
development, and also the marked difference in the texture of 


* “ Hypothesis follows hypothesis; the theoretical rubbish-heap accumulates ; 
and truth ever eludes us.”—FABRE (transl.). 


REMARKABLE TORTOISE, TESTUDO LOVERIDGII. 


‘||| ||| @aqsouagz fsyt:t:2 qua Surdpwopum f-------. soesceseees SAINGNS-plalys [BULIpId yy 
‘pervdunoo “q, “2ap)ayS.toy «pur “e “abpisaao) «7, Jo susutioods Suno KX 


‘1% NSY-4xXoT, 


36% 


518 MISS JOAN B. PROCTER ON THE 


the two forms of bone, seem to indicate that these are inde- 
pendent structures. It will also be remembered that in strip- 
ping the carapace from the young specimen of 42 mm., some 
of the neural plates adhered to the carapace rather than to 
the vertebra. Another point against endo-skeletal origin is the 
alternating level at which the costal plates arise on consecutive 
ribs, according to the arrangement of the epidermal shields 
above them. 

This brings up the question, also unsettled, regarding the eorre- 
lation of epidermal shields to the underlying plates. If the plates 
are of dermal or fascial (sub-dermal) origin, one presupposes such 
a correlation; if of endo-skeletal origin, there should be none. 
Various authors argue the case accordingly, H. H. Newman 
opposing any idea of correlation very strongly, whilst Gadow 
treats it as an accepted fact. G. H. Parker (1901), in his paper 
on the Sculptured Tortoise, attempts to explain the alternating 
arrangement of these two structures by applying Harrison’s 
observations on the regenerating tail of a tadpole (1899). 
He says (p. 23):—‘“* The tail of a developing tadpole is com- 
posed of an outer covering of ectoderm—which ultimately 
gives rise to the outer layers of the skin—and of a core of 
mesoderm. These two masses of tissue grow in very different 
ways, so that as the tail lengthens, the ectodermic covering, 
which is most actively produced anteriorly, slips posteriorly 
over the underlying mesoderm, whose region of growth is chiefly 
at the posterior end. Although this posterior migration of the 
ectoderm has been actually demonstrated only in the tadpole, 
there is reason to believe that it occurs in other vertebrates.” 

The present study points to a very marked and obvious correla- 
tion between shields and plates in TVestudo loveridgii, which, 
having its carapace in what one might term an incipient stage of 
development, throws much light on the subject. A careful study 
of many species of Testudo and Cinixys during the period when the 
plates are developing, brings forth a wealth of evidence that this 
correlation is an indisputable fact in these genera. In newly- 
hatched individuals of 7’. loveridgit (and also those of 7’. ibera, 
greca, horsfieldii, leithii, ete., etc.) the bony plates are lacking 
beneath the areolars, developing apparently as the shields them- 
selves commence to form their growth-rings. In specimens of 
different species the amount of plate which has accrued at the end 
of the season’s growth varies, but is at first 1 all cases greater 
beneath shield-sutures (that is around the shield-edges) than else- 
where. ‘The neural plates beneath a shield-suture are developed 
laterally, whilst the alternating ones are not. The costal plates 
beneath the costal shield-sutures are at least twice as broadly 
developed as those between; their proximal portions, however, 
alternate in breadth and narrowness inversely in strict accordance 
with the overlying shield-sutures. Moreover, in the male type- 
specimen the X-ray plate (vide Pl. III.) shows that the bony 
plates are in perfect harmony with the abnormal third vertebral 


REMARKABLE TORTOISE, TESTUDO LOVERIDGII. 519 


shield-sutures, and are thus adapted to support four instead of five 
vertebrals. This fact points to dermal origin of the plates, but is 
in opposition to Parker’s theory of ectodermic migration in 
Chelonia (@ propos of the tadpole’s tail), in which he points out 
that ‘‘any early local disturbing influence that affected both scute- 
and plate-producing tissues would leave its trace in the adult in 
the form of a region of modified scutes posterior to a region of 
modified bony plates.” At the same time, this does not alter the 
fact that the plates are adherent to the bones of the true skeleton, 
and are coincident with them regarding numbers etc. Yet 
development seems to be influenced and regulated chiefly by that 
of the overlying dermal areas corresponding to the epidermal 
shields. This is also seen in the Giant Tortoises known to 
have thin carapaces, such as 7’. abingdonii, microphyes, etc., in 
T. calcaratus, and in fossil forms such as the Pleistocene Marsh 
Tortoises, 7’. vosmaert from Rodriguez and J. indica from 
Mauritius. In these the form of the plates is similar to that of 
T. loveridgui, the essential network beneath the shield-sutures being 
of thick bone standing out im relief from the thin bone, which fills 
in what would be fenestra in the young or in loveridgii. 

In ordinary adult tortoises in which the carapace is complete 
and of equal thickness, this cannot be appreciated, but in these 
it can be studied in the initial stages of plate-development. 

The following quotation from Owen (1849, p. 161), whose paper 
T had not read “at the time of my own Geer vations, corroborates 
in every particular what I have found to be the case in so many 
species :— 

«A strong argument for regarding the costal plates as dermal 
ossifications rather than processes or continuations of the endo- 
skeletal elements, to which they are attached, may be drawn... . 
from the period of their ossification, and their relative position 
to the ribs with which they are connate. 

‘In the embryo Zestudo indica* the uniformly slender pleura- 
“pophyses are ossified nearly throughout their whole length 
before the ossification of the costal plates, usually regarded 
as their expanded tubercle, commences ; and the beginning of the 
superadded bone 7 is not at the same point in each rib, as might 
have been expected if it were the exogenous process called 
‘tubercle’ of the rib. The costal plates are situated in the 
young Testudo indica alternately nearer to and farther from 
the head of the rib; and their presence seems to be deter- 
mined rather by the angle of union of the superincumbent 
vertebral scutella with the lateral or costal scutella, than by 
the necessity for additional strength of the articulation of 
the ribs with the spine. Ossification commences at the point 
from which the three impressions radiate, and as this point is 


* [Probably 7. nigrita or elephantina, indica Schn. being an extinct species. ] 

+ “ This period, in its relation to the development of the neural arches and pleura- 
pophyses, corresponds precisely with that at which the dermal plates of the Crocodile 
begin to be ossified.” 


520 MISS JOAN B. PROCTER ON THE 


nearer the median line at the median apex of the costal scutellum 
than at the lateral apex of the vertebral scutellum, the resulting 
plates of bone are alternately further from or nearer to the middle 
line; and the first, third, and fifth costal plates have advanced 
along the proximal end of the rib so as to join the neural plates, 
whilst the second, fourth, and sixth costal plates leave a 
portion of the proximal end of the rib uncovered and crossing 
the space between the incipient costal plates and the neural 
plaueyeys et. 

‘‘When the partially ossified carapace of a young tortoise is 
dried, one cannot fail to be struck with the difference in the 
texture and external surface of the bones which unquestionably 
belong to the endo-skeletal vertebree, and of those which, notwith- 
standing their connection with neural spines and pleurapophyses, 
are developed in the fibrous substance of the corium. These 
nascent ‘ neural’ and ‘costal plates’ of the carapace havea granu- 
lar exterior and a coarse spongy texture, whilst the neural arches 
and pleurapophyses are compact, smooth, and with a polished 
external surface: the part of the pleurapophyses which passes 
beneath and is attached to the under surface of the ‘ costal’ plate 
(pls. i—viil.), contrasts strikingly with that superimposed dermal 
ossification. 

“The marginal plates present the same rough, coarse, 
granular character as the neural and costal plates..... their 
ossification has been governed by the presence of the marginal 
epidermal scutes, and, as in the case of the costal plates, 
by the points of junction of contiguous scutes; each marginal 
ossification is accordingly impressed by the lines indicating the 
junction of the marginal epidermal scutes with each other and, in 
the case of the middle ones, with the contiguous scutes of the 
plastron.” 

The whole of the foregoing evidence therefore points to the 
fact that the costal and neural plates are of dermal origin, not 
endo-skeletal; and are, in fact, similar in this respect to the 
nuchal, pygal, and marginal plates. 

Further, the corrrelation between the epidermal shield-sutures 
and the plates beneath, observed independently in the present 
study and previously by Owen, suggests that their growth is also 
correlated. 

The areas of dermis beneath each shield are well marked off 
from one another by a fold and an underlying groove in which 
the shield-edges are implanted. It is probable that these 
dermal areas grow in the same manner as their superimposed 
shields, and that the bony plates beneath, which are of dermal 
origin, have their growth similarly regulated. 


The following hypothesis is therefore suggested :-—— 


That the bony plates of the carapace develop from the dermis, 
primarily in the areas of growth activity. 


REMARKABLE TORTOISE, TESTUDO LOVERIDGII. 52] 


(a) Where this takes place concentricaily, as seen by such 
striations in the epidermal shields, the plates in the early stages 
of development are localised principally beneath the shield- 
sutures, as in Zestudo, Cinixys, etc. 


(6) Where growth is uniform throughout each area, develop- 
ment of the plates is also equalised, proceeding from the individual 
centres of ossification, as in Hmys etc. 


SUMMARY. 


Testudo loveridgii has an excessively depressed soft-shelled 
carapace, and is able to inflate itself to a certain degree. It 
possesses a bony carapace and plastron, but they are extensively 
fenestrated, incomplete, and similar in essentials to the juvenile 
stages of other species. 

Marginal plates five, six, and seven spread inwards in a unique 
manner, entering into the composition of the plastron and 
separating the hyo- and hypoplastrons from each other. Their 
upper portions are extraordinarily shallow. 

The ribs in 7’. loveridgit become (usually) entirely absorbed, 
apparently by the osteoclasts which are present beneath the 
periosteum. Asimulacrum of the capitular portion, soft like 
ligament, and formed chiefly of periosteum, persists for some 
while. 

The neural arch is vestigial and sometimes completely wanting, 
the neural plates being applied to the depressed centra to form 
the roof of the neural canal. Absorption probably takes place to 
some extent as in the ribs, but the arch is never more than a 
simple layer of bone, without spinous processes. 

The jaws, together with their investing horny sheaths, are 
denticulated with remarkable regularity. 

Fenestration in this and other species 1s caused by arrested 
development, and not, as previously supposed, by absorption with 
age. 

The development of the bony plates in T. loveridgw and the 
young of other species points to the newrals and costals being of 
dermal origin. The principal evidence is :— 

(1) That they are developing whilst the ribs and neural arches of 
the true skeleton are degenerating. (2) That the point of origin of 
each costal plate is alternately nearer to and further from the rib 
capitulum. (3) That the form of these plates in their early stages of 
development is in strict correlation with the borders of, or sutures 
between, the superineumbent epidermal shields. 

It seems possible that the development of the plates is regulated 
by that of the areas of dermis corresponding to the epidermal shields ; 
(1) forming a network as in T. loveridgii and other species when 
this growth proceeds concentrically, or (2) proceeding equally from 
each centre of origin where dermal growth takes place equally within 
each area as in Kmys etc. 


522 MISS JOAN B. PROCTER ON THE 


NOTES. 


The following field-notes and observations on the original 
specimens, and those collected on subsequent expeditions, are 
contributed by Mr. Loveridge :— 


“ Habitat. 


‘““The first specimen of this tortoise which I met with was 
found dead at the foot of a precipitous rock some 40 feet in 
height and situated near the top of a rocky kopje 500 feet 
above the surrounding dry thorn-bush country. From the 
flattened and broken remains, I concluded that it was a species 
unknown to me, and crushed by a rock having fallen upon it. 
(The rocks are grey granite, sheer precipiees on the kopjes and 
rounded boulders scattered on the plains around. Most of the 
specimens have been collected beneath the latter.) On December 
8th, 1918, my native collector brought in a small specimen which 
I at once assumed was T. tornieri Sieb., of which I had read brief 
notices but had never seen the original description. During 
succeeding days two batches of these tortoises were found in 
crevices or beneath rocks, but though I did not leave the district 
till December 28th, and had a native looking for them constantly, 
no more were found. 

“On January Ist, 1921, I sent the same boy who was with me 
for two years before, back to the same locality which lies south of 
Dodoma inarid country. Here, again, he found but two or three 
specimens, but, ranging round, came upon another kopje where 
they were more plentiful, though he alleged that without a single 
exception they were under the rocks, of which he had to remove 
a great many before being able to effect captures. He had one 
piece of good fortune in finding four young specimensall together 
beneath one boulder, with the exception of a slightly larger one 
already caught; these were the only young specimens taken. The 
smallest of these was unfortunately trodden upon and promptly 
died. The shell is much depressed as in the adult. When 
Mr. Boulenger described the species in 1920, he assumed a small 
dome-shaped tortoise, which I found preserved in a bottle (no 
data) in a German house near Morogoro, to be the young of 
1’. loveridgii; this, however, is disproved by the finding of young 
specimens with depressed shells. 

“In November 1921, I spent an afternoon with my boy looking 
for 7’. loverrdgu at Dodoma. He first took me to the place 
where he got the eleven tortoises, which is not on a kopje but on 
a huge rock 100 yards from a kopje. The rock is about 30 feet 
long and 10 feet wide, and slopes up from the ground to a height 
of 7 or 8 feet. On the knob is a flattish boulder, and beneath 
this he found them allin January (evidently estivating—January 
and February being our hottest months). The tortoises therefore 
climbed up the boulder : one would certainly never think of looking 
for them in such a spot. We then went to the place where he 


REMARKABLE TORTOISE, TESTUDO LOVERIDGII. 523 


found the three—in a cleft of a split boulder ; they had to climb 
the boulder and slide down an almost vertical cleft, where one 
would suppose they could not get out; the cleft was about 
18 inehes. In this cleft, amongst a litter of dead leaves, were two 
not half-grown individuals. We hunted about all the rest of 
the day, but found no more.... Their markings are very 
fine; Tabora specimens are noticeably darker than the Dodoma 
ones. 


“ Paring. 

“On December 12th, 1921, at 4.30 p.m., I observed two 
specimens paired for the first time. A native said they had been 
so approximately for half-au-hour. The shell of the male formed 
an angle of 45° with that of the female; his fore-feet were in the 
air; he kept moving his head in and out of his shell and had his 
mouth wide open. In April this female and others were heavy 
with eges.. On January 18th two more tortoises were paired at 
9 or 10 A.M. and from that date and onwards pairing took place 
daily, usually between 9 or 10 am. and 2 p.m. The female 
generally walked away with the male following; frequently the 
latter became impatient and snapped at her limbs, or crawled on 
her back, making vicious snaps at her head whenever it appeared. 
On a couple of occasions the males were so ill-tempered that they 
seized the edges of the females’ shells in their jaws and dragged 
them along, finally getting beneath them and apparently 
endeavouring to overturn them. 


“Habits of T. loveridgii compared with those of Cinixys belliana. 


“ As I had these species under observation for four months 
confined in the same enclosure, J found it very interesting to 
compare the habits of an exceptionally highly-specialised Box 
Tortoise with those of a retrogressive and soft-shelled species. 

“Owing to neither my native collector nor myself finding 
T. loveridgiti in the open, I had come to the conclusion that 
it was nocturnal; with a view definitely to ascertain if this were 
indeed so, I visited the enclosure at all hours of the night up to 
1 a.m., but with one or two solitary exceptions never found them 
out. At dusk they always retired beneath the heaps of stones and 
boulders provided for them, and there they piled themselves up, 
one atop of another, a great deal of noise being caused by the 
scratching of claws on shells. They did not again make an 
appearance until several hours after daybreak, which varied 
according to the dullness or brightness of the morning. 

“C, belliana, on the other hand, not infrequently stayed out at 
night, and particularly when rainy ; all seven of them would spend 
the night in trying to escape from the enclosure. Moreover, 
CO. belliana was always on the move at daybreak. On further 
reflection it was obvious that in their soft shells 7’. loveridgit 
would fall an easy prey to the carnivores roaming about after 
nightfall, and which are very plentiful in the Dodoma district 


524 MISS JOAN B. PROCTER ON THE 


(Leopard, Ratel, Civet, Mungoose). This was brought home to 
me very forcibly one night by a Civet entering an adjoining 
enclosure and eating four and a half young crocodiles in one night. 
The shell of C. belliana would resist most attacks; the thickness 
of their bony defence as seen in a sawn-through section is 
astonishing. 

“* 7’. loveridgit shows a fondness for climbing up and falling off 
the rockeries of its enclosure, which pastime, indulged in in its 
natural habitat, calls for a rapid recovery in turning over ; it also 
displays great agility in climbing up wire netting. One day 
eight specimens of the Soft Tortoise succeeded in escaping through 
a hole in the wire netting ; two were recovered in three days, 
being found on the second and third days at some huts 400 yards 
from the hill. They appear to feed oftener than C. belliana and 
are quite voracious. They feed well ona succulent grass here; the 
previous specimens taken home were fed on lettuce, and when 
this failed were induced to eat bread soaked in jam. 7’. loveridgu 
generally retires to its retreat when a shower comes on, but on 
oceasion I have known the whole lot turn out during heavy rain 
and feed voraciously, probably being thirsty. 

““T was at first disposed to think that 7. loveridgit could not 
swim, as the first half-dozen arrivals on being placed in water sank 
to the bottom of the bath and remained there without putting 
forth any effort. On seeing some young ones struggling in water 
and trying to swim, I retried a couple of females, which struck 
out well and kept themselves up when placed in a bowl of 
water.” 


Unfortunately nearly half of these valuable tortoises, kept alive 
by Mr. Loveridge, succeeded in escaping, several doing so en route 
to the coast on their way to me. Several spirit specimens also 
came to grief, but a sufficient number remain, reinforced by still 
later collections, including a beautiful series of young, to enable 
a thorough study of this most variable species to be made. 


BIBLIOGRAPHY. 


Agassiz, L. 1857. Contributions to the Natural History of the United States. 
Vol. i. Part ii. North American Testudinata; vol. 1. Embryology 
of the Turtle. Boston. (27 pls.) 

Baur, G. 1890. On the Classification of the Testudinata. Amer. Nat. vol. xxiv. 
pp. 530-536. 

1887. On the Morphogeny of the Carapace in Testudinata. Amer. Nat. 

vol. xxi. p. 89. 

J. F. van. 1896. Bemerkungen zur Phylogenie der Schildkréten. 

C. R. 3. Congrés Internat. de Zool. Leyde, pp. 322-335. 

Bosanus, L. H. 1819-21. Anatome Testudinis Europxe. Vilne, pp. 1-178 
(31 pls.). 

Boutencer,G.A. 1889. Catalogue of Chelonians, Rhynchocephalians, and Croco- 
dilians in the British Museum. 

1920. Une Tortue extraordinaire: Testudo loveridgii, sp. n. C. R. 
Acad. Sci., Paris, t. 170, pp. 263-265. 

Carus, C.G. 1827. Comparative Anatomy. Translated by R. T. Gore. Vol. i. 

p. 147. 


BEMMELEN, 


REMARKABLE TORTOISE, TESTUDO LOVERIDGII. 525 


Carus, C.G. 1834. Lehrbuch der vergleich. Zootomie. Bd. i. pp. 164-5 (2nd 
Edition). 
Cuaus, C.F. W. 1884-85. Text-book of Zoology, London, p. 226. (Transl. 
Sedewick and Heathcote.) 
Cooxer, R. E. 1905. “Gadow’s Hypothesis of Orthogenetic Variation in Chelonia.” 
Johns Hopkins Univ. Circular, no. 178. 
1910. Diversity in theScutes of Chelonia. Journ. Morph., Philadelphia, 
vol. xxi. pp. 1-75, 14 pls. (Original abstract in Science, n. s. vol. xxi. 
no. 632, 1905, p. 384:) : 
Cuvier, M. 1799. Lecons d’Anatomie Comparée. T. 1. p. 211. 
1809. Ossemens Fossiles. Annales du Muséum, Paris, t. xiv. p. 7. 
Gapow, H. 1899. Orthogenetic Variation in the Shells of Chelonia. Zool. 
Results, Willey, pp. 207-222 (pls. i—111.). 
1905. Orthogenetic Variations. Science, vol. xxii. pp. 637-640. 
1909. Cambridge Natural History, vol. viii. pp. 312-329. 
GrorFrroy St. Hrnarre. 1809. Ann. Mus. vol. xiv. p. 7. 
GreEenpaur, C. 1898. Vergleichende Anatomie der Wirbelthiere. Leipzig. 
Ba. i. 
Gortrx, A. 1899. Uber die Entwicklung des knocherner Riickenschildes (Carapax) 
der Schildkréten. Zeitschr. Wiss. Zool., Bd. lxvi. pp. 407-434 (pls.). 
Harrison, R.G. 1898. The Growth and Regeneration of the Tail of the Frog 
Larva. Arch. f. Entwickelungesmechanik d. Organismen, Bd. vii. 
pp. 431-485. 
Hay, O. P. 1892. The Batrachians and Reptiles of the State of Indiana. Indiana- 
polis. 
1898. On Protostega, the Systematic position of Dermochelys, and 
the Morphogeny of the Chelonian Carapace and Plastron. Amer. Nat. 
vol. xxx1i. Boston, pp. 929-950. 
———— 1901. The Composition of the Shell of Turtles. Sec. Biol. Acad. Sci. 
New York. 
1922. On the Phylogeny of the Shell of the Testudinata and the 
Relationships of Dermochelys. Journ. Morph., Philadelphia, vol. 36, 
pp. 421-441, pls. i. & ii. 
Haycrort, J. B. 1891. The Development of the Carapace of the Chelonia. 
Trans. Roy. Soc. Edinb. vol. xxxvi. pt. ii. pp. 835-342 (pl.). 
Horrmann, C.K. 1890. Bronn’s Klassen und Ordnungen des Tierreichs, Bd. vi. 
Abt. 3. 
Huxuey, T.H. 1871. The Anatomy of Vertebrate Animals. London, p. 201. 
Loénnzere, E. 1911-12. Reptiles, Batrachians, and Fishes collected by the 
Swedish Zool. Exped. to British East Africa 1911. Svensk. Vetensk. 
Akad. Handl., Upsala and Stockholm, t. 47, no. 6, p. 7. 
MecKEt, J. ee 1821-33. System der Vergleichenden Anatomie. Halle, Bd. ii. 
Abt. 1. 


Newman, H.H. 1905. The Significance of Scute and Plate ‘“‘ Abnormalities ”’ in 
Chelonia. (A cont. to the Evolutionary History of the Chelonian 
Carapace and Plastron.) Biol. Bull. Marine Biol. Lab., Woods Holl, 
Mass. vol. x. pp. 68-114. 


Owen, R. 1849. On the Development and Homologies of the Carapace and the 
Plastron of the Chelonian Reptiles. Phil. Trans. Roy. Soc. London; 
vol. exxxix. pp. 151-171 (pls.). 

Parker, G.H. 1901. Correlated Abnormalities in the Scutes and Bony Plates of 
the Carapace of the Sculptured Tortoises. Amer. Nat. vol. xxxv. 
pp. 17-24. 

Peters, W. % ae 1838. Observationes ad Anatomiam Cheloniarum. Berolini 

pls.). 

RatuKe, H. 1846. Sur le dévelopement des Chéloniens. Ann. Sci. Nat. Paris, 
Zoologie, sér. 3, t. 5, pp. 161-170. 

1848. Ueber die Entwickelung der Schildkréten. Braunschweig, 
pp. 1-268 (10 pls.) (wide pl. i11.). 


526 ON THE REMARKABLE TORTOISE, TESTUDO LOVERIDGII. 


SrEBENROCK, F. 1908. Testudo tornieri, nov. sp. Ak. Wiss. Wien, Math.-nat. 
Klasse, Bd. xxiv. (1v.) pp. 185-186. 


1903. Uber zwei seltene und eine neue Schildkréte des Berliner 
Museums. Sitz. Ber. Ak. Wiss. Wien, Math.-nat. Klasse, Bd. exi. 
p. 443 (vide pl. of T. tornieri). 


—--— 1909. Synopsis der rezenten Schildkréten, mit Berucksichtigung der 
in historischer Zeit ausgestorbenen Arten. Zool. Jahrb., Jena, 
Suppl. 10, h. 3, p. 538. 


TornieR,G. 1895-96. Deutsche Ost-Afrika, Bd. iti (Reptilien und Amphibien) 
Chelonia p. 2. 


Verstuys, J. 1914. Uber die Phylogenie des Panzers der Schildkréten und Uber 
die Verwandtschaft der Lederschildkréte (Dermochelys coriacea). 
Paleont. Zeitschr. Berlin, Bd. i. pp. 8321-347. 


Vouxer, H. 1913. Uber das Stamm- Gliedmassen- und Hautskelet von 
Dermochelys coriacea L. Zool. Jahrb. Bd. xxxiui. pp. 431-552, pls. 30- 
33. 


Waener, R. 1841. Icones Zootomice. Leipzig (80 pls.). 


EXPLANATION OF THEH PLATES. 
Pratt I. Vestudo loveridgii Blgr. (} nat. size). 


Pratell. Radiograph of 9 type-specimen (nat. size). Showing correlation between 
bony plates and epidermal shield-sutures (visible as a network of 
white lines). 


Puate III. Radiograph of ¢ type-specimen (nat. size). Showing correlated shield 
and plate abnormalities. 


~I 


ANATOMY OF CACOPUS SYSTOMA, 52 


27. Notes on the Anatomy of Cacopus systoma, an Indian 
Toad of the Family Engystomatide. By D. W. 
DEVANESEN, M.A.* 


{Received April 4, 1922: Read May 9, 1922. | 


(Text-figures 1-19.) 


CONTENTS. 

Page 
leplmtroductiowm) pst frank alee ee ie Ae bly ee eek oath hie BOT, 
2. The Viscera.. aa a Geer en eae, Van eS ROS 
3. Sub- entanbous mil Bite Fat. Thaities, a yyceene sbone, G0) 
4, Discussion on “the stout shape” of Cheap. corinne . 53l 
Seth epAialls keletom’ vce sccasece teeta cee Cee ee Be 
6. The Appendicular Skeleton AAR Sa0 cad eho LROCC TACO TOT A OS amELe 
(eeebieRMiursculature:. Aan tc teste an eae k ust ui ates canny Ach 
8. Summary........ 553 
9. Key-list of wssona ehirovierions ot lemained! clone muh 

AN So (PS TEA TDRES oh So ee senae ade ice sau Sao uee eonacsee Bait Noe em eta fie Es 15159 
10. References to Tikewnm, Wheod tenes fe retaanat (en ah Ao. 40 OOO) 


1. Introduction. 


Cacopus systoma is an Indian toad of the narrow-mouthed, 
edentulous and fossorial type belonging to the family Engy- 
stomatide, well-known for their habit of feeding upon ants and 
termites. Among the chief characters which distinguish the 
genus are the following: the palate has two dermal transverse 
folds more or less denticulated, one behind the vomers and the 
other in front of the cesophagus ; there are no pre-coracoids ; the 
sternum is a large cartilaginous plate; and the transverse pro- 
cesses of the sacral vertebra are rather strongly dilated. 

The following notes were compiled in India during what was 
intended to be a comprehensive study of the morphology of 
Cacopus systoma. Comparisons with the anatomy of certain 
other toads of the same family and comments thereon have been 
made wherever suitable. 

This work was done in the Zoological Laboratory of the Madras 
Christian College, Madras. Grateful acknowledgment is hereby 
made for suggestions and advice to Prof. W. Rae Sheriffs of the 
college mentioned, and to Dr. J. R. Henderson, F.L.S., formerly 
Superintendent of the Government Museum, Madras. To Prof. 
MacBride’s encouragement the communication of this paper is 
due; for this and for reading the manuscript, I thank him. 


* Communicated by Prof. E. W. MacBrips, D.Sc., LL.D., F.R.S., F.Z.S. 


\ 
528 MR. D. W. DEVANESEN ON THE 


2. The Viscera. 


The second dermal ridge on the palate of this toad is a con- 
spicuous structure stretching across from one eustachian opening 
to the other, its edge being thrown into well-marked denticula- 
tions. On laying open the pharynx*, one finds immediately 
behind this ridge and in close contact with it and occupying 
more or less the same level, a differentiated area of about 
80 sq. mm. (text-fig. 1). Transverse sections through the same 
revealed the fact that it consisted of closely-set, parallel, and 
longitudinal reduplications of the mucous membrane of the 
pharynx, each fold containing an extension of the basement mem- 
brane (text-fig. 2). A fold is on an average 1 mm. in height; 
the highest folds are found in the middle, and from this place 
the height of the folds on the sides gradually decreases. These 


Text-figure 1. 


’ 


“The Pharyngeal Organ of Cacopus systoma” exposed; the lower jaw and the 
floor of the mouth Ihave been cut away. L., lamelle arranged lengthwise ; 
R., the first, short, R.’, the second, long, denticulated ridge; H., eustachian 
opening; V., vomer; I.P., inner dark papilla on the toothless hind end of a 
vomer (V.); O.P., an outer one on the vestigial palatine (P.); P.H., pharynx 
cut open and deflected. 


folds show a faint tendency to anastomose, and their free edges 
hanging down into the cavity of the pharynx, in the fresh 
condition, appear swollen. This specialised area of the pharynx 
may, in my opinion, be given the status of an organ for two 
reasons: first, it 1s compact, having a certain definite outline, 
and it is restricted to a definite place, namely immediately 
behind the second dermal ridge; secondly, as the free end of the 
tongue lies, when in a state of rest, underneath this area, and 
since the presence of the intermaxillary glands is doubtful in 
this toad, the structure in question is probably a substitute for 
these glands, possessing a similar function; the cells of the mucous 
membrane in these folds are comparatively large and sacciform, 
which facts are perhaps suggestive of their glandular nature. 


* That part of the alimentary canal between the second dermal ridge and the 
stomach, I propose to call “‘ the pharynx,” as 1t is much wider than it should be for an 
cesophagus which is probably unrepresented in this toad. he cavity enclosed 
between this ridge and the jaws may be termed “the oral cavity.” 


ANATOMY OF CACOPUS SYSTOMA. 529 


I therefore propose to call this the ‘pharyngeal organ” of Cacopus 
systoma, It will be interesting to know if this structure is 
present in the other members of the family Engystomatide. 

On the outer surface of the pharynx, from each of the two 
points roughly corresponding to the two ends of the second dermal 
ridge, is given off a slip of tissue which is attached to the free end 
of that arm of the pterygoid meeting the cranium. This arrange- 
ment is presumably for the purpose of holding up the pharyngeal 
wall which, in this region, is particularly heavy with the dermal 
ridges and the “ pharyngeal organ ” described. 

The longitudinal folds of the mucous membrane of the stomach 
appear to be more numerous than in Rana, and in the convex 
region thereof they show a tendency to anastomose. The liver 


Text-figure 2. 


Sea 
Ss 
ee 


Wey FARIS NEIRES, 
ae Wg ipetale AS US ay 
be BR DoS gy GY 
7 “gp Sms Wetintats 
Es Sak we = 
Faces ee L. 


xe D. 

A transverse section through the middle region of “the pharyngeal organ.” 
L.M., outer lengthwise muscles of the pharynx; T.M., inner transverse ones 
of the same; 8.T., sub-connective tissue; B., basement membrane; L., lamella; 
E., ordinary stratified epithelial cells; E.’, the same epithelial cells enlarged 
and forming a sac; X., cell-mass resulting from the fusion of two adjacent 
lamellze; D., an incipient duct. 


consists of but two lobes. Among the numerous specimens I 
examined, I found eight—six females and two males—suffer- 
ing from stone in the bladder. The proximal part of each 
oviduct where it opens into the cloaca becomes enormously 
dilated into a sac, which may be distinguished as a uterus. In 
this bag probably the eggs are stored before they are finally 
passed out. It consists of a thick-walled, less capacious anterior 
portion and a membranous spacious posterior part. The internal 
lining of the former is thrown into numerous, densely-packed, 
villi-like processes, visible to the naked eye. The two bags or 
uteri are, along their inner sides, intimately connected by tissue ; 
I do not think that there is an actual fusion between their walls. 
The kidney possesses a simple lobe towards the hind inner end. 


530 MR. D. W. DEVANESEN ON THE 


3. The Sub-cutaneous and other Fat-bodies*. 


The occurrence of sub-dermal fat is by no means peculiar to 
this toad ; several species among the Anura are known to possess 
it, though its function in most of them still remains to be eluci- 
dated (Bouienger, C. L.,3). Accumulations of the fat in question 
are found in C. systoma chiefly in two situations: (1) in the 
neck-region above and below, in connection with the lymphatic 
maxillary septum; this lobular mass of fat may be termed “the 
cervical fat-body (text-figs. 3 and 13); and (2) in the inguinal 
region on the dorsal surface in relation with the lymphatic 
inguinal septum, which is very high and falls into folds; in the 


Text-figure 3. 


— 
" WN) 


= 
[\ 
EX. 
pp 
i 


Stk oe 


The skin on the back cut open and the sub-cutaneous fat-bodies exposed in a male 
C.F., cervical fat-body; S.F., supra-inguinal fat-body; F, fatty deposits on 
the head and neck; I.I., conspicuous inscriptiones tendine of the obliquus 
muscles, seen only among males. 


same region, but to a less extent, fat is developed in connection 
with the iliac sac and the posterior lymph-heart ; this extensive 
area of fat, in its totality, may be called “ the supra-inguinal fat- 
body” (text-fig. 3). Besides these two conspicuous sub-dermal 
fat-bodies, one oceasionally finds scanty deposits of the same 
substance under the skin in the parietal and occipital regions, 
in a small median area dorsally behind the head and on the 
under side of each arm near the brachial vein. I assume that 
these, too, are developed in relation to the lymphatic system. 
One also finds in this toad certain internal deposits of fat 


* Young specimens show neither an adequate development of these fat-bodies nor 
are they present in all the places mentioned. 


ANATOMY OF CACOPUS SYSTOMA. 531 


which presumably are sub-peritoneal fat. These are found in 
relation to the blood-vascular system and the vertebral column. 
The auricles and the truncus arteriosus are thickly covered with 
fat, and the ventro-auricular groove is filled with the same; this 
cardiac fat, it should be observed, is found on the inner layer 
of the pericardium. Small quantities of fat occur also on the 
walls of the inferior vena cava, where it opens into the sinus 
venosus. In close proximity to the vertebral column is situated 
a longitudinal patch of fat on both sides extending from the 
fourth vertebra to the ninth, and a small median deposit where 
the cranium articulates with the atlas. 

Among the other Indian frogs I examined, sub-dermal as well 
as sub-peritoneal fat in relation with the blood-vascular system 
was found in Rana hevadactyla, the analogous form in India to 
Rana esculenta, and in Rana breviceps, a burrowing species. Never- 
theless, in these instances, while fat under the skin is confined to 
the neck-region, even that does not occur so regularly. The 
same fat in the inguinal region has been noticed by other observers 
in several species of Bufo (3), Xenopus levis, and notably in 
Hemisus guttatum, an American Engystomatid (1); in the first- 
mentioned toad, however, it is ventrally situated, while in the 
last one there is a complicated development of the same in con- 
nection with the iliac sac which does not obtain in C. systoma. 

As this. Indian toad has a remarkable habit of burrowing 
underground when the dry-season commenced and of passing 
into a torpid state, it struck me that these fat-bodies might have 
a nutritive value to the animal during that estivating period. 
Therefore, in July and August, 1911 and 1912, I obtained by 
special efforts a considerable number and examined them. It 
was found that both the sub-dermal and sub-peritoneal fat-bodies 
had undergone, in accordance with my expectations, a great 
reduction in sizeand a change in colour from yellow to dull white, 
as compared with specimens examined during the wet-season. 

From the above considerations one may draw the following 
conclusions :—Fat occurs normally in several places in the body 
of C. systoma, both internally and under the skin. While the 
pre-renal fat-bodies may be looked upon as feeders to the gonads 
during the breeding season, the other fat-bodies in this toad may 
be regarded as assisting in the general nutrition of the body 
during estivation. It is noteworthy that in Bufo, Hemisus, and 
this toad the sub-dermal fat is developed in connection with the 
lymphatic system. 


4, Discussion on ‘the stout shape” * of C. systoma. 


A remarkable feature of the two toads of this genus is, as several 
observers have noted, the rotund shape of the body. This has 
been, till now, accounted for in various ways. Thus, Giinther (6) 


* Vide figure in Boulenger, G. A. (2), p. 496. 
Proc. Zoou. Soc.—1922, No. XX XVII. 37 


532 MR. D. W. DEVANESEN ON THE 


observed regarding C. globulosus that it was due to a certain 
fluid contained in the abdominal cavity and the extraordinary 
development of the ovaries in the female. But Pearson (7) 
remarks as follows:—‘ Gunther states that this distension is 
caused by a fluid contained in the abdominal cavity. <A brief 
examination shows that this is not thecase, and that the fluid- 
containing cavity is none other than the sub-cutaneous lymph- 
sinuses which are greatly enlarged in; this form. The dorsal 
sinus is especially spacious, and has a height of 15 mm. from 
floor to roof. (The length of the frog from mouth to vent is 
75 mm.) 

“In Giinther’s account he speaks of the distension of a female 
specimen being due to the growth of the ovaries, and his descrip- 
tion suggests that the ovaries grow into the large cavity on the 
back. This is not possible, as the sub-cutaneous lymph-sinuses 
are separate from the celom. There is only one other species in 
this genus, namely Cacopus systoma, which differs but slightly 
from C. globulosws, and which resembles it in the robust appear- 
ance due to the enlargement of the sub-cutaneous lymph-sinuses. 
This interesting character is not given in the diagnosis of the 
genus either by Giinther or Boulenger.” 

Thurston (8), with reference to the same species, says :—‘‘ On 
laying open the visceral cavity, the globular shape was found to 
be due to an enormous distension of the esophagus and stomach, 
the latter occupying nearly the whole of the abdominal cavity, 
and the remaining viscera &c. being compressed and lying 
posteriorly. There was no distension of the intestinal tract. The 
distension of the cesophagus and stomach was found on section 
to be caused by the presence in their cavities of a mass of winged 
white ants (termites) which when dried weighed 326 grains.” 

The descriptions of Giinther are based on slender evidence, 
namely the examination of two specimens in spirit. So far as 
I was able to observe in several fresh specimens of C’. systoma, 
I did not find an excessive quantity of the peritoneal fluid beyond 
what should generally be expected in the abdominal cavity. The 
supposition that the development of the ovaries is a source of 
the stoutness may at once be dismissed, as it can apply only 
to females, and again in them it is seasonal as in all batrachians. 
So also Thurston’s hypothesis, as food cannot be regarded as a 
permanent cause. 

I find that the stout shape of the body in this toad cannot be 
accounted for by any one character, but by a combination of 
features as the following :—The viscera as a whole exhibit a 
great development which is disproportionate to the small dimen- 
sions of the abdominal cavity. They therefore press against 
the sides of the abdomen and bring about a characteristic 
rotundity of the same. A feature which has not been noted as 
regards this toad is the inclusion of the proximal portion of the 
thigh into the trunk. The skin of the thigh is drawn along with 
that of the body, and consequently the contour of the latter 


ANATOMY OF CACOPUS SYSTOMA. 533 


begins posteriorly about the middle of the thighs*. This, I 
should think, improves upon the effect produced by the viscera. 
The enlarged lymph-sinuses, with the lymph and fat-bodies con- 
tained therein, enhance the stout shape. 

The purpose of the enlarged lymph-spaces of the two toads of 
this genus is perhaps to be sought in their fossorial habits. The 
liberally secreted lymph may serve to protect the internal organs 
from injuries incidental to animals with such habits. The muscles 
of the abdomen in C. systoma, chiefly the rectus abdominis, form 
a weak protection, as they are stretched to their utmost capacity 
by the pressure of the enormously-developed viscera. The 
presence of a fluid between the latter and the skin must, under 
such circumstances, serve to diminish the chances of injury from 
outside. This supposition is strengthened by the tact that larger 
quantities of lymph were found in specimens obtained during 
the dry weather than in those examined during the wet-season. 


5. The Axial Skeleton. 


An examination of the bones of the head revealed certain facts 
which, in my opinion, account for the narrow mouth of this toad 
in particular, and probably also for that of several other members 
of the same family in general. While the contour of the mouth 
which corresponds to the distal circumference of the skull, is a 
parabola in Rana, the same has been in this animal reduced to 
a mere are. Assuming that the skull-morphology in Rana is 
primitive, one can trace the narrow mouth in question to a 
generally vestigial condition of most of the bones of the suspen- 
sorium. Thus the squamosal has lost the anterior arm or the 
zygomatic process, what remains probably corresponding to the 
middle + and posterior arms of the same in Rana (text-fig. 4 A). 
The palatine, for another instance, is so inconspicuous, being 
about 3mm. long, that it can only be found in skulls very care- 
fully prepared. The remaining bones of the suspensorium also 
exhibit in a greater or a lesser degree the same degenerate state ; 
the cumulative effect of this, in all probability, was the forward 
displacement of the articular head for the lower jaw. For, a line 
drawn along the suture between the nasals and the fronto- 
parietals cuts the orbital fossee at about the middle and passes 
through the heads in question (text-fig. 40), whereas in Rana the 
same line lies far in front of the last (text-fig. 4D). This shifting 
of the articular heads necessitated by the vestigial condition of 
the bones of the jaws has probably led to a diminution in the 
contour of the mouth. 

The cranium as a whole is remarkable for its shortness and 


* This character is greatly exaggerated in C. globulosus, judging by the specimen 
in the Government Museum, Madras. Beddard (1) has recorded a similar condition, 
still more pronounced in Breviceps verrucosus, an Ethiopian Engystomatid. 

+ The presence of this middle arm is also doubtful, as the attachment with the 
‘pro-oti¢ is by means of a ligament which may, however, be taken to represent this 
arm. 


534 MR. D. W. DEVANESEN ON THE 
Text-figure 4 (A, B, C, D, and E). 


rae 


age 
eta 
BF 


S 


x ec) 
M 
CS 


A. A dorsal view of the skull of Cacopus systoma. The vestigial condition of the 
squamosal (Sq.) is remarkable. L., ligament connecting the latter with the 
pro-otic. For other letters consult the key-list of abbreviations. 

B. A ventral view of the same. P., vestigial palatine; T.F., trigeminal slit. For 
other letters consult the previous text-fig. (A.) and the key-list of abbreviations. 

C. A lateral view of thesame. L-M., a straight line drawn along the suture between 
the nasals and the fronto-parietals of the cranium. A-B, the plane in which 
the proximal circumference of the skull lies. C-—D, the plane in which the 
distal one of the same les. A.H., articular knob for the lower jaw; Fr.P., 
fronto-parietals ; Na., nasals. 

D. A lateral view of the skull of Rana hexadactyla. A brief comparison of the 
Cacopus skull with this will reveal the differences mentioned in the text. 
L.M., the same vertical line mentioned in the previous text-fig. (C). For other 
letters consult key-list. 

Hi. Lower jaw of Cacopus systoma, as seen from below. A., angulosplenial; M., 
Meckel’s cartilage; D., dentary; M.M., mentomeckelian; C., cartilaginous 
portion of the latter. 


ANATOMY OF CACOPUS SYSTOMA. 535 


relatively greater width, this being mainly due to the spheneth-— 
moid, which is extremely short, and it does not exhibit the 
prismatic shape usually seen in that of Rana. Hach mento- 
meckelian of the lower jaw carries at its posterior end a carti- 
laginous triangular piece which stretches behind and is free from 
the other component parts of the same jaw. ‘These two pieces 
are useful in so far as they provide partial attachments to the 
‘sub-mentalis muscle below and the genioglossus above (text- 
fig.4 E). 

The semilunar piece of cartilage in connection with the anterior 
cornua of the hyoid bone in Hemisws, described and identified 


Text-figure 95. 


The hyoid skeleton of Cacopus systoma viewed from below. Ex.H., “ extra-hyal’ 
A.C., anterior cornu; P.C., posterior cornu; B., basihyal or the hyoid plate; 
A.C.P., anterior cartilaginous process of the basihyal; P.C.P., posterior one of 
the same; A.B.B., anterior body-process of the basihyal or the proximal part 
of an anterior cornu; N., notch between the latter; S., thick cartilaginous part 
between the proximal ends of the two posterior cornua, probably belonging to 
the latter. 


as “the extrahyal” by Beddard (1, p. 909), appears to be repre- 
sented also in this toad (text-fig. 5, Ex.H). <A closer approxi- 
mation and fusion of the two extrahyals would complete the 
notch into a foramen corresponding to that of Henvisus. 

In the vertebral column of this toad one finds also a few 
interesting features. The transverse processes are comparatively 
short, taking the size of the centra and the neural arches into 
consideration ; those of the third vertebra are the longest ; next 
in order follow the ninth, fourth, second, eighth, seventh, sixth, 
and the fifth; the last three may be subequal. The spinous 
processes may be said to be absent in the first, sixth, seventh, 
eighth, and the ninth vertebra, while in the rest it is probably 
represented by a faint median ridge. The thick transverse 
processes of the second, third, and the fourth vertebra, the thin 


536 _ MR. D. W. DEVANESEN ON THE 


and slender ones of the fifth, sixth, seventh, and the eighth 
vertebra (those of the last three being directed conspicuously 
forward), the much dilated one of the sacrum, and the crestless 
urostyle combine to give a characteristic appearance to the 
vertebral column of this toad (text-fig. 6). It may be mentioned 
that the sacral diapophyses are remarkable for their uncommonly 
well-developed epiphysial cartilage at their free edges. 


Text-figure 6. 


Quee® 
I 
ay 


A dorsal view of the bones of the vertebral column of Cacopus systoma. 


I noticed an interesting abnormality in one specimen, namely 
the occurrence of two sacral vertebre, the‘ posterior one being 
fused with the urostyle. This condition is found normally to 
exist in Pelobates, Pipa, and Hymenochirus (5, p. 22). 


‘ 6. The Appendicular Skeleton. 


In the pectoral girdle the absence of the precoracoid probably 
necessitated the disappearance of the omosternum and the 
episternum. In the place of the sternum proper and the xiphi- 
sternum, one finds a broad, flat plate of cartilage generally calci- 
fied in the centre.’ This plate gives attachment to four different 


ANATOMY OF CACOPUS SYSTOMA. 537 


muscles, viz., the obliquus-internus, the inner portion of the 
sternohyoideus, “ sternocornuoideus,’ and the “abdominis-sternalis,” 
the last three to be described later in the section on muscles. 
This probably accounts for the unusually large size of this xiphi- 
sternal plate (text-fig. 7 A). 

The “ crista medialis,” a ridge of the humerus which, accord- 
ing to Ecker (4, p. 42), occurs among the males of three Huro- 
pean species of Rana as a sexual difference, is not to be found 
in the males either of 2. hexadactyla or of C. systoma. The 
wrist-bones of the latter show greater calcification than those of 


Text-figure 7 (A and B). 


A. The pectoral girdle of Cacopus systoma. X., xiphisternum; C., coracoid ; 
S., scapula; S.S., suprascapula; B., an apparently bony portion of the latter. 

B. The bones of a manus of Cacopus systoma. S., a sesamoid bone situated in a 
tendon; H., vestigial first finger. For other letters consult the key-list of 
abbreviations. 


Rana; they also exhibit certain differences in their shape and 
in the details of their articulation with one another. In addition 
to the three bones of the proximal row mentioned by Hecker, one 
very often finds a fourth bone, which is exceedingly small, 
situated above and between the os pyramidaleand the os lunatum. 
This may be a sesamoid bone, as it is placed in the course of a 
tendon (text-fig. 7 B). 

In the pelvic girdle the absence of the iliac crest is remarkable 
when one finds that none of those muscles connected with the 
same crest in Rana is absent. Whether this crestless condition 


538 MR. D. W. DEVANESEN ON THE 


of the ilia as well as that of the urostyle denotes a primitive 
feature, or whether it is a sign of degeneration, cannot be easily 
settled for the reason indicated above. At the free anterior end 
of each ilium there is found, attached to its outer side, a peculiar 
cartilaginous appendage. This has the shape of a plate, which, 
by folding on itself, encloses a groove into which is received the 
cartilaginous free edge of a sacral diapophysis. Thus the sacrum 
movably articulates with the ilium (text-figs. 8 & 10). In other 
words, the pelvic girdle, by means of the ilia, forms a gliding 
joint with the backbone through the sacrum. On the contrary, 
the sacral transverse processes in Rana are immovably attached 
by tissue to the anterior ends of the ilia, and consequently no 
movement in the manner mentioned above could take place. I 
was able to observe in emaciated specimens the ilia moving 


Text-figure 8 (A and B). 


A. The pelvic girdle of Cacopus systoma viewed from above. I., ilium; C.A., 
cartilaginous appendage of the latter. 
B. The same girdle seen sideways. P., pubis; I.S., ischium; A., acetabulum. 


obliquely backward and forward during the act of burrowing. 
I should therefore suppose that this gliding joint has something 
to do with the same habit of burrowing. 

Beddard (1, p. 12), with reference to a cartilaginous plate 
unconnected with the ilium but covering the sacral diapophyses 
in Breviceps verrucosus, remarks as follows :—“ I imagine that 
this cartilage belongs to the ilium, and that it is in consequence 
related to it as the suprascapula is to the scapula. We have, in 
fact, in this frog an exaggeration of the grooving which the 
anterior end of other forms shows at its line of articulation with 
the sacral vertebra. It is, moreover, interesting to observe that 
we find in this frog a kind of foreshadowing of the relations 
which the ilia bear to the sacrum in the higher Sauropsida, 


ANATOMY OF CACOPUS SYSTOMA. 539 


especially birds where the ilia are not merely attached to but 
cover the sacral vertebree.” In this Indian toad, however, the 
cartilaginous appendage in question not only covers the epiphysis 
of a sacral transverse process, but is also attached to an ilium. 
From the outer surface of the same appendage arises the «lio- 
lumbaris, a muscle which in Rana starts from the distal end of 
the ilium. ‘These facts probably indicate that it really belongs 
to the latter, in which case the remarks quoted above may be, 
with less diffidence, applied also to the same ilio-sacral relation 
in Cacopus. 

The pubis is interesting, as it exhibits signs of ossification 


Text-figure 9 (A and B). 
A. B. 


ne 
ne 
Bl 


A. A side view of the three bones of the pelvic girdle of an old adult Cacopus. It 
will be seen that the boundary of the pubis is indistinguishable; letters as in 
text-fig. 8 (A and B). 

B. A transverse section through the same pubis which suggested ossification. 
A., mass of degenerate cartilage at the symphysis: B., supposed network of 
bony tissue; C., degenerating cartilaginous matrix; V., probably a passage for 
a blood-vessel. 

which, if real, would be another anticipation of what occurs 

normally among the higher vertebrata. In older specimens the 

ilium and the ischium appear to gradually encroach upon it till, 
at last, in very old ones, it becomes indistinguishable from either 

of them, both in its area and in its appearance (text-fig.9 A and B). 

This suggests that, while there may be an ossification of the same, 

it has no separate centre, and that the process of substitution 

progresses from the adjacent boundaries of the ilium and the 
ischium. Sections through such a pubis after decalcification with 
picric acid show a structure which may be interpreted as an 


540 MR. D. W. DEVANESEN ON THE 


incipient stage in ossification by substitution (text-fig. 9B). No 
cartilage cell could be seen. The Haversian system may not be 
expected, as the pubis, if at all ossified, is more likely to become 
a spongy or cancellated bone. A median translucent mass (A.) 
found at the symphysis, I presume is a residue of the decaying 
cartilage not yet replaced. Jn this connection, | may mention 
that a similar phenomenon has been noticed in another toad, 
namely Pelobates (Gadow, p. 27). 

Among the bones of the hind-limb, the femur, the “os navi- 
culare,” and the prehallux deserve attention. There is, on the 
ventral side of the femur, a distinct crest stretching from the 
middle to the anterior end, stopping short of the head of the 
same. It is from this ‘‘ femoral crest” that the vastus internus 


Text-figure 10. 


A dorsal view of the ilio-sacral joint and a few muscles connected with the same. 
For lettering consult the key-list of abbreviations. 


muscle arises. To this crest is also attached a considerable 
number of fibres of the adductor longus, adductor brevis, and the 
pectineus (text-fig. 19 B and C). 

The os naviculare in the tarsus of this toad is a circular disc, 
convex in front and concave behind, and articulating exclusively 
with the astragalus. In front it is bound by tissue, and articu- 
lates as well with a certain piece of highly calcified cartilage 
which, as will be seen presently, is homologous with the prehallux 
of Rana. Though the tarsal bone in question has become so 
closely associated with the prehallux as to look like one of the 
component pieces of the latter, yet its identity could be recog- 


ANATOMY OF CACOPUS SYSTOMA, 54] 


nised with certainty by its connection with the muscle adductor 
longus digitt 1 (Keker, p. 109; & text-fig. 11 A). 
On the plantar surface of the foot in several forms of the Anura, 


Text-figure 11 (A, B, and C). 


A. The bones of a pes of Cacopus systoma. P.H., prehallux ; O.N., os naviculare ; 
A., astragalus ; C., caleaneum; O.C., os cuboideum, 

B. A side view of a pes of Cacopus, showing the relation of the muscle abductor 
hallucis A. to the prehallux P.H. P., plantaris; F.D., flexor digitorum; III, 
IV, and V, longus ; S., skin cut and deflected upward. 

C. The skin (Sk.) on the plantar surface of the same, cut and deflected sideways 
in order to expose the muscles connected with the tubercles in question. G. 
gastrocnemius, showing its relation to the “os sesamoide” (o0.s.) or the outer 
metatarsal tubercle; P., plantaris-inserted by means of a fascia on to the 
prehallux (P.H.) or the inner metatarsal tubercle. 


particularly in those with a fossorial habit, has been observed a 
certain prominent tubercle of a horny consistency. It would 


542, MR. D. W. DEVANESEN ON THE 


appear that till now this has been generally regarded as a 
cutaneous structure, rendered hard and horny in consequence 
of the nature of its function which is the scooping-out of the 
soil in suitable localities. This Indian toad has two such con- 
spicuous tubercles, called by Boulenger (2) the ‘inner metatarsal 
tubercle” and the “ outer metatarsal tubercle.” On maceration, 
it was found that not only these were not purely cutaneous 
developments, but also that they had their counterparts in Lana. 

It is a well-known fact that the Anura possess, in addition 
to the five toes normally present, an extra vestigial digit—the 
prehallux. Among the Indian frogs I examined, this digit con- 
sisted of four cartilaginous pieces in Rana hexadactyla, of two 
pieces in Rana tigrina, and of a single piece in Lana breviceps 
and Cacopus systoma. The first-mentioned frog is entirely aquatic 
in habit, and its prehallux forms a slight projection covered with 
ordinary skin, while in the three last-mentioned ones, which are 
burrowing forms, the same prehallux is converted into a con- 
spicuous shovel-shaped organ enclosed in a hard and horny dermal 
sheath (text-figs. 11 A & 12A, B,andC). The so-called “ inner 
metatarsal tubercle” of C. systoma is therefore none other than 
the distal piece of the vestigial toe of Rana which has acquired 
a large size and a shape suitable for its function in this toad. 
That this is so was confirmed by an examination of its connection 
with a muscle, viz. the abductor hallucis, which is inserted on to 
the prehallux—a muscular relation identical with what obtains 
in Rana (Kcker, p. 109). 

In the two terrestrial species of Rana just mentioned, one 
finds a gradual reduction in the number of pieces constituting 
the prehallux, assuming the condition in R&. hexadactyla to be 
the normal one. Simultaneously with this reduction, it would 
appear that in these forms there has been an increase in the size 
and a change in the shape of the distal piece of the prehallux. 
This reduction, however, may be regarded as advantageous, for 
the fewer the number of joints in an organ employed for digging, 
the greater will be its efficiency. Thus we find in C. systoma a 
single large piece constituting the prehallux*. 

it may be required why I homologise the single element of 
the prehallux of Cacopus with the distal piece and not with the 
proximal one in that of Rana. As the latter piece is in contact 
with the os naviculare, just as that of the prehallux of Cacopus 
is, one may be inclined to the alternative opinion mentioned. 
But a careful reference to the text-fig. 12 A, B, and C will show 
that my view is the more probable one; for it will be seen that 
while in Rana tigrina the distal piece numbered 4 undoubtedly 
corresponds to the same piece 4 either in Rana breviceps or in 
Cacopus systoma, in the latter two species the proximal piece 1 
of the former has disappeared. According to the alternative 
view, one will have to say that the distal piece 4 of Rana tigrina 


* T venture to suggest that the so-called spade of the “‘Spade-footed Toad” or 
Pelobates of Europe is also the prehallux modified. 


ANATOMY OF CACOPUS SYSTOMA. 543 


has disappeared in Rana breviceps and Cacopus, and the proximal 
piece 1 has acquired a shape similar to piece 4: this explanation 
appears more Ingenious than natural. Further, in accordance 


Text-figure 12 (A, B, and C). 


A 


O 
o 
oO 


A. The bones of a pes of Rana hewadactylu, Lettering same as in A of text-fig. 
11. It will be noticed that the prehallux consists of four pieces. 

B. The same of Rana tigrina. The prehallux is made up of only two pieces, 
viz. 4 and 1; 2 and 3 have probably disappeared. One may alternatively 
assume that 3 and 4 of R. hexadactyla have been omitted, and that 4 in 
R. tigrina corresponds to 2in the latter. Lettering same as in text-fig.11 A. 

C. The same of Rana breviceps. The prehallux is composed of but a single piece, 
like that of Cacopus. Lettering same as in text-fig.11 A. 


with the function of the prehallux in these frogs, it will be a 
distal piece which will come into direct play, and it will con- 
sequently lend itself to an adaptation in shape and size. 


544 MR. D. W. DEVANESEN ON THE 


in this toad a further specialisation of the digit in question 
seems to have been brought about in its relation to the tarsus. 
It is firmly attached to the os naviculare, as has been already 
described in connection with the latter, and through the inter- 
vention of the same it articulates exclusively with the astragalus. 
This articulation, I should think, in combination with the action 
of the muscles attached to it and to the tarsal bone mentioned, 
bestows a considerable advantage in the act of burrowing (text- 
fig. 11 BandC). In &. tigrina and R. breviceps the specialisation 
has not been carried to that extent; the same indirect articula- 
tion of the prehallux with the astragalus is only partial, as the 
first digit still continues to articulate with the latter indirectly 
through the os naviculare ; further, the prehallux and the tarsal 
bone connected with it are, in these frogs, much less calcified, 
and consequently they are less rigid and hard. Again, the 
plantaris muscle, which has no relation with the prehallux in 
Rana, has a connection with the same in this toad, in that it is 
indirectly inserted on to the latter by means of a fascia (text- 
1aye2, IIL (O))), 

It is exceedingly interesting to find the digit, which is generally 
believed to be a vanishing structure in the Anura, re-asserting 
itself under suitable conditions and becoming an organ of no 
mean importance to its possessor. This specialisation of the 
prehallux among the species of two different families, viz. the 
Engystomatide and the Ranide, may be regarded as a case of 
convergence. 

The “ outer metatarsal tubercle” of this toad, though smaller 
than the inner one, consists likewise of a single cartilaginous 
piece covered with a horny dermal layer. I identify this as 
the “os sesamoide” (Ecker, p. 106) of Rana, which is a carti- 
laginous element situated on the plantar side of the joint 
between the calcaneum and the fourth metatarsal bone. In this 
latter position exactly I found the cartilage of the “‘ outer meta- 
tarsal tubercle” situated in C. systoma, so that this also is no 
new structure developed by this toad, but is only a specialised 
and an exaggerated one of what is normally present in Rana. 
Nevertheless, it would seem that a remarkable specialisation had 
been brought about in its alliance with no less an important 
muscle than the gastrocnemius. 'To this ‘‘ os sesamoide” is exclu- 
sively inserted the narrow, attenuated yet strong band of the 
aponeurosis plantaris. This implies that an important calf muscle 
is concerned in the working of the “outer metatarsal tubercle” 
(text-fig. 11 C). 


7. The Musculature. 


The muscles of the hyoid apparatus and of the floor of the 
mouth are characterised by a great specialisation in this toad, 
a remarkable fact observed also by Beddard (1) in three other 
members of the same family. However, in the details of the 
specialisation, excepting perhaps in the sternohyoideus, one does 


ANATOMY OF CACOPUS SYSTOMA. 545 


not, find any features which could be distinguished as common to 
Cacopus and to those described by the authority mentioned. 

We will follow Beddard in calling the posterior portion of the 
sub-maxillaris the sub-hyoidews, and in restricting the original 
name to the anterior portion of the same muscle. Now this 
“ sub-hyoideus” is much larger than the sub-macxillaris—the 


Text-figure 13. 


ma 


= 


— 


Certain muscles on the ventral surface of the trunk and head of Cacopus systoma 
have been exposed. T., an inseriptio-tendina of the inner division of the rectus- 
abdominis; C.F., ventral extension of the “cervical fat-body”; I., the spot 

‘ where the outer division of the rectus-abdominis and that of the portio- 
abdominis ave attached to the skin; P.S., pectoral septum; S.M., swb- 
mawvillaris; S.H., sub-hyoideus, For other letters see key-list of abbreviations. 


reverse of what obtains in Lana (text fig. 13). The proportion 
which the former muscle assumes in the male is in conformity 
with the differentiation of a median vocal sac. It is a posterior 
portion of this muscle which, along with the skin in this region, 
becomes enormously expanded to form a large spherical resonator 


546 MR. D. W. DEVANESEN ON THE 


during the time of croaking (text-fig. 14). When at rest, this 
part of the muscle is withdrawn and stowed away in a con- 
spicuous double fold above the anterior portion of the same 
muscle. 

On removing the swb-maxillaris and the sub-hyoideus several 
inner muscles of the floor of the mouth come into view (text- 
fig. 15). It is seen that the transverse fibres of the swb-men- 
talis (S.M.) pass from the inner border of one mentomeckelian 
to that of the other, thus having apparently * no connection 
with the dentary. It is also seen that a greater part of the 
same muscle is connected with the cartilaginous piece borne by 
each mentomeckelian, which has been already described in the 
section on the axial skeleton. Of greater interest is the genio- 
hyoideus muscle, whose origin and insertion are very different 
from what obtains in Rana. The outer portion of this muscle 


Text-figure 14. 


A male Cacopus with the vocal sac in action; drawn from life; half natural size. 
The enormous distension of the vocal sac is remarkable; under natural 
surroundings it looked more like a float than a resonator. 


arises from the angulosplenial and passes on the outer side of 
the mentomeckelian cartilaginous piece and that of the sterno- 
hyoideus muscles to be inserted on to the posterior cornu (text- 
figs. 15 & 16 A). Though it is widely separated from the inner 
division, it is connected with the latter by some of its fibres, 
which diverge from its body at about the place of origin and 
become confluent with the same inner portion, so that on the 
right side of the toad this outer portion and that part of the 
corresponding inner division lying in the pre-hyoid part of 
the floor of the mouth have together the shape of the letter N. It 
will be noticed that a few of the connecting fibres are intercepted 


* It looks as though the cartilaginous part of the mentomeckelian has insinuated 
itself across the fibres of this muscle; for one finds between the outer side of this 
same part of the lower jaw several muscular fibres which I presume belong to the 
sub-mentalis (text-fig. 15, F.M.). 


ANATOMY OF CACOPUS SYSTOMA. 5AT 


by the cartilaginous portion of the mentomeckelian (text-fig. 16 A). 
The inner divisions of the two geniohyoideus muscles are still 
further remarkable, for,in the region of the basi-hyal, they fuse 

to form an extensive median sheet of muscle with two posterior 
offshoots, each of which, spreading on a posterior cornu, entirely 
conceals the Ayoglossus muscle in the usual position of the toad 


Text-figure 15. 


(if 
SE 


et 


A view of certain superficial muscles of the hyoid and deep-lying ones of the floor 
of the mouth. The lower jaw (L.J.) has been divaricated; the sawb-mawillaris 
and the “ swb-hyoideus ” have been removed; a right half of the xiphisternum 
has been cut away in order to expose at their origins the inner portion of the 
sternohyoideus (ST. I.) and the “ abdominis-sternalis” (A.S.). R.A., continua 
tion of the “inner division of the rectws-abdominis” as the “ outer portion of 
the sternohyoideus” ST.; I.3 and 1.4, third and fourth inseriptio-tendina of 
R.A.’ respectively; F.M., certain muscle-fibres between the lower jaw and the 
cartilaginous part of a mentomeckelian. For other letters consult the key-list 
of abbreviations. 


1 \ 
\ 


Th 


C.R II 
FM NY WN = DN 
hel \\' Wi \ * 
Sir ‘on i | \ 
R.A ! : (ihe | ay \ 
WZ / \ 
i | 
i i 
Pyetl 


during dissection. The insertion of the former, in consequence 
of its extensiveness, is extraordinary; for it consists of a peri- 
meter of the inner lateral edges of the two posterior cornua and 
the hind border of the basi-hyal enclosed between these two 
cornua, thus forming a parabolic curve (text-fig. 16 B, G.”’). 

The single ‘outer portion of the sternohyoideus” (or the 


* A condition similar to this has been observed by Beddard (1) in Breviceps. 


Proc. Zoot, Soc,—1922, No. XX XVIII. 38 


548 MR. D. W. DEVANESEN ON THE 


sternohyoideus proper) has differentiated into two separate 
muscles in this toad, one lying above the other, This differen- 
tiation starts at the fourtlr and the last inscriptio-tendina of the 
rectus-abdominis. A similar division of this muscle has been 
observed in Breviceps and Hemisus by Beddard (1). What corre- 
sponds to the “inner (median?) portion of the same muscle” 
is a strip which arises entirely from the xiphisternal plate (text- 
figs. 15, ST. I., & 16). Now, these three divisions of the sterno- 
hyoideus are separately inserted on to the body of the hyoid, 
one behind the other and in close succession; none of the fibres, 
however, is attached either to the posterior cartilaginous process 
of the same hyoid plate or to the posterior cornu. 

On removing the hyoid portion of the inner gentoglossus 
muscles, the hyoglossws pair comes into view which, in turn, 
conceals another muscle in the ordinary position of dissection. 
The thick, coarse fibres in the interior of the azygous portion of 
the hyoglossus are either sinuous or twisted like a corkscrew. 
As this muscle is the retractor of the tongue, this condition, at 
rest, of the fibres mentioned suggests that they are capable of 
being stretched to a greater extent than usual so as to permit 
the tongue to reach a comparatively longer distance. In Rana 
the paired portion of the same hyoglossus is the innermost 
muscle in contact with the lower surface of the posterior cornua ; 
but, in this toad, there intervenes a new pair of muscles between 
the two, which arises from the distal quarter of the latter, and 
is inserted to a strong fascia spreading in the space enclosed by 
the proximal parts of the anterior cornua and the anterior 
margin of the basi-hval (text-fig. 16 C,C.H.). Among the muscles 
connected with the hyoid skeleton, there are several which are 
solely attached to the posterior cornua on the one side and to 
the bones of the skull and of the pectoral girdle on the other, 
Now, it is likely that this new muscle, which we may term the 
cornuoideus-posterior, counteracts a probably too powerful pull 
of these muscles in question on the same cornua by bindimg the 
latter with the basi-hyal plate. One finds another new pair of 
muscles, not recorded in Kana, arising from the xiphisternum 
and running forward beneath the auricle to be attached to the 
posterior cornu (text-fig. 16 A,S.C.); this may be called the 
sterno-cornuoideus in accordance with its origin and insertion, 
The petrohyoideus muscles are distinguished by their robust 
development and by their numerous insertions. The peéro- 
hyoidews-anterior arises from the pro-otic by means of a single 
tendon, and spreads over the outer surface of the pharynx, to 
whose wall all its fibres are more or less adherent (text-fig. 16 A, 
B, C, and D). According to the several insertions of this muscle, 
one may recognise the following divisions: (1) a band, P.A. 1, 
inserted on to the fascia in relation with the proximal parts of 
the anterior cornua; (2)another anterior portion, P.A. 2, attached 
to the root of the same cornua; (3) a small narrow division, 
P.A. 3, attached to the lower surface of the basi-hyal; and (4) a 


ANATOMY OF CACOPUS SYSTOMA. 549 


Text-figure 16 (A, B, C, D, and E). 


A. The deep muscles of the hyoid and of the floor of the mouth. The sub- 
mentalis has been taken away. The sterno-hyoideus muscles on the right side 
have been severed at their origins and deflected while those on the left have 
been removed. O.G., an outer division of the geniohyoideus; I.G., an inner 
one of the same; [.G’., a fused median portion of I.G.; I.G.’’, a posterior 
offshoot of I.G.’ concealing a hyoglossus below; 8.C., sterno-cornuoideus ; S.C.', 
cut end of the same. For other letters consult the key-list of abbreviations. 


B. The deep muscles of the hyoid continued ; the geniohyoideus muscles have been 
removed excepting those fibres of I.G.’ and I.G.” at their insertion (G.") which, 
as will be seen, isa parabolic curve. ST.1 and ST.2, the two divisions of the 
“outer portion of the sternohyoideus” or the “ sternohyoideus proper.” P.A.1, 
an anterior division of the petrohyoideus; O.G.', cut end of the outer genio- 
hyoideus ; ST.’, cut ends of the sternohyoideus muscles ; F., fascia; H., hyo- 
glossus. Other letters as in A. 


‘C. The same muscles continued. The hyoglossus in the region of the basihyal has 
been cut away. C.H., “ cornuoideus-posterior”’; H'., cut end of the azygous 
hyoglossus; P.A., petrohyoideus-anterior; P.A.1, an anterior division of the 
latter. The remaining letters as in the other figures of this text-figure. 


D. The same muscles continued. The “cornauoideus-posterior”’ has been removed. 
P.A.2, P.A.3, and P.A.4, the several bundles into which the petrohyoideus- 
anterior separates itself before its insertion; P.P., petrohyoideus-posterior. 
For other letters consult the key-list of abbreviations. 

#. Certain muscles of the tongue viewed from below. The median and azygous 


part of the genioglossus (G.G.) has been cut in the middle in order to expose 
the muscle G.G.', which corresponds to the outer portion of the genioglossus, 
YQx - 


@ 


550 MR. D, W. DEVANESEN ON THE 


posterior thick division, P.A. 4, inserted on to the upper surface 
of the same plate. As regards the petrohyoideus-posterior (P.P.), 
while a small part of it clings to the cartilaginous free end of a 
posterior cornu, the bulk of the fibres pass above the latter and 
are inserted to the pharynx, a few fibres reaching also to the 
larynx. I could not find any trace of the omo-hyoid muscle, a 
negative feature observed also in Breviceps and Hemisus. 

Of the muscles of the tongue, it may be noted that the relative 
position of the two divisions of the genioglossus has undergone 
an alteration. The one corresponding to the outer part in Rana 
lies above the other corresponding to the median part of this 
muscle. Further, the former has shifted its insertion from the 
mucous membrane of the floor of the mouth to the body of the- 
tongue (text-fig. 16 E). 

Text-figure 17. 


SER...) aCe Mtoe 


View of certain muscles of the chest. On the right side the portio-sternalis- 
anterior and the portio-abdominis have been removed. S.R., sterno-radialis ; 
P.S., portio-scapularis or the vestigial deltoideus; P.S.A., portio-sternalis- 
anterior; P.S.A’., cut end of the same on the right; C.H., coraco-humeralis ; 
P.A’., outer division of the portio-abdominis; P.A.", an inner division of the 
same muscle; X., xiphisternum, 


It seems to me that the specialisation described above of the 
muscles of the hyoid and of the floor of the mouth is in some 
way connected with a more efficient method of obtaining food. 

The muscles of the chest as a whole, with the exception 
of the portio-sternalis-anterior and the portio-abdominis, exhibit 
a feeble development (text-figs. 13 & 17). This, in my opinion, 
is mainly due to the disappearance of the pre-coracoid, the 
omosternum, and the episternum. For instance, the sterno- 
radialis, which arises in Rana from the three elements of the 
pectoral girdle mentioned, has its origin shifted to the inner 
ventral border of the coracoid, which is less extensive; conse- 
quently this muscle is vestigial*. Again, the deltoideus exhibits 
a degeneration in the absence of one of its two divisions, 


* In this connection it may be mentioned that this muscle is absent in. 
Breviceps (1). 


ANATOMY OF CACOPUS SYSTOMA, 55 


viz. the pars-clavicularis. It would also appear that as the 
functions of these two muscles of the chest have become reduced 
to a minimum in this toad, a large size and an extensive origin 
for the same have been rendered unnecessary. According to 
Ecker (4), in Rana the sterno-radialis is a powerful flexor of the 
fore-arm, and the deltoidens draws the same limb forwards. These 
movements, though essential to a good swimmer, are in all 
probability of only a limited use to a terrestrial frog. Thus it 
becomes difficult to say whether the vestigial condition of the 
chest-muscles led to the disappearance of certain of the elements 
of the pectoral girdle, or the latter condition necessitated the 


Text-figure 18. 


A ventral view of the trunk and a thigh with the skin cut open in order to show 
the relation of certain muscles with the skin. I., several fibres of the rectus- 
internus-minor (R.1'.) arising from the skin (SK.) on the lower surface of a thigh ; 
A., the region of the skin from which the outer division of the portio-abdominis 
starts and to which that of the rectus-abdominis is inserted; S., sartorius ; 
R.L., rectus-internus-major; V.1., vastus-internus. 


former. It is more probable that the two degenerative processes 
might have proceeded concomitantly. 

On the contrary, one finds that the portio-sternalis-anterior 
and the portio-abdominis muscles have attained a great develop- 
ment, and the outer division of the latter has become partially 
converted into a cutaneous muscle (text-fig. 18). The attach- 
ment to the skin has, I should think, given additional purchase 
to this muscle in pulling the front imb backwards and towards 
the body, as happens while burrowing. These two chest-muscles 
are probably put to a greater use in this toad at the expense of 
the sterno-radialis and the deltoideus. 


552 _ MR. D. W. DEVANESEN ON THE 


Text-figure 19 (A, B, and C). 


ASN 


a ~N 
sy 


A. Certain muscles on the lower surface of a thigh. C.A., caput-anticum of the 
semi-tendinosus; A.L., adductor-longus; A.M., adductor-magnus; A.M"'., a 
hind portion of the same. Other letters as in text-fig. 18. 

B. Most of the muscles on the lower part of the thigh have been removed in order 
to show a few deep ones. A.L., adductor-longus; P., pectineus; A.B., 
adductor-brevis; F.,a crest of the femur; LS., iliac symphysis; P.S., pubic 
symphysis. - 

C. A view of the vastus-internus (V.I.) showing the details of its origin. C., 
capsule of the hip joint; F., a crest of the femur; T.S., a band of tendinous 
strips starting from a strong tendon (T.). 


ANATOMY OF CACOPUS SYSTOMA. 553 


The rectus-abdominis muscle of the trunk is remarkable for 
having shifted its origin backward from the pubes to the ischia ; 
its outer portion is, along the greater part of its length, adherent 
to the skin below. This is significant when viewed along with 
the absence of the cwtanews-pectoris in this toad. At the third 
inscriptio-tendina of each rectws-abdominis one finds a small tri- 
angular portion, separated off from its main body, passing forward 
and attached to the upper side of the xiphisternal plate (text- 
fig. 15, A.S.). This muscle, which we may call “the abdominis 
sternalis,” should be regarded as a differentiated part of the inner 
portion of the rectus-abdominis, for in Rana some of the fibres of 
the latter in the same situation are attached to the xiphisternum. 
While the rectus-abdominis muscles are generally thin and 
attenuated, the obliquus muscles are better developed and are 
thrown into transverse puckers, and bulge out prominently on 
the sides on account of the pressure of the enormously developed 
viscera. : 

Among the muscles of the thigh, the rectus-femoris-anticus is 
absent, the triceps-femoris, composed of three divisions in Rana, 
is made up of only two, and the origin of the vastws-internus is 
somewhat different. Though most of the anterior fibres of the 
last arise from the capsule of the hip-joint, the rest of them arise 
froma long tendon given off by the same capsule (text-fig. 19 C, T.). 
Further, this tendon is adpressed to the shaft of the femur by 
tendinous slips which spring from it laterally. On one side of 
the tendon, namely the inner lower side, these slips are so closely 
set as to form a band (T.8.). As regards the rectus-internus- 
minor, one finds that, on its lower side, a considerable number 
of fibres take origin on the skin of the thigh (text-figs. 
18 & 19A). This partially cutaneous origin of the muscle 
recalls a similar condition in the frogs described by Beddard (1). 
The semi-tendinosus is inserted by a narrow tendon which spreads 
in a triangular form over the anterior head of the tibio-fibula, 
in this respect differing from that of Rana, where the tendon in 
question unites with that of the sartorius to form an aponeurotic 
arch. The adductor-longus, the pectineus, and the adductor- 
brevis may be considered as forming a single muscle, for they are 
not distinctly delimited from one another. The greater portion 
of this huge muscle is hidden from view by the sartoriws and the 
adductor-magnus, and where it is inserted on to the shaft of 
the femur its fibres along the middle are firmly attached to a 
crest of the femut already mentioned (text-fig. 19 B). 


8. Summary. 


(1) In the pharynx a new structure is situated to which the 
status of an organ may be given, and it has been accordingly 
named “‘ the pharyngeal organ of Cacopus systoma.” 

(2) Sub-cutaneous as well as sub-peritoneal fat-bodies are 
normally present in this toad; while the former are developed 


554 MR. D. W. DEVANESEN ON THE 


in relation to the lymphatic septa, the latter are probably 
deposited in connection with the peritoneal membrane. Their 
function, as has been ascertained, is to assist in the general 
metabolism of the body during estivation. 

(3) The stoutness of this toad is, in the first instance, due to 
the enormously developed viscera. This condition is improved 
upon by the inclusion of a part of the thigh into the contour of 
the trunk and by the enlarged sub-cutaneous lymph-sinuses. 

(4) In the skeleton the vestigial state of several of the bones 
of the head is remarkable. This and the consequent engysto- 
matous mouth are, in my opinion, related in some unknown 
manner to the nature of the diet, which consists almost entirely 
of termites*, winged or wingless according to the season. 
Witness, in this connection, the similar effect produced by a 
similar diet on the Myrmecophagide among Mammals. 

(5) It is noteworthy that the pre-hallux of Rana, usually 
regarded as a degenerate digit, has in this toad become an organ 
with a definite function, viz. the so-called “inner metatarsal 
tubercle.” 

(6) While certain muscles, for example the sterno-radialis and 
the deltoideus, show a remarkable degeneration, other muscles, 
for instance the muscles of the hyoid and of the floor of the 
mouth, exhibit an equally remarkable specialisation. The 
muscles examined also show important differences in their rela- 
tive dimensions, origins, and insertions. Certain new muscles 
are present which have been already described, while three 
muscles are absent, viz. the cutanews-pectoris, omo-hyoid, and the 
rectus-femoris-anticus. Again, certain non-cutaneous muscles of 
Rana have acquired a partial connection with the skin, for 
example, the rectus-internus and the portio-abdominis. Assuming 
the museular anatomy of Rana to be typical of the Anura, one 
may trace the differences in the musculature of this toad to two 
factors, viz. the fossorial habit and the nature of its food. This 
creature passes the greater part of its life underground, seeking 
water only for breeding purposes. This implies that, while the 
same muscles called into frequent activity in an aquatic frog 
may suffer by the want of regular use, other muscles of a direct 
utility in burrowing are likely to be improved and exaggerated. 
At the same time, the curiously specialised muscles of the hyoid 
and of the floor of the mouth probably indicate a more efficient 
method of capturing insects. 

(7) Beddard (1) mentions three characters which may be 
common to the Engystomatide: these are (1) the partially euta- 
neous origin of the rectws-internus-minor, the specialisation of 
the muscles of the floor of the mouth, and the division of the 
“outer portion of the sternohyoideus.” One may say that in 


* Although it is true that insects form the chief food of frogs in general, yet a 
liberal diet of termites alone appears to produce the peculiar effect on the consti- 
tution which we have indicated. 


ANATOMY OF CACOPUS SYSTOMA. 555 


these respects Cacopus systoma also agrees with the three genera 
described by him. Nevertheless, in the details of the second 
feature mentioned there is hardly any striking resemblance, I 
should add that the omo-hyoid muscle is generally absent among 
the same family, as it has not been found either in this toad or 
in Breviceps and Hemisus. 


mop Key-list of certain of the Abbreviations mostly unexplained 
in the explanations to Text-Figures. 


A.C.—Anterior cornu, 
A.H.—Articular head for the lower jaw. 


‘C.P.—A cartilaginous part of the 
mentomeckelian. 
C.S.—Cartilaginous epiphysis of the 


sacrum. 
‘C.S.'"—Coccygeo-sacralis, 
C.1.—Coccygeo-iliacus. 
Ex.0.—Ex-occipital. 
Fr.P.—Fronto-parietal. 
I.G.—Inner division of a genio-hyoideus. 
I.L.—LIlio-lumbaris. 
L.D.—Longissimus-dorsi. 
M.—Marxilla. 
M.M.—Mentomeckelian. 
Na.—Nasal. 
0.C.—Occipital condyle. 
0.C.H.—* Os-capitato-hamatum.” 
O.E.— OBliquus-eaxternus. 
O.G.—Outer division of a 
hyoideus. 
O.1.—Obliquus-internus. 
0.L.—* Os lunatum.” 
O.N.—‘“‘Os naviculare”’ of a manus. 
O.P.—‘ Os pyramidale.” 
O.T.—* Os trapezium.” 
O.Ts.—* Os trapezoides.” 
Pr.—Pterygoid. 


genio- 


P.A.— Petrohyoideus-anterior. 

P.A.’—Inner division of a portio- 
abdominis. 

P.A."—Outer division of the same, 

P.M.—Premaxilla. 

P.P.—Petrohyoideus-posterior. 

Pro.O.—Pro-otic. 

P.S.—Parasphenoid. 

Q.—Quadrato-jugal. 

R.A.'—Inner division of a rectus-abdo- 
minis. 

k.A."—Outer division of the same. 

SQ.—Squamosal. 

ST.—“ Outer portion of the sterno- 
hyoideus”? or the  sterno- 
hyoideus proper. 

ST. 1 and ST.2.—Two divisions of the 


same. 

ST. L.—An inner portion of the sterno- 
hyoideus. 

S.H.— Sub-hyoideus.” 

S.M.—Sub-maxillaris. 

8.M.’—Sub-mentalis. 

T’.—Tongue. 

T,—Terminal epiphysis of the urostyle. 

V.—Vomer. 

X.—Xiphisternal plate. 


10. References to Literature. 


1. Bepparp, F. EH. (1908).—** On the Musculature and other 
points in the Anatomy of the Engystomatid Frog, 


Breviceps verrucosus.” 
(1908).—“ A Contribution to the Knowledge of the 
Batrachian, Rhinoderma darwini.” 
Soe. 1908, p. 678. 
(1908).—‘‘ Some notes on the Muscular and Visceral 


2? 


29 


Proc. Zool. Soc. 1908, p. 11. 


Proc. Zool. 


Anatomy of the Batrachian Genus Hemisus, with 
notes on the Lymph-Hearts of this and other 


Genera.” 


Proc. Zool. Soc. 1908, p. 894. 


2. Boutencer, G. A. (1890).—“ Reptilia and Batrachia” in ‘The 
Fauna of British India,’ 
3. Boutencsr, C. L. (1916).—‘“* On certain Sub-cutaneous Fat- 


Bodies in Toads of the Genus Bufo.” 
Soc. 1910; p. 379. 


Proce. Zool. 


556 ON THE ANATOMY OF CACOPUS SYSTOMA. 


4. 
5. 


Koxsr, A. (1889).—‘ The Anatomy of the Frog. Translated 
by G. Haslam. 

Gapow, H. (1901).—‘ Amphibia and Reptiles.’ Camb. Nat. 
Hist. 


. GintuER, A. (1864).—‘ Reptiles of British India.’ 
. Pearson, J. (1910).—“ On Cacopus globulosus.” Spolia Zey- 


lonica, vol. vil. 
Tuurston, EK. (1887).—‘‘ Extracts.” Proc. Zool. Soc. 1887, 
> We se) . Bay 
», ~ (1887).—‘ Catalogue of the Batrachians of South 
India.’ | 


IP, ZA. So 1OP2, OMLIWATIR, IPL, Il 


3 
CETACER OF THE NEM ZEAL ARITS Crag 


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CETACEA TOR WHE, NEW. ZEAEAN DE SEAS: 


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PaeZ Se 1922, OLIN ER kee e 


CETACEA OF THE NEW ZEALAND SEAS. 


THE CETACEA OF THE NEW ZEALAND-SEAS., DO 


28. A Review of the Cetacea of the New Zealand Seas.—1I. 
By W.R. B. Ottver, F.L.S., F.Z.8., Dominion Museum, 
Wellington, N.Z. 

[Received May 18, 1922: Read sian 18, 1922. 
(Plates I.-1V.*) 


Except the popular accounts by Waite, and by Hutton and 
Drummond, no general list of the New Zealand Cetacea has been 
published since Hector’s paper on the Delphinide in 1887. 
Having a number of notes made on various occasions to put on 
record, I take the opportunity of reviewing the Whales and 
Dolphins of our seas. The recorded knowledge of the group is 
very incomplete; and, moreover, some errors are apparent. The 
museums at Wellington and Christchurch contain a fair number of 
skeletons; those of Dunedinand Wanganui fewer. The only repre- 
sentatives of the Cetacea in the Auckland Museum appear to be 
three skulls. The specimens in Christchurch and Dunedin have, in 
the main, been recorded, but in Wellington and Wanganui there 
are some skeletons. of considerable interest which have not hitherto 
been described. These are mentioned in the present. paper. 
Besides cataloguing the specimens in the principal museums in 
New Zealand, I have included those specimens from New Zealand 
that I have been able to trace in the museums of Australia, 
America, and Europe. Many .of these I have examined. The 
short descriptions given with the lists refer mainly to variable 
characters ; and, where I myself have not examined the specimen, 
the author from whom I have quoted is given: An attempt has 
also been made to give the generic and specific names in 
accordance with the International Rules of Zoological Nomen- 
clature. The literature on the Cetacea is very extensive, but 
some of it has to be used with caution. This applies to many of 
the writings of Dr. J. EH. Gray, who unfortunately gave new 
names on quite insufficient grounds, sometimes founding genera 
and species on drawings and photographs he appeared not to 
understand. JI have quoted only the principal references, 
including those where new names are proposed or where there 
are descriptions based on specimens, 

I cannot claim.to have added a single species to the list of 
New Zealand cetaceans; on the contrary, I have omitted two, 
Delphinapterus leucas and Hyperoodon planifrons, and hope that 
the evidence that neither belongs to our fauna will be accepted. 

Tt will appear from the following account that there is much to 
be learned about. the distribution and habits of the cetaceans of 
our seas; moreover, collection of perfect: skeletons and descriptions 
of the external characters of the animals is much to be desired. 
In some cases the species are only known from skulls or skeletons. 


* For explanation of the Plates, see p. 585. 


‘558 MR. W. R. B. OLIVER: REVIEW OF THE 


It is hoped, therefore, that the present paper will be found useful 
in bringing together what is known on the subject, so that the 
gaps in our knowledge may be filled up. A great deal of 
information could be gained if, when a whale or dolphin is 
stranded, a description and, if possible, a photograph were sent 
to the nearest museum. By this means the animal might be 
correctly identified, and if belonging to a rare species, steps might 
be taken to preserve the skeleton. 

For supplying lists of specimens in the institutions under their 
charge, I have to thank the Director, British Museum (Natural 
History), London, the Secretary, United States National Museum, 
Washington, D.C., and the Director, Muséum d’Histoire Naturelle, 
Paris. 


BALENA. 


Balena Linné, 1758, Syst. Nat. ed. 10, p. 75; type, B. mysticetus 
L. Hubalena Gray, 1864, Proc. Zool. Soc. 1864, p. 201; type, 
Balena australis Desmoulins. Caperea Gray, 1864, |. c. p. 202 ; 
type, Lalena antipodarum Gray. 


BALENA AUSTRALIS. 


Balena australis Desmoulins, 1822, Dict. Class. Hist. Nat. 
vol. ii, p. 161 (Cape of Good Hope); Waite, 1909, Subant. Is. 
N.Z. vol. ii. p. 549. Balena antipodum Gray, 1843, in Dieffen- 
bach’s Travels in N.Z. vol. ii. p. 183 (New Zealand), pl. 1, 
lettered B. antipodarum.  Caperea antipodarum Gray, 1866, 
‘Cat. Seals & Whales Brit. Mus. p. 101; Proce. Zool. Soe. 1873, 
p. 143. Balena antipodarum Gray, Dietfenbach, Travels in N.Z. 
vol. i. p. 44; Van Beneden & Gervais, 1880, Ostéogr. Cet. p. 46, 
pl. 3. Hubalena australis Desm., Hector, 1872, Trans. N.Z. Inst. 
vol. v. p. 156; id. 1878,1].c. vol. x. p. 333. Balena hectori Gray, 
1874, Ann. Mag. Nat. Hist. ser. 4, vol. xii. p. 56. Macleayius 
australiensis Gray, 1873, Ann. Mag. Nat. Hist. ser. 4, vol. xi. 
p- 75; id. Proc. Zool. Soc. 1873, p. 130; id. 1874, Trans. N.Z. 
Inst. vol. vi. p. 90, pls. 16, 17. 


The Southern Right Whale, which is confined to the South 
Temperate and Subantarctic Oceans, appears to differ from the 
Nordeaper (4. glacialis) in the number of ribs and other osteo- 
logical features. It was formerly very abundant in the New 
Zealand seas, but has been almost exterminated by whalers. 
During the first half of last century immense numbers were killed, 
and of these usually more than haif were lost (see Hocken, Proc. 
N.Z. Inst. vol. iii. p. 69, 1871). The females visited the bays to 
ealve during the months of May, June, and July, and during this 
period both the cows and calves were killed. Later in the season 
the whaling-vessels cruised in the “ off-shore grounds.” Whaling- 
stations are at present established at Picton and Kaikoura, motor- 
launches being engaged for procuring the whales. Five Right 
Whales were brought into Picton in 1915-16 and one in 1917; 
from the Kaikoura station three were killed in 1917-18. A 


CETACEA OF THE NEW ZEALAND SEAS. 559 


whaling-station was formerly established at Campbell Island. 
Waite states that Right Whales are nearly always seen during 
the southern cruise of the Government vessels. 

Cook Strait.-—(a) Dieffenbach describes and figures (vol. ii. 
pl. 1) a Right Whale, stranded in Jackson’s Bay, Tory Channel, in 
1839. On this figure, Gray in the same work founded Balena 
antipodum ; the plate, however, was lettered Balena antipodarum. 
The tympanic bone later referred to this species (Gray, 1866) 
belonged to Veobalena marginata. Caperea was founded on 
B. antipodarum, and the tympanic bone mentioned. (6) Tory 
Channel. Scapula in Dominion Museum. Described by Gray as 
Balena heciori. Acromion distinct. 

Canterbury.—(a) Skeleton in British Museum sent by von 
Haast. Described by Gray (W/. australiensis). Vertebre 54. 
Cervicals and first dorsal fused into one mass by their centra. 
Sternum oblong, 645 in. Scapula with broad compressed 
acromion. (Gray.) (6) Akaroa Harbour. Skeleton with baleen 
in the Paris Museum. Taken by Berard and Arnoux, of the 
French corvette, ‘Le Rhin.’ Described by Van Beneden and 
Gervais, and by Hector (1878). Vertebre: C. 7, D. 15, L. 11, 
Ca. 21=54. Cervicals fused. Scapula with acromion represented 
only by aridge. Phalanges: 2,5, 6,5, 4. Distal end of first rib 
expanded and two-headed. Sternum triangular, sinuated, sides 
convex, articular facets at front angles. (From van Beneden and 
Gervais’s description and figure.) Wan Beneden in two places 
states the total number of vertebre to be 53, but figures for each 
portion of the vertebral column are also given as above stated. 

Otago.—Skeleton with baleen in the Otago Museum. Described 
by Hector (1878). Young specimen 29 feet long. Vertebre: C.7, 
D.14, L.11, Ca. 22=54. Six cervicals united by their centra. 
Caudal region imperfect. Ribs 14. Scapula with a well- 
developed narrow acromion. (Hector states that the dorsals 
number 15, and the ribs should probably be 15.) 


NEOBALANA, 


Neobalena Gray, 1870, Ann. Mag. Nat. Hist. ser. 4, vol. vi.. 
p- 155; type, Balena marginata Gray. 


NEOBALENA MARGINATA, (PI. I.) 


Belena margimata Gray, 1846, Zool. ‘Erebus’ & ‘Terror,’ 
p- 48, pl. | (West Australia). Balena antipodarum Gray, 1864, 
Proe. Zool. Soc. p. 202 (tympanic bone). Lalena marginata Gray, 
Hector, 1870, Trans. N.Z. Inst. vol. ii. p. 26, pl. 26; id, 1870, 
Ann. Mag. Nat. Hist. ser. 4, vol. v.p. 221. Neobalena marginata 
Gray, 1870, Ann. Mag. Nat. Hist. ser. 4, vol. vi. p. 156; id. 1873, 
l. ec, vol. xi. p. 108; id. 1871, Trans. N.Z. Inst. vol. iii. p. 123; 
id. 1874, 1. c. vol. vi. p. 938. Hecter, 1873, Trans. N.Z. Inst. 
vol. v. p. 155; id, 1875, l.e. vol. vii. p. 271, pls. 16, 17; id. 1878... 
1, ec. vol. x. p. 331. 


560 MR. W. R. B. OLIVER: REVIEW OF THE 


The Pigmy Right Whale has occurred a number of times on 
the New Zealand coast. Elsewhere it has been recorded from 
West Australia and South Australia. Salena marginata was 
founded on three plates of baleen now in the British Museum, 
A skull from Kawau was next recognized as belonging to the 
same species, and was described by Hector (1870, p. 26). This 
was constantly referred to by Hector and others (see Palmer, 
Index Gen. Mamm. p. 452, 1904) as the “‘ type” of the species, 
but this is obviously wrong. The type of Balena marginata is 
the baleen from West Australia in the British Museum. 

Kawauw.—Skull in Dominion Museum, Wellington. Described 
by Hector (1870). Hector afterwards referred to this as the type 
skull. On Hector’s description Gray separated the species from 
Balena under the generic name Neobalens. (Pl. I. figs. 2 & 3.) 

Oharia, Cook Strait.Skeleton in Dominion Museum, Welling- 
ton. Vertebre: C. 7, D.18, L. 2,Ca.14=41. Cervicals fused. 
Ribs 17, of which 2nd to 5th are two-headed. 

Canterbury Coast.—Skeleton in Australian Museum, Sydney. 

Stewart Island.—(a) Skeleton with baleen in British Museum. 
Described by Hector (1875, 1871). In this description the 
number of vertebre and ribs appears to be incorrectly stated. 
‘Compare the description by Beddard (‘ Book of Whales,’ p. 141, 
1900.) (6) Skeleton in Dominion Museum, Wellington. Skull 
symmetrical. Viewed from above, the general form is triangular, 
broadest behind where at the angles the supraoccipital and 
squamosals meet; from here it tapers at first gradually to the 
edge of the orbits, then suddenly to opposite the nasals; thence 
follows a long, tapering, pointed rostrum with slightly concave 
sides. Viewed from the side, the upper contour of the skull is 
nearly straight from condyles to vertex, then depressed at a broad 
angle and slightly bowed to the tip of the rostrum. Pre- 
maxillaries narrow, of nearly uniform width, their inner edges 
meeting as an acute ridge in front, but flattening out near the 
blow-hole and twisting so that the outer edges form ridges each 
side of it. Maxillaries broad behind, suddenly contracting to 
opposite the nasals, tapering to a point anteriorly, not extending 
as far forwardas the premaxillaries. The exposed lateral portions 
of the frontal form two bread quadrangular plates, overlapping 
the posterior portion of the maxillaries, and well below the level 
-of the supraoccipital, slightly concave centrally, but arched over 
the orbits. A narrow strip only of the frontal exposed between 
the supraoccipital and nasals. Wide deep lateral channels 
between upper and lower horizontal expansions of the supra- 
oecipital. On the palatal aspect of the skull the maxillaries form 
a sharp central ridge, between which the vomer is. exposed. 
Pterygoids meeting along the middle line, flat behind but 
anteriorly forming a ridge continuous with that of the maxillaries. 
Rami of mandible massive, compressed, angled above and below ; 
nerve foramina near the upper edge. (Pl. I. fig. 1.) Vertebre: 
‘C. 7, D. 18, L. 1, Ca. 14=40. Cervicals fused into a. single mass 


CETACEA OF THE NEW ZEALAND SEAS, 561 


by their centra and neural arches. Spinous processes forming a 
high acute vertical ridge; posteriorly the sides of the neural 
arches project as laminated plates. Seven nerve foramina on each 
side. Anteriorly there is a large vertical lateral process with 
three projecting knobs in a vertical line and a fourth behind the 
upper one. ‘The first dorsal has a forwardly-directed spinous 
process, short-pointed transverse processes, and bears no ribs. 
Transverse processes In the second dorsal angular, in the third 
and fourth wider and twisted, in the remainder very broad and 
flat, with the anterior sides projecting considerably in advance of 
the centra. Articular facets on the distal ends of the transverse 
processes of the 2nd to 10th dorsals. Spinous processes broad and 
flat, with lamellar metapophyses from the third dorsal backwards, 
Lumbar vertebre similar to last dorsal; the transverse processes 
longer in an antero-posterior than in a transverse direction. The 
first seven caudals bear chevrons. Transverse processes reduced 
to ridges on the fifth, last neural canal evident on the 8th caudal, 
which is the last of the compressed form. In all the vertebre 
behind the cervicals, the epiphyses are marked by distinct sutures. 
Ribs 17, articulated to the 2nd to 18th dorsals. First single- 
headed, broad and truncate distally, 2nd to 5th double-headed, but 
the capitula not reaching the centra. Remainder single-headed, 
with very broad, flat blades, except the last two pairs, which are 
small and narrow. The last twelve ribs are set at nearly a right 
angle to the transyerse processes of the vertebre. Scapula much 
wider than long, flat with a very slight spine; acromion directed 
forwards, flat, moderately broad, and tapering ; coracoid short and 
narrow. Sternum shield- -shaped, notched in front, one articular 
facet on each side for the first rib. 

Locality not known.—(a) Skull of young in Auckland Museum. 
Described by Hector(1875). (6) There is a skeleton or portion of 
one in the Paris Museum sent by Mr. Trail in 1879. It probably 
came from Stewart Island. 


BALZNOPTERA. 


Balenoptera Lacépéde, 1804, Hist. Nat. Cét. pp. xxxvi. 114; 
type, B. acuto-rostrata Lacép. 


To the four recent species constituting this genus, 18 generic 
names have been applied. Of these, eight, were founded by Dr. J. 
HE. Gray, whose prolific writings on the Cetacea resulted in more 
synonyms than valid names, and are the despair of later students. 


BALZNOPTERA MUSCULUS. 


Balena musculus Linné, 1758, Syst. Nat. ed. 10, p. 76 (Scotland) 
(not Balenoptera musculus auct.). Balenoptera sibbaldi Gray, 
1847, Proc, Zool. Soc. 1847, p. 92; Waite, 1912, Rec, Cant. 
Mus. vol. 1. p. 333, pls. 69-71 ; id. 1912, Guide Whales & Dolph. 
N.Z. p..10, pls. 1-3. Balenoptera Peedie Linné, Waite, 1909, 
Subant. ifs, N.Z. vol. ii. p, 550. 


562 MR. W. R. B. OLIVER: REVIEW OF THE 


The Blue Whale, which has a very wide range and is exceedingly 
abundant in the Antarctic Ocean, has only once been recorded 
with certainty on the New Zealand coast. 

Okarito.—Skeleton in the Canterbury Museum. ‘The dead 
whale came ashore near Okarito in February 1908. Described 
by Waite. Vertebe: C. 7, D. 15, L. 14, Ca. 28=64. Cervicals 
free. Ribs 15, 2nd to 5th two-headed. ‘Transverse processes of 
first lumbar vertebree with articular surfaces. Sternum 56 cm. 
long, with five processes; one posterior, long; two lateral, 
triangular ; twoanterior diverging, truncate. Pelvic bones about: 
30 em. in length, posteriorly bent downwards at nearly a right 
angle, each with a small detached bone about 2 cm. long above. 
Phalanges: 0, 7, 8, 8, 5. 

Coal Point, Otago.—A large whale came ashore here in 1873. 
Some plates of baleen only were preserved, and were described by 
Hutton (Trans. N.Z. Inst. vol. vii. p. 266, 1875). They probably 
belonged to this species. 


BALENOPTERA PHYSALUS. 

Balena physalus Linné, Syst. Nat. ed, 10, p. 75 (Kuropean: 
seas) (Balenoptera musculus auct.). Physalus australis Desm., 
Hector, 1875, Trans. N. Z. Inst. vol. vii. p. 257, pl. 18; id. 1878, 
l.c. vol. x. p. 336, Balenoptera australis Desm., von Haast, 
1883, Proc. Zool. Soc. 1883, p. 592. Balenoptera musculus (not 
Linné) Parker, 1885, Trans. N.Z. vol. xvii. p. 3, pl. 6. Steno- 
balena xanthogaster Gray, 1874, Ann. Mag. Nat. Hist. ser. 4, vol. 
xiv. p. 305; ‘Sulphur-bottom,” Hector, 1874, Ann. Mag. Nat. 
Hist. ser. 4, vol. xiv. p. 304. 

This is the common Finner of both northern and southern seas. 
It is occasionally cast ashore on the New Zealand coast. 

Nelson.—Stranded at entrance of Waimea River, 1883. 
Skeleton in Otago Museum.. Described by Parker. Vertebre: 
CO. 7, D. 15,15, Ca. 25=62. Ribs 15, of which 2nd and 3rd are 
two-headed, last rib less than half the length of the one in front. 
Sternum with posterior process longer than the lateral ones. 
Baleen slate-colour with white fibres. 

Cook Strait.—Stranded at Port Underwood, 10th June, 1874. 
Skeleton in Dominion Museum, Wellington. Described by 
Hector. On Hector’s description Gray founded the genus and 
species Stenobalena xanthogaster. Vertebre: C.7, D. 15, L. 15, 
Ca. 25=62. Ribs 15, of which 2nd and 3rd are two-headed. 
Sternum with posterior process shorter than laterals. 

New Brighton Beach.—Stranded 1881. Skeleton described by 
Haast. Vertebre: C.7, D. 15, L. 15, Ca.25=62. Pelvic bone 
13 in. long, 6 in. broad. Sternum with the posterior process. 
shorter than the laterals, (Von Haast.) 


BALANOPTERA BOREALIS. 
Balenoptera borealis Lesson, 1828, Hist. Nat. Cet. p. 342; 
Lillie, 1915, ‘Terra Nova’ Exped., Zool. vol. i. p. 117, pl. 7. 


CETACEA OF THE NEW ZEALAND SEAS. 563 


The Sei Whale does not appear to be common south of the 
Equator. The following are the only known occurrences of the 
species in New Zealand waters: 

Off Bay of Islands.—(a) Exterior described by Lillie. Baleen 
black, with fine greyish-white bristles. (5) A male specimen, 
14 m, long, was brought into the whaling-station at Whanga- 
mumu during my visit on the 5th December, 1911. It had 
been harpooned, the weapon passing right through the body 
and exploding in the water. Colour abeve slate, sides mottled, 
below silvery white; iris dark brown; baleen dark slate, 
upper inner angle streaked longitudinally with lighter, bristles 
white. Dorsal fin faleate. Flipper with lower edge nearly 
straight, upper rounded. Plaits between the fins 60, 7 or 8 
more on each side behind the eye. Plaits begin about 25 cm. 
from the tip of. flipper and anterior edge of fin. Contour 
of back nearly straight, suddenly depressed in front of flukes. 
Hairs on lower jaw in two rows; inside row with 12 on right 
side, 13 on left; outside row 1 on right, 2 on left side; total, 
28 hairs. There are some punctures behind the hairs, and others 
in front of the body-plaits. Measurements: Total length, 14 m.; 
snout to anterior edge of flipper, 3°80 m.; snout to anterior edge 
of dorsal fin, 9°40 m.; height of fin, 40 em.; length of upper 
edge of flipper, 115 cm.; its width, 38 em.; height of body, about 
180 cm. Baleen plates, 11°5 to 12°5 mm. apart; length of 
front plates, 8 cm., of longest 34 em. The mouth was filled with 
small crustaceans. The colours noted above were taken shortly 
after the whale was killed, and are lighter than those usually 
recorded. Lillie makes a similar observation. 

Cook Strait.—During a heavy gale at the end of January 1922, 
the carcase of an adult female of the Sei Whale floated through 
the entrance of Porirua Harbour and stranded opposite Paremata. 
It measured 53 feet in length. It had been dead some time, and 
caused a good deal of consternation among residents of the 
locality. The carcase was subsequently towed to sea and cast 
adrift. 


BALZNOPTERA ACUTOROSTRATA. 


Balenoptera acutorostrata Lacépede, 1804, Hist. Nat. Cét. vol. 1. 
p. 197; Lillie, 1915, ‘Terra Nova’ Exped., Zool. vol. i. pls. 4, 5. 
Balenoptera rostrata Fabricius, von Haast, 1881, Trans. N.Z. 
Inst. vol. xiii. p. 169, pl. 3; Benham, 1901, Proc. Zool. Soc. 
1901, vol. i. p. 278; id. 1902, Trans. N.Z. Inst. vol. xxxiv. p. 151. 
Balenoptera huitoni Gray, 1874, Ann. Mag. Nat. Hist. ser. 4, 
vol. xiii. p. 450; Hector, 1878, Trans. N.Z. Inst. vol. x. p. 337. 
Physalus antarcticus Gray, 1874, Ann. Mag. Nat. Hist. ser. 4, 
vol. xi. p. 316. 

The Pike Whale, like its congeners, is widely distributed All 
the records certainly identified from New Zealand have been 


from the South Island. In addition to the following specimens, 
Proc. Zoo. Soc.—1922, No. XX XIX. é 


564 MR. W. R. B. OLIVER: REVIEW OF THE 


however, a young Pike Whale was recorded as having been killed 
at Port Ahuriri, Hawke’s Bay, in 1885. Variations occur with 
regard to the vertebral formule. Apparently the nor rmal number 
isin, OME Mo sCan i438: 

Stunner , Canterbury.—Stranded 7th February, 1880. Skeleton 
in Canterbury Museum. Described by von Haast. Vertebre: 
C.7, D. 11, L. 12, Ca. 18=48. Ribs 11. Sternum longer than 
broad. Pelvic bones rod-shaped, about 22 em. long. 

Moeraki.—1885. Skull (incomplete) in Otago Museum. 

Otago Heads.—(a) Oct. 1873. Skeleton with baleen in British 
Museum. Described by Gray as the type of Balenoptera 
huttoni. Measurements given by Gray (p. 451). Vertebre: 
C.7, D. 12, L. 13, Ca. 16=48. Ribs1l. (Gray.) (0) Young 
female cast ashore, August 1900. Exterior described by Benham 
(1902). Larynx described by Benham (1901). Animal figured 
by Lillie. 

Locality not stated.—Skeleton in Dominion Museum, Wel- 
lington. Vertebre: C.7, D.11, L.13, Ca.17=48. Cervicals: 
2nd and 3rd fused by neural arch on right side; 4th and 5th 
separately fused by distal ends of upper lateral precesses on right 
side; otherwise free. Atlas with short transverse processes ; 
2nd to 5th with both upper and lower transverse processes ; 6th 
and 7th with upper transverse processes only. The 8th caudal 
bears the last spinous process and the last complete neural canal. 
Ribs 11. Sternum with the length about 4 mm. short of the 
breadth; posterior process long and of nearly even width. 
Scapula with long acromion widening towards the distal end; 
coracoid thick, less than half the length of the acromion. 


MEGAPTERA. 
Megaptera Gray, 1846, Zool. ‘Krebus’ & ‘Terror,’ Mamm. p. 16 ; 
type Balena longimana Rudolphi (= Bb. nodosa Bonnaterre). 


MEGAPTERA NODOSA. 


Balena nodosa Bonnaterre, 1789, Tabl. Kneyl. Méth., Cét. p. 5 
(New England, U.S.A.); Lillie, 1915, ‘Terra Nova’ Exped., 
Zool. vol. i, pp. 87-110, pls. 1-4. Megaptera nove zealandie 
Gray, 1864, Proc. Zool. Soc. 1864, p. 207; id. 1866, Cat. Seals 
& Whales Brit. Mus. p. 128, fig. 20; Hector, 1873, Trans. N.Z. 
Inst. vol. v. p. 1563; id. 1875, 1. c. vol. vii. p. 255; Van Beneden 
& Gervais, 1880, Ostéogr. Cét. p. 135; Gray, 1874, Ann. Mag. 
Nat. Hist. ser. 4, vol. xiil.p. 57. Megaptera lalandii Fischer, von 
Haast, 1883, Trans. N.Z. Inst. vol. xv. p. 214; Hector, 1878, 1. c. 
vol.x. p. 335. Megaptera boops (not Linné) Oliver, 1911, Trans. 
N.Z. Inst. vol. lxiu. p. 536. 

The Humpback is the most common whale frequenting the New 
Zealand seas, but no skeleton of an animal caught within this 
area has been adequately described. A satisfactory description, 
however, has been given by Lillie of the external and internal 


CETACEA OF THE NEW ZEALAND SEAS, 565 


features from specimens brought into the Whangamumu whaling- 
station. 

Humpback Whales regularly migrate, spending the southern 
summer south of New Zealand and the winter in subtropical 
seas. They appear at the Kermadec Islands in the latter part of 
August, but are most common in the group during October and 
November. They have their calves with them at this time. They 
appear off the Bay of Islands in the latter part of September, the 
main body passing southwards during October and November. 
By the middle of December no more are to be seen here. From 
this time until April they appear to be absent from New Zealand 
waters. They are hunted in Cook Strait from May to August, 
and appear off the Bay of Islands between the middle of April 
and the end of August, but are not seen at the Kermadec Islands 
until later in the year, as noted above. (Lillie; Oliver.) 

Kermadec Islands.—Frequent the group in large numbers with 
their calves from middle of August to end of November. 

Chatham Islands.—Baleen in the Canterbury Museum. 

Off Cape Brett.—Regularly shot by harpoon-gun from small 
steam-vessel and brought into the Whangamumu whaling-station. 
The yearly catch varies between 40 and 70. The exterior and 
anatomy have been well described by Lillie. 

Cook Strait—(a) Porirua Harbour. Skull in British Museum. 
Described by Hector (1875). (6) Tory Channel. Two scapulas 
in Dominion Museum, Wellington. Described by Gray (1874). 
(c) Common, May to August. Motor-launches are engaged in 
whaling at Picton, catching 35 to 40 Humpbacks annually. 

Kaikoura.—(a) Skull in Dominion Museum, Wellington. 
Described by Hector (1875). (6) A whaling-staticn is established 
here. Nine whales were brought in in each of the years 1918 
and 1919, 

Akaroa Harbour.—¥Yemale caught 6th May, 1875. Calf also 
killed. Described by von Haast (skeleton then in Canterbury 
Museum). Vertebre: C.7, D. 13, L.10, Ca. 21=51. Cervicals 
free. Sternum scarcely longer than broad. (Haast.) There is 
a skeleton in the British Museum, presumably this one. 

Otago.—Right tympanic and periotic bone in British Museum. 
Deseribed by Gray as the type of Megaptera nove zealandic. 


PHYSETER. 
Physeter Linné, 1758, Syst. Nat. ed. 10, p. 76; type, P. macro- 
cephalus Linné. 


PHYSETER MACROCEPHALUS, 

Physeter macrocephalus Linné, 1758, Syst. Nat. ed. 10, p. 76 
(Huropean seas); Hector, 1878, Trans. N.Z. Inst. vol. x. p. 337; 
Waite, 1909, Subant, Is. N.Z. p. 551. Catodon macrocephalus 
(L.), Hector, 1873, Trans. N.Z. Inst. vol. v. p. 157. Physeter 
catodon (L.), Lillie, 1915, ‘Terra Nova’ Exped., Zool. vol. i, 
p. 118. 


Bue 


566 . MR. W. RB. B. OLIVER: REVIEW OF THE 


It has been the fashion recently to use the name Physeter 
catodon for this species, its only claim being priority of place on 
the page. On the other hand, its identity appears doubtful. 
Gray, Cope, and others applied it to the Arctic White Whale, 
whereas the identity of P. macrocephalus is clear, and the name 
has been in general use since its publication. 

The Sperm Whale is found chiefly in tropical and warm 
temperate seas, though it occurs as far south as the south of 
New Zealand. It does not come close inshore like the Right 
Whale and Humpback, hence it is seldom cast ashore, although, 
strange to say, when this does happen, large schools are sometimes 
involved. It has been recorded off the Chatham and Campbell 
Islands. Lower jaws are preserved in the museums at Christ- 
church and Dunedin. There is also in the British Museum the 
lower jaw of a specimen taken between Cape Howe and New 
Zealand. 

Kaipara Coast.—(a) A school of 27 Sperm Whales—males, 
females, and young—stranded in 1895. (6) School of 25 stranded 
opposite Dargaville, March 1918. Length varied from 22 to 
45 ft. 

Cook Strait—Head cast ashore at Waikanae (Hector, 1873). 

South of New Zealand (Lat. 8. 44° 56’, Long. EH. 172° 53’).— 
School of 12 seen 31st March, 1912 (Lillie). 


Koeta. 
Kogia Gray, 1846, Zool. ‘ Erebus’ & ‘Terror,’ Mamm. p. 22; 
type, Physeter breviceps Blainville. 


KoaiA BREVICEPS. CEE ono.) 


Physeter breviceps Blainville, 1838, Ann. d’Anat. et Phys. vol. ii. 
p. 337 (Cape of Good Hope). Kogia breviceps Blainv., Hector, 
1878, Trans. N.Z. Inst. vol. x. p. 337. Cogia breviceps Blainv., 
Benham, 1901, Proce. Zool. Soc. 1901, vol. i. p. 278; id. 1901, 
PiCmvOl Ww ap. NOM: iG TOO2. Nc. voli. yp. 94) -side O02 .uirarass 
N.Z. Inst. vol. xxxiv. p. 155. Huphysetes pottsti von Haast, 
1874 (June), Trans. N.Z. Inst. vol. vi. p. 97, pl. 15; id. 1874 
(Oct.), Proc. Zool. Soc. 1874, p. 260. 


Though a rare whale, the Pigmy Sperm Whale has been taken 
in all seas. Schulte in 1917 (Bull. Amer. Mus. Nat. Hist. 
vol. xxxvil. p. 361) listed 21 occurrences, three of which were 
from New Zealand. ‘There are four skulls, without locality 
stated (but presumably from Lyall Bay and Petone), in the 
Dominion Museum, Wellington. Altogether at least 11 speci- 
mens have been cast ashore in New Zealand during the past 
40 years. . 

Napier.—1892. Skull in Otago Museum. 

Wanganui Beach.—(a) Skeleton and stuffed skin in the Wan- 
ganui Museum. Vertebre: C.7, D. 12, L. & Ca. 264+ =454. 
Cervicals fused. Capitulum of first rib articulates with centrum 


CETACEA OF THE NEW ZEALAND SEAS. 567 


of last cervical. Dorsal fin small, faleate. (6) On 3rd February, 
1909, I examined at Wanganui a specimen of Kogia breviceps 
which had been stranded at the Heads a few days before. 
Length, 180 cm.; flipper, 27 cm.; dorsal fin height, 9°5 cm. ; 
distance across flukes, 49 em. Black above, white below. The 
accompanying figure was made from a photograph taken at the 
Heads, where, it was reported, three individuals came ashore. 
(CEE EN ile S35) 

Cook Strait._(a) Port Underwood. Mandible in Dominion 
Museum, Wellington. Length of ramus, 32 em.; breadth at 
angle, 7°5 em.; symphysis, 6 em.; width between rami at base, 
28°5 em. Teeth: 14 on right side, 13 on left. (6) Lyall Bay. 
Cow and calf cast ashore, 1880. (c) Petone, Wellington Harbour. 
Cow and calf harpooned, 1884. (¢) Lyall Bay, 1887. Skeleton 
in Dominion Museum, Wellington. Teeth 14.14. Sternum of 
three segments. Scapula with broad backwardly - directed 
acromion and shorter and narrowly-directed coracoid, the borders 
overlapping for the distal half or more. Ribs 12, of which 7 are 
two-headed. Vertebre: C. 7, D. 12, L. & Ca. 20+=39+4+. 

Lyttelton Harbour.—Female stranded at Governor’s Bay, 17th 
August, 1873, Skeleton and half cast in Canterbury Museum. 
Exterior and skeleton described by von Haast as the type of 
Luphysetes pottsii. Teeth 13 in each ramus of mandible. Verte- 
bre: C..7, D. 12, L. 11, Ca. 20=50. Cervicals fused. Ribs 12, 
of which 6 are two-headed. Sternum of three segments, the 
first T-shaped. Phalanges: I, 10, 7, 5, 4. 

Purakanwi, Otago.—Stranded 23rd August, 1900. Skeleton 
in Cambridge University Zoological Museum. Descriptions by 
Benham—exterior and skeleton (1902); exterior and anatomy 
(Proce. Zool. Soc. 1901, p. 107); larynx (Proc. Zool. Soc. 1901, 
p- 278). Teeth 13 in each ramus of mandible, 1 on each side of 
upper jaw. Telvis absent. Vertebre: C. 7, D. 13, L. 9, Ca. 23 
=52. Ribs 13 pairs, the very last small, 6 are two-headed. 
Sternum of 3 segments. Phalanges: R. 2, 10,7, 6,3; L. 2, 9,7, 
6,2. (Benham.) 


BERARDIUS. 


Berardius Duvernoy, 1851, Ann. Sci. Nat. ser. 3, vol. xv. p. 51; 
type. B. arnuati Duvernoy. 

This genus comprises two species—one from New Zealand and 
the Argentine, and the other, B. bairdii, from the North Pacific. 


BERARDIUS ARNUXII. 


Berardius arnuxti Duvernoy, 1851, Ann. Sci. Nat. ser. 3, 
vol. xv. p. 51 (Akaroa, N.Z.); Gray, 1866, Cat. Seals & Whales 
Brit. Mus. p. 348; von Haast, 1870, Ann. Mag. Nat. Hist. ser. 4, 
vol. vi. p. 348; id. 1870, Trans. N.Z. Inst. vol. 11. p. 190; 
Hector, 1870, Ann. Mag. Nat. Hist. ser. 4, vol. v. p. 222; id. 
1873, Trans. N.Z. Inst. vol. v. p. 169; id. 1878, 1. c. vol. x. p 338, 


568 MR. W. R. B. OLIVER: REVIEW OF THE 


pl. 16; Van Beneden & Gervais, 1880, Ostéogr. Cét. pp. 377, 515, 
Atlas, pl. 21 bis, 23, 23 bis; True, 1910, Bull. U.S. Nat. Mus. 
73, p. 68. Berardius arnouwi Duvernoy, Flower, 1872, Trans. 
Zool. Soc. vol. viii. p. 212, pls. 27-29; Beddard, 1900, Book of 
Whales, p. 228. “Ziphid Whale,” Knox, 1871, Trans. N.Z. Inst. 
vol. ili. p. 125; Hector, 1871, ib. p. 129, pls. 16, 17. 


The Porpoise Whale has been recorded a number of times in 
New Zealand and once from La Plata (Marelli, Ann. Mus. Nace. 
Buenos Aires, vol. xxx. p. 411, 1920). The following eleven 
Specimens appear to be all the authentic New Zealand records. 
Besides these, a whale measured by Knox (1871) at Porirua prob- 
ably belonged to this species. 

Chatham Island.—Skull in Otago Museum. 

Coast near Wanganui.—Skeleton in Wanganui Museum. 
Skull—length, 115 em.; mandible, 104 em. Aveoli two in each 
ramus. WVertebre: C.7, D. 10, L. & Ca. 22+=39+. Three 
cervicals fused. Ribs 10, of which 7 are two-headed, 8th attached 
to centrum of 7th dorsal, 9th and 10th attached to transverse 
processes of the corresponding vertebre. Sternum: Ist and 2nd 
segments notched in front and behind, 3rd notched in front only; 
Ath of two lateral pieces, broad in front, united for 6 cm., free 
and obtusely pointed behind. 

Cook Strait.—(a) Worser Bay, Wellington Harbour. Stranded 
January 1870. Skull and other bones in Dominion Museum, 
Wellington. Measurements of animal given by Knox. Skull 
described by Hector (1871, 1873). Length of skull, 120 em. 
Only one pair of alveoli present in mandible. (6) Entrance to 
Wellington Harbour. Captured 12th January, 1877. Skeleton 
in Dominion Museum, Wellington. Exterior and skeleton de- 
scribed by Hector (1878). Teeth two in each ramus of mandible 
near apex. Vertebre: C.7, D.10, L.13, Ca. 17=47. First 
three cervicals fused. Ribs 10, of which 8 are two-headed. 
Sternum of 5 segments. Phalanges: 1, 6, 5, 4, 3. 

Saliwater Creek, north of Banks Peninsula.—Male, stranded 
27th December, 1873. Skeleton in Canterbury Museum. Teeth 
2.2. Vertebre: C. 7, D. 10, L. 13, Ca.17=47. Three cervicals 
fused. Ribs 10, of which 8 are two-headed. Sternum of four 
segments. Scapula with broad expanded acromion and narrow 
thick and but slightly broadening coracoid. Phalanges: 2, 5, 
4, 3, 3. 

New Brighton Beach Stranded 16th December, 1868. Skele- 
ton in Museum of Royal College of Surgeons, London. Exterior 
described by von Haast (1870). Skeleton described by Flower, 
Van Beneden & Gervais, and Beddard. Vertebre: O.7, D. 10, 
L. 12, Ca. 19=48. Three cervieals fused. Ribs 10, of which 7 
are two-headed. Phalanges: 2, 5, 4, 4, 3. 

Akaroa.—{a) 1846. Skull in Paris Museum. Described by 
Duvernoy as the type of Berardius arnuxii. Described and 


CETACEA OF THE NEW ZEALAND SEAS. 569) 


figured by Van Beneden and Gervais (p. 389, pl. 23). (6) Skeleton 
in British Museum. 

Otago Peninsula.—1840. Mandiblein Otago Museum. Teeth 
two in each ramus. 

New Zealand (without locality stated).—(a) Skeleton in Paris 
Museum sent by von Haast. Figured by Van Beneden and 
Gervais (pls. 21 bis, 23 bis). (6) Skull in U.S. National 
Museum, Washington. Measurements given by True. 


MESOPLODON. 


Mesoplodon Gervais, 1850, Ann. Sci. Nat. ser. 3, vol. xiv. p. 16; 
type, Delphinus sowerbensis Blainville. Dioplodon Gervais, 1850, 
Comptes Rendus, vol. xxxi. p. 512; type, Delphinus densirostris 
Blainville. Owlodon von Haast, 1876, Proc. Zool. Soc. 1876, p. 457; 
type, Uesoplodon grayt von Haast. Dolichodon Gray, 1866, Cat. 
Seals & Whales Brit. Mus. p. 353; type, Ziphius layardi Gray. 

For an account of the species of this genus, see Forbes, Proc. 
Zool. Soe. 1893, p. 216, and Flower, 1872, Trans. Zool. Soc. 
vol. vil. p. 211. 


MESOPLODON BowboInt. (PI. 111. figs. 1-3; Pl. IV. figs. 1-3.) 


Mesoplodon bowdoint Andrews, 1908, Bull. Amer. Mus. Nat. 
Hist. vol. xxiv. p. 203, pl. 13 (New Zealand). 


This species is now known from two skeletons :— 

(1) New Brighton Beach—1904. Skeleton in American 
Museum of Natural History, New York. Described by Andrews. 

Maxillary foramina approximately level with foramina in pre- 
maxillaries. Basirostral grooveabsent. Anterior margin of teeth 
posterior to hinder edge of mandibular symphysis. Vertebree : 
C.7, D. 10, L. 9, Ca. 20=46. First three cervicals ankylosed. 
Chevrons 9, Ribs 10, of which 7 are two-headed. Sternum of 
four segments. Scapula with superior margin of acromion over- 
lapping the coracoid border. Phalanges: 0, 4,3,3,2. (Andrews.) 

(2) New Zealand.—Skeleton in Dominion Museum, Wellington. 
This being the second specimen of the species known, a detailed 
description will be given :— 

Skull: Length 72 ecm., of rostrum (from outside anteorbital 
notch) 43°5 em.; breadth of skull at squamosals 33 em., at orbits. 
31:8 cm.; base of rostrum 12 cm. Temporal fosse 9°5 x 6°5 cm. 
Height from vertex to lower border of pterygoids 30-2 cm., of 
rostrum at anterior end of pterygoids 8'2em. (Pl. ILI. figs. 1-3.) 

Maxillary foramina 3 on right side, 2 on left, about | em. in 
advance of the premaxillary foramina (1 on each side). Rostrum 
solid. Contour of upper edge of rostrum straight for two-thirds 
of its length, then slightly depressed ; of lower edge parallel with 
upper for the first third, then sloping upwards for the middle 
third, at 14 em. from apex its height being 5 em., then gradually 
narrowing to apex. A prominent ridge on each side of the 


570 MR. W. R. B. OLIVER: REVIEW OF THE 


expanded portion of the maxille curving in anteriorly, and 
forming the lateral prominences at the base of the rostrum and 
dividing. the anteorbital notches. Proximal ends of premaxille 
strongly everted and overhanging the.nares; both directed 
obliquely backwards, the right 12°5 em., the left 7 em. long. 
Right nasal bone forming the vertex, the left smaller and lower. 
Mesorostral appearing between the premaxille throughout its 
length ; width at rostral base 2°3 em., at 14 em. in advance 1 cm., 
thence widening out towards the distal end. Its proximal end 
is a thin vertieal ridge of bone. Foramen magnum 5x4 cm., 
occipital condyles 7 x 4:2 em. No basirostral groove. Temporal 
fosse pyriform, above the level of the occipital condyles. On the 
lower surface of the skull the pterygoids diverge, showing the 
palatines as narrow divergent stripes on each side, about 10 cm. 
long and lem. wide. Between the palatines the two edges of the 
maxiile meet as a median ridge, the bones anteriorly diverging 
and exposing the vomcr. The palatines appear again above the 
pterygoids, their anterior ends being level with the pterygoid 
fosse. Pterygoids broken, but the exterior deeply concave and 
the basal portion parallel and meeting for a distance of 13 em. 

Mandible: Length of ramus 63 em., of symphysis 15 em.; tip 
to anterior end of left tooth, 13°5 em.; tooth-line along upper edge 
of ramus, 11 cm.; width of ramus at posterior end, 12°5 cm.; 
narrowest portion behind tooth 7 cm., at anterior edge of tooth 
65em. The upper margin curves upwards and slightly outwards 
(but is not swollen) to form the walls of the alveolus, the internal 
wall being much higher than the external wall, which in the 
present specimen falls, but may be broken. (PI. IV. figs. 1 & 2.) 

Tooth: The left tooth only is preserved. Its anterior margin 
at its junction with the upper edge of the ramus is 15 mm. in 
advance of the posterior end of the symphysis. Anteriorly the 
tooth is slightly concave, posteriorly strongly convex. The apex 
is directed slightly forwards. The base is formed of eight 
irregular fangs. Anterior margin, 13:8 cm.; base, about 10°5 
em.; thickness, 2°4 em. 

Hyoid bones: Thyrohyals separate; length, 17 em. Other 
bones not preserved. 

WVertebra: C! 7, D. 1051. 9, Ca. 1T5=41 © Atlas, axis) van 
third cervical fused by both centra and neural arches, 4th and 
5th fused oniy by the neural arch on the right side; 6th and 7th 
free. Atlas without diapophyses; two conical spines on crest. 
Diapophyses and parapophyses all separate. Seventh cervical 
with long diapophyses directed forwards, parapophyses reduced to 
knobs, above which are the articular facets for the tubercula of 
the first pair of ribs. Dorsals with centra concaveabove. Trans- 
verse processes of the anterior seven opposite the neural arches 
and bearing articular facets for the tubercula of the first seven 
ribs. The first six dorsals bear tubercles on the posterior edges 
of the centra for the articulation of the capitula of the 2nd to 7th 
ribs. The 8th, 9th, and 10th dorsals bear short transverse 


CETACEA OF THE NEW ZEALAND SEAS. 571 


processes ending in articular facets for the heads of the last three 
ribs. On the 8th and following vertebre the transverse processes 
and metapophyses are widely separated, the former being opposite 
the centra. Lumbars with strong inferior median carina. 
Neural spines truncated and expanded at their distal ends. 
Metapophyses lamellar, projecting forward and upward. Anterior 
11 caudals showing facets for chevron bones. ‘he last four are 
depressed oblong bones with no processes, but there appear to be 
some missing between the two series. ‘Transverse processes 
reduced to ridges on the 8th caudal, neural spine rudimentary on 
the 11th (the 12th and other caudals presumably missing). 

Ribs, 10 pairs. The first seven are two-headed, articulating by 
both capitula and tubercula. The first articulates with the 
centrum of the 7th cervical and the transverse process of the first 
dorsal, the 7th articulates with the centrum of the 6th and the 
transverse process of the 7th dorsal. The 8th, 9th, and 10th are 
single-headed and articulate with the transverse processes of the 
6th to 10th vertebre. The first two ribs broad and flat, with 
_the proximal ends strongly curved; the remainder with the 
proximal ends below the articular tubercles roughly triangular or 
quadrangular in section, the distal ends being flattened. 

The greatest length of each rib on the right side in a straight 
line is as follows :—Ist, 30°5 em.; 2nd, 46-8 cm.; 3rd, 54:7 cm. ; 
Ath. 59:0) em. ;* 5th, 60:3: cm.; 6th, 60:4 cm.; 7th, 60:0) em:; 
8th, 55°2 cm.; 9th, 50°5 em.; 10th, 42°5 cm. 

Sternum: The sternum consists of four segments. Three 
fenestra are formed at the junctions by notches in the middle of 
the anterior and posterior margins of the segments. There is a 
fourth fenestra in the centre of the posterior segment. On each 
side are five facets for the articulation of the costal ribs—one on 
the anterior segment, one on the posterior, and one at each of the 


junctions. 
(1) (2) (3) (4) 
mm. mm. mm. mm. 
Greatest length of segments ...... 182 116 90 130 
Greatest breadth of segments ...... 195 130 105 102 


Scapula: Breadth, 30-4 em., length (median), 20 em. Median 
length of acromion and coracoid, each about 9 cm. Acromion 
broad, lamellar projecting upward. Coracoid slender, expanded 
distally. The superior margin of the acromion is clear of the 
coracoid border. (PI. IV. fig. 3.) 

Comparison with type of JZ. bowdoint :—The present specimen 
differs from the type of J. bowdoini from New Brighton in no 
important respect. In the general proportions of the skull and 
mandible they appear to be within the range of individual 
variation, and agree in those characters separating J/. bowdoini as 
a species: these are the absence of a basirostral groove, short 
symphysis of mandible, position and size of tooth, relative positions 
of palatines and pterygoids, overhanging maxillary crest, and 


572 MR. W. R. B. OLIVER: REVIEW OF THE 


form of the rostrum. On the other hand, the anterior edge of the 
tooth in the New Brighton specimen is 15 mm. behind the 
posterior margin of the mandibular symphysis; in the Dominion 
Museum specimen it is about 14 mm.in advance of it. According 
to my description, the 8th to 10th dorsal vertebre have the corre- 
sponding ribs articulated to their transverse processes. Andrews 
states that the 8th rib of J. bowdoini is articulated to a facet on 
the posterior margin of the centrum of the 7th dorsal. The 
superior margin of the acromion in the type of M/. bowdons 
overlaps the coracoid border, whereas it is clear of it in the present 
specimen. 


MESOPLODON GRAYI. 


Mesoplodon grayt von Haast, Proc. Zool. Soc. 1876, p. 7 
(Chatham Island); Flower, 1878, Trans. Zool. Soc. vol. x. p. 417, 
pls. 71-73 ; Forbes, 1893, Proc. Zool. Soc. 1893, p. 216, pls. 12-15. 
Mesoplodon hectori (not Gray) Hector, 1874, Trans. N.Z. Inst. 
VOL VA. OO, pla lores vk 1376, lic. viola xm. Sos ap lemlele 
Berardius hectort (not Gray) Hector, 1875, 1. ¢. vol. vii. p. 262. 
Oulodon grayi von Haast, 1876, Proc. Zool. Soc. 1876, p. 457 ; 
id. 1877, Trans. N.Z. Inst. vol. ix. p. 450, pl. 26; Van Beneden 
& Gervais, 1880, Ostéogr. Cét. p. 516, pl. 62. AMesoplodon haasti 
Hlower el eney rans. Zool!Soc vol: xe". SIO plsaaig leis 
Mesoplodon australis Flower, 1878, |. c. p. 419, pls. 71-73; 
Lillie, 1915, ‘ Terra Nova’ Exped., Zool. vol. 1. p. 119. Meso- 
plodon know part (skull B) Hector, 1873, Trans. N.Z. Inst. vol. v. 
[Da LOG 


The New Zealand Scamperdown Whale appears to be fairly 
common in the New Zealand region. Elsewhere it has been 
recorded only from Patagonia. 

Chatham Islands.—(a) About 25 individuals stranded on 
Waitangi Beach, 1875. Three skulls sent to Canterbury Museum. 
One described by von Haast as the type of Mesoplodon grayi. On 
this specimen von Haast founded the genus Owlodon. It has 
also been described and figured by Forbes. A row of 17 small 
conical teeth in each side of upper jaw. The two remaining 
skulls in the Otago Museum. One has 17, the other 19 teeth in 
each side of the upper jaw. (6) Skeleton in U.S. National 
Museum, Washington. Vertebre: C.7, D.11, L.10, Ca. 20=48. 
Two cervicals fused. Chevrons 9. Ribs 11, of which 7 are two- 
headed; the last rib rudimentary, about 70 mm. long. (c) Skull 
and three rostra in British Museum. Described by Forbes. 
(Zz) Skeleton in Bishop Museum, Honolulu. Figured in Oce. 
Papers Bishop Mus. vol. i. no. 5, figs. 4-8, 1902. Ribs 9. 
Premaxillary behind maxillary foramina. 

North. Cape.—Skeleton found in Great Exhibition Bay, August 
1911. Skull and scapula in British Museum. Described by 
lillie. SBasirostral groove present. Foramina of fifth nerve one 
behind the other. Height of teeth, 3 in.; length, 23in. (Lillie.) 


CETACEA OF THE NEW ZEALAND SEAS. 573 


East Coast, North Island.—Male stranded, December 1875. 
Rostrum in Museum of Royal College of Surgeons, London. 
Described by Flower as the type of M. haasti. Measurements 
given by Forbes. Basirostral groove present. Foramina of fifth 
nerve one behind the other. 

Cook Strait.—(a) Lyall Bay. Cast ashore, January 1875. 
Skeleton in British Museum, Exterior and skeleton described by 
Hector (1875). Skeleton described by Flower as the type of 
M. australis. Skull measurements given by Forbes. Basirostral 
groove deep. Vertebre: C. 7, D. 9, L. 11, Ca. 20=47. Two 
cervicals united. Ribs 9, of which 7 are two-headed. (Flower.) 
(6) Wellington. Skeleton in American Museum of Natural 
History, New York. 

Kaikoura.—Mandible described and figured by Hector (1874). 

Saltwater Oreek, 30 miles north of Banks Peninsula.—Male 
stranded 15th December, 1876. ‘Two males and a female stranded 
29th December, 1876. Skeleton of one in Canterbury Museum. 
Skull measurements given by Forbes. Basirostral groove present. 
Vertebre: C. 7, D. 10, L. 11, Ca. 19=47. Two cervicals fused. 
The 8th and 9th caudals are joined by a bony growth. Ribs 10, 
of which 7 are two-headed. Sternum of 4 segments; Ist large, 
bifid in front, 2nd and 3rd notched in front and behind, 4th 
composed of three fused pieces, pointed posteriorly and with a 
central foramina. Phalanges: 1, 6, 6, 4, 3. Skeleton of another 
in Museum of Royal College of Surgeons, London. Deseribed by 
Flower and Forbes. Basirostral groove shallow. Vertebre: 
C. 7, D: 10, L. 11, Ca: 20=48. ‘Iwo cervicals fused. Ribs 10; 
of which 7 are two-headed. (Flower.) 

Kaiapot.—Skull in Canterbury Museum. Described by Hector 
(1873) and Forbes. 

New Zealand (locality not stated).—(a) Skullin Otago Museum. 
Figured by Hector (1878). (6) Skeleton in Paris Museum. 
Described by Van Beneden and Gervais. Skull measurements 
given by Forbes. Basirostral groove present. Foramina of fifth 
nerve one behind the other. 18 teeth in upper jaw on left, 17 on 
right side. Sternum of 5 segments. Vertebre: C. 7, D. 10, 
C. 11, Ca. 19=47. Two cervicals fused. Chevrons 10. Rubs 10, 
of which 7 aretwo-headed. Phalanges: 0,6,5,4,2. (c) Skeleton 
in Dominion Museum, Wellington. Rostrum narrow, deeper 
than wide at base. Basirostral groove deep. Foramina of fitth 
nerve one behind the other. Maxillary large, directed forwards; 
premaxillary small, behind and directed upwards. Length of 
skull 75 em., of rostrum 45 em,; width at squamosals, 28 cm. ; 
base of rostrum, 5cm. Mandible length, 67 cm.; symphysis, 
21:5 cm. This skull approaches closely the type of J/. australis. 
Ribs 10, of which 7 are two-headed. Sternum of 4 segments ; 
the first two free and notched in front and behind, the last two 
completely fused, the line of junction being denoted by a 
foramen. 


5A MR. W. R. B. OLIVER: REVIEW OF THE 


MeEsoPLODON LAYARDI. 


Ziphius layardi Gray, 1865, Proc. Zool. Soc. 1885, p. 358 (Cape 
of Good Hope). Dolichodon layardi Gray, Hector, 1873, Trans. 
N.Z. Inst. vol. v. p. 166, pl. 3. dlesoplodon layardi Gray, Hector, 
1873, Ann, Mag. Nat. Hist. ser. 4, vol. x1. p. 106; id. 1878, 
Trans. N.Z. Inst. vol. x. p.341; Flower, 1878, Trans. Zool. Soc. 
vol. x. p. 416; Turner, 1880, ‘Challenger’ Rep., Zool. vol. i. 
pt. 4, p. 2, pls. 1-3; Waite, 1912, Rec. Cant. Mus. vol. 1. p. 326, - 
pl. 62; id. 1912, Guide Whales & Dolph. N.Z. p. 16, pl. 4. 
Mesoplodon flowert von Haast, 1876, Proc. Zool. Soe. 1876, p. 478 ; 
id. 1877, Trans. N.Z. Inst. vol. ix. p. 442, pls. 25,26. Dolichodon 
traversu Gray, 1874, Trans. N.Z. Inst. vol. vi. p. 99. 


The Strap-toothed Whale has been recorded from South Africa, 
Australia, and New Zealand. 

Chatham Island.—Mandible described by Hector (1873) and 
‘Turner. 

— Great Barrier Island.—Skull in Canterbury Museum. ‘Teeth 
meeting over the upper jaw. 

Napier.—Teeth in Otago Museum. 

— Marlborough.—Skull in Dominion Museum, Wellington; 
acquired 1879. Length of teeth following curvature, 34 cm. ; 
greatest width, 60 mm. Denticle 7 mm. high, 10 mm. along 
base-line. 

Saltwater Creek, north of Banks Peninsula.—Stranded 16th 
March, 1874. Skeleton in Canterbury Museum. Described by 
von Haast as the type of MU. flowerr. Vertebree: C. 7, D. 10, 
L. 10, Ca. 19=46. Three cervicals fused. Ribs 10, of which 7 
are two-headed. Sternum of 4 segments. Scapula with broad 
acromion and narrow coracoid. Phalanges: 2, 5, 5, 4, 3. 

Lyttelton Harbour.—Specimen stranded at Allendale, 22nd 
March, 1912. Half cast in Canterbury Museum. Exterior 
described by Waite. 

Locality not stated.—(a) Skull in Dominion Museum, Wel- 
lington. Length, 94°5 em.; breadth, 41 em.; mandible length, 
80 cm.; symphysis, 22 em. Basirostral groove deep. (Teeth 
missing.) (b) Skelton in Dominion Museum. Basirostral groove 
shallow. Length of skull, 61 cm.; of mandible, 52 em. Verte- 
bre: C.7, D. 9, L. 12, Ca. 16=44. Two cervicals fused. Ribs 9, 
of which 7 are two-headed. Sternum of 3 segments. I have 
identified this specimen from cranial characters. It is from a 
young animal, and the teeth are small and triangular. They only 
differ from those of M. grayi in being strongly incurved. 


PAIKEA, gen. n. 


I propose this name for those species of Ziphioid Whales with 
the cranial characters of Mesoplodon and two terminal teeth in 
the lower jaw. Type, Berardius hectori Gray. The only other 
species that can be referred to this genus is J/esoplodon mirum. 


CETACEA OF THE NEW ZEALAND SEAS. 575» 


True, from the North Atlantic. The essential character is the 
position of the teeth. Other characters which these species have 
in common are the broad and deep rostrum and the long mandi- 
bular symphysis; also, the dorsal fin is placed far back and the 
teeth completely covered by the gums. In Mesoplodon, as it is 
now proposed to restrict it, the teeth are always placed near the 
posterior end of the symphysis so that the two species comprising 
Paikea apparently form a natural group in which the teeth, as 
pointed out by Dr. Harmer (Report Cetacea on British Coasts 
1918, p. 21, 1919), possibly correspond with the anterior pair of 
Berardius. 

The reduction of the teeth from four to two in the Ziphioid 
Whales is interesting. Birardius is perhaps the most primitive 
of the recent genera and has usually four teeth, but in a skull] in 
the Dominion Museum, Wellington, only the anterior pair is. 
present. In Ziphius and Hyperoodon it is the anterior pair that 
is retained, but in Mesoplodon the posterior pair only is present. 
Paikea differs from Mesoplodon in possessing only the anterior 
pair of mandibular teeth. 


PAIKEA HECTORI. 


Berardius hectori Gray, 1871, Ann. Mag. Nat. Hist. ser. 4, 
vol. vil. p. 117 ; Hector, 1873, Ann. Mag. Nat. Hist. ser. 4, vol. xi. 
p- 106. Berardius arnuxit (not Duvernoy) Hector, 1870, Trans. 
N.Z. Inst. vol. i. p. 27. “ Ziphid Whale,” Knox & Hector, 
1871, 1. c. vol. ii. p. 125, pls. 14,15. Mesoplodon knoxi Hector, 
1873, 1. c. vol. v. p. 167. Mesoplodon hectori Gray, Flower, 1878, 
Trans. Zool. Soc. vol. x. p. 416, pls. 71, 72; Forbes, 1893, Proc. 
Zool. Soc, 1893, p. 227. 


Known only from the following specimen :— 

Titaht Bay, Cook Strait.—Cast ashore January 1866. Skull’ 
in British Museum. All the references quoted refer to this. 
specimen. Basirostral groove absent. Foramina of fifth nerve 
level. 


ZIPHIUS. 


Ziphius G. Cuvier, 1823, Ossem. Foss. vol. v. p. 350, pl. ; type, 
4. cavirostris Cuv. Hypodon Haldeman, 1841, Proc. Acad. Nat. 
Sei. Philad. vol.i.p. 127; type, Delphinus desmaresti Risso (= H. 
cavirostris). 

If Aphius and Xiphias be considered variations of the same 
word, then Ziphius Cuvier, 1823, is preoccupied by Xiphias Linné, 
1758, a genus of fishes, and the next valid name to be used for the 
genus of whales is Hypodon Haldeman, 1841. Professor David 
Starr Jordan points out to me that Zphius seems to have 
been purposely so written by Cuvier perhaps to make it clearly 
different from Xtphias, which name was also used by the same 
author. 


576 MR. W. R. B. OLIVER: REVIEW OF THE 


AIPHIUS CAVIROSTRIS. 

Ziphius cavirostris G. Cuvier, 1823, Ossem. Foss. vol. v. p. 350 
(Mediterranean coast of France); Flower, Proc. Zool. Soc. 1876, 
. 4/7; Hector, 1878, Trans. N.Z. Inst. vol. x. p. 342; Turner, 
1880, ‘Challenger’ Rep., Zool. vol. 1. pt. 4, p. 27, pl. 2. Hpiodon 
chathamiensis Hector, 1873, Ann. Mag. Nat. Hist. ser. 4, vol. xi. 
p- 105; id. 1873, Trans. N.Z. Inst. vol. v. 164, pls. 4,5. Epiodon 
nove zealandie von Haast, 1877, Trans. N.Z. Inst. vol. ix. p. 430, 
pls. 24, 26. Ziphius nove zealandie von Haast, 1876, Proc. Zool. 
‘Soc. 1876, p. 466; id. 1880, Proc. Zool. Soc. 1880, p. 232; id. 
1880, Trans. N.Z. Inst. vol. xii. p. 241, pl. 8. Aiphius sp., 
Scott & Parker, 1889, Trans. Zool. Soc. vol. xii. p. 247, pls. 48-50; 
Beddard, 1900, Book of Whales, p. 235, pl. 12; Benham, 1901, 
Proc. Zool. Soc. 1901, vol. 1. p. 294. Hyperoodon planifrons (not 
Flower) Waite, 1913, Rec. Cant. Mus. vol. ii. p. 23, pls. 8, 9. 


The Goose-beaked Whale has been recorded from all seas. 

Chatham Island.—(a) Skull in Dominion Museum, Wellington. 
Described by Hector as the type of Hpiodon chathamuensis. (6) 
‘Teeth in British Museum. One figured by Hector, Trans. N.Z. 
Inst. vol. v. pl. v. figs. 2a, 2 6. 

East Cape, North Island.—Teeth and extremity of mandible 
in Canterbury Museum. Described and figured by Waite, who 
referred it with doubt to Hyderoodon planifrons, but his figure 
almost matches the anterior end of the mandible of the Petone 
specimen. | 

Cook Strait.—(a) Near Wellington. Skull described by Turner. 
(6) Petone Beach, Wellington Harbour. Stranded October 1919. 
Skeleton in possession of Mr. H. H. Travers, Wellington. 
Vertebre: C. 7, D. 9, L.-11, Ca. 19=46. First four cervicals 
fused, 5th and 6th separately fused by neural arch on left side. 
Last spinous process and neural canal on the 11th caudal. Ribs 9, 
of which 6 are two-headed. Sternum of 5 segments, the first 
4 are notched in front and behind, the last of two separate 
lateral pieces. Scapula with long acromion and coracoid. 

Kaiapot Beach.—¥emale, stranded 15th May, 1879. Skeleton 
in British Museum, Teeth described by von Haast (1880). 

New Brighton Beach.—Stranded 17th November, 1878. Ex- 
terior and teeth described by von Haast (1880). 

Lyttelton Harbour.—Stranded at Port Cooper, July 1872. 
Skeleton in Canterbury Museum. Described by von Haast as 
the typeof Ziphius nove zealandie. Vertebree: C.7, D.9, L. 11, 
Ca. 19=46. Four cervicals fused. Ribs 9, of which 7 are two- 
headed. Sternum of 5 segments, each notched anteriorly and 
posteriorly. Scapula with long, narrow acromion and coracoid. 
Phalanges: 2, 5, 5, 4, 3. 

Akaroa Harbour.—Female, stranded July 1873. Skull de- 
seribed by von Haast (1877). 

Warrington, Otago.—Came ashore alive, 6th November, 1884. 
Skeleton in Otago Museum. External characters, skeleton, and 


CETACEA OF THE NEW ZEALAND SEAS. RCL 


anatomy described by Scott and Parker; larynx described by 
Benham. Two sharp-pointed teeth at apex of mandible. Verte- 
bre: ©. 7, D. 10, L. 9. Ca. 20=46. Four cervicals fused. 
Ribs 10, of which 7 are two-headed. Sternum of 5 segments, 
each notched in front and behind. Pelvic bones about 9 cm. long. 
Phalanges: 1, 5, 5, 95, 2. 

Locality not stated.—Skull of young in British Museum. 


ORCINUS. 

Oréinus Fitzinger, 1860, Wiss.-populire Naturg. Saiug. vol. vi. 
p. 204; type, O. orca (= Delphinus orca Linné). (Orca Gray, 
1846, is preoccupied by Orca Wagler, 1830, a genus of Physe- 
teride.) 


ORCINUS ORCA. 


Delphinus orca Linné, 1758, Syst. Nat. ed. 10, p. 77 (European 
seas). Orca gladiator Gray, 1846, Zool. ‘Erebus’ & ‘Terror,’ 
p. 33; Hector, 1885, Trans. N.Z. Inst. vol. xvii. p. 208. Orca 
pacifica Gray, Hector, 1875, |. c. vol. vil. p. 260. 


The Killer Whale is found in all seas. Lillie has recorded it 
as far south as McMurdo Sound, and states that it is the 
commonest cetacean in the Ross Sea. It is occasionally seen off 
the New Zealand coast, especially about Cook Strait. Besides 
the following definite records, there is a skull of unknown locality 
in the Auckland Museum. 

Chatham Island.—Skull in Canterbury Museum. Teeth +4. _ 

Coast south of Wanganui.—Skeleton in Dominion Museum, 
Wellington. Vertebre: C.7, D. 11, L. 10, Ca. 24 =52. Four 
cervicals fused; 5th and 6th separately fused by neural arches 
only. Teeth 3. Ribs 11, of which 6 are two-headed. Sternum 
of one piece with five articular facets on each side. 

Otago Heads.—Skull in Otago Museum. Described by Hector 
(1875). Teeth 13. ‘ 


PSEUDORGA. 


Pseudorca Reinhardt, 1862, Overs. K. Danske Vidensk. Selsk. 
Forhandl. p, 151; type, P. crassidens (=Phocena crassidens 
Owen). 


PsEUDORCA CRASSIDENS. 


Pseudorca crassidens Owen, 1846, Brit. Foss. Mamm. p. 516 
(North Sea). Pseudorca meridionalis Flower, Hector, 1873, Trans. 
N.Z. Inst. vol. v. p. 163. 


The range of this species, which may be called the Tasmanian 
Blackfish, extends from the North Sea to the Chatham Islands. 
It is met with in large schools in New Zealand and Tasmanian 
waters. 


578 MR. W. R. B. OLIVER; REVIEW OF THE 


Chatham Island.—In March 1906 a school of several hundreds 
was stranded on Long Beach, Petre Bay. These were referred to 
by Waite (Subant. Is. N.Z. p. 551, under Globiocephalus melas), 
who visited the locality in February 1907. In December 190$ 
the remains of many of them were still to be seen, and I was able 
to identify the species as Pseudorca crassidens. ‘Teeth in upper 
jaw 8 on each side. 

Lyall Bay.—Cranium in Dominion Museum, Wellington. 
Described by Hector. This was at one time referred by Hector 
to Beluga (Ann. Mag. Nat. Hist. ser. 4, vol. ix. p. 438, 1872). 


GLOBICEPHALA. 


Globicephala Lesson, 1828, Hist. Nat. Mamm. et Ois. vol. i. 
p. 441; type, Delphinus deductor Scoresby (=D. melas Traill). 


GLOBICEPHALA MELENA, 


Delphinus melas Traill, 1809, Nicholson’s Journ, vol. xxii. p. 21 
(North Atlantic). Globiocephalus macrorhynchus Gray, Hector, 
1870, Trans, N:Z. Inst. vol, ii. p. 28; id. 1873, l.c. vol. v. p. 164; 
Hel, WeSi7s, os wWwolls vam, jos ASIN) ale IOAN Ave saiohy IMS7 A yillb Gs wolls tx. 
p. 481, pls. 18, 134; id. 1870, Ann. Mag. Nat. Hist. ser. 4, vol. v. 
p- 222. Globicephalus melas Traill, Hector, 1885, Trans. N.Z. 
Inst. vol. xvii. p. 209; Waite, 1909, Subant. Is. N.Z. p. 551. 


The Blackfish is found in all tropical and temperate seas. Like 
its relative the Tasmanian Blackfish, it goes about in schools. It 
is occasionally reported stranded in considerable numbers, but as 
the term “blackfish” is popularly applied to several species, the 
specific identity is not always certain. Skeletons and skulls are 
common in New Zealand museums. 

Chatham Islands—Skull in Dominion Museum, Wellington 
(according to Hector, Ann. Mag. Nat. Hist. ser. 4, vol. x1. p. 105, 
1873). 

Cook Strait.—(a) Two skulls and other bones in the Dominion 
Museum. Described by Hector (1870, p. 28; 1873). One of the 
skulls is very massive, measuring 67°5 cm. in length and 47°5 cm. 
in breadth. Rostrum: length, 34 cm.; breadth at base, 26:5 cm. 
Pterygoids approximating to within 3 or 4mm. Teeth 2. 
Intermaxillaries broad, expanding a little in front, approaching 
to within 2 cm. of the edge of the maxillaries, upper surface 
rugose anteriorly. Six cervical vertebrae fused. (6) Lyall Bay. 
School ran ashore, January 1876. Ten skeletons were secured. 
One now mounted in Dominion Museum. Exterior of animal 
and skeleton described by Hector (1877). Teeth 1°. Vertebree: 
C27. D. 11) kL. 13, Ca, 29-60. > Six cervicals fused?” Ribs Msror 
which 7 are two-headed. Phalanges: 4,13,10,3,1. Two skeletons 
in the British Museum are perhaps from this school. (c¢) Skeleton 
in Otago Museum. Teeth =. Vertebre: C. 7, D. 11, L. 12, 
Ca. 26=56. Four cervicals fused. Ribs 11, of which 6 are 


CETACEA OF THE NEW ZEALAND SEAS, SSF 


two-headed. Sternum of 3 segments, the first two perforated, 
the last divided longitudinally. » 

Otago.—Skeleton in Canterbury Museum. Pterygoids sepa- 
rated along the middle line of about lem. Teeth 19. Ver- 
tebre: C.7, D. 1], L. 14, Ca. 24 = 56. Six cervicals. fused. 
Pelvic bones curved, rod-like, about 15 cm. long. Ribs 11, of 
which 6 are two-headed. Sternum of 3 segments, the anterior 
one notched in front, perforated in the centre, and with the sides 
projecting outwards and backwards. Scapula with broad L-shaped 
acromion, coracoid with distal end expanded. 

Stewart Island.—(a) Skull in Otago Museum. (6) Captured 
January 1874. Skeleton in British Museum, exterior described 
by Hector (1875). 

Campbell. Island.—(Filhol, Mission d’Ile Campbell, Zool. p. 34, 
1885, G. macrorhynchus). 

Locality not stated.—(a) Skull in Auckland Museum. Figured 
by Hector (1875, pl. 16). (6) Skeleton in Dominion Museum. 
Teeth §. .Vertebre: C. 7, D. 11, L. 13, Ca. 25=56. Five cer- 
vicals fused. . Ribs 11, of which 6 are two-headed. Sternum of 
3 segments. 


GRAMPUS. 


Grampus (§ Delphinus) Gray, 1828, Spicilegia Zoologica, vol. i. 
p: 2; type, Delphinus griseus Cuvier. 


GRAMPUS GRISEUS. 


Delphinus griseus Cuvier, 1812, Ann. Mus. vol. xix. p. 14 
(Brest, France). Grampus richardsont Gray, Hector, 1873, Trans. 
N.Z. Inst. vol. v. p. 163. Grampus griseus Cuv., Waite, 1912, 
Rec. Cant. Mus. vol. i. p. 328, pl. 63; id. 1912, Guide Whales & 
Dolph. N.Z. p. 19, pl. 5. 


Risso’s Dolphin has been recorded in such widely sundered 
regions as the North Sea and the coast of New Zealand. It isa 
rare species wherever found. Only twice has it been recorded 
from our seas. 

(1) Manawatu Beach.—Mandible in Dominion Museum, Wel- 
lington. Described by Hector. Length of ramus 44°5 em., of 
symphysis 6 cm. ‘Teeth 3 on each side opposite the symphysis, 
blunt, conical, the posterior one 8 mm. in diameter and project- 
ing 8 mm. above the dried gum. 

(2) Pelorus Sound.—Figured by Waite. It is strange that 
the famous cetacean known as ‘ Pelorus Jack” should not have. 
been mentioned in scientific literature until Waite in a short 
note confirmed the identification of the Rev. D. C. Bates, and 
published a photograph showing the contour of the dorsal fin 
and flukes. It has an extensive literature in newspapers and 
magazines. A special booklet by J. Cowan has been devoted to 
it, while an article on ‘‘ Pelorus Jack—Tuni-Rangi,” by W. T. 
Downes, appears in the Journal of the Polynesian Society, 
vol. xxiii. p. 176, 1914. Both these give the Maori legend. 

Proc. Zoo. Soc.—1922, No. XL. 40 


580 MR. W. R. B. OLIVER. REVIEW OF THE 


“‘ Pelorus Jack” has been described as “ bluish-white tinged 
with yellow and purple, and with irregular brown-edged seratch- 
like lines in all directions. His flippers are blackish and mottled 
with grey” (Cowan). Length about 14 ft. Dorsal fin high and 
faleate. This famous cetacean for many years frequented the 
waters outside the entrance to Pelorus Sound, and was in the 
habit of meeting steamers on the route between Nelson and 
Wellington. It accompanied the vessels for some miles, and 
played from side to side of the bows, keeping just in front 
and frequently leaping out of the water. It was protected by 
an Order in Council. ‘‘Pelorus Jack” has not been seen since 
oie 


CEPHALORHYNCHUS. 


Cephalorhynchus (§ Delphinus) Gray, 1846, Zool, ‘Erebus’ & 
‘Terror,’ Mamm. p. 36; type, D. heavisidi Gray. 


‘CEPHALORHYNCHUS HECTORI. (PI. II. figs. 1 & 2.) 


Electra hectori Van Beneden, 1881, Bull. Roy. Acad. Belg. 
ser. 3, vol. i. p. 887, pl. 2. Lagenorhynchus clangulus (not Gray) 
Hector, 1870, Trans. N.Z. Inst. vol. ii. p. 27; id. 1872, Ann. 
Mag. Nat. Hist. ser. 4, vol. ix. p. 436, fig. Hlectra clangula (not 
Gray) Hector, 1873, Trans. N.Z. Inst. vol. v. p. 160, pl. 12; 1d. 
1877, l. c. vol. ix. p. 350, pl. 11. Cephalorhynchus hectora Ben., 
Hector, 1885, Trans. N.Z. Inst. vol. xvii. p. 209; True, 1889, 
Bull. U.S. Nat. Mus. xxxvi. pp. 112, 177, pl. 32. Cephalorhyn- 
chus albifrons True, 1899, 1. c. pp. 111, 177, pl. 32. 


Confined to New Zealand seas. It is the Common White-nosed 
Porpoise, never seen far from the coast. 

North Coast.— Specimen captured. Skeleton in Paris Museum. 
Described by Van Beneden as the type of Hlectra hectori. Teeth 30. 
Vertebre: OC. 7, D. 14, L. 15, Ca. 27=63. Two cervicals fused. 
Ribs 14. 

Bay of Islands—Skull in Dominion Museum, Teeth 30. 
Length of skull 30-5 em., of rostrum 14:5 em.; width of rostrum 
at base, 7°5 cm. 

Wanganwi Coast.—(a) Skeleton in WanganuiMuseum, Sternum 
of two segments; four pairs of ribs articulate with the anterior 
and one pair with the posterior segment, which is perforated. 
Two cervicals fused. (6) Stuffed skin in Wanganui Museum. 
Specimen stranded alive, Castlecliff Beach, May 1921. Mr. G. 
Shepherd, Curator of the Wanganui Museum, has kindly supplied 
me with a description which enables me to give the following 
particulars, Forehead grey. Snout and a margin round the 
grey of the forehead, black. Dorsal surface slaty black, shading 
into slaty grey on the sides and then into black bordering the 
white under surface. Tip of lower jaw, flippers, dorsal fin, and 
tail black. A white spot in the axil of each flipper. Under sur- 
face white, interrupted by an isthmus of black connecting the 
two flippers. The white extends backwards to midway between 


CETACEA OF THE NEW ZEALAND SEAS. 581 


the vent and the flukes, dividing into three equally long pointed 
(PI. II. figs. 1 & 2) bands, one central and one extending obliquely 
along each flank. An elongated dusky patch surrounds the vent. 
Total length of specimen 1-90 m. An interesting point to observe 
is that, according to Mr. Shepherd, the forehead is grey and not 
white. This intermediate colour probably accounts for the vary- 
ing deseriptions given to this animal, resulting in it having been 
listed under two names. 

Oook Strait.—(a) Outside Wellington Harbour. Described by 
Hector (1870). Vertebre: C.7, D. 14, L. & Ca. 48=69. (6) Off 
-Cape Campbell. Two shot from a large school. Exterior de- 
seribed by Hector (1872, 1873). (c) Wellington Harbour. Skull 
in Dominion Museum. ‘Teeth 34. Length of skull 31:2 cm., of 
rostrum 16 em. ; width of skull at squamosals 16 em., of rostrum 
at base 8 cm. (d@) Half model in Otago Museum. There is 
also a skeleton, perhaps of this specimen, but locality not 
given. 

Canterbury.—(a) Skeleton from Banks Peninsula in Canter- 
bury Museum. Vertebre: C.7, D.13, L.15, Ca. 29=64. Two 
cervicals fused. Ribs 13, of which 4 are two-headed, last rib 
less than half the length of the 12th. Sternum of one piece. 
(6) Skullin U.S, National Museum, Washington. 


LAGENORHYNCHUS. 


Lagenorhynchus Gray, 1846, Zool. ‘ Krebus’ & ‘Terror,’ Mamm, 
p. 30; type, Delphinus acutus Gray. 


LAGENORHYNCHUS OBSCURUS. 

Delphinus obscurus Gray, 1828, Spic. Zool. p. 2 (Cape of Good 
Hope). Clymenia obscura Gray, Hector, 1873, Trans. N.Z. Inst. 
vol. v. p. 160, pl. 1; id. 1877, l.c. molec pl li. ade 1TSoarlec. 
vol. xvii. p. 211; Hutton, 1877, l.c. vol, ix. p. 349. 


The Dusky Dolphin is common round the coast. It isapparently 
widely distributed, being recorded from the Cape seas and in the 
Southern Ocean as far south as 58° 8. lat. (Lillie.) 

Wanganui Coast.—(a) Skull in Dominion Museum, Wellington. 
Described by Hector (1873). Teeth = Length of skull, 
35:5 em.; breadth at squamosals, 16-7 cm.; rostrum length, 
19°3 cm.; breadth at base, 85 em. (6) Two specimens in 
Wanganui Museum. 

Cook Strait-—Skeleton in Dominion Museum, Teeth 2?. 
Vertebre: CO. 7, D. 13, L. 19, Ca. 32=71. Two cervicals fused. 
Ribs 13, of which 6 are two-headed. Phalanges: 2, 9, 6, 2, 1. 
Sternum of 3 segments. 

Banks Peninsula.—(a) Lyttelton Harbour. Skull in Canter- 
bury Museum. (0) Akaroa. Cast in Canterbury Museum, 

There are two other skeletons of unknown locality in the 
Dominion Museum. 


40* 


582 MR. W. R. B. OLIVER: REVIEW OF THE 


LISSODELPHIS. 


Lissodelphis Gloger, 1841, Hand- u. Hilfsb. Naturg. vol. 1. 
pp. xxxiv, 169; type, Delphinus peronii Lacépéde. (Zursio 
Wagler, 1830, is preoccupied by Z'ursio Fleming, 1822, a genus. 
of Physeteride.) 


LissoDELPHIS PERONII. 


Delphinus peronii Lacépéde, 1804, Hist. Nat. Cét. p. 316 (south 
of Tasmania). Zwrsio perontt Lacép., Lillie, 1915, ‘Terra Nova” 
Exped., Zool. vol. i. p. 121. 


This species is included in the New Zealand fauna on the 
authority of Lillie, who records seeing two specimens in lat. 8. 
47° 04', long. E. 171° 33' (south-east of Nugget Point, Otago). 
The species is easily recognised on account of its peculiar 
coloration. It has been recorded in the Southern Ocean from 
Tasmania to Chile. 


STENELLA. 


Stenella ($ Steno) Gray, 1866, Proc. Zool. Soc. 1866, p. 213; 
type, Steno attenuatus Gray.  Prodelphinus Gervais, 1880, 
Ostéogr. Cét. p. 604; type, P. marginatus Duvernoy (=P. euphro- 
syne Gray). 


I have adopted Stenella as the earliest valid name for this 
genus, passing over Sousa, proposed earlier on the same page, 
because it is probably only a variation of Susw (preoccupied by 
Lesson, 1828). It may also be added that Susu or Sousou is a 
Hindu word, and therefore the type of Sowsa, if not hitherto- 
fixed, should be the Indian species S. lentiginosus. 


STENELLA EUPHROSYNE. 


Delphinus ewphrosyne Gray, 1846, Zoo]. ‘ Krebus’ & ‘ Terror,’ 
p-. 40, pl. 22. Clymenia nove zealandie (not Quoy & Gaimard) 
Hector, 1873, Trans. N.Z. Inst. vol. v. p. 159, pl. 2; id. 1877, 
le Ga \wolls ab:<5 Falls JkIe 


A large species, recorded from the North and South Atlantic. 
Its claim to be included in the New Zealand fauna rests on the 
following skull :— 

Waitkanae Coast.—Skull in Dominion Museum, Wellington. 
Described and figured by Hector. The proportions of this skull 
and the position of the nerve foramina agree with the type of 
D. euphrosyne as figured by Gray (Zool. ‘ Erebus’ & ‘Terror,’ 
pl. 22). Teeth 7 Length of skull 48 cm., of rostrum 28:4 em. ; 
breadth of skull at squamosals, 25:3 em.; breadth of rostrum at 
base, 12 cm. Pterygoids approximate to within 2 mm. 


CETACEA OF THE NEW ZEALAND SHAS. 583 


STENELLA PSEUDODELPHIS. 


Delphinus pseudodelphis Schlegel, 1841, Abhandl. Gebiete Zool. 
Heft 1, p. 22. Clymenia attenuata Gray, Hector, 1885, Trans. 
N.Z. Inst. vol. xvii. p. 211. 


This Dolphin has been recorded from the South Atlantic Ocean 
and Cape seas. Probably it should be united with S. malayanus 
Lesson (1826) and S. frenatus F. Cuvier (1836), and the distri- 
bution given as Atlantic, Indian, and Pacific Oceans. Its inclusion 
in the New Zealand fauna rests on a single skull. There is also 
in the Dominion Museum a mandible which I refer to this 
species. The tip is broken, but there are 36 alveoli present. 
Length of ramus 34 em., of symphysis 6 cm. 

Parapara.—Skull in Dominion Museum. Hector doubted this 
skull being obtained in the New Zealand area. It, however, bears 
the legend ‘“ Loc. Parapara. Hector 1879. Steno attenuatus 
Gray” written on the premaxille. The proportions of the skull 
and the contour of the premaxille are identical with the type of 
D. attenuatus figured by Gray (Zool. ‘Hrebus’ & ‘ Terror,’ pl. 28), 
but there are high fronto- supraoccipital ridges (like the figure of 
D. doris in Gray’: swork). It is a large skull, and the high’ ridges 
may be. a sign of age. Pterygoids ‘in contact. Teeth “missing, 
alveoli 39, but end of rostrum not quite perfect. Length of skull 
39-2 cm., of rostrum 24 cm.; breadth of skull at sypuamosals 
16-2 cm., “at orbits 14-7 em. ; breadth of rostrum at base 9 em. 


DELPHINUS. 
Delphinus Linné, 1758, Syst. Nat. ed. 10, Pp Oh wee, D. delphi 


Linné, 


DELPHINUS DELPHIS. 


Delphinus delphis Linné, 1758, Syst. Nat. ed. 10, p. 77 
(European seas). Delphinus nove zealandie Quoy & Gaimard, 
1830, Voy. ‘Astrolabe,’ Zool. vol. i. p. 149, pl. 28; Hutton, 1877, 
Trans. N.Z. Inst. vol. ix. p.349. Delphinus forsteri Gray, Hector, 
1872, Ann. Mag. Nat. Hist. ser. 4, vol. ix. p. 438.; id. 1873, 
Trans. N.Z. Inst. vol. v. p. 158, pls. 2, 3; 1d. 1874, 1. c. vol.. vi. 
Peco vad LS, Vey voli ple . 


The Dolphin is found in all seas; it is common round the New 
Zealand coasts. 

Bay of [slands.—Skull in Dominion Museum. 

Hauraki Gulf.—1 have seen a skull from Motutapu Island. 

East Coast, North Island.—Specimen captured by the ‘Astrolabe’ 
off Tolaga Bay. Described by Quoy and Gaimar das the type of 
Delphinus nove zealandic. 

Wanganui Beach.—Skull described by Hector (1873). 

Cook Strait.—{a) Lyall Bay. Skeleton in Dominion Museum. 
Exterior described by Hector (1874). Teeth $$. Vertebre: 
C.7, D. 14, L. 17, Ca. 37=75. Two cervicals fused. Ribs 14, 


584 MR. W. R. B. OLIVER: REVIEW OF THE 


of which 3 are two-headed. Costal ribs 8, of which 5 articulate 
with the sternum. Sternum of one piece. Phalanges: 2, 9,7, 2, 1. 
(6) Porirua. Skull im Dominion Museum. Described by Hector 
(1872). 

Lytielton Harbow.—Skeleton in Canterbur y NiEnsecne from 
Governor’s Bay, 1896. Vertebre: C.7, D. 15, L. 20, Ca. 33=75. 
Ribs 15, of which 5 are two-headed, the last pair small, half the 
length of the 14th. Costal ribs 8, of which 4 appear to articulate 
directly with the sternum. Sternum of one piece, apparently of 
three fused segments. Pelvic bones curved, rod-like, about 
8 cm. long. 

There is a skeleton from New Zealand in the British Museum. 
True (Bull. U.S. Nat. Mus. 36, p. xxxvi, 1889) gives the measure- 
ments of the skull. 


TURSIOPS. 


Tursiops Gervais, 1855, Hist. Nat. Mamm. vol. ii. p. 323; 
type, Delphinus tursio Fabricius (= D. truncatus Montagu). 


TURSIOPS TRUNCATUS. 


Delphinus truncatus Montagu, 1815, Wern. Trans. vol. iui. pl. 5. 
Tursio metis Gray, Hector, 1873, Trans. N.Z. Inst. vol. v. p. 162; 
Gla Weds je we Wolle ib jo: 477, pls. DW 2 Elitton Sion eieme: 
vol. vii. p. 180. Tursiops tursio (Fabr.), Hector, 1885, 1. c¢. 
vol. xvil. p. 210. Beluga kingii (not Gray) Hector, 1873, Ann. 
Mag. Nat. Hist. ser. 4, vol. xi. p. 105; id. 1873, Trans. N.Z. Inst. 
vol. v. p. 163. Delphinapterus leucas (not Pallas) Hector, 1885. 
Me vol. xvii. py 209) 


The Cowfish appears to be widely distributed, having been 
recorded from localities as distant as the North Sea and New 
Zealand. In the latter place it has been recorded from both 
east and west coasts of the South Island. 

Lyttelton Harbour.—Skeleton in Canterbury Museum, from 
Governors Bay, 1895. Teeth 24. Vertebre: C.7, D. 13, L. 17, 
Ca. 27=64. Two cervicals fused. Ribs 13, with a vestige about 
4 cm. long on the right side only of a 14th. Costal ribs 9, of 
which the first is rather broad and thick, five articulate directly 
and four indirectly with the sternum. Sternum of 4 segments, 
the anterior curved upwards and notched in front and with a 
minute perforation in the centre. 

Otago Harbour.—1913. Skull and cast in Otago Museum. 

Dusky Sound.—(a) Skull in Otago Museum. 1898. (6) Skeleton 
in Otago Museum. Captured in Useless Bay, 10th May, 1875. 
Exterior and skull described by Hutton. Teeth 23. (c) Skeleton 
in Dominion Museum. Captured in Useless Bay, “10th May, 1875, 
Described by Hector (1877). Teeth $8. The anterior pair, upper 
and lower small, not projecting above the alveoli. Vertebre: 
Cian 2 le: 7, Ca, 28=64. Two cervicals fused. Ribs 12, 
of clad 4 are umeiantled) Costal ribs 10, of which 5 articulate 


CETACEA OF THE NEW ZEALAND SEAS. 5B5 


with the sternum. Sternum of 4 fused segments. (d) Skull in 
Dominion Museum, labelled ‘‘ Dusky Bay, 6/1/73.” 

Locality not stated.—(a) Skeleton in Wanganui Museum. 
Teeth 22. Vertebre: C.7, D. 12, L. 17, Ca.22+ =58+. Ribs 12, 
of which 5 are two-headed. ‘Two cervicals fused. (6) Skull 
(incomplete) in Dominion Museum. Labelled in Dr. Knox’s 
handwriting “ Beluga kingii Gray. The Australian Beluga. 
From Swainson Coll.” This is merely an imperfect cranium ; 
the whole of the rostrum and mandible are missing. It is from 
a young animal, as the frontal is quite separate and the cranial 
sutures are open. It appears to agree in all essential characters 
with the skulls of Twrsiops truncatus in the Dominion Museum. 
The main point of difference which gives the present specimen a 
distinct appearance is the absence of a transverse supraoccipital 
ridge, but this is obvionsly due to age, and could not be expected 
to be pronounced in a young animal. This specimen was referred 
to under the name Beluga kingit by Hector (1873), and is 
responsible for the introduction into New Zealand literature of 
the ‘‘ White Whale, Delphinapterus leucas,” which finds a place in 
Hutton and Drummond’s ‘ Animals of New Zealand,’ but has 
been treated more cautiously by Waite, who omitted it altogether 
in his ‘Guide to the Whales and Dolphins of New Zealand.’ . 


EXPLANATION OF THE PLATES. 
Puate I. 
Neobalena marginata. 


Fig. 1. Skull, lateral view. 
2. Skull, superior view. 
3. Skull, inferior view. 


Puate II, 


Cephalorhynchus hectori. 


ee 
a 


ig. 1. Side view. 

2. Inferior view. 

Kogia breviceps. 
Fig. 3. Specimen stranded at Wanganui, 
February 1909. 


Prats III. 
Mesoplodon bowdoini. 


Fig. 1. Skull, superior view. 
2. Skull, inferior view. 
3. Skull, lateral view. 


Prate IV. 


Mesoplodon bowdoini. 


Fig. 1. Mandible. 
2. Left ramus of mandible. 
8. Scapula. 


DENTAL CHARACTERS OF AUSTRALIAN RATS. 587 


29. On the Dental Characters of certain Australian Rats. 
By Prof. F. Woop Jonus, Adelaide University. 


[Received May 22, 1922: Read June 18, 1922.] 
(Text-figures 1-12.) 


The observations recorded in the present paper, although very 
limited in their scope, may possibly prove to be of some value 
when they are extended over a wider range of types than is avail- 
able to the author. 

The business of diagnosing the specific characters of the 
material which he studies does not, as a rule, fall within the 
province of the anatomist, but there are times when even 
the teacher of human anatomy has to enlarge his field of enquiry, 
and enlarge it so that he comes within measurable distance of 
being a systematic zoologist. The attempt to assign a specific 
name to an Australian Murine is an enterprise no amateur 
should engage in, and it must not be thought that the present 
author is ‘responsible for the determination of the species men- 
tioned here. But short of giving an actual name toa species, it 
may happen that one who is not a systematic zoologist may want 
to track down, as nearly as possible to its proper position, some 
animal into the structure of which he is enquiring. 

Among the characters which bulk large in the differential 
diagnosis of various Murines are the crown patterns of the molar 
teeth. It is quite certain that not a tithe of the literature 
dealing with the molar patterns of the rats has been reviewed 
during a search of the works and periodicals available here in 
Adelaide; but enough has been studied to convince the author 
that although it is a simple thing to diagnose the crown occlusal 
pattern of the molars of a young animal, it is ditticult or even 
impossible to say what may have been the pattern when once the 
molars are worn down in an old or an aged specimen. ‘Text-fig. 8 
illustrates the condition of the left upper molar series in three 
individuals belonging to one species, and it is’easy to see that in 
the oldest individ ral ¢ a diagnosis of the original occlusal pattern is 
a matter of considerable uncertainty. Consideration of the inter- 
esting problem of the relation of crown-pattern to root-formation 
has prompted the author to turn to the root-patterns in order to 
see if they presented any constant or useful features. 

A series of circumstances has led up to this little investigation 
of the root-patterns of a few Australian rats. In the first place, a 
number of skulls and certain cranial fragments were found upon 
an island,—Franklin Island in Nuyt’s Archipelago,—the living 
rats inhabiting which had been already properly identified. The 
skulls were all of aged individuals, and no diagnosis could be made 
from the molar patterns; and yet it was of some importance 
that the identity of the fragments should be sufficiently well 


588 : PROF. F. WOOD JONES ON THE 


established to make their relation to the living rats of the 
island clear. 

In the next place, upon another island (Goat Island) in the 
same Archipelago the tracks of a rat were seen upon the sea- 
beaches, but despite every effort no rat could be caught or shot or 
even seen during a brief visit to the island. Nevertheless the 
fact that the tracks were in reality those of rats was proved by 
the finding of skull fragments in the dejecta of some birds of prey. 
These fragments were all of lower jaws and lacked teeth (see 
text-fig. 1). Save for a single humerus no other portion of the 
rats was recovered. It was a matter of some interest to see if 
the rat of Goat Island, and the living rat of Franklin Island, and 
the dead rat of Franklin Island were or were not identical. 


Text-figure 1. 


Fragment of left lower jaw taken from the pellets of birds of prey on Goat Island; 
Nuyt’s Archipelago. The edentulous alveolus has a characteristic root-pattern. 


Still more recently the author was confronted with the remains 
of so many rats that the deposit formed by their bones constituted 
so vast a bulk as to be exploited as a commercial undertaking, 
though situated some 40 miles from the railroad and some 
200 miles from the place to which the deposit could be sent by 
rail. These bones were in a system of caves (Buckalowie), and 
among the millions of rats which had gone to the formation of 
the deposit not one seems to be represented by a whole skull. 
Jaws and fragiments of jaws devoid of teeth were to be had by 
the thousand (see text-fig. 2), and at once the question arose— 
Could the fragments be identified by an examination of the jaws 
from which the teeth had been lost ?¢ Whatever the rats were, 


DENTAL CHARACTERS OF AUSTRALIAN RATS. 589) 


they were certainly ancient and they were mixed with the remains 
of many other animals, amongst which were Zhylacinus and 
Thylacoleo. Even the accident of the loss of teeth in the jaw 
fragments composing the deposit on the floor of the caves provided 
a partial answer to the question, for it was at once evident from 
an examination of the alveolar cavities for the upper molars that 
two quite different root-patterns were represented. 

In order to determine the affinities of these root-patterns the 
author extracted teeth from such properly identified skulls as he 
could obtain, and the present paper is merely a record of the 
findings. If the results are capable of no further extension, at 
least they permit one to say that the past and present rats of 


Text-figure 2. 


Typical fragments from bone débris of Buckalowie Cave deposit. A. Portion 
bearing left upper molar root-cavities. B. Left lower molar root-cavities. 


Franklin Island are the same animal, that the murine contem- 
porary of Z’hylacoleo which formed the cave-deposit at Buckalowie, 
was a creature a good deal like the Franklin Island rat; but that 
the unknown rat which lives on Goat Island is not the same sort 
of rat at all, but is like a rat whose remains make a small and 
recent addition to the cave-deposits and like existing members of 
the genus Rattus. 


Hypromys Geoff., 1805. 


The species examined is that known as the ‘‘Golden-bellied 
Water-Rat,” H. chrysogaster Geott. It is now a comparatively 
rare animal, and in South Australia (and in some of the other 
States) it is becoming increasingly difficult to obtain. The speci- 
mens examined were trapped either on the Onkaparinga River in 
the Mount Lofty Range or on the River Murray at Tailem Bend. 
The animal is so thoroughly distinctive that its specific diagnosis 
needs no authority. 


590 PROF. F. WOOD JONES ON THE 


Upper Molars. 


The upper molar series when examined from its occlusal sur- 
face presents a remarkably simple pattern in the young animal 
when practically no signs of attrition are present, and the same 
simplicity is preserved in the adult when attrition is well advanced 
(see text-fig. 3, A). The anterior molar consists of three very 
distinct masses arranged in the antero-posterior axis of the tooth. 


Text-figure 3. 


Left upper molar series of Hydromys chrysogaster. A, showing the occlusal 
surface of the two molars; and B, the alveolar cavities. The outlines of the 
anterior molar are superimposed on its alveolar cavities in B. The cavity 
marked X is not present in some specimens. 


Each of these divisions shows a central depressed area when 
viewed from the occlusal surface. 

The posterior molar consists of a small anterior lingual portion 
and a larger posterior mass. The total area of the posterior 
molar is less than half that of the anterior tooth. (See text- 
fig. 3.) In advanced age the wear of the tooth involves the raised 
rim of the individual crown masses, and beyond a flattening of 


DENTAL CHARACTERS OF AUSTRALIAN RATS. 591 


the occlusal surfaces no marked change of crown-pattern takes 
place with increasing age. 

Upon removing the two upper molars in Hydromys a very 
curious arrangement of the alveolar cavities is seen. There are- 
either 8 or 9 separate sockets for the reception of roots, small or 
large, of the anterior tooth, and two cavities for the roots of the 
posterior tooth. (See text-fig. 3, B.) 

Of the 8 or 9 root-cavities for the anterior tooth 2 belong to 
the anterior lamina of the crown, 5 or 6 belong to the middle 
lamina, and | to the posterior lamina. 


Text-figure 4. 


AX 
Z mngual 


Anterior left upper molar of Hydromys chrysogaster to show the relation of the 
multiple roots to the crown divisions. In this example only two lingual roots 
are present. A =anterior, P=posterior extremity of molar. 


The anterior root is large and shows at its extremity a tendency 
to be itself subdivided into 3. Four very minute labial cavities 
belong to four small labial roots of the medial lamina and one or 
two far larger lingual root-cavities belong to the same subdivision. 
The small posterior portion of the anterior molar has one large 
alveolus for a single large root which shows a marked tendency to 
be bifid. (See text-figs. 3 & 4.) 

The two roots of the posterior tooth are arranged in the long 
axis of the jaw and thus both tend to be elongated from side to. 
side and to be bifid. 


592 PROF, F. WOOD JONES ON THE 


Lower Molars, 


The lower molar series consists of two dental masses with the 
same simple type of occlusal surface as that displayed by the 
upper teeth. The anterior molar is divided into two main parts, 
and in some specimens, but not in all, a very small posterior shelf 
is present. The posterior molar is also divided into two main 
masses, but the whole area of the tooth is considerably less than 
that of the anterior one. (See text-fig.5, A.) Again there is 
the same complexity of the alveolar cavities and roots. Again 
9 root-cavities belong to the anterior molar mass. The anterior 
portion of the anterior molar has a large anterior root and two 


Text-figure 5. 


Left lower molar series of Hydromys chrysogaster. A. Occlusal surface. 
B. Alveolar cavities, the outlines of the anterior molar being shown. 


small postero-lingual and two small postero-labial roots. The 
posterior portion has two small labial, one small lingual, and a 
large, transverse, and partially sub-divided posterior root. There 
are again two rather tortuous root-cavities, situated antero- 
posteriorly, for the posterior molar. (See text-fig. 5, B.) 
Looking at the whole of the peculiar features of this upper and 
lower molar series, with its strange collection of roots and alveoli, 
it is difficult to avoid conjecturing that, despite the simple crown- 
pattern of the anterior molar it is in reality very far from being 
a simple tooth. It is difficult to avoid speculating as to whether 
Hydromus has in reality simplified its dentition by the loss of a 


DENTAL CHARACTERS OF AUSTRALIAN RATS, 593 


molar. If the line of the palatal suture, or the origin of the 
posterior zygoma root, should prove to be at all stable points in 
cranial architecture, it would look as though the anterior lamina 
of the upper anterior molar occupied somewhat the same 
anatomical position as the whole of the anterior molar in more 
typical forms. 


Rattus. 


(1) &. greyi. Specimens from Mount Compass, South Australia. 
Identity established by Mr. Oldfield Thomas (Ann. & Mag, Nat. 
Hist. series 9, vol. viii. p. 425, October 1921). 


Upper Molars. 


The crown-pattern of the upper molars is shown in text- 
fig. 6, A, 


Text-figure 6. 


Left upper molar series of Rattus greyi. A, showing the occlusal surface; 
and B, the alveolar cavities. 


It is typical and needs no description. 
The roots are small, their alveoli being a series of clean-cut 


594 PROF. F. WOOD JONES ON THE 


holes in the jaw-margin, and the tooth itself is not to any extent 
received in a depressed area of the jaw. 

The anterior molar has 5 roots, a large single median anterior 
root and 4 others arranged in pairs behind it. 

The second molar has 4 roots arranged as an anterior pair and 
a posterior pair. The third molar has an anterior pair of roots. 
arranged labio-lingually and a single median posterior root. In 
the whole root series there is a single anterior median root, and a 
single posterior median root, and 5 lingual and 5 labial roots 
arranged in pairs (see text-fig. 6, B). : 

What might be termed the root-formula for the upper molars 
would therefore be 5. 4. 3. 


Lower Molars. 


The occlusal surface of the lower molars shows two sub-divisions 
of the crown of the two posterior molars and three sub-divisions 
of the anterior molar,.the anterior and middle sub-divisions of 
the anterior tooth being often partially fused together (see text- 
LIC PAD)s 

The root-cavities are well developed and clear cut, like those of 
the upper molars. The anterior tooth has 4 roots, consisting 


Text-figure 7. 


Left lower molar series of Rattus greyi. A, the occlusal surface; 
and B, the alveolar cavities. 


of an anterior median rounded root followed by a pair of roots 
(lingual and labial), followed again by a single root elongated 
from side to side. 

The second and third teeth have 3 roots each arranged as a 
pair of anterior rounded roots and a single elongated posterior 
root (see text-fig. 7, B). The lower root-formula could therefore 
be written as 4. 3. 3. 

(2) &. terre-regine shows exactly the same condition as R. greyi 


DENTAL CHARACTERS OF AUSTRALIAN RATS. 595 


and the introduced forms 2. raétus and R. norvegicus are identical. 
Evidently the unknown rat which lives on Goat Island is akin, 
as a comparison of text-figs. 1 & 7, B, will show. 


Leporitius Oldfield Thomas, 1906. 


The species examined is that recently named ZL. jonesi (Ann. & 
Mag. Nat. Hist. ser. 9, vol. vill. p. 618, Dec. 1921). All the 
examples come from Franklin Island in Nuyt’s Archipelago, on 
which island the type was taken. 


Upper Molars. 


The upper molar occlusal pattern is shown at text-fig. 8. The 
molars are large and are deep set in the jaws. The alveolar 
cavities show a rather noteworthy sinking of the whole tooth 
into the alveolar area of the jaw; the cavities for the indi- 
vidual roots opening from an area which is itself somewhat 
below the general surface of the surrounding bone. (See text- 


fig. 9.) 


Text-figure 8. 


Left upper molar series of Leporillus jonesi. Three specimens, A, B, and C, 
showing the alteration of the occlusal pattern, with varying degrees of attrition. 


The first molar has 3 roots, each having a distinct socket in 
the jaw. Of these three roots one is anterior and median, and 
the other two are posterior lingual and labial; the postero- 
lingual root in some specimens shows a tendency to be bifid. 

The second molar also has 3 roots, but the lingual root is 
further advanced in the jaw so as to be more truly lingual, and 
not so postero-lingual in position. 


Proc. Zoou. Soc.— 1922, No. X LI. 4] 


596 PROF. F. WOOD JONES ON THE 


The third molar has 2 roots, of which the anterior is elongated 
from side to side and shows a tendency to bifureation ; possibly 


Text-figure 9. 


Alveolar cavities of the left upper molar series of Leporillus jonesi. 


it represents the anterior and lingual roots of the second molar. 
The root formula would therefore be 3. 3. 2. 


Lower Molars. 


Of the lower molars the large anterior member shows three 
lamin on its occlusal surface, the second molar two laminz, and 
the third also two. The root-pattern shows the same reception 
of part of the crown into a depressed area of the jaw as is seen 
in the case of the upper teeth. (See text-fig. 10, B.) Hach 
tooth is provided with 2 roots, and anterior rounded root, and a 
posterior one elongate from side to side. In the case of the first 
molar there is a tendency towards lateral bifurcation of the 
posterior root. The lower root-formula is therefore 2. 2. 2. 

The rats whose remains have gone in such countless numbers 
to constitute the bone-doposits in the Buckalowie caves, had the 
molars sunk into the jaw in the same fashion as is seen in the 
Franklin Island Leporillus. They had the same root-formula 
above and below (see text-fig. 2 and compare text-figs. 9 & 10); 


DENTAL CHARACTERS OF AUSTRALIAN RATS. 597 


they also seem to have had the same crown pattern, certainly 
they had not the crown pattern which is distinctive of Conilurus. 
The measurements of fragments of their skulls are practically 
those of the Franklin Island Leporillus; and it is probably not 
fav from the truth to say that a rat very like the modern insular 


Text-figure 10. 


Left lower molar series of Leporillus jonesi. A. Occlusal surface. 
B. Alveolar cavities. 


form was a contemporary of Zhylacoleo and Thylacinus in the 
northern parts of South Australia. No traces of living rats were 
seen at Buckalowie, nor do the guano-miners know of the 
presence of any in the district. 


Notromys Lesson, 1842. 


The only example of this genus that I have been able to 
examine is JV. cervinus, the specimen coming from Miller’s Creek 
to the west of Lake Hyre. 

The specimen is one which shows very little wearing of the 
occlusal surfaces of the molars. There is seen the same tendency 
for the tooth to be received in a depression of the jaw as was 
noted in Leporillus. The root-pattern is also reminiscent of that 
genus, for the upper molars 1 and 2 are 3-rooted teeth, the 
roots being one anterior and two posterior. The third molar, 
however (in the only specimen examined), showed a single 
cavity which contained a partially subdivided root. (See text- 
mies WL) 

- The root-formula would therefore be 3. 3.1. 

The lower molars are all 2-rooted teeth, but there is a well- 

marked tendency towards reduction of the posterior root of the 


third molar. The root-formula is 2. 2. 2. 
4|* 


598 DENTAL CHARACTERS OF AUSTRALIAN RATS. 


Text-figure 11. 


Left upper molar series of Notomys cervinus. A. Occlusal surface. 
B. Alveolar cavities. 


Text-figure 12. 


Lower molar series of Notomys cervinus. A. Left upper molar crown-pattern. 
R. Right lower molar root cavities. 


ConcLusion.—It is suggested that the study of the molar root- 
pattern, and the expression of what might be termed a root- 
formula, are of some importance in systematic work among the 
Murines. Unfortunately it is not possible in Adelaide to carry 
out even a partial survey of the Australian rodents, for the 
material is not obtainable, but it is hoped that such workers as 
possess material will record the root-patterns of Condlurus, 
Mastacomys, and other interesting Australian forms. 


2, Z S> 922, GARTER, Pil. 


STRUCTURE 


OF THE ENAMEL 


SSSR. WRAS 


IN THE PRIMATES. 


—_ 


Py Zo Se WG, CARTIER, JE I. 


STRUCTURE OF THE ENAMEL IN THE PRIMATES. 


DA, S, 1922, CARTER, Pi, i. 


SHRUCHURE OR Tne ss NAMEE SING iE me RiMAGiE Ss 


P. Z. S. 1922, CARTER, PI. IV. 


STHRUCHKURE Ob Hib ENAMEL IN dudes PR UMAGKE'S: 


Vv. 


1922, CARTER, Pl. 


lee Aa Se 


INSEE eR MAMES: 


STRUCTURE OF THE ENAMEL 


I> Zo Ss, IQP2, CARER, Plo Wl. 


SMOG S, ON ASUS IBINVNWUS IL, WN WIENS PRIMATES, 


IP Zo Se 1922 CARWEIR, Jl Wil. 


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STRUCTURE OF THE ENAMEL IN THE PRIMATES. 


STRUCTURE OF THE ENAMEL IN THE PRIMATES. 599 


30. On the Structure of the Enamel in the Primates and 
some other Mammals. By J. THorRNroN CARTER, 
Hon. Research Assistant, University College, Uni- 
versity of London. 


[Received May 24, 1922: Read June 13, 1922. ] 
(Plates I. to VIT. *) 


During the past forty years the theory of the evolution of the 
crowns of mammalian molar teeth, which is associated with the 
names of Cope and of Osborn, has attracted the attention of a 
large number of workers, so that a voluminous literature on the 
subject has accumulated. 

Whilst so much work has been devoted to the gross forms of 
the teeth, it is surprising to find that little attention has been 
given to the minute anatomy of the dental tissues as evidence in 
throwing light on the problem of affinity. 

In the year 1850,Sir John Tomes, F.R.S., submitted to the Royal 
Society a contribution, entitled ‘‘The Structure of the Dental 
Tissues of the Order Rodentia” (Phil. Trans. 1850), in which he 
gave a detailed description of the pattern of the enamel in various 
genera, and from their comparison drew the interesting con- 
clusion that the various families of Rodents possess a ‘“ constant 
and exclusive character in the structure of the enamel”; further, 
that there is a “different and distinct character in each of the 
larger groups, and that the variety of structure is constant 
throughout members of the same group: we may take, for 
example, the Sciuride, the Muride, and the Hystricide, in each 
of which the structure of the enamel is different and in each is 
highly distinctive”... ... ‘“‘the varieties in the structure of the 
dental tissue, with a few isolated exceptions, justify and accord 
with the arrangements of the order into the several divisions 
proposed by Mr. Waterhouse and deduced by him from the 
relations of the several parts of the skull.” 

Yet a year earlier than the publication of his observations on 
the histology of the enamel-in Rodents, Tomes had contributed to 
the Philosophical Transactions a paper ‘‘On the Dental Tissues 
in Marsupial Animals” (Phil. Trans. 1849, vol. exxxix.), in which 
he demonstrated a marked feature in their enamel in the form of 
a system of tubes continuous with the dentinal tubes, this being 
a constant character of the teeth of all Marsupials, excepting 
only those of the Wombat. This character “attains its utmost 
development in the Macropodide, and is more feebly present 
in some Dasyuride, whilst in Myrmecobius and yet further in 


* For explanation of the Plates. see p. 607. 


600 MR. J. THORNTON CARTER ON THE 


Tarsipes 1t is reduced to small limits” (Tomes, Dental Anatomy, 
p- 43). 

The significance and value of these contributions lies in the 
fact that Tomes demonstrated clearly that the enamel pattern in 
Rodents provides a specific criterion of affinity, and that the 
structure of the enamel in Marsupials, taken in conjunction with 
other anatomical characters, may be employed to demonstrate 
Marsupial relationship. 

Seventy years have passed since the publication of these 
researches, but, in so far as I am aware, during that period no 
one has extended Tomes’s work and published the results of the 
examination of the enamel structure in all the representative 
members of a diversified order; and further, with the exception of 
Sir Charles Tomes, F.R.8., the distinguished son of a distinguished 
father, no one has applied the results obtained to the determt1- 
nation of affinities or relationship in the case of extinct mammals. 
In a paper on the “‘ Minute Structure of the Teeth of Creodonts, 
with special reference to their suggested resemblace to Mar- 
supials” (P. Z. 8. 1906), Sir C. Tomes employed the character of 
tubular enamel as a test of Marsupial relationship, and found 
that in none of the Creodont teeth which he examined was this 
character present. Included with this Creodont material was a 
portion of a tooth of Borhycena, one of the Sparassodonta, in the 
enamel of which Tomes found no trace of tubes. But the 
Marsupial characters of this group were so numerous that Prof. 
D. M.S. Watson, F.R.S., asked me to undertake a further exami- 
nation of the enamel in other members of the suborder, and I 
was fortunate enough to discover this Marsupial character in the 
teeth of Cladosictis, Pharsophorus, and of Borhyena (Journ. 
Anat. 1919). 

Following the provision of the material for the examination of 
Sparassodont teeth, the authorities of the American Museum of 
Natural History handed over to me a rich and representative 
collection of teeth and jaws of Hocene mammals, comprising some 
eighty genera, which had been formed for the purpose of obtain- 
ing a set of sections with the hope that, in the words of 
Dr. Matthew, “the investigation thereof will provide a new line 
of evidence for the aflinities of mammals, as distinct from the 
skeleton or the teeth or the soft anatomy”..... and to “ have 
a cross check on relations of the same kind as the cross check 
between teeth adaptations and feet adaptations.” Already several 
hundreds of sections have been prepared from this material, and 
the results obtained encourage the hope that Dr. Mathew’s expec- 
tations will be fulfilled. To obtain the utmost value from such a 
collection, it is desirable to possess a complete series of sections of 
teeth of every genus of existing mammalia, and, in consequence, 
I have laboured to build up a collection of ground sections which 
should embrace not only representatives of each genus, but also, 
in so far as is possible, sections of each tooth in the individual 
dentition. 


STRUCTURE OF THE ENAMEL IN THE PRIMATES. 601 


Included in the material supplied by the American Museum 
are teeth of Kocene Primates and Menotyphla: this material 
was found to be highly fossilized and very friable, and before 
examining it thoroughly, I decided to complete my collection of 
tooth sections of recent Primates. A large number of old-world 
and new-world monkeys have been examined, and some 200 
sections have been prepared from teeth of practically all the 
members of the suborder Lemuroidea. 

The results obtained from this microscopical examination 
disclose certain new features, which, taken in conjunction with 
other anatomical characters, should be of value. 

The first striking feature to be recorded is that throughout the 
whole suborder Lemuroidea a constant feature in the enamel is a 
regular system of tubes continuous with the dentinal tubes. 
Chiromys provides a solitary exception, and thus occupies a 
position similar to Phascolomys amongst the Marsupials. The 
richness of penetration varies greatly both in the teeth of 
different families and also in different areas of the enamel of 
individual teeth *. 

In the Lemuriformes the amount of penetration is usually 
sight, seldom exceeding the extent shown in PI. II. fig. 2, where 
the tubes (¢) are seen to pass through about half the thickness of 
the enamel: over the apex of the unworn incisor teeth and in the 
cusps of the molars the tubes often pass outwards to within a 
short distance of the free surface, as shown in PI. III. fig. 2. As 
one passes downwards towards the neck of the tooth and the 
enamel becomes thinner, the penetration becomes more sparse, 
most of the dentinal tubes splitting up into a fine brushwork just 
beneath the enamel surface. Such tubes as pass over into the 
enamel traverse a layer of uncalcified tissue shown in PI. il. 
figs. 1 & 2 and designated by the letters 2.0. 

In the Indrisine the tubular penetration is extremely rich, 
rivalling the extent found in the Macropodide; for over the 
apices of the cusps of all the teeth there are tubes in the enamel 
continuous with almost every dentinal tube, and these pass out- 
wards through the greater thickness of the enamel. The illus- 
trations shown in Pl. I. figs. 1 & 2 give a clearer idea of this 
condition than can be conveyed by a verbal description. 

Towards the necks of the teeth the penetration is still quite 
rich, but in the sulcuses of the crowns of the molars the pene- 
tration does not proceed to any great extent. 

In Propithecus the degree of penetration is markedly less than 
in the Indrisine, but in certain areas of the enamel it is a con- 
spicuous feature (Pl. II. fig. 3). 

In the Loriside there is a very rich penetration, attaining its 


* For simplicity I retain the terms “tube” and “penetration,” since they are 
employed in all: text-books in the description of this condition in the enamel, but I 
consider the so-called tubes to be fibrils, epiblastic in origin, and though continuous 
with the dentinal fibrils, they are not a penetration of these structures into the 
enamel (vide Carter, Phil. Trans. Roy. Soc. ser. B, ceviil. 1917). 


602 MR. J. THORNTON CARTER ON THE 


greatest development in the enamel of the tooth-cusps. With a 
view to economy of illustration, I have not included a photo- 
micrograph of their enamel, resembling as it does the enamel of 
the Galagos, illustrated in Pl. ITT. fig. 3. 

In Perodicticus potto (Pl. VI. fig. 4) the penetration is not 
nearly so rich as in the Lorises and the Galagos, whilst in 
Nycticebus (Pl. VI. fig. 3) it is but slight, many of the tubes in 
the enamel of the molars ending just within the enamel in the 
form of small bulbous spindles similar to those shown in the 
enamel of Callithriv (Pl. IV. fig. 3). 

In Tarsius, concerning whose position in the classification of 
the Primates so much has been written in recent years, the 
penetration attains a degree of richness even greater than is 
found in the enamel of Galago, and rivals in extent the condition 
shown in /ndris (Pl. III. fig. 1). 

The actual structure of the enamel in the Lemurs also presents 
features of great interest, inasmuch as one finds that in sections 
of the entire tooth, the general pattern, or form and arrangement 
of the prisms, in the Asian and African forms differs from that 
of the Mascarene Lemurs, and that the enamel of Z'arsius is to all 
intents and purposes identical with that of Galago. 

The pattern of the Mascarene genera is shown in the photo- 
micrograph of a molar of /ndris (Pl. V. fig. 1), and this pattern 
is constant for all the genera, including Chiromys (Pl. V. fig. 2). 
Immediately beneath, on the same plate, is a photo-micrograph 
of a portion of the enamel of Semnopithecus (Pl. V. fig. 3), and it 
will be seen that the general structure is closely similar to the 
two lemurs, the enamel being composed of prisms or rods of a 
somewhat granular structure and with straight, even margins, the 
prisms being separated one from another by a slight amount of 
calcified interstitial substance. 

The enamel pattern in the Asian and African Lemurs is quite 
distinct from that seen in the Mascarene forms. It is composed 
of rods or prisms with undulating margins, which in longitudinal 
section present a sharply-defined picture showing the wavy out- 
line of the margins of the rods (Pl. IV. fig. 1). In transverse 
section (Pl. VI. fig. 2) these are seen to be cylindrical rods (e.r.), 
separated one from another by a considerable amount of the 
calcified interstitial substance usually designated by workers on 
the histology of enamel as the interprismatie material (7.p.m.). 

A pattern of enamel identical with this obtains in the teeth of 
the Hapalide and the Cebide. In Callithrix, which Gregory 
regards as a very primitive form, there is a slight degree of 
penetration of the enamel, the prolongation of the dentinal 
tubes terminating in the enamel (e) in the form of little 
bulbous spindles (6) (Pl. IV. fig. 3). The close similarity of the 
enamel pattern of Callithrix, which is identical with that of all 
other Platyrrhines, to the pattern of Galago and of Tarsius 
will be apparent on comparing the micro-photographs figured 


GR) Ii) 


STRUCTURE OF THE ENAMEL 1N THE PRIMATES, 603 


I have prepared sections of teeth of Megaladapis grandidiert 
and of Paleopropithecus from the Pleistocene of Madagascar. 
In the former penetration was almost absent, but in Palceopropi- 
thecus a system of tube is present, developed to an extent which 
equals the condition seen in the teeth of Propithecus itself 
(PG. figs 3): 

In pattern both forms have an enamel identical with that of 
the recent Mascarene forms. 

The microscopical examination of the enamel in Primates 
therefore discloses certain features which have not been noted 
before. They are :— 

(1) The enamel in all Lemurs, with the exception of Chiromys, 
possesses a system of tubes continuous with the dentinal tubes. 
In Jndris and Propithecus the penetration is very rich, but in the 
Lemurine it is feebly developed. In the Galagos, the Lorises, 
and in Perodicticus potto the penetration is rich also, but in 
Nyceticebus it is slight, so slight, in fact, that except in the apices 
of the cusps, the penetration is reduced to a degree which is but 
little greater than one finds in the teeth of the South American 
monkeys. 

(2) There is a marked and constant difference between the 
enamel pattern in the Mascarene forms and the Asian and 
African forms, the enamel in the former having a pattern 
identical with that found in the Catarrhines, whilst the Asian 
and African forms present a pattern identical with that seen in 
the teeth of the Platyrrhines. The Lemuroidea, therefore, parallel 
the Anthropoidea in enamel structure. 

(3) Tarsius has an enamel indistinguishable in sections from 
that of Galago. 

Amongst living mammals the penetration of the enamel by 
tubes is, as already stated, a character found in all Marsupials, 
Phascolomys excepted. No other order possesses this character 
as a constant feature of all its genera. 

In the order Insectivora, however, there are a number of 
genera which do possess it. In the Menotyphla one finds an 
extremely rich penetration in the enamel of the Macroscelide 
(Petrodromus and Rhyncocyon were examined), but in sections 
prepared from a large number of teeth from skulls of Z’upara 
there is no trace of any penetration. In the Lipotyphla all teeth 
of Soricine and of Crocidurine have enamel which is richly 
tubular. Hrimaceus has a moderate degree of penetration, but 
Gymnura has none. The Talpine also do not disclose any 
penetration. 

The two genera comprising the suborder Hyracoidea have an 
enamel which is richly tubular. 

The only other living mammal which possesses this character 
in the enamel is the Jerboa, one solitary creature out of the large 
number comprising the order Rodentia. 

In the enamel of the teeth of all fossil Marsupials which I have 


604 MR. J. THORNTON CARTER ON THE 


examined, extending back to Peradectes (from the Tiffany beds of 
Colorado (Lower Kocene)), there is a penetration. 

In the suborder Multituberculata (Allotheria), which Dr. 
Matthew tells me he regards as in the Metatherian stage of 
evolution, but separate from the Marsupials and Placentals, 
I have also discovered a rich penetration and a distinct pattern 
of enamel, which will be alluded to later in this communication. 

With the presentation of fresh details of the minute anatomy 
of the teeth the first question to arise is that of their 
significance. 

Is the tubular system of the enamel to be regarded as an 
heritage derived from remoter ancestors, or has it some adaptive 
significance in relation to the present needs of its possessor 2 

To throw light on the question, I have prepared sections of 
teeth of several primitive Primates. 

In the enamel of Wotharctus and of Pelycodus, undoubted 
Lemuroid Primates from the Lower Kocene, I have been able to 
discover no trace of penetration by tubes, although I have pre- 
pared a number of sections from teeth of different specimens. 

In Hemiacodon, one of the Anaptomorphide from the Middle 
Kocene, there is marked penetration, well shown in Pl. VI. fig. 1, 
where, in the enamel over the apex of the tooth, the tubes (¢) 
are clearly shown passing some distance into the enamel, though 
a little further down in the crown of the same tooth all trace of 
structure, in both enamel and in dentine, had become obliterated 
in the process of fossilization. 

In Phenacolemur, concerning whose affinities there has been 
some discussion (Broili and Schlosser including it amongst the 
Insectivores, whilst Dr. Matthew, in his list of Eocene material 
supplied to me, includes it amongst the Menotyphla), the enamel 
is also tubular. 

Nothodectes (Plesiadapis), whose affinities also are doubtful, 
discloses no sign of penetration, and the structure of its enamel 
lends no support to the views of Stehlin that it is closely akin to 
the Chiromyide, but rather supports the opinion of Matthew 
that it is nearer to the Menotyphla, and is a primitive synthetic 
type intermediate between Tupaiaoids and Lemuroids. 

With regard to the microstructure of the enamel in these 
extinct Primates, it is interesting to record that Hemiacodon 
possessed an enamel identical in pattern with that of Tarsius of 
the Asian and African Lemurs, and therefore with that of the 
Platyrrhine, whilst Votharctus and its forerunners Pelycodus 
disclose a structure similar to that found in the Mascarene forms, 
living and extinct, and to the Catarrhine. 

It is obvious, therefore, that, in the examination of a fragment 
of a tooth, the discovery of the presence of a system of tubes in 
the enamel, unsupported by other evidence, would not afford a 
precise test of affinity; but, taken in conjunction with the pattern 
of the enamel, the demonstration of tubes would enable one to 
state, with a degree of certainty, as to whether such a fragment 


STRUCTURE OF THE ENAMEL IN THE PRIMATES. 605 


had been derived from a Maultituberculate, a Marsupial, an 
Insectivore, a Lemur, or whether the tooth was that of a Jerboa ; 
and further, as the evidence adduced in this paper shows, in the 
case of a Lemur one could differentiate between the Mascarene 
and the Asian and African forms. 

It remains for me to give a very brief description in general of 
the tubular enamels of those mammals apart from the Lemurs to 
which I have alluded. 

The enamel pattern of the Multituberculates is quite distinctive, 
and differs fundamentally from all others which I have examined. 
If a section of a tooth of Polymastodon be ground and examined 
(Pl. VII. fig. 5), tubes (¢) will be seen passing from.the dentine 
(d) into the enamel (e): by careful illumination a faint pattern 
may be discerned, which can be developed by washing the section 
for a time in acid alcohol until such an image as is shown in 
Pl. VII. figs. 2, 3, and 4 is disclosed. 

Here one sees a series of horseshoe-shaped bodies (h.s.), which 
become smaller and more widely separated as the outer edge of 
the enamel is approached. In whatever plane the section may be 
cut these structures never become complete circles, the two ends 
remaining apart and terminating in bulbous slightly recurved 
enlargements. Such an image would be seen in transverse 
section of any spiral structure. 

Any further description of the minute structure of this peculiar 
enamel would be out of place in this paper, but it may be men- 
tioned that Péilodus also has an enamel which is richly tubular 
and an enamel pattern similar to, but by no means identical with, 
that of Polymastodon. 

The Marsupials possess a very distinctive general pattern, as is 
shown in Pl]. VII. fig. 1, where the rods or prisms (p) are seen, 
in transverse section, to be arranged in rows which sometimes 
merge: these rows are separated by a very definite area of inter- 
prismatic material (7.p.m.), so that, as Dr. Mummery has pointed 
out, in teased preparations of developing enamel this substance 
splits up into lamine. The dark dots (¢) are the so- -ealled tubes 
seen in section. A section showing the enamel prisms in such 
rows and presenting the  tube- Bacneteion is peculiar to 
Marsupials. 

I have not figured the richly tubular enamal of the Jerboa, but 
it is the only rodent which shows any trace of tube- -penetration, 
and its enamel pattern presents the criss-cross arrangement of 
the rods which Sir John Tomes showed to be a character peculiar 
to the Rodents. 

With regard to the significance of the presence of a system of 
tubes in the enamel, the variability of its degree and of its 
distribution and the fact that, whilst richly developed in one 
creature it may be totally absent in a closely-related form, would 
seem to indicate that it is connected with some adaptive process, 
and that, taken alone, it has little or no value as evidence of 
affinity or line of descent. 


606 MR. J. THORNTON CARTER ON THE 


The microstructure of the enamel, however, appears to be par- 
ticularly constant. In the long and well-authenticated series of 
creatures comprising the ancestry of the Horses, the microstructure 
of the enamel does not change, in spite of the modification of the 
teeth from the brachydont form in Lohippus through all the pro- 
gressive stages leading up to the complex hypsodont molar teeth 
of recent horses. 

In the Marsupials the pattern of the enamel in recent and 
fossil Australian forms appears to be identical with that of the 
teeth of Peradectes from the Tiffany beds of Colorado (Lower 
Eocene). 

Certain fossil rodents from the EHocene exhibit a close 
similarity in enamel structure to recent forms, and where, as in 
Chiromys, « Lemur has evolved a rodent dentition, the structure 
of its enamel still retains its Lemurine character. 

It was my intention to hold over a description of the minute 
structure of the teeth of Primates until I had published an 
account of the structure of the teeth in Insectivores, recent and 
fossil, and of the primitive Creodonts, the material for which has 
been examined, together with that of a large number of other 
Kocene mammals. But Professor J. P. Hill, F.R.S., and Professor 
D. M. S. Watson, F.R.S., were of the opinion that the facts 
presented in this communication would be of immediate interest 
to the Fellows of the Society, the more so as recent paleonto- 
logical discoveries have provided fresh evidence of the antiquity 
of the Higher Primates. 


In conclusion, I have to express my deep gratitude to my 
colleague, Major G. S. Sansom, and to Mr. F. J. Pittock for 
their kindness in preparing the micro-photographs used in the 
illustrations, and to Professors Hill and Watson for constant 
encouragement, 

Above all I have to thank Dr. W. D. Matthew, F.R.S., for 
providing invaluable material, and for most valuable information 
and suggestion in conversation during his recent visit to Kurope. 

Most of the recent material employed was acquired by pur- 
chase, but I am deeply indebted to the Zoological Society through 
Mr. R. I. Pocock, F.R.S., for the gift of teeth of Chiromys, 
and to Professor J. Elliot Smith, F.R.S., for a mandible of 
Tarsius. 


Summary. 


A microscopical examination of the teeth of Primates discloses 
the fact that all members of the suborder Lemuroidea possess, in 
a varying extent, the character of a penetration of tubes into the 
enamel. 

In the Lemurine this penetration is slight, but in the Indrisine 
it is very rich. 

The Galagos and Pottos also have a rich penetration, but in 
Nycticebus it is but feeble. 


STRUCTURE OF THE ENAMEL IN THE PRIMATES. 607 


Tarsius has an enamel richly tubular, and in its structure 
essentially Lemurine. 

A further interesting feature is that the pattern of the enamel 
in the Mascarene Lemurs differs from that of the Asian and 
African forms, and that the former has a structure identical with 
that found in the enamel of all Catarrhines, whilst the structure 
of the enamel in the Platyrrhines is identical with that found in 
the Asian and African Lemurs. 

Two primitive Lemuroid Primates, Pelycodus and Notharctus, 
from the Kocene show no penetration, but in the structure of 
their enamel resemble the Mascarene Lemurs. 

Henmiacodon, a Tarsioid Primate, also from the Eocene, has an 
enamel which is tubular, and in structure appears identical with 
the enamel of the recent Jarsius. 

The enamel pattern affords a useful test of affinity if taken in 
conjunction with the character of a tube-penetration, and enables 
one to discriminate between Multituberculates, Marsupials, and 
Placentals. 


EXPLANATION OF THE PLATES. 


The following 1s a list of reference letters common to the various figures :— 


6. bulbous spindles. | h.s. borseshoe-shaped bodies. 
d. dentine. i.p.m. interprismatic material. 
e. enamel. u.l. uncalcified layer. 
er. enamel rods. | 
Prate I. 


Fig. 1. Longitudinal section through the incisor of Indris, showing the presence in 
the enamel (e) of tubes (¢) continuous with those of the dentine. x 180 

. Longitudinal section through the protocone of an upper molar of Indris, 
showing a similar condition to that seen in fig. 1. 180. 


rej 

ie 
gg 

bo 


RramE lle 


Fig. 1. Longitudinal section of a lower incisor of Lemur catta, showing the form and 
structure of the enamel rods or prisms (p), and at the surface of the 
dentine (d@) next the enamel a layer of uncalcified material (w.l.). > 350. 

. Longitudinal section of a lower meisor of Lemur, showing the passage of 
tubes (¢) from the dentine (d) into the enamel (e); the tubes traversing 
the layer of uncalcified material (w./.) are shown in fig. 1. X 350. 

Fig. 3. Section of a lower premolar of Propithecus, showing the presence of tubes (¢) 

in the enamel (e). > 300. 


a 
8 
bo 


Prats II]. 


Fig. 1. Section of a tooth of Tarsius, showing the junction of the tubes (¢) in the 
enamel (e) with those of the dentine (d). X 450. 

Fig. 2. Section of a lower incisor of Microcebus. X 250. 

Fig. 3. Section from the coronal surface of a lower molar of Galago. X 350. 


Prats LV. 


Fig. 1. Section of a tooth of Tarsius, showing the general pattern of the enamel (¢) 
with the undulating margins of the enamel rods (e.7.) and the tubes (¢) in 
the enamel. X 350. 

. Section of a tooth of Galago, presenting an appearance identical with that 
shown in fig. 1. XX 340. 

. Section of a premolar tooth of Callithrix (Callicebus), showing an enamel 
pattern similar to Tarsius and to Galago. ‘The dentinal tubes terminate 
a very short distance within the enamel as little bulbous spindles (8). 
X 350. 


ie?! 

ae 
gg 

i) 


ee 
iQ 
oo 


608 


Fig. 


Oe 


bo 


ON THE STRUCTURE OF THE ENAMEL IN THE PRIMATES. 


PLATE V. 


. Section of the occlusal surface of the left upper molar of Indvis; the photo- 


micrograph taken from the same tooth as the illustration shown in Pl. I. 
fic. 2. ‘The tubes (¢) pass but- a short distance into the enamel (e). 
The pattern of the enamel is clearly shown, the enamel reds (e.r.) having 
straight margins and showing a fine granular structure. X 450. 


. Section of a molar of Chiromys. The dentinal tubes terminate at the amelo- 


dentinal junction. The pattern of the enamel is similar to that of the 
other Mascarene Lemurs. 500. 


. Section of enamel of Semnopithecus. X 450. 


Prats VI. 


. Section of a cusp of a molar of Hemiacodon, showing the presence of tubes 


(¢) in the enamel (e). X 275. 


. Transverse section of an incisor of Galago, showing the enamel rods (e.7.) cut 


across and having a circular form separated one from another by a con- 
siderable amount of interprismatic material (7.p.m.). X 300. 


. Section of a molar of Nycticebus, showing the slight amount of penetration 


of the “tubes” (¢) into the enamel, whilst many end in a bulbous spindle 
(6) similar’ to those shown in the enamel of Callithrix (Pl. IV. fig. 3). 
<< 300. 


. Section of an incisor of Perodicticus potto, showing the passage of tubes (é) 


and an enamel pattern identical with that seen in Galago and in Tarsius. 
x 340. 
Prats VII. 


. Section of an incisor of Macropus ruficollis, showing the enamel rods (e.r.), 


cut transversely, lying in rows which are separated by sharply-defined 
areas of interprismatic material (i.p.m.). The ‘‘tubes ” of the enamel are 
seen as black dots (¢).. X 1500. 


. A transverse section of the enamel of a molar of Polymastodon, etched to 


show the structure. The horseshoe-shaped bodies with bulbous ends (h.s.) 
would suggest sections through an elongated spiral since they never form 
complete rings. XX 275. 


. Transverse section through the enamel of the lower incisor of Polymastodon, 


showing a similar condition to that seen in fig. 2, but demonstrating a 
finely granular structure of the areas embraced within the horseshoe- 
shaped bodies. X 400. 


. Longitudinal section of the enamel of a molar. X 480. 
. Unetched section of a molar of Polymastodon, showing the passage of tubes 


(¢) from the dentine (d) into the enamel (e). 300. 


ID Zo So IGLD, POIRS IME RICOOP EIR, IP, il. 


PROBOSGIDIA from BALUCHISTAN. 


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Seu: 
Theyre 
Pare i) 
; 


P. Z. S. 1922, FORSTER-COOPER, PI. Il 


PROBOSCIDIA from BALUCHISTAN. 


2, Za So IG2P, TORS WEIR COOPEIR, il, Wi. 


PROBOSCIDIA from BALUCHISTAN. 


chirb 


ao 


15 74 So G22; HORSE COORS2, Il. I 


PROEOSCIDIA frorn BALUCHISTAN. 


ON MIOCENE PROBOSCIDIA FROM BALUCHISYAN. 609 


31, Miocene Proboscidia from Baluchistan. By C. ForstEr 
Coopgr, M.A., F'.Z.8., Superintendent of the University 
Museum of Zoology, Cam prrlee: 


[Received May 3, 1922: Read May 23, 1922. | 
(Plates I-LV.* and Text-figures 1-12. 


The present paper contains a description of some Proboscidian 
remains obtained during two expeditions to the Lower Miocene 
deposits of Dera Bugtiin Baluchistan. 

‘These fragments are of interest, as they throw some further 
light on the earliest known Indian Elephants which have been 
described partly as a variety of Bunolophodon angustidens T and 
partly as belonging to a genus, Hemimastodon t, not known else- 
where. ihevar are also earth ol a fragment of a lower jaw of 
a small Dinotherium. 

Schlesinger §, in his beautifully illustrated account of the 
Mastodons in the Natural History Museum of Vienna, has figured 
and. described two forms of B. angustidens. The criteria for 
distinguishing these two varieties lie, according to him, in the 
structure of the molars, and his descr iption.. seems to refer 
especially to the intermediate teeth as being the more commonly 
found remains. In both forms the ridges consist of. a large 
round outer cusp on the outer side of the tooth, with a smaller 
cusp: pressed against its inner border and with one. or -more 
additional cusps in the valleys which separate the ridges, » On the 
inner side of the tooth the ridge. is continued by two similar 
main cusps—a larger on the outer side of the tooth and-a smaller 
on the median side, but without any accessory cusps in the 
valleys. 

Of the two varieties, one, which Schlesinger calls var. typica, 
shows these cusps as round in section and rather separate from 
one another, the surface of wear of the cusps being cireular and 
flat with the general surface of the tooth; the other, var. swb- 
tapiroidea, differs in having the cusps more elliptical.in section 
and less separate from one another, the accessory columns are 
less developed, and the areas of wear are oval and their planes 
slope from the general level of the tooth. While: making. these 
distinctions, Schlesinger at the same time denies the validity of 


* For explanation of the Plates, see page 626. 

+ Lydekker, ‘Paleeontologia Indica,’ ser. 10, vol. vii. pt. 4, pp. 28-25 (1884). 
T use this generic name after Schlesinger, with whose work the chief comparisons in 
this paper are made and without prejudice to other names. The British Museum 
uses the name Tetrabelodon. My friend Professor H. F. Osborn writes to me since 
this paper was set up in type that he prefers the name T’rilophodon. 

t Pilgrim, ‘ Paleontologia Indica,’ n.s. vol. iv. pt. 2, p. 17 (1912). 

§ Schlesinger, Mitt. Geol. Geselt. Wien, Bd. xi. p. 133 (1918). 


610 MR. C. FORSTER COOPER ON MIOCENE 


Lydekker’s variety palwindica, which was founded chiefly on a 
third lower moiar and which he states is within the range of 
variation of var. typica. The specimens now to be described 
tend, if anything, to reverse Schlesinger’s view and in part to 
confirm Lydekker’s. 

The material in the collection consists of two palates of 
B. angustidens of slightly different age, part of a lower jaw 
belonging to the older of the two palates, and some separate upper 
and lower molar and premolar teeth. 

The younger of the two palates (text-fig. 1) has on each side 
the last premolar and first molar in moderate wear, and the 
second molar erupting and unused. In front are the broken 
roots of the third premolar. Beyond this point and behind the 
second molars the specimen ends. The fourth premolars are 
weathered, but show a square outline and four subequal cusps a 
little elongated in the transverse direction. There is a shelf in 
front and low talon behind, and there are no intermediate cusps. 
They are very similar in size and shape to a correspoding tooth 
figured by Schlesinger for B. longirosire. ‘There is no sign of a 
successional tooth underneath, so that it is not likely that they 
are imperfect milk teeth. 

The first molars are three-ridged, with a very small talon. The 
first two ridges are worn, the third as yet untouched. They 
agree more with those figured by Schlesinger as var. subtapi- 
roidea * than as var. typica t. 

The second molars, again, agree best with the figures of szb- 
taptroidea ¢, but here the cusps seem more separate from each other, 
a difference which may be more apparent than real, and due in 
large measure to the tooth being unworn; an internal accessory 
cusp on the third ridge is more prominent than his figures show. 

The other palate is rather older in that the second molar is 
now in wear on the first two ridges. The specimen on its right 
side has an open socket, from which the third premolar has fallen 
out; the fourth premolar and first molar are well worn, the second 
is partly in wear, and the front ridge of the third is preserved in 
the alveolus. It is clear that this animal had two premolars and 
two molars in simultaneous wear, which isa longer time than 
Andrews$ suggests for this species. The present specimen 
appears to show that these four teeth remain until the third molar 
erupts, when it finally pushes the premolars and first molar 
forward and out. This is consonant with the view, supported by 
other reasons, that the stage of ‘angustidens represented by these 
Indian forms is earlier than that of the more typical French 
forms from Sansan etc., and is possibly as early as any yet 
described. 

The structure of the teeth in this second palate is essentially 
the same as in the other, except that in the second molars the 


* Schlesinger, Mitt. Geol. Geselt. Wien, Bd. xi. pl. iii. fig. 2. 
+ Loe. cit. pl. i. fig. 2. { Loe. cit. pl. vir. fig. 3: 
§ Andrews, Phil. Trans. R. 8. vol. xciv. p. 108 (1908). 


PROBOSCIDIA FROM BALUCHISTAN, . 611 


division of the various cusps in the ridges is less noticeable. 
This, again, is apparently due to wear ; the grooves between the 


Text-figure 1, 


», $ NN 


Bunolophodon angustidens. Palate with second molars just erupting. }. 


cusps are often not more than shallow wrinkles in the thick 
enamel, and very little use would be sufficient to obliterate 
them. 


Proc. Zoou. Soc.—1922, No, XLII, 42, 


612 MR. C. FORSTER COOPER ON MIOCENE 


In connection with these palates, the third upper molar (PI. I. 
fig. 1) may be described, of which there are two separate speci- 
mens, a right and left, both unworn and of similar pattern. 
They differ in shape from corresponding teeth from France in 
the collection of the British Museum in being broad in front and 
in getting narrow behind so rapidly that the ground-plan of the 
tooth is that of an isosceles triangle instead of the more usual 
elongated form. 

A tooth figured by Schlesinger * (var. subtapiroidea) is ap- 
parently nearer the present one in shape, but, being tilted up in 
the figure, his specimen cannot be closely compared. In structure 
the two are very similar, except that in the Indian specimens the 
fourth ridge (talon) consists of three cusps only instead of four, 
and that anything in the nature of valley cusps is conspicuous by 
its absence. In Schlesinger’s form the cusps of the main ridges 
are three on each side, except on the last ridge where they are 
double only; in the present form they are partly double, but 
here and there show either the treble condition or the beginning 
of it. In his form, however, there is a pair of accessory cusps 
which, while not very conspicuous, can be seen clearly in the 
first two valleys. These are absolutely unrepresented in the 
Indian form, with the sole exception of the first valley in one of 
the two teeth, which has a low but distinct cusp on the outer 
side connected to the posterior flank of the first ridge. It is 
possible that the “ subtapiroid” condition is the more primitive, 
and small mutations in the direction of greater complication of 
the valleys are therefore to be expected. 

These third upper molars, while nearly as broad as Schlesinger’s 
specimen, are shorter, and are considerably more so than French 
specimens. 

Another separate upper tooth, in quite unworn condition, is 
here figured (Pl. I. fig. 2). It is apparently a second molar of 
the left side, but is rather larger than the corresponding teeth 
in the two palates already described. It is similar to them in all 
respects, except that the arrangement of the cusps is much more 
that of the “ typica” form. ‘This is especially noticeable in those 
of the inner side and on the second ridge, where as many as 
eight cusps of different sizes can be counted. At first sight this 
tooth might be mistaken for a very small third molar on account 
of its rather pointed end, large talon, and absence of posterior 
pressure mark, If this were really so, it might have some bearing 
on Pilgrim’s genus Hemimastodon. The pointed appearance is 
due to the thick enamel being broken away on each side, and is 
therefore purely artificial; if this part be restored, the tooth has 
the square end of a second molar. The large talon is due to the 
tooth being in germ and becomes resorbed when the third molar 
begins to press against it; in each of the two palates in which the 
third molar is not yet in contact with the second the latter has a 
large talon. 

* Schlesinger, loc, cit. pl. vii. fig. 3, 


PROBOSCIDIA FROM BALUCHISTAN,. 613 


Of upper premolars there are two specimens, both unworn. 
One, apparently a third premolar of the left side, consists of four 
cusps arranged in pairs, but not joined into ridges, and a small 
talen. This i isa small tooth (PI. LV. fig. 3) about 30 mm. long and 
20 mm. in width. The other, a fourth premolar (Pl. LV. fig. 2) 
45 mm. long and 36 mm. wide, is formed of two separate cusps in 
front, while the hinder cusps are subdivided each into about four 
smaller ones, and make a nearer approach to a ridge; the talon is 
fairly well-marked. 

It may be stated at this point that no part of the material in 
the present collection gives any evidence in support of the genus 
Hemimastodon *, which was originally described from specimens 
from the same beds in Baluchistan. Schlesinger t has criticised 
the validity of this genus somewhat adversely, and has pointed 
out that, with the exception of the type specimen (a third upper 
molar), all the other specimens fall within the limits of known 
variation of BL. angustidens. The type tooth has only three 
ridges instead of four, and is of a more pronounced triangular 
shape than the third molars just described. Schlesinger has 
made the tentative suggestion that it may be a four-ridged tooth 
with the front ridge broken away +, which has been misinterpreted 
as a complete three-ridged one. This, of course, is possible, and 
fractured specimens of proboscidian teeth with extraordinarily 
little evidence of the fractured surface are not rare. On the 
other hand, the broken edges of these teeth all show a convex 
anterior border, whereas the presumed line of fracture in the 
tooth under discussion is decidedly concave. The normal front 
border of these teeth seems to be convex in unworn specimens, 
which in fully-developed teeth may become flat or concave by 
pressure of the tooth in front. If Pilgrim’s specimen be an 
entire one, there remain alternative explanations, Hither it is 

an abnormal specimen, or Pilgrim was correct in making a generic 

distinction for it and keeping it apart from the genus Buwno- 
lophodon. In neither case is there any proof for or against 
his view, except the negative evidence of two seasons’ collecting 
over the same ground, which has produced nothing to confirm 
the genus Hemimastodon, while the more ordinary Tomatilone of 
Buonolophodon angustidens occurred in fair quantity. 

The right and left fragments of the lower jaw are in pr ecisely 
the same stage of wear as the older of the two palates, and are in 
all probability parts of the same animal. The fourth premolar 
and first molar are well worn, the second has the first two ridges 
in wear, and a complete third molar lies in the alveolus. 

Compared with a specimen in the British Museum, the 
mandible (text-fig. 2) is not only smaller, but gives the impression 
of having been considerably shallower than i is usual; it compares 


* Pilgrim, loc. cit. t Loe. cit. p. 48. 
TA third lower molar minus the first ridge would give a shape very close to 
that tigured by Pilgrim, 
42* 


MR. C. FORSTER COOPER ON MIOCENE 


614: 


EX 


“SNTOAATL Of} UT ABTOU PALY YIM Mel 1aMoT 


'G OANSY-9X9T, 


‘suaprysnbun uopoydojoung 


Se 


= 


PROBOSCIDIA FROM BALUCHISTAN, 615 


more closely with Schlesinger’s figure of a specimen *, a rather 
older animal, which is named var. subtapiroidea, but which is 
noted as not typical. 

The third lower molar of this present specimen agrees, however, 
neither with those figured by Schlesinger for var. typica tT, nor 
for var. swbtapiroidea?, nor with any of the corresponding teeth 
of angustidens in the British Museum collection or at Cambridge. 

On the other hand, there are four specimens of third lower 
molars in the present collection which agree in a certain point 
among themselves and also with that described by Lydekker as 
the type tooth of var. palewindica. 

There are the usual four ridges and small talonid. Hach ridge 
is formed of two cusps, each cusp being unequally divided into a 
large outer and small inner one, the two smaller cusps lying towards 
the middle line of the tooth, and thefour cusps which constitute the 
ridge are arranged in a straight line at right angles to the long 
axis of the tooth. The only exception to the division of the cusps 
is that the inner cusp of the fourth ridge is practically undivided. 
So far these characters approximate to those of swbhtapiroidea ; 
the first and second valleys, however, have each a large cusp 
standing up separately in unworn specimers from the ridges, but 
showing in worn specimens as being more connected to the 
anterior ridge than the posterior; the third valley in this speci- 
men has no cusp visible. This arrangement does not appear to 
be that of Schlesinger’s var. tapiroidea. A reference to his 
figures shows that it should have the valley cusps small or 
absent { and the main cusps clearly divided, while in var. typica 
the main cusps are less clearly divided and the valley cusps are 
prominent. There are, moreover, two cusps to each valley, the 
anterior being derived from the ‘posterior flank of the external 
cusp of the anterior ridge, and the posterior from the anterior 
flank of the corresponding cusp of the posterior ridge §. 

A specimen of a small form of typiea from Sansan, in the 
Cambridge Museum, is figured for comparison with the Indian 
roruas) (Ea otis. 1): 

Hither of these forms can be matched by specimens from the 
Middle Miocene of France in the British Museum collection. 
The valley cusps may be large or small, but always show the same 
tendency to be two in each valley. The Indian specimens all 
show single cusps in the valleys, large in the first, smaller in the 
second, and in the third valley may even be absent. 

Lydekker || states that var. palwindica is separated on account 
of a “somewhat greater development of the accessory columns.” 


* Schlesinger, loc. cit. pl. v. figs. 1 & 2 

seOC CUES pl. vill. 

t This is best shown in his fig. 1 of plate viii. 

§ This arrangement can clearly be seen in his fig. 4 of plate viii., where a string, 
used to support the specimen, runs between the two cusps in the second valley of the 
tooth. 

|| Cat. Foss. Mamm. British Museum, vol. iv. p. 29. 


616 MR. C. FORSTER COOPER ON MIOCENE 


It is, however, not so much that the cusps are necessarily more 
developed, as that their comparative isolation from the ridges is 
the point that strikes the eye. And a greater character is the 
fact that they are single in each valley. An absolutely unworn 
third lower molar similar to the tooth in the jaw described above 
is here figured (Pl. II. fig. 1 and text-fig. 3), but being in better 
condition, it shows the characters more clearly. 

Of European forms in the collection of the British Museum, 
the nearest approach in general appearance to the Indian teeth 
is a damaged specinten with the first ridge and front part of 
the second broken away (Pl. II. fig. 2). The main cusps were 
probably divided as in the Indian specimen, but the divisions 
have become obliterated by slight wear. The third ridge shows 
the presence of the second valley cusp, which is small but enough 
with the larger anterior one to close the valley. The third 
valley is similar to the Indian specimen in having only one cusp. 
The talon of the Indian specimen is very smal] and consists of 


Text-figure 3. 


IMR 


A a. 


ond 


\WN ee 
i Th a saa 


a 


) 
( 


| f 
a au gunn DMS “2. Nn er 


Bunolophodon angustidens. Third lower molar in side view. > 3. 
two cusps only, and this is rather a feature of the Indian teeth, 
which have either two or never more’ than three cusps on this 
ridge. French forms seem to have three or four cusps as in the 
specimen figured, while Schlesinger figures * four molars, two of 
subtapirordea with two cusps, one of typica from France with two, 
and one of typica with three. On the whole, the Indian lower 
teeth seem to come more under Schlesinger’s heading of typica, 
while the upper teeth approach that of subtapiroideat. 

A larger and more worn third lower molar (text-fig. 4) is 
figured which shows a distinct curvature of the lateral borders. 
This curvature Lydekker has quoted as one of the characters of 
var. paleindica, but the presence of perfectly straight teeth in 
these deposits shows that it has no great significance. On the 
other band, it shows very clearly that the valley cusps are single. 

* Schlesinger, foc. cié. pl. viil. 


_t In one case the upper and lower teeth belong to one individual, which renders 
his distinction of less value. 


PROBOSCIDIA FROM BALUCHISTAN. 617 
It is not unlike Lydekker’s figured * specimen (text-fig. 5), except 
that in the latter the cusps of the transverse ridges are not 
straight across, but are bent up in the middle line in the third and 


Text-figure 4. 


Q 


t 
XN 


= \ 


WN XY 
SS} 
eae 


Zs 


: gfe BBE: = 
pei 


s= 


I 
Aig 
S es 


Bunolophodon angustidexs. Third lower molar. X 4. 


especially in the fourth ridges, After making allowance for this, 
his form can be seen to fall in with the other Indian teeth here 
described. As this complete singleness of valley cusps does not 
seem to occur elsewhere, it seems that Lydekker’s view that the 


* Mem. Geol. Surv. Ind. vol. ili. pl. 4, fig. 3. 


618 MR. G. FORSTER COOPER ON MIOCENE 


Indian specimens form a separate variety on this character is to 
some extent justified *, and has at least as much to commend it, 
in the present state of our knowledge, as have the varieties 


Text-figure 5. 


Bunolophodon angustidens. Outline of the type-specimen of var. paleindica, 
figured by Lydekker. X 3. 


typica and subtapiroidea, which appear to have intermediate 
stages between them. 


“Species” and “ subspecies” have little significance in paleon- 


* A close approach, however, to the Indian teeth is figured by Wegner (Palaon- 
tographica, vol. lx. 1913, pl. xv. figs. 8 & 4). This specimen is more advanced in 
its larger talon, and has a number of accessory cusps in the first valley. The second 
and third valleys, however, are like the Indian form except that the cusps of the 
fourth ridge are fully divided into two each. Wegner’s specimens, an upper as well 
as a lower molar, come from the Upper Miocene of Doppeln, and though more 
advanced than the Indian form, are constructed on somewhat thesame pian. Wegener 
notices the same difference between his forms and specimens from Sansan as is 
noticed here. a 


PROBOSCIDIA FROM BALUCHISTAN, |, 619 


tology except for descriptive purposes, and as other material 
becomes available from new sources, all these distinctions will be 
merged in the mass of small variations from which new mutations 
will eventually arise. Moreover, but few specimens of angustidens 
have, as yet, been obtained from India, and additional material 
is required to show whether the differences here noted are con- 
stant or not. 


[IX 


Text-figure 6. 


Nn a Sees Ses Wy 


if 


Bunolophodon angustidens. First lower molar in side view. 1. 


Of the other lower teeth, the second molar is rather different 
from a French specimen from Simorre (B.M. No. 42720) in being 
longer and narrower in proportion, in the more widely open valleys, 
and in the greater prominence of the talonid ridge, where the two 
cusps stand up very prominently (text-fig. 2). Of first lower 
molars there are several specimens, one of them (text-fig. 6 and 


Text-figure 7. 


\ 
\ ™~ 
la on tale 
\ er ES Zn 


Bunolophodon angustidens. First lower molar (B.M. No. 29671) from Simorre, 
in side view. X 1. 


PI. LV. fig. 1) absolutely unworn. Compared with two specimens 
(text-fig. 7), also unworn, from Sansan and Simorre respectively, 
the Indian tooth is rather longer for its breadth and is widest at 
the last ridge; the valleys, as in the other lower molars, are 
less blocked by the accessory cusps. The talonid is formed of two 


620 MR. C. FORSTER COOPER ON MIOCENE 


well-marked cusps, a feature of all the Indian teeth, whereas of 
the EKuropean forms here compared, one has three cusps and the 
other a low ridge of eight or nine wrinkles. The valleys are 
more open and the ridges lower. The accessory cusps in the 
second valley are much lower. 

There remain for a brief notice several ends of lower teeth, 
apparently third molars, which are of unusual form. They all 
resemble the one figured (PI. III. fig. 2) in the buckling up of 
the last ridge and in the curious structure of the talonid, which 
is formed of four cusps of different sizes pressed together into a 
conical hillock. ‘The valley between the last ridge and the talonid 
is widely open and without any accessory cusp. This condition 
Ihave not been able to match at all closely, although some 
specimens of 5. longirostre approach it. The teeth are the 
average size of Bb. angustidens. Whether they represent another 
form or are within the range of variation of the latter species 
cannot at present be told. 


DINOTHERIUM. 


This genus was first described from India by Falconer *, who 
named a specimen from Perim D. indicum. Its specific characters, 
according to him, lie in the shape of the jaw, there being no 
difference from D. giganteum in the teeth except a greater thick- 
ness of the enamel. 

A second form, smaller than the first, from Attock was men- 
tioned, but not definitely named by him. This was subse- 
quently called D. pentapotamie by Lydekker 7. In the Catalogue 
of Fossil Mammalia in the British Museum, presumably his 
final opinion on the subject, he included the two forms under the 
name indicum, and stated that many of the differences previously 
used by him had proved to be individual variations. 

Another ‘ species,” D, sindiense (Lydekker), is too fragmentary 
to afford any evidence of a specific character. 

More recently Pilgrim has mentioned this genus. In the 
Records § of the Indian Geological Survey he names a form from 
Baluchistan D. naricwm, there stating that ‘it differs very 
markedly from the other known species,” and giving certain 
characters. In his Memoir||, however, he withdrew this name 
and made the form a variety, gajense of D. pentapotamic, although 
Lydekker had already shown cause for making the latter the 
same as D. indicum. 

Presumably Pilgrim’s reasons for making his form a variety 
were those by which he was at first inclined to make it a species, 
i.e. the fact of the ridges in the last lower molar not being parallel 


* © Walconer’s Memoirs,’ vol. i. p. 408. 

+ Pal. Ind. ser. x. vol. 1. pt. 2, p. 72, pl. ix. 

+ Pt. iv. p. 11, and footnote. 

§ Pilgrim, Rec. Geol. Sury. Ind. vol. xxxvii. p. 156 (not vol. xxxviii. as quoted by 
Pilgrim in his Memoir). 

|| Pilgrim, Mem. Geol. Surv. Ind., n. s, vol. iv. no 2 (1912). 


PROBOSCIDIA FROM BALUCHISTAN. 62] 


to one another, but diverging from within outwards, ete. In the 
same paper he gives characters in which he considers the Indian 
forms differ from D. giganteum. These are :— 
1. The longitudinal ridge connecting the transverse crests in 
the molars is very much stronger in the Indian forms. 
The present writer is unable to see this difference in speci- 
mens in the present collection, nor can the published figures 


x. 


Text-figure 8. 


Lower jaw of left side in side view. 


Dinotherium sp. 


‘jh : an 


Wire 
Yy ; 


Yj al 
0 thule if lye 


i 
ut L Pa 


a 


ain 
ae cn 


of Indian forms be said to show any great difference in this 
respect from Huropean specimens in the British Museum 
collection. 


622 MR. C. FORSTER COOPER ON MIOCENE 


2. The transverse valley in the second upper molar is com- 
pletely blocked on the inner side, and 

3. The inner posterior column of the third upper premolar is 
more isolated than in gigantewm, im which a ridge in continuation 
of this column extends right across the valley. 

These two characters seem to be within the limits of varia- 
tion of European forms. <A third upper premolar from 
Darmstadt (B.M. collection, No. M. 3494) has the column 
quite isolated. 


$m 


Text-figure 9. 


Lower jaw of left side in surface view. 


<= 


<== 


Dinotherium sp. 


oS 
“Wo S 
ff aK Wes 
LU 
y. 


The material of the present collection from the Bugti beds is, as 
is the common experience, both scanty and fragmentary, and 
consists only of part of a lower jaw and some separate upper and 
lower teeth. The lower jaw (text-figs. 8 & 9) belongs to a small 


PROBOSCIDIA FROM BALUCHISTAN. 623 


form, and in size and structure compares closely with D. hobleyi *. 
The third ridge of the last molar is not parallel to the front ones, 


but slopes at an angle similar to that found in D. hobleyi and 
D, levius. 


Of the separate teeth, none shows any great difference from 
specimens of European forms with which it has been compared. 
An upper molar, probably a second, is here figured (text-fig. 10). 


Text-figure 10. 


———— 
ee 


anrne: ——oae 
QQ’ f — 


Dinotherium sp. Second upper molar. A. Surface view. B. View from 
inner side. C. External side. I. Posterior view. X 3. 


There is nothing in it which can be construed as a “ longitudinal 
ridge,” nor is there anything in the nature of a ‘‘ cusp blocking 
the valley” which has not been found within the range of 
variation of Kuropean specimens. A third upper premolar 
(Pl. IV. fig. 6) is also figured to show the separation of 
the inner cusps said to be characteristic of the Indian forms, 


* Andrews, P. Z. S. 1911, 


624 MR. C. FORSTER COOPER ON MIOCENE 


but which, as has just been stated above, occurs also in 
D. gigantewm. ‘he remaining teeth in the collection are not 
distinguishable from the smaller forms from Hurope, and call 
for no comment except for three specimens. Of these, one appears 
to be an anterior lower milk molar (Pi. IV. fig. 4 and text-fig. 11), 
a tooth which apparently has not so far been noticed. It is 
24 mm. long and 21 wide, and consists of two ridges, one running 
from the middle of the inside border round the front and along 


Text-figure 11. 


Dinotherium sp. Anterior lower milk molar, internal view from side. X 1. 
the whole outside border, but being interrupted by a small notch 
towards the front of the outer border; the second ridge runs 
transversely across the hinder part of the tooth from a high cusp 
on the inner side towards the outer border. The tooth is pointed 
in front, wide behind, and triangular in ground-plan. Posteriorly 
there is a small cingulum shelf and anteriorly a small cingulum 
cusp. 

Text-figure 12. 


Sp. incert@ sedis. Side view of tooth, the anterior end pointing to the right. x 1. 


The next tooth (Pl. IV. fig. 5) is apparently a third upper 
premolar, but of unusual form. It is partially worn, and shows on 
one side two low cusps not touched by wear; opposite these 
are two large irregular areas of wear. There is no cingulum 
around these cusps, but a small one at each “end” of the tooth. 
In the plate the two low cusps are placed in the same position as 
the ectoloph of the more normal premolar (broken in this speci- 


PROBOSCIDIA FROM BALUCHISTAN. 625 


men) (Pl. LV. fig. 6) when the main differences appear to be the 
low ectoloph and general deficiency of cingulum. Probably it is an 
abnormal tooth. The third specimen (PI. LV. fig. 7 and text- 
fig. 12) is even more problematic. It is bilophodont, 36 mm. 
long and 28 mm. broad; each ridge consists of two cusps joined 


Measurements of specimens, in millimetres. Approximate 
jigures are marked with an asterisk. 


{ 
| Length. | Breadth. 


{ 


BUNOLOPHODON ANGUSTIDENS. 
Lower teeth. 


3rd molar. Baluchistan (text-fig. 3 and Pl. II. fig 8 Db ayeke erent 148 | 64 
< a (Gextafloy4) ae: : Heuee wanes 162 67 
vi . (text-fig. 2), in jaw Aen eRe ie reece shh el O Sila ns Ose 
fi damayed specimen Pont ianire coach 143* | 64% 
¥ Auvergue. B.M. No. M5975 SE Sapecee ee A ba LOO. 68 
‘ Sansan. BM. No. 40729 . S OeAT BOC UE Oto Ses CoM ENS CONT eo 68 
y bs B.M. No. 37242 a ... 180 73 
a Es Cambridge Museum (PI. TOE, ‘fig. ee Pas 135 64 

2nd molar. Baluchistan (text-fig. 2), IN jaw oe eee | 99 55 
i Siamegam, | IBWS NGS APD “a.cccocspcotonasacsbonouedteadl| Og I GB 
a - IAW INO, COPED acoscoecoive .obsede ona sokauseuel | Gey Whe) GIL 
i 3 Bev No ySvo42ah we ey 05) In 162 

stimoelars Baluchistant (GextatloaG)ieeaceeeseree ses see cece seeeeeene | 78 | 45 
i a PODBRED ER Ceca eoO Senet remot ol; We Main Mets 
“ Shimane | Iai MLS IN@s4O7Gs— aon ccocnoeas ssdntocbeonobes « 82 | 49 
Fe Simo, BIW IN@s BOGAl sonosaacpaonnocosscs Gcecdoconll CL AB} 
- ? India. BEM INOg MAD SOS ie met ets oho eae ee Ake aie 

Upper teeth. | | 

3rd molar. Baluchistan (Pl. I. fig. 1) . a aa MOB 7) 
re Shes Eo OSS Ges ee et GD I FO 
ee Chevilly. B.M. No. 7424, a cast . Aesop lleme napa ie dUsseh i iiT he) 

Ynd molar. Baluchistan, older palate ............... 2.00. ecg 95 5 
A 4s HOMO? WHEE soason sob osonbobedosouda séee 98 57 
a i (BIST akon: SPR Tiiate oI Tie 66 

Ist molar. Baluchistan, younger palate .................cc0..--2 00 a9 49 


The width of the palates between the front of the second 
molar is about 65. 


DINOTHERIUM sp. | | 

Third lower molar .... Peat welll 56 
Length of fragment. of lower jaw v (text-f figs. 8 & i 9)... alts OO) im| 

First upper molar . Sree te 64 52 

Second upper molar ‘(text- fig. 10) . ass aed ct as to dL ae 63 62 

Avan) Whey eee WACIENP! /c.coace. apidboboodsboriubs soa noaben\iaenone she seanabede Bomace TO. 1. BB 


together transversely. ‘The anterior ridge is thick and somewhat 

Rated in front, the posterior ridge is Sain and slightly concave 
in front, the enamel i is smooth and there is no trace of cingulum. 
It has been compared with all forms of bilophodont teeth in the 


626 ON MIOCENE PROBOSCIDIA FROM BALUCHISTAN. 


collection of the British Museum without result*. It is certain 
that it does not belong to any known form of Dinotheriwm, and 
yet it is to teeth of that genus alone that it can be compared. 


SUMMARY. 


The Proboscidia of the Bugti beds of Baluchistan consist of 
a form of Bunolophodon angustidens and a small Dinotheriwm. 

The Indian B. angustidens is of a small type more primitive 
than the French forms from Sansan, nearer to Schlesinger’s form 
subtapiroidea than his form typica, but not identical with it. 
Lydekker’s subspecific name var. palwindica may be retained for 
the present until intermediate forms are found to occur. The 
genus Hemimastodon is not supported by material in the 
collection. ) 

The Dinotherium is not found to differ from the smaller 
European forms nor from D. hobleyi of Africa, and it is doubtful 
in the present state of knowledge whether the Indian Dinotheres 
are separable from the European. 


EXPLANATION OF THE PLATES. 


Prate I. 
Fig. 1. Bunolophodon angustidens. Third upper molar. 
2. mn Second upper molar. 
” pp 
Both figures are reproduced about } natural size. 
Prats II. 
Fig. 1. Bunolophodon angustidens. Third lower molar. 
2. . 5 Incomplete specimen from Sansan. (B.M. No. 
32533.) 
Both figures are reproduced about $ natural size. 
Prate III. 
Fig. 1. Bunolophodon angustidens. Third lower molar from Sansan. (Cambridge.) 
2. iy (angustidens*). ‘Terminal part of a third lower molar. 
Both figures are reproduced about 2? natural size. 
Prats LY. 
Fig. 1. Bunolophodon angustidens. Unworn first lower molar. 
2. uv A Fourth upper premolar. 
3. ES 60 Third upper premolar. 
4. Dinotheriwm sp. Anterior milk molar. 
5. » Third upper premolar. 
6. » » 2 Third upper premolar of unusual type. 
7. Sp. incert@ sedis. ? An anterior premolar. 


All figures are reproduced a shade under natural size. 


_* Such as Listriodon ete. India has produced animals with very unusual denti- 
tions such as Tetraconodon; the present specimen may belong to yet another 
strange form, 


I ZA, So W922, TSUN RIMOS IR, JP lls 


CEPHALORHYNCHUS COMMERSONII. 


3 


CEPHALORHYNCHUS COMMERSONII AND C. 


HEAVISIDEI. 


P. Z. S. 1922, HARMER, PI. Ill. 


SPECIES OF CEPHALORHY NCHUS. 


SIR SIDNEY HARMER ON COMMERSON’S DOLPHIN. 627 


32. On Commerson’s Dolphin and other Species of Cephalo- 
rhynchus. By Sir Stpney F. Harmer, K.B.E., Se.D., 
F.R.S., F.Z.S8., Director of the Natural History Depart- 


ments of the British Museum *. 


(Plates I-III. 7) 
[Received May 17, 1922; Read May 23, 1922. ] 


The literature of the Cetacea includes numerous specific names 
which are of little use to science, owing to a want of knowledge 
of the animals to which they were originally applied. Many 
descriptions are based on external features alone, while others 
depend exclusively on osteological characters. The correlation 
of these two kinds of description is difficult and often impossible, 
and there is probably no group which includes a larger proportion 
of doubtful species than could be found in a complete list of the 
names which have been given to Cetacea. 

Under these circumstances I have felt much satisfaction in 
being able, as I think, to rescue one of these doubtful species 
from its present position. In 1804, Lacépéde (pp. xliv, 317) gave 
a brief description of a Dolphin which had been observed by 
Commerson, near Tierra del Fuego and in the Straits of Magellan, 
during Bougainville’s voyage round the world. _Commerson’s 
MSS., addressed to Buffon, by whom they were sent to Lacépede, 
included a diagnosis of this Dolphin :—‘‘Tursio corpore argenteo, 
extremitatibus nigricantibus.” It is further stated that’ the 
black colour appears only on. the “‘ extremities,’ and that the 
back and almost all the surface of the animal shine like a 
polished surface, white and, so to speak, silvered. These 
Dolphins, somewhat inferior in size to the Common Porpoise, 
were observed, during the Southern summer, playing round 
the vessel, and were described as being among the most 
beautiful inhabitants of the sea. They were given the name of 
‘* Le jacobite,” obviously in allusion to their striking black and 
white colour. 

Quoy and. Gaimard (1824, p. 87) mention a Dolphin, ‘“ moitié 
blanc, moitié noir, 4 museau peu aiongé,” which they saw at the 
Falkland Islands. A specimen was killed, but it sank imme- 
diately. Lesson (1827, p. 181) states that during the voyage of 
the ‘ Coquille’ they only once observed, at the Falkland Islands, 
the Black-and-white Dolphin of Quoy and Gaimard, and that every- 
thing authorised him to think it was the Delphinus commersonii. 


*° Published by permission of the Trustees of the British Museum. 
+ For expianation of the Plates, see p. 638. 


Proc. Zoon. Soc.—1922, No. XLII. 43 


628 SIR SIDNEY HARMER ON COMMERSON’S DOLPHIN 


Other authors of the early part of the last century are unable to 
do more than quote the original account, although Gray (1846, 
p- 30) surmised that this Dolphin might be identical with his own 
Beluga kingit. 

The external characters of the species which forms the special 
subject of this paper were admirably figured by Moreno (1892, 
pl. ix.), who described it (p. 385) under the name of Lageno- 
rhynchus floweri, and gave satisfactory figures of the skull 
(pl. viii.). It was first observed by him, in October 1874, in the 
bay of the River Santa Cruz, which is on the Patagonian coast, 
almost in the same latitude as the Falkland Islands. It was 
seen in hundreds on certain occasions, but no specimen could at 
first be obtained, although a female, the original of the three 
figures of the larger individual shown in pl. ix., was secured from 
the same locality in December 1876, and two other specimens 
were obtained from Tierra del Fuego or Santa Cruz in 1884, 
The colour-markings were identical in all the specimens. The 
individual whose external measurements are recorded was 138 em. 
in length, and the total length given for the skull is 28 em. 

It is unfortunate that Moreno’s specific name, which was based 
on a good and well-illustrated account of external and cranial 
characters, cannot be accepted. It must be regarded, I think, 
as a synonym of Delphinus commersonu, which, although 
described inadequately and without illustrations, appears to be 
perfectly recognisable in the light of the evidence now available. 
Not only were Commerson’s and Moreno’s specimens from 
substantially the same locality, but the silvery colour, with black 
confined to the ‘‘ extremities,” described by the earlier observer, 
is equally noticeable in Moreno’s figures, which represent an 
animal mainly white, with the head, tail, flippers, and dorsal fin 
black. 

The only other actual description of this animal with which I 
am acquainted is that of the late Dr. W. 8S. Bruce (1915, p. 500, 
pl. i.), who shows the external characters of a specimen, 4 ft. 4 in. 
in length, which was obtained by him in 1893 at Port Stanley, 
Falkland Islands. It was referred, doubtfully, to Lagenorhynchus 
eruciger (d’Orb. & Gerv.), and it was styled the ‘ Piebald 
Porpoise.” Dr. Bruce records a number of positions to the 
south of the Rio de la Plata at which Dolphins believed to be 
‘‘ Piebald Porpoises ” were observed from the ship. 

Tt is shown by the records of Moreno and Bruce, as well as by 
the new evidence now submitted, that a Porpoise characterised 
by its strongly contrasted black and white colours arranged in a 
very definite pattern, is common in the neighbourhood of the 
Falkland Islands and of Tierra del Fuego. I think there can 
be no doubt that this Porpoise belongs to the same speeies as 
the ‘ Jacobites” which played round Bougainville’s ship in the 
Straits of Magellan, and were observed later at the Falkland 
Islands by Quoy and Gaimard, and by Lesson. 


AND OTHER SPECIES OF CEPHALORHYNCHUS. 629 


Liouville (1913, p. 165), in a discussion, under the heading of 
Lagenorhynchus fitzroyi, of certain Dolphins which have been 
recorded as L. cruciger or by other names, comes to the conclusion 
that a number of conspicuously marked Southern Dolphins, 
recorded under various specific names, are colour-varieties of a 
single species. Whether this view be correct or not, the con- 
cordant evidence derived from several distinct sources, with 
regard to Commerson’s Dolphin, shows that this species at least 
is remarkably constant in the position of its areas of black and 
white. Jt may be suggested that these striking markings have 
the protective value which has been ascribed to the “ disruptive” 
type of coloration in animals. As seen in the water, the white 
area probably divides the body into two parts which seem to 
have little, if any, connection with one another. The Dolphinis, 
in fact, effectively camouflaged, and perhaps the protection is 
specially successful in water lable to contain floating ice. The 
principal enemy of Commerson’s Dolphin is probably Orcinus, 
the Killer Whale. 

The British Museum has recently received the following 
material, referable to the genus Cephalorhynchus, in which 1 
place Delphinus commersoni Lacép. (= Lagenorhynchus flowert 
Moreno and ZL. eruciger Bruce, nec auctt.) :— 


A, An eviscerated male specimen (in salt), 4 feet 6 inches long, 
captured in Stanley Harbour, Falkland Islands; presented by 
Mr. J. HE. Hamilton, Magistrate of West Falkland, who stated 
that the stomach Contained “krill” (the Norwegian whalers’ 
name for the small Crustacea which form the food of Whalebone 
Whales) and the pens of cuttlefish. The vertebral epiphyses are 
quite free, and the animal was accordingly immature. 

B. A skull, with lower jaw, picked up on the shore of Byron 
Sound, West Falkland ; presented by Mr. Rupert Vallentin. 

C. Two photographs of a specimen stranded in the Falkland 
Islands on another occasion; presented by Mr. Rupert Vallentin, 
who stated that these Porpoises are fond of basking in the beds 
of Macrocystis when the sun is shining, and that he knew of no 
other species in that locality. 

D. A note-book, lent by Mr. Lionel E. Adams, containing 
observations on Dolphins observed by him from a ship in the 
Straits of Magellan, together with a small photograph showing 
two of them in the air during a jump out of the water. The 
note-book included an excellant sketch, completely agreeing with 
A and ©, and the passage “Jet black and pure white (no 
‘““shades). I did not see any variation in the markings of one of 
“the hundreds that played about the ship.” 


The following notes were made on Mr. Hamilton’s specimen 
(Pl. I. figs. 1 & 2) before its skeleton was prepared. The skin, 
although somewhat abraded, showed the external coloration 
perfectly distinctly. 

43* 


630 SIR SIDNEY HARMER ON COMMERSON’S DOLPHIN 


Colour.—Jet-black and pure white, the principal white area. 
shading to some slight extent through grey to the black areas in 
front and behind (as in Moreno’s pl. ix. fig. 1), but the junctions 
of the two colours otherwise very sharply marked. The black 
vecurred on the whole of the head and lower jaw (interrupted 
ventrally by a pear-shaped, median, white area on the throat), 
extending backwards obliquely from the sides of the head to the 
pectoral fins, which were black on both surfaces, and was 
continued across the ventral side as a broad band, behind the 
white marking on the throat, this band giving off an acutely 
pointed median prolongation backwards, between the flippers. 
The tail, including the flukes on both surfaces, was black all 
round, for nearly a foot from its emargination, and this colour 
was continued forwards obliquely on the back’ beyond the dorsal 
fin, but separated by a white interval even in the mid-dorsal line 
from the black of the head. A large heart-shaped black area 
surrounded the reproductive opening, and was observed by 
Mr. Hamilton in another male specimen examined by him. The 
white part of the skin consisted of (a) the median ventral area on 
the throat, (6) the main white area encircling the body obliquely, 
much more developed ventraily than dorsally, and including 
most of the ventral and lateral regions of the body. 

Mr. Vallentin’s photographs (Pl. I. fig. 3; Pl. I. fig. 1), one 
of which shows that the specimen was a male, correspond in 
every essential detail with the above description. Mr. Adams's 
sketch (D), which represents animals seen in the sea, agrees closely 
with the others, but from the conditions under which it was 
made, it is not surprising that it does not show any black area 
round the reproductive opening. His photograph shows two 
living animals in the air, as seen from the ship, and it suggests 
the idea of a Porpoise “ moitié blanc, moitié noir,” as described 
by Quoy and Gaimard. 

Neither Moreno nor Bruce shows a broad black marking round 
the reproductive opening. Moreno’s ventral view indicates, 
however, a narrow, longitudinal, black mark in this situation, its 
posterior third being nearly constricted off from the front 
portion. The specimen was a female, and the question arises 
whether the individuals of this sex normally have a ventral black 
marking narrower than that of the male. One of Bruce’s figures 
(the lowest in the Plate) of an animal .of unrecorded sex gives 
some indication of a constricted, narrow marking like that shown 
by Moreno. 

External form.— The head of A had no beak distinctly outlined 
from the remaining portion, but was nearly conical. The dorsal fin 
had an elongated base, and was low and rounded, with but little 
indication of a faleate shape. ‘The flippers were also more 
rounded at their ends than in most Dolphins. These characters 
point to Cephalorhynchus (cf. True, 1889, pp. 108, 176), and this 
conelusion is confirmed by the cranial characters. 


AND OTHER SPECIES OF CEPHALORHYNCHUS. 631 


Eeternal measurements of Mr. tLamilton’s specimen (A) :— 


Cn. 
Rota e bla ein cee a sciacasas epee eset 13 
Dorsal fin (middle) to tip of snout ......... 81 
Ge) LO emanroimahionof tails | Odo 
Fe siti llerak@ iano LOR NSIS" Kecadernecbaseamuedan 22°8 
Flipper, length, HAIN PMUAh toePeadaneosood ser 16:5 
45.0 INDUICIENOIP NOVACIEE |” Spe soeeneune 23°5 
Gape, length of, from tip of upper jaw ... 16°5 
Snout to blow- Ti] =e hic eset a vote od 20°3 
Tip of lower jaw to (middie of) umbilicus.. 64 
Umbilicus to reproductive aperture f ...... 14 
Reproductive aperture te anus............... 18°5 
Anus to emargination of tail ............... 40°5 


The characters of Cephalorhynchus Gray were re-defined by 
Flower, in his well-known paper on the Delphinide (1883, p. 473), 
the type-species being Delphinus heavisidii Gray, 1828. In his 
Review of the Delphinide, True (1889, pp. 108, 176) recognises 
four species of the genus—the type-species from the Cape of Good 
Hope, C. albifrons True and C. hectori Van Beneden from 
New Zealand, and C. ewtropia Gray from Chili. In 1893 and 
1896 several new Dolphins from Chili were described by R. A. 
Philippi. His two papers were later criticised by True (1903), 
who expressed the opinion that Phocena or Tursio albiventris 
(Perez, MSS.) Philippi, 1893, 1896, is a synonym of C. eutropia, 
and that Tursio platyrrhinus Philippi, 1896, is probably the 
same species. Tursio? panope t Philippi, 1896, was not thought 
by True to be referable to Cephalorhynchus, but it seems possible 
that it may also belong to this genus. 

The external char Meiers have been described 1n Cephalor hynchus 
heavisider, C. albifrons, C. hectori, and C. albiventris. C.eutropia, 
C. plaiyrrhinus, * C. panope, and ? 0. chiloénsis are known from 
cranial characters alone. 

The British Museum possesses the type-specimens of C. ew- 
tromias and C. heavisidet. The skull of the latter was inside the 
mounted skin when Flower wrote his revision of the Delphinide 
in 1883. It has since been removed, and although defective 
in the occipital region, it is otherwise in good condition. 
The following table gives measurements of the skulls in the 
Museum :— 


* The measurement recorded by Moreno is 188 cm., and that given by Bruce is 
132°2 cm. 
+ The distances between the apertures were in each case measured from their 
peddle points. 
£ Philippi (1901) later described a new Dolphin, Tursio ? ehiloénsis, from Chili, 
Ww hich he stated to have resemblances to 7’. NODE: 
§ Figured by Gray 1846, pl. xxxiv. figs. 1, 2; see also Gray, 1866, p. 262. 


632 SIR SIDNEY HARMER ON COMMERSON’S DOLPHIN 


C. commersonii. 


Cf. Flower. 


| P.Z.S. 1883, 


a 
iva} se 
Dade ie p- 506. 
aS 3s -a ai 
Sh hee Wee he 
Setar ice oe 
ares & & aa isi 
OF | ais ae fee 
Met. peciauienrs= 
| 
i otal lenethi(centimetres)).....065-24..-2.41 | 80%0 29°8 29:5 | 29:0 
Do INOSAAADT, WOVEN: si oonodd sbooao ond abodaoven sacl, LUOes) 14:0 140 13:2 
3. a width at base .. al SES 73 65 6°8 
4. = » Opposite “2nd tooth | 
trom hind end.. [Deal 65 6:2 59 
5. a » at middle .. 7:0 53 5:2 51 
6. a vertical depth, at middle “of | 
WOOWETRONY Gonsabiodends con nb 55> alee 16 16 11855 
7. Tooth-row, length of . le USES} 12:3 12:3 116 
8. Maxilla, right, posterior expansion “of, 
UGTYESID arolons coe 105 10°9 11:2 10°4, 
9. Do. do. width (middle 
of orbit to nares) 65 672 60 | 51 
10. Do. width at posterior end of | 
tooth-row.. 22 16 IES) 3: 
11. Premaxilla, right, width at posterior 
end of tooth-row Ne7/ 18 a | 1°8 
12. » Width behind nares .. Me ae a 12 
13. Orbital width (at front of eh bacedean awl 12°7 12:4 12:9 
14. Glenoid width ......... Ho AGP 7/ 14°6 14°38 14°5* 
15. Greatest height of skull .. 14°83 13°2 13°2 13°0* 
16. Mandible, length .........00-000000c. | 30:0 23°0 23:0) Wile 2258 
lyfe Bs condyle to posterior end of | 
tooth-row ..... sana MALES 108 1a One 11:0 
18, 4s depth, at middle of ae Pane: |) 688 4:8 4:6) | 43 
19. a3 » at posterior end of | 
toobh=vow, 50. -2. +: | Bea 33 2:9 2°6 
20. AF » at middle of tooth- | | 
row |. 22 1:9 17 15 
PERCENTAGES. 
11 ANayeenll Mkesayed tl Nisan sopeuaroonccsek sobon somo dete | 100°0 1000 100:0 | 100°0 
D, ROTHPOT, NEVA 1... s6yb5udee v0 doboso neo sus oc: 63-71 47-0 475 | 45°5 
4. a width opposite 2nd tooth | 
fron behind gees ae) Rome? 21°8 21:0 | 20°4 
7 Mooth-rows lengthiorh eae an) 46:3 41-2090) Ale 7a OO 
8. Maxilla, right, posterior expansion, | | 
Sasa EN he Ca Ne Gea) 365 38:0 | 35°9 
9. Maxilla, right, “posterior expansion, | 
width... 179 20°8 |) 20d aalaaG 
18. Mandible, » depth ‘middle of. f part b ‘behind 
teeth . ; 16:0 16:1 ssX6) | 148 


* Estimated (skull imperfect). 


The external characters of Mr. Vallentin’s Dolphin were not 
definitely known, although he himself had no doubt that the 
skull belonged to ‘the species shown in his photographs. Com- 


AND OTHER SPECIES OF CEPHALORHYNCHUS. 633 


parison of the skull with the other Falkland Is. specimen 
confirms this conclusion. The only difference of importance is 
that the rostrum of Mr. Vallentin’s specimen (B) is distinctly 
narrower at its base than is that of Mr. Hamilton’s skull (A). 
Flower (1883, p. 469) stated that the rostrum becomes longer 
and wider, in proportion to the brain-case, in older Dolphins. 
The skull B has an unfinished appearance in the region of the 
base of the rostrum, as if growth were there incomplete; and 
the notches in the maxille are wider than in A. The two skulls 
are practically alike in their orbital width, and by an addition to 
the rostrum at its base, B would become similar to A in the width 
of this structure and in the reduction of the maxillary notches. 

C. eutrepia and Philippi’s three species albiventris, platyr- 
rhinus, and panope agree with one another in having a rostrum 
which in length exceeds 50 per cent. of the total skull-length. 
The tooth-row is also relatively long in the same species, the 
percentage in C. ewtropia (type) being 46°3, as shown by the 
preceding table, and ranging from 46:0 to 51°5, as shown by 
Philippi’s measurements, in the other three species. C. commer- 
sonti agrees with C. heavisidei, and differs from the other four 
species, in having a rostrum less than half the length of the 
skull, and a shorter tooth-row, not exceeding 42 per cent. of the 
skull-length. The skulls of the two species are also absolutely 
smaller (29-30 cem.), C. ewtropia having a skull more than 36 cm. 
long, and those of Philippi’s three species ranging from 33:0 to 
39°0 cm. 

Although C. commersonti has a considerable resemblance to 
C. heavisidet in cranial characters, | have noted the following 
differences in the skulls available for comparison (PI. IT. figs. 2 
& 3). The skull of C. heaviside: has previously been figured by 
Schlegel and by Van Beneden and Gervais, as pointed out below; 
and that of C. commersonii by Moreno (1892, pl. viii.). 

Premaxilla.—The outer edge of this bone, in Cephalorhynchus 
generally, forms a prominent elevated ridge in front of the 
anterior nares (Flower, 1883, p. 473). In C. commersonii the 
left bone is not prolonged behind this ridge, that of the right 
side forming a slender splint passing towards the nasal of its 
side, but not reaching it. Moreno’s fig. 1a seems to agree in 
this respect. In C. heavisidet this posterior process is consider- 
ably larger in the left premaxilla, and still larger in the right 
bone, where it is 12 mm. wide and touches the nasal, passing 
back to about the middle of its length. C. ewéropia resembles 
C’. heavisidei in these respects, except that the right premaxilla 
does not reach the nasal, from which it is 11 mm. distant. The 
floor of the depressed triangle in the premaxille, in front of the 
anterior nares, is flatter in C. commersonii than in C. heavisider, 
where it is slightly concave. The width of the premaxille in the 
front half of the triangle is distinctly greater in C. heavisider 
than in the other species, and the maxille are here corre- 
spondingly narrower. 


634 SIR SIDNEY HARMER ON COMMERSON’S DOLPHIN 


Mawilla.—C. commersonii seems to be characterized by the 
specially large posterior expansions of the maxille. Measuring 
the length of these expansions from the bottom of the maxillary 
notches along a line parallel with the middle line of the skull, 
and the breadth as indicated in the table of measurements, these 
parts are absolutely longer in C. commersonu than in the con- 
siderably larger type-skull of C. ewtropia. In C. heavisided they 
are relatively narrow, and do not completely cover the orbit, the 
whole upper margin of which is visible in a dorsal view, with 
part of the postorbital process, which slopes a good deal 
outwards. 

In C. commersonii, the front of the orbit is completely con- 
cealed by the maxillary expansions, the only part visible dorsally 
being the postorbital process, which is more vertical than in the 
other species. Rather more of the upper surface of the orbit is 
shown in Moreno’s figures, particularly in fig. 2, which is 
described as representing a younger specimen; and the extent of 
the overgrowth of the orbit by the maxilla is perhaps dependent 
on age. 

In ©. heavisidet there is a slight constriction of the outer 
margin of the rostrum, 20 mm. in front of the maxillary notches, 
so that a small lateral lobe is partially marked off at the base of 
the rostrum, on each side. ‘These lobes do not occur in 
C. commersonie. 

Nasal.—These bones, though no doubt variable, show certain 
features which may be distinctive. In C. heavisidec they are 
subdiscoidal, the central part the thickest and the margin thinner, 
and they are in contact with one another, on the upper side only, 
for about 7 mm., leaving a small triangle of the. frontals un- 
covered between their lower borders and the mesethmoid. The 
greatest diameter is 25mm. In C. ewtvopia each bone is quite 
twice as wide as long (25, 11 mm.), and has’a sharply marked, 
nearly vertical, anterior face, which almost reaches the meseth- 
moid. The left nasal is wanting in skull B of C. commersonii, 
and the right nasals are not altogether alike in the two skulls. 
The shape seems to be somewhat intermediate between those 
of the other two species, being more quadrangular than in 
CO. heavisidet, and rather longer in proportion to the width than 
in C. ewtropia, the two bones having a median suture nearly 
reaching the mesethmoid. The right nasal of skull A measures 
22 mm. in width and 17 mm. in length. Moreno’s fig. la gives 
similar evidence. 

Mandible (P1. ILI. fig. 2).—While differing in certain respects, 
all three species show, more or less distinctly, the outwardly bowed 
rami, corresponding with the expanded part of the rostrum, 
given by True (1889, p. 108) as a generic character. That of 
C. eutropia (c) is much larger and heavier than those of the other 
species, but it resembles the mandible of C. commersonii (6) in 
general form and in the shape of its coronoid process, which is a 
blunt triangle with its apex directed nearly vertically and lying 


AND OTHER SPECIES OF CEPHALORHYNCHUS. 635 


well in front of the condyle. In C. heavisidei (a) the coronoid 
process is directed backwards, so that its apex lies but little in 
advance of the condyle, while from its front end the upper edge 
of the jaw slopes down to the teeth, instead of remaining at first 
horizontal, as in the other two species; and the jaw is slighter 
and has less vertical depth than in them. 

Pterygoid.—The importance of these bones in distinguishing 
genera in the Delphinidz was pointed out by Flower (1883), who 
relied on Van Beneden and Gervais’ figure (pl. xxxvi. fig. 1a) 
for evidence as to Cephalorhynchus, and reproduced it on p. 473. 
There can be no reasonable doubt that the figure was drawn from 
a skull with incomplete pterygoids, and it is thus to some extent 
nusleading. The type-skull of C. heavisidet resembles the figure 
as reproduced by Flower, in respect of the pterygoids, but a 
comparison with the two skulls of OC. commersonii, in both of 
which these bones are complete, shows that they are much 
longer than would be inferred from that figure. They resemble 
in shape the pterygoids shown by Flower (1883, p. 471) in Globi- 
cephala (Globiceps), but their posterior margin is more oblique, or 
less nearly transverse. In skull A (Pl. II. fig. 1), the palatal 
surface of each pterygoid measures 36 mm., from the point where 
it leaves the vomer, to its posterior tip. In their free, posterior 
portions the two bones are separated by a narrow, f-shaped 
interval, 18 mm. long and widening to 8 mm. across, at the com- 
mencement of the posterior oblique borders. Skull B is similar, 
except that the interval between the free parts of the pterygoids 
is A-shaped, and the length to the tip is only 30 mm. The 
pterygoids are incomplete in Moreno’s see 


Teeth.—Skull A (C. commersonit) has = om = 


out cutting away any of the gum. One or two small ones may 
perhaps be present, beneath the gum, at the anterior end of the 
series. In skull B, in which many of the teeth are wanting, 
the number is about 29 on each side of each jaw. In C. heavisidet 


Eaten visible with-~ 


(type), the number is = = 


30-31 on each side of each jaw. 

The osteological features of the Cetacea are so variable that it 
is often impossible to be sure how far characters selected from an 
examination of a limited amount of material are of any value. 
It is thus important to point out that, in the features enumerated 
above, the two skulls of C. commersonii are in substantial agree- 
ment with one another, except where attention is called to a 
difference. There is evidence, moreover, that other specimens 
of C. heavisidei agree with the type-skull. The skull of this 
species has been figured and briefly described by Schlegel (1841, 
Poo leg pl. it etioss IAL ple tvs tie. 6)))s ; and by Van Beneden and 
Gervais (1868-1879, p. 999, pl. xxxvi. figs. 1-16), from a skull in 
the Paris Museum. The etter authors: were inclined to regard 
the specimens of Cephalorhynchus from various southeri localities 


while in C. eutropia (type) it is 


636 SIR SIDNEY HARMER ON COMMERSON’S DOLPHIN 


as belonging to a single species. I have to thank Prof. R. 
Anthony for the information that the skull figured in the 
‘ Ostéographie’ (figs. 1, 1 @) is from the Cape of Good Hope, 
Dussumier, 1827, although he thinks that the lower jaw figured 
(16) may not be from the same specimen. It may no doubt 
be assumed that Dussumier’s specimen at least belonged to 
C. heavisidet. 

The following may be noted as points of agreement between 
the type-skull of C. heavisidet and Schlegel’s figures :—The small 
lateral lobes at the base of the rostrum on each side; the narrow 
posterior expansions of the maxille, not completely covering the 
orbit; the shape of the nasals, which do not quite touch one 
another and leave a triangular part of the frontals exposed between 
themselves and the mesethmoid; and the broad posterior ends of 
the premaxille, both of which nearly reach the nasals. In 
Van Beneden and Gervais’ figure most of these resemblances are 
also noticeable, and attention may be specially directed to the 
right premaxilla, which reaches the nasal behind. 


CEPHALORHYNCHUS Gray, 1850 (see Flower, P. Z.S. 1883, p. 473, 
and True, 1889, Buil. U.S. Nat. Mus. no. 36, pp. 108, 176). 


C. comMERSoNII Lacép. 

Delphinus commersonti, *‘ Le Jacobite,” Lacépéde, 1804, Hist. 
Nat. Cét. pp. xliv, 317 (referred to by Quoy and Gaimard, 1824, 
Voy. ‘Uranie,’ p. 87; and by Lesson, 1827, Voy. ‘ Coquille,’ 
jos MSs 

Delphinus commersoniti Desmarest, 1822, Mamm. 2° Partie, 
[Oo Ole 

Phocena commersoniit Lesson, 1827, Man. Mamm. p. 414. 

Lagenorhynchus flowert Moreno, 1892, Rev. Mus. La Plata, 
ill. p. 385, pls. viil., ix. 

Lagenorhynchus cruciger® Bruce, 1915 (nec auctt.), ‘ Piebald 
Porpoise,’ Scotia Rep. vol. iv. p. 500, pl. i. 


Known localities.—Tierra del Fuego, Straits of Magellan, coast 
of Patagonia, Falkland Islands. 

Haternal characters.—Length at least 140 em. Head conical, 
without distinet beak. Dorsal fin bluntly triangular, not falcate. 
Flippers rounded at the end, not faleate. Most of the sides and 
ventral surface silvery white, this colour extending completely 
across the back, between the head and the dorsal fin. A large 
white, median, pear-shaped area, widest behind, on the throat. 
Head, tail, caudal, pectoral, and dorsal fins jet-black (the fins on 
both surfaces), a narrow dorsal band of black passing obliquely 
from the tail to just beyond the dorsal fin, a broad black band 
extending across the ventral side between the flippers, produced 
backwards into a median, acuminate process. A black area 
surrounding the reproductive opening. White and black areas 
very sharply defined. Visible teeth 29-30 on each side of 
each jaw. 


AND OTHER SPECIES OF CEPHALORHYNCHUS, 637 


Skull.—Total length at least 29-30em. Rostrum about 47 per 
cent. of the total, at base half as wide as long, without pro- 
jecting lateral lobes posteriorly. Tooth-row 41-42 per cent. 
Posterior lobes of maxille large, almost completely covering 
the orbits; the postorbital process nearly vertical and hardly 
visible from above. Premaxille little prolonged behind the 
anterior nares, the floor of the depressed triangle flat. Nasals 
subquadrangular, meeting in a median suture, the width much 
less than twice the length. Pterygoids separated behind by a 
N- or A-shaped interval; the posterior border oblique, its tip 
30-36 mm. from the vomer. Upper border of mandible 
horizontal, behind the teeth, the coronoid process bluntly tri- 
angular, 


Literature. 


BENEDEN, P. J. VAN, and Gesryais, P. 1868-1879. ‘Ostéo- 
graphie des Cétacés.’ 

Brucr, W.S. 1915. “Some Observations on Antarctic Cetacea.” 
Scotia Reports, vol. iv. p. 491. 

Frower, W. H. 1883. “On the Characters and Divisions of the 
Family Delphinide.’’ Proc. Zool. Soc. 1883, p. 466. 

Gray, J. KE. 1846. Zool. ‘Erebus’ and ‘Terror.’ Vol. i. ‘ Ceta- 
ceous Animals.’ 

Gray, J. E. 1866. ‘Catalogue of Seals and Whales’ (British 
Museum). 

LackprpE. 1804. Hist. Nat. Cétacées. 

Lesson, R. P. 1827. Voy. ‘ Coquille,’ Zool. vol. 1. 

Liouviniz, J. 1913. ‘Cétacés de ’Antarctique.’ 2° Exp. Ant- 
arct. Frang. 1908-1910. 

Moreno, F. P. 1892. “ Noticias sobre algunos Cetaceos fésiles 
y actuales de la Reptiblica Argentina.” Rev. Mus. 
La Plata, ili. p. 383. 

Puiurppr, R. A. 1893. “ Los Delfines de la Punta Austral de la 
America del Sur.” An. Mus. Nac. Chile, 1? Secc. 
Zool. no. 6. 

Puiuiprr, R. A, 1896. ‘‘ Los eraneos de los Delfines Chilenos.” 
An. Mus. Nac. Chile, 14 Sece. Zool. no. 12. 

Puiuipri, R. A. 1901. “ Kine neue Art chilenischer Delphine.” 
Arch, f. Naturg. 1901, Bd. i. p. 276 (previously 
published in 1900, see Zool. Record, 1901, Mammalia). 

Quoy and GaAimarD. 1824. Voy. ‘ Uranie’ et ‘ Physicienne.’ 
Zoologie. 

ScHLEGEL, H. 1841. ‘“ Beitriige zur Charakteristik der Ceta- 
ceen.” Schlegel’s Abhandl., 1 Heft, p. 1. 

True, F. W. 1889. “A Review of the Family Delphinide.” 
Bull. U.S. Nat. Mus. no. 36. 

Trur, F. W. 1903. “ On species of South American Delphinide 
described by Dr. R. A. Philippi in 1893 and 1896.” 
Proc. Biol. Soc. Washington, xvi. p. 133. 


638 SIR SIDNEY HARMER ON COMMERSON’S DOLPHIN. 


EXPLANATION OF THE PLATES. 


Prats I. 
Cephalorhynchus commersonii. 
Figs. 1&2. Mr. J. E. Hamilton’s eviscerated specimen (A), lateral and ventral 
THOS 6 6 ge 


Vig. 3. Ventral view of another individual (C). From a photograph sent by 
My. R. Valentin. 


(Prats II. 
Cephalorhynchus commersonii. 


Rig. 1. Side view of specimen C. From a second photograph sent by Mr. R. 
Vallentin. 


Vig. 


bo 


. Skull of specimen A, dorsal view. X about +. 


C. heavisidet. 


Wig. 3. Skull of the type-specimen, dorsal view. X about 


CoH 


PrarE LT. 
Cephalorhynchus spp. 
Hig. 1. C. commersonii. Skull of specimen A, ventral view. > about i. 
ig. 2. Lower jaws, from the right side. X about 4. 
a. C. heavisidei. Type-specimen. 


b. C. commersonii. Specimen A. 
c. C. eutropia. Type-specimen. 


THE ANATOMY OF THE TONGUES OF THE MAMMALIA. 639 


33. The Comparative Anatomy of the Tongues of the 
Mammalia.—VII. Cetacea, Sirenia, and Ungulata. 
By CHarues IF. Sonntag, M.D., F.Z.8., Anatomist to 
the Society. 
[Received April 18, 1922: Read May 9, 1922.) 


(Text-figures 25-30.) 


ConTENTS. 

Page. 

Introduction eer rey ee doa bes eee oe eRe Ae at OOO 
Orders@etacedia stray tamara creo h ee ta Mey eASe ODD 
cp) HSUDCELSUEIW Seaiestncivg Gactena neon bentutes tobueeee sa Monee mar sec aame Rh O45 
doy a OfnaVes WUE KH tea near Seay eacieaneun soo dee adcane ees Ae ne arma ere ARM MEL Ss 4 
Summ anysan dC onc lustonsmersers een ee ee i ceee CeEeeeee een COA 
Bibliography merece eee: teacree eee cage tee Soe ace ee nee GHG 

IytrRopuctTiIon. 


In papers * on the tongues of the Primates, I showed that the 
mobility is well marked, the gustatory and secretory organs are 
well developed, and the papilla are not specialised for one kind 
of food. In the remaining mammalian orders one or more of 
these characters is highly developed, or greatly diminished, in 
accordance with the nature of the diet and mode of feeding. As 
the literature contaims many descriptions of individual tongues, 
the remaining papers of this series will be limited to general 
descriptions of the different types, and special attention will be 
paid to physiology and classification. 

The present paper is based on the examination of specimens in 
the Society’s Prosectorium, the British Museum (Nat. Hist.), 
and the Museum of the Royal College of Surgeons. 


Order CETACEHA. 
List of specimens examined. 
Suborder Mysracocrri (Baleen Whales). 


Pieces of tongue of Balena and Balenoptera. 


Suborder Oponroceri (Toothed Whales). 


Sperm Whale (Physeter macrocephalus), Sowerby’s Beaked 
Whale (Jesoplodon bidens), Beluga (Delphinapterus leucas), 
Porpoise (Phoceena communis), Cephalorhynchus eutropia, Risso’s 
Dolphin (Grampus griseus), Common Dolphin (Delphinus delphis), 
Bottle-nosed Dolphin (Zursiops tursio), White-beaked Dolphin 
(Lagenorhynchus albwrostris). 


* P.Z.S. 1922, pp. 1-29, 277-322, 497-524, 741-767. 


640 DR. C. F, SONNTAG ON THE COMPARATIVE ANATOMY 


The tongues of the Odontoceti differ greatly from those of the 
Mystacoceti, and both differ considerably from those of the other 
mammalian orders. Some are so simple that they resemble the 
tongues of fishes. 

Size:—In many Cetacea the tongue fills the space between 
the halves of the mandible, but it does not do so in the Narwhal 
(Monodon monoceros), Balenoptera borealis (29), and Delphina- 
pterusleucas. Barclay (2) showed that the food must pass far back 
to reach the tongue, if it is an organ of taste in the latter. In 
Balenoptera boops (19) it forms a large mass, projecting upwards 
between the baleen plates like an intermolar eminence. The 
bulk may be so increased by gaseous decomposition after death 
that the tongue protrudes from the mouth. This protrusion, 
however, does not take place in the Odontocete tongue. 

Consistence :—In the Odontoceti the tongue is firm, hard and 
muscular, and the upper surface feels like parchment. In the 
Mystacoceti, on the other hand, it is soft, from the presence of a 
large amount of oil or fat which, according to Owen (27), sepa- 
rates the mucous membrane from the muscles. Schulte (29) has 
also shown that masses of fat separate the musculi genioglossi in 
Balenoptera borealis. The oi] will exude from the cut surface 
of the tongue for a long period in preserved specimens. Some 
have, in fact, likened the tongue to a sac of blubber. Rawitz (28) 
and Eschricht (13) described the fat in adult animals, and 
Kiikenthal (21) saw it in a 117°5 ecm. fetus of Balenoptera 
musculus. 

Mobility :—-John Hunter (20) showed that the tongues of the 
Odontoceti are more muscular and mobile than those of the 
Mystacoceti, and attributed the difference to the methods of 
feeding. In the former they are organs of prehension, but they 
are passive in the latter, for the food flows into the open mouth. 
Scoresby (31) described the mode of feeding in Balena mysticetus 
as follows :—‘‘ When the whale feeds, it swims with considerable 
velocity under water, with its mouth wide open; the water enters 
by the fore part, but is poured out again at the sides, and the 
food is entangled and sifted, as it were, by the whalebone, which 
does not allow anything to escape.” 

In the Odontoceti the mobility varies. In Orcella brevirostris 
(1) it is great, for the free part extends back as far as the fourth 
interdental space. In Platanista gangetica the apex is bound to 
the mandibular symphysis by a fold of mucosa, but the edges are 
free and mobile. The animal is blind and burrows in the mud 
at the bottom of rivers for small fishes and crustacea, which con- 
stitute its diet. So the sensitive edges of the tongue may be 
organs of exploration. In Mesoplodon bidens, according to 
Turner (34), the tip is mobile from side to side. 

In both suborders the tongue is more mobile in the new-born 
animal than in the adult. 

Shape:—The tongue is large and shapeless in the adult 
Balenoptera boops (10), but it is broad and squat in the feetal 


OF THE TONGUES OF THE MAMMALIA, 641 


Balenoptera borealis (29). In the majority of the Odontoceti it 
has the usual mammalian form, but it is slipper-shaped in 
Grampus griseus and Lagenorhynchus albirostris. 

The apew varies considerably. It is full and rounded in 
Balena and Balenoptera. In the Odontoceti, on the other hand, 
it is not so full and rounded, and it may or may not have pro- 
cesses and warty growths. In Globicephalus melas (25), Cephalo- 
rhynchus eutropia (text-fig. 25 B), and Physeter macrocephalus 
(text-fig. 25.A) it is plain and pointed. In Grampus griseus 


Text-figure 25. 


Tongues of the Cetacea. A: dorsum of the tongue of a foetus of Physeter macro- 
cephalus ; B: lateral view of the same, showing the loose wrinkled frenum ; 
C: tongue of Cephalorhynchus eutropia, 


(text-fig. 26 A), Lagenorhynchus albirostris (text-fig. 26 B), and 
Delphinus delphis (text-fig. 26 C) it is broader and smooth, but 
Carus and Otto (8) described and figured it as covered with 
tubercles in the latter. The apical lobules are small in Phocena 
communis (text-fig. 27 A), large in Delphinapterus leucas (text- 
fig. 27 B), and in two rows in Vursiops tursio (text-fig. 27 C). 
The lateral borders are immense and massive in Balenoptera 
boops, and thin in the foetal B. borealis. They are very variable 
in the Odontoceti. In no Cetacean have they any lateral organs. 


642 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY 


Owen (27) stated that they are plain in the Mystacoceti, but 
Wyman (35) described lobules in B. borealis. Schulte (29), how- 
ever, did not figure them in the feetus of that species. 

Tt is difficult to decide where the oral and pharyngeal parts of 
the tongue meet in many species, for vallate papille are frequently 
absent. In Orcella fulminalis the base is delimited by a sulcus, 
whose ends correspond to the angles of the mouth. And many 
glands open into the sulcus (1). 

Sulet :—Median dorsal and median ventral sulci are absent in 
most cases. But many fine longitudinal and transverse sulci 
may be present. In Orcella they feel gritty to the touch. 

The tongue may be smooth and plain all over, as in the feetal 
Balenoptera borealis. It is wrinkled all over in Orcella brevi- 
rostris. In many species the posterior part of the dorsum is 


Text-figure 26. 


Tongues of the Cetacea. A: Grampus griseus; B: Lagenorhynchus albirostris ; 
C: Delphinus delphis; s: glandular sac. 


divided into areas by sulci. The inferior surface is more or less 
corrugated, and may rest on a cushion formed by folds of the 
mucosa of the floor of the mouth. The degree of corrugation 
varies at different ages, for Anderson (1) showed that the tongue 
in the young Platanista gangetica is smooth, but its root is 
corrugated in the adult. In Cephalorhynchus eutropia a thick 
fold surrounds the tongue below the apex. 

Glands:—The most marked features on the tongues of the 
Cetacea are the orifices of innumerable glands, and nearly every 
account records their presence. They vary greatly in extent and 
prominence, and they are more numerous than in all other 
Mammalia. 

* Genus Orcella :—In O. fulminalis many racemose glands open 


® 
OF THE TONGUES OF THE MAMMALIA, 643 


into the basal limiting sulcus. In 0. brevirosiris there are no 
glands on the inferior surface, but the whole dorsum has patulous 
orifices. Those on the base of the tongue are very large. 

Genus Platanista:—The glands are numerous, but not as long 
as those in Orcella. Some open into sacs. 

Genus Mesoplodon (text-fig. 28):—Numerous large and small 
glandular orifices are present, and there are five large sacs with 
linear orifices. 

Genus Lagenorhynchus (text-fig. 26 B):—The orifices cover 
the posterior two-thirds of the tongue, and increase in size from 
before backwards. The central ones lie on elevations. No sacs 
are present, and there are no glands on the inferior surface. 

Genus Tursiops (text-fig. 27 C) :--Many small orifices surround 
the edges of the anterior part of the tongue, and there are large 
clusters beneath the tip. 


Text-figure 27. 


Tongues of the Cetacea. A: Phocena communis; B: Delphinapterus leucas ; 
C: Tursiops tursio. 


Genus Phocena (text-fig. 27 A):—No sacs are present, and 
many minute orifices crowd the posterior part of the dorsum. 

Genus Delphinus (text-fig. 26 C) :—Many small orifices, lying 
in the centre of small areas crowd the posterior part of the 
dorsum. And there are clusters of pores on each side of a 
median ventral elevation. ‘T'wo sacs (s) are present on the base 
of the tongue. 

Genus Delphinapterus (text-fig. 27 B):—No sacs are present, 
and the glands are restricted to the posterior part of the dorsum. 

Genus Cephalorhynchus (text-fig. 25 B):—Innumerable small 
elevations with minute, but patulous, orifices cover the posterior 
part of the tongue and pharynx. No orifices are present on the 
inferior surface, and no sacs are present. 

Genus Physeter (text-fig. 25 A):—The surface of the fetal 
tongue is pitted all over. But the nature of the specimen at 

Proc. Zoon. Soc.—1922, No. XLIV. 44 


644 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY 


my disposal did not permit histological examination being 
made. 

The lingual glands are tubular or branching, and vary in 
length. And authors who have described true vallate papille 
mention that glands open into the fosse. Murie writes as 
follows of the tongue of Globicephalus melas (25): “It exhibits 
numerous glandular papille and depressions, probably the 
representatives of papille fungiformes; other larger and much 
deeper furrows behind may be circumvallate cavities or mucous 
glands.” 


Text-figure 28. 


The tongue of Mesoplodon bidens showing glandular orifices and 
the five large sacs. 


Papille :—In most Cetacea, papille are scanty or absent, and 
those which are present are usually tactile or mechanical in 
function. The sense of taste is very slight or absent, and in no 
other mammalian order is it so deficient. 

Papille are most numerous in Orcella and Platanista. In the 
former the oral part of the tongue has filiform papille, and the 
pharyngeal part has pedunculated and sessile papiile, arranged 
singly or in pairs at the mouths of large racemose glands. In 
the latter the free part is thick with filiform papille divided into 
processes. 


OF THE TONGUES OF THE MAMMALIA. 645 


Grampus griseus (text-fig. 26 A) has neither filiform nor fungi- 
form papille. At the junction of the oral and pharyngeal parts 
of the tongue there are two rows of deep narrow slits in V-for- 
mation, but there is no mesial suleus. Hach row has six fissures. 
In the specimen in the Museum of the Royal College of Surgeons 
they are absent. 

Owen (27) described four large fossulate papille in Hyperoodon. 
But Turner (84) recorded many crypt-like depressions and 
papille, and a vallate V in Mesoplodon bidens. In my specimen 
of MW. bidens there are five large sacs, probably glandular in 
‘character. 

In Cephalorhynchus eutropia (text-fig. 25 B) there are no 
filiform or fungiform papille. Between the oral and pharyngeal 
parts of the tongue are five fissures in V-formation. 

In Delphinus delphis (text-fig. 26 C) the slits have closed lips. 
In Delphinapterus lewcas (text-fig. 27 B) they are longer, and 
the lips of one are opened to disclose a row of ylobular bodies. 
Phocena communis (text-fig. 27 A) has eight small fissures placed 
end to end in V-formation. 

Neither papille nor fissures are present in J/onodon monoceros, 
Lagenorhynchus albirostris (text-fig. 26 B), Delphinus phocena, 
and Balenoptera borealis. 

Tn no Cetacean is there any trace of lateral organs. It appears, 
therefore, that the gustatory function 1s practically absent. 

The Inferior Surface of the Tongue is usually folded, both 
longitudinally and transversely, and its mucosa is usually soft 
all over. But there is a firm bounding zone in Grampus griseus, 
Lagenorhynchus albirostris, and Physeter macrocephalus. Some 
forms have glandular pits beneath the apex. In no case did I 
‘see any trace of a sublingua or plice fimbriate, but Schulte (29) 
described a small triangular sublingua in the feetal Balenoptera 
borealis. 


The frenum is absent in Delphinapterus leucas and Mesoplodon 
bidens. It is slight in Grampus griseus, Lagenorhynchus albi- 
rostris, Cephalorhynchus eutropia, and Phocena communis. 
Schulte (29) said it is absent in the feetal Balenoptera borealis. 
Anderson (1) described it in the fetal Orcella brevirostris, but 
said it is absent in the adult. I observed a very marked frenum 
in the new-born Physeter macrocephalus (text-fig. 25 A), These 
observations would show that the Cetacea require a frenum 
while suckling, but not when they lead an independent existence. 

No Cetacean has salivary papille or plice fimbriate. 


Summary. 
1. The tongues of the Cetacea have their glandular organs 
better developed, but their gustatory and mobile functions are 


less, than in other Mammalia. 
44* 


646 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY 


2. The tongues of the Mystacoceti agree with those of the 
Odontoceti as follows :—1. Filiform papillw are scanty or absent. 
2. The mucosa is more or less corrugated. 3. There is no trace 
of foramen cecum, lytta, frenal lamella, lateral organs, and 
apical gland of Nuhn. 

3. The tongues of the Mystacoceti and Odontoceti differ in the 
following points :— 


Mystacoceti. Odontoceti. 
Tongue soft. Tongue firm and hard. 
Intermolar elevation present. Absent. 
Much oil in the tongue. Absent. 
Apex massive. Not so. 
Absent. Marginal lobules present. 
Lateral borders ill-defined. Well-marked. 
Glands less numerous. Glands very numerous. 
Muscles slight. Muscles well-developed. 
Mobility slight. Mobility variable. 


Order SIRENIA. 


The tongues differ considerably from those of the Cetacea, 
and their characters approximate to those of the tongues of the: 
Ungulata. 


Text-figure 29. 


Tongues of the Sirenia. A and B: dorsum and lateral aspect of the tongue of 
Halicore indicus ; C: lateral view of the tongue of Manatus americanus. 


The tongues (text-fig. 29) are firm and hard, but not very 
mobile. That of MJanatus thickens progressively from before. 


OF THE TONGUES OF THE MAMMALIA, 647 


backwards, but there is a distinct intermolar eminence. In 
Halicore, on the other hand, the posterior two-thirds are greatly 
elevated as in some Ungulata. 

The apex is rounded and entire in both genera, and the lateral 
borders are entire and devoid of lobules which characterise the 
Cetacean tongue. Neither notches nor sulci are present. 

The mucosa on the eral part is plain, but that on the 
pharyngeal part has many folds. The base has glandular openings 
arranged singly or in pairs. There are no glands on the inferior 
surface, and no apical gland of Nuhn is present. The glands are 
less developed than in the Cetacea. 

Papille :—In both genera there is, behind the apex, a cluster 
of retroverted cuticular spines. And as the tongue is not very 
mobile they are of great assistance in cropping the vegetation on 
which the animals live. Behind that cluster the dorsum is plain 
in Manatus, but covered with a velvety pile of small papille in 
Halicore. Owen (27) figures a plain dorsum behind the spines 
in the latter. 

In Manatus, according to Owen (27), there are many vallate 
papille. In Halicore they are represented by clusters of pits. 

Lateral Organs:—In Manatus (4) these are well-developed 
and appear as large cushions with numerous fissures. In Halicore 
these are absent. 

The Jnferior Surfuce has many large orifices in Manatus, and 
many embedded cylindrical bodies in Halicore, but I was unable 
to examine the latter microscopically. 

The /renum is slight, and there is no frenal lamella, foramen 
cecum, lytta, sublingua, or plice fimbriate. No comb-like 
structures are found on the infero-lateral aspect. 


Order UNGULATA. 
Suborders PERIssoDACTYLA and ARTIODACTYLA. 


In most species the tongue is long, comparatively narrow, and 
very mobile. But itis broad and flat in Rhinoceros. It has the 
greatest mechanical power in Giraffa. 

The apex is truncated, pointed or rounded, and may or may 
not have a notch. In many species it has clusters of hard 
mechanical papille, asin the Sirenia. It is free in all forms, 
and this gives it considerable mobility. 

The lateral borders are full and rounded. ‘They may be com- 
paratively smooth, or covered with prominent conical and 
fungiferm papille. But Sus is the only genus with lateral 
lobules similar to those in the Cetacea. 

The anterior part of the tongue is flat and very mobile. 
Posterior to that is an intermolar eminence which raises the food 
up to the molar teeth. Most posteriorly is a flat, thin, more or 
less glandular part. 

The intermolar elevation is present in all families. It is entire 


648 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY 


in all except the Rhinoceros in which it is cleft. And it is 
covered with papille belonging to one or more of the conical, 
fungiform, and vallate series. 

In the Perissodactyla it is well-marked, but low and flat. It 
has only conical papille in Hquws caballus. In Tapirus indicus 
it has both conical and fungiform papille. In Rhinoceros clusters 
of vallate papillee cover its halves. 

In the Suina it is also flat. And both conical and fungiform 
papille are present. 

The Camelidz have well-marked eminences. And they possess 
large vallate and very hard projecting conical papille im the 
Llama. 

In the Cervide and Bovide the prominent eminence has very 
hard conical and fungiform papille, and the former vary con- 
siderably in size and shape. Clusters of vallate papille may 
invade the sides of the eminence. I did not examine a suffi- 
ciently large series of tongues to draw conclusions as to their 
value for purposes of classification. 

The eminence is prominent in Zragulus, but flat in Hyomoschus. 

When the posterior third of the tongue is examined it is seen 
how glandular orifices are numerous in the Perissodactyla, but 
few or absent in all other forms. And the characters of these 
structures in the different Mammalian orders, with special 
reference to the relation between their size and that of other 
parts of the oral glandular apparatus, have already been 
described. 

It is seen in this paper that the Cetacea, which have no salivary 
glands, have large lingual glands; the Sirenia, which have small 
salivary glands, have fewer glands; and the Ungulata, with good . 
salivary glands, have few lingual glands. Also the Perissodactyla 
have larger lingual glands than the Ruminantia. 

Circumvallate Papille (text-fig. 30 A):—The number and 
arrangement vary, and the following patterns were observed by 
myself :— 


1. No papille. 

2. A pair of papillee. 

3. Several papille in a straight line, or wide-angled V. 
4, Rows of papille on each side of the tongue. 

5. Clusters or fields of papillee. 


In the Perissodactyla there are two large papille in Hquus 
caballus, H. asinus, and EH. chapmanni, but several observers 
noted three in H. caballus. Mayer (22) described a pair in 
Tapirus americanus, but I noted several in a straight line or 
wide-angled V in 7%. americanus, 1. indicus, and 1’. bairdi. In 
Rhinoceros there is a field of ten papille on each side of the 
tongue. 

In the Suina there is a pair of papille in Sus scrofa, S. babi- 
russa, Potamocherus penicillatus, Phacocherus eliani, and Dicotyles 
torquatus. Schwalbe (80) recorded three papille in Sus scrofa. 


} 
OF THE TONGUES OF THE MAMMALIA. 649: 


The Tylopoda have the largest vallate papille, and they are 
arranged in two converging lines on the narrow intermolar 
eminence. In Camelus dromedarius there are seven on each side 
in a single chain. But Mayer (22) stated that the seven in 
C., bactrianus are in two rows on each side—an inner one with 
three papille, and an outer one with four. In Auchenia the 
numbers of papille are not identical in each row. 

The Tragulide have types of papille which are not found in 
any other group. There is a pair of long furrowed papille 
surrounded by a patulous fossa. in both Tragulus and Hyomo- 
schus; but Flower (14) described many small papille in the 
latter. 

The tongue in Giraffa has more papille than that of any other 
mammal. Miinch and Tuckerman counted fifty, and Owen (27) 
described two fields, each with 15-20 elements. 

In the Cervidee the papille are usually arranged in two rows 
on each side. And the following numbers were observed by 
myself, or recorded by others :— 


Muntiacus muntiac ............ 6 on each side. 
(CPO ACV ODOIS Fae ah ae abceats 26-28 (26). 
es SIUC SE ein INI, HN eRe Y 15-20 =, hn 
os | GOTOOTS EO eotecsaanshhees DOT Re rf 
SONALI LUISA LOT aisat anew, LOS i 
Capreolus cUpred ..........0.00. T2234 fisd vi (24). 
Rangifer tarandus.............4. D6 ots af (22). 
AUIGES TOGO DUGS oeBes-4-e7 soe saanen 18-20... if (33). 
Cariacus virginianus ......... Lee 7 (33). 
oi LOLECCUS MY Ere nee LOST aye ah (33). 


In the Bovide there are rows of small papille on each side, 
and the number of rows are shown in the classification given 
below. 

The following list contains the number of papille :— 


[KOS UTMPUIS.. «gan dbp Sonawenodseee: 10--17 on each side. 
Bison americanus ............... 18 be 4 

Path OTL USUUSa tt eceiiat mec: 11 on one side, and 6 on the 

other (33). 
/BOUOOS GOCNGUS  Loosbadoesseosoe sr 17-19 on each side (33). 
Budorcas taxicolor ..........:. (ARE ee Aero): 
Conmochwtes guns. sn 20 ares feel ( Le AE shag 
Cephalophus maxwelli ......... bat ts 
iA QOTSUVISI ee oe WW id 

Antilope mergens ............... 18-20 , ee (GN: 
Rupicapra rupicapra ......... UO Eon etheldel Gs) 
Antilocapra americana ...... 36 (33). 
CODECRERCUS pero tee De ee a 

PIE TUCIO Te te ty Meee tac tte ora: i 
DOU CUES: SASS ee ME 12 


A PUROSRUGUS Bose eacbdpoacdoose een Bay 


‘650 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY 


It is frequently difficult to determine by the naked eye 
whether a certain papilla is of the vallate or fungiform variety. 

It appears, therefore, that the papillary patterns are distri- 
buted as follows :— 


INOW oan anlles) Teaebicsencasuce onodee Hyracoidea. 
AS pairorpapillee! yee: ese Equide, Suide, Phaco- 
cheride, Tragulide. 
Papille in a line or V ....... Tapiride. 
Papillleeyim OWS) 22.2256. 405-7 Camelidee, Cervide, and 
Bovidee. 
iPapillesimrtield sis caucee eee Rhinocerotidee and Giraffidee. 


The papille are oval, cylindrical, or conical with the bases ef 
the cones projecting beyond the vallums. And the surface is 
smooth, granular, or lobulated (text-fig. 30 B.). The fossa is 
closed or patulous, and the vallum varies in prominence. ‘Taste- 
buds are usually well-marked. 

Fungiform Papille (text-fig. 30 C.H.F.):—The distribution on 
the dorsal and ventral surfaces varies in the different families. 
In appearance they are hemispherical, or almost pedunculated, 
and the surface is smooth, granular, or covered with processes. 
Many have rich supplies of taste-buds. 

In the Perissodactyla they are not very numerous, but have 
the usual mammalian arrangement in clusters and rows; and 
those on the lateral borders are very numerous. In /quus there 
are none on the intermolar elevation, but there are prominent 
ones. there in Tapirus. In neither genus is there a marked 
ventral papillary zone. 

The tongues of the Suina have many papille on the dorsum, 
but few on the ventral surface. And those on the lateral borders 
may be very prominent. 

In the Tylopoda the papillae are not numerous on the dorsum, 
but they form a very wide ventral papillary zone. And in no 
other family is the latter so large. 

In the Cervidee there is a prominent cluster of papillae behind 
the apex. Between it and the anterior extremity of the inter- 
molar eminence there is an area possessing very few papille, but 
the latter is bounded laterally by papillary bands. The ventral 
papille are numerous, but small. 

In the Bovide there is no thick apical, dorsal cluster, and the 
papille stretch right back from the apex to the intermolar 
eminence. They are only absent from a thin central strip of the 
dorsum. They are very regularly arranged. They are not 
numerous inferiorly in Bison, but they are numerous, smali, and 
closely packed in Antilope, Capra, and Ammotragus. 

In the Tragulide the papillae are numerous on the dorsum, 
and have the usual arrangement. They are absent only from a 
narrow central strip. 


OF THE TONGUES OF THE MAMMALIA. . 651 


Conical Papille:—The teeth in the Ruminants are assisted by 
the action of the hard conical papille comminuting the food 
against the prominent palatal ridges. 

In the Perissodactyla they are innumerable, slender, silky, 
and set very closely together. And their characters are similar 
over the entire dorsum. 

In Sus the lateral borders of the tongue have innumerable 
long, club-shaped conical papille, as in the Cetacea. But no 
other genus of the Suina possesses them. In Potamocherus the 


Text-figure 30. 


eet? 2066 
@eS82G269 


3 oe h. bE A 


“AL A Pata tiy: 
AA AMAA. 
Ah bod Mb hfe, 
AEA. 
GMA AAA IME 


Ne 


Tongues of the Ungulata. A (a-i): vallate papillary patterns; B(a-j): naked 
eye appearances of the vallate papille: C (a—-c): fungiform papille; D: conical 
papilles; E: fungiform papille of the Bovide; KF: fungiform papille of the 
Cervide ; I.M.E: intermolar eminence ; G: sublingual combs. 


conical papille- on the base are very large, pointed and directed 
backwards. In Dicotyles all the papille are very minute. The 
characters of the conical papille and lateral organs are useful for 
classifying the Suina (page 653). 

In the Camelide the anterior part of the tongue has minute 
closely-set papille. But those on the intermolar elevation are 
large, flat, hard, and separated into two groups by a smooth 
central strip. 


652 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY 


The Bovide and Cervide have papille similar to those in the 
Camelidz, but those on the eminence are smaller, more numerous, 
and not divided into two groups. 

In the Tragulide the conical papille on the dorsum are all 
small in TVragulus, but in Hyomoschus those on the base of 
the tongue are large, pointed, closely set, and directed back- 
wards. ‘Those on the base are not so disproportionately large in 
Tragulus. 

The ventral papillary zone is narrow in the Perissodactyla, 
but wide in the Artiodactyla, especially the 'Tylopoda. 

The conical papille are shown highly magnified in text- 
fig. 30 D. 

Lateral Organs:—As Oppel (26) has collected the various 
published accounts, it is only necessary to show here their value 
for purposes of classification. They are frequently absent, and 1 
would suggest that the lateral rows of circumvallate papillee 
replace them in these cases. 

In the Perissodactyla they are absent, according to Boulart 
and Pilliet (5), in Vapirus americanus, Hquus caballus, and 
Rhinoceros. But they are well-marked in all the Tapiride- 
examined by myself. Complete accounts of the organs have been 
published by Sertoli (32) and Honigschmeid (19). 

In the Suina they are present in Sus, Phacocherus, and 
Potamocherus, but they are absent in Dicotyles. The Babirussa 
has circular organs, but those in the other forms consist of rows. 
of laminee and sulci. 

In the remaining Artiodactyla they are present in the Giraffide, 
Tragulide, and Antilope mergens, but they are absent in all others. 
examined, 

The Lytta is represented by a median ventral ridge, which 
varies in width and prominence, but it is not at all like that in 
the Carnivora. And sections show that it has a central core. 

In Tapirus indicus it 1s narrow, prominent, and firm. But it 
is wide, flat, and softer in Hgwws; and the structure in the latter 
has already been described by Briihl (7). Owen (27) recorded 
its presence in Lhinoceros. In all Perissodactyla it does not. 
widen much from before backwards. 

In Dicotyles it forms a long, narrow isosceles triangle. 

_ The Tylopoda, as exemplified by the Llama, have short, pro- 
minent crests with very thick mucosa. 

In the Tragulide it is broad and flat, especially in Hyomoschus. 
But it is most variable in the Cervide and Bovide. In these it 
is narrow and sharp, low and broad, or absent. The appearances, 
however, are of no value for purposes of classification. 

On the infero-lateral aspects of the tongue there may be long 
bands with divided free edges, or rows of separate processes. 
They may help to keep the interstices between the teeth clear, 
or they may help to mix the food and saliva. They have not 
been named, so I suggest the term ‘sublingual combs” for 
them. ‘They are restricted to the attached part of the inferior 


OF THE TONGUES OF THE MAMMALIA. 653 


surface of the tongue. The following arrangements have been 
observed :— 


1. Combs absent——Perissodactyla, Suina, Hyracoidea. 

2. Edges divided into triangles—Tylopoda, Tragulide. 

3. Long, more or less separate processes—Bovide and 
Cervidee. 

Some forms are shown in text-fig. 30. 

The frenum is always present, and permits the tongue to be 
very mobile. But there is no trace of a frenal lamella. 

There is no trace of a foramen cecum, sublingua or plice 
fimbriate, and the apical gland of Nuhn is present only in 
Ovis aries. 

It has been the object of the above summary of the characters 
of the tongues of the Ungulata to show their value for purposes. 
of classification. And they have been arranged schematically as 
follows. From that system it will be seen that the characters of 
the tongue are a useful addition to the external and skeletal 
characters at present in use. 

Classification.--The following characters are valuable for 
purposes of classificatlon :--1. Character of the orifices of glands 
on the base. 2. Nature of the intermolar eminence and its 
papille. 3. Arrangement of the vallate papille. 4. Lateral 
organs. 95. Sublingual combs. 6. Distribution of conical and 
fungiform papille. 


Suborder PERISSODACTYLA. 


Numerous orifices of glands on base. No combs. Conical 
papille silky and closely-set. Ventral papillary zone very small 
or absent. 


A. Intermolar eminence cleft. Vallate o uep in fields. No 
lateral organs ......... Witisscs LIMOGES. 
B. Eminence low, flat, and entired 
a. A pair of vallate papilla. No lateral organs. No fungiform 


papille on eminence ........ eae aeeraeeaen | LLUQKLUS: 
6. Vallate papille form a row or v. Tatars @ organs present. 
Large fungiforms On eminence............ccceeee eee «=Lapirus: 


Suborder ARTIODACTYLA. 


Few or no orifices of glands on base. Combs variable. Conical 
papille hard and not so closely set. Ventral papillary zone 
well-marked. 


A. No sublingual combs. Two vallate papille. 


a. Hdges of tongue have long processes...............06cec0eseeeees. SUS 
6. No long processes present. : 
1. Papillee on base immense ... 0.0... ...ce. ccc eee cesses seeeseneess. Potamocherus. 
il. Papille on base not immense. 
a’. Lateral organs present ..............:c0cccseeeeeceeeersenee. Phacocherus. 


> Wo Jeiiorel CEAINS "nes cocebdosdpoocosnspstocabson soneasBeg nonene.  ¢DOGTOICSE 


654 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY 


B. Sublingual combs present. 
a. A pair of long, narrow vallate sie 
. Basal conical papille small.. RpHcneccagreaonnodacobaaaca baat ie A ReKG TOUS 
il. Basal conical papillae very eck Sins ube teseseseeeeeee Sd yomoschus. 
6. A single row of immense vallate papille on each side... Camelus, Auchenia. 
ec. One or more rows of small vallate papilla on each oe 


i. Fungiform papille absent from an extensive area in 
front of the intermolar eminence. Two rows of vallate 
ORION pcsuogoodcees A aaeenence. | OBB VLD zs 
i. Fungiform papille Pretent right ie tne eminence, but 
are absent from a central strip on the anterior part of 
the dorsum. 


a’. One row of vallate papille on each side ................08 Cephalophus. 

Hs INGO ROWS Oil CAEN, BNW) ” Sonooa000 cco0c0 ode covaosonancs 046 Bos, Ovibos, Bison, 
Connochetes, Strepsiceros, Antilope. 

G5, J HOOP TRONS ON GAYOIN EHCIE). dcanccincn oso seosesoos cnn ees badadn des coe Budorcas, Ovis. 


Suborder HyRACOIDEA. 


The tongue in Hyraz has a low, flat elevation similar to that: 
in Halicore, and the lateral organs are well-marked. ‘There is 
no vallate papilla in several specimens examined by myself, and 
by Briicher (6), Gmelin (16), Munch (24), and Tuckerman (33). 
And there is no trace of a lytta. All conical papille are 
minute. The basal lingual glands have few orifices on the 
surface. It is evident, therefore, that the tongue has some 
affinities with those of the Sirenia. No sublingual combs are 
present. 


Suborder PROBOSCIDFA. 


The tongue is short and wide, and it corresponds to the inter- 
molar elevation of the other Ungulata. The apex is bound down 
to the floor of the mouth, as in the Cetacea. In Z#. indicus 
Mayer (22) observed six papille, but Miall and Greenwood (28) 
recorded four. In the specimen examined by myself there were 
two on the right side and one on the left. In 4. africanus 
Forbes (15) saw four papille on the right side, and three on the 
left, but Mtinch (24) observed three on each side. The lateral 
organs are well-developed, but their contained taste-buds are 
sometimes very few. No sublingual combs are present. 


SUMMARY AND CONCLUSIONS. 


1. The Cetacea have simpler tongues than any other mammals. 

2. The tongues do not support the view of some authorities 
that there are affinities between the Cetacea and Ungulata. But 
the differences between them are due to the nature of the diet 
and the mode of feeding. And the nature of the food causes 
the Sirenian tongues to resemble those of the Ungulata in some 
points. 


OF THE TONGUES OF THE MAMMALIA. 655: 


The essential differences between the Cetacea, Sirenia, 
Artiodactyla, and Perissodactyla are :— : 

Cetacea—Mobility slight. No gustatory organs. Glands. 
numerous. An intermolar eminence is only present in some 
Mystacoceti. Few or no mechanical papille. 

Sirenia—Mobility greater. Slight gustatory organs. Glands 
fewer. Small intermolareminence. Mechanicai papille present. 

Artiodactyla and Perissodactyla—Mobility great. Well-marked 
gustatory organs. Glands variable, being numerous in the latter 
and few in the former. Intermolar eminence well-marked, but 
variable in size. Mechanical papille well-marked. 

3. The characters of the tongue have several points in common 
in the Artiodactyla and Perissodactyla, but those of the 
Hyracoidea and Proboscidea are such as to separate them into 
groups by themselves. Of these the Hyracoidea have some resem- 
blance to the Sirenia. 

4. The Cetacea are the only mammals in which the tongues. 
may lie far back in the mouth. 

5. The tongues of the Mystacoceti differ from those of the 
Odontoceti. 

6. The tongues of the Mystacoceti contain large quantities of 
oil, but this is absent in all other mammals. 

7. The excessive development of cuticular spines on the 
anterior part of the tongue in the Sirenia allows great use to. be 
made of the moderate mobility. 

8. The Perissodactyla differ from the Artiodactyla in the 
characters of the glands on the base of the tongue. 

9. Sublingual combs are only present in the Camelide, 
Tragulide, Cervide, and Bovide. 

10. It is impossible to distinguish the genera of the Cervide 
from one another by the characters of the tongue. 

11. The distribution of the fungiform papille is a good means. 
of telling the Cervide from the Bovide, 

12. The Hyrax and Cetacea are the only mammals in which I 
was unable to detect vallate papille. 

13. The sheep (Ovis aries) is the only.mammal below the 
Primates which possesses the apical gland of Nuhn. 

14. The characters of the fungiform papille distinguish the 
deer from the antelopes. 


BIBLIOGRAPHY. 


1. AnpERson, J.— Zoological and Anatomical Researches of the 
two Expeditions to Western Yunnan. London, 1868-75. 

2. Barcuay, J.—‘‘ On the Beluga Whale.’ Wernerian Trans- 
actions, vol, 111. 

3. Baur, C. J.—‘“ Ueber den Bau der Zunge.” Deutsch. Arch. 
f. d. Physiol. hrsg. von J. F. Meckel, 1822, pp. 350-358. 

4. Bepparp, F. E.—‘: Notes on the Anatomy of a Manatee.” 
Proce. Zool. Soc. 1895, pp. 47-53. 


69 


St 


6 


DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY 


. Bounart and Priurer.—‘‘ Note sur l’organe folie de la langue 


des Mammifeéres.” Journ. de Anat. et de la Physiol. 
1885, pp. 337-345. 


§. Bricuer, C.—Deutsche Zeitschr. f. Tiermed. u. vergl. 
Pathol. 1884, pp. 93-111. 
7. Broun, K. B.—Gelegenheitsschrift vorgelegt dem Professorem- 
Kollegium d. K. K. Wiener Tierarznei-Instituts, 1850. 
8. Carus and Orro.—‘‘ Erlauterungstafeln zur verg]. Anatomie,” 
vol.iv. Leipzig, 1835. 
9. CHatin, J.—‘‘ Les Organes des Sens dans la Série Animale.” 
Paris, 1880. 
10. Cuvier, F.—Art. ‘“Cetacea” in Todd’s Cyclopedia of Anatomy 
and Physiology,” vol. 1. 
11. Epyer, V. v.—Kolliker’s Handbuch der Gewebelehre des 
Menschen. Leipzig, 1899. 
12. Extenpercer and Miiier.— Handbuch der vergleichenden 
Anatomie des Haustiere. Berlin, 1896. 
13. Escuricnut, D. F.—Zoologisch-anatomisch-physiologisch 
Untersuchungen iiber die nordischen Waltiere. Leipzig, 
1849. 
14, Fuowrer, W. H.—‘“On the Organs of Digestion in. the 
Mammalia.” Medical Times and Gazette, 1872, pp. 319- 
322 and 427. 
15. Forses, W. A.—‘“ On the Anatomy of the African Elephant.” 
Proce. Zool. Soc. 1879, pp. 180-195. 
16. Guetin.—“ Zur Morphologie der Papilla vallata und foliata.” 
Arch. f. mikr. Anat. 1892, pp. 1-28. 
17. Gruxpier, C. A.—De Camelo Dromedari observata quedam 
anatomie. Diss-Inaug. Tubinge, 1817. 
18. Gurut, E. F—Handbuch des vergleichenden Anatomie der 
Haussiiugethiere. Berlin, 1844. 
19. H6énicscuMiEeD, J.—Several papers enumerated by Oppel (see 
no. 26). 
20. Hunter, J oHN.—‘ Observations on the structure and economy 
of Whales.” Phil. Trans. Roy. Soc. 1787, pp. 371-450. 
91. KiikenrHaL, W.—Several papers enumerated by Oppel (see 
no. 26). 
92. MAYER, a A.—‘* Uber die Zunge als Geschmacksorgan.” 
Nova Acta Acad. Leop.-Carol. 1844, pp. 721-748. 
93. Mratuand Greenwoop.—Journalof Anatomy and Physiology, 
1878, pp. 17-29. 
94, Mincu, F.—‘ Die Topographie der Papillen der Zunge des 


Menschen und der Saiugethiere.” Morphol. Arb. 1896, 
pp. 605-690. 


. Muriz, J.—‘* On the Organisation of the Caaing Whale, 


26. 


Globicephalus melas.” Trans. Zool. Soe. vol. viii. pp. 235- 
301. 

Opret, A.—Lehrbuch der vergleichenden Mikroscopischen 
Anatomie, vol, iii. pp. 325-344. 


27. 
28. 
29. 


30. 
31. 


32. 


33. 


34. 


35. 
36. 


OF THE TONGUES OF THE MAMMALIA. 657 


Owen, R.—The Comparative Anatomy of Vertebrates, 
vol. i. pp. 193-197 (1868). 

Rawitz, B.—‘ Uber Megaptera boops.” Arch. f. Natur- 
geschichte, 1900, pp. 71-114. 

Scuunre, H.—“ Anatomy of a fetus of Balenoptera borealis.” 
Mem. Amer, Mus. Nat. Hist. 1916, pp. 389-502. 

Scuwatse, G.—Arch. mikr. Anat. 1868, pp. 154-187. 

ScoresBy.—‘ On Balena mysticetus.’ Wernerian Trans- 
actions, vol. i. p. 578. 

Servroui, E.—Molleschott’s Untersuchungen zur Naturlehre, 
1876, pp. 403-415. 

TuckerMAN, F.—Journal of Morphology, 1890, pp. 152- 
193, and 1891, pp. 69-94. 

Turner, W.—‘The Anatomy of a second specimen of 
Sowerby’s Whale.” Journal of Anatomy, vol. xx. 
pp. 144-188 (1886). 

Wyman, J.—Bost. Jour. Nat. Hist. vol. vii. p. 603 (1864). 

Lanper, K. F.—‘‘ Some Points in the Anatomy of the Takin 
(Budorcas taxicolor whitei).” Proc. Zool. Soc. 1919, 
pp. 203-218. 


THE EXTERNAL CHARACTERS OF SCARLTURUS. 659 


34, The External Characters of Scarturus and other Jerboas, 
compared with those of Zapus and Pedetes. By R. I. 
Pocoexn, F.R.S:, F.Z:8. 

[| Received May 9, 1922: Read May 23, 1922. | 
(Text-figures 29-37.) 


Page 
TiAROCIAGHIOM | scidopooodseons onccbosonoacwboooacHoHdaoDaKedeos. HABE) 
Deere] FAN OEENETINED ccotscocoocosetcosoacboseees cap onocencca , Le) 
The Rhinarium and Mouth .............................. 661 
Theor) WHORES. godee0 cnoseanksaoo con oas cco ceunaeocessacuocs — (lshs) 
TTS RES An ee, i ee pn Fa AK Wane NRO A A ah eee NOGA 
UMS aD GnAe MINOT Bas dor caeenen eek beeen nen etenocodtesendesnendes ASOT 
Days) 15 GME 9 DoyoXe soo Gabe Nen naan seou soo eadsbodtenon coosdacan i OFAD) 
Amuseangs dixternale Genitalia snesscssete ceases ONO 
The Skull of Scarturus.. Rees and Remar ret ai S]7A 
The Genus Scirtopoda Brande Beier seayaenaeeh DONO 
Ce dns Nea bul GOT Hin Hill On, 680 


INTRODUCTION. 


At the Scientific Meeting of the Zoological Society held on 
May 24th of last year (P.Z.S. 1921, p. 645), Major Flower ex- 
hibited examples of three species of Eg gyptian Jerboas (Jaculidee), 
including a pair of the very rare four-toed form, Scarturus 
tetradactylus, which had never been previously imported to 
Kurope alive, and was almost unknown in museums, apart from 
the original example in Berlin, which was described as Dipus 
tetradactylus by Lichtenstein in 1823. These examples were 
exhibited in the Zoological Gardens for some months, but died 
in the winter and came into ny hands for examination *. ‘lo 
compare with them I had several specimens of the two common 
Egyptian Jerboas (Jaculus gaculus and Scirtopoda orientalis) t ; 
and also one of the five-toed genus Allactaga (A. indica), a pair 
of examples of Zapus hadsonianus or a nearly related species, 
and one male specimen of Pedetes caffer—the last-mentioned 
examples having been preserved for many years in alcohol in the 
Society’s collection. ‘The specimens above mentioned form the 
material upon which this paper is principally based. 


* Although apparently fully adult when received, all the skeletal tissues at the 
time of death were exceedingly frail. Even the tips of the incisor teeth were 
unealcified and pliable—a condition I never before observed in any mammal. 

+ Good figures, drawn from life, and descriptions of these, as well as a brief 
account of S. tetradactylus, may be found in Anderson and de Winton’s ‘ Mammalia 
of Kgypt.’ My reasons for adopting the name Scirtopoda are given below (p. 679). 


Proc. Zoou. Soc.—1922, No, XLV. 45 


660 MR. R. I. POCOCK ON THE EXTERNAL | 


That Zapus belongs to the same group of Rodents as Jaculus 
has long been admitted, and is disputed, so far as | am aware, by 
no one acquainted with this order. ‘lhe evidence rests mainly 
upon certain cranial characters connected more particularly with 

the zygomatic arch and the preorbital foramen, the structure 
of which distinguishes that genus from the Muride, despite the 
venerally murine shape of the skull. It is also conceded that 
Sicista, formerly known as Sminthus, is tolerably closely related 
to Zapus. With regard to Pedetes, my pretext for including 
that genus in this paper is its former ascription to the same 
section * as Zapus and Jaculus, although now recognised as 
representing a quite distinct family, sometimes associated with 
the Hystricomorpha, together with the family Anomaluride. 


External Appearance. 


The general appearance of Sicista and Zapus may be described 
as murine. They have a long mouse-shaped head and compara- 
tively small diurnal eyes, and, although the hind legs of Zapus 
are elongated and saltatorial, they are not very much longer rela- 
tively than in our common woed-mouse, Apodemus sylvaticus. 'The 
Jerboas, on the other hand, are not the least like typical murine 
rodents in appearance. The head is very large, wide, and, asa 
rule, short especially in the snout, which is deep from above 
downwards, and the eyes are relatively enormous and nocturnal. 
The hind legs are of great length and the fore limbs are short. 
A Jerboa with the fore limb, measured from the elbow, of 
approximately the same length as that of a Zapus has the foot, 
measured from the heel, about twice as long as that of Zapus. 
Hence, when progressing on all fours, the gait of a Jerboa is 
exceedingly awkward, owing to the great elevation of the hind 
quarters. They seldom adopt that mode of progression, usually 
employing their hind legs alone. When moving at speed they 
take prodigious hops, like a Kangaroo; but they frequently go at 
a more leisurely pace, trotting or striding along like an ostrich 
or a human being, an action of which Kangaroos, and, pre- 
sumably, Zapus also, are quite incapable. 

The great var iation in the size of the ears affects the appearance 
of different species of Jerboas when compared with one another. 
They also differ in the shape of the muzzle. As above stated, the 
muzzle is typically short and truncated; but in Huchoreutes, 
hence named mnaso, it is relatively long. Presumably also 
Allactaga, although I have never seen a living example, must 
have a differently shaped muzzle from Jaculus on account of the 
prominence of the upper incisor teeth, which form an angle 
instead of, as in Jacwlus, a continuous curve with the nose. 
Scarturus is intermediate between the two genera just mentioned 
with respect to the direction of these teeth. 


* See Flower and Lydekker’s ‘ Mammals, Living and Extinct,’ p. 480 (1891), 


CHARACTERS OF SCARTURUS AND OTHER JERBOAS. 661 


The Rhinariwm and Mouth. 


In Zapus the rhinarium is well defined and marked by a 
distinct vertical groove, formed by the thickened bulging upper 
and inner margin of the nostrils. Above the thickening of the 
upper rim there is a short transverse area. The area below 
the rhinarium, extending from the nostrils to the edge of the 


Text-figure 29, 


A. Head of Zapus hudsonianus. 


B % Jaculus jaculus. 
Cary Scirtopoda orientalis. 
D Hy Scarturus tetradactylus. 


upper lip, is covered with shorter hair than the rest of the 
muzzle. There is no sign of a groove or philtrum dividing 
the upper lip in the middle line. (Text-fig. 30, A.) 

From Miller’s description of the rhinarium of Siczsta loriger 
(Cat. Mamm. Western Europe, p. 537, 1912), it seems that the 
muzzle resembles in general features, at all events, that of Zapus, 


4n* 


662 MR. R. I. POCOCK ON THE EXTERNAL 


He describes the anterior border of the nostril as thickened, 
with asmall wart beneath it [? the nostril], and the space between 
the nostrils as narrow, marked by a well-developed vertical ridge, 
and the upper lip as having no groove in front. 

In Jaculus, Scirtopoda, and Scarturus the rhinarium is very 
like that of Zapus, showing a similar thickening ef the upper and 
inner rim and a median groove, but the area below the rhinarium 


Text-figure 30. 


= Raa WH 
¥ Yi i 
WN 


Ge 


D S 


. Muzzle and rhinarium of Zapus hudsonianus. 


A 

B. ” 9 Scirtopoda orientalis. 
C. The same, contracted. 
D 
E 


. Muzzle and rhinarium of Jaculus jaculus, to show especially the papille, 
slightly enlarged and cleared of hair, carrying the long stout vibrissee. 
. Muzzle and rhinarium of Pedetes caffer. 


in the middle line is naked and constitutes a kind of shallow 
philtrum. Of the muzzle of this genus, Anderson and de Winton 
said :—‘“ Over the bare area of the nose of Jaculus there is a 
thickened fold of skin capable of being drawn forwards, so as 
almost to cover the nostrils, this structure serving to protect them 
when the animal is using its broad snout to push out the earth 


CHARACTERS OF SCARTURUS AND OTHER JERBOAS. 663 


when digging its burrow” (Mamm. Egypt, p.304). The rhinarium 
when thus partially concealed also contracts very perceptibly, the 
nostrils are approximated and assume a more vertical appearance, 
and the philtrum is obliterated by the median juxtaposition of 
the two halves of the upper lip. This device probably serves to 
exclude fine blown sand, as well as the soil when digging is in 
operation. (Text-fig. 30, B, C.) 

The rhinarium of Pedetes is quite unlike that of Zapus and 
Jaculus. It is covered above and in front with fine short hair; 
but the nostrils, which are elongated and appear to be valvular, 
are surrounded by a narrow rim of naked skin. The median 
portion of the rhinarium below the nostrils in front is angled, 
but there is no trace of groove or philtrum dividing the deep 
upper lip, which is covered uniformly with longish hair (text- 
fig. 30, E). 

Both in Zapus, Jaculus, Scirtopoda, and Scarturus, the mouth 
is provided with well-developed palatal and lingual lobes jutting 
inwards from the cheek. The palatal lobes meet along the middle 
line of the palate, but are separable and do not fuse in’ the 
middle line, the epithelium of the palate being smooth. The 
lingual lobes, which are long, meet in the middle line over or 
in front of the tongue, forming a secondary floor to the mouth. 
Zapus 1s provided with cheek-pouches opening just inside the 
corner of the mouth. The Jaculide are without them, and, so 
far as I am aware, they have never been recorded in Sicista. 

In connection with the mouth, reference may be made to the 
presence in some genera of grooves on the incisor teeth. Their 
incidence is remarkable and puzzling. They are absent in Sicista, 
the least specialised genus of the group; but are present in Zapus, 
which seems to be much more nearly akin to Sicista than it is to 
any genus of Jaculide. Within the Jaculide, the grooves are 
present in Dipus, Jaculus, Scirtopoda,and Cardiocranius ; absent | 
in Allactaga, Scarturus, Huchoreutes, and Pygeretmus. Their | 
presence, as recorded by Satunin, in Cardiocranius—a genus 
resembling Pygeretmus in possessing five toes on the foot and a — 
specialised broad lanceolate tail—suggests that the character has ~ 
no great systematic value. 


Facial Vibrisse. 


The mystacial vibrissee are well developed in Zapus. One or 
two long superciliary vibrisse are also present; but in the long 
hairs clothing the cheek I failed to find the genal tuft. The 
interramal is represented by two very fine vibrisse in one example. 
In the Jaculide the interramal tuft is apparently absent, 
and the superciliaries and genal tufts are of inconstant occur- 
rence, and represented at most by one or two fine vibrissee each. 
But the mystacials are well developed, one of them on each side 
set low down being stouter and longer than the rest. This 


664 MR. k. 1. POCOCK ON THE EXTERNAL 


peculiarity is not so well marked in Alluctaga and Scarturus as 
in Jaculus; but the two common Kgyptian Jerboas differ some- 
what in this respect. In Scirtopoda orientalis this long bristle is 
distinctly stouter and stiffer than the rest, and rises from a small 
papillary root, whereas in J. jacwlus the differ ence is considerably 
greater, and the papilla from which it arises is much larger. 
(Text-fig. 30, D.) 

In Pedetes the mystacial vibrissee are comparatively few in 
number and short, although stout. A few longish superciliaries 
are present. The interramal tuft is apparently absent; but the 
genal is represented by two or three long vibrissee set high up 
the cheek a little behind and below the posterior angle of the eye 
as in the Hystricomorpha. 

By the characters mentioned above the genera may be arranged 


as follows :— 


a. Rhinarium ill defined, covered with hair except immediately round 
the nostrils, which are not visibly swollen and have no median 
groove between them; superciliary and genal vibrisse well de- 
veloped; mystacials stout and comparatively short . WiWhskee! Pedetes: 
. Rhinarium well defined, naked, nostrils with ssiollen i rim anal a 
groove between them; superciliary and genal vibrissz few or 
absent, mystacials long and slender for the most part. 
}. Head narrow and low, murine in shape; no median groove on 


upper lip; mouth with cheek-pouches (? in Sicista) ...... Zapus, Sicista. 
. Head broad and high; upper lip. divided by philtrum, no cheek- 
pouches ......... lictiteseee. Jaculus, Scirtopoda, Scarturus*. 


The Har. 


In Zapus the eav is moderately large and expanded, and arises 
from the head by a broad base which is not tubular. The anterior 
edge is normally overfolded, and inferiorly about on a level with 
the supratragus it is produced into a lamina which is normally 
folded inwards out of sight. The lower edge of this curves into 
the cavity of the ear and ceases on its floor. The tragus is 
represented by a small movable lobe,and the antitragus by a large 
tufted lamina which, when the ear is folded, projects forwards 
and closes the lower portion of the cavity. The orifice is situated 
just below and on the inner side of the deep intertragal notch. 
(Text-fig. 31, A-C.) 

Miller describes the ear of Sicista loriger as ‘‘ moderately large, 
extending a little beyond the eye when laid forward, its outline 
simple, ovate, its posterior border with large thickened lobe at 
base. capable of completely closing the ieatus, its anterior border 
with minute though evident sub-basal notch and projection ; 
both surfaces of ear densely clothed with short hairs; basal lohe 
conspicuously tufted.” This description might have been taken 


* JT have not seen a sufficient number of specimens of Allactaga and other Asiatic 
genera of Jaculide to know whether the cleavage of the upper lip is a constant 
character in this group or not. 


CHARACTERS OF SCARTURUS AND OTHER JERBOAS., 665 


from the ear of Zapus, with the anterior flap concealed, as is 
normally the case, by being folded inwards. 


Text-figure 31, 


. Ear of Zapus hudsonianus, expanded. 

. The same, contracted. 

. Base of ear of the same, with basal portion cut open between tragus 
and antitragus. 

. The same of Scarturus tetradactylus. 

. The same of Scirtopoda orientalis. 

. Expanded ear of Pedetes caffer. 

. Folded ear of the same. 

. Base of the ear of the same, cut open. 


Qnh> 


Torneo 


1.. infoided lamina of anterior border; ¢., tragus; at., antitragus; s., supratragus ; 
o., orifice of ear; ¢., columnar process. 


In the Jaculide the ears vary greatly in length. In Hucho- 
reutes, the longest-eared form, they considerably overlap the tip 
of the exceptionally long snout, when turned forwards, In 


666 MR. R. I. POCOCK ON THE EXTERNAL 


Scarturus, which has a short snout, they surpass it by a little. 
In Allactaga indica they veach about to the end of the muzzle, 
and in most species assigned to this genus they appear, judging 
from Lichtenstein’s figures, to reach well beyond the eyes at all 
events. In Sciréopoda orientalis they overlap the eyes bya little ; 
in J. gaculus they just cover them. In other species figue ed by 
Lichtenstein they are as short as in the typical species or shorter. 
But despite their length they all appear to be similarly con- 
structed. They are more or less tubular at the base, the orifice 
being sunk deep down within the cavity. There is no trace of 
tragal lobe or antitragal flap, such as are seen in Zapus and 
Sicista; but just within the anterior edge, on a level with the 
Supratvagus, there is a delicate membranous lamina descending, 
as 1n Gaps, into the tubular basal portion and normally folded 
in out of sight. The lower end of this lamina curves backwards 
and inwards, and just inside its lower end there is a deep cavity 
bounded above by the anterior end of the supratragus, which is a 
slender delicate ridge running backwards towards the posterior 
edge of the ear, exactly as in Zapus. The orifice is situated 
a little below ‘thts anterior cavity, and is Separated from it by a 
thickening. There are minor differences in the structure of the 
ear which may prove to have systematic importance, but my 
material is not sufficient in quantity to establish the point. In 
Scirtopoda orientalis, for mstance, the anterior infolded lamina 
is larger than in Scarturws, and the area round the auditory 
orifice is more swollen. In Scartwrus, on the other hand, the 
lower end of the passage leading to the anterior pit is much more 
swollen than in S. ortentalis, and the tubular portion of the ear 
is provided with a fine delicate lamina ascending from a point 
behind the auditory orifice up to the position of the aborted 
antitragus. This is not so noticeable in S. orientalis. Sclater’s 
Brief deste iption of the ear of Huchoreutes contains nothing to 
suggest any difference between that genus and the three I have 
examined in the structure of the organ in question. (Text- 
fig. 31, D, EH.) 

From what has been said above it will be clear that the kinship 
claimed to exist between Zapus and the Jaculide on cranial 
grounds is completely borne out by the ear. The ear of the 
Jaculide may be regarded as a derivative of the type seen in 
Zupus, specialised by its more tubular base, with which the dis- 
appearance of the tragus and antitragus is correlated. In the 
Jaculidee, too, the anterior basal pit is deeper and its inferior edge 
1s closer to the auditory orifice. 

The ear of Pedetes is very different from those described above, 
although in its tubular base and its length it recalls in appear- 
ance the ear of some Jaculide, It shows, however, in its posterior 
border behind the tip, a shallow but long emargination, similar 
to the emargination so common in many of the Hystricomorpha. 
The anterior edge is not overfolded and there is no inturned lamina 
at the base. There is no antitragus, but from the inner side of 


GHARACTERS OF SCARTURUS AND OTHER JERBOAS. 667 


the outer wall of the tubular base there arises a compressed 
process which projects upwards as a free outgrowth some little 
distance above the margin of the basal tubular portion. When 
the ear is expanded this process, which possibly represents the 
tragus, inclines forwards along the edge of the ear in front of it. 
The supratragus is a well-defined ridge running upwards and 
backwards, but stopping abruptly in the middle “of the ear far 
away from its posterior edge; and the wall of the ear between 
the posterior end of: the supratragus above and the auditory 
orifice below is folded and creased. (Text-fig. 31, F—H.) 

The reason for the termination of the supratragus in the middle 
of the ear is this. When the ear is folded, it doubles up exactly 
lengthwise along the middle line, so that the anterior and posterior 
margins are evenly in contact throughout their length, except 
close to the basal tubular portion; but the space that is left 
between them at this spot is filled by the process above described, 
and that appears to be its function. The grooving of the wall 
of the lower portion of the ear is also subservient to this peculiar 
method of folding, which could not be effected if the rigid supra- 
tragus extended across the cavity of the pinna. 

By the structure of the ear the genera discussed fall into the 
following three headings :— 

a. Kar long, folding exactly along the middle line, so that its anterior 

and posterior edges coincide; a columnar process rising from its 


basal tubular portion; supratragus a well-defined thickened ridge. 
No inturned flap at the base of the anterior edge in front ......... Pedetes. 


a’. Kar long or comparatively short, not folding so that the two edges 
of the pinna exactly coincide ; no columnar process rising from its 
basal portion ; a distinct inturned flap at the base of the anterior 
edge in front. Supratragus comparatively weak. 
b. A small tragus and large antitrazus, base of pinna not tubular. 
Zapus, Sicista. 


b’. No tragus or antitragus ; base of ear more or less tubular. 
Jaculus, Scirtopoda, Scarturus, Allactaga. 


The Fore Foot 


In. Sicista subtilis, judging from Tullberg’s figure, the fore foot 
is comparatively broad, short, and unmodified ; the four main 
digits are subequal in length, ‘ne third and Roweealh projecting a 
little beyond the second and fifth, the latter being a little shorter 
than the second; the first, or pollex, is quite short, and springs 
from the middle of the side of the foot close to the distal edge 
of the carpal pad. The digital pads are moderate in size and 
elliptical, and the claws are small. The plantar pad consists of 
three moderately large interdigital elements, subtriangular in 
shape and in contact or nearly so. The carpal pads are large, 
covering about half the area of the sole, the internal element 
being about twice the area of the external, which is in contact 
with it. (Vext-fig. 32, B.) 

The fore foot of Zapus is comparatively long and narrow. The 
carpal pads and pollex are tolerably similar to those of Svcista, 


668 MR. R. I. POCOCK ON THE EXTERNAL 


but the three interdigital elements of the plantar pad are small, 
obscurely defined, circular tubercles widely separated from each 
other. The four main digits, moreover, are longer and thinner 
than in Sicista aud unevenly spaced, the third and fourth being 
close together at the base, but the second is some distance from 
the third, and the fifth fromthe fourth. The claws are compara- 
tively long, and the sole of the foot is granular. ‘he foot isa slight 


exaggeration of the type seen in A podemus sylvaticus, the common 


Text-figure 32. 


A. Right fore foot of Scarturus tetradactylus. 
B. The same of Sicista subtilis (after Tullberg). 
’. The same of Zapus hudsonianus. 
D. The same of Apodemus sylvaticus, for comparison with C. 
KH. Right hind foot of Sicista subtilis (after Tullberg), 
F. The same of Zapus hudsonianus. 
G. The same of Apodemus sylvaticus, for comparison with F. 


English wood-mouse, the third and fourth digits being conjointly a 
little more removed from the second and fifth. (Text-fig. 32. C.) 

The fore feet of the Jerboas (Jaculidz) seem to resemble very 
closely those of Sicista in a general way, except that the claws 
ave much longer, being generally very long, and the interdigital 
elements of the plantar pad greatly reduced in size. In Scarturus 
they are represented by three widely separated circular papille ; 


GHARACTERS OF SCARTURUS AND OTHER JERBOAS. * 669 


in Huchoreutes, according to Sclater, there are only two, corre- 
sponding apparently to the median and outer of Scarturus ; m 
Allactaga indica I can find only the median remaining, whereas 
in Scirtopoda orientalis and J. jaculus all definite trace of them 
seems to have disappeared, the sole of the foot in front of the 
carpal pads being merely marked by integumental creases. As 
in Sicista and Zapus, the carpal pad is double, the outer or ulnar 
element being comparatively small, while the inner or radial is 
very large, especially in Allactaga indica, where it occupies 
practically the whole of the posterior half of the sole. The 
pollex is a mobile cylindrical process, covered above with a large 
nail in Jaculus, Scirtopoda, Scarturus, and Allactaga, but in 
Euchoreutes, according to Sclater, it is armed, like the other 
digits, with a long claw. This author also described the digits 
of Huchoreutes as long, but, judging from the figure, they are no 
longer than in other genera of Jaculidee. As in several Rodents, 
notably the Squirrels, objects are held in the fore foot by folding 
the four digits back upon the large carpal pad, which plays the 
part, as it were, of athumb. (Text-fig. 32, A.) 

The fore foot of Pedetes is short and broad, and has five well- 
developed digits, all armed with long, stout, nearly straight claws 
—the pollex being like the others, but shorter. The digital pads 
are scarcely defined; and the under sides of the digits and the 
short area of the sole distal to the carpal pads are naked and 
granularly roughened. No trace of plantar pads remains. The 
carpal pads are remarkable and, so far as I know, unique. They 
are very large. The inner pad is an elevated thickened cushion, 
beset on the summit with minute spicules. The inner pad is 
laminate and normally fits closely against the outer pad, but is 
independently movable like a valve, so that a deep space separates 
the summits of the two pads. Its free edge, moreover, bears a 
fringe of long hairs. This modification suggests that the carpal 
pads are prehensile and aid in the grasping of food that is 
gripped by the digits with their long claws. These claws are no 
doubt also fossorial. (Text-fig. 35, A—C.) 

From the following summary of the characters of the fore foot 
it will be seen that Pedetes stands apart from the other genera. 
It may be noted, however, that Sicista falls with the Jerboas 
and not with Zapus :— 

a. Fore foot with well-developed pollex armed with a claw as long as 


on the other digits; claws very long, stout, and straight; carpal 
pads separable and prehensile, the movable outer pad with fringe 


OislOn SH AITS Herc Me eae menace OPER Eason oar esac ae aetna mek CULE LCS: 
a'. Fore foot with short pollex armed with a nail or slender claw; claws 

of digits 2 to 5 long or short, but slender and curved ; carpal pads 

normal, confluent, the outer immovable and not fringed. 
6. Foot elongated and narrow, digits unevenly spaced, 3 and 4 in 

EXSKVANINGS) Ok) Bat Gc vocnbuoausoup edd eus eee dedBhmeeeeediae donbcd sosecuadbiubeiie my Aew ave 

b/. Foot short and broad, digits 2 to 5 evenly spaced. 

ec. Pollex armed with small claw (sec. Sclater) ..................... Huchoreutes. 

ec’, Pollex armed with broad blunt nail ......... Sicista, Jaculus, Scarturus, 


Scirtopoda, Allactaga. 


670 ; MR. R. Ll. POCOCK ON THE EXTERNAL 


The Hind Foot. 


The hind foot in Sicista subtilis, according to Tullberg’s figure, 
is much broader and shorter than in Apodemus sylvaticus, but 
closely resembles in those particulars the foot of Ratius norve- 
gicus. The hallux is shorter than in that species and closer to 
the second digit, and the second, third, fourth, and fifth digits 
are subequally spaced and close together, as in the fore foot, the 
third and fourth projecting a little beyond the second and fifth. 
The claws are short and the digital pads normal end semi- 
elliptical. The plantar pad is represented by four separated 
interdigital elements, the median being comparatively large and 


Text-ficure 33. 
tas} 


\ yy! 


A. Right hind foot of Allactaga indica. 
B. The same of Scarturus tetradactylus. 
C. The same of Jaculus jaculus, with hair cut short. 


triangular, the others much smaller and subcircular, the one at 
the base of the hallux being remote from the rest and only a 
little in advance of the external metatarsal pad, which is also 
quite small and subcircular, while the internal metatarsal pad is 
elongated and situated near the inner edge of the foot halfway 
between the heel and the hallucal element of the plantar pad. 
(Text-fig. 32, E.) 

The hind foot of Zapus is very different from that of Sicista. 
It is very long and slender, the second, third, and fourth digits 
being well in advance of the first and fifth, with the second 


CHARACTERS OF SCARTURUS AND OTHER JERBOAS. 671 


farther from the third than the latter is from the fourth, these 
two being closely united proximally. The hallux is moderately 
long and slender, and rises a little behind the origin of the fifth ; 
it reaches to the base of the second, which is shorter than the 
third approximately by its digital pad ; the fifth reaching only 
to the distal end of the narrow webbing tying the third and 
fourth closely together proximally. The claws are long and 
slender, and the digital pads elliptical. The plantar pad is repre- 
sented by four small, isolated, subcircular, interdigital elements 
set on the granular sole in accordance with the disposition of 
the digits. The metatarsal pads are represented merely by the 
internal element, which is a small subcircular pad set one-third 
of the distance along the metatarsal area between the hallux 
and the heel. (Text- ‘fig: 32, F.) 

The hind feet of the Jaculide differ profoundly from those of 
Zapus in the disposition of the digits and other particulars. 
They are markedly perissodactyle, the third digit being the 
longest, set in the middle line, and symmetrically tanked on each 
side by the second and fourth, which are equal in length and 
rise at the same level. Their digital pads are moderately or very 
large, deep, compressed, and laterally grooved, with lobulate 
distal edge. The plantar pad is represented by a larger or smaller 
soft median papilla, pointed apically. The metatarsus is much 
longer than in Zapus, and, as is well known, the three bones 
corresponding to the three main digits are completely fused into 
one. The first and fifth digits may be absent or present. When 
present they rise a long way back on the metatarsus, and, since 
they do not normally touch the ground, have been described as 
functionless. They are slender, with a small normal digital pad 
and thin longish claw. The metatarsal bone proper to each 
remains distinct from the median ‘‘cannon” bone, and, when 
both these digits are retained, they spring from the side of the 
metatarsus at the same level. Progressive reduction in the 
number of digits from five to three is shown in the family. In 
Pygeretmus, Allactaga, and Huchoreutes the first and fifth are 
retained. In Scarturus the first is lost and the fifth retained. In 
Jaculus both are lost. ‘The under side of the digits varies greatly 
in hairiness. In the two Egyptian species, Jéculus jaculus and 
Scirtopoda orientalis, they are thickly fringed on ‘each side with 
long hairs, which curve inwards over the under surface of the 
digits and form a soft hairy cushion. Close to the pads these 
hairs are exceptionally long. They quite conceal the median pad 
from the lateral aspect, but the pads of the second and fourth 
digits are not entirely covered by them. In these two hairy- 
footed species the median plantar papilla is very small and soft, 
and the digital pads are comparatively small. In Scarturas, 
which occurs in the same localities as Seir topoda orientalis, on 
the contrary, the under sides of the digits have no long fringes, 
and the digital pads are very large and deep, and more strongly 
grooved and marginally lobate thi an in Scirtopoda orientalis quad 


672 MR. R. I. POCOCK ON THE EXTERNAL 


Jaculus jaculuws. They conceal the claws from the lower aspect . 
of the foot, but these claws are smaller than in J. jaculus and 
S. orrentalis. he plantar papilla, moreover, is a large conical 
eminenee. ‘The three main digits of Allactaga indica resemble 
those of Scartwrus tetradactylus in the comparatively scanty 
development of hairs on the under side; but the pads do not 
appear to be so large*. Nevertheless, the plantar papilla is a 
large conical eminence as in Scartwrus. (Text-figs. 33, 34.) 

In the four species of this group above described the claw of 
the median digit is considerably smaller than those of the lateral 
digits. 


Text-figure 34. 


SSS STS 
SSS ES 


a 
Taos SS 


SESS 


A. Right hind foot of Scarturus tetradactylus, from outer side. 
B. Lower view of same, with digits closed together. 

C. Right hind foot of Allactaga indica, from outer side. 

D. The same of Seirtopoda orientalis. 

E. Lower view of the same, with digits closed. 


The hind foot of Pedetes, although saltatorial, has the meta- 
tarsus much shorter in relation to the length of the digits than 
in the Jaculide ; but it resembles the hind foot of the latter in 
having the third digit the longest, set in the middle line, and 
flanked on each side by the second and fourth; and, as in the 
Jaculide, the under side of the digits and mostly of the meta- 
tarsus is naked, but covered with the hairs which overlap them from 
the sides. As in Scarturus, there are four digits, the first having 
entirely disappeared, while the fifth is retained. But, apart from 


* The only specimen examined had been for many years in alcohol, and possibly 
the pads were shrunken in size, 


CHARACTERS OF SCARTURUS AND OTHER JERBOAS. 673 


these features, the foot is very different from that of all the genera 
of Jaculide. For instance, the fifth digit forms a uniform series 


Text-figure 35. 


= 


SSS 


SSS 
BZ teyZZ 
LE 0 


nif ad Vasey yee 


Jijéz-e 
SSW 


at va \ 
ZS 
SKK 


Sy 


: SSSVQ 
eZ 


— 


ay Se 
So 


SSA SSS == 2 
MARY: Oca 


N 


\\ 


iy 
\ 


\ 1 
RAS 


: 
< 


WD 


SS 
SSssss= 


A. Left fore foot of Pedetes caffer, from outer side. 

B. Right fore foot of the same, from below. 

C. Two carpal pads of the same, spread (on rather larger scale). 
D. Right hind foot of the same, with hairs cut short. 

q. Left 0 3 with hairs entire. 

F. The same from outer side, showing fourth and fifth digits. 


les] 


c., outer movable carpal lobe with fringe of hair; 4, fifth digit. 


with the next, the space between it and the fourth being equal 
to that between the fourth and third or third and second, 


674 MR. R. I. POCOCK ON ‘THE EXTERNAL 


Again, the claws are broad and strong like pointed hoofs, and 
beneath are completely fused with the digital pads, which are 
represented merely by an ill-defined area constituting the heel 
of the hoof. I can find no trace of plantar pads, although ‘Tull- 
berg describes one at the base of each digit. All that I can find 
is an elevation at the base of each digit formed by the underlying 
bone. That, however, does not constitute a pad. The middle 
area of the foot behind the base of the digits is provided with a 
mat of long hairs which functionally takes the place of the 
plantar pad. Behind this the sole is naked up to and including 
the heel. (Text-fig. 35, D-F.) 

Good characters for distinguishing the families, subfamilies, 
and the principal genera are supplied ‘by the hind feet -— 

a. Claws broad and hoof-like; digital pads scarcely developed, merely 


forming an ill-defined heel to the hoofs; plantar pads suppressed, 

their place taken by a mat of hairs above the base of the digits ... Pedetes. 
a’. Claws sharp and curved, not hoof-like; digital pads normally or 

exceptionally developed ; plantar pad present, the middle of the 

foot above the base of the digits naked. 

6. Foot with separated metatarsals, ‘murine,’ artiodactyle cr nearly 
so; plantar pad represented by four interdigital papille; hallux 
much smaller than fifth digit and higher up the toot, the fifth 
at least reaching the base of “the fourth ; digital pads normal. 

c. Foot short with digits 2 to 5 subequal, subequally spaced, and 
terminally almost in a line; ee and metatarsal Pople 
comparatively large .......... .. Sicista. 

e’. Foot long, with third and founch Aeite) dereninale anal ilose 
together, the second farther from the third, and the fifth still 
farther from the fourth; plantar and metatarsal papillee 
TUOVODIN eceonaccossnocncoco00 ‘ sdoecsuansose CABPDUISs 

6’. Foot with aires neocon ener ecils sell very mone: nae etre” 
markedly perissodactyle; plantar pad represented by a single 
median conical papilla ; hallux, when present, as large as the fifth 
digit and set on the same level, so high up the foot that their tips 
fall far short of the bases of the adjoining digits; digital pads of 
second to fourth large, deep, compressed, and deeply grooved 
laterally. 

d. Fitth digit retained; short fringes of hair on digits 2 to 3; 
plantar papilla very large and exposed *. 

Gs Wve chyeats (hela) WENN be snoccoeandascobceasdcassconecancseda © AUNGICHAGE. 
ef. Mirstidicita(halliux) absentee eercaceecenacerec ee tease ean SCOURS: 

d’, Fifth digit absent ; long fringes of hairs on digits 2 to 3; plantar 
papilla very small, normally concealed by overtolded fringes 
(VE INEHDP Tne oceans dousco cos S00 sao sboconoonosaceg oD0cas 39 counee, GKMCUMIS, SCUPIOAOTG. 


Tails. 

In Sicista and Zapus the tail is long, subcylindrical, tapering, 
and covered with scales and short hairs as in typical murine 
Rodents, the hairs at most forming a slight terminal pencil. In 
the majority of the Jaculide it is more definitely quadrate, 
covered with short hairs, not distinctly scaly, and is provided 
with a distinct terminal tuft, which, however, varies considerably 
in size. In Jaculus jaculus, Scirtopoda orientalis, and Scarturus, 


* The hairiness of the feet in Allactaga may vary specifically according to 
habitat, 


CHARACTERS OF SCARTURUS AND OTHER JERBOAS, 675 


for instance, the hairs of the tuft are long and show a marked 
tendency to a bilateral, distichous, or feather-like arrangement, 
apparently to give support and prevent the end sinking into soft 
sand when used as a prop; and probably for use as a guide-mark 
in the dark, the tuft is usually conspicuously black or brown and 
white, the distal half being white and the proximal black or deep 
brown, and frequently additional white hairs precede the dark- 
coloured portion. The size of the tuft, however, varies even 
within specific limits. It is, for example, very large in Allactaga 
saliens Gm. (= jaculus Pall.) and quite small in Alactagulus 
acontion Pall. and Allactaga elater Pall., judging from Lichten- 
stein’s figures. In the two first-mentioned, moreover, it is 
conspicuously coloured, whereas in the last it appears to be uni- 
formly or nearly uniformly brown. In Huchoreutes, according to 
Sclater, it is moderately developed and bicoloured, but ‘the 
hairs do not seem to be arranged in so distichous a manner as in 
Alactaga.” 

Two genera, however, of this family—namely, Pygeretmus and 
Cardiocranius—have the tail of a very different shape. It is 
narrow and cylindrical at the base, then broadened, depressed, 
and lanceolate, with the apex attenuate, but not tufted at the 
end as in typical Jerboas. Thus two distinct types of tail are 
found in the Jaculide. 

In Pedetes the tail is long and bushy throughout, being 
uniformly and thickly covered with long hairs from base to 
apex. 


Summary of the Characters of the Tails. 
a. Tail subcylindrical or subquadrate. 
6. Tail uniformly and thickly covered throughout with long hairs... Pedetes. 


b’. Tail covered for the most part with short hairs, often tufted at 
the end. 
e. Tail covered with short hairs and rows of scales, at most 
pencilled at the end . Sroiaets See ee ea SUCTSEO ZU DUSs 


. Tail covered with haies,b pat not visibly aenly. with larger or 
smaller terminal tuit . Recetas gies Taeulus: Scirtopoda, 
‘Scarturus, Allactaga, Euchoreutes. 


a’, Tail broad, flattened, and lanceolate, covered with short hairs, 
apically pencilled, not visibly scaly ............ Pygeretmus, Cardiocranius. 


Anus and External Genitalia. 


In Zapus the anus, as in most Rodents, opens in the centre of 
an area of naked skin above the genital orifice, the two orifices 
being separately visible on superficial inspection. In the female 
the vulva is, however, just below the anus and the naked: skin 
surrounding it is continuous above with that surrounding the 
anus. There is no elongated, conical, preputial excrescence and 
no visible external clitoris. In the male the penis, normally 
retrospective, 1s separated by a fringe of hair from the anus 

Proc. Zoou. Soc.—1922, No. XLVI. A6 


676 MR. R. J. POCOCK ON THE EXTERNAL 


Text-figure 36. 


yi 
bie 
Mi 


wl? 
G 


A. Rear end of Scirtopoda orientalis, from behind, with the anal lobe raised 
to show the anal orifice (a.) above the penis (p.). 

B. The same from the side, with the anal lobe in natural position and the 
penis (p.) partially protruded. 

C. Upper side of glans penis of Scirtopoda orientalis, with spikes lying 
forwards. 

D. Lower view of glans penis of the same, with apex dilated. 

E. Dilated tip of glans penis of the same from the front, with spikes 
raised. 

¥. Side-view of glans penis of the same, with spikes partially raised. 

Upper view of glans penis of Jaculus gaculus. 

. Lower view of the same. 

. Tip of glans penis of the same, from the front. 

. Upper side of glans penis of Scarturus tetradactylus. 

. Rear end of S. tetradactylus, with glans partially protruded. 

. Rear end of Pedetes caffer, from behind, with crescentic anus above 

penis. 


SrA eo 


a., anus; g., glandular pit in penis of Scirtopoda orientalis; o., genito-urinary 
orifice; p., penis. 


CHARACTERS OF SCARTURUS AND OTHER JERBOAS, 677 


above, and its glans is slender and subcylindrical, according to 
Tullberg *. 

In all the specimens of Jaculide examined the anal and gene- 
rative orifices are so closely applied that on superficial inspection 
there appears to be a single orifice only, situated upon a thick 
naked excrescence a little distance below the root of the tail. 
In the females, indeed, the vulva and anus are slightly insunk, 
and are closed by a common sphincter. In the males the tip of 
the penis in its natural position is pressed up against the lower 
side of the anal prominence which carries the anal orifice, con- 
cealed from view, on its underside. This prominence is exceed- 
ingly large in Scirtopoda orientalis (text-fig. 36, A, B). 

The penis of Scirtopoda orientalis was first deser ibed by Sonninit 
and Olivier =. The glans is large and oval in shape, and is thickly 
covered with many backwardly directed short spines; but, in 
addition, it is armed with a pair of long slender stiliform spikes, 
like those in the pouch of the glans of Celogenys, Cavia, and 
some other Hystricomorphs. These spikes are rigid, except at 
the tips, which are soft. According to Sonnini, these tips form 
a small yellow button like the anthers of certain flowers; but 
in the specimens examined by me they were pointed and only 
distinguishable by their softness from the rest of the spike. 
Normally these spikes lie forwards upon what is the upper side 
of the penis when this organ is extended from the prepuce and 
directed forwards, and their tips overlap the apex of the glans, 
but they are capable of being elevated or even directed back- 
wards. Beneath the narrowed apex of the glans the large orifice, 
which has an upper and a lower lip, opens, and on each side of 
it there is a shallow, probably glandular, pouch. The lower side 
of the glans is provided with a median, probably glandular. 
cutter, the edges of which, when brought together, form a 
narrow lineal crease (text-fig. 36,C-F). In Jaculus jaculus the 
glans penis is very different, as Anderson and de Winton pointed 
out. It has no spikes, and the small spines seen in Scirtopoda 
orientalis are represented by minute pits, in each of which there 
is sunk a scale with serrulate free edge. There is a median 
thickening on what is the upper side of the penis when this 
organ is turned forwards, and there is a narrow median groove 
below. There is a curved groove above the upper lip of the orifice 
and a small prominence on each side of it, but no glandular 
depression. (Text-fig. 36, G-I.) 

In Scartwrus the glans is oval or cordate from above or below, 
pointed from the side-view, and the terminal orifice is simple. 
It is armed with sharp recurved spines, which are fewer in number 
and relatively larger than in Scirtopoda orientalis, but there is 


* In the specimen I examined the tip of the glans was covered with an extra- 
neous thickening of hard material closely adherent to it. 
+ Voy- en Egypte, i. pp. 173-174 (18900). 
~ Bull. Soc. Phil. Paris, ii. no. 40, p. 121 (1801). 
A46* 


678 MR. R. I. POCOCK ON THE EXTERNAL 


no trace of the pair of long spikes seen in the latter species. 
According to the descriptions of Olivier and Tullberg the glans 
penis of Allactaga saliens closely resembles that of Scartwrus. 
(Text-fig. 36, K, L.) 

The anus and genitalia of the male of Pedetes caffer were 
described and ficur ed by Calori*. To his account Tullberg had 
nothing to add. The anus is situated a long way below the root 
of the tail, much farther than in the Jaculide, and the prepuce 
of the retroverted penis ferms a conical prominence beneath it. 
Superficially the anus appears as a crescentic slit, with the 
concavity downwards; and this appearance results from the 
downward growth of the skin of its upper border forming a 
flap over the anus itself. Calori described a gland-opening in 
the middle line of the perineum between the anus and the penis. 
This does not appear to me a very appropriate description, 
because no orifice is visible on the area named. There is, how- 
ever, the orifice of a gland-opening within the prepuce just ‘above 
the urino-genital orifice when the penis is retroverted and at 
rest. The penis itself is elongated, compressed, and covered 
with minute spicules; but there is no trace in it of the 
glandular pouch which is so characteristic of the Hystricomorphs. 
(Text- -fig, 36, M.) 

a. A large gland-opening within the prepuce of the male on the anal 


side of the genito-urinary orifice ; 5 anus in the form of a crescentic 
slit some distance above the penis . See .. Pedetes. 


a’. No preputial gland on the anal side oP ie pantie -urinary or nines oe 
the penis; anus not crescentic in shape. 
6. Anus espored above the oes of the penis, which is not tucked up 
against it.. Beane Or 3 PaenR Aap Ra RSD 8 CALI DUIS: 


b’, Anus Seen beneath: an excrescence, rhe ine andes ae of which 
the tip of the penis is applied. 
ce. Glans penis spiny with two immense erectile spikes ............  Scirtopodax 


’. Glans penis without spikes. 
he Glans penis elongated, parallel-sided, covered with minute 
pits, each provided with a serrate scale ..csc.csccesseceesves .. dJaculus. 


d’. Glans penis ovate, armed with spines............. Seartwrus, Allactaga. 


The Skull of Scarturus. 


The skull and dentition of Scartwrus agree in essentials with 
those of Allactaga, and differ in many points from those of 
Jaculus—that is to say, the incisors are ungrooved and slightly 
projecting, though not nearly so strongly as in A. mdica, for 
example ; the minute upper premolar is retained ; the zygomatic 
arch is comparatively slender where it forms the anterior wall of 
the orbit ; the channel for the nerve is formed by a deep notch in 
the floor of the antorbital foramen and not a closed tube as 
in Jaculus; the auditory bulle are much less inflated than in 
Jaculus, despite the large size of the ears, and the mastoid is not 
visibly inflated. I cannot, indeed, find any generic characters to 


* Mem. Accad. Sci. Bologna, v. pp. 297-298 (1854), 


CHARACTERS OF SCARTURUS AND OTHER JERBOAS. 679 


distinguish the skull of Scartwrus from that of Allactaga, unless it 
be the lesser prominence of the incisor teeth and the presence of 


Text-figure 37. 


(CEN 


A. Skull of Scarturus tetradactylus, from the side. 

B. The same, from below. The molar teeth which were out of the skull 
are diagrammatically represented. 

C. The same, from above. 


a longish median process projecting backwards from the palate 
over the mesopterygoid fossa, a character observed in the one 
skull of Scartwrus examined. 


The Genus Scirtopoda Brandt. 


In his selections of the type-species for some of the genera and 
subgenera introduced into the family Jaculide by the older 
authors, Thomas cited Dipus mauritanicus Duv. as the typical 
representative of Brandt’s genus Scirtopoda and also of its sub- 
genus Haltomys (Ann. Mag. Nat. Hist. (8) 1. p. 308, 1908). He 
also stated that mawritanicus is identical with or closely allied to 
the large Egyptian Jerboa described by Olivier as Jaculus gerboa, 
the species quoted as Jaculus orientalis in the preceding pages of 
this paper. Except that the ears are relatively and the bodily size 
actually larger in this species than in Jaculus jaculus and the 


680 MR. RB. I. POCOCK ON THE EXTERNAL 


colour slightly different, I can find no difference between the two 
in external characters to which I should attach generic value. 
In dentition and cranial characters they are also much alike. 
But the external genitalia of the male are profoundly different. 
In my opinion these differences are of greater systematic im- 
portance than the cranial and dental characters employed for 
separating generically such forms as Jaculus jyaculus and Dipus 
sagitta*. I propose, therefore, to resuscitate the genus Scirtopoda 
Brandt, separating it from Jaculus by the peculiar armature of 
the glans penis. 

Tam not in a position to give an opinion as to the suggested 
identity between S. mauritanicus and S. orientalis; but Duvernoy’s 
and Lereboullet’s figures and description of the penis of mawzi- 
tanicus (Mém. Soc. Mus. Strasbourg, ili. no. 2, p. 48, pl. iv. fig. 12, 
1842) show that that organ is of precisely the same nature as in 
orientalis. 'The spikes on the penis appear, indeed, to be shorter 
In mauritanicus than in orientalis, but that may be merely an 
individual difterence. 


Conclusions. 


The analytical keys published in the foregoing pages to show 
the incidence of the genera according to the characters described 
attest the isolation of Pedetes from the rest of the genera, and 
thus bear out the prevalent opinion of the present day that that 
genus is In no way related to the Jerboas and Jumping Mice. 
Pedeies, therefore, need not be considered further in the present 
connection. 

The keys also bear witness to the affinity nowadays admitted 
to exist between the Jerboas and the Jumping Mice, but show 
that the two may be sharply distinguished by the structure of 
the hind feet. 

In 1901 Lyon (Proe. U.S. Nat. Mus. xxiii. p. 666), using 
skeletal (mainly cranial and dental) characters, classified the 
Jerboas and Jumping Mice as follows :— 


Family ZAPODID&. 
Subfamily Sminraina (Sminthus, ? Homys). 
a ZAPODINE (Zapus, Napeozapus, Hozapus). 


Family DIPODID#. ; 
Subfamily Diropina (Dipus, Aliactaga, and probably Platycer- 
conus). 
- Hucnorevtin&® (Huchoreutes). 


* [am aware that the use of soft parts for differentiating genera of Mammals 
may be an annoyance to modern systematic mammalogists, who, following the 
fashion set in the United States, like to work from dried skins and skulls. But, 
although the zoo-geographical and other results obtained from that method are of 
great value and interest, 1 am sure that the sooner the swing of the pendulum carries 
us back to the method in vogue thirty years ago of working from fresh or spirit- 
preserved material, combined with skins, the better it will be for the science of 
mammalogy. 


CHARACTERS OF SCARTURUS AND OTHER JERBOAS. 681 


Apart from the alterations necessitated by the nomenclature 
now in vogue and the addition of several recently proposed or 
recently restored generic names to the Jerboa family, I have no 
modifications of this classification to suggest, except the severance 
from the rest of the Jerboas of the two genera Pygeretmus 
(Platycercomys) and Cardiocranius*, which agree in having broad, 
flattened, lanceolate tails, distinguishing them at once from the 
rest of the family. They may constitute the subfamily Pygeret- 
mine, which appears to be a specialised offshoot of the five- 
toed <Allactaga-group. The two genera differ at least in the 
following characters :—Pygeretmus has lost the anterior upper 
premolar, which Cardiocranius retains; the latter has grooved, 
the former ungrooved upper incisors. 

Lyon divided his Dipodine into two groups: (1) Dipus, with 
its subgenera, possessing three toes, a considerably inflated 
mastoid, the upper incisor grooved, the anterior upper premolar 
absent, vamd the antorbital canal for the nerve complete. 
(2) Allactaga, with its subgenera, has more than three digits, the 
mastoids not much inflated, the upper incisors without grooves, 
the small anterior upper premolar present, and the antorbital 
canal for the nerve incomplete. 

ff the above-mentioned characters held good there would be 
strong reasons for separating Allactaga and its kindred from 
Dipus and its allies as a special subfamily Allactagine. But 
there seem to be too many intermediate forms to make such a 
course advisable. In the number of the toes, for instance, 
Scarturus is precisely midway between Dipus or Jaculus and 
Allactaga. In the Asiatic three-toed Jerboa, Dipus, the 
mastoids are much’ less inflated than in the African forms, 
Jaculus and Scirtopoda; Dipus thus connects those genera with 
Allactaga in that particular. In Dipus, too, the anterior small 
premolar is retained as in Allactaga, whereas it is lost in Jaculus 
and Scirtopoda. On the other hand, the small five-toed Asiatic 
Jerboa, Alactagulus acontion, resembles the three-toed African 
genera, Jaculus and Scirtopoda, in the loss of this molar. 

Of the characters mentioned by Lyon, therefore, there remain 
as distinctive only the presence or absence of the grooves in the 


* Satunin, Ann. Mus. Zool. St. Petersb. 1902, p.582. The genus Oardiocranius 
has an extraordinarily wide and inflated cranial portion of the skull, which, as com- 
pared with the facial portion, is much larger even than in Huchoreutes. I associate 
this genus with Pygeretmus on the assumption that the peculiarly modified tail 
has not been developed twice independently within the group. 

+ For the sense in which this genus is here used, sagitta being its type-species, 
see Thomas, Ann. Mag. Nat. Hist. (8) 11. p. 808 (1908). Trouessart seems to have 
been unaware of this paper by Thomas Lhe he published his ‘ Faune des Mamm. 
d’Europe’ in 1910. At all events, he proposes the new name Dipodipus for the 
group of Asiatic three-toed species, with sagitta as the type, for which Thomas 
showed the old name Dipus to be available. Satunin seems to have come in- 
dependently to the same opinion on this point as Thomas (Mitth. Kankas. Mus. 
1907, p. 72). Trouessart also resuscitated Scirtopoda in quite a different sense 
from that in which Thomas employed it (cf. supra, p. 679). But, since Thomas 
was the first reviser of the nomenclature of these Jerboas,. I follow his decisions. 


682 THE EXTERNAL CHARACTERS OF SCARTURUS. 


upper incisor teeth and the closed or open canal for the antorbital 
nerve. Iam not aware whether these are constant differences. 

- Adopting Lyon’s classification with the modifications above 
mentioned, the families and subfamilies of Jumping Mice and 
Jerboas will be as follows :— 


I. Family ZAPODIDA. 
1. Subfamily Sicistina *= SminrHin# (Sicesta). 
2. is ZAPODINE (Zapus, Napeozapus, Hozapus). 


II. Family JacuLip# or DIPoDIDA. 
1. Subfamily Jacutin# or Diropine® (Jaculus, Scirtopoda, 
Dipus, Scarturus, Allactaga, Alactagulus). 
2. 59 PYGERETMINE[ (Pygeretmus, Cardiocranius). 
3 as HuUCHOREUTINAE (Huchoreutes). 


The chief distinguishing characters of the Zapodide and 
Jaculide and of the Sicistine, Zapodine, and HKuchoreutine may 
be found in Lyon’s paper. Those of the Pygeretmine are stated 
above. With regard to the Jaculine, the genera which appear 
to me to be admissible, though there may be others, may be 
distinguished as follows :— 


a. Five or four toes on the hind foot; incisors ungrooved; antorbital 
nerve-canal open above. 
6. Five toes on foot, hallux present. 


ce. Small upper premolar retained .............e cece Allactaga. 
clsSmaill tipper, prem OlarslOstipucanuuncsscerece steer ee iectaises- er ecene Alactagulus. 
b’. Four toes on foot, hallux lost + (skull and dentition as in 
Allactaga) .. Sen sseosete-s | NCORTURUS. 


a’. Three toes aes on a Jeeta Soave incisors anomie ‘conor itel canal 
closed above (? always). 


d. Small upper premolar retained; mastoids less inflated ............. Dipus. 
d’. Small upper premolar lost ; mastoids much inflated. 
e. Glans penis without spikes, with insunk scaly armature ...... Jaculus. 
e’.Glans penis with a pair of large erectile spikes and spinous 
PHBT TE RADUTOH ec Konan dudledo md padoAd Sédoc. Boo bo AeoponsdanaoLoa Usobeddoeoaces | OOMUA/ROCG. 


* See Miller, Cat. Mamm. Western Europe, p. 536 (1912). 


3 


REVISION OF THE GENUS LIGIA ( FABRICIUS ). 


nN on 
Viana 
mye dee 


rie 


: ; ee 
PZ.08. 1922, JACKSON, Flo liam 


REVISION OF THE GENUS LIGIA ( FABRICIUS). 


ON THE ISOPOD GENUS LIGIA. 683 


35. A Revision of the Isopod Genus Ligia (Fabricius). By 
Harotp Gorpon Jackson, M.Sc., F.Z.8.; Reader in 
Zoology in the University of London, at Birkbeck 
College. 

[Received May 11, 1922: Read June 13, 1922.] 
(Plates I. & 11.*) 


ConTENTS. 

Page 

(YER reracenne Mia ae aik ser NAW) Ot tas auc OO 

(2) Introduction.. Ua Rn oe Meet HU: eater eecieeen AG OE 

(8) Surualf@naraciers Ba SAU RPE One at aA RAEN. Se eS: 

(4) Generic and Specific Chimactee saadeekeucnenanedt 685 

(BH) Wind ers CRUG Shs ccocooaco nbc oppdcacodece dos | EM 

(6)Keyatoitheicenus)Wigcal msn t sere e desea OOO 

(7) MUNG RBIS ICAGOG ono 6oecrsiooqeeosney doe eencce aso sccose EXD 
The TDAGIG, COQHHEO Gene socouc 20s Aadnan sadbanoodenanne | ES)IL : 

Oh, TD GG Ed GOUUGISOG uo o3600.000 pxcesconaeecboudeoase | RIL 

8}. TG Get GUAINIROTEED oeboocee eoeee codons pebecoaso. | Ee 

A. Digia occidentalis 2.0.0.1 .....-.2.-2.-----) 692 

5. Tgua cinenascens’ si. ceeey ates) 89S 

Gh ILAGNG CROLCRO scoanacnanonoce cvonseensaseccnoce ORB} 

Ue IDEGTO QU fORSOG, coo ade nda eos dannabaoo ccsopeecosce OWE 

Sb TEGAG CUBUECG), “Geo ochoedcouescoasapnseqonbercscn ) OLE) 

), TAGE GPREKEOUGECS odancnaccsoones sannncesasee ngs etele) 

NO! IOGOL IXHUGHCSIS sss sb che cea caoseeeconnoses, WORLD 

ll. Ligia nove-zealandie@ ..............-.... 697 

U2 wibigea CaUdinranG v0 here see OOS 

13. Te iment tens aciacericd oe catasae UL 

WA), ILAG EE FBCPEGOSBco000° ono dobsaerne ovoadesuantoece OEY!) 

15. Ligia CAELAP DES we ee hen aU eke OLED 

Species described by other aire: Leer a OL 

Doubtful or insufficiently descr heat species ....:. 701 

Bibliography .. Me ven UE net amber ger an sem gO 

Tienlyrationloeiensibintes! AO PRN A ed 7103 


(1) Preface. 


This paper on the genus Ligia is intended to be the first of a 
series dealing with the Terrestrial Isopods. The opportunity of 
undertaking the work was given to me by Dr. W. T. Calman, 
F.R.S., who placed at my disposal the material, now in the 
possession of the British Museum, which belonged to the late 
G. Budde-Lund. The author of ‘Isopoda Terrestria’ evidently 
intended to use this material, which comprises specimens of most 
known species of terrestrial Isopods, for the revision of his great 
work on which he was engaged at the time of his death. He was 
only able to complete the genera Hubelum, Armadillo, andthe sub- 
family Spherillonine, but he left notes of observations on other 
genera which will be referred to in their proper place. I propose 


* For explanation of the Plates see p. 703. 


684 MR. H. G. JACKSON ON THE 


to endeavour to carry on, from the place at which it was left by 
Budde-Lund, the task of making a critical revision of the ‘Isopoda 
Terrestria,’ with the aid of his collection and other material in 
the Museum. 

I should like here to pay tribute to the generosity with which 
Dr. Calman has always put his unrivalled knowledge of the 
Crustacea at my disposal during this investigation ; my grateful 
thanks are also due to my wife for many of the figures illus- 
trating this paper and a great deal of other tedious work, and to 
Mr. Frank Forty for his excellent figures of LZ. pigmentata and 
L. perkinsi. 

(2) Introduction. 


Budde-Lund recognised seventeen species in the genus Ligia, 
five of which he had not seen and therefore did not describe. 
Since the publication of ‘Tsopoda Terrestria’ several new species 
have been created, three of which still stand, some of the doubtful 
species have been set on their feet, and the genus Geoligia has 
been created to contain two purely terrestrial Ligiide. Budde- 
Lund’s descriptions were always in Latin and, valuable as they 
are, were often extremely sketchy; and no serious attempt was 
made to describe thoroughly any species of Zigia until Sars 
in 1899 gave his classic diagnosis and figures of Ligia oceanica. 
Later, in 1901 and 1916, Chilton did an equally admirable 
service for Liaia nove-zealandie and exotica. Not all modern 
descriptions have improved greatly on their predecessors, although 
Miss Richardson has given careful accounts of the American 
species of Ligia. 

Having had the privilege of examining an extended series of 
forms, I have been ina very favourable position for weighing the 
relative importance of various characters in discriminating 
between species. In descriptions of new species, characters are 
again and again cited as specific which are found on examination 
to be generic, but not referred to—either being considered as un- 
important or not having been noticed—by previous authors. For 
example, such a careful and reliable writer as Chilton gives 
“transverse depression just posterior to the angle of the eye, 
interrupted in the middle” as a specific character of Ligia nove- 
zealandice, when, in fact, there is no Ligia of any species I have 
seen in which this “ depression ” is absent. 


(3) Seawal Characters. 


The very marked: sexual differences have been a stumbling- 
block in the recognition of genuine species. 


The secondary sexual characters in which the sexes differ are 
as follows :— 


1. The antenne of the male are usually relatively longer than 
those of the female and are frequently stouter, though not in all 
species. 


ISOPOD GENUS LIGIA. 685 


2. The perzopods often possess well-marked processes or spines 
in the male only, and also frequently bear a plate-like expansion 
covered with file-like serrations on two or more segments of the 
first two or three legs, as has already been remarked by Chilton 
in Ligia oceanica (1899). The feature is constant enough to 
justify the conjecture that it is of value in holding the female 
im pairing. 

3. The sutures between coxal plates and terga are frequently 
well marked on the dorsal surface of three or more segments of 
the thorax of the female, but in the male there is rarely more 
than the faintest indication of the suture. This character will 
be referred to in detail in the description of each species and in 
the discussion on Geoligia. 

4. The male is usually larger than the female, but frequently 
relatively narrower; doubtless this difference is associated with 
the brood-pouch of the latter. 


It follows from this that descriptions of species founded on any 
of the above characters, in which the sex is not named, must be 
looked upon with suspicion. As far as possible I have always 
noted in the following pages characters in which the sexes differ, 
or have stated on which sex the description has been based. 


(4) Generic and Specific Characters. 


Nearly every external character possessed by Ligia has been 
used from time to time for systematic work. As these characters 
are of very unequal value, I will pass them in review and 
endeavour to justify the characters on which I have based my 
descriptions. 

The proportions of the body and the size vary with age and sex, 
but bearing that in mind, they may be a valuable guide. 

The shape of the somites of the body differs very little in the 
different species. As a rule the hinder margins of the first 
three thoracic somites are transverse and the remainder pro- 
gressively more and more curved, and the coxal plates are drawn 
out into progressively longer backwardly. directed spines. The 
first three or four somites of the thorax are about equal in length 
and the others successively shorter. Unless there is any strongly 
marked difference, | have therefore omitted these points in my 
descriptions. 

The general shape of the body cannot be described in other 
terms than “elongate-oval” or the like, so it is unnecessary to 
repeat it for each species. 

The colour in the case of preserved specimens, especially if they 
have been kept in spirit a long time, is almost valueless. 

The surface of the body is always covered with minute 
granulations, and the free margins of the somites are beset with 
minute sete, but in some species the surface is notably smoother 
than in others, 


686 MR. H. G. JACKSON ON THE 


The cephalon has a transverse ‘groove on its hind border and 
two somewhat pear-shaped post-orbital pits (‘interrupted de- 
pression” of Chilton). ‘here are three transverse grooves on the 
epistome in all species. 

The eyes of the same species have been described by different 
authors as ‘moderate,’ ‘‘large” or “small.” Compared with 
some nearly related genera, the eyes of any Zigia would be 
large: it is therefore unnecessary to do more in a specific descrip- 
tion than compare one species with another. 

The abdomen (metasome) has the first two somites covered by 
the last thoracic and without the drawn-out pleural plates possessed 
by the remaining somites. ‘The difference in width between 
abdomen and thorax is more marked in some species than in 
others, but I cannot consider it a good character, as it is distinctly 
affected by the preservation of the specimen ; when the body is 
flaccid and the anterior abdominal somites are more exposed, the 
difference is exaggerated. In many species there are indications 
of lateral oblique carine on the third, fourth, and fifth abdominal 
somites. I have noted the character when it is conspicuous. 

The terminal segment (telson a 6th abdominal sonaite) varies 
slightly with every species and is, therefore, a valuable cha- 
racter. The pesterior margin may be produced into spines or 
blant processes. It may bear a median process which may be 
acute or almost obsolete; the corners may be drawn out into 
postero-lateral processes. On each side above the point of insertion 
of the uropod the margin may bear two processes (sometimes 
little more than undulations), the dner and outer accessory 
processes (PI. IT. fig. 10). 

The key given on p. 689 is based in the first place on the shape 
of the posterior margin between the two inner accessory processes. 
If a median process is present, the margins on either side of it, 
between it and the inner accessory processes of each side, are 
always, however slightly, concave. Long usage has sanctioned for 
this type of telson the inaccurate term “ triangulate.” 

On the other hand, an ‘“‘arcuate” margin is smoothly convex 
between the inner accessory processes. 

It is conceivable that cases may arise which are intermediate 
between the two types, but the material described below presents 
no difficulties. 

The appendages are of great systematic value. 

The first antenne are vestigial throughout the genus. 

The second antenne differ greatly in length, in the character of 
the segments of the flagellum and in the relative length of the 
pedunele and flagellum. I have followed Miss Richardson in 
describing the length of the appendage in relation to the somites 
of the body, but I cannot follow her in attaching importance to 
the relative length of the segments of the peduncle. The 
number of segments of the flagellum is useful as a general indi- 
cation, but as 1t sometimes differs on the two sides of the same 
animal, it must not be taken too seriously. 


ISOPOD GENUS LIGIA. 687 


The mouth-parts differ slightly throughout the genus. It is 
interesting to note that in the majority of species the maxillipede 
(whose palp has always five separate or indicated joints) and 
second maxilla differ from those of Ligia oceanica, the type of the 
genus. The first maxilla and the mandible are, however, very 
constant in structure, on which account I have considered it 
unnecessary to deal with them under each species. 

The perwopods are always biunguiculate. Inthe males of certain 
species the inner edges of the first and sometimes second and 
third leg are deeply suicate. I have not enough material to 
pronounce on the value of this character, but as I have found it 
present in some and absent in other specimens collected at the 
same spot and obviously identical in all other particulars, I feel it 
necessary to regard it with caution until its significance is known. 
The sexual differences have been alluded to above. 

I have not found the pleopods to be of any systematic value. 

The wropods are frequently distinctive both in shape and size. 


(9) The genus Geoligia. 


This genus was founded by Dollfus in 1893 on a single damaged 
male specimen obtained from Valencia (Venezuela) at a height 
of 1200 metres above sea-level. The character on which the 
genus was separated from Ligia was the absence of indication of 
separation of coxal plates from terga on the thorax, added to 
the unusual habitat of the specimen. 

In 1900 the same author described specimens from the Hawaiian 
Islands, obtained at heights varying from 2000 feet to 4000 feet, 
as a new species of the same genus. These differed from the 
holotype in many particulars, and one male specimen was in 
possession of a complete uropod of extraordinary form, the rami 
being composed of several segments. Dollfus accordin gly added 
to the characters of the genus the possession of jointed uropods, 
and his diagnosis is as follows :— 


““Coxopodites (épimeres) non ou trés peu distincts. 
Appendices des uropods articulés. Lereste, et notammement 
les parties buccales, comme dans le genre Ligia—espéces 
terrestres.” 


An examination of the degree of separation of the coxal plates 
from the terga throughout the genus Ligia reveals that it would 
be a most treacherous character on which to found a species, 
let alone a genus. It is variable, even within a species, but 
a generalisation that would be true for the majority of cases 
might be stated thus :—The sutures between the coxal plates and 
terga are always visible to a greater or lesser extent in the 
female on the second, third, and fourth thoracic somites; they 
may be present or absent on the other Poe In the male the 
sutures may be altogether obliterated (e.g. ZL. olfersii) or well 
marked on each thoracic somite (eng. i; occaniea). 


688 MR. H. G. JACKSON ON THE 


The original specimens of the Hawaiian G'eoligia perkinsi are 
preserved in the British Museum. An examination of the 
material shows that any generic distinction based on this 
character must be abandoned, as the females all have this suture 
well marked on the second, third, and fourth somites. 

An examination of the single uropod cannot but raise doubts 
as to the genuine nature of the “joints.” These are at most 
irregular intervals, and show under a high magnification no 
arthrodial modification of tissue such as would be found in a 
natural structure. 

It is therefore suggested that the segments of the rami were 
produced accidentally. Among two dozen specimens of Ligia 
exotica—a species distinguished by the length and slenderness 
of the uropods—I found two specimens which showed “joints” 
in the rami, identical in every respect with those of Geoligia 
perkinst Further, when placed side by side with a uropod which 
had been purposely bent, no difference could be detected. The 
probability that the jointed uropod of Geoligia is of accidental 
origin is therefore exceedingly strong, and the last anatomical 
difference between Geoligia and Ligia is thereby removed. 

If G. simoni and G. perkinsi are compared, it will be found that 
they differ widely from one another in relative size of eyes and 
head, length of antenne, shape of telson, and other points. The 
artificial nature of the genus is shown by the fact that they differ 
more from each other than each differs from certain species of 
Ligia, and ditticulty arises in finding not generic but specific 
differences between these species and various species of Ligia. 

Tn 1915, Pearse (p. 549) described under the name of Ligyda 
richardsone, specimens he collected in Cincinnati, Colombia, at 
3800 feet elevation. This species greatly resembles Geoligua 
simont, and occurs at no great distance from it, and it is very 
probable that coliection of more material from Venezuela and 
Colombia will show that both must be referred to the same 
species. If habitat is to count for everything in the delimitation 
of genera, then Pearse should have made a new genus for 
L. richardsone or have placed it in Dollfus’s genus, of which he 
seemed unaware. His action in retaining the genus Ligia 
for his new species supports my contention that bionomical con- 
siderations should have no weight in the separation of this genus 
from Ligia. 

It may be assumed that these three species have arisen from 
littoral species of Ligia which have migrated inland and become 
slightly modified in so doing. Thus Geoligia perkinsi is almost 
identical with the littoral LZ. hawaiensis, but G. stmoni and 
L. richardsone present « more difficult problem, as the adjacent 
littoral species that have been recorded (L. baudiniana and 
L. olfersii) do not bear any obvious relation to them. Hither 
they are modified from the baudiniana or from an unrecorded 
New World species resembling ZL. ttalica, from which they can 
only with difficulty be distinguished. 


ISOPOD GENUS LIGIA. 689 


I propose, therefore, on the grounds I have given above, to 
unite Geoligia with Ligia, and regard G. simoni and G. perkinsi 
as species of the latter genus. 


(6) Key to the genus Ligia. 


The following species are omitted from the key as being 
insufficiently described or of doubtful validity :—Z. ehrenbergii 
Brandt, LZ. cajennensis Koch, L. australiensis Dana, L. cursor 
Dana, L. vitiensis Dana, and ZL. malleata Pfeffer. 

The general distribution of each species is given as a possible 
aid to identification. The key is to be considered as artificial, and 
not necessarily indicating relationship of forms to one another. 


Posterior margin of telson triangulate ...........0..0cc0 ccc ece see eee ene 2. 
‘ a margin of telson arcuate 9. 
A Process on propodite of 1st leg of male Sh 
i i process on Ist lee of male 6. 


ie longer thankbhonaxinanaley yy eee eeteeeeicnene 


Antennae yshouten than thonaxsmemalelsy, secs nesses hese se uaeceee: 


Or 


Telson with blunt median process and blunt accessory processes. 


hawaiensis (Pacific Islands). 
Telson with acute median process and two acute accessory processes. 


exotica (warm seas). 


Flagellar joints of antenna long and slender. 


¢ 

( Flagellar joints of antenna short and squat... cinerascens (Japan). 

Bi 

( occidentalis (tropical W. America). 


(avait comb of long setz on carpus and merus of 1st leg of male. 
5 ; baudiniana (tropical W. America). 
( Without Comlbsonbilistalectotum all creas sate tana eee ee 7 


meee de 


Antenne as long or longer than thorax ............... 00 ccccse ane eee 8. 
1. Antenne not reaching further than 5th thoracic segment. 
pigmentata (Suez). 
First leg of male deeply grooved ................ olfersii (tropical E. America). 
& First leg of male without grooves ............... perkinsi (Hawaiian Islands— 
terrestrial). 
5 i Wadich of body more than half its length ........0.c.c..00e ves cee eee 10. 
( Wiath Ot bodyAlessithanwhalfeits length’ ss...csseseseceeseeeee eee gle 
Carpus and merus of 1st leg swollen ............. pallasii (N.E. America). 
a has and merus of Ist leg deeply grooved... dilatata (Cape Peninsula). 
le { Antenne as long or longer than whole body .................0...0., 12, 
; UMaine Shorbenithamvbodsya Hueco casas nese cece ee ee ae eeteeeeG 13. 
ie ¢ Uropods longer than body ...................... gracilipes (S. Africa). 


Wiurenad not so long as body ..................... natalensis (Natal). 


690 MR. H. G. JACKSON ON THE 


ace processes of telson produced to sharp points...... 14, 
13. 
Postero-lateral processes of telson obsolete, or small and blunt . 15. 
Antenne as long as or longer than thorax, setose. 
14, nove-zealandie (New Zealand, Chile), 
Antenne shorter than thorax, not setose ...... oceanica (Hurope). 
Eyes large and separated by less than their horizontal length... 16. 
15. 
Eyes small and separated by more than their horizontal length. Wie 
Flagellum of antenna with about 20 long and slender joints. 
P richardsone (Colombia—terrestrial). 
Woh Flagellum of antenna with about 17 short and setose joints. 
glabrata (Cape). 
Flagellum of antenna with about 20 joints ... italica (Mediterranean). 
17. Flagellum of antenna with about 16 joints ...  simoni (Venezuela— 
terrestrial). 


(7) The genus Ligia, Fabricius, 1798. 
I quote Sars’s diagnosis of the genus :— 


“‘ Body regularly oval, or oblong oval, moderately convex 
above, with the metasome not abruptly contracted; last 
segment rather broad, with distinct epimeral plates. Eyes 
large and convex. Antennule very small, with the last 
joint rudimentary, nodiform. Antenne rather strong and 
elongated. Mandibles with a ciliated lappet and numerous 
penicils behind the cutting-part. Maxillipeds comparatively 
short and stout, with the terminal part rather expanded, 
epignath rounded. Legs gradually increasing in length 
posteriorly, dactylus distinctly bi-unguiculate. Opercular 

late of pleopoda sub-branchial. Uropoda more or less 
elongated, basal part not produced inside; rami narrow, 
styliform, subequal, each with a single apical spine.” 


The only point that will not hold for the whole genus is that 
relating to the contraction of the metasome. It may or may not 
be as broad as the mesosome. 

T have thought it desirable to include in the following 
descriptions, for the sake of uniformity, complete accounts of all 
species that I have examined, although three of them have been 
fully described elsewhere. When full synonymy and figures have 
already been published, I have been content to refer to them when 
LT have been satisfied as to their correctness. 

The order in which the characters of each species are dealt with 
is based on convenience in examination, and does not pretend to 
any natural or logical order. Although the males and females of 
a species may differ considerably, I have considered it safe to 
re-describe somé species, and describe a new species from the female 
only, as, with the data given above, it may be predicted with some 
certainty in what characters the male may be expected to differ 
from the female, and to found the specific diagnosis on common 


characters. 


ISOPOD GENUS LIGIA. 691 


. 1. Lieia oceantca (Linneus). 


Ligyda oceanice Richardson (1905), p. 684 (q. v. for complete 


synonymy). 

Ligia oceanica Dahl (1916), p. 32. 

Male specimen described. 

Length 25mm. Breadth 12 mm. 

Surface moderately coarsely granulated. Hyesof moderate size, 
rounded and separated by twice their horizontallength. Antenne 
moderately long and stout ; flagellum reaching as far as the hind 
border of the 4th thoracic somite, the peduncle half-way down 
the 2nd somite. Flagellum with 12 to 14 very short and stout 
segments. Oowal plates divided by very distinct grooves from 
terga of every thoracic somite in both sexes. Abdomen not 
abruptly contracted. TZelson arcuate; the postero-lateral processes 
acute and nearly as long as the middle of the hind border ; 
accessory processes not produced, but the inner forming a 
prominent undulation. Mouth-parts: 2nd maxilla bilobed, with 
two hairy bristles on inner side. Maxillipede palp clearly divided 
into five joints. Ist perwopod of the male only has merus, carpus, 
and propodus produced on inner side to a flat. plate-like expansion, 
with free border fringed with short sete. The surface of this 
expansion is covered by oblique serrations. Uropods 7 mm. 

Disiribution. Hammershus ; Cuxhaven; Portsmouth ; Northern 
Seas. (See Richardson (1905) for complete list.) 

Figured in full by Sars (1899). 


2. Liera PALLASII Brandt. (PI. I. fig. 4.) 


Ligyda pallasti Richardson (1905), p. 682 (q. v. for synonymy). 
Ligyda pallaswi Richardson (1909), p. 125. 


Male specimen described. 

Length 35 mm. Breadth 20 mm. 

Surface coarsely granulated. yes of moderate size, rounded 
and separated by twice their horizontal length. Antenne 
moderately long and stout; flagellum reaching as far back as hind 
border of 5th somite, peduncle to hind border of 2nd somite; 
flagellum with 15 short and stout segments, without large sete 
but densely covered with exceedingly minute sete. Cowal plates 
divided by deep and distinct grooves on every thoracic somite in 
both sexes. Abdomen not abruptly contracted, with prominent 
oblique carine on each side of drd, 4th, and 5th somites. Velson 
arcuate; postero-lateral processes acute, produced as far as or 
beyond middle of hind border ; accessory processes not produced, 
but the inner forms a prominent undulation. Telson twice as 
broad aslong. Mouth-parts as in L. oceanica. Ast, 2nd, and 3rd 
perwopods with carpus and merus expanded as in oceanica. 2nd 
and 3rd legs with small process on propodus not reaching beyond 
the dactyl in both sexes. Uropods 8 mm. 

Proc. Zoou. Soc.—1922, No. XLVIT. 47 


692 MR. H. G. JACKSON ON THE 


Distribution. Litycha; Alaska; California; British Colombia. 
(See Richardson (1905) for complete list.) 

Figures in Richardson (1905). 

This species is nearly related to the foregoing. 


3. LietA GLABRATA Brandt. (PI. I. fig. 5; Pl. I. fig. 6.) 


Ligia glabrata Budde-Lund (1885), p. 263. 
2Ligia dilatata Collinge (1920), p. 475. 


Female specimen described. 

Length 13mm. Breadih 6mm. (Another imperfect specimen 
measured 19 mm. by 9 mm.) 

Surface very slightly granulated or nearly smooth. yes 
of small size, separated by about one and one-half times their 
horizontal length. Antenne of moderate length; flagellum 
reaching as far back as hind border of 5th thoracic somite, 
peduncle to hind border of 2nd segment; flagellum with 15 
to 17 slightly setose, short, and squat segments. The cowal 
plates are divided by deep grooves on the 2nd, 3rd, and 4th 
thoracic somites. Abdomen not abruptly contracted. Telson 
arcuate; postero-lateral processes blunt and little produced; 
accessory processes almost obsolete. Mouwth-parts: 2nd maxilla 
not bilobed, though a crease indicates the usual division; without 
hairy bristles on inner side. Maxillipede with Ist and 5th joints 
of palp distinct; remainder only indicated by deep marginal 
indentations. Percopods typical in form. Uropods 4:5 mm. 

Distribution. Cape of Good Hope; Table Bay. 

Collinge (1920) surmises that ZL. glabrata is possibly an 
immature form of JL. dilatata. These specimens are, however, 
undoubtedly mature, and must be identified with the former of 
Brandt’s species. I have seen no male of this species. 


4, LictA OCCIDENTALIS Dana. (PI. II. figs. 7 & 8.) 


Ligyda occidentalis Richardson (1905), p. 681 (q. v. for 
synonymy). 

Male specimen described. 

Length 25mm. Breadth 11 mm. 

Surface minutely granulated. Wyes large and quadrangular, 
and separated by less than the horizontal length of one eye. 
Antenne moderately long and slender; flagellum reaching as far 
back as hind border of 6th thoracic somite, peduncle to hind 
border of 2nd. Flagellum with 22 long and slender segments. 
Division of cowal plates lightly marked on all thoracic somites. 
Abdomen not abruptly contracted.  Telson very obtusely tri- 
angulate; postero-lateral processes are as long as or longer than 
median process; accessory processes very small. Mowth-parts: 
2nd maxilla weakly bilobed, with no hairy bristles; maxillipede 
with five distinctly marked joints. Propodus of Ist perwopod 
with prominent process on inner side of distal end, which projects 
forwards by the side of the dactyl. Carpus and merus of Ist and 


ISOPOD GENUS LIGIA. 693 


2nd legs and carpus of 3rd flattened and striated as in oceanica. 
Uropods 10 mm. 

Distribution. California; San Francisco. (See Richardson(1905) 
for complete list.) 

Figures in Richardson (1905). 


5. Licia cineRAsceNS Budde-Lund. (PI. II. fig. 9.) 


Ligia cinerascens Budde-Lund (1885), p. 265. 
Ingyda cinerascens Richardson (1909), p. 126. 


Male specimen described. 

Length 27 mm. Breadth 12 mm. 

Surface minutely granular, rather smooth. Hyes large and 
nearly round, separated by their horizontal length or a little less. 
Antenne moderately long and stout; flagellum reaching as far 
back as half-way across the 6th somite; the peduncle half-way 
across the 2nd; flagellum with 25 short and stout segments. 
Coxal plates faintly indicated on all the thoracic somites of the 
male and very distinctly in the female. Abdomen not abruptly 
contracted. Telson broadly triangulate, the median process 
obtuse ; postero-lateral processes produced to acute points reaching 
no further backwards than inner accessory processes; accessory 
processes slight but sharp. J/outh-parts: 2nd maxilla strongly 
bilobed, with two small hairy bristles on inner side ; maxillipede 
with the five joints entirely separate. First yperwopod with 
prominent inner process on propodus in male. Uvopods 10 mm. 
+ (rami broken). 

Msiribution. Hakodate, Japan; Manila or Chile. 

This species is nearly related to Z. occidentalis, but differs from 
it in the telson and the character of the antenne. The segments 
of these are more numerous in L. cinerascens, but short and squat 
so that the whole appendage is relatively shorter than in ZL. occi- 
dentalis. 


6. Liaia exotica Roux. (PI. II. fig. 10.) 


Ligia dentipes Budde-Lund (1885), p. 268. 

¢Ligia malleata Pfeffer (1889), p. 36. 

Ligia exotica Budde-Lund (19086), p. 303. 

Ligyda exotica Richardson (1905), p. 676 (g. v. for complete 
synonymy). 

Ligyda exotica Richardson (1909), p. 125. 

Ligia exotica Budde-Lund (1912), p. 391. 

Ligia exotica Chilton (1916), p. 462. 


Male specimen described. 

Length 27 mm. Breadth 17 mm. 

Surface moderately roughly granulate. Hyes large and separated 
by less than their horizontal Jength. Antenne very long and 
slender; flagellum reaching as far back as hind border of 5th 
abdominal somite, pedunele to hind border of 3rd thoracic somite. 
Flagellum with 28 or more very long and slender segments. The 

i AT* 


694. MR. H. G. JACKSON ON THE 


antennais shorter in the female. he wholeisas long as the thorax, 
and the peduncle reaches half-way across the 3rd somite. Cowal 
plates distinctly marked on all segments in both sexes. Abdomen 
not abruptly contracted. Zelson with acute median process ; 
postero-lateral processes drawn out to equal or exceed the median 
in length; accessory processes long and acute. If outh-parts : 
2nd avail la bilobed, but without hairy bristles on inner side. 
Maxillipede with five joints of palp not clearly divided. First 
perceopod with prominent inner process on propodus in male. I 
find the first three legs of the male to have similar expansions on 
the carpus and merus to the Ist leg of L. oceantea, but Chilton 
does not mention the point, so it may not be universal or present 
at all seasons of the year. Uropods 17-5 mm. 

Distribution. ‘‘ Widely distributed on the warmer shoves of the 
Atlantic, Pacific, and Indian Oceans, and it has been recorded on 
the American coast as far south as Chili and Puntarenas” (Chilton 
(1906)). Budde-Lund’s specimens are as follows:—Aden; 
Bagamoja, Shellah-Lama, KH. Africa; Bissao; Brazil; Colombo ; 
Malacca; Nagasaki; Rio de Janeiro; Singapore; Pulo Milo, 
Tonga (Malay Archipelago). (See Richardson (1905) for further 
list. 

eesttend describes under the name L. denéipes a species to 
which he gives no character that distinguishes it from ewotica. A 
careful examination of his specimen reveals no cause why this 
species should be any longer separated from L. exotica. 

Figured and described 1 in detail by Chilton (1906). 


7. Licta otFersit Brandt. 
Ligyda olfersti Richardson (1905), p. 674 (q. v. for synonymy). 


Female specimen described, owing to mutilated condition of 
the males in my possession. 

Length 19 mm. Breadth 9 mm. 

Swrface minutely granular, rather smooth. yes of narrow- 
oblong shape, separated by less than their horizontal length. 
Antenne long and slender ; flagellum reaching as far back as hind 
margin of thorax, peduncle to hind margin of 2nd thoracic somite. 
Flagellum with 26 (broken) long and narrow segments. Cowal 
plates divided by deep grooves on 2nd, 3rd, and 4th thoracic 
somites. In the male there is no indication of the line of fusion. 
Abdomen not abruptly contracted. Telson broadly triangulate ; 
Bestenor ater processes acute, but not longer than inner accessory 
processes ; accessory processes strongly marked. Mouth-parts : 
2nd maxilla weakly bilobed, without hairy bristles; maxillipede, 
five divisions only indicated on margin and not going right across 
the palp. Percwopods of female typical in form. In the male the 
carpus and merus of the Ist and 2nd are deeply excavated on the 
inner side. Uropods 11 mm. 

Distribution. St. Thomas, Trinidad; Florida; Brazil. 

Figures in Richardson (1905). 


ISOPOD GENUS LIGIA. 695 


8. Liera rratica Fabricius. (Pl. 1. figs. 11 & 12.) 


Tagia italica Budde-Lund (1885), p. 269 (q. v. for synonymy). 
Ligia ttalica Budde-Lund (1908 a), p. 11. 


Male specimen described. 

Length 85 mm. Breadth 3-5 mm. 

Surface nearly smooth; scattered minute granules. Myes of 
moderate size, somewhat rectangular and separated by horizontal 
length of one eye or a little less. Antenne long and slender ; 
flagellum reaching as far as hind border of 6th thoracic somite, 
peduncle to the 2nd. Flagellum with 18-20 long and slender 
segments. Cowal plates either exceedingly faintly separated or 
the division is apparently obsolete. In the female the separation 
is marked, but not very distinctly, on the 2nd, 3rd, and 4th 
thoracic somites. They are but little produced backwards on any 
somite. Abdomen abruptly contracted ; lateral margins of somites 
only very slightly produced backwards. Telson arcuate ; postero- 
lateral processes obsolete; only slight undulations mark the 
accessory processes. Mowth-parts: 2nd maxilla strongly bilobed, 
without hairy bristles; maxillipede with five joints indistinctly 
separated. DPercwopods typical in form. Uvropods 4 mm. 

Distribution. ‘* A common species on the sea-shore in almost all 
the countries adjacent to the Mediterranean Sea, also found in 
the Madeira and Canary Islands” (Budde-Lund (1908 @)). 

I agree with Budde-Lund in looking upon Brandt's ehrenbergit 
as a synonym, but, also with him, | am doubtful of the identity 
of the specimen deseribed by Dana under that name (1852). The 
matter is further discussed under /. gracilipes. 


9. Ligeia eRaciuirEs Budde-Lund. (PI. II. figs. 13, 14, & 15.) 


Tagia gracilipes Budde-Lund (1885), p. 270. 
¢Ligia ehrenbergui Dana (1852), p. 738. 
¢Ligia dilatata Collinge (1920), p. 475. 


The somewhat macerated condition of the specimens makes 
it impossible to give a complete description of any one. The 
following account applies to the female, and as I cannot be positive 
that any of the specimens were males, [ have not referred to the 
charaeters of that sex. 

ee 7mm. Sreadth 3°5 mm. 

- Surface minutely granulated. yes large and separated by less 
than their horizontal length. Antenne extraordinarily long and 
slender. Whole appendage one-quarter as long again as thorax 
and abdomen together; peduncle reaching as far back as hind 
border of penultimate abdominal somite. Flagellum with 22 
(Budde-Lund) very long and slender segments. (A separate 
antenna had 28 segments; each segment was about five times as 
long as broad.) Cowal plates separated by deep grooves on 2nd, 
3rd, and 4th thoracic somites, and distinctly marked on the 5th. 
Abdomen abruptly contracted. Telson arcuate ; the postero-latera] 


696 MR. H. G. JACKSON ON THE 


processes blunt and but little produced; accessory processes 
almost obsolete. Mouth-parts: 2nd maxilla without any trace of 
division into two lobes; without hairy bristles on inner side. 
Maxillipede indistinctly divided into five segments. Perwopods 
very slender, but otherwise typical in form. The Usvopods were not 
attached to any specimen, but a separate appendage was 12 mm. 
in length, the base being 3 mm. and the rami exceedingly long 
and slender. They are stated by Budde-Lund to be longer than 
the body. 

Distribution. §. Africa, ‘“ e ‘ Laudana’ exampla pauca in museo 
Simon asservantur ” (Budde-Lund). 

Although the material from which this species was made is 
poor, there is no doubt, I think, that it isa good one. Collinge, 
(1920), who had only Budde-Lund’s short description to guide him, 
discussing the point, suggests that it is a “young form of some 
species,” admitting, however, that young specimens have, as arule 
short antenne, whereas the gracilipes is notable for the great 
length of the antenne. He seems to overlook the quite extra- 
ordinary length of the uropods. Dana (1852) identified provision- 
ally as L. ehrenbergit Brandt, specimens found at Madeira. His 
description and figures of these specimens apply to L. gracilipes in 
every particular but size and locality. The size he gives as 
between 17 mm. to 19 mm. long, and 7 mm. to 8 mm. broad— 
more than double the dimensions of Budde-Lund’s specimens. 
The locality is N.W. Africa as against 8. Africa of the 
gracilipes. It seems as if Dana would have been justified in 
creating a new species for the reception of the specimens, but as 
he did not do so, they must be identified with Z. gracilipes if 
further material should prove them to be the same. I do not 
think they should be so identified in the present state of our 
knowledge. 


10. Licta HAWAIENSIS Dana. (PI. II. fig. 16.) 


Ligia hawaiensis Dana (1852), p. 740. 
Ligia hawaiensis Budde-Lund (1885), p. 271. 
Ligia vitiensis Stebbing (1900), p. 646. 


Male specimen described. 

Length 19° mm. Breadth 7-5 mm. 

Surface minutely granular. Hyes large and quadrangular and 
separated by less than their horizontal length. Antenne very 
long and slender ; flagellum reaching as far back as hind border 
of 3rd abdominal somite ; peduncle half-way across 3rd _thoraeic 
somite; flagellum with 30 long segments, each about twice as 
long as it is broad. The antenne in the female reach to the 
hind border of the thorax, and the flagellum has about 26 segments. 
Coxal plates scarcely, if at all, separated. In the female, deep 
erooves mark the distinction on the 2nd and 3rd thoracic somites, 
but indications are absent or exceedingly faint on other somites. 
Abdomen abruptly contracted. Telson triangulate ; median process 


ISOPOD GENUS LIGTA. 697 


well marked ; postero-lateral processes acute but short; accessory 
processes well marked. Mouth-parts: 2nd maxille weakly lobed, 
without hairy bristles on inner side; maxillipede with palp deeply 
lobed on margin into five portions, but segments otherwise in- 
distinetly marked. First perwopod with prominent inner process 
on propodus in male only. Expansions similar to those of 
oceanicw on carpus and merus of this leg. Uropods (separate) 
(15) rosea 

Distribution. Hawaiian Islands; Molokai; Matadona, China 
Straits, British New Guinea. 

T have examined the specimens of the Willey Collection labelled 
L. vitiensis, and identify them, without possible doubt, as this 
species. 

Figured by Dana (1852). 


11. LigtA NOV #-ZEALANDIA Dana. 


Tigia novi-zealandie Dana (1852), p. 739. 

Ligia cursor Budde-Lund (1885), p. 265. 

Ligia nove-zealandie Chilton (1901), p. 106 (¢. v. for complete 
synonymy). 

Tigia nove-zealandie Chilton (1909), p. 287. 

Ligia nove-zealandice Chilton (1911), p. 568. 


Male specimen described. 

Length 12mm. breadth 5°5 mm. 

Surface minutely granular. Hyes rather small and quadrangular, 
and separated by twice their horizontal length. Antenne very 

long and slender ; flagellum reaching as far back as hind border 
of 4th abdominal somite; peduncle to hind border of 3rd thoracic 
somite. Flagellum with 20 small and setose segments. The 
antenne are shorter in the female. The whole is as long as the 
thorax, and the peduncle reaches half-way across the 3rd somite. 
The segments of the flagellum are more setose than in the male. 
Coxal plates very faintly marked or absent. In the female they 
are marked by deep grooves on the 2nd, 3rd, and 4th thoracic 
somites. Abdomen abruptly contracted. TZelson arcuate ; postero- 
lateral processes acute and produced, but shorter than middle of 
hind border ; accessory processes marked by slight undulations. 
Mouth-parts: 2nd maxilla with no trace of division into two 
lobes; without hairy bristles on inner side. Mawillipede with Ist 
and 5th joints only, completely separated, remainder only 
indicated by indentations on inner side. Ist and 2nd perwopods 
with carpus much swollen in male only; subchelate. Uropods 
5 mm. 

Distribution. Coast of New Zealand; Steward Island ; Sunday 
Island, Victoria; Chile; Juan Fernandez. 

I found that Budde-Lund’s specimens, labelled Ligia cursor, 
from Juan Fernandez (and described by him under that name 
(1885)) agreed in every particular but length of antennz with 
Chilton’s description (1901) of JZ. nove-zealandie. These 


698 MR. H. G. JACKSON ON THE 


specimens were all females. An examination of Chilton’s 
original specimens of novee-zealandie showed that the antenne 
were shorter in the female than in the male, and that the females 
agreed entirely with Budde-Lund’s cursor. Fragments of a 
specimen, labelled “ Z. cursor (male),” from Chile, in Budde-Lund’s 
collection, proved to have subchelate Ist legs like those of the 
male of the nove-zealandiw. The cursor described by Budde- 
Lund is therefore synonymous with Dana’s nove-zealandie. 

Dana’s original account of LZ. cursor (1852) figures and describes 
the telson as triangulate. Budde-Lund was niiilled by the locality 
of his specimens into crediting Dana with an error on that point, 
and hence gave a description of L. nove-zealandic from S. America 
(specimens of which he thought he had not seen) under the 
name of LZ. cursor. The latter species has not apparently been 
found since Dana described it from ‘‘ Valparaiso.” 

The matter is of considerable interest, as Chilton has already 
had occasion to observe (1909 a & 1915) the similarity of fauna of 
South America and New Zealand in connection with other 
terrestrial and aquatic forms. 

Figured and described in detail by Chilton (1901). 


12. Ligra BAUDINIANA Milne-Edwards. (PI. II. figs. 17 & 18.) 


Ligyda baudiniana Richardson (1905) p: 1 OSI (ae acaitOr. 
synonymy). 


Male specimen described. 

Length 20mm. Sreadth 9 mm. 

Surface rather coarsely granular. Hyes large and separated by 
less than their horizontal length. Antenne long and slender ; 
flagellum reaching as far back as hind border of last thoracic 
somite, peduncle to hind border of 2nd. Antenne slightly 
shorter in the female. Flagellum with 29 small and slender 
seoments. Coxal plates very faintly marked on 2nd, 3rd, and 4th 
thoracic somites. In the female they are very distinct on the 
corresponding somites. Abdomen not abruptly contracted. 
Telson triangulate; median process bluntly rounded; postero- 
lateral processes acute, but not projecting as far back as median 
process ; accessory processes represented by undulations. J/outh- 
parts: 2nd maxiila bilobed, but without hairy bristles on inner 
side. Maxillipede palp with five distinctly separate segments. 
Propodus and carpus of 1st perwopod (in male only) each produced 
on inner side to a flat plate-like expansion ; free borders each 
fringed with a comb of exceptionally long and markedly equal 
sete. The plates are obliquely striated. Uropods (separate) 
12-5 mm. ‘The uropods on the female specimens were shorter in 
proportion than these, which possibly, however, did not belong to 
the specimen described above. (In an 18mm. female the uropods 
measured 8°5 mm.) 

Distribution. Bermudas; Mexico; Yucatan; the Bahamas ; 
Progriso, Colon. 


ISOPOD GENUS LIGIA. 699 


Budde-Lund (1885), without having seen a specimen, regarded 
this species as a synonym of JZ, exotica. ‘The features which dis- 
tinguish it from ZL. exotica were fully pointed out by Richardson 
(1902), with additional evidence for its separation from that 
species. Chilton (1916), who had not seen a specimen, remarks 
that it ‘“‘ may be difficult to find characters that will distinguish 
between them in all cases.” 

Among Budde-Lund’s material, I found a tube bearing a name 
of an undescribed species of Ligia which he evidently intended 
to describe in the course of his revision of ‘ lsopoda Terrestria.’ 
An examination of the specimens showed at once that they were 
identical with Milne-Edwards’s baudiniana, which had obviously 
been overlooked by Budde-Lund. This furnishes a further proof 
of the complete independence of this species from the exotica, as 
upheld by Richardson. 

Figures in Richardson (1905). 


13. Licta PiemenTATA, sp.n. (PI. I. figs. 2 & 3.) 


The material consisted of a single female specimen and a few 
appendages which may have belonged to a male. 

Length 14mm. Breadth 5 mm. 

Surface slightly granulated, nearly smooth.  Colowr cream, 
with scattered minute black spots. Hyes large, quadrangular and 
separated by their horizontal length. Antenne of moderate 
length; flagellum reaching as far back as half-way across 5th 
thoracic somite, peduncle half-way across 2nd somite. Flagellum 
with 18 short and broad segments. Cowal plates with suture lines 
exceedingly lightly marked on 2nd, 3rd, and 4th thoracic somites 
only. They are produced but little backwards on any somite. 
Abdomen not abruptly contracted. T'elson triangulate ; median 
process bluntly pointed ; postero-lateral processes deute but short 
inner accessory processes small and blunt. Mouwth-parts: 2nd 
maxilla strongly bilobed; without hairy bristles on inner edge. 
Maxillipede palp with joints completely separated. Percopods 
typical in form. <A separate one (male?) had a thick bunch: of 
sete on the dactyl, overthe uneuis. Uropods6mm. Base 2:5 mm., 
rami 3°D mm. 

Distribution. Suez. 

Budde-Lund had labelled the tube containing this specimen 
with the aboye specific name, but had apparently published no 
description or figures of it. 


14, Liata perKinsi (Dollfus). (PI. I. fig. 1.) 
Geoligia perkinsi Dollfus (1900), p. 525. 


Male specimen described. 

Tength 18mm. Breadth 8 mm. 

Surface almost smooth. WHyes large and separated by their 
horizontal length ora little less. Antenne very long and slender ; 


700 MR. H. G@. JACKSON ON THE 


flagellum reaching as far back as hind border of 4th abdominal 
somite ; peduncle half-way across 3rd thoracic somite. Flagellum 
with 30 very long and slender segments. Cowal plates show no 
sign of division from terga, but in the female the sutures are 
strongly marked on the 2nd, 3rd, and 4th thoracic somites. 
Hinder thoracic somities and free abdominal somites with postero- 
lateral angles much drawn backwards and very acute. Abdomen 
not abruptly contracted. Telson triangulate; median process 
blunt; postero-lateral processes acute and drawn out, but not as 
far back as inner accessory processes. Both accessory processes 
represented by undulations. Mouth-parts: 2nd maxilla weakly 
bilobed and without hairy bristles. Maxillipede with five separate 
joints to the palp. Perwopods typical in form. Uropods 141mm. 

Mstribution: Hawaiian Islands, Karrai 6000 ft., Oloa 2000 ft. 

Although this species is apparently purely terrestrial, it shows 
no modification whatever from the typical littoral Ligiide. The 
reasons for removing it from Dollfus’s genus are given in the 
beginning of this paper. 


15. Ligta NATALENSIS Collinge. 
Tigia natalensis Collinge (1920), p. 474. 


Male specimen described. 

Length 10mm. Breadth 4:5 mm. 

Surface minutely granular, rather smooth. Hyes moderately 
large and separated by their horizontal length. Antenne ex- 
ceedingly long and slender; whole appendage longer than body in 
male, and as long as body in female; peduncle reaching as far as 
hind border of 4th thoracic somite in male, and half-way across 
that somite in female. The flagellum has 20-23 long, slender, and 
distinctly setose segments. Cowal plates show no sign of division 
from terga, but are faintly marked on 2nd, 3rd, and 4th thoracic 
somites of female. Abdomenabruptly contracted. Zelson arcuate; 
postero-lateral processes only very slightly produced ; the acces- 
sory processes obsolete, MJouth-parts: 2nd maxilla not divided 
into two lobes, without hairy bristles on inner side. Maxillipede 
with Ist and 5th joints of palp separated; remainder only 
indicated by deep marginal indentations. Perceopods slender, but 
of typicalform. Uropeds: No specimens of this size had uropods 
attached. Ina male specimen 7 mm. in length they measured 
4 mm.; a female 10°5 mm. had uropods of 4 mm. 

Distribution. Umklali, Winkle Spruit Beach, South Coast, 
Natal. 

This species is nearly related to Z. gracilipes, but is distinguished 
from it by the more massive form of body and appendages and the 
smaller uropods. Iam unable to confirm Collinge’s observation 
that the flagella of the antennez of 7 to 9 mm. specimens of nata- 
lensis have fewer joints than those of gracilipes. In most cases, 
specimens of that size were as mature in form as larger specimens. 


Figured and described by Collinge (1920). 


ISOPOD GENUS LIGIA. 701 


I have not seen specimens of the following species :— 


16. Ligta pILATATA Brandt. 


Tigia dilatata Brandt (1833), p. 171. 

Lygia dilatata Krauss, Siidafrik. Crust., 1843, p. 62. 

Ligia dilatata White, List Crust. Brit. Mus. 1847, p. 98. 

Ligia dilatata Budde-Lund (1885), p. 262. 

Ligia dilatata Stebbing, Ann. South African Mus. 1910, vi. 

p. 437. 

Ligia dilatata Collinge (1920), p. 475. 

Distribution. Cape Peninsula. 

Collinge has re-described and figured this species in the last- 
named work, but has added little to Budde-Lund’s previous 
description. Reasons for regarding L. glabrata and L. gracilipes 
as independent species and not as shown in Collinge’s synonymy 
are given above. 


17. Lica RICHARDSON (Pearse). 
Ligyda richardsone Pearse (1915), p. 549. 


Distribution. Sierra Nevada de Santa Marta, Colombia, at 
3800 ft. elevation. 


Fully described and figured by Pearse. 


18. Liera stmont (Dollfus). 

Geoligia simoni Dollfus (1893), p. 343. 

Distribution. Valencia, ‘‘ vers 1200 métres d’altitude, dans les 
foréts.” 

This species only appears to differ from the preceding in the 
greater relative breadth of the body and the larger size of the eyes. 
Dollfus’s description is based on a single male specimen, however, 
and occurring as they do so elose together, it is probable that 
collection of further material will show them to be identical. 

Figured and briefly described by Dollfus. 


The following species are insufficiently described or of doubtful 
validity :— 

19. Lieia CAJENNENSIS Koch (1847). 

20. Lica AUSTRALIENSIS Dana (1852). 

21. Licia cursor Dana (1852). 

22. Lieta viTrENsis Dana (1852). 

23. Licta MALLEATA Pfeffer (1889). 

Distribution Bagamoyo. 

The description of this species and the locality in which it was 


found suggests its identity with /igia ewotica. Pfeffer does not, 
however, mention the character of the Ist leg nor the sex of the 


702 MR. H. G. JACKSON ON THE 


species he described. If it was a male it differs from L. exotica 
in the absence of a process on the propodus, but if a female 
it is probably synonymous with L. exotica. 


Bibliography. 

Branpt, J. F. 1838. Conspectus monographie Crustaceorum Oniscodorum 
Latreillii. Bull. dela Société Imp. des naturalistes de Moscou, vi. 
15) eke = 

3UDDE-LuND, G. 1885. Crustacea Isopoda Terrestria. Copenhagen. 

Buppn-Lunp, G. 1908a. Terrestrial Isopods. Swedish Zool. Exped. to Egypt and 
the White Nile, 1901, p. 11. 

Buppg-Lunp, G. 19086. Isopoda von Madagaskar und Ostatrika. Voeltzkow. 
Reise in Ostafrika in den Jahren 1903-1905, Bd. ii. p. 303. Stuttgart. 

Bupper-Lunp, G. 1912. Terrestrial Isopods. Trans. Linn. Soc., Zool. 2nd ser. 
vol. xv. p. 391. London. 

Critton, C. 1899. Notes on the Sexual Characters of Ligia oceanica. Ann. & 
Mag. Nat. Hist. ser. 7, vol. iii. March, p. 197. London. 

Cuitton, C. 1901. The Terrestrial Isopods of New Zealand. Trans. Linn. Soc., 
Zool. 2nd ser. vol. viii. part 4, May, p. 99. London. 

Cutttron, C. 1909a. The Crustacea of the Subantarctic Islands of New Zealand. 
The Subantarctic Islands of New Zealand, pp. 793-807. Wellington, 
N.Z. 

Cuitton, C. 19096. Additions to the Terrestrial Isopoda of New Zealand. Trans. 
N.Z. Inst. vol. xlii. p. 287. Wellington, 1910. 

Cuintton, C. 1911. The Crustacea of the Kermadec Islands. Trans. N.Z. Inst. 
vol. xliii. 1910, p. 568. Wellington. 

Critton, C. 1915. Deto,a Subantarctic genus of Terrestrial Isopods. Journ. Linn. 
Soc., Zool. vol. xxxii. no. 219, p. 485. London. 

Cuinton, C. 1916. Fauna of the Chilka Lake. Some Terrestrial Isopods from the 
shore of the Lake. Mem. Indian Mus. vol. y. p. 461. 

CoutincE, W. EH. 1920. Contributions to a Knowledge of the Terrestrial Isopoda of 
Natal. Part 1. Annals Natal Mus. vol. iv. pt. 2, November. 

Daunt, F. 1916. Die Asseln oder Isopoden Deutschlands, Jena, p. 32. 

Dana, J.D. 1852. Crustacea: U.S. Explor. Exped. vol. xiii. pt. ii. Philadelphia. 

Doutrus, A. 1898. Isopodes Terrestres. Voyage de M. E. Simon au Venezuela. 
Ann. Soc. Ent. Fr., Décembre, p. 343. 

Dotirus, A. 1900. Crustacea Isopoda: Fauna Hawaiiensis, vol. ii. pt. v. p. 525. 
Cambridge. 

Kocn, C. L. 1847. System der Myriapoden, mit den Verzeichnissen u. Berichtig- 
ungen zu Deutschlands Crustaceen etc. Heft. 1-40, Regensburg, 
p. 212. 

Prarsz, A. S. 1915. An account of the Crustacea collected by the Walker 
Expedition to Santa Marta, Colombia. Proc. U.S. Nat. Mus. vol. 49, 
p. 649. Washington. 

PrEFFER, G. 1889. Ubersicht dervon Herrn Dr. Franz Stuhlmann in Agypten, auf 
Sansibar und den gegentiberliegenden Festlande gesammelten ... und 
Krebse. Jahrbuch der Hamburg. Wiss. Anstalt. vi. Jahrgang, Zweite 
Halfte, 1888, p. 36. Hambure. 

RicHaRrpson, Harriet. 1902. The Marine and Terrestrial Isopods of the 
Bermudas, with descriptions of New Genera and Species. Tvrans. 
Connecticut Acad. Sciences, vol. xi. Jannary, p. 806. 

RicHaRpson, Harrier. 1905. A Monograph on the Isopods of North America. 
Bull. U.S. Nat. Mus. no. 54, p. 673. Washington. 

RicHarpson, Harrinr. 1909. Isopods collected in the North-West Pacific in 1906. 
Proc. U.S. Nat. Mus. vol. 87, October, p. 125. Washington. 

Sars, G.O. 1899. Crustacea of Norway. Vol. ii. Isopoda. Bergen. 


Steppine, T. R. R. 1900. On Crustacea brought by Dr. Willey from the South 
Seas. Zoological Results, Part v., December, p. 646. Cambridge. 


Fig. 


ISOPOD GENUS LIGIA. 


EXPLANATION OF THE PLATES. 


Prate I. 

1. Dorsal view of Ligia perkinsi. X65. 

2. - » tL. pigmentata, sp.n. X95. 
3. Telson of L. pigmentata, sp.n. X13°5. 
4. LZ. pallasii. 2nd leg of male. 5. 

5. L. glabrata. ‘Telson and uropods. 

Prats If. 

6. Ligia glabrata. Antenna. 9. 

7. LL. occidentalis. Telson. 7. 

8. . Antenna. 6. 

9. LZ. cinerascens. Antenna. X65. 
10. LZ. exotica. 'Telson. X10. 
ll. 2. italica. Antenna. X77. 
12. S Telson. 16. 
13. LL. gracilipes. Antenna. 7-7. 
14. oy Uropod. 7. 
i165, 5 Telson. X17. 
16. L.hawaiensis. ‘Velson. X13. 
17. L. baudiniana. Velson. 11. 
18. 5 Ist lez of male. © x9. 


<M 


703 


ee he 

aa, can Wises: acs bi 
ee ey xe uy 
rh F 


THE HYMENOPTEROUS FAMILY STEPHANIDA. 


705 


36. Monograph of the Hymenopterous Family Stephanidee. 


By Ernest A. Euusorr, F.Z.8., FES. 
[Received March 31, 1922: Read May 9, 1922. | 
(Text-figures 1-8.) 


IND EX. 
Page 
BorHRIocERUS. Fanatopus (cont.). 
americanus Sichel.................. 736 brevicollis Hderl...........0000008 
europeus Sichel......... .......... 730 celebensis Szépl. eae 
ane carpe ae Wi astaaaneeii 
serrator Bab. ............ce0e00eereee 718 conradti Hnder Simian 
fasciatus Szépl. .............2-----+- 
DIASTEPHANUS. fernandopoensis Schultz ......... 
alutaceus Morley ER Ee Vie fon flavicollis Cam. ed aie cee 
lonboneeynits JOIWMOEP aso soonos soooDec6G 805 formosanus Hinderl. .........06c05- 
birdi Szépl... 804 Suscinervis Cam. BeEeee 
brevipetiolatus ‘Ender eal MD velae 816 HAVEN bIc! TAREE sbbotossodoocoooussed 
carinifrons Hnderl. oo kent indicus Enderl. drs a a 
celebensis Szépl. son ooo noGrba.0an oro 822 var. sumbana Taaledl. LY a ee 
chinensis HiUM@oté ..005. 202... 20 -- 814 var. sulcaticollis Enderl....... 
dohrni Hnderl. . “AM asreaseneh hisee | TNR VAG OLN Wangs dneovabeesees 
elegans ELT OG Ge as anne ts 810 iridipennis Elliott.. 
fasciatus Szépl. Fa SOR BeIdOGEOdGoe 826 lacte1pennis Ries. 
flaviceps Hlliott..................... 824 longicauda Elliott ............... 
flavidentatus Hnderl. spsbog,) 0S) lonsicollisiCancmees teen sees 
flavifrons Hilliott ...............-.. 825 natalicus Westw. ............ 
flavomaculatus Hnderl. ......... 807 nigripes Szépl. ; 
flavonotatus Elliott odaDosooounaoDe 826 nova-guineeners Szépl. . 
frontilinea Morley................-- 813 ocellatus Huliott.. Shae 
fuseidens -Kieff. .................-... 804 piceicornis Chie me 
fuscinervis Cam. .............00... S24 DUNChabUseHULOCG ane ee 
CARCHINS IPGIA | /Saso8on dso neada sss cocdec 805 | ruficeps Smith tet eee oo 
leucodon Kieff. PEGE Ot SOD: rugiceps LORE OA Seales eae 
leucodontus Schletté. ............... 810 schlettereri Hinder. 
maculifemur Hnderl. 819 similis Szépl. . 
nigripes SZépl.\--.....+.--.--.-..--- 812, simpsoni Kieff. . 
nova-guineensis Szépl. ............ 807 sulcaticollis Hhderl. . 
pallescens SCRUCEE Ee en Ud sumbanus Hnderl.. es 
parviceps Enderl. ...........0.000-- 812 togoensis Stadlm. .............0066 
quadridens Elliott ............... 818 turcomanorum Semenow ......... 
rothkirchi Schalthess ...........- 808 TEWPNIG(S0YS WEHIUAOAP toc ahsiens doceceban 
ruficollis Hnderl. ................. 819 
salomonis Morley .. 891 | Fanus diadema Wab. ............... 
similis Szépl. . 824 | HemisrEPHANUS 
simillimus Elliott 823 peel oollie ‘Baan 
sulcatus Elliott .. 820 AOI RGHIOEE. ouch 
szépligetii indendy, nh enso8 SIEGE) TUCO, cock 
terebellus Finderl................... 818 pi Beis Wilesdlest ae Crue 
tertianus Morley ...........-....-- 806 CGE aca ame gaat 
t erythrocephalus Cam. ......... 
togoensis Enderl. Die EA SLUT | elabricoxis Roman 
trilineatus Hlliott.................. 813 | EN EE HILAL Mle bee 
trilobatus Elliott .................. 816 | pea tomae din Bee pl Bs aah 
Fa@NnatoPus. limpidipennis Schlett. ............ 
albomaculatus Cam................ 781 | MACKTULUSHScHlerta ee ee eee 
annulitarsis Enderl............... 787 | maculipennis West. 
aratifrons Hinder. .:....0...2--. ++ 799 | marginalis Schlett. ............4.. 
ROME IG ie paabon cob gscagauboees WSR) pehilkei Hndernt: oo... vei. cc-e. ++. 2s 
birdi Széplees reece 804 peruanus Huderl. ...........c.000> 


706 MR. BE. A. ELLIOTT ON’ THE 


HEMISTEPHANUS cae): 
ruficeps Cam. . 
submaculatus Weestw: Bee 
UIE ISQIPUGIAES bogesuuboosuebe cadcod sns 
WEWIOSINS SA BUIATpioodoasepacs sea 548036 
wiistneil Schlett. 


IcHNEUMON. 
COROMCIAS JORIS nos sod conoon od0590 050 
Sapamaiaayp LANs acs cna coobos obo eoncogdac 


M&GIscHus. 

acutus Lep. J Serv. .............+ 
americanus Sichel...............--- 
annulator Brull. 5 
antinori Giribodo ............-+-+>- 
lOOMMASISIS SWISS” scoscaotande saac00 
brasilianus Sichel...........--.-+:- 
PRMMTANS: CRASS saaccsocsoos seesdeac 
CEHORGIEMETIS JDIGUS. ‘soe caciene Sodda0 e650 
claripennis Cam. 

coronator Brad. 


cylindricus Westw. ..............- 
ducaliswifestausseseececckeeeecer 
erythrocephalus Cam. ... 

EULOPeUSHSCCREL se ee renee ssee ces 


floridanus Davis He ates 
InKOFER NH (OlGT> Seeacassoacdescs 958 o52 
furcatus Brull. 
indicus Westvw. 
insidiator Smith : 
iMSULANIS SMMAth 30... 2202. se. ee ee 
longicaudatus Costa ............--- 
long reollisiC agence eee ea ere 
MACULIErONsS Oasis concede 
maculipennis JFestzw. satis 
TAO. SHU Doadnodbooe a3 stoke anoSenace 
migricauda Sachel .cccscs: tee cmse ss 
pallidibansis Gat -. sees 
THUNELGA Ns (CIHTOs Gon oddgddscd dooce Sdo0s6 
ruficeps Sauss. 
ruficollis Cam. Bee es Te, Seg, 
SJNO MENIOVE S700, ‘coos 206 05da00 aoced 
submaculatus Weséw. ..........- 
TANESAUDIS) (SIDA Docs soaateeseces 63006060 
UENEENADIS SATAN : sob sored convescsdoae 
tarsatus var. Sichel ..........-...- 
texanus Cress. SER Hoe ao ATE 
GACT PIG SSMU Sa5h04 Be ccacsaceeboocke 
violaceipénnis Cam. 


MEGISELEIUS read NEES: 


NEOSTEPHANUS. 
albomaculatus Cam. ..........:.--- 
alluaudi Kieff. Seep aa cere ce 
camerunus Hnderl. .....c.s....-5: 
Crassipes Bischopfe ss. 5. eseeeaeee 
globiceps Hnderl. .........0.. sce 
MNEHEAIIIS) ISLC Aton) Bactbosedacusen600 
natalious Wnderl. ...............05: 
TYSON JOS “bo sqsaceton see casdec 


PARASTEPHANELLUS. 
Moxa VS LAMONT saporcasnaceebe rao se 
brevistigma Hinderl. ............... 
claripennis Cam. 
damellicus Westw. 


Page 


774 
773 
767 
775 
768 


718 
718 


742 
736 
736 


PARASTEPHANELLUS (cont.). 
eburneus Morley 


lee JDO RG ssc 000 020 cx d000Ne sac | 
IEevaQollbis WG RKGQ) sooccancetes o3ticce 
maculitrons" Cat eeeeeeeerertaas 


malayanus Cam. 


martini Stadlm.......... cae 
OH OMEVNS JBYPHES, 362 grossa nop cosccee 


palliditarsis Cam. 


TUG NES LOH Me con concedes aose00 507 
pysmeus Haderl. oes. esse see 


rubripictus Hlliott 


rufidornatus Morley............ .. 
TULO-OMNAbUSeOOUe eee eee 
GCMIADIS) OIG conacasnnacs boobas saelsee 


Pimps coronator Fab. ............ 


SCHLETTERERIUS. 


cmetipesiC7:css sess eae eee: 
TWOHIOES S07 cdo gon covton don aeasoa sas con 


STEPHANUS. 
acutus Lep. S Serv. . 
andinus Schletté.. re 
annulator Brull., “Westw.. 
anomalipes F's¢. . 


antinori Gribodo . vik ci eee 

bicoloxaifestcnseeeeee eee eee 
DORNEENSISUSa7Sse eee eee eee 
brasalvensis Sauss....esesieece- see 
brevicollis Enderl...... ............ 
brevipetiolatus Ender]. ....... 
lovepbnAe Ie! OFAASSS . 53 sds cnc cascan see a<1 
capitatus Schiett. ...i.0s.-.5:-0-- 
ceylonicus | Oamsnee ered 


cinctipes Cress. .... 


collarifer Schilethcte.A pas ane 
Comm aeMonleyenns tenes eee 


conradti Ender]. 


(COMOMENKOVE TOS saoqonasaccscondece 
coronatus PanzZ. .....0s.--..--cese 
crassicauda Morley ............- 


cylindricus Schlett 


damellicus End. nec Westw. 


damellicus Westw. 


diadema Flab., Westw. ... Hele 


diversus Schlett. 

ducalis Westw. ...... 
erythrocephalus Schlett. 
europzeus Sichel.. 


Sflavomaculatus Enderl. ....... 


froggattii, Cam.. 


frontalis Klug., Westw. soc 


furcatus Lep. & Serv. 


GRY SDAA soceha ste acdene ee 


globiceps Enderl. ......... 


hematipoda Montrz.............. 

hormianus)Enderlae eee pear 
OKO \MSIIWS codassesadeesuss a: 
imsignis Schlett.........065. -.n--e- 
intermedius Szépl................ 2. 
Janceolatus)A@iejnuene cee ere: 
lacteipennis Schlett................ 
leucodontus Schlett. ............... 
limpidipennis Schlett. ... ........ 
COUCTATIRS VAP oops dee Soadoaons S3aace 


HYMENOPTEROUS FAMILY STEPHANID, 707 


Page’ | Page 
STEPHANUS (cont.). | STEPHANUS (cont.). 

macrurus Schlett. ........0000.. 770 |  sehletterert Ender. ............... 788 
maculipennis Schlett. ............ 773 | SEONIOP JOA, Bon cccaaetasoosdcaccocss ks) 
malayanus Cam: n..--:.....-..... (60 Sickmanmil Schletterat tare LIE, 
marginalis Schlett. .............. 764 spoliaton) Schletiy sca eeeesseeecee O2O 
martini Stadim. ............4.... 748 submaculatus Schlett. ............ 773 
natalicus Westw. ......:.s.02 56+... 793 sulcitrons| Schlett. .-2css sce ---see-- 724 
TIGRE SOOO 54s sua0vs aed sonsnoeedone | Lats) TASES ISCAS | yan ssasoaceanogcsaes CAS) 
TUGOOAINS GOA soc casccododenaacsdbe Wee) Generischiletibaeyeaee cee eee Ol 
MIC MICAUGANSZCLEL ee ees acest aa) terebellus Enderl, .................. 818 
pachylomerus Schlett. ......:..... 723 | texanusi Oessumem asa eee en WoO 
mallescens schlett. -..-c-c4-..:.4) S03 ||» tibiator Sehlett. io... Jostens) DM 
(DUO JHWOG scacobsdoddancooeosnas 9 GES | VOGMCOSISISNEWUNID, Joo bedososas cbanoey | LAE 
pygmeus Hinderl. .................. 760 | HORUS J LOALAT 5.5 s00n00H00 pn09 96 boa | LBL 
THIVOODS NIG sognecgodasessenassue | COU turcomanorum Semenow.......... 731 
rubripes Morley .................. 734 | unicolon ScHlero ame eeeeeeee: 725 
FOREPOS SDAP oacncardo uke ndnicogongeicgen 0 GAs wadosus Schilett.... ie. ei eee 775 
rutofemoratus Szépl................ 748 MUONS GAYS | anocanadacasevonaaeos C8} 
mMUrO-ornabus Cay. Ay tcecaans) LDS | violaceipennis Cam. ............... 746 
TAS OSS JBVUIOUE ssa anecs0 cencddsavccs 25 | wustnert Schiett. ...............-0 768 
salomonis Westw. .................. 821 | xanthocephalus Cam. ............ 724, 
sausswret Schultz ................. 737 | Xorrpxs coronatus Lam. ......... 718 


STEPHANIDE. 


The position of this family remains undetermined, as it 
appears to form a transition between the Ichneumonide and 
Evaniide, with some leanings towards the Braconide. Different 
authors have taken very varied views as to its affinities. Cresson 
includes only Stephanus and the synonymous Megischus ; Cameron 
adds Monomachus; while Ashmead unites Stephanus and 
Megalyra. “ven Schletterer, the monographer of the family, 
declined to express any decided opinion. Some authors unite 
Stephanus and Stenophasmus as members of this family, though 
Ashmead places the latter next to Doryctes, and they are now 
usually considered to be Braconide. The two may be easily 
distinguished :— 


Antenne 30-40-jointed, shorter than body; anterior ocellus 
surrounded by five frontal tubercles; mandibles recurved, 
parrot-like; mesonotum not sulcate: scutellum tripartite ; 
posterior femora armed with two or three large teeth 
beneath. Fore wing with one cubital cell, hind wing with- 
out cell, or with one only ; abdomen convex beneath. 

STEPHANIDA. 


Antenne with about 70 joints, longer than body; frontal 
tubercles wanting, or not more than two; mandibles not 
recurved ; mesonotum with two longitudinal sulci; scutellum 
entire; posterior femora mutic; fore wing with two cubital 
cells ; hind wing with two or three complete cells ; abdomen 
more or less concave beneath. . . . . STENOPHASMIDA. 


Roman (Arkiv for zoologi, xi. 1917) states that the structure 
of the body, especially of the abdomen and legs, is sufficiently 
characteristic to entitle them to rank as a family, and considers 

Proc. Zoou. Soc.—1922, No. XLVITI. 48 


708 MR. E. A. ELLIOTT ON THE 


them to be nearest to the Evaniide, especially the Aulacine, 
which have similar habits. Evidence of this is adduced from the 
general form and the mode of insertion of the antenne, the 
tendency of the posterior margin of the head to become collar- 
like, the strong pleural sulcus for the reception of the middle 
legs, and possibly also the tendency of the neuration of the wings 
to become more simple. The special character of the five frontal 
tubercles occurs in the same manner in the Oryssini, and Roman 
would place the Stephanidee as the lowest member of the great 
group of Ichneumonide, between the Evaniide and Aculeata. 

For convenience, especially as an aid to identification, I accept 
Enderlein’s subfamilies, with the addition of Meostephanus 
Kieffer, while admitting the possibility of transition forms. 

Morley (Entom. 1917, p. 106) has drawn attention to the 
analogy of their structure with that of the Pimplid tribe Xorides, 
to which Lamarck actually ascribed Stephanus serrator under the 
name of Yorides coronator. 


GENERAL DESCRIPTION. 


The head is large and globose, with five frontal tubercles 
surrounding the anterior oceilus, from which the family takes 
its name (Greek: stephanos, a crown or wreath). The anterior 
tubercle is frequently larger than the rest, very rarely obsolete; 
the posterior pair are almost invariably smaller than the others, 
and have been quite overlooked by some authors, who describe 
species as trituberculate only. 

The powerful mandibles protrude rectangularly, are externally 
basally angular, anteriorly compressed, pointed and bent down- 
wards) like the beak on a parrot, with long grey to golden 
pubescence beneath. The maxillary palpi are long, slender, and 
five-jointed ; basal joint short, second twice as long as first, the 
others of about equal length. The labial palpi are short and 
four-jointed ; the three basal joints clavate, the apical cylindrical 
and longer. The eyes are large and roundish elliptical. Straight 
lines joining the ocelli would form an equilateral triangle with the 
base behind ; the posterior ocelli are close to the interior margin 
of the eyes. The 30-40-jointed antenne are placed very low 
down, the face being very short, cheeks normally as long as scape, 
and the frons proportionately 1 arge. Temples usually smooth 
and shining. he scape is incrassate, and the other joints, 
especially towards the middle, indistinetly discreted, gradually 
shorter and more slender towards the apex, distinctly pubescent 
except the basal ones; second flagellar jot normally twice as 
long as first; third as long as first and second together. 

The frons is more or less coarsely arcuately, transversely, or 
irregularly striate; the occiput has usually several transcarine 
close to the posterior tubercles, posteriorly often transversely 
rugose and laterally irregularly punctate rugose. Temples 


HYMENOPTEROUS FAMILY STEPHANID. 709 


usually polished smooth, with a few scattered punctures, rarely 
distinctly rugose. The cheeks are always distinctly developed 
and usually about the same length as the scape. The posterior 
margin of the head is either simple (serrator), bordered (furcatus), 
or produced into a more or less distinct collar (collarifer). 

The pronotum consists of an anterior narrowed neck, which is 
usually more or less distinctly transrugose, and a posterior 
erescentic part—herein called the semiannular—which is either 
entirely smooth or more or less finely rugose, with a narrow polished 
posterior margin. The mesonotum is curvate in front, rugose or 
punctate, rarely smooth; it has a central longitudinal row of 
punctures, sometimes almost obsolete, on each side of which is a 
smooth space, beyond which are two lateral impressions or rows 
of punctures. ‘the scutellum is divided into three sections by 
distinct crenulate sulci, is almost smooth, with a few large 
punctures on the margins of the lobes, of which the lateral ones 
are often more or less punctate. The mesoplure are deeply 
impressed above, somewhat smooth and shining, distinctly sculp- 
tured below. The metapleure are often separated from the 
median segment by a row of punctures, a sulcus or a carina, or 
rarely by a sulcus and a carina; less commonly confluent. Of 
the metanotum proper only a very narrow band, laterally broader 
triangular, is seen and is longitudinally striate. The median 
segment is large, usually with large, shallow punctures; some- 
times the interstices are alutaceous, and rarely, as in gigas, it 1s 
rugose, anteriorly often diffusely punctate, and more or less grey 
pubescent. The abdomen is inserted near the apex of the median 
segment, not far from the hind coxe. The petiole is rarely 
sessile, as in Schlettererius, usually nearly as long as the remaining 
segments together; it is veryrarely smooth (maculipennis), usually 
more or less finely trans-striate. The remainder of the abdomen 
is usually smooth, with a few dull spots, due to microscopic 
sculpture, more rarely entirely dull. There are seven segments 
in the male, six in the female, the posterior ones being short and 
indistinctly discreted. The terebra in female is usually as long 
as or longer than the whole body, the spicula ferruginous or red, 
its sheaths entirely black, or white or pale-banded before the 
apex, rarely rust-red (insignis). 

The anterior legs are short, their femora and tibize compara- 
tively slender, the tarsi five-jointed, the penultimate joint very 
short and furnished with pencils of hair. The hind legs are very 
elongate, their coxe stout, usually transrugose, often with finer 
striation between coarse transverse ridges. The second joint of 
the trochanters is indistinct. The hind femora are strongly 
fusiform, smooth and polished, rarely finely sculptured ; on the 
under side they bear two or three large teeth and a varying 
number of smaller serrations. The hind tibiz are compressed 
either in the basal third, or as far as, or even beyond, the middle 
and constricted. The hind tarsi are usually five-jointed in the 

48* 


710 MR. EB. A. ELLIOTT 


ON THE 


male and three-jointed in the female; only in Schlettereriws and 
Stephanus serrator they are five-jointed in the female and three- 


jointed only in S. tebzator male. 


Text-figure 1. 


Parastephanel/us. 


I 


Hem istep Nanus. 


= 
Foenatopus. 
— 
Diastephanus. 


1. Neuration complete. 


2. Discoidal cell 4-4 of cubital 
cell, not touching the submedian, 
and more or less petiolate. Ex- 
ternal submedian cell open behind. 
Radius not extending to margin 
of wing. 


3. Discoidal cell about as large 
as cubital, touching submedian 
cell. External submedian cell 
open behind. Radius extends to 
margin of wing. 


4, Cubital and discoidal cells 
wanting ; external submedian cell 
open behind. 


5. External submedian cell is 
indicated by a part of the median 
nervure only, 


6. External submedian cell en- 
tirely wanting. 


The neuration of the wings shows five, apparently constant 
forms, which have been utilized as the foundation of as many 


subgenera, 


HYMENOPTEROUS FAMILY STEPHANID, 711 


Schultz (Spolia Hymenop. 1906, p. 273) writes: ‘I cannot 
accept Menatopus Sm. or any of the ‘newly-erected genera of 
Stephanus Jur., as they are purely artificial, and useful at most 
for distinguishing groups of species in this rich genus.” Are not 
most of our genera artificial, and is not their object exactly that 
stated above? Cameron, again, in Ann. Soc. Ent. Belg. lvi. 1912, 
p- 358, makes the curious statement: ‘There are three longitudinal 
nervures, which is one of the points separating Stephanus from 
Parastephanus.” ‘There are always three such nervures, the 
difference being in the development. 

When the neuration is complete, as in Stephanus, s. str., there 
ave three basal cells: costal, which is very narrow and often in- 
distinct, median and inner submedian ; an outer series of three: 
cubital, discoidal, and external submedian; and a long, some- 
what narrow radial cell. The type of this subgenus is Stephanus 
serrator Fab. 

In the second form the neuration is similar to that of the 
second, but the discoidal. cell is only about one-fourth of the size 
of the eubitai, and is petiolate, not touching the inner submedian. 
They are Indo-Australian. Type: Parastephanellus pygmceus 
Enderl. 

In the third form the submedian or posterior nervure is 
abbreviated, not extending beyond the apex of the inner sub- 
median cell; the external submedian cell is thus open behind. 
The discoidal cell is about the same size as the cubital, and 
touches the inner submedian cell on a longer or shorter base. 
This subgenus, Hemistephanus Enderl., is exclusively Neotropical. 
The type is H. macrurus Schlett. 

In these three subgenera the stigma is opaque and the 
bounding nervures indistinct. In the three following subgenera 
the stigma is smaller, translucent, the bounding nervures distinct, 
the cubital and discoidal cells wanting. 

In the fourth form, Veostephanus Kieff., there are three com- 
plete basal cells; the median nervure extends to the full length of 
the outer submedian cell and the second transverse nervure is 
present, the outer submedian cell being thus bounded on three 
sides. Type: Weost. alluaudi Kieff. ! 

Form five, Menatopus Smith, differs in having only a short 
prolongation of the median nervure beyond the basal cells, with- 
out any indication of the second transverse nervure. ‘Type: 
Ff. indicus Westw. 

Form six, Diastephanus Ender]., has the neuration still further 
reduced, no nervure extending beyond the basal cells. Type: 
D. flavomaculatus Ender. 

In these three forms the radius does not extend to the margin 
of the forewing, the radial cell being apically incomplete. 

The colour in the majority of species is mainly black, frequently 
inclining to brownish; the head is often ferruginous or red, 
antenne and mandibles basally and the legs more or less red. 


mae MR. E. A. ELLIOTT ON THE 


A few species are fulvous, as F’. natalicus Westw. The size 
varies enormously within the same species, e.g. S. coronator, 
20-40 mm. The terebra in female varies greatly in different 
species, but appears to retain the same proportional length in 
each species. ‘The males are, as a rule, smaller and more slender 
than the females, although individuals may exceed the size of the 
smaller females. 

Very little appears to be known as to the life-history of these 
insects. Jurine writes of S. serrator as living in dry wood; 
Montrouzier observed S. hematipoda flying from trunk to trunk 
on the edge of a wood; the Cuban S. brunneus was taken in some 
numbers and in both sexes coming out of the same hole ina 
diseased tree, called Jalia. Roman (Arkiv for zoologi, 1917) 
considers it to be certain that they are forest insects and parasites 
on wood-boring larve. He found them always round fallen or 
dead trees, which were infested by the larve of Rhyncophora, 
Anthribide, Longicornia, and Buprestide. He hazards a conjec- 
ture that the special hosts belong to the Brenthidee, which, like, 
the Stephanide, are slender, elongate creatures, occurring in all 
warm countries. 

There can be little doubt that the family is more widely dis- 
tributed and the individuals more numerous than at present 
supposed. No collector had paid special attention to them till 
Roman did so in 1914-15 in Brazil, where he took 98 specimens, 
representing 7 species, all belonging to the subgenus Hem- 
stephanus. It is somewhat remarkable that H. vadosus accounts 
for 68 specimens, and that his two new species, H. angulicollis 
and glabricoxis, are represented each by one female only. The 
formation of the prothorax in these is very characteristic. 

One difficulty, inseparable from every attempt to compile a 
monograph without having access to authenticated specimens of 
every species, lies in the fact that every author has his own 
methods of description, and lays especial stress on different 
characters. Schletterer first gave scientific descriptions, including 
details of the posterior margin of the head, proportions of the 
basal flagellar joints, relative length of petiole to the remaining 
segments, and general sculpture. 

Enderlein desires to emphasize the value of the microscopic 
sculpture of the central tergites, while Roman finds in the 
sculpture of the pronotum valuable characters for the determina- 
tion of species. 

Smith and Cameron have given us many quite worthless 
descriptions, based largely on colour only, but worst of all are 
Westwood’s notes, one cannot call them descriptions, of S. diadema 
and frontalis in the Trans. Ent. Soc. Lond. 

What may be the effect of the Great War on the various 
collections, especially in Belgium, cannot yet be known, but it is 
certain that the German lust of destruction has caused heavy loss 
to science in all non-militant branches. 


HYMENOPTEROUS FAMILY STEPHANIDA, 713 


Table of Genera. 


(2.) 1. Hind wing with basal cell; abdomen sessile......... 1, Schlettererius. 
(1.) 2. Hind wing without basal cell; abdomen petiolate. 
(8.) 3. Cubital and discoidal cells present; stigma thick, chitinized ; 
the bounding nervures indistinct ......... ........ (STEPHANING.) 
(5.) 4. Neuration complete ..... ..... vecesseee 2. Stephanus. 
(4.) 5. Median vein incomplete, nate Cabeedan eal open behind. 
(7.) 6. Discoidal cell about 4+ of the size of the cubital cell, does not 
touch the submedian cell, but is petiolate ; stigma broad. 
5. Parastephanellus. 
(6.) 7. Discoidal cell nearly as large as the cubital, and touches the 
submedian on a longer or shorter base; stigma narrow, 
Lonowyandy pounbeds ses tss-d-.-5. a seee Hea esiaa ete: Hemistephanus. 
(3.) 8. Cubital and discoidal baile Latah ; Stiemis hyaline, bounding 
nervures distinct .............. eataeaeee eee es HIGH: ATOPODIN®.) 
(12.) 9. External submedian cell preeene panel 
(all) 210); 59 Bi », open behind only ......... 5. Neostephanus. 
(a@))) al 35 ms » open apically and behind. 6, Fanatopus. 
(9.) 12. i: x » entirely wanting ......... 7. Diastephanus. 


SCHLETTERERIUS Ashm. 


Schlettererius Ashmead, p. 150; Stephanus Say, p.61; Cresson 
(2), 1880, p. xviii. 


This genus appears to agree with Stephanus Jur. in the 
neuration of the fore wings, the formation of the mandibles and 
the insertion of the antennz, but differs in having one complete 
cell in hind wing, abdomen sessile, the basal segment being not, 
or but little, longer than the second. 

Only one species, S. cinctipes Cress., has hitherto been 
recognized, but the description of S. rufipes Say, though 
defective, suffices to place it here. I give the original 
descriptions. 


1. RUFIPES Say, l.c. 
Stephanus Jur. 


“8 rufipes. Black; abdomen sessile; thorax not remarkably 
attenuate before. Inhabits Pennsylvania. 

‘“ Body somewhat sericeous; palpi pale yellowish; scutellum 
with a groove on each side, rough ; metathorax rough, and with 
two slightly elevated longitudinal distant lines ; wings hyaline ; 
a large triangular fuscous carpal spot; feet rufous ; posterior pair 
of tarsi dusky ; abdomen a little rough at base; oviduct as long 
as abdomen. 

“ Length one-fifth of an inch. 

«Although the arrangement of the wing nervures agrees 
precisely with S. coronatus Jur., yet the form of the body differs 
materially, the thorax not exhibiting the remarkable attenuation 
before, and the abdominal petiole is not visible.” 


14 MR. E. A. ELLIOTT ON THE 


2. CINCTIPES Cress. 


Stephanus cinctipes Cress. (2), p. xviii; Schlett., p. 156. 
Schlettererius, Ashm., 1. c. 


“@. Black; labrum, narrow band at base of all the tibie, 
and apical third, except extreme tip of the ovipositor sheaths, 
white; tarsi testaceous, paler at base; trochanters, apex of first 
abdominal segment above, and most of the second and third seg- 
ments, ferruginous ; in front of ocelli a sharp semicircular carina, 
toothed in the middle and on each side; face transversely rugose ; 
cheeks nearly smooth; immediately back of ocelli a series of 
sharp transverse ridges (sometimes this part is tinged with dull 
ferruginous) ; “nigee OTE finely, transversely wrinkled, the im- 
pressed longitudinal lines composed of deep pits; pleure and 
metathorax roughly punctured, the former less so and shining ; 
middle of scutellum smooth and polished; tegule dull testaceous ; 
wings pale fuscous towards tips, an angular subhyaline band 
commencing at base of stigma, apex of wings paler than beneath 
stigma ; all tarsi 5-jointed, the penultimate joint with a long 
tufted process at the tip beneath; anterior tarsi double the 
length of their tibiz and very slender ; posterior coxe large and 
toothed above near the apex, their femora with two large teeth 
beneath, and a number of small unequal teeth between and on 
either side of them; their tibie not much thickened toward tip 
and not dilated; their tarsi about two-thirds the length of the 
tibie, with the first joint rather longer than the second, which 
is about equal in leneth with the third; abdomen smooth and 
polished, except the first segment, which is finely roughened 
and not longer than the posterior cox; ovipositor about 
double the length of the body. Length :55--75 inch.” 

Habitat: Washington Territory (Morrison). 

Schletterer (J. ¢.) gives the habitat as “ Subreg. 3. United 
States, Massachusetts, Pennsylvania, Washington, New York,” 
and adds that S. cinctipes is connected with the Hurepean 
S. serrator by its 5-jointed hind tarsi, and partly by the style of 
its sculpture, but differs in having two large teeth on the hind 
femora (serrator has three), in the sheaths of the terebra being 
white-banded before the apex, ete. 

Ashmead (Jd. c.) says: “ Posterior tarsi in both sexes normal, 
unarmed.” This appears to be a misprint for ‘‘femora,” and is 
incorrect since cinctipes has the hind femora armed with two 
large teeth beneath. 

S. coronatus Panz., Jur., etc. is a synonym of S. serrator Fab. 
The two species differ in size, colour, and especially in the rela- 
tive length of terebra; both occur in Pennsylvania. 

Say (J. ¢.) states that the neuration of his rufipes agrees pre- 
cisely with that of S. coronatus Jur., but does not especially 
mention the basal cell in hind wing. 


HYMENOPTEROUS FAMILY STEPHANIDA. TAD 


STEPHANUS. 


The designation Stephanus Jur., originally applied to the 
whole family, was restricted by Enderlein (2), p. 473, to the 
species possessing ‘‘ complete” neuration, comprising three basal 
cells, the costal, median and inner submedian, and three outer 
cells, cubital, discoidal, and external or outer submedian, together 
with a closed radial cell; and I follow him in calling this group, 
so characterized, the subgenus Stephanus. 

Unlike other subgenera, it is distributed throughout the 
whele range of the family, and to it belong the few known 
European species, 

There does not appear to be any character, other than the 
neuration, by which the species of this subgenus can be distin- 
guished from those of any other. As a rule they are large 
insects, mainly black, often with a red head, but size varies in 
the females from 10-40 mm, 

The terebra varies from slightly shorter than the body to 
twice its length. The males are mostly smaller than the 
females, though in some species the few specimens known of 
both sexes are nearly of the same size. Temples smooth and 
shining, unless otherwise stated. 

The type of the subgenus is S. serrator Fab., which is 
unfortunate in some respects, as it has five-jointed hind tarsi in 
both sexes and tridentate femora; the former character is 
unique and the latter is shared only by S. borneensis Sauss. 
S. tibiator Schlett., from Aden, is exceptional in having the hind 
tarsi three-jointed in the male. 

The characters utilized for differentiation of species in the 
following table are: Ist, number of joints in hind tarsi; 2nd, 
sculpture of temples; 3rd, colour of sheaths of terebra. Other 
important points are: relative proportion of petiole to rest of 
abdomen; sculpture of head and thorax; length of basal 
antennal joints: and colour of wings. 

The colour of the head (black, red, or, in one case, yellow) is a 
good distinction ; here, as among Coleoptera, it is often found 
the black shows a tendency to become rufescent, but can never 
be mistaken for a true red. 


Table of Species. 


OF 
(2.) 1. Hind tarsi 5-jointed ; hind femora tridentate; temples finely 
rugose-punctate. S-l7 mm. ....................... 1. serrator Fab. 
(1.) 2. Hind tarsi 3-jointed ; hind femora bidentate (except No. 25). 
(8.) 3. Temples coarsely rugose-punctate. 
(5.) 4. Antenne very short; temples with a smooth space behind 
eyes; head bordered ; mesonotum entire......... 2. antinorii Gribodo. 
(4.) 5. Antenne normal. 
(7.) 6. Temples with smooth raised space behind eyes; prothorax 


normal; hind femora slender....................... 8. gigas Schlett. 


716 


(GQ) 


(3.) 8. 
(22.) 9. 
(11.) 10. 
(10.) 11. 
(15.) 12. 
(14.) 13. 


—_ 
() iS 
See N) 
nN — 

bt po © w wv 
or E ew vw 


2 


(39.) 40. 


(88.) 41. 


(23.) 42. 
(60.) 43 


MR. E. A. ELLIO'L ON THE 


Temples without smooth space behind eyes; prothorax with 
an inflated ring between neck and semiannular; hind 
femora very incrassate........0....:0000 esc eseesseees 4. tibiator Schlett. 


Temples smooch. 

Sheaths of terebra entirely black. 

Frons coriaceo-granulate, legs entirely smooth ... 5. nigricauda Sichel. 
Frons arcuate or trans-rugose, legs distinctly sculptured. 

Frons arcuate rugose. 


Petiole shorter than rest of abdomen ; terebra about one-third [Schlett. 
longer than body; head red . tcessseees oss 6. Pachylomerus 


|. Petiole longer than rest of siidlomons anelbia half as long [Cam. 


again as ‘body ; ; head pale yellow, black banded. 7. xanthoc ephalus 


. Frons coarsely, irregularly, or transversely rugose. 
. A distinct sulcus across vertex and part of occiput; terebra 


nearly half as long ee as ae wings infumate, 
centrally darker ............. Wiens. 8. swleifrons Schiett. 


. No distinct sulcus on head or oc sine 
. Dark chestnut-brown; sheaths of terebra blue-black. 


9. brunneus Cam. 


19. Head black ; body chiefly black. 


20. Petiole and hind coxz coarsely transrugose; wings infuscate. 


10. unicolor Schlett. 


. Petiole finely trans-striate ; hind coxie smooth; wings sub- 


hy alimesinidescentaas aay eee eee 11. hornianus Ender. 


. Sheaths of terebra white-banded before apex. 
. Posterior margin of head simple, not bordered or reflexed. 


Terebra about as long as body. 


. Head black. 
. Second flagellar joint twice as long as first, third little shorter 


than first and second together. 


. Mesonotum cone Pena hind femora shining, 


STMNOOWN nos con daecorce sania . 12. andinus Schlett. 


. Mesonotum irreg salen rugose- etter Tita oanemn dull. 


13. sickmanni Schlett. 


. Second and third flagellar jou of equal length and little 


longer than first.. ee eee eee iL ewcrassicauda Moneys 


. Head not black. 
. Head, anterior femora and tibize mostly, and all tarsi red. 


15. tarsatus Sichel. 
Head yellow, body dull ferruginous .................. 16. texanus Cress. 


3. Terebra distinctly longer than body. 
. Petiole basally smooth, head granulate, mesonotum coriaceo- 


punictates eee eee eee nL cmeunopawsasichels 


. Petiole basally niet. Theat and mesonotum otherwise 


sculptured. 


. Mesonotum smooth, 2nd and 3rd flagellar joints of eaual 


» length and little longer than first ............... 18. éortus Morley. 


. Mesonotum more or less punctate. 
. Mesonotum rugose-punctate. 
. Median segment centrally densely punctate, laterally diffusely. 


Head and thorax black, abdomen brown, petiole rufescent. 
19. anomalipes Fst. 

Median segment evenly rugose-punctate throughout. Head 

and thorax ferruginous, abdomen red-brown, petiole 


nigrescent .............. wusssessesss. 20. Gicolor Westw. 
Mesonotum cites aurea, smedtion coon coarsely 
mebicwlatey nae sepleg . 21. nigricans (Sichel). 


Posterior margin of head honiles ea or ie aigilexcdl, 


3. Posterior margin of head bordered but not reflexed. 


HYMENOPTEROUS FAMILY STEPHANIDA, falvfi 


(49.) 44, Terebra about as long as or very little longer than body. 
(46.) 45. Petiole much shorter than rest of abdomen, mesonotum finely 
trans-striate and diffusely punctate............... 22. niger Smith. 
(45.) 46. Petiole as long as or very slightly shorter than rest of 
abdomen, mesonotum not trdns-striate. 
(48.) 47. Wings brown, centrally darker ........................ 23. froggatti Cam. 
(47.) 48. Fore wings hyaline, with steely iridescence......... 24. ceylonicus Cam. 
(44.) 49. Terebra distinctly longer than body. 
(51.) 50. Hind femora tridentate, forewings fuscous, with darker spot 
orifascia imimiddle ye. :)2....c tes sts seeeeeeneseseees) 2On OOMIEENSts) SAUSS. 
(51.) 51. Hind femora bidentate. 


(55.) 52. 
(54,) 53. 


(53.) 54. 


(57.) 58. 
(56.) 59. 
(43.) 60. 
(62.) 61. 
(61.) 62. 


(64.) 63. 


Wings evenly infumate. 
Frons arcuate rugose ; metapleurz coriaceous above, diffusely 
punctate beneath ; 8 “entirely black or black-brown. 
26. furcatus Lep.& Serv. 
Frons irreg gully rugose, ee densely punctate, 
head red . i Sopobeadbssods soa ello FOI MEGTES ISEIMUSS) 


. Wings centrally or Giese daneer 
. Wings centrally darker, petiole shorter than rest of abdomen. 
57. Head not impressed behind vertex, semiannular smooth, 


mesonotum posteriorly confluently punctate ... 28. coronator Fab. 


Head compressed behind vertex, semiannular transrugose, 
mesonotum posteriorly coarsely transrugose ... 29. ducalis Westw. 


Wings infumate, external median and submedian cells and 

then to margin of wing dark brown, cea as long as rest 

of abdomen ......... SROBB HEE _ 30. lanceolatus Kieff. 
Posterior margin of feed literal, celine like. 
Wings strongly ee ae darker ; eae shorter | Montrz. 

than rest of ‘abdomen .......... aie: . 31. hematipoada 


Wings slightly infuscate or She Aline, petiole! as nee as rest 
of abdomen. 
Head black, cheeks pale marked, median segment cribrate 
punctate ern e- . 82. acutus Lep. & Serv. 


(63.) 64. Head red, median segment ‘confluently ‘punctate. . 38. villosus Kieff. 
é. 
(2.) 1. Hind tarsi 8-jointed, hind femora bidentate ...... 4, tibiator Schlett. 
(1.) 2. Hind tarsi 5-jointed. 
(6.) 3. Hind femora tridentate. 
(5.) 4. Mesonotum irregularly rugose, metapleuree rugose; wings 
hyaline, centrally and apically infuscate......... 1. serrator Fab. 
(4.) 5. Mesonotum cribrate punctate, metapleure densely punctate ; 
fore wings with transverse dark band!............ 25. borneensis Sauss. 
(3.) 6. Hind femora bidentate. 
(10.) 7. Posterior margin of head produced, collar-like. 
(9.) 8. Posterior margin of head translucent but not reflexed. Meta- 
pleuree and median segment partly confluently punctate. 
33. villosus Kietf. 
(8.) 9. Posterior margin of head reflexed. Lateral lobes of scutelluim 
coarsely punctate; metapleuree and median segment not 
confluently punctate ............. Beta 36: pilosus Elliott. 
(7.) 10. Posterior margin of head bordered, Te not collar like. 
(12.) 11. Wings subhyaline green and red iridescent ; outer orbits are 
whiteemarked ...00cccccecceceeessesssesees seve. 11. hornianus Ender. 
(11.) 12. Wings not iridescent. 
20.) 18. Wings infumate, centrally darker. 
( g > J 


(17.) 14. 


Head red. 


718 MR. E. A. ELLIOT! ON THE 


(16.) 15. Head impressed behind vertex; semiannular transrugose; 
mesonotum centr ally suacoulh, "metapleure coarsely rugose. 
29. ducalis Westw. 
(15.) 16. Head not impressed behind vertex; semiannular smooth ; 
mesonotum densely, partly confluently punctate; meta- 
pleure densely punctate................0.......02..:. 28. coronator Hab. 
(14.) 17. Head black. 
(19.) 18. Second and third flagellar joints of equal length and little 
longer than first; median segment densely punctate; 
petiole shorter than rest of eden trans-striate 
(EIVROWUIEIOVONDES, 3s ceo secconec ee ...seseee-e 23. rubripes Morley. 
(18.) 19. Second flagellar joint twee as shlong as Ree ‘hin distinctly : 
longer than second; median segment diffusely punctate ; [Szépl. 
petiole smooth, only basally rugose............... 34. rafofemoratus 
(13.) 20. Wings not centrally darker. 
(28.) 21. Wings more or less infumate. 
(25.) 22. Thorax confluently punctate. 


(24.) 23. Petiole trans-striate, rest of abdomen smovth and shining. 
Dark chestnut-brown . COO ete Lcsssssssee. 9. brunmeus Cress. 


(23.) 24. Petiole very finely tr ciculate. ee of abdomen finely 
alutaceous. Black; head and anterior legs red. 35. comma Morley. 


) 25. Thorax otherwise sculptured. 
.) 26. Vertex transrugose; median segment scrobiculate reticulate ; 
hind cox trans-striate. Black; head red...... 15. tarsatus Sichel. 
(26.) 27. Vertex arcuate striate; median segment densely, partly con- 
fluently punctate; hind coxz finely striate between coarse 


scale-like rugosities. Black to dark brown, cheeks white- [Serv. 

ANAT Mcrae as enNstareee duonbbedennoosadboogsoare en amano: IED es 
(21.) 28. Wings clear hyaline; Le anal oc Cee ey 

carinate .. DAU en aie Mn Ua aati 7 SAREE One . 37. rugosus Elliott. 


1. SERRAYTOR Fab. 


Ichnewmon No. 193, Zschah, i. p.60, 2; tab. Synistata, fig. 1938, 
% ,1789. Ichnewmon serrator Fab. (1), p. 224, 2 ,1798. Gigoieanvnns 
coronatus Panz., pl. et fig. 18,1801, 9; Jur., tab. : 2; Lep.& Serv., 
p- 489, pl. 376, fig. 2a, Gy De ‘Nees ab IDS we Deroy, oat, OD 3 
Blanch., p. 23. Bracon serr ator Fab, (2), p. 108, Or Tchnewmon 
coronatus Latr., p.179, 9. NXorides coronatus Lamarck, p. 135); 
i.c. iv. Hd. 2, 1835, p. 347. Stephanus serrator Brull., p. 537, 2; 
Westw. (4), p. 227, 2; Sichel (2), p. 472, 9 ¢; Schlett., p. 91, 
ONSe Andre np ole Onse 


2 ¢. Head with frons moderately coarsely and regularly, 
occiput more finely and irregularly rugose, arcuately near the 
posterior ocelli. Temples finely rugose-punctate, with a small 
smooth raised space behind eyes. Posterior margin of head 
simple. Second flagellar joint twice as long as first, third longer 
than second, but shorter than first and second together. Neck 
coarsely transrugose, semiannular more finely. Mesonotum 
coarsely and irregularly rugose, more finely to nearly smooth in 
front, with a distinct central longitudinal channel and two lateral 
divergent rows of small punctures. Mesopleurz coriaceous 
rugose, with coarser reticulations behind; scutellum centrally 
smooth, laterally longitudinally rugose; metapleure rugose, but 
less coarsely than the median segment, from which they are 


HYMENOPTEROUS FAMILY STEPHANIDA, 719 


separated by a distinct sulcus. Petiole stout, irregularly rugose, 
basally more coarsely, much shorter than the remaining feebly 
shining part of abdomen. Terebra longer than body, its sheaths 
entirely black. Hind legs with the cox finely coriaceous ; 
femora tridentate, tibiz constricted beyond middle and longer 
than femora; hind tarsi five-jointed in both sexes. 

Black ; antenne basally brown; mandibles basally ferruginous, 
cheeks white-marked; abdomen rufescent, apically black, or 
entirely black; legs brown, rufescent beneath, their joints 
whitish. Wings basally hyaline, centrally and apically slightly 
infuscate. 

Length 8-17 mm. 

Habitat : Germany, France, and Central Europe. 

The tridentate hind femora and five-jointed hind tarsi will 
suffice to distinguish this species 9, and the only other ¢ with 
the former character (borneensis Sauss.) is nearly twice as large. 


2. ANTINORIL Grib. 


Megischus antinori Gribodo, p. 346, 2. Stephanus ant. Schlett., 
fos HOSS Vivo vale eolomnlla(@)) sos IKoy Ge 


2. Frons moderately shining, densely and irregularly, not 
arcuately, reticulate rugose. Five frontal tubercles (not six as 
stated by Gribodo in his original description), of nearly equal 
size, a triangular space in front of the anterior ocellus shining 
sianrornl Vertex densely, coarsely, and deeply rugose- punctate, 
with five curved carinz in front and a longitudinal impression 
behind. Occiput coarsely but diffusely and not deeply rugose- 
punctate, without longitudinal impression. Posterior margin of 
head sharply bordered. Cheeks rather shorter than scape and 
as long as second flagellar joint, shining smooth and diffusely 
punctate. Temples with a smooth shining protuberance near 
eyes (as in gigas), above it coarsely and deeply punctate, below 
densely and coarsely rugose-punctate. Antenne unusually short, 
32-jointed, reaching only to the base of median seement; second 
flagellar joint two and a half times as long as first, third only as 
long as second. Neck weakly transrugose, separated by a right- 
angled impression from the semiannular, which is densely and 
coarsely rugose-punctate, with a broad smooth posterior margin. 
Prosternum shining, anterior third finely rugose-punctate, remain- 
der smooth with very sparse and deep punctures. Mesonotum 
entire, centrally almost smooth, otherwise very coarsely, but not 
deeply or densely rugose-punctate. Scutellum shining, with a 
few punctures on the lateral lobes, and the borders of central 
lobe finely striated. Mesopleure densely and coarsely reticulate, 
with a smooth shining space below tegule and above middle 
coxe. Metapleurze prominent, coarsely reticulate rugose, with a 
broad, deep, and curvately carinate impression in front and a 
similar smoother impression behind. Median segment discally 
extremely coarsely reticulate, laterally rugose, separated from 


720 MR. E. A. ELLIOTT ON THE 


the metapleure by a fine longitudinal carina, behind which is a 
narrow sulcus. Petiole basally transrugose, then finely trans- 
striate, apically smooth, shorter than rest of abdomen. Terebra 
nearly as long as body, ferruginous, sheaths entirely black. 
Hind legs with coxze stout, with setiferous punctures in front, 
more sparse towards middle, apically finely trans-striate ; femora 
inflated, microscopically finely and densely alutaceous and 
punctate. 

Head, thorax laterally, legs including tibize and petiole beneath 
with long, sparse, coarse hairs; similar but shorter and finer 
hairs on abdomen. 

Wings hyaline, apically infumate, stigma opaque, nervures 
pitch-brown. 

Black; mandibles and antenne red-brown ; cheeks and base 
of anterior femora red-yellow. Wings faintly iridescent. 

Length 25 mm. Terebra 23 mm. (Gribodo: 26 mm.; terebra 
20 mm.) 

Habitat: Mahal Uonz, Shoa, East Africa, 1 2 taken by 
Marquis Orazio Antinori. Described from the unique type in 
the Museum at Genoa by W. A. Schulz. 

The extremely short antenne and six frontal tubercles, as 
described by Gribodo, inclined Schletterer to doubt this species 
being a true Stephanus, but the latter character proves to be a 
mistake, the five frontal tubercles being quite normal; and 
Schulz states that the right antenna is complete, the apical 
joint is normally pointed and 13 times as long as the penultimate, 
not showing any signs of being abnorinally formed or dwarfed. 

This species resembles S. gigas and S. tibiator Schlett. in the 
very coarse sculpture of the body and especially of the temples, 
but differs from both in the unusually short antenne. From 
S. gigas 1t may be further distinguished by the berdered posterior 
margin of the head and by the broadly smooth posterior margin 
of prothorax ; from S. tibiator it differs in the absence of the 
longitudinal suleus on occiput, in the formation and sculpture of 
the "prothorax, etc. No other specimen having been yet taken, 
it must remain uncertain whether the short antenne are truly 
characteristic of the species. 


3. GIGAs Schlett. 
Stephanus gigas Schlett., p. 96. 


Frons very coarsely and irregularly rugose; occiput arcuately 
rugose near the posterior ocelli, behind this centrally trans- 
versely, laterally irregularly rugose. Temples covered with 
dense, rather deep and partly confluent punctures, excepting a 
smooth space behind ocelli. Posterior margin of head simple. 
Second flagellar joint fully twice as long as first, third only as 
long as second. Neck coarsely transrugose, semiannular coarsely 
rugose-punctate throughout, excepting an indistinctly rugose 
impression near tegule. Mesonotum centrally somewhat smooth, 


HYMENOPTEROUS FAMILY STEPHANID®, 721 


with a central longitudinal row of broad and deep punctures, and 
two deep, divergent sulci, laterally very coarsely rugose-punctate. 
Central section of scutellum smooth, with a few marginal 
punctures ; lateral lobes densely, coarsely, and partly confluently 
punctate. Metapleuree and median segment very coarsely and 
reticulately rugose, separated by a deep rugose sulcus. Hind 
legs with the coxe short and stout, transrugose, more finely 
Denna ; femora comparatively eral and slender, polished 
smooth, with two large teeth; tibiz much longer than femora 
and compressed only in basal third. 

Black ; mandibles basally and hind coxee ferruginous, antennee 
with basal half brownish. 

Habitat : Schiras, Persia. Type in Imp. Nat. Hist. Museum, 
Vienna. 

This description was taken from a specimen without abdomen, 
and as no mention is made as to the number of hind tarsal 
joints, the sex cannot be determined. It appears to resemble 
closely S. tibiator Schlett. from Aden, in the coarse sculpture of 
head and temples, though differing in the structure of pronotum 
and in the slender hind femora. 


4. TIBIATOR Schlett. 


Stephanus tibrator Schlett., p. 2938, 29 g; Morley (1), pp. 34 & 
[1 Oe 


@ 3. Frons very coarsely and irregularly to reticulately 
rugose ; vertex obliquely arcuate rugose; occiput very irregularly 
punctato-rugose, with a central longitudinal impression. Temples 
coarsely and deeply punctate, posteriorly more diffusely and 
cleanly, near the cheeks more densely and confluently. Cheeks 
feebly punctato-rugose. Second flagellar joint twice as long as 
first, third slightly more slender then second and but little 
longer. Neck very short, laterally coarsely rugose, above with 
two large foveze separated by a longitudinal carina; then follows 
a coarsely rugose-punctate inflated part, separated by a distinct 
constriction from the semiannular, which is apically coarsely 
rugose, centrally and laterally densely and partly confluently 
punctate, with a polished smooth posterior margin. Mesonotum 
centrally diffusely punctate, apically and laterally rugose ; 
scutellum smooth with diffuse punctures, its three lobes defined 
by crenulated sulci. Mesopleure rather coarsely and irregularly 
rugose in the upper impressed part, irregularly to reticulate 
rugose beneath, with a smooth, shining, and diffusely punctate 
space near the tegule; metapleure very prominent, coarsely 
and irregularly rugose, separated by a deep sulcus from the 
coarsely reticulate rugose median segment. Petiole trans-striate, 
apicaliy smooth and shining. ‘erebra shorter than body, 
sheaths entirely black. Hind legs with coxee basally densely 
aciculate punctate, apically transrugose; femora strikingly 


nie MR. E, A. ELLIOTT ON THE 


incrassate, bidentate, smooth and shining, with diffuse setiferous 
punctures: tibiz as long as femora, compressed to about middle ; 
tarsi three-jointed in both sexes. 

Black, with a tendency to brownish, especially in the cheeks 
and tarsi. Antenne ferruginous in 9, black in ¢. Wings 
hyaline. 

In ¢ the rugosity of the semiannular is more pronounced 
than the puncturation; the lateral lobes of the scutellum are 
more rugose than punctate; the abdomen is longer and more 
slender in 9. 

Length 9, 21-24 mm.; abdomen 16-18 mm.; petiole 10-113 
mm.; terebra 20-22 mm.; dg, 21 mm. 

Habitat: Aden; Ruaha River, late German Hast Africa. 

This species appears to differ from S. gigas Schlett. in the 
formation of the prothorax and im the strikingly incrassate hind 
femora 

In some respects it resembles S. pachylomerus Schlett., from 
which it may be distinguished by the coarsely punctate temples, 
formation of the prothorax, and by the shorter petiole and 
terebra. 

Tt resembles S. antinori in size, length of terebra, and incras- 
sate hind femora—this last is not uncommon among the African 
species. It differs in the longitudinal occipital sulcus, structure 
and sculpture of prothorax, formation of mesothorax and 
sculpture of mesopleure 

S. tibiator S is distinguished from all others yet known by the 
hind tarsi being three-jointed, and is the only known exception 
to the rule, that the males of this family have those tarsi five- 
jointed. 

S. serrator 2 alone has the hind tarsi five-jointed, otherwise 


they are invariably three-jointed. 


5. NIGRICAUDA Sichel. 

Megischus nigricauda Sichel (2), p. 479, 2. Stephanus n. 
Schlett., p. 106, 9. 

@. Frons coriaceo-granulate, vertex arcuate rugose; occiput 
with basal third trans-striate, remainder smooth polished. Pro- 
notum smooth and shining, neck subquadrate; mesonotum 
almost impunctate, smooth and shining; with an anterior 
longitudinal impression, separated from the smooth polished 
sorcelliora by a row of punctures; median segment serobiculate 
reticulate. Petiole finely trans-striate, remainder of abdomen 
smooth polished. Terebra as long as body, sheaths black. Legs, 
especially the hind ones, enrol and shining ; hind femora 
bidentate. 

Black; head red, vertex black, cheeks pale marked ; mandibles 
basally and two or three basal antennal joints red or rufescent ; 
anterior tarsi rufous, middle tarsi white-banded, base of hind 


HYMENOPTEROUS FAMILY STEPHANIDA, (23 


metatarsus and whole of following joints white, claws black. 
Wings flavescent, costa beyond stigma black; nervures fuscous. 

Length of body and terebra 15-163 mm. Wings 8-84 mm. 

Habitat: Manila (Luzon). Prof. Semper. 29? 9°. 

Sichel remarks on the variability of the serrations between the 
large teeth. In one specimen one of these, near the apex of 
femora, amounts almost to a third tooth, while on the other 
specimen this is present on one femur only. 

The coriaceo-granulate frons, the very smooth pro- and meso- 
notum, and the apparent absence of all sculpture on the legs 
suffice to distinguish this species. 


6. PACHYLOMERUS Schlett. 
Stephanus pachylomerus Schlett., p. 98, 9 ; Morley (1), p. 111. 


@. Frons coarsely arcuate rugose; vertex very convex and 
arcuate to transverse rugose; occiput transrugose. Posterior 
margin of head simple. Temples centrally prominent, very 
shining; cheeks shorter than scape. Second flagellar joint more 
than twice as long as first, third only as long as second. Neck 
very short, with a deep oblique impression, and superficially 
punctate-rugose ; semiannular polished, with very diffuse fine 
punctures. Mesonotum with medium-sized and partly confluent 
punctures. Scutellum smooth, margins of lobes obliquely striate. 
Mesopleure with the upper impressed part oblique rugose in 
front, smooth behind, the lower convex part diffusely punctate. 
Metapleurz coarsely and irregularly rugose, separated from the 
cribrate-punctate median segment by a deep, slightly rugose 
sulcus. Petiole finely transrugose, distinctly shorter than the 
rest of abdomen ; second segment shining smooth, basally feebly 
rugose, the remainder rather dull. Terebra longer than body, 
its sheaths entirely black. Hind legs with coxe strongly shining, 
diffusely punctate; femora very incrassate, smooth, diffusely 
punctate, bidentate ; tibie slightly longer than femora, compressed 
nearly to middle. 

Black; head, base of mandibles, scape, third segment entirely 
and legs partly red; apical abdominal segments more or less 
rufescent. Wings subhyaline, discal and external submedian 
cells infuscate. és 

Length 28 mm.; terebra 35 mm. 

Habitat: West Africa (Gaboon); type in Nat. Hist. Museum, 
Hamburg. Gold Coast; in coll. British Museum. 

This species resembles S. coronator and ducalis in shape and 
colour, but may be easily distinguished by the entirely black 
sheaths of the terebra, prominent temples, weaker puncturation 
of mesothorax and more incrassate hind femora. From S. sud- 
cifrons, which it also resembles, it may be known by the absence 
of the sulcus on vertex and by the simple posterior margin of 
head. It differs from S. vanthocephalus Cam. in the short pro- 
notum, sculpture of prothorax and hind coxe, and in coloration. 


Proc. Zoou. Soc.—1922, No. XLIX. AY 


(24 MR. E. A. BLLIOTT ON TSE 


7. XANTHOCEPHALUS Cam. 
Stephanus xanthocephalus Cameron (10), p. 358, 2. 


2. Frons strongly arcuate striate; vertex irregularly arcuate 
rugose, occiput finely and closely trans-striate; posterior margin 
of head simple. Pronotum densely and rather strongly trans- 
striate; mesonotum shining, irregularly striate and punctate, 
basal half centrally smooth. Scutellum smooth, apically sparsely 
punctate. Pleurze smooth, white pubescent. Median segment 
with large, diffuse punctures, apically irregularly trans-striate. 
Petiole finely and closely striate, apically more strongly, longer 
than rest of abdomen. ‘Terebra half as long again as body, 
sheaths entirely black. Hind legs with coxe densely striate, 
femora with two large teeth, one medium and two very small. 

Black; head pale yellow, with broad black band from occiput 
to posterior tubercles; scape testaceous; antennz black; third 
segment partly, ventral surface and apical segment entirely 
rufous; apices of femora, anterior tibize and tarsi, apex of hind 
tibie, and hind metatarsus rufo-testaceous. Wings hyaline, 
posterior discoidal celi infumate, stigma and nervures black. 

Length 28 mm.; terebra 43 mm. 

Habitat: Dima, West Africa. A. Koller. Type in Congo 
Museum, Tervouren. 

The pale yellow, black-banded head is very distinctive. 


8. SULCIFRONS Schlett. 
Stephanus suleifrons Schlett., p. 110, 2. % Megischus insidiator 
Smith (3), p.7 d. 


@. Frons coarsely and obliquely to irregularly rugose, vertex 
transrugose, the rugosity gradually passing into puncturation on 
occiput, and all sculpture disappearing at back of head; a very 
distinct longitudinal sulcus across vertex and part of occiput; 
posterior margin of head strongly bordered; cheeks rather shorter 
than scape. Second flagellar joint twice as long as first, third 
shorter than first and second together. Neck coarsely and 
obliquely rugose; semiannular rather closely and partly con- 
fluently punctate, its posterior margin polished smooth. Meso- 
notum diffusely and coarsely punctate. Scutellum smooth, with 
a few conspicuous marginal punctures. Mesopleure feebly rugose 
and shining in the upper impressed part, punctate below; meta- 
pleuree very coarsely and densely punctate, separated from the 
coarsely cribrate-punctate median segment. by a deep and smooth 
suleus. Petiole transrugose, apically smoother, shorter than rest 
of abdomen, which is basally shining, otherwise dull. Terebra 
much longer than body, sheaths entirely black. Hind legs with 
coxe coarsely transrugose; femora entirely smooth, bidentate, 
the tibiee longer than femora, compressed to beyond middle. 

Black; head, two basal antennal joints, anterior legs and hind 
tarsi red or ferruginous. Wings slightly infuscate, centrally 
darker. 

Length 23 mm.; terebra 30 mm. 


HYMENOPLTEROUS FAMILY STEPHANIDA. 725 


Habitat : Mindanao, Philippines. Type in Museum at Ham- 
burg. 
3 (2). ML. insidiator, Smith. ‘ Wale. Length 9 lines. The 
head and base of the antenne ferruginous; the former trans- 
versely striated, with the posterior margin of the vertex smooth 
and shining, or with a few punctures. Thorax strongly, but not 
closely punctured ; wings fusco-hyaline, the anterior and inter- 
mediate tibizw rufo-testaceous, with the femora obscurely so; the 
posterior femora with two stout teeth beneath and six minute 
ones between them; the posterior tibiz at their apex, and the 
tarsi pale rufo-testaceous. The abdomen elongate, lanceolate at 
the apex, and entirely smooth and shining. Hab. Mysol.” 

This species may be best known from the very similar S. coro- 
nator K. by the sulcus on vertex and entirely black terebral 
sheaths; the latter character and the smaller size will distinguish 
it from 8. ducalis Westw. Smith’s description of his insidiator 
is defective, but it may be the male of swleifrons, which it appears 
to resemble. 


9. BRUNNEUS Cress. 

Megischus brunneus Cresson (1), p. 84, 2 3. 

2. Head coarsely rugose, vertex deeply trans-striate. Thorax 
coarsely and confluently punctate; apex of median segment trans- 
rugose. Petiole trans-striate, remaining segments shining 
smooth. Terebra longer than body, spicula brown, sheaths blue- 
black. Hind femora bidentate, the anterior tooth largest; hind 
tibize compressed in basal third ; hind tibie 3-jointed, metatarsus 
much dilated. All tarsi yellow pubescent beneath. 

Dark chestnut-brown, antenne blackish, scape brown, legs 
brown, knees yellowish, posterior femora and tibie exter nally 
nigrescent. Wings subhyaline, tinged with fuscous; stigma and 
nervures black, former with basal pale spot. 

Length 1-11 lines (14-23 mm.); _ terebra 9-134 lines 
(18-27 mm.) ; fore wings 93-143 lines (19-29 mm.) 

6. Similar to 9, but more slender; the two teeth on hind 
femora equal in size, hind tibie less dilated, all tarsi five- -jointed, 
hind metatarsus not dilated. 

Length 9 lines (183 mm.); alar expanse 93 lines (19 mm.) 

Habitat: Cuba. 

Collection Ent. Soc. Philad. Two 2 9,two ¢ ¢ specimens. 
Both ¢ and Q of this species were taken by Dr. Gundlach in 
considerable numbers, coming out of the same bole in the trunk 
of a diseased tree, called Jalia in Spanish.” 


10. un1coLor Schlett. 

Stephanus unicolor Schlett., p. 104, 2. 

@. Frons rather coarsely and irregularly to ar cuately rugose ; 
occiput transrugose, more coarsely near the ocelli; posterior 


margin of head with raised border; cheeks as long as scape. 
49% 


126 MR. E. A. ELLIOTT ON THE 


Second flagellar joint one and a half times as long as first, third 
fully twice as long as first, but shorter than first and second 
together. Neck very short, laterally coarsely rugose, bifoveate 
above; semiannular smocth and shining, with a few anterior and 
lateral superficial punctures. Mesonotum diffusely punctate, with 
a distinct row of central punctures. Scutellum polished smooth, 
with a few lateral punctures. Mesopleurze smooth above, feebly 
rugose and diffusely punctate beneath; metapleure coarsely 
reticulate to irregulary rugose, separated by a smooth shining 
sulcus from the median segment, which is reticulate to cribrate 
punctate. Petiole moderately coarsely transrugose, little shorter 
than the rest of abdomen, the central segments more or less dull, 
the basal and apical segments shining smooth. 'Terebra as long 
as body, sheaths entirely black. Hind legs with coxe coarsely 
transrugose; femora stout and polished, bidentate; tibie 
compressed to beyond middle. 

Black; the head shows a tendency to become red on the frons, 
pale yellow on cheeks and temples; scape ferruginous, anterior 
legs brown to rufescent. Wings slightly infuscate. 

Length 14 mm. 

Habitat: Mindanao, Philippines. Type in Nat. Hist. Museum, 
Hamburg. 

This species is somewhat similar to 8. sulcifrons Schlett., but 
only half as large and wanting the distinctive vertical sulcus. 
It also resembles F’. indicus West., D. lewcodontus Schlett., and 
D. pallescens Schlett., from all of which it may be known by 
the neuration. 


11. HoRNIANUS Enderl. 
Stephanus hornianus Enderl. (5), p. 12, 2 3. 


2 3S. Frons coarsely transrugose; vertex with four trans- 
carine ; occiput irregularly rugose. Second flagellar joint shorter 
than third. Neck trans-striate ; semiannular smooth polished, 
with a row of punctures before posterior margin. Propleure 
trans-striate, mesopleurze smooth, lower half punctate, apex sub- 
rugose; metapleurz widely reticulate, separated from the median 
segment by a very deep, smooth, and shining suleus. Median 
segment widely reticulate, basally diffusely punctate, apically 
densely transrugose. Petiole finely trans-striate, as long as, or 
slightly shorter than remainder of abdomen. Terebra in 9 
rather longer than body. Hund legs with coxe shining smooth, 
diffusely punctate, transrugose in front and behind; femora 
diffusely punetate, bidentate. 

Black; a yellow spot on outer orbits; petiole and legs except 
coxe and hind femora ferrugineus; last tarsal joint black. 
Wings subhyaline, with red and green transverse iridescence on 
outer margin; stigma and nervures black. 


Length ? Habitat: Ceylon. 


HYMENOPTEROUS FAMILY STEPHANID#., 127 


Enderlein appears not to have given the size of this species, 
but as he compares it with S. ceylonicus Cam. and S. hematipoda 
Montrz., it is probably about 28-30 mm. The former of the 
above species is said to resemble the present one in colour, with 
the same spot on the outer orbits, but the sculpture of the meso- 
and meta-pleure differs, the petiole is black, and the wings have 
a steely iridescence. 


12. AnpINus Schlett. 
Stephanus andinus Schlett., p. 138, @. 


Q. Frons somewhat coarsely and irregularly rugose; occiput 
transrugose, coarsely in front, more regularly towards the eyes. 
Temples finely but distinctly obliquely rugose, with a smooth 
space behind eyes. Posterior margin of head simple. Second 
flagellar joint twice as long as first, third a little shorter than 
first and second together. Neck rather coarsely transrugose ; 
semiannular finely but distinctly obliqueiy rugose. Mesonotum 
coarsely and confluently punctate, the usual median row of 
punctures wanting and the two lateral impressions represented 
by indistinct rows of punctures. Scutellum coarsely and diffusely 
punctate. Mesopleurz above finely alutaceous and dull, beneath 
slightly shining and diffusely punctate; metapleure dull, finely 
and diffusely punctate, separated from median segment by a deep 
and dull suleus. Median segment diffusely but deeply punctate, 
apically transrugose and with distinct lateral sulci. Petiole 
trans-striolate, basally more irregularly rugose, shorter than rest 
of abdomen, which is discally dull, laterally shining. Terebra 
as long as body, sheaths white-banded before apex. Hind legs 
with coxe trans-striolate; femora shining smooth, bidentate ; 
tibize constricted not quite to middle. 

Black ; cheeks pale marked; the anterior legs, especially the 
tarsi, tend to become brownish. Wings hyaline. 

Length 19-20 mm. 

Habitat: Colombia. Type in Royal Nat. Hist. Museum, 
Berlin. 

This species differs from S. acutus Lep. & Serv. in the simple 
posterior margin of head, coarser puncturation of mesonotum, and 
shorter petiole. From S. niger Smith it may be known by the 
finer sculpture of the frons, obliquely rugose temples, and rugose 
semiannular. 


13. SICKMANNI Schlett. 

Stephanus sickmanni Schlett, p. 152, 2. 

@. Frons coarsely and irregularly rugose; occiput anteriorly 
arcuately, posteriorly transversely, and laterally irregularly 
rugose. Temples with a few apical punctures. Posterior margin 
of head simple. Cheeks slightly longer than scape. Basal flagellar 


728 MR. E. A. ELLIOTT ON THE 


joints normal. Neck coarsely transrugose, semiannular distinctly 
transrugose and finely punctate, with smooth posterior margin. 
Mesonotum coarsely and irregularly rugose-punctate. Scutellum 
centrally smooth, laterally with diffuse to dense and partly con- 
fluent punctures, lateral lobes smooth in front, densely and 
confluently punctate behind. Mesopleure above distinctly 
alutaceous, shining smooth, with diffuse punctures below; meta- 
pleure diffusely punctate, more densely in front, separated from 
median segment by a very shallow and alutaceous sulcus. Median 
segment rather coarsely and diffusely punctate, longitudinally 
rugose round insertion of petiole, marginal furrows indistinct. 
Petiole centrally smooth, basally and apically somewhat rugose, 
shorter than remainder of abdomen, which is apically dull. Terebra 
about as long as body, sheaths white-banded before apex. Hind 
legs with coxee coarsely transrugose above, laterally more irregu- 
larly ; femora subglabrous, bidentate; tibie compressed not quite 
to middle. 

Black; cheeks pale marked; antenne, mandibles basally and 
the legs, especially about the joints, tend to become rufous. 
Wings almost hyaline. 

Length 24-25 mm. 

Habitat: Georgia, U.S.A. Type in Royal Nat. Hist. Museum, 
Berlin. Named after Prof. Franz Sickmann, of Iburg, Hanover. 

This species resembles small individuals of S. furcatus Lep. & 
Serv., from which it differs in the shorter petiole and terebra, 
and sculpture of prothorax and mesonotum. For differences 
between this species and S. bicolor Westw., see the latter species. 
It bears also a great resemblance to H. marginalis Schlett., but 
is easily distinguished by the neuration and by the reflexed 
posterior margin of head in latter. 


14. crassicAuDA Morley. 
Stephanus crassicauda Morley (1), p. 106, 9. 


2. Head coarsely and irregularly rugose, a central longitu- 
dinal trans-striate suleus on occiput ; posterior margin of head 
simple ; temples finely punctate, prominent and glabrous above. 
Second and third flagellar joints of equal length and a little 
longer than first. Neck short, with three strong trans-carine ; 
semiannular finely trans-striate, its posterior margin narrowly 
smooth; mesonotum coarsely punctate; scutellum smooth, with 
diffuse and rather fine puncturation. Meso- and meta-pleure 
alutaceous and dull, with a few punctures, latter separated from the 
median segment by a suleus. Median segment scabriculous and 
centrally trans-striate. Petiole trans-striate, as long as rest of 
abdomen ; apical segments discally emarginate. Terebra slightly 
longer than body, sheaths unusually stout and abruptly pointed, 
ferruginous, with a white band before the equally broad black 
apex. Hind legs with coxe elongately pilose and sparsely trans- 


pie a 


HYMENOPTEROUS FAMILY STEPHANID®. 729 


carinate; femora smooth and pilose, with two very long and 
slender teeth ; tibiae constricted in basal third and again beyond 
middle. 

Black ; a white spot on cheeks ; base of flagellum ferruginous ; 
anterior legs, hind trechanters, constricted part of tibie, their 
apex, and the hind tarsi fulvous. Wings slightly infumate, 
external submedian cell darker, with a pale space beyond it. 

Length 26 mm.; abdomen 16 mm.; petiole 8 mm.; terebra 
27 mm. 

The type, in the British Museum, bears a label “ Australia, 
1868 ” only. 

The cotype, in same collection, is much smaller: body 205 mm.; 
abdomen 13 mm.; petiole 6 mm.; terebra 19 mm. It was taken 
by F. P. Dodd, 5. 111. 08, at Townsville, Queensland. 


15. TARSATUS Sichel. 


Megischus tarsatus Sichel (2), p. 476, pl. 10, figs. 4 et 5, 2. 
Stephanus tarsatus Schlett., p. 107, 2 3. 


@. Face below antenne obliquely striate, the strie descending 
from the sides to the central line; frons transrugose; vertex 
coarsely transrugose; occiput with a small central part trans- 
rugose, laterally irregularly reticulate. Neck elongate, trans- 
rugose; mesonotum apically smooth, centrally diffusely punctate 
with a few transrugosities; scutellum smooth, with deep and 


- coarse but diffuse punctures; median segment scrobiculate 


reticulate. Abdomen slightly pubescent; petiole trans-striate. 
Terebra as long as body, white-banded before apex. Hind cox 
trans-striate; hind femora bidentate. 

Black; head except apex of mandibles, scape, sometimes also 
basal flagellar joint, apex of pronotum, anterior femora, tibice 
mostly, and the tarsi red; antenne fusco-piceous. Wings in- 
fumate, nervures fuscous or rufo-fuscous. 

Length 38 mm. Expanse of fore wings 18 mm. 

S. Similar to @ ; head more strongly transrugose, occiput 
centrally more broadly trans-striate, metathorax more rugose, 
petiole and coxe more finely sculptured, serrations on hind 
femora finer; abdomen fusco-piceous. 

Length 24 mm. Expanse of fore wings 11 mm. 

Habitat: Manila, Philippines; Bachian, Molucca Islands. 

Schletterer (J. c.) tentatively synonymizes this with MW. tarsalis 
Smith, from Bachian, remarking that it corresponds in the 
infumation of the wings, length and colour of terebra, colour of 
body, sculpture of petiole, and, to some extent also, in sculpture 
of head and thorax. It appears, however, to differ in having the 
head coarsely sculptured, black, not red as in ¢arsatus, the thorax 
coarsely punctate, and is only 9 lines (18 mm.) long. I consider 
it a distinct species, but the. description is insufficient for certain 
determination. 


730 MR. BE, A. ELLIOTT ON THE 


16. TEXANUS Cress. 
Megischus texanus Oress. (2), p. 190, 2. 


2. Frons coarsely reticulate; vertex with two trans-carine ; 
occiput finely transrugose. Pronotum smooth; mesonotum 
coarsely rugose; metanotum (median segment) centrally and 
apically rugose. Petiole as long as rest of abdomen. Terebra as 
long as body, basal half of sheaths ferruginous, apical half black, 
white-banded before apex. Legs shining; hind femora bi- 
dentate; hind tibize constricted above middle. Wings ‘too 
much damaged to be described.” 

Opaque, dull ferruginous; head yellowish, mandibles apically 
black; antennze apically darker; hind tibize with constricted 
part yellow; frontal tubercles apically and abdominal segments 
from second apically fuscous. 

Length 10 mm. 

Habitat: Comal Co., Texas. 1 @. 

In the absence of details as to the wings, I place this species 
in the subgenus Stephanus. It appears to be characterized by 
the sculpture of the head and by the colour, especially of the 
terebral sheaths. 


17. EBUROP@US Sichel. 


Bothriocerus europeus Sichel (1), 1860, p. 759, 9. Megischus 


eur. Sichel (2), p. 484, 9. Stephanus eur. André, p. 486; 
Schlett., p. 95, ©. 


@. Head granulate, frons near tubercles irregularly rugulose, 
occiput above regularly transrugose. Pronotum irregularly 
trans-striate, its extreme posterior margin smooth; mesonotum 
coriaceo-granulate, obscurely trans-striate, with central longitu- 
dinal row of punctures; median segment coriaceo-granulate and 
apically obscurely trans-striate. Petiole trans-striate, basally 
smooth, remainder of abdomen smooth and shining. ‘Terebra 
longer than body, its sheaths apically white-banded. Legs 
smooth. 

Black; mandibles and antenne basally rufo-testaceous ; 
tegule, petiole, base of second segment, trochanters, anterior 
tibiee, hind tibize basally, and all tarsi red or rufescent. Wings 
subhyaline, stigma black, nervures fuscous. 

Length 13 mm.; terebra 17 mm. 

The unique specimen was taken in Sicily in Sept. 1859. 

In the absence of details as to the relative lengths of the 
antennal joints and of the hind legs, it is impossible to place 
this species with certainty. It appears to be very near anoma- 
lipes Fst., from which it differs chiefly in the basally smooth 
petiole. The granulate sculpture is probably the same as that 
otherwise described as coarsely punctate. A similar case occurs 
where the same sculpture is described by one author as “ coarsely 
striate” and by another as “ carinate,” 


HYMENOPTEROUS FAMILY STEPHANIDA., 731 


18. tortus Morley. 
Stephanus tortus Morley (1), p. 84, °. 


@. Frons reticulately, but not transversely rugose ; the 
three anterior tubercles prominent, acute; occiput laterally 
coarsely reticulate, centrally transaciculate; posterior margin of 
head simple. Second flagellar joint rather longer than first, 
third only as long as second. Neck transrugose, semiannular 
glabrous. Mesonotum smooth, central row of punctures distinct, 
lateral rows subobsolete. Mesopleursze smooth above, rugulose 
beneath; metapleursze rugulose, separated from the median 
segment by a strong, glabrous sulcus. Median segment deeply 
and rather diffusely punctate, apically transaciculate. Petiole 
shorter than rest of abdomen, finely trans-striate throughout, all 
remaining segments smooth shining. Terebra about one-third 
longer than body, sheaths white-banded before apex. Hind legs 
with coxe discally glabrous and finely granulate, below and 
inside obsoletely trans-striate; femora bidentate, shining smooth, 
sparsely punctate ; tibize compressed in basal third, centrally 
excised and apically inflated. 

Black; mandibles and a spot near their base red; petiole 
dark red, rest of abdomen piceous; anterior legs, except basally, 
red ; hind tibiz piceous, centrally red. Wings slightly infumate. 

Length 26 mm.; abdomen 19 mm.; petiole 9 mm.; terebra 
34 mm. 

Habitat: Nedunkernie, N.P. India. Taken by O.S8. Wickwar. 

Type in coll. Claude Morley, Monk Soham, Suffolk. 

The proportions of the flagellar joints, granulate hind coxe, 
and structure of hind tibize distinguish this species from all 
others with white-banded terebral sheaths. 


19. ANOMALIPES F'st. 


Stephanus anomalipes Forst., p. 228, 9 ; Schlett., p.93; André, 
p. 487, 2. 


Q. Frons rather coarsely and irregularly rugose; occiput 
diffusely and arcuately rugose behind the tubercles, posteriorly 
transversely, laterally irregularly rugose. Temples with a few 
shallow and confluent punctures in front; posterior margin of 
head simple. Second flagellar joint twice as long as first; third 
longer than second, but shorter than first and second together. 
Neck rather coarsely and transversely, semiannular more finely 
and less regularly rugose. Mesonotum rather coarsely, 
posteriorly more densely transrugose punctate, with a distinct 
central longitudinal line of punctures and two lateral impressions. 
Scutellum centrally shining smooth, laterally finely and densely 
punctate. Mesopleurs rather coarsely coriaceous throughout ; 
metapleure coarsely and irregularly rugose, separated by a 
distinct and slightly alutaceous sulcus from the median segment, 
which is coarsely and densely rugose-punctate, Petiole rather 
slender, finely coriaceous, basally indistinctly transrugose, 


732 MR. E. A. ELLIOTT ON THE 


shorter than the remaining segments together, also shorter than 
the hind coxe, trochanters and femora together ; rest of abdomen 
basally rugose, thence dull to apex. Terebra longer than body, 
its sheaths white-banded before apex. Hind legs with coxe 
plainly trans-striate above, laterally coriaceous and shining ; 
femora internally shining, externally and abeve almost shining, 
slightly rugose behind, bidentate; tibiz longer than femora, 
compressed in basal third. 

Black; cheeks pale marked, mandibles and antenne basally 
ferruginous, temples tending to brownish; abdomen brewn, 
petiole rufescent; legs more or less rufescent. Wings slightly 
infumate throughout. 

Length 17 mm.; terebra 21 mm. 

Habitat: Hungary. Forster's type is in the State Museum 
at Budapest. 

Distinguished from serrator Fab. by the three-jointed hind tarsi, 
white-banded terebral sheaths, ete. From S. sickmanni Schlett. 
it may be known by the rugose, not punctate metapleure 
coriaceous petiole, and longer terebra. 


20. BIcoLOR Westw. 


Stephanus bicolor Westw. (2), p. 5385; (8), p. 276; Schlett., 
p. 154, @. | 


2. Frons arcuately to irregularly rugose; occiput anteriorly 
arcuately, posteriorly rather coarsely transversely, laterally more 
irregularly rugose. Temples very diffusely punctate. Posterior 
margin of head simple. Cheeks a little longer than scape. Basal 
flagellar joints normal. Neck rather “coarsely transrugose ; 
semiannular finely transrugose and sparsely punctate. Meso- 
notum rather coarsely punctato-rugose. Scutellum smooth, 
with a few punctures on outer margin only. Mesopleurze above 
alutaceo-rugose, below somewhat densely punctate; metapleure 
irregularly rugose, separated from the median segment by a 
nearly smooth sulcus. Median segment rather densely punctate 
centrally, laterally more diffusely, oblique to transverse rugose 
round insertion of petiole, crenulate marginal sulci distinct. 
Petiole basally irregularly, otherwise transversely striate, shorter 
than remainder of abdomen, which is entirely dull. Terebra 
evidently longer than body, sheaths white-banded before apex. 
Hind legs with the coxe laterally coarsely and irregularly, above 
indistinctly, apically decidedly transrugose; femora slightly 
shining, bidentate ; tibize compressed to middle. 

Badious; head and thorax ferruginous, remainder of body red- 
brown ; petiole more or less nigrescent. Wings slightly infumate. 

Length 15-22 mm. Habitat: Kentucky, Georgia, U.S.A. 

This species may be known from S. sickmanni Schlett., which 
it much resembles, by the longer terebra, sculpture of meso- and 
meta-pleuree and of petiole, which is rugose throughout, not 
centrally smooth. The colour is also distinctive, being a decided 
red-brown or chestnut, S. sickmanni being deep black. 


HYMENOPTEROUS FAMILY STEPHANID#®. 733 


21. nraricAns (Sichel), Schlett. 


Megischus tarsatus var. nigricans Sichel (2), v, p. 476, 9. 
Stephanus nigricans Schlett., p. 109, 9. 


@. Face below antenne irregularly rugose reticulate, frons 
transrugose; occiput centrally broadly transrugulose, laterally 
coarsely and deeply reticulate. Neck elongate, transrugose ; 
mesonotum diffusely punctate; scutellum centrally smooth, 
laterally diffusely punctate; median segment coarsely reticulate. 
Petiole trans-striate; terebra longer than body, broadly white- 
banded before apex. Hind coxe trans-striate; hind femora 
bidentate. 

Black; tegule apically rufescent; anterior legs red, front 
femora externally black. Wings infumate, nervures fuscous, 

Length 33 mm.; fore wings 12 mm.; terebra 35 mm. 

Habitat: Ceylon. One @ in coll. Sichel. 

Sichel considers this a variety of S. tarsatws Sichel, which it 
strongly resembles, but the different sculpture of face and occiput, 
black head, and longer terebra entitle it to specific rank. 


22. NIGER Smith. 
Megischus niger Smith (4), i. p. 44; Cam. (1), p. 420, pl. xviii. 
fig. 19, 2. Stephanus niger Schlett., p. 135, °. 


Q. Frons coarsely and irregularly rugose; occiput anteriorly 
arcuate-rugose, posteriorly transversely, laterally irregularly 
rugose; cheeks scarcely as long as scape. Posterior margin of 
head strongly bordered. Second flagellar joint two-and-a-half 
times as long as first, third as long as first and second together. 
Neck with a distinct transverse impression, behind which it is 
finely and sparsely rugose; semiannular polished, with very fine 
and diffuse punctures. Mesonotum finely trans-striate and 
diffusely punctate, the usual central row of punctures and the 
two lateral impressions obsolete. Scutellum smooth, with a few 
marginal punctures. Mesopleurze smooth, with diffuse coarser 
and finer punctures; metapleure finely and irregularly rugose, 
separated by a deep and smooth sulcus from the median segment, 
which is diffusely and irregularly punctate, with distinct 
marginal furrows. Petiole very distinctly trans-striate, much 
shorter than remainder of abdomen, but nearly as long as the 
hind cox, trochanters, and femora together. Terebra scarcely 
longer than body, its sheaths white-banded before apex. Hind 
cox trans-striate between stronger rugosities ; hind femora dull, 
bidentate ; tibize compressed not quite to middle. 

Black; cheeks pale marked and the anterior legs with a 
tendency to brown. Wings hyaline. 

Length 23-25 mm. 

Habitat: Panama, Guatemala. Type in British Museum. 

This species is distinguished from the similar S. marginalis 
Schlett. by the bordered but not reflexed posterior margin of 


(a4 MR. E. A. ELLIOTT ON THE 


head, shorter petiole, and by the sculpture of prothorax and 
median segment. 

S. sickmanni Schlett. has the posterior margin of head simple, 
neck of prothorax short, semiannular rugose-punctate, and meso- 
pleurz punctate. 


23. FROGGATTIE Cam. 


Megischus froggattii Cameron (9), p. 357, 9 5d. Stephanus 
froggattti Cam. (11), p. 98; Elliott (2), p. 28, 29 d. Stephanus 
rubripes Morley (1), p. 107, 9d. 


@ 6. Frons strongly transrugose; vertex arcuately and 
occiput transversely rugose ; cheeks shorter than scape ; posterior 
margin of head bordered. Second and third flagellar joints of 
equal length and rather longer than first. Neck short, cen- 
trally longitudinally impressed, with one strong basal carima ; 
semiannular widely aciculate, extreme base smooth. Mesonotum 
coarsely punctate, with central smooth space; mesosternum flat 
and polished. Scutellum smooth, basally bounded by a crenulate 
furrow. Propieure diffusely punctate; mesopleurze smooth 
above, with three or four irregular rows of punctures below ; 
metapleure closely rugulose throughou t, separated by a smooth 
sulcus and a carina from the median segment, which is rather 
densely but superficially punctate. Petiole transaciculate, a 
little shorter than the remaining segments together; base of 
second segment feebly rugose, “igs emaonlh and shining. Terebra 
as long as, or very “slightly longer than body, sheaths black, 
broadly white-banded before apex. Hind legs with coxe slender 
and strongly trans-striate; femora smooth and shining, bi- 
dentate ; tibize longer than femora, constricted in basal third ; 
tarsi five-jointed in ¢, three-jointed in 9 ; all tibize andthe hind 
femora elongately pilose. 

Black ; mouth-parts and genal orbits testaceous ; anterior legs 
except coxe and trochanters bright red; hind femora and tarsi, 
except apically, and apical two-thirds of hind tibie red. Wings 
brown, centrally darker, serge and nervures Dleds 

Length, 2, 33 mm.; abdomen 22 mm. » peels 10 mm. ; 
terebra 35 imMmON 2. hy UG mm: ; abdomen 11 mm. ; petiole 45 mm. 
Another ¢, 12, 8, 32 mm. Taken by W. W. Fr oggatt, Solomon 
Islands, July—Aug. 1909. 

@ 3, length 22 mm.; abdomen 13 mm.; petiole 6 mm.; 
terebra in 9, 21} mm. Taken by C. M. Woodford, Solomon 
Islands, 1887. 

There are also in the British Museum 5 9 9,2 ¢ 6, their 
dimensions lying between those given above. 

Cameron states that the abdominal petiole is a little longer 
than the rest of abdomen, but actual measurement shows it to be 
shorter. He also gives the colour of the wings as fusco- 
violaceous ; it is now brown ; possibly faded. 


HYMENOPTEROUS FAMILY STEPHANIDA. ax) 


This species much resembles S. rufo-femoratus Szépl., also 
from the same locality, but differs in the relative length of the 
basal joints of the antennz, sculpture and length of petiole, and 
the strongly trans-striate hind coxe. 

Cameron considers it to be near his Megischus violaceipennis 
from New Britain, in which the sculpture of the body tends to 
be reticulate. 

Mr. ©. IT. Brues found in the Museum of Harvard College 
422,246 6, from Auki, Tulagi, Fulakora, and Malaili. The 
larger females, 26-31 mim., have the femoral teeth more strongly 
developed than in the smaller individuals of 21-23 mm. The 
males measure from 20-21 mm. 


24, CEYLONICUS Cam. 
Stephanus ceylonicus Cam. (4), p. 102; Morley (1), p. 34, @. 


2. Frons irregularly transrugose, arcuately above; vertex 
aud occiput arcuate rugose; posterior margin of head finely 
bordered. Second flagellar joint twice as long as first, third 
little longer than second. Neck closely trans-striate, with 
central apical row of punctures, semiannular transcarinate, 
laterally diffusely punctate, a narrow posterior margin smooth. 
Mesonotum diffusely punctate; scutellum only laterally punctate. 
Propleure closely obliquely striate; mesopleure almost im- 
punctate; metapleure smooth above, coarsely and almost 
reticulately transrugose beneath. Median segment diffusely 
punctate, apically transreticulate. Petiole closely trans-striate, 
as long as rest of abdomen, which is smooth. Terebra slightly 
longer than body, sheaths white-banded before apex. Hind 
cox transrugose, hind femora bidentate. 

Black; a pale spot below eyes; anterior legs and base of hind 
tibiee rufo-testaceous. Wings hyaline with steely iridescence ; 
stigma and nervures black. 

Length 28 mm. 

Habitat: Trincomali, Ceylon (Yerbury). Kandy and Gal- 
gamua, Ceylon (Morley.) 


25. BORNEENSIS Sauss. 
Megischus borneensis Sauss., p. 202, 2 3. 


“Od. M. ruficeps simillimus, ejusdem pictura; capite 
tamen toto rufo; tantum mandibule apice nigris. Antenne 
nigre, primo articulo rufo. Caput sicut in specie, laudata 
insculptum sed superne totum et occipite transverse arcuato- 
strigatum. Facies rugoso-punctata, nonnunquam tamen utrin- 
que strigis nonnullis, valde oblique conferens, vel toto transverse 
obsoleta arcuata -strigata. Mesonotum politum, cribrosum, 
antica utrinque punctis confluentibus crasse strigatus; scutello 
polito. Coxe postice transverse elevato-rugate ac striate. 
Femora postica, denti usque 3 armata (1 ante medium, 2 prope 


736 MR. E. A. ELLIOTT ON THE 


apicem). Ale fusco nebulose, in medio vitta transversa vel 
maculis fusca. 

‘“‘ Variet. a. Pedibus fusco-rufescentibus. 

6. In individuis minoribus et in maribus soul peu 
incomplete explicata. 

“ O, Long. 20-31; thoracis 4°8-7°5; petioli 6-10; oviposit. 
32°5-40 mm. 

“3. Long. 25; thoracis 4°6; petioli 6°5; ale 12 mm.” 

Habitat: Borneo. 

Very similar to furcatus Lep. & Serv. and to ruficeps Sauss., 
but distinguished from both by the tridentate hind femora and 
the central dark mark or band in fore wing; from the latter also 
by the sculpture of the frons, and the longer terebra. 


26. FuRcATUS Lep. & Serv. 

Stephanus furcatus Lep. & Serv., p. 1835, 2 ; Schlett., p. 130, 
2d. Stephanus brasiliensis Westw. (1), p. 413, pl. 66, fig. 3; 
Ae (3), p. 227, d. Megischus annulator Brull., p. 539, pl. 40, 
fig. 5; Sichel (2), p. 482, 9; Cam. (1), p. 419, pl. 18, fig. 158 
Senter annulator Westw. (8), p. 228, @. Boiiietocerns 
americanus Sichel(1), p.761,2. Megischus americanus Sichel (2), 
p. 480, 2. Megischus brasilianus Sichel (2), p. 483, 3. 

Q 3. Frons very coarsely irregularly to arcuately rugose ; 
the three anterior tubercles very large; occiput coarsely rugose, 
arcuately in front, transversely behind, laterally irregularly. 
Temples apically feebly rugose; cheeks as long as _ scape. 
Posterior margin of head strongly bordered. Second flagellar 
joint fully twice as long as first, third longer than second but 
shorter than first and second together. Neck coarsely trans- 
rugulose, semiannular smooth, slightly transrugose in front and 
with a few lateral punctures. Mesonotum with a central row of 
very coarse punctures, a smooth, diffusely punctured space on 
each side bounded by divergent impressions, beyond which it 
is laterally transrugose, with a more or less extensive smooth 
space. Scutellum polished and diffusely punctate. Mesopleurze 
finely coriaceous above, obliquely rugose in front, smooth and 
diffusely punctate beneath; metapleure with a few coarse 
rugosities in which are some large punctures, separated from the 
median segment by a shallow but distinct and almost smooth 
sulcus. Median segment punctate, more densely and partly 
confluently centrally and laterally, apically transrugose. Petiole 
trans-striate, shorter than rest of abdomen, also aineraiee than the 
hind cox, trochanters and femora together; second segment 
shining smooth, basally rugose, remaining segments rather dull. 
Terebra longer than body, white-banded before apex. Hind legs 
with coxe finely transrugose between coarse, scale-like rugosities; 
femora shining smooth, bidentate ; tibize compressed not quite to 
middle; tarsi five-jointed in ¢, three-jointed in 9. 


HYMENOPYEROUS FAMILY STEPHANIDA. tol 


Black to dark brown, cheeks sometimes pale marked. Wings 
infumate. 

Length 25-39 mm.; terebrain 9, 38-52 mm. 

Habitat: Brazil; Guatemala. 

The sculpture of the median segment is variable. In some 
specimens in the British Museum the punctures are more diffuse, 
only apically and laterally confluent, forming transverse rugosities; 
in others the punctures are more confluent, the greater part of 
the segment being coarsely transrugose. The punctures appear 
to be more distinet in the ¢ ¢ than in the Q 9. 

S. annulator Brull., 9 S$; the types are in the British 
Museum, and certainly belong to the present species. 9: length 
25 mm.; terebra 27 mm.; abdomen, anterior legs, and hind 
tibize rufescent. <¢: length 23 mm.; the front legs and hind 
tibiee red. 

S. brasiliensis Westw., 9. Type in British Museum. This 
is a very small specimen, only 17 mm. long, terebra 20 mm., 
with abdomen and all tarsi rufescent. I cannot find any 
specific difference from furcatus Lep. & Serv. 

This species is said to be the least rare of the family. It can 
be distinguished from SS. sickmanni Schlett. by the raised 
posterior margin of the head, sculpture of thorax, and longer 
terebra. It also superficially resembles Hemistephanus mar- 
ginalis Schlett., but may be at once distinguished by the 
neuration of the wings. 


27. RUFICEPS Sauss. 
Megischus ruficeps Sauss., p. 201, 2. Stephanus saussuret 


Schulz (2), p. 322, 2. 
‘‘ Ba Chieu, prov. Saigon, 1886; Cambodia, 1886. 


“©. Validus, cinereo-hirtus, J/. furcati Brull. statura et ill 
similime. Caput rufus, ore nigro, palpis fuscis. Antenne 
nigre, basi superne rufe, primo articulo rufo. Caput anterius 
rugato-punctatum, haud arcuato strigatum, supra crassissime 
areolato-punctatum, in vertice rugis transversis elevatis 1-4 
instructum ; occipitis vitta longitudinali tramsversa striata. 
Frontis depressis subrhomboidalis, rugosa, per turbiculos rugi- 
formes 5 nigros cireumseripta, marginibus infero-lateralibus 
vittam transversa strigatum notatis. Pronoti collum crasse trans- 
verse plicato strigatum. Mesonotum crasse cribro-punctatum, 
haud strigatum. ‘Tegule rufide. Mesopleuree dense punctate. 
Scutellum et postscutellum rugulosa, basi et apice polita. 
Metanotum crassissime foveolato punctatum. Abdominis petiolus 
ut solitum confertim transverse striolatum. Abdomine de 
reliquo levigatum, sericans, basi petiolari rugulosa. 'Terebra 
nigra, annulo ante apicem longiusculo albido. Pedes nigri, politi, 
partibus rufo-testaceis. Coxe postice plus minus crasse 
subsquamoso rugose, ac punctate, superne apice ieee 
strigate. Femora postica subtus crenata, dentibus-que 2 armata. 
Ungues rufee. Alze nebulose, venis nigris. 


738 ' MR. E, A. ELLIOTT ON THE 


“Long. 19-27; thorax 45-6; petioli 6°5-9°5; ale 10:5-16°5; 
oviposit. 26°5-34 mm.” 

M. de Saussure tentatively suggests as males for this species 
two very small insects, taken in Siam. They differ in having 
the head entirely red, the occiput almost striate, as in 
S. borneensis Sauss., the sculpture of the body less pronounced, 
and the pronotum scarcely striate in one specimen. Length 
10 mm. 

This species appears to closely resemble S. furcatus Lep. & 
Serv., but the face is rugose-punctate, not arcuate rugose; the 
mesonotum, median segment, and mesopleurz densely punctate, 
and the head red. It also resembles S. coronator Fab., but 
differs in the red head and the sculpture of head and mesopleure. 

Schulz (/. ¢.) alters the name of this species to sausswrei, on 
the ground of ruficeps being preoccupied by Cameron’s Central 
American insect. As the latter falls into the subgenus 
Hemistephanus, I reinstate the original name. 


28. CORONAWOR Fab. 


Pimpia coronator Fab. (2), p. 118, 2. Stephanus coronator 
Boisd., p. 656, pl. 12, fig. 7, 2 ; Schlett., p. 114, @ g ; Hnderl. (4), 
p- 306, 9 ¢. Megischus cor. Brull., p. 538, 9 ; Sichel (2), 
p: 477, 23. Megischus insularis Smith (1), p. 120, 2d. 
Megischus vidwus Smith (2), p. 1388, 5. AMegischus longicaudatus 
Costa, Ach. (2), p. 271, 2 6; (1) p. 81, 2 Gd, pl. iv. figs. la ¢, 
Lo QQ. 

®. Face coarsely arcuate rugose ; occiput immediately behind 
tubercles arcuately, behind this centrally transversely, laterally 
obliquely rugose. Posterior margin of head sharply bordered. 
Cheeks distinctly longer than scape. Second flagellar joint twice 
as long as first, third a little longer than second. Neck very 
coarsely transrugose; semiannular polished, with a few punctures 
in front. Mesonotum coarsely, densely, and posteriorly partly 
confluently punctate, anteriorly more diffusely, with a distinct 
central longitudinal row of punctures and indistinct lateral 
impressions. Scutellum centrally smooth, lateral lobes diffusely 
punctate. Mesopleure almost smooth and shining above, rugose- 
punctate beneath; metapleurse coarsely and densely punctate, 
separated by a slightly rugose suleus from the coarsely eribrate 
punctate median segment. Petiole basally coarsely, apically more 
finely transrugose, little shorter than remainder of abdomen, 
which is basally rugose, otherwise feebly shining. Terebra rather 
longer than body, sheaths white-banded before apex. Hind legs 
with cox coarsely transrugose behind; femora polished smooth 
and posteriorly extremely finely punctate, bidentate; tibize much 
longer than femora, compressed in basal third; tarsi three-jointed. 

g. Similar to @ but more slender; petiole is to rest of 
abdomen as 5: 6; hind tarsi 5-jointed. 


HYMENOPTEROUS FAMILY STEPHANID &. - 739 
3 


Black; head, mandibles except apices and the scape ferru- 
ginous; anterior legs chestnut-brown. Wings infumate, with 
darker spot in middle. ing 

Length 20-40 mm.; males average about 5 mm. less than 
females. 

Habitat: East India; Sarawak; Java; Bachian; Ceram; 
Amboina; Kaissa; New Guinea; Australia; Sumatra. Dr. Dohrn. 

M. imsularis Smith. Types in British Museum. From 
Sarawak. Rather small, but typical specimens of ‘SS. coronator 
Fab. 

Length ¢,19mm.; 9, 20 mm.; terebra 25 mm. 


Text-fig, 2. 


Stephanus coronator. 


Stephanus coronator. 


This species greatly resembles S. ducalis Westw., from 
which it may be distinguished by the smooth semiannular 
and by the sculpture of mesonotum, mesopleure, and median 
segment. 

From S. hematipoda Montrz. it can be known by its red head 
with less prominent posterior margin, and by the sculpture of 
head, median segment, and hind coxe; from S. sulcifrons Schlett. 
it may at once be separated by the absence of the sulcus on 
vertex and by the white-banded terebral sheaths, which latter 
character will also distinguish it from the African S. pachylomerus 
Schlett. 

Proc. Zoou. Soc.—1922, No. L. 50 


740 MR. E. A. ELLIOTT ON THE 


29. pucALIS Westw. 


Megischus ducalis Westw. (4), p. 2389, 9. Stephanus ducalis 
Schlett., p. 112, 2. Stephanus lucidus Szépl., p. 532, 3. 


®. Frons coarsely transrugose; vertex coarsely arcuate 
rugose; occiput centrally distinctly impressed, arcuate rugose in 
front, becoming transverse towards posterior margin of head, 
which is sharply bordered; cheeks shorter than scape; flagellar 
joints normal. Neck coarsely transrugose, semiannular less 
coarsely, with shining smooth posterior margin. Mesonotum 
centrally smooth, diffusely punctate in front, with a rather 
indistinct central longitudinal row of punctures and two shallow 
lateral impressions, laterally and posteriorly coarsely rugose- 
punctate. Mesopleurze smooth above, rugose-punctate beneath ; 
metapleure coarsely rugose-punctate, separated by a deep trans- 
rugose sulcus from the cribrate punctate median segment. 
Petiole basally irregularly, apically finely and transversely 
rugose, shorter than rest of abdomen; 2nd segment basally 
coarsely rugose, its apex and the last segment shining, remaining 
segments dull. Terebra longer than body, sheaths black, white- 
banded before the apex. legs sparsely pubescent; hind legs 
with coxee coarsely transrugose, femora smooth and shining in 
front, finely punctate behind, bidentate; tibiz longer than 
femora, compressed not quite to middle; tarsi three-jointed. 

Black; head, scape, and base of mandibles ferruginous. Wings 
infuseate, centrally darker. 

Length 20-36 mm.; terebra 25-4] mm. 

Westwood’s type is in the British Museum. 

3S. Differs from the @Q chiefly in the occiput being more 
obliquely rugose, petiole only basally rugose, 2nd segment basally 
rugose as in 9, but remainder of abdomen smooth and shining ; 
hind tarsi five-jointed ; anterior femora partly, their tibize and all 
the tarsi often dark red. 

Length 20-23 mm. Szépligetti’s type (S. lwcidws, 3) is in 
the Hungari ian National Museum at Budapest. 

Habitat : widely distributed in the Indo-Australian Region ; 
East India, Java, Sumatra, Borneo, Pulu-Penang, Celebes, 
Sumba, Key Island, Bachian, and New Guinea. 

Schulz (2), p. 19, mentions a very large 9, 36°5 mm., in 
which the posterior angles of median segment, metapleure, 
petiole, and hind coxe are all of a metallic steel-blue or violet— 
colours not previously observed in this family. 

This species differs from S. coronator Fab. chiefly in the head 
being impressed behind the vertex; the rugose semiannular, and 
the coarse sculpture of mesonotum, mesopleurze, and median 
segment further distinguish it. From S. hematipoda Montrz. 
it may be known by the impressed head, the arcuately rugose 
vertex, less prominent posterior border of head, and by the 
sculpture of the median segment. ‘The absence of the sulcus on 
vertex and the white-banded terebral sheaths will suffice to 
separate this species from S. suleifrons Schlett. 


HYMENOPLEROUS FAMILY STEPHANIDA, 741 


30. LANCEOLATUS Kieff. 
Stephanus lanceolatus Kieffer (3), p. 277, 2. 


@. Head from antenne to the sharply bordered posterior 
margin coarsely arcuate rugose; anterior frontal tubercle scarcely 
larger than the lateral ones, posterior pair smaller, the space 
between tubercles centrally smooth. Flagellar joints normal. 
Pronotum elongate, with apical transcarina, posteriorly trans- 
striate, laterally smooth and shining. Mesonotum and median 
segment smooth, coarsely and diffusely punctate. Scutellum 
smooth and shining, coarsely punctate on margins of lobes. 
Propleurz nearly smooth; meso- and meta-pleure coarsely and 
diffusely punctate, former anteriorly subrugose. Petiole coarsely 
and closely trans-striate, as long as rest of abdomen; second 
segment smooth and shining, four times as long as broad; 
remainder transverse, shagreened, and shining. Terebra about 
half as long again as body, sheaths white-banded before apex. 
Middle and hind coxe smooth, with whitish pubescence ; hind 
femora smooth, bidentate ; tibie compressed to middle. Stigma 
very long and pointed. 

Black; head except apices of mandibles, two basal antennal 
joints, front legs except basal third, and hind tarsi red; inter- 
mediate legs black-brown. Wings infumate, distally darker, 
external median and submedian cells and thence to posterior 
margin brown ; stigma and nervures black. 

Length 24 mm. ; terebra 37 mm. 

Habitat: Obidos on the Amazon (P. Kihler), 

Distinguished from S. maculipennis Westw., especially by the 
coarsely trans-striate petiole without lateral tubercles; from 
S. rujiceps Sauss. it may be known by the arcuate rugose head 
and darker wings. 


31. HaMaAtTIPODA Montrz. 
Stephanus hematipoda Montrz., p. 114; Schlett., p. 116, 2. 


2. Frons coarsely and irregularly rugose, somewhat obliquely 
near the eyes; occiput arcuate rugose behind the tubercles, 
posteriorly centrally transrugose, laterally more irregularly. 
Temples diffusely and finely punctate. Posterior margin of head 
with slightly reflexed border. Neck very coarsely trans-striate, 
semiannular smooth, laterally coarsely punctato-rugose. Meso- 
notum centrally diffusely punctate, with a longitudinal row of 
conspicuous punctures and indistinct lateral impressions, re- 
mainder coarsely and subtransversely rugose-punctate. Scutellum 
polished smooth with a few marginal punctures. Mesopleure 
smooth above, slightly rugose in front and behind, coarsely 
rugose- punctate beneath, separated by a deep rugose suleus from 
the median segment, which is very coarsely cribrate punctate, 
laterally and apically indistinetly transrugose. Petiole coarsely, 
anteriorly irregularly, otherwise distinctly piety rugose, 

50* 


742 MR, E. A. ELLIOTT ON THE 


a little shorter than remainder of abdomen and as long as hind 
coxee, trochanters, and femora together ; second segment basally 
rugose, apically smooth, the remainder dull. ‘Terebra white- 
banded before apex. Hind legs with coxe very coarsely trans- 
rugose; femora smooth, bidentate; tibiz much longer than 
femora, compressed not quite to middle. 

Black; cheeks marked with pale red; anterior legs ferruginous, 
their coxe black and tarsi pitch-brown ; apical inflated part of 
hind tibie ferruginous. Wings strongly infumate, centrally 
darker, 

Length 30 mm. 

Habitat: Woodlark Island, NewGuinea. Bachian, Jobi Island. 

Two specimens of this species are in the Royal Nat. Hist. 
Museum in Dresden; in both the antenne are wanting and 
terebra broken; Montrouzier states the latter is white-banded. 

Schletterer considers this species very similar to S. coronator 
Fab. and S. ducalis Westw., in both of which the head is red 
and the sculpture very different. 


32. acutus Lep. & Serv, 
Megischus acutus Lep. & Sery., p. 489; Brull., p. 540, 2. 
Stephanus acutus Schlett., p. 137, 9. 


@. Frons moderately coarsely arcuately rugose; occiput 
very coarsely arcuate rugose near tubercles, otherwise irregularly 
to transrugose. Temples smooth and shining. Posterior 
margin of head elongate, reflexed collar-like. Cheeks shorter 
than scape. Basal flagellar joints normal. Neck coarsely trans- 
rugose, less distinctly basally; semiannular smooth, with coarse 
but diffuse punctures. Mesonotum shining, laterally densely, 
partly confluently punctate, a median longitudinal row of 
punctures, on each side of which is a smooth space, followed by a 
row of broad punctures. Scutellum polished, with a few marginal 
punctures. Mesopleure shining, diffusely punctate; metapleure 
densely pubescent, somewhat rugose, separated by an almost 
smooth sulcus from the median segment, which is cribrate 
punctate and apically transrugose. Petiole finely trans-striolate, 
about as long as remainder of abdomen, which is shining, with 
a few dull spots, due to microscopic sculpture. ‘Terebra a little 
longer than body, white-banded before apex. Hind legs with 
coxe trans-striolate between coarse rugosities; femora smooth, 
bidentate; tibisee compressed not quite to middle. 

Black; cheeks and mandibles pale marked ; anterior legs have 
a tendency to become brownish. Wings slightly infuscate. 

Length 21-30 mm. 

Habitat: Brazil; Colombia; Kansas, U.S.A. 

This species resembles S. niger Smith, but has the posterior 
margin of head reflexed, face arcuate rugose, mesonotum punctate, 
not rugose, and longer petiole. 


HYMENOPTEROUS FAMILY STEPHANIDA. 743 


33. VILLosuS Kieff. 

Stephanus villosus Kieffer (1), p. 4, 2 3. 

©. Face and frons irregularly and coarsely rugose, space 
between the tubercles strongly impressed; vertex with three 
arcuate transcarine ; occiput coarsely and densely longitudinally 
rugose; posterior margin of head produced into a_ broad 
horizontal translucent collar. Second flagellar joint two and a 
half times as long as first, third more than twice as long as first 
and second together and about two and a half times as long as 
second. Prothorax coarsely and rather diffusely transrugose, 
basally more strongly and more diffusely. Mesonotum smooth, 
with median line of punctures and laterally transrugose ; 
seutellum with two anteriorly convergent irregular rows of 
punctures and slightly rugose, anterior lobes diffusely punctate. 
Mesopleurze white pubescent, the impressed part transrugose. 
Metapleure and median segment with large and partly confluent 
punctures, former white pubescent. Petiole finely trans-striate, 
about as long as rest of abdomen, which is basally striate. Terebra 
as long as body, sheaths black, white-banded before apex. Hind 
legs with coxe finely trans-striate, as long as the bidentate 
femora; the tibize longer than femora, constricted in basal 
third and compressed beyond middle; tarsi three-jointed, with 
long grey pubescence beneath. 

Shining black ; head except apices of mandibles, scape, and 
second joint of hind trochanters red; anterior legs black-brown. 
Thorax and legs with long, sparse, whitish pilosity, the hairs on 
meso- and meta-pleure inserted on granular prominences. 
Wings hyaline. 

Length 16 mm. 

3. Similar to 2, but second flagellar joint not twice as 
long as first; third shorter than first and second together. 
Mesonotum red-brown ; hind tarsi five-jointed, red, less densely 
pubescent beneath ; genital organs testaceous. 

Length 12 mm. 

Habitat: Jataney, Prov. Goyaz, Brazil. Coll. M. de Saule. 

The unusual preportions of the basal flagellar joints In Q are 
very characteristic. 


34, RUFOFEMORATUS Szépl. 

Stephanus rufofemoratus Szépl., p. 532, 3. 

3. Frons coarsely rugose, vertex transrugose, occiput coarsely 
rugose; posterior margin of head bordered. Flagellar joints 
normal. Neck rugose, semiannular smooth, diffusely punctate. 
Mesonotum coarsely punctate, shining; scutellum smooth and 
shining. Mesopleure closely punctate and shining. Median 
Beemient diffusely and coarsely punctate and shining. Petiole 
only basally rugose, remainder smooth and shining, as long as 
vest of abdomen. Hind legs with coxe shining, diffusely rugose ; 
femora smooth, bidentate; tarsi five-jointed. 


44 MR. E. A. ELLIOTT ON THE 


Black; legs from femora red; basal half of hind tibiz black. 
Wings brown, centrally darker; hind wings with mdications of 
cubital and radial nervures. 

Length 22 mm. 

Habitat: Rubiana, New Georgia, Solomon Islands. Type in 
the Hungarian National Museum, Budapest. 

Hasily distinguished by the smooth petiole and red femora. 

This species closely resembles S. rubripes Morley, from which 
it differs in the smooth petiole, proportions of basal flagellar 
joints, the sculpture of semiannular, mesopleure, and hind.coxe. 


35. comMA Morley. 
Stephanus comma Morley (1), p. 105, 3. 


¢. Head semicircularly rugose in front, trans-suleate below 
scrobes. Tubercles small and transverse; vertical carina 
prominent; occiput trans-striate; posterior margin of head 
truncately bordered. Antennze very slender and elongate, 
21 mm., second flagellar joint half as long again as first, third 
half as long again as second, shorter than first and second 
together. Yhorax superficially but confluently punctate, the 
rather short prothorax transcarinate, 1ts posterior margin smooth. 
Scutellum smooth, the basal suleus crenulate. Mesopleuree 
longitudinally bisuleate; metapleure rugulose, bisuleate above. 
Petiole minutely transaciculate and apically explanate, shorter 
than rest of abdomen, which is finely alutaceous and somewhat 
shining. Hind coxe (4 mm.) rather shorter than femora (5 mm.), 
which are bidentate, with four small teeth before the central tooth 
and another beyond it. Lower nervure of external median cell 
in fore wing deflexed in the form of a comma at base, apically 
wanting ; hind wing with costal nervure only, emitting a slight 
spurious nervure from its apex below the three frenal hooks. 

Black; head except apices of mandibles, the tubercles and the 
occiput centrally, base of antennz, second and fourth segments 
laterally basally, whole of third segment, anterior legs except 
cox, and apical third of hind femora including the teeth bright 
brick-red. Hund tibiz basally black, relique desunt. Wings 
slightly infumate hyaline; stigma and nervures black ; costal and 
external submedian cells conspicuously darker. 

Length 27 mm.; abdomen 18 mm.; petiole 8 mm. 

The type was taken by Dr. W. M. Graham “in forest on 
trunk of tree” at Pbuasi, Ashanti, on 29 May, 1907. In coll. 
British Museum. 


36. PILosus Hlliott (4), p. 256, 3. 


3. There are sparse white hairs on the head, longer and 
denser on the petiole and apex of abdomen. The pleure are 
covered with dense white pilosity, and on the legs, the hairs, 
though less dense, are longer and more conspicuous. Frons, 


HYMENOPTEROUS FAMILY STEPHANID®. 745 


vertex, and occiput strongly arcuate rugose; one strong carina 
between the posterior ocelli and two behind them; ocellar space 
depressed. Posterior margin of head broad and slightly reflexed. 
Temples with a few setiferous punctures. Second flagellar joint 
fully twice as long as first; third a little longer than second. 
Prothorax transcarinate, semiannular smooth and laterally hairy. 
Scutellum centrally smooth, lateral lobes coarsely rugose-punctate. 
Metapleurz and median segment cribrate punctate, separated by 
a strong carina, which is anteriorly somewhat rugose, posteriorly 
smooth and shining. Petiole trans-striate, basally more coarsely, 
shorter than rest of abdomen, which is smooth. Hind coxe 
stout, coarsely transrugose ; femora smooth, bidentate, the teeth 
long and acuminate ; tibie as long as femora and trochanters, 
strongly compressed in basal third. 

Black; head, scape, base, and apex of anterior femora red; 
frontal tubercles apically black. Wings faintly iridescent ; 
fore wings centrally, hind wings apically infuscate ; stigma and 
nervures black-brown. 

Length 22 mm. ; abdomen 144 mm.; petiole 6 mm. 

Habitat: Palaboehan, Ratoe. M.E. Walsh (ex Frisby), 22 Feb., 
1916. Type in coll. Claude Morley. 

In the coarse sculpture and conspicuous pilosity, this species 
much resembles S. villosus Kieff. 


37. RUGOosUS Elliott (4), p. 256, 3. 


3. Frons arcuate rugose; vertex and occiput with strong 
longitudinal carine, occiput regularly, vertex less regularly 
arcuate striate ; three strong carinee behind posterior ocelli. 
Posterior margin of head bordered. Cheeks slightly shorter 
than seape, smooth. Basal flagellar joints normal. Neck trans- 
carinate, semiannular smooth, with a few punctures. Propleurze 
finely trans-striate. Mesonotum apically smooth, central row of 
punctures distinct, the smooth space on each side with a single 
puncture before and behind, laterally strongly punctate, 
separated from scutellum by two carine. Scutellum centrally 
smooth, laterally strongly rugose-punctate. Metapleure and 
median segment cribrate punctate, separated by a strong carina. 
Petiole strongly transrugose, shorter than the remaining smooth 
segments. Hind coxe coarsely trans-striate; femora smooth, 
bidentate; tibie slightly longer than femora, compressed to 
middle. 

Black; head, except apices of mandibles, and apex of thorax 
above red; anterior legs more or less rufescent. Wings hyaline. 

Length 16 mm.; abdomen 10 mm.; petiole 44 mm. 

Habitat: Java; ; Preanger Regency, near Seekoboemi. M. E. 
Marsh, 1918. Type and Go-types in coll. Claude Morley. 

This species bears a superficial resemblance to both 8. wnicolor 
Schlett. and S. ducalis Westw., but differs considerably in 
sculpture. 


746 MR. E. A. ELLIOTT ON THE 


38. VIOLACEIPENNIS Cam. 
Megischus violaceipennis Cameron (2), p. 225, 9. 


‘“‘Niger; pedibusrufis; coxis, trochanteribus, femoribus posticis 
_basique tibiarum posticarum nigris; alis fusco-violaceis, nervis 
TACT OO 

Long. 24 mm. 

Antenne black, slender. Front rugosely punctured above ; 
the sides above stoutly obliquely, the lower part transversely 
striated, the ocellar region irregularly reticulated, the keels above 
them mostly curved; above the reticulated upper portion are 
four long curved keels, the vertex behind them is irregularly 
rugosely striated ; the ocellar region is distinctly depressed ; the 
three tubercles form a triangle. The base of the pronotum is 
smooth and shining; behind on the sides are two stout curved 
keels; the rest of the pronotum is opaque and irregularly trans- 
versely striated. The base of the mesonotum is smooth and 
shining, the rest of it bears mostly large and deep punctures, 
except a space on either side of the centre; on the sides and 
base the punctures tend to become confluent. Scutellum some- 
what triangular in shape, smooth, shining and impunctate. 
Median segment closely, stoutly irregularly punctured ; behind, 
the punctures run into reticulations. Propleure smooth and 
shining; the mesopleurz opaque, irregularly punctured, and 
striated; the metapleure rugosely punctured. The four front 
legs are rufous, with the coxe darker; the hinder black, except 
the apical two-thirds of the tibie and the tarsi, which are rufous 
at the base, blackish towards the apex; the hind coxe are, as 
usual, irregularly striated ; the two teeth on the hind femora are 
acute; the apical one is longer and narrower than the hinder. 
Wings fusco-violaceous; the nervures and stigma are black. 
The petiole is longer than the rest of the abdomen united; it is 
opaque and closely striated. The ovipositor is annulated with 

' white and is not quite as long as the body. The recurrent 
nervure in the fore wing is interstitial.” 

Habitat: New Britain (Dr. Arthur Willey). ‘ype in Cam- 
bridge Museum. 

Said by Cameron to be very like his S. froggattii, but, in the 
absence of details as to the posterior border of head and relative 
length of the basal antennal joints, I am unable to place it 
definitely in my table. 


PARASTEPHANELLUS Enderl. 


In the (2) Zool. Anz. xxxill. pp. 453-475, Dr. Gunther 
Enderlein erects a new subgenus, Parastephanus, which he 
defines as having that part of the median nervure beyond the 
outer submedian cell obsolete, and the cell consequently open 
behind, the size of the discoidal cell and the breadth of the 
stigma being variable. He thus includes species from the Indo- 
Australian and Neotropical Regions. 


HYMENOPTEROUS FAMILY STEPHANIDA. TAT 


In the (4) Stett. ent. Zeit. 1906, p. 301, the name is altered to 
Parastephanellus, Parastephanus being preoceupied by Haeckel 
for Protozoa, The apparently constant difference between the 
relative sizes of the discoidal and cubital cells and of the stigma 
in the above-named two regions led to a further subdivision, 
removing the Neotropical species to a new subgenus, Hemiste- 
phanus. 

Parastephanellus, confined to the Indo-Australian Region, has 
the discoidal cell petiolate and only about one-fourth as large as 
the cubital cell; the radius does not extend to the margin of the 
fore wing, and, as a rule, the stigma is short and broad. ‘The 
sheaths of the terebra are black, except in No. 6, in which they 
are rufescent. 


Table of Species. 


(6.) 1. Hind femora tridentate. 
(3.) 2. Length15-20mm. Head red, apical abdominal segments dull. 
1. martini Stadln. 
(2.) 3. Length 7-8 mm. 
(5.) 4. Head black, pronotum transrugose; median sagan with 
large diffuse puNcthurespe see sna. 2. orbitalis Brues. 
(4.) 5. Head rufescent, white-marked ; pronotiam elattons! ; median 
segment without visible sculpture estes oc beeps 3. glaber Elliott. 
(1.) 6. Hind femora bidentate. 
(16.) 7. Petiole distinctly shorter or longer than rest of abdomen. 
(18.) 8. Petiole distinctly shorter than rest of abdomen. 
(10.) 9. Yellow-brown. Frons finely punctate, mesonotum rugose. 


A, pygmeus Ender. 
(9.) 10. Black. 


(12.) 11. Frons evenly trans-striate; neck smooth, terebra longer than 


ods Asset arses Y cces. 0. Levicollis Morley. 
(11.) 12. Frons subarcuate dates cantnetly te neck trans- f 
striate; terebra shorter than body ................ 6. albiceps Elliott. 
(8.) 13. Petiole distinctly longer than rest of abdomen. 
(15.) 14. Frons punctate, ae transrugose ; _mesopleuree smooth, 
Stigma) blackeeyesamesceee: Pes auydek oaeen ted maculifrons Cam. 
(14.) 15. Frons transrugose, ecu Aoee ‘Geass striate ; mesopleur 
trans-striate, stigma pale ......................... 8. eburneus Morley. 


(7.) 16. Petiole about same Jength as rest of abdomen. 
(20.) 17. White line on face below eyes. 
(19.) 18. The two posterior frontal tubercles subobsolete; posterior 

margin of head bordered ; antennz long and slender. 

9. damellicus Westw. 

(18.) 19. All frontal tubercles preminent; posterior margin of head 

simple; antenne short .......:.........-...-++-.--.. 10. pictipes Roman. 
(17.) 20. Head otherwise marked. 
(26.) 21. Frons arcuate striate. 
(23.) 22. Stigma very short. Head ochre-yellow ; aga trans- 

SURIAILCL WH Auer eerie aman Mees! hat Len Gn . LL. brevistigma Ender). 


(22.) 23. Stigma normal. 
(25.) 24. Frons closely subarcuate striate; pronotum trans-striate. 
12. palliditarsis Cam. 
25. Frons strongly arcuate striate; pronotum smooth. 18. seitus Elliott. 


v2) 
.) 26. Frons not arcuate striate. 


lord 
748 MR. E. A. ELLIOTT ON THE 


(28.) 27. Frons trans-striate; intertubercular space unusually long. 


14. clar ipennis Cam. 
(27.) 28. Frons rugose. 


(82.) 29. Frons rugose punctate. 


(81.) 30. Neck of pronotum eee trans-striate; median segment 
pUNCtate) ese eee: sd acitustechec fpyiling rufo- ornatus Cam. 


(30.) 31. Neck longitudinally ctnfetiar cedliem eeoaramt reticulate rugose. ; 
16. rubripictus Elliott. 


(29.) 32. Frons rugose but not punctate; metapleuree smooth; median 
segment diffusely punctate ...............0::00.. 17. malayanus Cam. 


1. MARTINI Stadlm. 
Stephanus martini Stadelmann (1), p. 236, 29 d. 


®. Frons transrugose, coarsely punctate below; occiput 
coarsely transrugose ; posterior margin of head sharply ‘bordered. 
Basal flagellar joints normal. Neck coarsely and obliquely 
rugoss; semiannular coarsely punctate with polished posterior 
margin. Mesonotum diffusely punctate. Scutellum smooth, 
with a few diffuse marginal punctures. Mesopleure shining, 
apically punctate; metapleurz densely punctate, separated by a 
deep and shining sulcus from the coarsely cribrate-punctate 
median segment. Petiole basally longitudinally, apically trans- 
versely rugose, and as long as the remaining segments together ; 
apical seements dull. inereen very little longer than body, 
sheaths entirely black. “Hind coxe tranerusoseh hind femora 
finely trans-striate and tridentate; tarsi three-jointed. 

Black; head, scape, and first flagellar joint red; anterior legs 
nigro-fuscous ; all tarsi testaceous. Wings subhyaline. 

3. Similar to 9. Nigro-fulvous, head red, antennz basaily 
red, apically nigro-ferruginous; base of hind femora ferru- 
ginous; tarsi five-jointed. Length, 2, 17-20 mm.; ¢, 15 mm: 

Habitat: Dele, Sumatra. Named after the captor, Hofrat 
Dr. med. L. Martin. _ Type in Berlin Museum. 

Stadelmann states that this species closely resembles S. sulez- 
Jrons Schlett., which has the neuration of the subgenus Stephanus; 
he states also that the neuration is complete, but his figure 
(J. c. pl. 6. fig. 4) shows the external submedian cell open behind 
as in Parastephanellus. 

Other differences are said to be: the sulcus on vertex indistinct 
in 9, wanting in ¢ ; third flagellar joint longer, petiole longer 
and more coarsely sculptured; hind femora trans-striate and 
tridentate, but smooth and bidentate in sulcifrons. 

Easily known by the tridentate hind femora and large size. 


2. ORBITALIS Brues. 
Parastephanellus orbitalis C. L. Brues, p. 98, 3. 


3. Frons and vertex centrally transrugose; cheeks longer 
than scape; posterior margin of head bordered but not reflexed. 
Basal flagellar joints normal. Pronotum transrugose, the 
rugosities extending obliquely to the pleure; a punctate area on 
each side before the smooth posterior margin. Mesonotum 


HYMENOPTEROUS FAMILY STEPHANIDA, 749 


sparsely foveate punctate, with punctate median line ; scutellum 
smooth, with coarse lateral punctures. Mesopleure pubescent in 
front, smooth and shining behind; metapleure irregularly 
rugose. Median segment with Hteaces large, and shallow 
punctures, separated “from the metapleuree by a smooth sulcus 
and a reticulate ridge. Petiole two-thirds as long as thorax, 
trans-striate above, irregularly rugose laterally Amel below ; 
remaining segments polished. Claspers short, basally smooth, 
apical half punctate and hairy, apex rounded, with dense fringe 
of short hairs. Hind coxe trans-striate, with a smooth area 
externally near the base. Hund femora tridentate. 

Black; face to just above antenne and inner orbits narrowly 
pale yellowish, outer orbits more widely above white. Palpi and 
antenne piceous, bage of flagellum fuscous; propleure below 
rufous ; front legs anteriorly brown-yellow ; all knees and apices 
of tibie are whitish, all tarsi honey- yellow. Wings slightly 
infuscate; stigma about half as wide as radial cell, petiole of 
discoidal cell short. 

Length 7-8 mm. 

Habitat: Ful kora, Solomon Islands | F. W. Mann). Type and 
paratype in the Museum of Comparative Zoology at Harvard 
College, U.S.A. 

The black head and the sculpture of pronotum and median 
segment will distinguish this species from the only other of 
similar size with tridentate hind femora (glaber Elliott). 


3. GLABER Elliott. 
Parastephanellus glaber Elliott (2), p. 129, ©. 


©. Frons arcuate rugose, vertex strongly transrugose, 
occiput becoming gradually smoother towards the slightly raised 
posterior margin, and with distinct longitudinal impression ; 
carine between the posterior ocelli very inconspicuous, scarcely 
to be distinguished from the transverse strie; anterior frontal 
tubercle very prominent. Cheeks shorter than scape ; basal 
flagellar joints normal. Prothorax glabrous. Neck rather short ; 
mesonotum smooth ; scutellum with marginal punctures only ; 
pro- and meso-pleure very finely alutaceous, metapleuree almost 
smooth and median segment with scarcely any visible sculpture. 
The pleure, abdomen beneath, and the legs with erect white 
hairs. Petiole very finely trans-striate, slightly longer than rest 
of abdomen, which is smooth. Terebra longer than body, 
sheaths black. Hind legs with coxe finely trans-striate, femora 
smooth, tridentate ; tibia compressed to middle. 

Black ; head rufescent, with white line under eyes as in 
P. damellicus Westw.; a testaceous mark on frons just below 
the anterior tubercle, which is pale rufescent. Legs rufescent, 
base and apex of tibize paler and all metatarsi whitish. 

Length 8 mm.; abdomen 43 mm.; petiole 24 mm.; terebra 
11 mm. 


Habitat: Singapore. ‘‘F. Sm. Coll.” 


750 MR. E. A. ELLIOTT ON THE 


The almost entire absence of visible sculpture is characteristic. 

The colour of head and legs and the sculpture will easily 
distinguish P. orbitalis Brues, which is the only other species of 
similar size with tridentate hind femora. 


4, pyemMamus Hnderl. 


Stephanus pygmeus Hnderl. (1), p. 197, 2 go. Parastepha- 
nellus pygmeus Knderl. (2), p. 472 et seqg.; (4), p. 289 et seqg. 


9 3. Frons extremely finely and densely punctate; all five 
frontal tubercles acuminate, the posterior pair smaller; vertex 
and occiput obsoletely transaciculate ; posterior margin of head 
sharply bordered. Antenne very slender, second flagellar joint 
scarcely one and a half times as long as first; third shorter than 
first and second together. Neck almost smooth, its posterior 
margin polished. Mesonotum rugose; scutellum polished 
smooth ; metanotum (the small visible part of the true meta- 
notum) longitudinally striate. Mesopleuree smooth, pubescent, 
diffusely punctate in front only; metapleure coarsely and 
deeply rugose, separated by a shining smooth space and a row of 
punctures from the densely punctate median segment. Petiole 
finely rugose and shorter than the remaining segments together. 
Terebra about two-thirds of the length of body; spicula red- 
brown, sheaths entirely black. Legs covered with fine erect 
hairs, hind femora bidentate. 

Yellow-brown ; apical two-thirds of antenne, apex of man- 
dibles, meso- and meta-thorax, pleure and median segment 
darker, yellow-brown to black-brown. Wings hyaline, external 
submedian cell infumate ; stigma short and broad, dark brown. 

Length, 9, 7-8 mm.; 6, 5-7 mm. 

Seven ¢ ¢ and 3 2 Q in Berlin Zoological Museum ; collected 
by Nicholitz at Milne Bay, New Guinea. 

Dark specimens of the present species greatly resemble lighter 
ones of P. damellicus Westw., also from New Guinea, but are 
easily known by the shorter and broader stigma, infumate spot 
in wing, shorter petiole, and, in @, shorter terebra. 


5, LH#VICOLLIS Morley. 
Parastephanellus levicollis Morley (1), p. 111, 2 ¢o. 


Q. Head finely and evenly trans-striate; vertex coarsely 
transcarinate ; posterior margin of head bordered ; cheeks smooth 
and a little longer than scape. Second flagellar joint half as 
long again as first; third half as long again as second. Neck 
very short and smooth, semiannular smooth, with a few fine 
punctures, as is also the mesonotum. Scutellum quite smooth 
and separated from mesonotum by crenulated lines. Mesopleure 
and metapleurz obsoletely shagreened, not punctate, the latter 
separated from metanotum by a basally obsolete carina. Median 
segment centrally smooth, apically rugulose, and basally with a few 


HYMENOPTEROUS FAMILY SYTEPHANIDA, 751 


large punctnres. Petiole very finely transaciculate, shorter than 
the remaining segments, which are smooth; second segment 
basally somewhat elongately attenuate. Terebra longer than 
body, sheaths entirely black. Hind coxe transaciculate, their 
femora smooth with a few setiferous punctures, bidentate; tibize 
longer than femora, constricted to middle. 

Black; antennz and palpi infuscate; base of antenne, 
mandibles except apices, clypeus and frons fulvous; face and 
whole external orbits and all tarsi whitish. Wings hyaline, 
stigma and nervures piceous. 

¢ differs in having the whole head except the whitish external 
orbits dull red ; base of occiput nitidulous; basal flagellar joints 
longer, though in the same proportions; propleure dull testa- 
ceous; anterior femora and tibiz piceous. 

Length 9 mm.; abdomen 5 mm.; petiole 2 mm.; terebra in 
9 llmm. A single pair, now in British Museum, taken in the 
Ding-Ding Island in the Indian Ocean during 1896 and 1900. 


6. ALBICEPS Elliott. 
Parastephanellus albiceps Elliott (2), p. 130, 2. 


@, Frons subarcuate striate, the strie broken into granules, 
vertex and occiput finely granulate striate ; second flagellar joint 
one and a half times as long as first, third slightly shorter than 
first and second together. Pronotum very finely trans-striate 
throughout, neck very short; mesonotum subglabrous; scutellum 
normal; propleure finely, obliquely striate; mesopleure aluta- 
ceous; metapleure punctate, separated by a deep crenulated 
sulcus from the alutaceous and punctate median segment. 
Petiole trans-striate, shorter than the rest of the smooth 
abdomen. ‘Terebra rather shorter than body, rufescent, apically 
black. Hind legs with cox trans-striate, femora finely 
alutaceous, bidentate; tibie compressed to middle. 

Black; mandibles basally flavous, all orbits broadly white ; 
antenne basally, anterior legs except middle coxze rufescent, hind 
tibie and tarsi much paler. Wings hyaline; stigma and 
nervures brown. 

Length 93 mm.; abdomen 53 mm.; petiole 24 mm.; terebra 

2 sea, 
“Habitat: Victoria, Australia. R. E. Turner, 1907. 
Easily known by the extremely broadly white orbits. 


7. MACULIFRONS Cam. 
Megiseleus maculifrons Cam. (3), 1902, p. 32, 3. 


3g. Frons closely punctate; vertex and occiput transrugose ; 
posterior margin of head bordered. Second flagellar joint twice 
as long as first, third shorter than first and second together. 
Prothorax short, nearly smooth and shining, neck with a 
transverse carina; mesothorax closely and finely punctate, dull ; 


Mea, MR. E. A. ELLIOTT ON THE 


scutellum smooth. Mesopleuree nearly smooth; metapleuree 
transrugose, separated by a sulcus from the cribrate-punctate 
median segment. Petiole trans-striate, longer than rest of 
abdomen, which is feebly shining, discally depressed. Hind legs 
with coxe trans-striate above; femora alutaceous, shining, 
bidentate ; tibize constricted to beyond middle. 

Black; head red, upper part of frons, the vertex, and occiput 
black, with a broad red mark behind the posterior ocelli; outer 
orbits yellowish ; scape and basal flagellar joint rufous. Wings 
subfuscous, nervures black. 

Length 12 mm. 

Habitat: Baram District, Low Country, Sarawak. Robt. 
Shelford. Type in British Museum. 

The peculiar generic name “ Megiseleus” appears to be a 
printer’s error. If the writing were indistinct, the ‘‘ ch” might 
easily be mistaken for ‘‘ ele.” 


b) 


8. EBURNEUS Morley. 
Parastephanellus eburneus Morley (1), p. 35, 2 og. 


¢@. Frons rugose, trans-striate. The three anterior frontal 
tubercles acuminate, front one longest and recurved; posterior 
pair smaller and rounded. Occiput finely trans-striate. Posterior 
margin of head bordered. Second flagellar jomt one and a half 
times as long as first; third shorter than first and second 
together. Neck short, smooth, and shining; semiannular finely 
rugose. Mesopleurze obsoletely trans-striate and dull; meta- 
pleuree smooth and shining, separated by a deep sulcus from the 
median segment, which is smooth, with large, diffuse punctures. 
Petiole with basal half finely trans-striate, longer than remainder 
of abdomen, which is dull and obsoletely punctate. ‘Terebra 
shorter than body, sheaths black. Hind legs with coxe obsoletely 
transaciculate ; femora nearly smooth and shining, bidentate; 
tibiee compressed to middle. 

Black; face, clypeus, mandibles except apically, and a streak at 
the inner orbits testaceous white; a broad stripe from base of 
mandibles to near occiput ivory-white ; scape and basal flagellar 
joint testaceous; base and apex of intermediate tibie white. 
Wings clear hyaline, stigma pale. 

Length 6 mm.; terebra 5 mm. 

¢. Similar to @; petiole piceous, genital valvule testaceous. 

Length 33 mm. 

The @ was taken by O, 8. Wickwar at Candy, Ceylon, Feb. 
1910; the ¢ by Green, on a leaf infested by a Coccid, Hemi- 
chionaspis brasiliensis, at Pradenula, Ceylon, June 1909. Types 
in British Museum. i 

Similar to the Australian P. rufo-ornatus Cam., 3, but the 
petiole is longer, the sculpture of median segment is different, 
and the coloration is very distinctive. 


HYMENOPTEROUS FAMILY STEPHANIDA, 153 


9. DAMELLICUS Westw. 


Stephanus damellicus Westw. (5), p. 126, pl. xxiv. fig. 2; 
Schlett.,p.116, 2. Hemistephanus damellicus Morley (1), p. 112. 
Parastephanellus d. (2), p. 29, 2. 


Westwood’s original description is as follows :— 

“Niger; capite et antennarum basi piceo-rufis; pedunculo 
abdominis pedibusque rufis; capite antice trituberculato verticeque 
carina parva transversa instructo, linea tenui alba utrinque sub 
oculos; antennis gracillimis, pone medium fuscis; collari trigono- 
truncato, collo levi, metanoto basi serie striolarum brevissimarum 
notato, pedunculo abdomen longitudine equanti, transversim 
levissime striolato; abdomine ovali nitido, oviductu longitudine 
caput et corpus totum equanti; pedibus posticis crassis, coxis 
elongatis transversim carinatis, femoribus clavatis, subtus serratis 
dentibusque tribus majoribus armatis; tibiis pone medium subito 
dilatis; tarsi posticis 3-articulatis; alis hyalinis venis distinctis 
nigris, stigmate nigro basi lutescenti.— Hab. Austral. In Mus. 
Hopeano Oxoniz. 

“The mandibles are robust, their inner margin (seen from 
above) has two obtuse teeth, seen laterally they are more 
irregular, The marginal cell is slightly open atthe tip, the vein 
forming the hinder margin of the incomplete second submarginal 
cell is abbreviated, and the second discoidal cell is incomplete, 
the vein forming its hinder margin being obsolete. 

“Species proxima Stephanus hematipoda Montrouzier, Ann. 
Soc. Agric. Lyon, T. vil. 1, p. 113. Hab. Woodlark Island.” 

Having examined the beautifully preserved type, I am able to 
correct and complete the above description. 

Frons anteriorly rugose-punctate, transrugose above, vertex 
with avery conspicuous transverse carina, behind which the 
head is areuate rugose, becoming trans-striate towards the 
strongly bordered posterior margin. The three anterior tubercles 
very prominent, the two posterior subobsolete. Antenne very 
slender ; basal flagellar joints normal. Neck smooth, semiannular 
transrugose. Mesonotum coarsely but not closely rugose- 
punctate; metanotum (the small strip visible) longitudinally 
striate. Mesopieure finely punctate; metapleure punctate and 
separated from the rugose median segment by a line of 
punctures. Petiole finely striate throughout, slightly shorter 
than remainder of abdomen, which is smooth and shining. 
Terebra slightly shorter than body. Hind legs with coxee strongly 
trans-striate; femora bidentate; tibiz compressed to slightly 
beyond the middle. Black; the anterior tubercles and the carina 
on vertex are bright red, remainder of head and base of antenne 
dull red, with a broad white line on each side, starting from the 
posterior margin, passing under the eyes, and meeting over the 
mouth. Neck of pronotum, mesonotum partly, petiole, and legs 
red. Wings hyaline, stigma rufescent, basally paler, nervures 
black. 


754 MR. E. A, ELLIOTT ON THE 


Length 84 mm.; abdomen 5 mm.; petiole 2 mm.; terebra 
8 mm. 

The “carina parva transversa” is unusually large, and its 
bright red colour makes it very conspicuous. The small visible 
portion of the true metanotum is longitudinally striate, but the 
median segment is rugose. The hind femora are furnished in 
the anterior half beneath with two comparatively long and very 
sharp spine-like teeth, and can only be called bidentate. There 
is a slight blunt projection near the base, more pronounced on 
the right femur than on the left, but these do not count as teeth. 
The white line on the head, extending, as one might say, from 
ear to ear, is very characteristic. 

Enderlein ((1), p. 196) describes both sexes of a Paraste- 
phanellus, which he considers to be identical with the present 
species, differing in colour only, The neck coarsely longitudinally 
striate, the semiannular slightly rugose in front, smooth behind ; 
the petiole as long as remainder of abdomen and apically smooth. 
No mention is made of the conspicuous carina on vertex, nor of 
-the white line on head, also petiole and legs are differently 
coloured. They are closely-allied species, but that of Westwood 
comes from Australia, whereas Enderlein’s was taken in New 
Guinea, Bismarck Archipelago, and Borneo. For this species— 
rubripictus Elliott—see p. 759. 

The measurements given by Westwood in lines are approxi- 
mately correct, and the petiole, remainder of abdomen, and 
terebra appear to be of equal length. Accurate measurements 
show slight differences, which may be ignored for purposes of 
identification, as in my table. 

The present species is closely allied to, and may even be 
identical with P. pictipes Roman. 

The British Museum has of this species 7 9 2, taken by 
R. E. Turner at Mackay, Queenlsand, and 1 Q from the Swan 
River. They vary in size from 63 mm., terebra 7 to 13 mm., 
terebra 144 mm. The only difference from the type is in the 
slightly longer terebra, which is shorter than the body in the 
type but longer in these. I do not consider this sufficient to 
create a new species. 


10. pricTIPEs Roman. 
Parastephanellus pictipes Roman (1), p. 1, ©. 


@. Head seen laterally scarcely higher than long.  Frons 
strongly but scarcely transversely rugose ; the five tubercles rather 
prominent but obtuse, the vertex behind them furnished with a 
short carina; occiput towards base more coarsely and more 
transversely striate, with indistinct longitudinal impression ; 
posterior margin of head emarginate, not bordered. Antenne 
short, scarcely reaching beyond thorax, 23-jointed; pedicellus 
sub-elongate, half as long as scape; first flagellar joint scarcely 
longer than scape, second one and a half times as long as first ; 


HYMENOPTEROUS FAMILY STEPHANID®, 755 


third and fourth of equal length and little longer than second. 
Thorax sub-depressed, pronotum closely and finely trans-striate, 
neck apically deeply emarginate and bifoveolate. Mesonotum 
coarsely rugose-punctate, with anterior central longitudinal line 
of punctures, a strongly crenulate line before scutellum, which is 
rugose, apically subglabrous, alutaceous. Mesopleure tumid, 
scabrous, with broad and smooth posterior margin; metapleurze 
prominent, irregularlyscabrous. Sternum smooth and shining, very 
finely trans-striolate, the punctate mesoleus open behind. Median 
segment depressed, coarsely rugose-reticulate, basally laterally 
move finely sculptured. Petiole about as long as remainder of 
abdomen, centrally slightly incrassate, trans-striate, with smooth 
apical margin; second segment half as long as first, shining, 
microscopically reticulate; remaining segments transverse, third 
to seventh finely and closely trans-striolate, eighth exserted, 
coriaceous. Apical margins of ventral segments sparsely pale 
pubescent ; hypopygium apically compressed and subacuminate. 
Terebra shorter than body, sheaths black. Hind legs with cox 
trans-striolate inside ; femora bidentate, reticulate, basally exter- 
nally smooth and shining; tibiz reticulate, half as long again as 
femora, with a central, shining, acute-angled impression behind 
(2 constricted to middle and there impressed). Wings extending 
beyond apex of petiole; in fore wing, stigma nearly five times as 
long as its breadth, costa not extending beyond it; in hind wings 
the costa extends to within a third of the apex of wing. 

Black; basal half of flagellum, a straight line on cheeks from 
base of mandibles, touching the eyes, to the temples—not reach- 
ing the posterior margin of temples—apices of coxe, trochanters 
partly, extreme base and apex of anterior femora, anterior tibiz 
chiefly, posterior tibiz basally, and all tarsi red; intermediate tarsi 
basally white. Wings hyaline, stigma and nervures black, former 
basally narrowly pale; radix rufescent, tegule nigro-fuscous. 

Length 10-11 mm.; terebra 9°5 mm. 

Taken by Dr. E. Mjdberg’s Swedish Scientific Expedition to 
Australia, at Derby, Kimberley Division, West Australia, 
OctalkOLO oe 

This is possibly a dark form of P. damellicus (Westw.), 
Enderl., as both have the same unusual streak of colour on the 
cheeks. But Westwood describes his species as having three 
large teeth on the hind femora, whereas the present species 
has only two. Enderlein (Arch. Naturg. 1901) describes a 
supposed damellicus Westw., from New Guinea, the Bismarck 
Archipelago, and Borneo, which has two femoral teeth but not 
the long pale streak on cheeks, and also differs from pictipes in 
the sculpture of the pronotum. The present species is now the 
only Parastephanellus in the Stockholm Museum; the nearest 
species with which it might be compared is the somewhat unlike 
Hemistephanus texanus Cresson, A detailed description has 


been given so that this species may in future be clearly under- 
stood. 


Proc. Zoou, Soc.— 1922, No, Li. 5] 


756 MR. E. A. ELLIOTT ON THE 


Westwood’s incorrect description of his damellicus as having 
the hind femora tridentate has caused much confusion. The 
present species appears to differ chiefly in the five frontal 
tubercles being prominent, posterior margin of head not bordered, 
and pronotum somewhat differently sculptured. The peculiar 
coloration of the head points to close connection between the 
two, but I do not venture to synonymize them for the present. 


11. BRrevistigmA Enderl. 
Parastephanellus brevistigma Enderl. (8), p. 203, &. 


3. Frons finely arcuate transrugose ; the three anterior frontal 
tubercles acuminate, the two posterior flatly rounded; vertex with 
three trans-carine, the space between them polished; occiput finely 
and closely trans-striate. Antenne slender and about 25-jointed. 
Pronotum finely trans-striate, its lateral angles sharply trans- 
rugose, posterior margin polished smooth, neck short. Mesonotum 
coarsely rugose-punctate; scutellum smooth. Mesopleure smoeth, 
very finely striate ; metapleure smooth, reticulately punctate in 
front above, separated by a row of deep punctures from the 
median segment, which is coarsely, but not reticulately punctate. 
Petiole slender, finely and irregularly transrugose, about as long 
as remainder of abdomen. Hind coxe finely trans-striate, often 
granulate ; femora bidentate. 

Head clear ochre-yellow, vertex slightly nigrescent ; apical 
two-thirds of antenne brown; prothorax ferruginous, posterior 
half black above; remainder oor thorax, abdomen, hind coxe, 
femora, and tibice black. Base of second segment and whole of 
anterior legs ferruginous. Wings hyaline, faintly infuscate ; 
stigma # mm. long, and, like the nervures, brown. 

Length 6:4 mm.; abdomen 3°9 mm.; petiole 1:9 mm.; fore 
wing 4:4 mm. 

Habitat: Hankau (Koshun), Formosa; 28th April, 1912. 
Type in Dahlen Museum. (H. Sauter.) 


2. PALLIDITARSIS Cam. 


Megischus palliditarsis Cam. (8), p. 56, 2. I. ruficollis Cam. 
LCE Oils ie 


© $. Frons closely subarcnate trans-striate; vertex more 
finely trans-striate and occiput nearly smooth. Three frontal 
tubercles short and bluntly rounded in 9, more pointed in ¢. 
Postertor tubercles distinct, but broad and blunt. Posterior 
margin of head simple. A strong carina between the posterior 
ocelli and two smaller ones behind it. Second flagellar joint a little 
longer than first and about as much shorter than third. Pro- 
notum, pro- and meso-pleuree smooth; scutellum smooth, with a 
few diffuse punctures; the bordering furrows weakly crenulated 
in 9,smoothin ¢. Metapleure smooth in 9°, slightly aciculated 


a 


in ¢. Petiole closely trans-striate throughout, longer than rest 


HYMENOPTEROUS FAMILY STEPHANID. 757 


of abdomen, whichis smooth. Hind legs with coxe closely trans- 
striate, femora smooth and bidentate; tibize constricted to beyond 
middle in Q, not quite sofarin d¢. Terebra in 2 longer than 
body. 

@. Black: head ruto-testaceous, outer orbits broadly white, 
propleure and prosternum rufous, base of second segment and 
the anterior legs rufo-testaceous; middle tibiw basally broadly, 
apically narrowly white; hind legs black. Front tarsi partly, 
middle tarsi entirely, and hind metatarsus except at apex white. 

3g. Similar to 9, but head darker red, face and oral region 
pale; prothorax entirely red, basal antennal joints paler, anterior 
legs testaceous, femora darker, hind legs black with knees rufous, 
constricted part of tibiz and all tarsi white. Wings, in both 
sexes, hyaline, nervures and stigma rufous in 9, darker, with 
pale base to stigma in ¢. 

Length, 9,6 mm.; abdomen 33 mm.; petiole 2mm.; terebra 
9mm, Length, d,7mm.; abdomen 33 mm.; petiole 2 mm. 

There is no doubt that the two species of Cameren are ¢ and 
2 of the same. His statement that the hind femora of MW, ruji- 
collis are trans-striate is an evident mistake for coxe. There are 
few things more deceptive than the relative proportions of petiole 
and rest of abdomen. Cameron gives the petiole in this species as 
“twice longer”; the actual measurements are 2 and 14 mm. 

Habitat: Kuching, Borneo. John Hewitt. Types in British 
Museum. 


13. scrrus Elhott. 
Parastephanellus scitus Hlliott (2), p. 163, 3. 


3. Frons strongly arcuate striate, vertex and occiput trans- 
striate, posterior frontal tubercles subobsolete, two carinze between 
the posterior ocelli; posterior margin of head bordered. Scape 
longer than cheeks; second flagellar joint one and a half times as 
long as first; third rather longer than second. Pronotum trans- 
striate, the extreme base smooth, neck short; mesonotum finely 
punctate; scutellum normal; propleurz obliquely striate; meso- 
pleuree smooth; metapleurz coarsely punctate and separated by a 
sulcus from the closely and finely punctate median segment. 
Petiole finely trans-striate, as long as rest of abdomen, which is 
smooth and shining, Hind coxe coarsely, their femora finely 
trans-striate, the latter somewhat strongly inflated, bidentate; 
tibiz compressed to a little beyond middle. 

Black; head dark red, antenne pale rufescent, becoming 
gradually darker towards the apex; extreme base of pronotum, 
front legs, middle tibize and tarsi, hind tarsi, and base of second 
segment rufescent. There is a scarcely noticeable paler line 
under the eyes. Wings hyaline, slightly iridescent; stigma 
rufescent, basally pale; nervures blackish. 

Length 8 mm.; abdomen 5 mm.; petiole 24 mm. 

Habitat: Pusa, Bengal. (T. B. F.) 21.vi.11. 

51* 


758 MR. E. A. ELLIOTT ON THE 


This is the first specimen of this subgenus I have seen from 
India proper, all the others being from more southern localities ; 
one from Ceylon, one from Ding- Ding Island in the Indian 
Ocean, and several from Borneo and Mnetaralte, It differs from 
all other males yet described in the sculpture of the head and 
hind legs, and in colour. 

The males are decidedly rarer than the females, and the mate 
of this species should be discovered ere long. 


14. CLARIPENNIS Cam. 
Megischus claripennis Cam. (8), p. 56, @. 


@.+Frons finely and closely transrugose, vertex subarcuate 
striate, occiput finely striate, almost smooth behind. The anterior 
frontal tubercle longest, and separated from the next pair by a 
greater distance than these are from each other, the enclosed 
space being unusually elongate. Three carine between the 
posterior ocelli, the front one strongest. Posterior margin of 
head strongly bordered. Second flagellar joint half as long again 
as first; third longer than second, but shorter than first and 
second together. Py onotum entirely smooth and shining; meso- 
notum shagreened, and with longitudinal furrow; scutellum 
elongate and smooth. Propleurse smooth; mesopleure aciculate ; 
metapleure obliquely striate. Median segment strongly reticu- 
late punctate. Petiole longer than rest of abdomen, closely 
trans-striate; remainder of abdomen smooth. Terebra half as 
long again as body, sheaths black. Hind coxe closely trans- 
striate; hind femora smooth, bidentate, their tibize compressed to 
a little beyond middle. 

Black; the outer orbits broadly, the anterior tarsi and the 
middle tibiz testaceous, hind metatarsus basally white; head 
rufo-testaceous, vertex blackish, four basal antennal joints pale 
rufo-testaceous; the second segment basally rufous. Wings clear 
hyaline; stigma and nervures almost black. 

Length 8 mm.; abdomen 33 mm.; petiole 2 mm.; terebra 
13 mm. : 

Habitat: Kuching, Borneo. John Hewitt, B.A. Type in 
British Museum. 


15. RUFO-ORNATUS Cam. 


Stephanus rufo-ornatus Cam. (5), p. 45, 3. Parastephanellus 
rufo-ornatus Klliott (1), p.92, d. P. rufid-ornatus Morley (1), 
pp. 35 et 112, 29S. 


3. Frous closely, rugosely, and almost reticulately punctate ; 
vertex closely trans-striate; arcuate stris between frontal 
tubercles, and a distinct genvreal longitudinal suleus on vertex. 
Second flagellar joint one and a male times as long as first. 
Pronotum ‘short, finely and closely trans-striate. Mesonotum 
coarsely rugose; scutellum smooth, with two short lateral 


HYMENOPTEROUS FAMILY STEPHANIDA. 759 


Impressions at apex, between which it is striate. Pro- and 
meso-pleure obscurely striate; apical half of metapleure 
irregularly reticulate. Median segment finely and closely 
punctate, apically centrally striate. Petiole finely and closely 
trans-striate, as long as rest of abdomen. Hind femora bi- 
dentate. 

Black; a broad yellow line from base of antennze to beyond 
eyes; mandibles pale yellow; antenne, frontal tubercles, 
scutellum, apex of petiole, anterior legs, hind cox partly, 
trochanters, apices of femora, the tibiz and tarsi rufous. Wings 
clear hyaline; stigma fuscous, basally pale; nervures brown. 
Abdomen from second segment sometimes red. 

Length 5-10 mm. 

 ®. Similar to g. Petiole slightly shorter; terebra longer 
than body. 

Length 13-14 mm.; abdomen 83 mm.; petiole 4 mm.; terebra 
15 mm. 

Habitat: ¢. Burnett River, Queensland(R.Semon); Kuranda, 
Queensland (Dodd), var. Kuranda, 1904. 92. Mackay, Queens- 
land ; Swan River, West Australia. 

The 9 @, together with the ¢ ¢ from Kuranda, are in the 
British Museum. 

Kasily distinguished from P. brevistigma Enderl. by the sculp- 
ture of metapleure and median segment, and by the colour. 


16. ruBRipictus Elliott. 


Stephanus damellicus Enderl. (nec Westw.) (1), p. 196, 2 3. 
Parastephanellus rubripictus Elliott (1), p. 129, 2 3. 

@ 3. Frons rather coarsely rugose-punctate, laterally trans- 
rugose; the three anterior tubercles very acuminate, posterior 
pair small; vertex and occiput transrugose ; posterior margin of 
head sharply bordered. Antenne very slender; basal flagellar 
joints normal. Neck coarsely longitudinally striate; semiannular 
feebly rugose, with polished posterior margin. Mesonotum 
diffusely and coarsely rugose-punctate. Scutellum centrally 
smooth, with marginal puncturation. Metanotum longitudinally 
striate. Mesopleurz pubescent, finely and diffusely punctate ; 
metapleurz separated by a row of punctures from the coarsely 
and reticulately rugose median segment. Petiole finely rugose, 
apically more or less smooth, as long as the remaining segments 
together. Terebra in 2 as long as body, spicula red-brown, 
sheaths black. Legs finely pubescent; hind coxee densely trans- 
rugose; hind femora bidentate, their tibie compressed almost to 
middle. 

Black; face, mandibles except apices, cheeks, 4 or 5 hasal 
antennal joints, anterior legs and hind tibie red-brown; petiole 
and hind femora sometimes brownish or rufescent. Wings clear 
hyaline; stigma long and narrow, brown. 

Length, 2, 11-12 mm.; ¢, 6-11 mm. 


760 MR. E. A. ELLIOT! ON THE 


Habitat: Milne Bay (Micholitz); Bismarck Archipelago 
(Dr. Dahl); Borneo (Grabowsky). 

This species is distinguished from all others in this subgenus 
by the long and narrow stigma; from P. pygmeus Enderl. it 
differs also in the longer basal flagellar joints and entirely hyaline 
wings. 

Enderlein (/.¢.) considers this species to be merely a dark 
variety of P. damellicus Westw. The sculpture of the pronotum 
is entirely different, the petiole and hind legs are black, the 
characteristic white line on face is wanting, and the stigma is 
longer and narrower. These points, taken in conjunction with 
the different localities, appear to justify me in regarding them as 
separate species. 


17. MALAYANUS Cam. 
Stephanus malayanus Cameron (6), p. 43, 2. 


Q. Frons closely and coarsely transrugose; vertex arcuate 
striate. Anterior frontal tubercle very large; temples broad and 
inflated. Second flagellar joint half as long again as first. 
Pronotum striate at apex of basal fovea; propleurze smooth 
above, closely striate beneath ; mesopleure nearly smooth; meta- 
pleure apically coarsely reticulate. Scutellum smooth, with 
large basal transverse fovea, from which two lines of foveate 
punctures run to apex. Median segment smooth, with large 
diffuse punctures. Petiole closely trans-striate, as long as 
remainder of abdomen. ‘Terebra slightly longer than body, 
sheaths black. Hind coxe irregularly trans-striate; hind femora 
bidentate. 

Black; outer orbits, front tubercle, face, frons, and scape more 
or less obscurely red; anterior legs brown, their tarsi paler ; 
basal joints of hind tibize testaceous. Wings clear hyaline; 
stigma and nervures black. 

Length 11 mm.; terebra 12 mm. 

Habitat: Mankwari, New Guinea (May). 

Cameron states that this species much resembles P. maculifrons 
Cam., from Borneo, but is more slenderly built, with hyaline 
wings and a fovea at base of scutellum, maculifrons having the 
wings infumate, no fovea at base of scutellum. 


HEMISTEPHANUS. 
Enderlein (4), pp. 189 et 301. 


Dr. Giinther Enderlein defines this subgenus as having that 
part of the median nervure beyond the external submedian cell 
obsolete and the cell open behind; the discoidal cell almost as 
large as the cubital and always touching the submedian cell; 
stigma long and narrow. Type: H. macrurus Schlett. 

This group, or subgenus, appears to be confined to South 
America, being at present known chiefly from Brazil, with a few 


HYMENOPTEROUS FAMILY STEPHANID&. 761 


from Colombia, Peru, and Chili. It seems probable that this is 
not the true distribution, or rather, that they are spread through- 
out the whole of the warm parts of the Southern Continent, their 
apparent absence being due to the absence of collectors. 

‘The males are poor iy represented, only 6 out of 17 species here 
recorded being known in that sex. Szépligeti briefly mentions 
S. (71.) wiistenti, 3 , stating merely “ 34, tarsi four-jointed, meta- 
tarsus brown” (p. 534). The statement that the hind tarsi are 
four-jointed is repeated with respect to several other species, and 
seems inexplicable, seeing that he correctly gives others as five- 
jointed. 

As a general rule, they are black insects with red head and 
nearly hyaline, or evenly infuscate wings. The known exceptions 
are: H. marginalis Schlett. with black head, and H. maculipennis 
and submaculatus Westw. with centrally darker wings; the types 
of the two last are in the British Museum. 

Roman works out this subgenus in (2), largely based on material 
collected by himself in the Amazon Region in 1914-15. He 
does not consider the difference in neuration of the fore wing to 
be of sufficient value to justify the erection of a new genus—I 
prefer to call it a subgenus—angl proposes to call it merely a 
group of the genus Stephanus. ‘Three main reasons for this are 
advanced :—lst. That they differ from Stephanus, s. str., only in a 
single wing character, and that there is in the Stockholm 
Museum a transition form in which the external submedian 
cell is only slightly open. It is not unusual to find in Stephanus, 
s. str., that the apical transverse nervure bounding this cell does 
not actually join the median nervure, thus leaving the outer lower 
angle of the cell slightly open, but the median nervure, never- 
theless, extends to the full length of the cell. 2nd. As far as is 
known, all the 8. American Stephanus, s. str., are 25 mm. or 
more in length, Hemistephanus all less than 25mm. I am 
unable to decide as to the value of this fact. 3rd. Megischus 
tecanus Oress., originally described from a specimen with 
mutilated wings, is said to be represented in the Stockholm 
Museum by a “topotype and to be a Hemistephanus, but differing 
in colour and sculpture from the S. American species, showing 
strong aftinities te the N. American species of Stephanus, s. str, 
T should be not inclined to withdraw the subgenus on this evidence 
alone. There may be a mistake in identification, or the northern 
representatives of the group may have their special characters. 
For the present I place that species in Stephanus, s. str. 

Practically nothing is known of the life-history of these insects. 
They are always found on or around dead or dying wood, and 
are certainly forest dwellers, parasitic on wood-boring larvie. 
Roman (J. c.) suggests that their hosts are to be found among the 
Brenthide, their larvee being elongate and apparently suitable for 
those of slender elongate insects like the Stephanide ; also they 
are both found in all warm countries. Buprestide may also be 
considered, but Brenthidze he thinks most probable. 


762 


2. 
(18.) 1. Posterior margin of head produced eollar-like. 
(18.) 2. Petiole distinctly longer than rest of abdomen. 
(4.) 3. Petiole twice as long as rest of abdomen; pronotum poste- 
riorly rugose punctate; abdomen from second segment 
shining smooth; terebra about 14 times as long as body. 
1. collarifer Schlett. 
(3.) 4. Petiole not more than half as long again as rest of abdomen. 
(6.) 5. Head black; neck indistinctly transrugose, pronotum oblique, 
ADIOS [SING CooyaocooccobososdouSean ese boooHPEbocdo ces  $o TNCUROT OCIS SOME, 
(5.) 6. Head red. 
(8.) 7. Neck transverse, hind coxes smooth beneath ...... 3. glabricoxis Roman. 
(7.) 8. Neck longer than broad; hind coxe trans-striate beneath. 
10.) 9. Neck twice as long as broad; pronotum smooth, diffusely 
punctate behind only, median segment with punctures of 
varied size, interstices partly punctate-rugose... 4. peruanus Enderl. 
(9.) 10. Neck not more than 1} times as long as broad. 
(12.) 11. Neck of pronotum iaterally angulated ; punctures on median 
segment uniform, interstices smooth ............. 5. angulicollis Roman. 
(11.) 12. Neck of pronotum not laterally angulated; interstices on [Cam. 
median segment alutaceous PA See 6. erythrocephalus 
(2.) 13. Petiole at most as long as rest of nbdomens 
(15.) 14. Terebra scarcely longer than body ; median segment centrally 
narrowly, laterally broadly impunctate ......... 7. tener Schlett. 
(14.) 15. Terebra at least one-third longer than body. 
(17.) 16. Petiole as long as rest of dome ; terebra half as long again 
as body .. Re liseesteesee. 8. wtistneit Schlett. 
(16.) 17. Petiole Bionter “dhe est OF aedenieny terebra one-third [Schlett. 
longer than body... sk Be Ron 9. limpidipennis 
(1.) 18. Posterior margin ‘yoedlenall ¢ or nat Jondlonadl but not produced 
collar-like. 
(26.) 19. Posterior margin of head bordered. 
(21.) 20. Terebra twice as long as pee semiannular smooth and 
shining ........ a . 10. macrurus Schlett. 
(20.) 21. Terebra not more hea ele. as sou again as body. 
(23.) 22. Median segment confluently punctate. 
(226.) 22a. Scutellum smooth, with marginal punctures only. 27mm. 
11. intermedius Szépl. 
(22a.) 226. Scutellum closely punctate throughout. 18mm. lla. granulatus Hlhott. 
(22.) 23. Median segment diffusely, not confluently punctate. 
(25.) 24. Semiannular striate; metapleuree coarsely alutaceous and 


MR..E. A. ELLIOTT ON THE 


Table of Species. 


punctate; hind coxe closely rugose; wings hyaline. 
12. pehlkei Enderl. 


. Semiannular smooth; metapleure irregularly rugose ; hind 


coxe with sparse scale-like rugosities, wings subhyaline | Westw. 
centrally slightly darker......................::...... 18. submaculatus 

(19.) 26. Posterior margin of head simple, not bordered. 

(28.) 27. Semiannular and petiole except basally smooth; wings sub- | Westw. 


(31.) 32. 


hyaline, centrally darker ........................... 14. maculipennis 
J b] J 


. Semiannular more or less rugose ; wings not tenn ally darker. 
. Terebra more than half as long again as body; petiole as long 


as rest cf abdomen .......... tessssssscetes. 15. nuficeps Cam: 


. Terebra little or not longer her Dee 
. Terebva a little longer than body; petiole shorter than rest 


of abdomen ; wings clear hyaline .................. 16. vadosus Schlett. 
Terebra only as long as body; petiole slightly shorter than 
rest of abdomen; wings infumate ............... 17. eylindricus Westw. 


HYMENOPTEROUS FAMILY STEPHANID A, 763 


3. 
(6.) 1. Posterior margin of head produced collar-like. 
(3.) 2. Petiole shorter than rest of abdomen; collar short. 7. tener Schlett. 
(2.) 38. Petiole longer than rest of abdomen. 
(5.) 4. Petiole nearly twice as long as rest of abdomen... 1. collarifer Schlett. 
(4.) 5. Petiole a little longer than rest of abdomen; neck Jong and 
Slenderareeetesares ceereteeereee 4 peruanus Emnderl. 


(1.) 6. Posterior margin of head arin ae bordered: 
(8.) 7. Semiannular coarsely, obliquely rugose; 4th tergite finely 
longitudinally parallel striate; wings slightly infumate. 
17. cylindricus Westw. 


(7.) 8. Semiannular quite or nearly smooth. 
(10.) 9. Wings clear hyaline; 4th tergite div aurea eee daa 
striate ......... qOOPR UN . 16. vadosus Schlett. 
(9.) 10. Wings centrally dlandeae. nouadesdose cposcgocooansopccdens, Lal, qaaKenMe/oeInIS 
[ Westw. 


1. COLLARIFER Schlett. 
Stephanus collarifer Schlett., p. 141, 29 3. 


Q@. Frons arcuate to oblique rugose; all five frontal tubercles 
prominent; occiput coarsely longitudinally to obliquely rugose ; 
posterior margin of head reflexed and produced into a long, 
translucent collar. Basal flagellar joints normal. Neck with a 
strong transverse fold, behind which it is shghtly transrugose ; 
semiannular transrugose, laterally finely punctato-rugose. Meso- 
notum centrally smooth, with indistinct longitudinal row of 
punctures and two divergent, superficially punctate impressions, 
remainder transrugose and diffusely punctate. Scutellum smooth, 
with a few isolated but deep marginal punctures. Mesopleurze 
finely alutaceous and shining above, sparsely punctate beneath ; 
metapleurz coarsely reticulate rugose, not separated from the 
median segment by any sulcus. Median segment with diffuse, 
shallow punctures, between which it is finely alutaceous; marginal 
sulci distinct and sometimes an indistinct central impression. 
Petiole transrugose, nearly twice as long as the smooth and 
shining remainder of abdomen, and almost longer than the hind 
cox, trochanters, and femera together. Terebra longer than 
body, its sheaths white-banded before apex. Hind legs with 
cox trans-striate between coarse, scale-like rugosities; femora 
polished smooth, bidentate; tibize compressed to beyond middle ; 
tarsi three-jointed. 

Black; head entirely, mandibles and antenne basally ferru- 
ginous. Wings subhyaline; ‘‘neuration complete, excepting 
that part of the median nervure outside the submedian cell.” 

3. Similar to the 9, but head and thorax more finely sculp- 
tured, median segment closely cribrate punctate ; hind tarsi five- 
jointed. 

Length, 9,18 mm.; terebra 24 mm.; ¢,12 mm. 

Habitat: Brazil (Tefe on the Amazon and San Paulo Prov.) ; 
Surinam. Type in the Royal Nat. Hist. Museum, Berlin ; 
co-type in coll. Wiistnei, in Sonderburg. 


764 MR. BE. A. ELLIOTT ON THE 


This species is distinguished from all others by the extremely 
elongate posterior margin of the head, and by the great com- 
parative length of the petiole, which is twice as long as the rest 
of the abdomen. 


2. MARGINALIS Schlett. 
Stephanus marginalis Schlett., p. 133, 2. 


2. Frons and occiput coarsely and irregularly rugose, a few 
arcuations immediately behind frontal tubercles. Posterior 
margin of head reflexed collar-like. Second flagellar joint three 
times as long as first; third as long as first and second together. 
Neck indistinctly finely rugose, especially centrally; semiannular 
finely obliquely rugose. Mesonotum centrally polished smooth, 
anteriorly and laterally rather coarsely transversely punctato- 
rugose, the median line of punctures obsolete. Scutellum 
polished smooth, with a few very conspicuous punctures. Meso- 
pleure almost smooth, with large, diffuse punctures ; metapleuree 
coarsely reticulate rugose, separated from the median segment 
by a polished smooth suleus, which is broad in front, narrow 
behind. Median segment anteriorly smooth, centrally and 
posteriorly finely transrugose, coarser towards apex, with diffuse 
superficial punctures. Petiole basally distinctly, apically in- 
distinctly transrugose, very little longer than rest of abdomen: 
second segment basally rugose, remainder of abdomen dull. 
Terebra scarcely longer than body, white-banded before apex. 
Hind legs with cox distinctly transrugose between widely 
placed scale-like rugosities ; femora polished smooth, bidentate ; 
tibie compressed to beyond middle. 

Entirely black ; wings slightly infumate. Length 25 mm. 

Habitat: Bahia, Brazil. Type in Imp. Nat. Hist. Museum, 
Berlin. 

This species strongly resembles Stephanus furcatus Lep. & Serv., 
niger Smith, and sickmanni Schlett., from all of which it may be 
distinguished—in addition to the neuration—by the reflexed 
posterior margin of the head and the elongate basal flagellar 
joints. The latter character, and the sculpture of the mesonotum 
will separate it from its nearest allies, H. collarifer Schlett. and 
H. wistnew Schlett. 


3. GLABRICOXIS Roman. 
Stephanus (Hemistephanus) glabricoxis Roman (2), p. 9, 2. 


@. Frons arcuate rugose, space between tubercles longitu- 
dinally rugose; vertex with 2-3 transcarine; occiput more 
longitudinally rugose; posterior margin of head collar-like, 
translucent. Scape as long as second flagellar joint; third joint 
rather longer than first and second together, fifth slightly shorter 
than fourth and as long as sixth. Neck transverse, apically 
deeply emarginate, transverse fold broad and deep; an oblique 
costa runs on each side of the fold, behind which the pronotum 


HYMENOPTEROUS FAMILY STEPHANIDA. 765 


is polished smooth; mesonotum anteriorly and laterally coarsely 
transrugose, posteriorly centrally smooth; impressed part of 
metapleure striate; median segment rugulose, especially laterally, 
the umbilical punctures do not extend to the sides; stigmal sulcus 
obsolete in front. Petiole slender, transrugose, longer than 
remaining segments; second basally transrugose, slightly acicu- 
late beyond middle; third basally finely transreticulate, otherwise 
finely aciculate; fourth transverse, basal half transreticulate, 
apical half polished. Terebra one-third longer than body, its 
black apex more than half as broad as the subapical white band. 
Hind legs with cox polished smooth externally and beneath 
except towards apex, furnished with a few transverse piliferous 
eranules, the usual close striation only visible above and inside; 
femora apically above and externally finely reticulate; tibie 
compressed to beyond middle; metatarsus about four times as 
long as broad. Fore wing not reaching apex of second segment. 

Black; head except apices of mandibles, two basal antennal 
joints, and hind metatarsus red. Wings strongly infumate, 
stigma and nervures black, the former basally pale, radix and 
tegule black. 

Length 18°5 mm.; terebra 24 mm. 

Habitat: Cururuzinho, Rio Autaz, Brazil ; 20.x.1914. One 2 
only. 

Very similar to . angulicollis Roman, differing especially in 
the sculpture of the short neck, the chiefly smooth coxe and 
darker wings, as well as in minor details. 


4, pERUANUS Enderl. 


Hemistephanus peruanus Enderlein (4), p. 302, 9 ; Roman (2), 
DG Qse 


2 3S. Head rather large; frons transrugose, straightly in 
front, arcuately behind. All five frontal tubercles well developed, 
the anterior one produced into a carina in front. Vertex and 
occiput arcuate rugose, posteriorly smooth, without central 
impression; posterior margin of head broad and translucent. 
Second flagellar joint about two and a half times as long as first ; 
third about as long as first and second together. Pronotum 
elongate, polished smooth, with a few punctures on semiannular 
and a strong apical carina. Mesonotum anteriorly with a median 
row of punctures, on each side of which is a smooth space, 
bounded by two divergent rows of punctures, outside of which 
are four strong rugosities, dorsally smooth, with large diffuse 
punctures. Scutellum smooth. |Mesopleuree coarsely and 
diffusely granulate rugose, separated from the median segment 
by a wavy longitudinal carina. Median segment with large and 
small shallow punctures on a partly rugose punctate sround, a 
short longitudinal impression before spiracles, laterally indistinctly 
rugose, anterior lateral angles smooth polished. Petiole basally 
finely transrugose, apex broadly smooth, longer than remainder 


766 MR. BE. A. ELLIOTT ON THE 


of abdomen; second segment with extreme base finely rugose, 
then microscopically longitudinally aciculate: third to beyond 
middle and fourth basally finely shagreened and dull, remainder 
smooth. Terebra rather longer than body, white-banded before 
apex. Hind legs with coxe finely transrugose throughout ; 
femora bidentate smooth. 

Black; head ara & 3-4 basal antennal joints ferruginous, thorax 
basally dark red-brown. ‘Tarsi densely gold pubescent beneath. 
Wings strongly infumate, nervures black-brown, stigma slende 

Length, 9, 194 mm.; abdomen 133 mm. ; petiole 74 mm. 
terebra 214 mm.; fore wing 1S am Ss", 91-14 mm. 

Habitat : Proy. Chanchamayo (Rio Toro}, Peru; Rio Autaz and 
Manaos, Brazil. 

This species is best known from all others by the elongate, 
unsculptured pronotum. It resembles Parastephanellus collarifer 
Schlett., from which it may be distinguished by neuration of 
wings, sculpture of median segment and abdomen, and by the 
colour. 


5. ANGULICOLLIS Roman, 
Stephanus (Hemisteph.) angulicollis Roman (2), p. 7, 2. 


@. Frons arcuate rugose, space between the stout anterior 
tubercles longitudinally rugose, the posterior pair squamiform ; 
vertex with 3-4 transcarine; occiput anteriorly more or less 
arcuately rugose, remainder smooth, centrally longitudinally 
aciculate, the rugose and smooth parts forming an obtuse angle 
with each other. Posterior margin of head produced into a 
translucent collar. Scape as jong as second flagellar joint, third 
rather longer than first and second together, fifth as long as 
fourth and rather longer than sixth. Neck elongate, about one 
and a half times as long as broad; in front of the transverse fold 
is a broad sulcus, on each side of which isa strong raised costa 
running forwards and suddenly bent outwards, followed by others 
which become smaller in the direction of the fold; semiannular 
coarsely rugose before and below its polished posterior margin. 
Mesonotum with distinct central row of punctures, parapside 
coarsely subtransversely rugose; impressed part of metapleuree 
striate. Median segment with umbilicate punctures of about 
even size ona smooth ground, lateral impressions from spiracles 
to anterior margin. Nodemen more than twice as long as head 
and thorax, petiole slender, strongly trans-striate throughout, 
longer than the remaining segments together; extreme base of 
segment two transrugose, remainder polished and closely acieu- 
late; third slightly longer than its apical breadth, basally finely 
transreticulate, rest aciculate, with broadly smooth apical margin ; 
3rd to 6th slightly apically emarginate; fourth not longer than 
its basal breadth, basal third tr ansreticulate, centrally transacicu- 
late, apically polished ; remaining segments strongly transverse. 
Terebra about one-third longer than “pody, the black apex about 


HYMENOPTEROUS FAMILY STEPHANIDE. 767 


half as broad as the subapical white band. Hind legs with coxee 
closely trans-striate, subobsoletely above, a few  piliferous 
granules. Femora towards apex above and externally very finely 
reticulate; the tibiz compressed to beyond middle; metatarsus 
five times longer than its maximum breadth. Fore wings not 
reaching apex of second segment. 

Black; head except apices of mandibles, scape, first flagellar 
joint partly, and hind metatarsus obscurely red; palpi fuscous. 
Wings nearly hyaline; stigma black, basally narrowly pale; 
nervures and radix black, tegulee pale, basally black. 

Length 17°5 mm.; terebra 22 mm. 

Habitat: Apipica, Rio Autaz, Brazil, 14.1x.1914. One ©. 

Apparently isolated by the peculiar structure of the neck, but 
near /7. peruanus Enderl. in the elongate neck and small trans- 
verse fold. 


6. ERYTHROCEPHALUS Cam. 


Megischus erythrocephalus Cameron (1),p.421, 2. Stephanus e. 
Schlett., p. 140, 9. 


2. Frons transrugose; occiput longitudinally rugose, with 
faint central sulcus ; posterior margin of head broadly bordered. 
Neck longitudinally sulcate, laterally obliquely carimate; semi- 
annular trans-striate, posterior margin narrowly smooth. Meso- 
thorax trans-striate throughout; scutellum smooth, with three 
large marginal punctures on each side; meso- and meta-pleure 
and median segment alutaceous, with diffuse large punctures, the 
two latter separated by a sulcus. Petiole trans-striate, longer 
than rest of abdomen; second segment basally coarsely trans- 
rugose, thence and basal two-thirds of third discally longitudi- 
nally aciculate and dull, remaining segments smooth. Terebra 
slightly longer than body, sheaths white-banded before apex. 
Hind legs with coxe coarsely trans-striate; femora~- smooth, 
bidentate ; tibize constricted in basal third, apicai third felted 
beneath. 

Black; head red. Wings hyaline, stigma and nervures black. 

Length 19 mm.; abdomen 13 mm.; petiole 7 mm.; terebra 
20 mm. 

Habitat: Buguba, Panama. Type in British Museum. 

Closely allied to 4. collarifer Schlett., but sufficiently distin- 
guished by the less developed posterior margin of head and 
shorter terebra. 


7. TENER Schlett. 
Stephanus tener Schlett., p. 142, 9 ; Roman (2), p. 7, 2 ¢. 


@. Frons finely arcuate rugose; occiput finely longitudinally 
to obliquely rugose, more transversely near posterior ocelli; 
the three anterior frontal tubercles very prominent; posterior 
margin of head produced into a short but distinct and trans- 
lucent collar ; cheeks scarcely as long as scape. Second flagellar 


768 MR, BE. A. ELLIOTT ON THE 


joint twice as long as first, third longer than first and second 
together. Neck elongate and very slender, posteriorly rugose, 
especially laterally ; semiannular polished smooth, with a few 
fine punctures and oblique striations. Mesonotum laterally 
deeply punctato-rugose, with a central line of small but distinct 
punctures, on each side of which is a smooth space bounded by a 
row of broad punctures. Scutellum smooth, with a few con- 
spicuous marginal punctures. Mesopleure rather smooth and 
shining above, dull beneath, with diffuse and shallow punctures ; 
metapleure anteriorly finely and irregularly, posteriorly coarsely 
reticulate rugose, not separated by any sulcus from the median 
segment, which is punctate, except narrowly in the middle and 
broadly at the sides. Petiole transrugose, as long as remainder 
of abdomen; second segment smooth, basally rugose ; third and 
fourth finely wavily transrugose, third centrally feebly longitu- 
dinally aciculate (in small individuals nearly smooth). Terebra 
as long as body, white-banded before apex. Hind legs with 
coxe extremely finely transrugose between coarse, scale-like 
rugosities ; femora polished smooth, bidentate; tibize compressed 
to beyond middle. 

Black ; head, scape, first flagellar joint, and mandibles basally 
ferruginous. Wings byaline. 

Length 93-145 mm. The type in the Hungarian National 
Museum at Budapest is 13 mm. long. 

Habitat: type from Obidos on the Amazon, Brazil; Rio 
Autaz. 

3. Similar to ¢, but segment 4 subtransverse. Length 
9-10°5 mm. 

Habitat: Rio Autaz, Amazon District, Brazil; taken by 
Roman in 1914-15. 

The elongate neck and shorter terebra will Suites to distinguish 
this species from H. collar fer, limpidipennis, and wiistneit Schlett., 
in all of which the neck is normal and the terebra longer than the 


body. 


8. wistNEtI Schlett. 


Stephanus wiistneti Schlett., p. 149, @. 

@. Frons irregularly rugose; vertex transrugose; occiput 
rugose-punctate. Cheeks shorter than scape. Posterior margin 
of head produced into a very distinct collar. Basal flagellar 
joints normal. Neck with a few deep rugosities ; semiannular 
polished smooth, with diffuse punctures. Mesonotum with a 
central row of punctures, on each side of which is a smooth 
space, laterally rather densely rugose-punctate. Scutellum 
smooth, with a few fine marginal punctures. Mesopleurz almost 
smooth and slightly shining above, dull and rather coarsely and 
densely rugose-punctate beneath; metapleure reticulate rugose, 
more coarsely behind than in front, not separated from the 
median segment by any suleus, Median segment densely 


HYMENOPTEROUS FAMILY STEPHANIDA, 769 


cribrate punctate, with a distinct crenulate suleus on each side. 
Petiole very finely and indistinctly trans-striate, as long as the 
remaining segments ; second segment basally striate, rest smooth 
and shining, with a few dull spots due to microscopic sculpture. 
Terebra half as long again as body, sheaths white-banded before 
apex. Hind legs with coxe finely striated between coarse 
rugosities, distinctly transrugose beneath; femora polished 
smooth, bidentate ; tibiz compressed to beyond middle. 

Black ; head, mandibles basally, and scape ferruginous. Wings 
hyaline. 

Length 12 mm.; terebra 18 mm. 

Habitat: Massanary, on the Amazon, Brazil. ‘Type in collec- 
tion of Herr Wiistnei, after whom it is named. 

Schletterer states that this species is deceptively like 1. limpidt- 
pennis, but the sculpture of face is different, petiole and terebra 
longer, the former less distinctly sculptured and the median 
segment more densely punctate. From /. marginalis it may be 
distinguished by smaller size, shorter basal flagellar joints, shorter 
petiole, and shorter terebra. 

Szépligetti, p. 533, states briefly of this species:— ¢. Tarsi 
A-jointed; metatarsus brown. Q. Length 14 mm.; terebra 
18mm. Habitat: Fonteboa, on the Amazon, Brazil. 

This error as to the ¢ ¢ having 4-jointed hind tarsi is 
repeated by him in other cases; they are always 5-jointed. 


9. LIMPIDIPENNIS Schlett. 
Stephanus limpidipeninis Schlett., p. 147, @. 


®. Frons rather coarsely and subarcuately rugose; vertex 
transrugose; occiput more coarsely punctato-rugose. Temples 
with a few punctures near eyes. Posterior margin of head pre- 
duced into a long translucent collar. Cheeks decidedly shorter 
than scape. Basal flagellar joints normal. Neck of prothorax 
with two coarse transverse folds, between which it is deeply 
impressed and shining, behind them lightly punctato-rugose ; 
- semiannular rather densely, partly confluently punctate. Scutellum 
entirely smooth with a few marginal punctures. Mesonotum 
laterally rather coarsely and densely rugoso-punctate. Meso- 
pleurz smooth above, dull beneath, with large diffuse punctures ; 
metapleure in front lightly and indistinctly, behind coarsely and 
deeply reticulate rugose, not separated from the median segment. 
Median segment shining smooth, with large diffuse punctures, 
centrally closer, marginal sulci distinct. Petiole trans-striate, 
basally irregularly rugose, distinctly shorter than remainder of 
abdomen, which is basally finely rugose, otherwise smooth with a 
few dull spots due to microscopic sculpture. Terebra evidently 
longer than body, white-handed before apex. Hind legs with 
ooxe finely trans-striate between very coarse rugosities; femora 
smooth, bidentate; tibiz compressed to beyond middle. 


770 MR. E. A, ELLIOTT ON THE 


Black; head, base of mandibles, and scape ferruginous. Wings 
extremely clear hyaline. 

Length 14-15 mm. ; terebra 19-20 mm. 

Habitat: Massanary, Brazil. Typein Hungarian Nat. Museum, 
Budapest. 

This species differs from HH, wiistneii chiefly in the more 
arcuate rugose head, shorter and more coarsely sculptured petiole, 
and more diffusely punctate median segment. From H. collarifer, 
with which it agrees in size, colour, posterior margin of head, and 
length of terebra, it may be best known by the rugose- punctate 
occiput, shorter petiole, deeper sculpture of the semiannular, and 
by the very clear wings. 


10. macrurus Schlett. 
Stephanus macrurus Schlett., p. 128, 9. 


@. Frons rather finely arcuate 1ugose; occiput in front 
arcuately to transversely rugose. Posterior margin of head 
strongly bordered. Cheeks shorter than scape. Basal flagellar 
joints normal. Neck of prothorax apically finely transrugose, 
the remainder, together with the whole semiannular, smooth. and 
shining. Mesonotum sparsely punctate, centrally smooth, with a 
longitudinal row of broad punctures, laterally finely transrugose. 
Seutellum polished smooth, diffusely and finely punctate. Meso- 
pleurze above finely plutaceous aud shining, beneath dull, with 
large and partly confluent punctures ; metapleuree dull, finely 
and diffusely punctate, separated from median segment by an 
indistinct and finely rugose sulcus. Median segment with diffuse 
large punctures, interspersed with finer ones. —° Petiole trans- 
rugose, basally more coarsely, a little shorter than rest of 
abdomen, which is basally rugose, then shining, with a few dull 
spots. ‘Lerebra twice as long as body, sheaths white-banded 
before apex. Hind legs with coxe shining, finely trans-striate ; 
femora polished smooth, apically finely sculptured and subopaque, 
bidentate ; tibiz longer than femora, compressed not quite to 
middle. 

Black; head ferruginous, mandibles red, apically brown ; 
antennee pitch-brown, apically ferruginous; neck of prothorax 
brown or rufous; hind tarsi rufescent. Wings hyaline. 

Length 23-24 mm. 

Habitat: Surinam. Type in Royal Natural History Museum, 
Berlin. 

This species may be distinguished from H. maculipennis 
Westw. by the s sculpture of petiole and hind coxe, longer terebra, 
and hyaline wings. From H. collarifer Schlett. and limpidi- 
pennis Schlett., which it resembles in form and colour, it may at 
once be known by the much less prominent posterior margin of 
head and by the longer terebra. 


HYMENOPTEROUS FAMILY STEPHANID®. Wik 


11. INTERMEDIUS Szepl. 
Stephanus intermedius Szépl., p. 533, Q. 


@. Frons coarsely and indistinctly arcuately rugose; vertex 
arcuately, occiput transversely rugose ; cheeks short, only half as 
long as scape; posterior margin of head sharply bordered. Basal 
flagellar joints normal. Pronotum short, irregularly trans-striate, 
its posterior margin polished. Mesonotum coarsely and irregularly 
rugose; scutellum smooth, with coarse marginal punctures. 
Mesopleurz in front finely punctate, a few coarse punctures 
behind and transrugose below; metapleure pubescent, with a 
few isolated punctures, separated by a deep smooth sulcus from 
the confluently punctate median segment. The petiole trans- 
rugose, rather shorter than rest of abdomen. ‘Terebra rather 
longer than body, white-banded before apex. Hind legs with 
cox trans-striate; femora smooth, bidentate; tibiz compressed 
in basal third. 

Black; head, scape, and hind tarsi ferruginous. Wings 
infumate, light brown. 

Length 27 mm.; terebra 30 mm. 

Habitat: Tonantins, Brazil. 

This species comes very close to H. macrurus Schlett., from 
which it may be distinguished by the confluent puncturation of 
the median segment, infumate wings, and much shorter terebra. 


Illa. GRANULATUS Elhott. 
Hemistephanus granulatus Elliott (2), p. 130, 2. 


@. Head, excepting the smooth temples, granulate rugose, 
vertex and occiput more finely; posterior margin of head 
bordered; anterior tubercle prominent; three very conspicuous 
carine between the posterior ocelli. Basal flagellar joints normal. 
Pronotum rather short, trans-striate, with smooth posterior 
margin; mesonotum centrally trans-striate, laterally punctate ; 
scutellum closely punctate throughout, marginal punctures deep, 
Mesopleure finely and closely punctate; metapleure coarsely 
punctate, separated by a crenulate sulcus from the median 
segment, which is confluently, almost granulate punctate. Petiole 
trans-striate, slightly shorter than rest of abdomen; base of 
second segment trans-striate, remainder smooth, feebly shining. 
Terebra very nearly as long as body, rufescent. Hind legs with 
cox trans-striate ; femora smooth, bidentate ; tibize compressed 
to middle. 

Rufescent ; all metatarsi flavous to white. Wings infumate. 

Length 13 mm.; abdomen 74 mm.; petiole 3}mm.; terebra 
133 mm. 

Habitat: Swan River, West Australia. 

The closely punctate scutellum and general granulate sculpture 
will distinguish this species. 

If the label “Swan River” is correct, this is the first, and 
Proc. Zoou. Soc.—1922, No. LIT, 52 


772 MR, E. A, ELLIOTT ON THE 


up to the present, the only species of this subgenus known 
from the Australian Region. The large size of the discoidal 
cell places it here, but all other known examples come from 
South America. 


12. praLKet Enderl, 


Hemistephanus pehlkei Enderl. (4), p. 304; Roman (2), pp. 10 
ep 13,79). 


2. Frons closely, posteriorly arcuately rugose; space between 
tubercles transrugose in front, obliquely behind. Vertex strongly 
arcuate rugose; occiput in front arcuate rugose, becoming finer 
and straighter behind, with a broad, indistinct median impression ; 
posterior margin of head finely bordered. Second flagellar joint 
normal, third longer than first and second together. Neck 
rather short, with four transverse carine; semiannular 
moderately transrugose, the narrow posterior border smooth, and 
with lateral very fine oblique striation. Mesonotum anteriorly 
irregularly punctate rugose, with a median line of confluent 
punctures, centrally smooth, laterally coarsely punctate. 
Scutellum polished smooth, laterally coarsely punctate. Meso- 
pleuree rugose reticulate and diffusely punctate; metapleurz 
similarly, but more coarsely punctate, separated from median 
segment by an anteriorly obsolete carina. Median segment 
smooth, with microscopic reticulation and diffuse punctures which 
do not touch the middle or sides; marginal sulci wanting. 
Petiole trans-striate, apically more finely, shorter than rest of 
abdomen, Second segment basally rugose, towards apex longitu- 
dinally aciculate; third to fifth basally granulate aciculate ; 
fourth apically longitudinally aciculate, remainder very finely 
aciculate. Terebra one-third longer than body, sheaths black, a 
subapieal white band as broad as the black apex. Hind coxee 
transrugose ; hind femora smooth, bidentate. 

Black; head except apices of mandibles and 3 or 4 basal antennal 
joints ferruginous; hind tarsi gold pubescent beneath. Wings 
clear hyaline; stigma black, nervures black-brown. 

Length (type) 203 mm.; abdomen 143 mm.; petiole 6 mm. 
terebra 28 mm.; fore wing 103 mm. (Enderlein). 15-204 mm, 
(Roman). 

Habitat: Rio Magdalena, Colombia (Pehlke); Rio Autaz and 
Curururuzino, Amazon District, Brazil (Roman). 

Roman states that this species is specially distinguished by 
the extremely fine oblique striation on the semiannular and at 
the lower end of the pleural furrow, also by the divergent longi- 
tudinal striation en the fourth tergite. The finely bordered 
posterior margin of the head and the carina between metapleure 


and median segment are characteristic, also the whole sculpture 
of the abdomen, 


HYMENOPTEROUS FAMILY STEPHANIDA. es 


13. sUBMACULATUS Westw. 


Megischus submaculatus Westw. (4), p. 230. 2, Stephanus s. 
Schlett., p. 126, 2. : 

Q. Frons arcuate rugose;. occiput anteriorly arcuately, 
posteriorly transversely rugose. Posterior margin of head very 
finely bordered. Second flagellar joint one and a half times as 
long as first, third as long as first and second together. Neck 
very finely trans-striate; semiannular entirely smooth. Meso- 
notum very diffusely and finely punctate; scutellum smooth, with 
a few marginal punctures. Mesopleure shining smooth above, 
diffusely punctate below; metapleure irregularly rugose, 
separated by a narrow but distinct suleus from the median 
segment, which is very diffusely punctate, apically irregularly 
transrugose. Petiole trans-striate, a little shorter than rest of 
abdomen, with two very distinct lateral tubercles before middle ; 
remaining segments smooth, apex of abdomen duli. Terebra 
longer than body, sheaths white-banded before apex. Hind cox 
with diffuse scale-like rugosities ; hind femora polished, bidentate; 
hind tibiz compressed to middle. 

Black; head and scape red. Wings subhyaline, scarcely darker 


centrally, 
Length 18 mm.; terebra 33 mm. 


Habitat: Para, Brazil. Type in British Museum, 

Very similar to maculipennis Westw., but differs in the almost 
smooth neck of pronotum, more diffuse puncturation of mesonotum 
and median segment, trans-striate petiole with much larger 
tubercles, and the subhyaline wings, not darker centrally. 


14. MACULIPENNIS Westw. 

Megischus maculipennis Westw. (4), p. 229, d. Stephanus m. 
Sehlett., p. 127, ¢ ; Szépl., p. 533, 9 o. 

2 3. Frons and vertex arcuate rugose; occiput regularly, 
laterally irregularly transrugose. Posterior margin of head 
simple. Second flagellar joint one and a half times as long as 
first; third normal. Neck very coarsely, basally more finely 
transrugose ; semiannular polished smooth. Mesonotum coarsely 
and diffusely, laterally more densely and partly confluently 
punctate, the median row of punctures and lateral divergent 
impressions very distinct. Scutellum smooth, with a few con- 
spicuous marginal punctures. Mesopleurz above shining smooth, 
below finely and diffusely punctate; metapleurz coarsely and 
irregularly rugose, separated by a narrow but distinct sulcus 
from the median segment, which is coarsely and diffusely 
punctate, apically transrugose. Petiole smooth, basally finely 
trans-striate, with a small tubercle on each side before the 
middle, shorter than rest of abdomen, which is subnitidulous, 
apically dull. Hind coxe with diffuse coarse rugosities, only 
near posterior margin densely transrugose ; hind femora polished 
smooth, bidentate ; hind tibize longer than femora and compressed 
to middle. 

D2 


od 


174 MR. E. A. ELLIOTT ON THE 


Black; head entirely, frons and hind legs centrally ferru- 
ginous; mandibles and antenne basally rufescent. Wings 
subhyaline, centrally strongly infumate. 

Length 25-33 mm.; terebra 35-50 mm. 

Habitat: Para, Brazil. Type of 2 in British Museum. 

Szépligetti (J. c.) states:—“ 2. Length 25-30 mm.; terebra 
35-50 mm.; sheaths white-banded before apex; posterior margin 
of head sharp; petiole transrugose; hind tarsi 3-jointed ; 
otherwise as ¢. Yurimaquas, Peru, and Prov. Piauchy, Brazil. 


Text-figure 4. 


Hemistephanus maculipennis. 


“Var. 1. ©. Hind tibie entirely black. La Merced, Brazil. 

“Var. 2. 9 3. Median segment transrugose. centrally punc- 
tate. Wings pale brown to hyaline, centrally dark marked. Only 
basal half of front legs more or less red. Petiole of 2 rather 
lightly transrugose. Length 33 mm.; terebra 50mm. _ Vilca- 
nota, Peru.” 

In the type-form the smooth petiole, centrally dark wings, and 
sculpture of prothorax will distinguish this species from sub- 
maculatus Westw. and macrurus Schlett., both of which it 
resembles. The transrugose petiole of Szépligetti’s specimens 
appears to be merely a variety. 


15. RUFICEPS Cam. 


Megischus ruficeps, Cameron (1), p. 420, pl. 18, fig. 9, 9. 
Stephanus capitatus Schlett., p.151, 9. Hemistephanus ruficeps 
Morley (1), p. 112, @. 


HYMENOPLEROUS FAMILY STEPHANIDA. 775 


“ Niger, capite rufo; alis fere hyalinis, 9 long. 19 mm. ; 
terebra 25 mm. Hab. Panama, Volcan de Chirique, 2500-4000 
feet.” (Cameron.) 

2. Head coarsely rugose, vertex more finely; posterior 
margin of head simple. Antenne very slender, about two- 
thirds of length of body, scape elongate; basal flagellar joints 
normal. Neck coarsely, semiannular more finely trans-striate. 
Scutellum smooth, with large and coarse marginal punctures. 
Mesopleure aciculate, rather dull; metapleure rugose and dull. 
Median segment nitidulous, with large and diffuse punctures. 
Petiole irregularly but closely trans-striate, as long as remainder 
of abdomen. ‘Terebra half as long again as body. Hind legs 
with coxe striolate, more strongly towards apex, inner side 
densely pilose; femora smooth, bidentate ; tibie and tarsi 
covered with dense, shining, fulvous pubescence. 

Black with red head. Wings subhyaline. 

Length 19-21 mm.; terebra 25-35 mm. 

Habitat: Panama (Cameron); Amazon (Bates). Type in British 
Museum. 


The present species may be best separated from cylindricus 
Westw. by the much longer terebra and by the sculpture of the 
pro- and meso-thorax and of the hind coxe. It also resembles 
H, wiisineri Schlett., from which it may be known by the simple 
posterior margin of the head, coarser puncturation of the 
scutellum, and by the greater size. : 

In 1861, Smith described a Fenatopus ruficeps, and subse- 
quently withdrew his genus Yanatopus as synonymous with 
Stephanus, On these grounds Schletterer (/.c.) altered the 
specific name to capitatus. More recent investigation has 
resulted in the original genus Stephanus being divided into a 
number of subgenera, in each of which it appears permissible to 
repeat a specific name; hence I restore the original name 
ruficeps given by Cameron. 


16. vaposus Schlett. 
Stephanus vadosus Schlett., p. 146, 2 ; Roman (2), p. 11, 2d. 


Q. Head irregularly rugose, frons finely, occiput more 
coarsely and transversely in front; posterior margin of head 
simple. Second flagellar joint barely twice as long as first ; 
third longer than first and second together. Neck feebly trans- 
rugose, semiannular polished smooth above, obsoletely rugose 
below. Mesonotum with irregular to transverse superficial 
rugosity, central row of punctures distinct, lateral ones indistinct. 
Scutellum polished smooth, marginally punctate. Mesopleure 
shining above, dull beneath; metapleure irregularly rugose, 
not separated from median segment, which is smooth shining, 
very diffusely punctate, with distinct marginal sulci. Petiole very 
finely trans-striate, little shorter than rest of abdomen; second 
seement basally finely rugose, apical third longitudinally striate, 
fourth longitudinally divergently striate. Terebra longer than 


776 MR. E. A. ELLIOT ON tHE 


body, white-banded before apex. Hind legs with coxe finely 
trans-striate between coarse ridges; femora smooth, bidentate ; 
tibiz compressed about to middle. 

Black; mandibles and antenne basally and hind metatarsus 
ferruginous; anterior legs brown, with a tendency towards 
rufescent. Wings entirely hyaline. 

Length 12-20 mm.; terebra 15-24 mm. 

6. Differs from 2 in having 3rd abdominal segment usually 
distinctly elongate, 4th transverse or rarely quadrate. Length 
8-11°5 mm. 

Habitat: Brazil, Manaos, Rio Autaz. Type in National 
Museum, Budapest. 

This species appears to be one of the most frequent in the 
Amazon region, yet, until 1914, it had been only once taken, and 
the ¢ was not known. It comes close to H/. cylindricus Westw., 
from which it may be distinguished chiefly by the finer sculpture 
of pro- and meso-thorax, longer terebra, and entirely hyaline 
wings. From //. limpidipennis Schlett., wiistnreii Schlett., and 
tener Schlett., all of which it resembles, it is at once separated by 
the simple posterior margin of the head. 


‘17. eyLinpRicus Westw. 

Megischus cylindricus Westw. (4), p. 230. Stephanus cylin- 
dricus Schlett., p. 144; Enderlein (4), p. 306, 9 ; Roman (2), 
Wo Oy Qos 

@. Head entirely finely and moderately irregularly rugose, 
more transversely near posterior ocelli; three anterior frontal 
tubercles very prominent; posterior margin of head simple. 
Second flagellar joint fully one and a half times as long as first ; 
third longer than first and second together. Neck coarsely 
rugose, anteriorly transversely, posteriorly more irregularly ; 
semlannular smooth above, laterally posteriorly obliquely rugose. 
Mesonotum rather coarsely rugose-punctate, a small central 
smooth space with indistinct longitudinal row of punctures, the 
lateral ones more distinct. Scutellum polished, with diffuse 
marginal punctures. Mesopleuree shining smooth above, dull 
and diffusely punctate beneath; metapleure finely and diffusely 
punctate in front, irregularly rugose behind. Median segment 
strongly shining, very finely and diffusely punctate, marginal 
sulci obsolete. Petiole basally irregularly rugose, remainder 
distinctly trans-striate, a little shorter than rest of abdomen, also 
slightly shorter than the hind cox, trochanters, and femora 
together ; second segment basally subrugose, third basally trans- 
versely, apically longitudinally striate, rest smooth, with a few 
dull spots due to microscopic sculpture. Terebra as long as 
body, sheaths white-banded before apex. Hind legs with coxe 
finely striate between coarse rugosities; femora smooth, 
bidentate; tibiee compressed to middle. 

Black; head entirely, mandibles and antenne basally ferru- 
ginous, hind tarsi badious. Wings slightly infuscate. 

Length 13-15 mm. 


HYMENOPTEROUS FAMILY STEPHANIDA. 777 
3. Differs from 9 in the stouter petiole, third segment not, 
or scarcely longer than its basal width, fourth strongly transverse 
(subelongate in @), its extreme base longitudinally striate. 
Length 19-14°5 mm. 

Habitat: Rio Autaz, Apipica, Cururuzinho, Brazil; Peru, Dept. 
Chanchamayo. 

This species seems by no means plentiful. It resembles 
H. vadosus Schlett., from which it may be known by the coarser 
sculpture of pro- andi meso: -thorax, shorter terebra, and colour of 
head. The simple posterior margin of the head and longer third 
flagellar joint will separate it from H. limpidipennis, tener, and 
wustnew Schlett. 


NEOSTEPHANUS. 
Kieffer (1), p. 4, gen. n. 


Smith’s genus Kenatopus was defined as having the cubital 
and discoidal cells wanting, or only indicated by indistinct 
nervures. 

Kieffer (/. ¢.) subdivided this genus, placing in a new subgenus, 
Neostephanus, those species in which the apical margin of the 
external submedian cell is marked by a transverse nervure, while 
retaining in Yenatopus those in which this nervure is absent. 

The species of the subgenus Veostephanus appear to be confined 
to Central and South Africa and Madagascar, to which latter 
island the type-species, V. alluaudz Wieff., belongs. 


Table of Species. 


(2.) 1. Hind femora tridentate; frons irregularly rugose; vertex 
with four trans-carin®@ ............c0ce cee lL. alluaudi Kietf. 
(1.) 2. Hind femora bidentate. 
) 


(4.) 3. Temples rugose and dull except shining raised space near 
eyes; pro- and meso-notum closely rugose ...... 2. insignis Schlett. 
(3.) 4. Temples smooth and shining. 
(6.) 5. Head large and globose; frons arcuate punctate... 38. globiceps Endevl. 
(5.) 6. Head ordinary; frons not arcuate punctate. 
(10.) 7. Frons rugose. 
(9.) 8. Frons obliquely rugose; median segment diffusely punctate. 
4. camerunus Enderl. 
(8.) 9. Frons subarecuate rugose; median segment coarsely and 


partly confluently punctate ........................ 5. erassipes Bischoff 
(7.) 10. Frons reticulate. 


(12.) 11. Vertex with one transcarina; 2nd and 3rd flagellar joints 
equal; mesopleurze smooth; black, with white-marked 
head and abdomen ......... Levees. 6, albomaculatusCam. 


(11.) 12. Vertex bicarinate; 3rd fagellar aan as long as first and 
second together ; mesopleure granulose and dull; red- ‘ 
brown, abdomen *yellow-marked secssseectersecaee 7. penthert Kieff. 


1. aLLUAUDI Kieff. 
Neostephanus alluaudi Kieffer (1), p. 1, 2. 


@. Face and frons irregularly rugose, becoming arcuate 
towards the frontal tubercles, the space between which is not 


778 Mi. E. A. ELLIOTT ON THE 


impressed; vertex with four transcarine; occiput finely and 
densely trans-striate. Posterior margin of head simple. Second 
flagellar joint about one and a half times as long as first; third as 
long as first and second together. Neck transcarinate, semi- 
annular finely and densely trans-striate, posterior margin smooth 
and shining. Mesonotum irregularly rugose, with three longitu- 
dinal impressions; scutellum shining sanouiih, with afew amare ier 
punctures, the lobes separated by rows of br oad punctures. Median 
segment finely shagreened, shining and diffusely punctate. Petiole 
finely trans-striate, longer than rest of abdomen. Terebra shorter 
than body, sheaths rufescent, apically black. Hind legs with 
coxe as long as trochanters and femora together, strongly trans- 
rugose ; femora very finely shagreened, tridentatel: tibiee con- 
stricted beyond the middle, covered internally with dense erect 
red pubescence; tarsi densely pubescent beneath, metatarsus a 
listle longer than the following joints together. 

Black ; head except apices of mandibles, basal third of antenne, 
the front coxe and legs, inflated part of hind tibie and all tarsi 
red; apical third of antenne and the intermediate legs brown 
Wings subhyaline. 

Length 15 mm.; abdomen 9 mm.; terebra 12 mm. 

Habitat: Madagascar, south of the Bay of Antongil and St. 
Marie de Madagascar. Three specimens taken by M. Ch. 
Alluaud. 

(? Gold Coast, N. Territories, Yahi. Dr. J. J. Simpson, x1. 1916). 

It is unfortunate that the type of the genus is the only one yet 
known with tridentate hind femora. 


2. INSIGNIS Schlett. 
Stephanus insignis Schlett., p. 100, 9. 


@. Head rather coarsely and irregularly rugose throughout. 
Temples subrugose and dull, except a raised smooth and shining 
space near the ocelli. Posterior margin of head simple. Cheeks 
longer than scape. Second flagellar joint twice as long as first ; 
third shorter than first and second together. Neck rather coarsely 
irregularly to transversely rugose; semiannular coarsely irregularly 
rugose. Mesonotum rather coarsely and irregularly punctato- 
rugose. Central section of scutellum anteriorly dull; behind this 
a small smooth polished space, laterally diffusely punctate, lateral 
lobes rather coarsely and densely punctate. Mesopleure polished 
smooth above, laterally rugose with rather dense indistinct 
puncturation; metapleuree coarsely and irregularly rugose, 
separated by a smooth shining sulcus from the median segment, 
which is somewhat densely punctate. Petiole as long as, or 
slightly longer than rest of abdomen, trans-striate, remainder of 
abdomen smooth and shining. Terebra scarcely as long as body, 
spicula red, the sheaths ferruginous, apically black. Hind legs 
with coxee and femora much incrassate, former coarsely and 
irregularly rugose, more transversely towards apex ; femora finely 


HYMENOPTEROUS FAMILY STEPHANIDA. 779 


rugose-punctate and dull, bidentate; tibiz constricted to beyond 
middle. 

Black, with a tendeney towards red; cheeks pale marked, 
mandibles and antenne basally and frontal tubercles rufescent, a 
curved red line round anterior ocellus and all legs more or less red. 
Wings entirely hyaline. 

Length 14 mm. 

Habitat: South Africa. Type in Nat. Hist. Mus., Berlin. 

This species may be best known by the rugose and dull temples, 
the dense and irregular sculpture of pro- and meso-notum, dense 
puncturation of scutellum, and median segment and red sheaths 
of terebra. 


3. GLOBICEPS Enderl. 
Stephanus globiceps Enderl. (1), p. 200, 2. 


Q. Head large and globose, its longitudinal axis very elon- 
gate, temples and occiput very long and convex. Frons arcuate 
punctate; all five frontal tubercles distinct ; vertex coarsely trans- 
rugose; occiput finely arcuate rugose, with distinct longitudinal 
impression ; posterior margin of head sharply bordered. Basal 
flagellar joints normal. Neck elongate, smooth, laterally obliquely 
rugose; semiannular trans-striate, posterior margin smooth. 
Mesonotum strongly transrugose and punctate. Scutellum cen- 
trally polished, laterally strongly punctate.. Mesopleurze pubescent, 
smooth, apical half diffusely punctate; metapleure alutaceous and 
dull in front, coarsely punctate behind, separated from median 
segment by a row of punctures. Median segment punctate, 
apically more densely. Abdomen wanting in the unique specimen. 
Hind coxee finely trans-striate, their femora elongate, tarsi three- 
jointed. 

Black; head ferruginous yellow; apex of mandibles and a 
transverse band between eyes black; five basal antennal joints, 
anterior legs, apices of hind femora with their tibie and tarsi 
ferruginous. Wings hyaline, very faintly infuscate; stigma and 
nervures dark brown. 

Length cirea 14 mm., the proportions of the remaining parts 
being about as in WV. insignis Schlett. 

Habitat: Johann Albrechtshohe, N.Cameroons. March 1896; 
L. Conradt. © 

The shape, sculpture, and colour of head appear distinctive. 


4, CAMERUNUS Enderl. 
Neostephanus camerunus Enderl. (4), p. 299, 2 3. 


2 gd. Head medium; frons coarsely obliquely rugose, more 
transversely in front, almost circularly between the tubercles ; 
vertex strongly, occiput less strongly arcuate rugose, with distinet 
posterior median impression. Posterior margin of head bordered. 
Basal flagellar joints normal. Neck short, coarsely obliquely 
rugose; semiannular smooth, posterior margin polished. Meso- 


780 MR. E. A. ELLIOTT ON THE 


notum apically punctato-rugose, dorsally alutaceous between deep 
but diffuse punctures; scutellum polished smooth, laterally . 
diffusely punctate, a row of about 10 subquadrate punctures on 
each side between it and the dorsum. Mesopleure with small 
and diffuse punctures and microscopic shagreening; metapleuree 
widely reticulate, smooth above, separated from median segment 
by a carina, which passes into a row of punctures above in front. 
Median segment with diffuse, sharply-detined punctures, anterior 
lateral angles nearly smooth, apex rugose-punctate round the 
insertion of petiole. Petiole slender, finely trans-aciculate, 
extreme apex polished, longer than rest of abdomen in <, shorter 
in @; second segment basally rugose, remainder smooth. 
Terebra in 9 twice as long as body, ferruginous brown, sheaths 
brown, broadly white-banded before apex. Hind coxe rather 
slender, finely reticulate outside, trans-aciculate inside; femora 
bidentate. Radius in fore wing sharply geniculate and extending 
nearly to margin of wing. 

Black; head—except mandibles apically, vertex, and space 
between frontal tubercles,—five basal antennal joints, all 
trochanters, tibie and tarsi, apices of hind femora, petiole 
laterally, apical segment dorsally only in @, entirely in d, and 
the male genitalia ferruginous; anterior femora brown. Wings 
hyaline, apically slightly infuscate; stigma and nervures brown. 

Length, 9,183 mm.; abdomen 13 mm.; petiole5?mm.; fore 
wing 11 mm.; terebra 37 mm. 6,13 mm.; abdomen 8 mm.; 
petiole 5 mm.; fore wing 11 mm. 

Habitat: Barombi, Cameroons. 1 9,1 $3 L. Conradt. 


5. CRASSIPES Bischoff. 
Neostephanus crassipes Bischoff, p. 329, 2. 


Q. Frons subarcuate rugose; vertex transrugose; occiput 
punctate-rugose with central impression; posterior margin of 
head bordered. Antenne normal. Prothorax smooth, laterally 
obliquely rugose; mesonotum irregularly punctate rugose; 
scutellum smooth, laterally strongly punctate. Mesopleure 
diffusely punctate: metapleure coriaceous in front, rather closely 
punctate behind, separated by a row of strong punctures from 
the median segment, which is coarsely, densely, and partly 
confluently punctate. (Details of petiole wanting.) Terebra 
much longer than body, sheaths black, white-banded before 
apex. 

"Tela to black-brown ; face below anterior tubercles, temples, 
cheeks, occiput laterally, four basal antennal joints, second segment 
laterally, third basally and ventrally, last segment entirely, hind 
trochanters, apices of hind femora, all tibiz and tarsi more or less 
yellow-brown. 

Length 19 mm.; terebra 27 mm. 

Habitat: Kasindi, north of Lake Albert Edward. Jan. 1908. 

Details as to the sculpture etc. of abdomen and as to hind legs 


HYMENOPTEROUS FAMILY STEPHANIDA. 781 


are wanting in the original description, and the unique specimen 
of WV. globiceps was without abdomen, while the only details as to 
the hind legs are that the coxe are trans-striate and the femora 
elongate. 

Apparently very like WV. globiceps Enderl., but frons less 
arcuately rugose, space between tubercles trans-striate, sculpture 
of occiput coarser and the impression less deep, puncturation of 
metapleure and median segment confluent. 


6, ALBOMACULATUS Cam. 
Fenatopus albomaculatus Cam. (7), p. 18, @. 


@. Frons closely and regularly reticulated; vertex duil, 
irregularly reticulate, with a transcarina behind tubercles; 
occiput irregularly closely striate, with an indistinct longitudinal 
impression. Second and third flagellar joints of equal length. 
Neck irregularly trans-striate, semiannular lightly reticulate ; 
mesonotum basally reticulate, laterally obliquely striate; scutellum 
smooth and dull. Mesopleuree smooth; metapleure finely 
reticulate. Median segment with round, clearly sepasated 
punctures, and microscopic sculpture. Petiole as long as thorax; 
terebra as long as body, sheaths black. Hind coxe dull, coarsely 
aciculate, basally reticulate, apically striate; hind femora with a 
blunt, rounded basal tooth and two other long-pointed teeth. 

Black; clypeus and a large spot on malar region whitish 
testaceous; mandibles basally and the cavina on vertex rufescent : 
base and apex of petiole-and a transverse spot at base of second 
segment white. Wings hyaline; stigma and nervures black. 

Length 12 mm.; terebra 12 mm. 

Habitat: Sebakwe, S. Rhodesia. 


7. PENTHERI Kieff. 
Neostephanus pentheri Kieffer (2), p. 101; Morley (1), p. 111, ¢. 


¢. Frons dull, reticulate; vertex with two transcarine ; 
occiput finely rugose. Antenne normal. Prothorax trans- 
rugose ; mesonotum very short and transverse, finely shagreened, 
as is also the scutellum. Mesopleure closely granulose and dull ; 
metapleure trans-striate and shining, separated by a carina from 
the median segment, which is as long as broad, reticulate and 
diffusely punctate. Petiole finely trans-striate, as long as re- 
maining segments. Hind legs with cox transrugose; femora 
bidentate ; tibizee compressed in basal half; tarsi five-jointed, meta- 
tarsus longer than the four following joints. 

Red-brown ; mandibles black, antenne basally pale, base and 
apex of petiole and a basal spot on each side of third segment 
yellow. Wings subhyaline; stigma black, basally white, 

Length 10 mm. 

Habitat: Zerua,S. Africa. Taken by Dr. Penther in November. 

The British Museum has a specimen taken in Angola in 1878 
by J. J. Monteiro. 


782 MR. E. A. ELLIOTT ON THE 


Fanarorus Smith. 


This subgenus was erected in 1860 by Smith, to include all 
those species of Stephanus having reduced venation, but was 
restricted by Enderlein (Zool. Anz. xxviii. 1906) to those having 
only the basal cells complete, and a portion of the median nervure 
beyond the transverse nervure representing the upper boundary 
of the submedian cell. In Diastephanus this part of the median 
nervure is either entirely wanting or reduced to a mere stump. 
Although these subdivisions of the Stephanide may be said to be 
purely artificial, yet, since some of them appear to be confined to 
certain regions, and no transition forms are known, they seem to 
be founded on permanent differences, and are of the highest value 
in identifying the numerous species. 

The number of teeth on the hind femora and the length of the 
petiole as compared with the rest of the abdomen, and, in the @, 
the length and colour of the terebra and its sheaths, are among 
the best characters for differentiation. 


Table of Species. 


Or 
(25.) 1. Hind femora bidentate. 
(16.) 2. Petiole distinctlylonger or shorter than rest of abdomen. 
(12.) 3. Petiole distinctly longer than rest of abdomen. 
(5.) 4. Sheaths of terebra black; head arcuate striate ... 1. ocellatus Elliott. 
(4.) 5. Sheaths of terebra pale-banded before apex. 
(7.) 6. Sheaths of terebra white-banded ..................... 2. indieus Westw. 
(6.) 7. Sheaths of terebra yellow-banded. 
(9.) 8. Terebra only as long as body............ 0.0.0. .ce ses eee 3. sumbanus Enderl. 
(8.) 9. Terebra much longes than body. 
(11.) 10. Mesopleurz closely a 2nd segment basally smooth 
and shining ........ lececceeee 4. formosanus Ender]. 
(10.) 11. Mesopleurx very chinese punetates 2nd segment basally 
granulate and dull . a tesesecsseee 5. Sulcaticollis Enderl. 


(3.) 12. Petiole distinctly Norte Ghant rat de abdomen’ 
(14.) 13. Terebral sheaths ferruginous, broadly yellow-banded, apex 
black. Head bale marked ; middle tibiae and hind meta- 
tarsus white.. wean ceseeeeeeeee 6. annulitarsis Enderl. 
(13.) 14. Terebral sheaths ot Coifomipandede: vertex transrugose. 
(15.) 14a. Sheaths ferruginous, apex black; mesonotum irregularly 
coarsely rugose; scutellum smooth and shining; ; metapleure 
aud median segment alutaceous, separated by a carina. [Schultz. 
7. fernandopoensis 
(14a.) 15. Sheaths brown, apex black; mesonotum and scutellum 
densely rugose-punctate, latter with central smooth space ; 
metapleure and median segment densely superficially 
punctate, not separated ........0..... cere 8. schlettererit Enderl. 
(2.) 16. Petiole about as long as abdomen. 
(18.).17. Pronotum very short, densely rugose-punctate; terebra 
shorter than body, sheaths yellow-brown ...... 9. brevicollis Ender]. 
(17.) 18. Pronotum not very short. 
(22.) 19. Pronotum normal. 


(21.) 20. Neck oblique rugose, semiannular rugose-punctate; meso- 
pleuree punctate; terebra as long as body, sheaths red- 
brown, apex black ............... vesseseseeeee 10. conradti Tinderl. 
(20.) 21. Neck trans-striate, semiannular anal inevopleute smooth and 
shining ; terebra shorter than body, black, white-banded [Schlett. 
[DROME GOES cooconmoncenbeaso caseosonatosoedannnpoacocds ty (Ke Aa oy OCTOLOIS 
(19.) 22. Pronotum very elongate. 
(24.) 23. Neck trans-striate, semiannular and mesopleure smooth. 
12. longicollis Carn. 
(23.) 24. Pronotum transrugose throughout. Rufo-piceous, abdomen  [Semenov. 
white-marked .1.............c...ceeesecsesseeeessvesese LB. Curcomanorum 
(1.) 25. Hind femora tridentate. 
(29.) 26. Neck of pronotum very elongate. 
(28.) 27. Neck of prothorax alutaceous ; hind femoral teeth black. 
14. simpsoni Kieff. 
(27.) 28. Neck of pronotum finely striate; middle and apical hind 
femoral teeth white........... 02.0... cece... 15. variidens Eliott. 
(26.) 29. Neck of pronotum not elongate. 
(31.) 30. Neck of pronotum very short; petiole shorter than rest of 
abdomen ; terebra as long as body ............... 16. arcuatus Kieff. 
(30.) 31. Pronotum of ordinary length. 
(35.) 32. Frons striate, but not arcuately. 
(84.) 33. Frons finely trans-striate, occiput alutaceous ; petiole shorter 
than rest of abdomen ; terebra shorter than body. 
17. natalicus Westw. 
(33.) 34. Frons rugosely trans-striate, occiput smooth ; petiole longer 
than rest of body ; terebra 13 times as long as body. 
18. longicauda Elliott. 
(32.) 35. Frons arcuate rugose. 
(87.) 36. Frons sparsely, almost granulately rugose; pronotum strongly 
trans-striate; legs rufescent ; basal and central femoral 
teeth white, the apical black........................ 19. rugiceps Elliott. 
(86.) 37. Frons moderately arcuate striate to rugose; vertex and 
occiput trans-striate. 
(41.) 38. Neck of prothorax smooth. 
(40.) 39. Pronotum entirely smooth; terebra shorter than body, black. 
20. piceicornis Cam. 
(39.) 40. Rest of pronotum trans-striate; terebra half as long again 
as body, white-banded .....................4.... 21. iridipennis Elliott. 
(38.) 41. Neck of pronotum striate or rugose. 
(43.) 42. Pronotum strongly transrugose; lateral lobes of scutellam 
GHOSE? OUND ccassosonss2c0sdosseso0n cau node ooboss 22. punctatus Elliott. 
(42.) 43. Neck of pronotum more or less coarsely obliquely striate, 
remainder alutaceous. 
(45.) 44. Metapleuree and median segment confluent, coarsely and 


(44.) 45. 


HYMENOPTEROUS FAMILY STEPHANIDA, 783 


superficially punctate. Terebraas long as body, ferruginous. 
23. togoensis Stadlm. 
Metapleure and median segment separated by a line of 
punctures, diffusely punctate; terebra shorter than body, 
LENG Sees sapacaamaucne sconneceancpeactes . 24. claripennis Elliott. 


3. 


. Hind femora bidentate. 
. Scutellum depressed, coarsely punctate; petiole shorter than 


rest of abdomen; 2nd segment basaily longitudinally 
striate. Black; head bright rufo-testaceous... 25. aratifrons Enderl. 


3. Scutellum not depressed, only laterally punctate; petiole 


longer than rest of abdomen; 2nd segment basally smooth. 


784 MR. E. A, ELLIOTT ON THE 


(5.) 4. Frons and occiput finely transrugose ; pronotum transrugose 
throughout; 2nd segment apically dull. Black; head 
yellow, vertex and occiput black . veceessesese 4, formosanus Enderl. 


(4.) 5. Frons coriaceo-rugose, vertex and Reciott transrugose; neck 
elongate, transrugose, semiannular smooth; 2nd segment 
basally smooth and shining. Black; head red, pale marked. 
2, indicus Westw. 
(1.) 6. Hind femora tridentate. 


(10.) 7. Anterior frontal tubercles normal, larger than the posterior. 


(9.) 8. Frons transrugose, occiput alutaceous; pronotum trans- 
striate; petiole longer than rest of abdomen. wufo- 
testaceous, median seement apically black ...... 16. natalicus Westw. 


(8.) 9, Frons arcuate rugose, occiput finely transrugose and 
impressed; neck oblique striate; semiannular alutaceo- 
punctate; petiole as long as, or slightly shorter than rest 
of abdomen. Black; head, thorax, and legs ferruginous. 

22. togoensis Stadlm, 


(7:) 10. Anterior frontal tubercles smaller than the posterior ; vertex 
arcuate striate; mesonotum irregularly striate. Black ; 
face, outer orbits, vertex, prothorax except central black 
line, meso- and meta- pleur z, and anterior legs testaceous. 
26. flavicollis Cam, 


1. ocetuatus Elliott. 
Fenatopus ocellatus Elliott (2), 4 p- 13, 2. 


2. Head arcuate striate, occiput more finely; three strong 
cavine between the posterior ocelli; posterior margin of head 
bordered ; second flagellar joint one and a half times as long as 
first ; third as long as first and second together. Neck elongate, 
trans-striate, remainder of pronotum subglabrous; mesonotum 
coarsely punctate; pro- and meso-pleure glabrous, metapleurz 
and median segment cribrate punctate. Petiole trans-striate, 
longer than rest of abdomen; terebra shorter than body, black. 
Hind legs with cox trans- stri iate; femora smooth, bidentate ; 
tibize compressed to middle. 

Black ; head and base of antenne rufescent, ocellar region pale, 
and the ocelli deep, shining black; basal third of middle tibiz 
and basal half of their metatarsi white. Wings hyaline, stigma 
yellow-brown. 

Length 13 mm.; abdomen 8 mm.; petiole 43 mm.; terebra 
10 mm. 

Habitat: Mount Matang, Sarawak. 18.1.1914; G. EK. Bryant. 

The deep black ocelli are strikingly conspicuous against the pale 
rufescent background. 


2. InDIcUS Westw, 

Stephanus indicus Westw. (2), p. 588; (3), p. 277; Schlett., 
105 WAS), QS Megischus indicus Westw. (4). KFenatopus 
ruficeps Smith (2), p. 58, 9. Fenatopus indicus Enderl. (3), 

. 290, 9, 4), p..297.; Morley (1), p. 36, 2d. 

2. Frons finely coriaceo- rugose ; occiput finely trans- -striate, 
with a shallow central longitudinal impression. Posterior margin 
of head bordered. Second flagellar joint one and a half times as 


HYMENOPTEROUS FAMILY STEPHANID. 785 


long as first; third nearly as long as first and second together. 
Neck elongate, very slender and finely transrugose above; 
semiannular shining smooth, laterally subrugose. Mesonotum 
lightly transrugose. Scutellum smooth and shining, with a few 
marginal punctures. Mesopleure polished smooth above, other- 
wise diffusely punctate ; metapleuree and median segment super- 
ficially cribrate punctate, not separated by any sulcus, Petiole 
slightly longer than rest of abdomen, rather shorter than hind 
cox, trochanters, and femora together, very finely trans-striate, 
remaining segments entirely smooth. Terebra slightly longer 
than body, sheaths white-banded before apex. Hind legs with 
cox slender, finely transrugose; femora dull, bidentate; tibie 
longer than femora, compressed to beyond middle. 

Black ; head red, frons and temples pale marked, anterior legs 
more or less brown, intermediate tibize and tarsi white-marked. 
Wings hyaline. 

3. Similar, but more coarsely sculptured, especially on median 
segment. 

Length 11-14 mm. 

Habitat: East India, Ceylon, Malabar, Philippines, Borneo, 
Sumbava, Celebes. Sumatra, a specimen measuring 19 mm. 
recorded by Dr. Dohrn. 

Enderlein (/.c.) records specimens of this species, taken by 
Dr. Dohrn in Sumatra, having the impression on occiput more 
distinct, neck of pronotum more coarsely transrugose, semiannular 
transaciculate in front, basal two-thirds of second abdominal 
segment polished smooth, remainder of abdomen dull, and hind 
femoral teeth black. Length 19-20°5; terebra 21-23 mm. This 
may prove to be a distinct species. 

“Var. (?) 2. Face finely alutaceo-rugose. Certainly a 
different species.” Szépligetti, Termés. Fiiz. xxv. p. 531 (1902). 

Habitat : Borneo. 


3. SUMBANUS Ender]. 


Fenatopus imdicus var. sumbana Enderl. (4), p. 298; F. 
sumbanus Enderl. (3), p. 209, 9. 


2. Frons finely coriaceo-rugose, occiput finely trans-striate ; 
posterior margin of head slightly bordered. Second flagellar 
joint about one and a half times as long as first; third nearly as 
long as first and second together. Neck deeply transrugose, 
laterally finely and closely trans-striate, semiannular smooth 
above, without trace of sculpture, laterally finely and closely 
trans-striate, posterior angles strongly transrugose. Mesonotum 
irregularly to transversely rugose. Scutellum smooth, laterally 
finely aciculate with marginal punctures. Mesopleure rather 
smooth, diffusely punctate, with a central rugose and closely 
trans-striate space. Metapleurz and median segment with large 
eribrate punctures. Petiole very finely and closely trans-striate, 
longer than rest of abdomen; second segment basally rugose, 


786 MR. E. A. ELLIOTT ON THE 


centrally smooth, apically microscopically sculptured and dull. 
Terebra as long as body, sheaths black, with yellowish subapical 
band. Hind coxe trans-striate; hind femora bidentate. 

Black; head flavo-ferruginous, frons and vertex darker ; basal 
half of second segment rufescent; anterior coxe and legs, hind 
tibie and tarsi ferruginous, femoral teeth rufescent. Wings 
hyaline, stigma very long and narrowish, brown; nervures dark 
brown. 

Length 163 mm.; petiole 5-4 mm.; terebra 163 mm.; fore 
wing 9 mm. 

Habitat: Sumba, Malay Archipelago. 1 9 ; Grelak. Typeat 
Stettin. 

Differs from /’. indicus in the sculpture of prothorax, shorter 
terebra, and colour. The femoral teeth are rufescent, whereas in 
I. indicus they are black (Enderlein), or white (Schlett.); the 
black apex of terebral sheaths is more than half as broad as the 
pale subapical band in swmbanws, but very much less in indicus. 


4. rorMosANUS Ender]. 
Fenatopus formosanus Ender]. (3), p. 207, 9 ¢. 


2 6. Frons rather finely and irregularly transrugose, the 
ridges outwardly forked ; occiput more finely rugose, with indis- 
tinct longitudinal median impression. Pronotum transrugose, 
more densely towards base, and intermixed with coarse punctures, 
lateral margins smooth, a broad posterior margin polished smooth 
and in front of it two roundish lateral spaces very finely and 
densely striate, having a velvety appearance, posterior angles 
strongly rugose. Scutellum polished smooth, laterally sparsely 
punctate. Mesopleure rather closely punctate, posterior margin 
very finely punctate and sparsely yellow pubescent. Petiole finely 
and densely trans-striate; second segment basally smooth and 
shining, apical half dull. Terebra in 2 much longer than body, 
sheaths black, with a broad yellow band before apex. No details 
as to the hind legs are given. 

Black; in @ frons black-brown, with three longitudinal rufo- 
testaceous lines, vertex and occiput black, remainder of head 
ferruginous yellow; in ¢ head light yellow, vertex and occiput 
black. Antenne black, two basal joints yellow, third brown. 
Anterior legs from femora red-brown, tibie and tarsi in ¢ paler. 
Hind tarsi, sometimes also hind tibie, dark red-brown. Wings 
hyaline, stigma pale brown, nervures black-brown, apical nervure 
of median cell more or less pale. 

Length, 9, 12:4-17°75 mm.; petiole 45-7 mm.; terebra 16°5— 
24-5 mm.; fore wing 7°5-10°75 mm. 6, 103-163 mm.; petiole 
4-6-6 mm.; fore wing 5:3-9 mm. 

Habitat: Formosa, many localities, by H. Sauter, 1909-12; 18 
29,12 6 Ss. Co-types in Dahlen and Stettin Museums. 

One large 9, taken by H. Sauter in Hankau in 1912, has the 
nervure bounding the median cell pigmented throughout. Length 


HYMENOPYEROUS FAMILY SYTEPHANID#. 787 


21°5 mm.; petiole 7 mm.; terebra 29 mm.; fore wing 12°6 mm. 
Enderlein states that the development and pigmentation of the 
nervure mentioned varies with the size of the individual, being 
shortest and palest in the smallest. 


5. SULCATICOLLIS Enderl. 


Fenatopus indicus var. sulcaticollis Enderl. (4), p. 298. /. 
sulcaticollis Enderl. (3), p. 210, 2. 


@. Frons finely coriaceo-rugose; occiput finely trans-striate, 
temples smooth polished; posterior margin of head slightly. 
bordered. Second flagellar joint about one and a half times as 
long as first; third nearly as long as first and second together. 
Pronotum not rugose, but finely and densely trans-striate 
throughout, except the rugose posterior lateral angles and the 
smooth posterior margin. Mesonotum transrugose; scutellum 
smocth, with diffuse coarse marginal punctures. Mesopleure 
smooth, very diffusely punctate; metapleuree and median segment 
cribrate punctate.  Petiole finely and densely, but unusually 
superficially trans-striate, slightly longer than rest of abdomen ; 
second segment with basal sixth granulate, apical half dull, 
central part smooth and shining. Terebra longer than body, 
sheaths black-brown with broad yellow subapical band. Hind 
coxe more strongly trans-striate than the petiole; hind femora 
bidentate. 

Black; head and three basal antennal joints ferruginous; 
apices of tergites 2-7 brownish ; anterior coxe dark ferruginous, 
anterior tarsi paler, hind tarsi black-brown. Wings hyaline, 
stigma very narrow, brown ; nervures dark brown. 

Length 18 mm.; petiole 64 mm.; terebra 23 mm.; fore wing 
93 mm. 

Habitat: Sumatra (Dr. Dohrn). Type in Stettin Museum. 

From /’. indicus Westw. and J’. sumbanus Enderl. it differs in 
sculpture of pronotum and petiole as well as in colour, and from 
the latter also in the longer terebra. 


6. ANNULITARSIS Enderl. 
Fenatopus annulitarsis Enderl. (3), p. 206, ¢. 


2. Frons, vertex, and occiput arcuate rugose. Pronotum 
strongly transrugose, laterally finely aciculate, posteriorly longi- 
tudinally striate; scutellum coarsely and densely punctate, 
posterior margin finely aciculate. Mesopleure smooth, closely 
punctate, posterior margin finely trans-striate, upper half yellow 
pubescent. Petiole finely and densely trans-striate; second 
segment smooth, basally rugose, remainder of abdomen dull. 
Terebra shorter than body, sheaths ferruginous, with indistinct 
yellow-brown band before the black apex. Basal half of hind 
coxe transrugose; hind femora bidentate. 

Black; face, cheeks, and temples ferruginous yellow ; hind 
legs partly dark ferruginous; middle tibize and metatarsus 

Proc. Zoou. Soc.—1922, No. LILI. 53 


788 MR. E. A. ELLIOUVT ON THE 


basally, hind metatarsus entirely whitish yellow. Teeth on hind 
femora black. Wings hyaline, stigma pale brown. 
Length 9-143 mm.; petiole 2°4-3°6 mm.; terebra 7:8- 
11:6 mm. ; fore wing 5°7-7°8 mm. 
‘Habitat: Taihorin, Kankau, Hoozan, Formosa. 1910-12; 
H. Sauter. Co-types in Dahlen and Stettin Museums. 


7, FERNANDOPOENSIS Schultz. 
Fenatopus fernandopoensis Schultz (1), p. 273, @. 


2. Frons finely and densely arcuate rugose, dull, with 
subobsolete longitudinal carina; vertex coarsely transrugose ; 
occiput finely and irregularly rugose; cheeks finely and sparsely 
punctate. Posterior margin of head finely bordered. Basal 
flagellar joints normal. Pronotum alutaceo-rugose, extreme 
posterior margin polished smooth, laterally and apically diffusely 
punctate. Mesonotum irregularly and coarsely transrugose ; 
scutellum centrally smooth and shining, laterally finely punctate 
and dull. Mesopleuree with posterior larger half closely punctate. 
Metapleure and median segment finely alutaceo-punctate, latter 
diseally diffusely and coarsely punctate, separated by a carina. 
Petiole finely and densely trans-striate, shorter than rest of 
abdomen. Terebra shorter than body, spicula and sheaths ferru- 
ginous, latter apically black. 

Black; mandibles except apex, face and cheeks rufo-testaceous ; 
ferruginous are five basal antennal joints, anterior legs (coxe, 
trochanters, and femora nigrescent), hind coxz apically, hind 
trochanters chiefly, basal half and apex of hind tibie and the 
tarsi, basal band on second and third abdominal segments, apex 
of third tergite obscurely and apex of last sternite. All the last 
tarsal joints and the claws black. Wings hyaline. 

Length 9 mm.; terebra 7 mm.; fore wing 5 mm. 


Habitat: Fernando Po. Type in coll. W. A. Schultz. 


8. SCHLETTERERI Enderl. 


Stephanus schletterert Fnderl. (1), p. 201, 2. Fanatopus schl. 
(2), p. 474. 


@. Frons transrugose, with indistinct central carina; vertex 
coarsely transrugose ; occiput irregularly and finely rugose- 
punctate ; temples smooth, subrugose near the eyes. Posterior 
margin of head sharply bordered. Neck almost smooth; semi- 
annular polished smooth, laterally punctate. Mesonotum and 
scutellum densely rugose-punctate, latter with a small smooth 
central space. _ Metanotum not longitudinally striate. Meso- 
pleure smooth, diffusely and superficially punctate ; metapleure 
and median segment not separated, densely and _ superficially 
punctate. Petiole finely trans-striate, shorter than remainder of 
abdomen. ‘T'erebra rather shorter than body, sheaths yellow- 
brown, apically black. Hind coxe finely trans-striate. 


HYMENOPTEROUS FAMILY STEPHANIDA, 789 


Black ; mandibles except apex, basal half of antenne, a streak 
on temples along eyes, the tegule, tibize and tarsi yellow. Wings 
colourless hyaline. 

Length 10-11 mm. 

Habitat: Johann Albrechtshohe, N. Cameroons. IL. Conradt ; 
1895-96. 

Differs from /. brevicollis Knderl. in the formation of the pro- 
pleure, reticulate transrugose face, longer and smoother pronotum, 
and shorter petiole. 


9. BREVICOLLIS Enderl. 

Stephanus brevicollis Knderl. (1), p. 201, 9. Fanatopus b. 
(2), p. 474. 

Q. Head densely granulate rugose; vertex transrugose ; 
occiput densely punctate rugose. Posterior margin of head 
sharply bordered. Second flagellar joint a little longer than 
first; third and fourth as long as scape and about as long as first 
and second together. Pronotum short, densely punctate rugose ; 
mesonotum and scutellum densely punctate, latter without any 
central smooth space; metanotum finely transrugose. Propleure 
separated from prothorax by a distinct acute-angled sulcus, 
produced into free-standing edges, which, seen laterally, appear 
like two sharp points. Mesopleurze above smooth and shining, 
lower half punctate; metapleure and median segment separated 
only by a smoother streak, and both closely punctate. Petiole 
extremely finely trans-striate, as long as rest of abdomen. 
Terebra rather shorter than body; the spicula and sheaths, 
except black apex, yellow-brown. Hind coxe finely trans- 
striate. 

Black; mandibles except apex, three basal antennal joints, 
tegulee, anterior tibie and all tarsi yellow-brown. Wings hyaline. 

“Length 7-10 mm. 

Hepitat.: Bismarcksburg, Togo. LL. Conradt; 1895. 

This species appears to differ from all others in the peculiar 
formation of the propleure. From /. schlettereri Ender]. it further 
differs in the shorter prothorax and coarser sculpture of pro- and 
meso-notum. 


10. conraptt Enderl. 


Stephanus conradtt Ender]. (1), p. 202, 2. Hanatopus c. (2), 
p. 474. 


@. Frons very densely arcuately longitudinally rugose; vertex 
with three or four coarse ridges; occiput densely irregularly 
rugose, with subobsolete central channel. Temples subrugose 
near the eyes. Posterior margin of head slightly bordered. Basal 
flagellar joints normal, third slightly shorter than fourth and as 
long as scape. Neck obliquely rugose, remainder diffusely 

53* 


790 MR. E. A. ELLIOTT ON THE 


punctate. Mesonotum indistinctly transrugose; scutellum 
nitidulous, with fine and diffuse small punctures. Metanotum 
relatively elongate, sharply longitudinally striate. Propleure 
separated from pronotum by a narrow sulcus; mesopleure 
rugose-punctate ; metapleurz rugose, apically trans-striate below, 
sparsely aciculate above. Median segment not very closely 
punctate, separated from the metapleure by a shining, curvate 
impression. Petiole finely striate, about as long as rest of 
abdomen. Terebra slightly shorter than body, sheaths red-brown, 
apically black. Hind coxe finely trans-striate. 

Black ; mandibles basally, six basal antennal joints, anterior 
legs, apices of hind femora, basal half of hind tibie and hind tarsi 
red-brown or chestnut. Wings hyaline, nervures brown. 

Length 15 mm. 

Habitat: Bismarcksburg, Togo. LL. Conradt; March 1893. 


11]. LACTEIPENNIS Schlett. 
Stephanus lacteipennis Schlett., p. 119, 2. 


2. Frons rather finely and arcuately punctato-rugose; vertex 
arcuately rugose, occiput gradually more irregularly. Posterior 
margin of head simple. Basal flagellar joints normal. Neck 
finely but distinctly trans-striate; semiannular smooth and 
shining, apically and laterally finely trans-striate. Mesonotum 
rather coarsely transversely punctato-rugose. Scutellum with 
larger section smooth, lateral lobes diffusely punctate. Meso- 
pleuree smooth and shining, rugoso-punctate in front; meta- 
pleure and median segment very coarsely cribrate punctate, 
separated by an indistinct sulcus. Petiole finely trans-striate, 
about as long as remainder of abdomen and distinctly shorter 
than the hind cox, trochanters, and femora together; second 
segment basally finely rugose, rest of abdomen dull. Terebra 
slightly shorter than body, sheaths black, white-banded before 
apex. Hind legs with coxe distinctly trans-striate; femora 
anteriorly shining smooth, posteriorly finely trans-striate and 
vather dull, bidentate; tibize not longer than femora, compressed 
to beyond middle. 

Black; head, mandibles basally, and scape ferruginous; a pale 
streak from cheeks along eyes; anterior legs chestnut-brown, 
their joints pale marked. Wings milky white. 

Length 22 mm. 

Habitat: Borneo. Type in National Museum, Budapest. 

The peculiar miiky-white wings and short tibie are distinctive 
of this species. 


12. LONGICOLLIS Cam. 


Megiseleius longicollis Cam. (8), p. 32, 2. 


2. Frons coarsely and closely striate, obliquely above, trans- 
versely below ; vertex and occiput closely obliquely striate, latter 


HYMENOPTEROUS FAMILY STEPHANIDA. 791 


with indistinct impression. (No details as to the proportions of 
the antennal joints are given, and these are broken off in the 
eo-type in the British Museum.) Prothorax elongate, deeply 
incised at apex, the incision basally rounded; neck trans-striate, 
semiannular smooth. Mesonotum basally coarsely and irregularly 
reticulate, apically centrally impressed and laterally irregularly 
punctate. Scutellum smooth, laterally punctate. Mesopleurs 
smooth, basally pilose; metapleurs rugose. Median segment 
closely and coarsely punctate rugose. Petiole trans-striate, about 
as long as the smooth remainder of abdomen. Terebra longer 
than body, white-banded before apex. Hind coxe closely but 
not strongly striate; hind femora bidentate. 

Black; head pale rufous, orbits yellowish, three basal antennal 
joints, anterior legs, and base of second segment rufescent ; pro- 
thorax brownish. Wings clear hyaline, stigma and nervures 
black. 

Length 18-20 mm.; terebra 20-22 mm. 

Habitat: Sarawak (Robert Shelford). 


13. TURCOMANORUM Semenow. 
Stephanus turcomanorum Semenow, p. 435; André, p. 488, 9. 


@. Face and frons coarsely subtransversely rugose; vertex 
with two transcarine enclosing a falcate space; occiput trans- 
rugose; temples finely subrugose above, with a shining raised 
space near external margin of eyes, beneath which they are 
coriaceous; posterior margin of head simple but sharp. Third 
flageliar joint longer than second, but shorter than first and second 
together. Pronotum regularly transrugose, neck elongate. 
Mesonotum in front irregularly punctato-rugose, behind more 
feebly and subarcuately rugose. Mesopleure densely coriaceo- 
rugose; metapleure and median segment reticulate rugose, 
scarcely separated. Petiole finely and densely trans-striate, about 
as long as rest of abdomen. ‘Terebra about as long as body, 
sheaths unicolorous nigro-piceous. Hind legs with cox regularly 
transrugose; femora finely coriaceous, dull, bidentate; tibia 
constrieted in basal two-thirds, and at the base of the clavate 
‘part irregularly and strongly impressed as if mutilated. 

Rufo-piceous ; head, thorax, and legs chiefly rufescent; base 
and apex of petiole and two round spots on basal third of second 
segment white. Wings hyaline, stigma and nervures fuscous, 
former basally white. 

Length 10 mm. 

Habitat: Transcaspia; one 9 taken near the military road to 
the station of Tedshe on the river of that name. Type in Mus. 
Zool. Acad. Cesar Scient. at Petropolsky. 

This species is not unlike /. indicus Westw., but distinguished 
by the coarse sculpture of head and temples, and by the white- 
marked petiole and second segment. 


792 MR. E. A. ELLIOTT ON THE 


14. srupsoni Kieff. 
Fenatopus simpsoni Kieffer (6), p. 233, 9. 


2. Head subglobose, dull; frons coarsely and irregularly 
rugose, reticulate between tubercles, with median longitudinal 
carina; vertex transcarinate; occiput reticulate. TVirst flagellar 
joint half as long as scape, second one and a half times as long as 
first; third and fourth of equal length and each twice as long as 
second, Pronotum elongate, dull, alutaceous, more strongly 
behind; mesonotum reticulate; propleure and sternum coria- 
ceous; meso- and meta-pleure, scutellum, and median segment 
alutaceous, with large and diffuse punctures. Petiole densely 
trans-striate, as long as rest of abdomen. ‘Terebra shorter than 
body, ‘‘composed of two red filaments, apically black and spatu- 
late, without sheaths.” The absence of the sheaths must have 
been due to accident. Hind coxe closely trans-striate; hind 
femora tridentate, finely and indistinctly trans-striate; hind 
tibize only apically slightly inflated. 

Black; mandibles except apices, cheeks, 4-5 basal antennal 
joints, a distal spot on each side of third segment, tibiz and tarsi 
red; a spot on temples, one on vertex and the trochanters dull 
ved; anterior femora and apical third of hind tibie brown-black ; 
hind femoral teeth black; middle metatarsus whitish. Wings 
hyaline, stigma linear, yellowish hyaline. 

Length 11 mm.; abdomen 6 mm.; petiole 5 mm.; terebra 
10 mm. 

Habitat: Ion, South Nigeria; 13.11.1910; J. J. Simpson, 
Entom. Research Committee. 


15. VARIIDENS Elliott (4), p. 257, °. 
Fenatopus variidens Elliot (4), p. 257, 2. 


@. Frons and ocellar space very finely striate; vertex and 
occiput transversely, almost arcuately striate; posterior margin 
of head strongly bordered. Cheeks slightly shorter than scape. 
Second flagellar joint one and a half times as long as first; third 
nearly as long as first and second together. Prothorax finely 
trans-striate, neck elongate, apically above deeply incised. Meso- 
notum irregularly rugose-punctate, central row of punctures 
distinct; scutellum centrally smooth, lateral lobes punctate. 
Mesopleurze smooth; metapleurz and median segment cribrate 
and confluent. Petiole finely trans-striate, slightly shorter than 
rest of abdomen. ‘Terebra longer than body, with a 3 mm. broad 
white band before the 1 mm. broad black apex. Hind cox 
trans-striate; femora smooth, tridentate; tibie slightly longer 
than femora, compressed almost throughout. 

Black; head ferruginous, frons darkest, inner orbits and 
anterior frontal tubercles paler; vertex, including ocellar space 
and occiput, black. Front tibie and tarsi rufescent; base of 
intermediate femora, their tibie externally and apical tarsal 
joints rufescent, metatarsus white; hind legs entirely black, 


HYMENOPTEROUS FAMILY STEPHANIDA. 193 


except the middle and apical femoral teeth, which are white. 
Wings iridescent hyaline, nervures pale brown, stigma centrally 
translucent. 

Length 16 mm.; abdomen 94 mm.; petiole 44 mm.; terebra 
18 mm. 

Habitat: Paleboehan, Ratoe. M. EK. Walsh (ea Frisby); 22nd 
February, 1916. Type in coll. Vlaude Morley. 

The colour of the femoral teeth is distinctive. In F. rugiceps 
Hiliott the basal tooth is white, the apical black; in the present 
species this is reversed. 


16. arcuatus Kieff. 
Fenatopus arcuatus Kieffer (4), p. 118, 2. 


@. Head subglobose; frons irregularly rugose, almost reticu- 
late; vertex transcarinate; occiput irregularly rugose, with 
longitudinal sulcus; temples centrally prominent; posterior 
margin of head bordered. Second flagellar joint about one and a 
half times as long as first; third as long as first and second 
together. Neck very short, thorax cribrate punctate above 
throughout; median lobe of scutellum centrally slightly sha- 
greened. Mesopleure diffusely cribrate punctate. Petiole 
densely transaciculate, rather shorter than rest of abdomen. 
Terebra as long as body, sheaths brown, apically black. Hind 
legs with coxve trans-striate; femora tridentate; tibize compressed 
rather beyond middle. 

Dull black; head entirely red, or occiput black-brown; five 
basal antennal joints red; anterior femora brown; all tibize and 
tarsi pale brown or brownish white. Wings hyaline, stigma and 
nervures brown. The radius curvate, not angulate, emitted from 
apical third of stigma. 

Length 10 mm. 

Habitat: Cameroons. Type in National Entom. Museum, 
Berlin. (Conradt.) 


17. NaTALIcUS Westw. 


Stephanus natalicus Westw. (5), p. 127, pl. xxiv. fig. 8: Schlett., 
p. 102, 93. WNeostephanus natalicus Enderl. (2), p. 474. 
Fenatopus natalicus Morley (1), p. 110, 2 6 


@ g. Frons finely transrugose, vertex transcarinate; occiput 
alutaceous. Posterior margin of head simple. Second flagellar 
joint one and a half times as long as first; third nearly as long as 
first and second together. Prothorax trans-striate. Mesonotum 
diffusely punctate; scutellum smooth. Meso- and meta-pleure 
nearly smooth, former pubescent, latter confluent with median 
segment, which is superficially and diffusely punctate. Petiole 
shorter than rest of abdomen, transaciculate, remainder smooth 
and feebly shining. Terebra shorter than body, fulvous, apically 
black. Hind legs with coxe apically transaciculate; femora 
tridentate ; tibiz constricted to beyond middle. Tarsi five-jointed 
in 6, three-jointed in female, 


794 MR. E. A. ELLIOTT ON THE 


Fulvo-testaceous; antenne basally nearly white; apex of 
median segment discally black; central abdominal segments 
apically nigrescent. Wings fuscous or fulvo-hyaline, iridescent, 
stigma pale. 

Length, 9,10 mm.; abdomen 6 mm.; petiole 23 mm.; terebra 
Sy omying 5 ay onan, 

Habitat: Port Natal, 5. Africa. Type in Hope Museum, 
Oxford. 

In the British Museum are an old pair from F. Smith’s 
collection with label *‘ Port Natal,’ and a female with a green 
label from the same locality; all three were probably taken by 
the Swedish collector, Gueinzius, about 1859. 

This species bears a strong superficial resemblance to Dvaste- 
phanus alutaceus Morley, but, in addition to the venation, 1t may 
be distinguished easily by the sculpture of vertex and pronotum, 
and by the tridentate hind femora. 


18. Lone1cAUDA Elliott. 
Fenatopus longicauda Elliott (2), p. 30, 2. 


@. Frons transrugose, vertex finely trans-striate, occiput 

smooth and shining; frontal tubercles short and blunt; three 
stout carinz between the posterior ocelli. Posterior margin of 
head simple. Basal flagellar joints normal. Prothorax finely 
trans-striate ; mesonotum cribrate punctate; mesopleure smooth, 
with a few large punetures in the middle; metapleure coarsely 
punctate above, smooth below, separated by a crenulate furrow 
from the cribrate punctate median segment. Petiole trans- 
striate, a little longer than rest of abdomen, which is smooth and 
shining. Terebra half as long again as body, with a sharply- 
defined subapical white band, 2 mm. wide and 14 mm. at apex, 
black. Hind coxe trans-striate, basally more coarsely; hind 
femora very finely alutaceous and rather shining, tridentate ; 
hind tibize compressed to a little beyond middle. 

Black ; mandibles except apex and face pale testaceous; anterior 
legs rufescent, tibie paler; hind tarsi rufescent. Wings basally 
infusecate, nervures black. 

Length 15 mm.; abdomen 10 mm.; petiole 53 mm.; terebra 
22 mm. 

Habitat: Nilgiri Hills, India, 3000 ft.; 14th April, 1888 Sir 
George Hampson. Type in British Museum. 

Very like 7. wridipennis Elliott, but easily known by the infus- 
cate wings and smooth occiput with simple margin. 


19. RueicEers Elliott. 
Fenatopus rugiceps Klliott (2), p. 29, 9. 


@. Frons granulate ruguse, vertex and occiput subarcuate 
rugose. Frontal tubercles rather small and blunt, the space 
between them rugose; three carine between the posterior ocelli. 


HYMENOPTEROUS FAMILY STEPHANIDA. 795 


ior margin of head strongly bordered. Scape longer than 

Posterio f head strongly bordered. Scape longer tl 
cheeks, Second flagellar joint not quite half as long again as first ; 
third about as long as first and second together. Prothovax 
strongly trans-striate, with rather broad smooth posterior margin ; 
mesonotum coarsely punctate, apically smooth; propleuree lightly 
punctate; mesopleurz basally punctate, apically smooth ; meta- 
pleurz coarsely punctate, separated by a row of punctures from 

lan segment, which has large and diffuse punctures. 
the med g t, which has larg 1 diff punct 

rans-striate, as long as rest of abdomen, which is fine 
Petiole t triate, as long t of abdomen, which is finel 
alutaceous. Terebra shorter than the body, basally rufescent, 
the colour gradually passing into an ill-defined subapical yellowish 


Text-figure 5. 


Fonatopus rugiceps. 


ring, extreme apex black. Hind coxe coarsely transrugose, their 
femora tridentate, the basal tooth broad triangular, the central 
longer and more rounded, the apical narrow and very sharply 
pointed ; hind tibie constricted in basal two-thirds. 

Black ; head rufo-testaceous, frons and vertex nigrescent, face 
and base of mandibles testaceous. Front legs rufo-testaceous, 
femora darker ; middle legs lighter, with base of tibiee and rather 
- more than basal half of metatarsus white; hind legs rufescent, 
the basal and central femoral teeth white, the apical black, apex 
of tibize and the metatarsus except its apex white. Wings feebly 
infuseate, nervures blackish. 


796 MR. E. A. ELLIOTT ON THE 


Length 12 mm.; abdomen 7 mm.; petiole 33 mm.; terebra 
10 mm. 

Habitat: South India. F. Smith coll. Type in British 
Museum. 

The type bears a label with the MS. name “Vf. ruficeps 
Smith.” A red head is very common in this subgenus and in 
no way distinctive. The rugosity of the head seems more 
characteristic. This species may be easily known by the colour 
of the femoral teeth, which appears to be a very constant 
character. 


20. PICEICORNIS Cam. 
Fenatopus piceicornis, Cam. (10), p. 360, ¢. 


2. Frons subarcuately, vertex more distinctly arcuately rugose; 
occiput finely and closely trans-striate. Pronotum smooth; 
mesonotum in basal half strongly, irregularly reticulate, laterally 
coarsely punctate ; scutellum smooth, laterally coarsely punctate. 
Median segment coarsely and diffusely punetate. Petiole finely 
trans-striate, as long as rest of abdomen. Terebra slightly shorter 
than body. Hind femora tridentate. 

Black; outer orbits broadly yellow, mandibles apically rufo- 
testaceous, antennee piceous, apically darker; anterior tibie and 
tarsi pale testaceous; hind trochanters, apex of tibie and the 
tarsi rufo-testaceous. Wings clear hyaline, stigma and nervures 
black. 

Length 13 mm.; terebra 13 mm. 

Habitat: Dima, West Africa (A. Koller). Type in Congo 
Museum, Tervueren. 

The pale apex of mandibles is unusual, they are usually 
darker there. 


21. IRIDIPENNIS Elliott. 
Fenatopus iridipennis Elliott (2), p. 30, 2. 


@. Frons arcuate rugose, vertex and occiput finely and 
distinctly trans-striate; two short carine between the posterior 
ocelli; posterior margin of head bordered. Basal flagellar joints 
normal. Neck of pronotum smooth, rest trans-striate, with narrow 
smooth posterior margin. Mesothorax with large, well-separated 
punctures. Propleure finely striate; mesopleure finely alutaceous, 
metapleure and median segment cribrate punctate, separated by 
a carina. Petiole trans-striate, extreme apex smooth, as long as 
rest of abdomen; base of second segment trans-striate, remainder 
smooth. Terebra half as long again as body, white subapical 
band 2 mm. broad. Hind coxee finely trans-striate, their femora 
tridentate; tibize constricted in basal two-thirds. 

Black; head red, vertex black; the anterior tubercle has its 
front half red, the posterior half black. Anterior legs and hind 
tarsi rufescent, base of middle tibiwe and their metatarsus testa- 


HYMENOPTEROUS FAMILY STEPHANIDA, 197 


ceous. Wings hyaline, strongly iridescent throughout, nervures 
black. 

Length 15 mm.; abdomen 10 mm.; petiole 5 mm.; terebra 
22 mm. 

Habitat: Dehra Dun, India. Nov. 1907; Lt.-Col. F. W. Thom- 
son, I.M.S. 

The sharply-defined red colour on the frons, including just the 
front half of the anterior tubercle, is unusual. May be distin- 
guished from /’. longicauda Elhott by the striate occiput, smooth 
neck of pronotum, and bordered posterior margin of head. 

A male, tentatively placed with this female, differs in the finer 
sculpture of the head and striate metapleure. The base of the 
antenne and the head are testaceous, with the vertex and part of 
occiput black. A very slender insect. 

Length 11 mm.; abdomen 7 mm.; petiole 4 mm. 

Habitat: Kangra Valley, 4000 ft. July 1889; Dudgeon. 


22. punctatus Elliott. 
Fenatopus punctatus Elliott (2), p. 73, 9. 


2. Frons arcuate rugose, vertex and occiput finely trans- 
rugose; posterior margin of head bordered; three carine between 
the posterior ocelli. Scape slightly longer than cheeks; basal 
flagellar joints normal. Pronotum strongly transrugose, with 
broad smooth posterior margin; mesonotum transrugose ; central 
lobe of scutellum smooth, with a few large punctures, the lateral 
lobes closely punctate. Petiole finely trans-striate, shorter than 
rest of abdomen, which is smooth and shining. 'Terebra as long 
as body, white-banded. Hind coxe transrugose; femora trans- 
rugose, tridentate; tibie compressed to middle. 

Rufescent ; a broad white band on each side under the eyes to 
base of mandibles. Wings hyaline, nervures rufescent. 

Length 133-163 mm.; abdomen 83-11 mm.; petiole 4-5 mm.; 
petiole 134-163 mm. 

Habitat: Burma. Two specimens in British Museum; the 
smaller with label ‘“ Birmah, F. Smith, coll.”; the larger 
‘“* Birmah ” only. 

The puncturation on the lateral lobes of scutellum is unusually 
dense. 


23. TOGOENSIS Stadlim. 


Stephanus togoensis Stadlm. (2), p. 80, ¢ ; Enderl. (1), p. 203, 
9S. Var. fasciatus Ender). 1. ¢., 2. Diastephanus togoensis 
Enderl. (2), p. 476; Morley (1), p. 113. 


a 


g. Frons arcuate rugose; vertex with three or four coarse 
ridges; occiput rather more finely transrugose, with subobsolete 
central impression. Posterior margin of head slightly bordered. 
Second flagellar joint one and a half times as long as first; third 


798 j MR. E. A. ELLIOTT ON THE 


slightly shorter than first and second together. Neck of pronotum 
laterally obliquely rugose, remainder alutaceo-punctate ; meso- 
notum coarsely punctate; scutellum smooth, with marginal punc- 
tures; metanotum longitudinally striate. Mesopleure diffusely 
punctate, upper margin polished; metapleurz and median segment 
confluent, coarsely and superficially punctate. Petiole finely trans- 
striate, as long as, or slightly shorter than, remainder of abdomen. 
Hind coxe trans-striate; hind femora tridentate. 

Black; head, base of mandibles, five basal antennal joints, pro- 
thorax, tegule and legs, except hind coxe and apex of hind 
femora, ferruginous. Wings hyaline, nervures dark brown. 

Length 8-11 mm. 

2. Similar to 3 ; terebra as long as body, spicula red-brown, 
sheaths ferruginous, apically black. Length 11 mm. 

Habitat: Bismarcksburg, Togo. LL. Conradt, 1893. 

Var. fasciatus Knderl. More slender than the type-form; 
band across basal half of third segment rufescent; posterior 
femora basally dark. 

Habitat: Johann Albrechtshohe, North Cameroons. L. 
Conradt, 1895-96. — 

Dr. Enderlein (/.¢.) places this species in his subgenus Diaste- 
phanus, but Stadelmann expressly states that the neuration is 
exactly as in /’. indicus Westw., and it is so closely similar to 
F. conradti that I incline to consider it merely a colour variety 
of that species. The different sculpture of scutellum and the 
absence of the slight smooth depression between metapleurz and 
median segment appear to be the only differences. No mention 
of the femoral teeth is made in the description of 7’. conradti. 


24, CLARIPENNIS Elliott. 
Fenatopus claripennis Elliott (2), p. 81, 2. 


2. Frons arcuate rugose, vertex subarcuate trans-striate, with 
distinct central longitudinal furrow, occiput more finely striate. 
Posterior margin of head bordered. A long, stout carina just in 
front of the posterior ocelli and two shorter ones between them. 
Seape as long as cheeks. Second flagellar joint one and a half 
times as long as first, third nearly as long as first and second 
together. Neck of pronotum coarsely, obliquely striate, the 
remainder alutaceous; mesonotum moderately punctate. Meso- 
pleurze alutaceous, basal half diffusely punctate; metapleure 
nearly smooth, with diffuse, large punctures, separated by a line 
of punctures from the median segment, which is alutaceous and 
diffusely punctate. Petiole trans-striate, its extreme apex smooth, 
as long as rest of abdomen. ‘Terebra shorter than body, black. 
Hind legs with coxe trans-striate, femora tridentate and very 
finely alutaceous, tibiee constricted to middle. 

Black; head rufescent, vertex darker; tegule, basal half of 
petiole and its apex broadly, and the second and third segments 


HYMENOPTEROUS FAMILY STEPHANID®. 799 


basally red. Anterior legs rufescent, femora centrally and tibiz 
partly darker; middle metatarsus yellowish; hind legs black, 
apex of femora, the tibie and tarsi rufescent. Wings very clear 
hyaline, apical half iridescent. 

Length 15°mm.; abdomen 9 mm.; petiole 43 mm.; terebra 
13 mm. 

Habitat: Nyassaland, Chiromo, Ruo River. R.C. Wood, 1916. 
Type in British Museum. 

The long red carina in front of the posterior ocelli and the 
sculpture of the prothorax sufliciently distinguish this species. 


25. ARATIFRONS Enderl. 
Fenatopus aratifrons Enderl. (8), p. 207, 3. 


3S. Frons finely and densely, vertex and occiput more strongly 
transrugose. Pronotum with fine striation between coarser 
rugosities, laterally anteriorly alutaceo-granulate, posteriorly 
closely longitudinally striate. Scutellum depressed, coarsely 
punctate. Mesopleure sparsely punctate, posterior margin smooth, 
with fine trans-striation, upper third with sparse yellow hairs. 
Petiole densely trans-striate, second segment smooth, basally 
longitudinally striate, apically dull. Hind coxe irregularly 
rugose, posterior fourth more regularly ; hind femora bidentate. 

Black; head bright rufo-testaceous; front coxee and anterior 
legs ferruginous yellow ; hind tarsi ferruginous, metatarsus pale 
yellow. Wings hyaline, stigma apically rounded, ochre-yellow ; 
nervures brown. 

Length 10:2 mm.; petiole 25 mm.; fore wing 6 mm. 

Habitat: Kankau, Formosa. H. Sauter. Type in Dahlen 
Museum. 


26. FLAVICOLLIS Cam. 
Fenatopus flavicollis Cam. (10), p. 359, 3. 


3. Anterior frontal tubercle smaller than the posterior, space 
between them obliquely striate; vertex arcuate striate. Meso- 
notum basally depressed and there bounded by a curved carina, 
and apical trans-suleus, central part bounded by shallow oblique 
sulci and irregularly striate. Scutellum smooth, with irregular 
marginal punctures. Metanotum (median segment) diffusely 
punctate. Petiole dull and closely striate. Hind coxe closely 
striate; hind femora tridentate. 

Black; face, clypeus, outer orbits chiefly, scape, 3-4 basal 
flagellar joints, prothorax except central black line, meso- and 
meta-pleure, and anterior legs testaceous; vertex, base, and apex 
of hind tibiz rufo-testaceous. Wings hyaline, stigma pale, ner- 
vures dark testaceous. 

Length 8 mm. 

Habitat: Dima, West Africa. A. Koller. Type in Congo 
Museum, Tervueren. 


rf 
800 MR. E. A. ELLIOTT ON THE 


DiAsrePpHANUS Enderl. 
Enderlein (2), p. 473. 


This subgenus, as defined by Enderlein, has the lowest form of 
neuration, consisting of three basal cells and a widely open radial 
cell; the median nervure rarely extends beyond the transverse 
nervure, and then only as a short stump, whereas in Yenatopus 
it extends to the full length of an outer submedial cell, or closely 
approximating that length. It occurs in the Ethiopian, Oriental, 
and Australian Regions, but has not yet been recorded from the 
Western Continent. 

The species vary in size from 6 to 29 mm. in length, and the 
colour is, on the whole, brighter and more varied than in the 
other subfamilies; a white or flavous frons with rufescent lines 
is very common. 

In preparing a table of species, I have experienced much diffi- 
culty in consequence of the want of details in some descriptions, 
in which, for instance, such important points as the sculpture of 
the head and the proportional length of the basal antennal joints 
are omitted. The colour of the terebral sheaths forms a good 
distinction in many cases, but these being broken in one species, 
renders it useless in that section. The colour of the terebra must 
always be understoed to refer to the sheaths; the spicula is 
usually rufescent. 


Table of Species. 


. Hind femora tridentate. 
Petiole as long as rest of abdomen. 
. Hind femoral teeth white. 
. Femoral teeth milk-white; frons arcuate striate, mesonotum 
and median segment coriaceous and diffusely punctate ; 
terebral sheaths Fae with broad subapical yellow 
band . ii pocancbeatasaagoacsacsseccan dle daaxmodlorn IKivett, 
(4.) 5. Femoral etn Ww ietahe ‘fronts finely rugose, mesonotum 
coarsely irregularly rugose-punctate, median segment 
cribrate punctate, terebral sheaths black......... 2. pallescens Schlett. 
(3.) 6. Hind femoral teeth not all white. 
(8.) 7. Posterior margin of head produced into a collar; femoral 
teeth brown, median segment alutaceous ...... 3. fuscidens eff. 
(7.) 8. Posterior margin of head bordered, but not collar-like. 
(10.) 9. Sheaths of terebra black; neck Ie hind femora punctate, 
the teeth black.. uy popedace Sogandusnaos.. Clb Wamu tsvasyolle 
(9.) 10. Sheaths of terebra BRR Ioomnledl aol elongate; hind femora 
trans-striate, central tooth white .................. 5. bilineatus Elliott. 
(2.) 11. Petiole distinctly longer or shorter than rest of abdomen. 
(17.) 12. Petiole distinctly shorter than rest of abdomen. 
(14.) 13. Body unusually slender, pronotum elongate and shining, 
frons densely; striate) 7... vege-nesesaieeecaceessteee Os omacules Kaett. 
(13.) 14. Body normal, prothorax not llonneseites more or less dull. 
(16.) 15. Head finely reticulate, posterior margin simple... 7. tertianus Morley. 
(15.) 16. Head rugose, posterior margin bordered ............ 8. mova-guineensis  . 
(12.) 17. Petiole distinctly longer than rest of abdomen. [Szépl. 


— 
a 
— 

me oo to 


(19.) 18. 
(18.) 19. 


(1.) 20. 
(38.) 21. 
(23.) 22. 


(22.) 23. 


(27.) 24. 
(26.) 25. 


(25.) 26. 


(24.) 27. 


(31.) 28. 


(30.) 29. 
(29.) 30. 


(28.) 31. 


(33.) 32. 


HYMENOPTEROUS FAMILY STHEPHANIDA. 801 


Metapleurze and median segment finely reticulate-rugose ; { Enderl. 
terebra shorter than body, sheaths red-yellow . 9. flavomaculatus 


Metapleurze and median segment irregularly coarsely rugose; {Schulthess. 
terebra as long as body, sheaths red-brown ... 10. rothkirchi 


Hind femora bidentate. 
Sheaths of terebra with subapical pale band. 


Sheaths broadly yellow-banded; head small; second seg- 
ment finely punctate and dull; femoral teeth yellowish. [ Enderl. 
11. flavidentatus 
Terebral sheaths white-banded. 
Hind femoral teeth white. 


Anterior tubercle obsolete, frons transrugose, neck elongate, 
slender, median segment centrally basally smooth 
12. leucodontus Schlett. 


Anterior tubercle normal, frons arcuate striate, neck normal, 
median segment cribrate punctate throughout . 13. elegans Elliott. 


Hind femoral teeth unicolorous with femora. 
Frons rugose. 


Frons with strong longitudinal carina, neck normal, meso- 


pleuree and hind femora smooth .................. 14. carinifrons Ender. 
Frons not carinate, neck short and broad, mesopleurx aciculate- 
punctate, hind femora aciculate .................. 15. nigripes Szépl. 


Frons not rugose. 

Head very small, frons alutaceous, neck granulate aciculate, 
metapleure and median segment with large hexagonal 
PUNCHINESHeetecereeeeeeeeeeci ee cee eee eee eee eee Opamurceps Emncderls 


(32.) 38. Head normal. 
(35.) 34. Frons reticulate, neck transcarinate, metapleurz finely acicu- 

late, petiole longer than rest of abdomen ...... 17. frontilinea Morley. 
(34.) 35. Frons trans-striate. 
(37.) 36. Frons arcuate striate, terebra shorter than body... 18. titdineatus Elliott. 
(386.) 37. Frons trans-striate, but not arcuately ; terebra longer than 

[NXOGKy pabaeoesbonsec: Peete eee eee oa chinensiseElliotts 
(21.) 38. Sheaths of terebra siiteaToRONE, 
(40.) 89. Body rufo-testaceous, femoral teeth white; terebra apically 

black . np odd ass cbadeda9c.n0o osancaEcHaceAsabedcdonce LAU GHAAIOGUS Ilona. 
(39.) 40. Body Shield. pinch) 
(42.) 41. Petiole only half as long as rest of oor head small; [| Enderl. 

sheaths of terebra DrOWN ...ssececcscseeeceeeecseeee Qk brevipetiolatus 
(41.) 42. Petiole much more than half as avn as eae 
(46.) 43. Terebra about as long as body. 
(45.) 44. Frons finely trans-striate; first and second fees vents of 

equal@lenctheareeee entree sue 22. trilobatus Elliott. 
(44.) 45. Frons transrugose: second eeelIne ie oneand a = times 

as long as first .. Briana tou essen . 23. dohrni Ender. 
(43.) 46. Terebra distinctly Tenger or hor ee (een oder 
(48.) 47. Terebra longer than body ; frons arcuate rugose; two pro- 

minences on posterior half of hind femora strongly 

developed .. ppdedacadaqca pea cdusenosHedobononcadanes PEE GRuerRGAS INTO. 
(47.) 48. Terebra shorten than eae 
(50.) 49. Terebra only half as long as body, Head small, frons arcuate 

rugose, median segment aciculate, laterally punctate 

25. terebellus Ender. 

(49.) 50. Terebra much more than half as long as body. 


(52.) 51. 


Anterior femoral teeth yellow, the posterior black. Head 
small; petiole very slender 2nd segment punctate. 
26. ruficollis Enderl. 


802 MR. A. E. ELLIOTT ON THE 


(51.) 52. Femoral teeth unicolorous. 
(56.) 53. Femoral teeth white. 
(55.) 54. Second flagellar joint one and a half times as long as first, 

third longer than first and second together ; mesopleure 

smooth above; median segment strongly punctate {| Ender]. 

throughouteerers teen Aran . 27. maculifemur 
(54.) 55. Second flagellar joint Laie as one as finn: canal only as 

long as first and second together ; mesopleure trans- 

striate, median segment cribrate punctate, with elongate 

triangular central smooth space ................. 28. sulcatus Elliott. 
(53.) 56. Femoral teeth unicolorus with femora. 
(60.) 57. Neck of prothorax elongate. 
(59.) 58. Second flagellar joint one and one-third as long as first, 

petiole longer than rest of abdomen; second segment 

TUATNOSE conve cede A eae enn) salomonis Westw. 
(58.) 59. Second flagellar costa fully ee as 1 ones as cc petiole 

slightly shorter than rest of abdomen ; 2nd “segment 

SATO AY cooondoodcod ce sed Sav0b0 oon aDOGeocOHoon dU nGosedsen BOs CAOGROES Sasol, 
57.) 60. Neck of BD rothorae nor sh 
2.) 61. Frons irregularly rugose; abdomen from 2nd segment finely, 

densely punctate, dull... ...e.. eee eeeeeeeeeeees) OL. Szépligetti Hnderl. 

(61) 62. Frons finely trans-striate, abdomen from 2nd segment smooth 

and polished errce-tnseeereeeeeeetece eee eee ee Sa ISUNUULLONU ICS MEN oun 


(6.) 1. Hind femora tridentate. 

(5.) 2. Frons alutaceous, first and second flagellar joints of equal 
length ; anterior legs yellow. 

(4.) 3. Posterior margin of head bordered ; neck finely rugose, semi- 
annular coarsely rugose; petiole about as long as rest of — 
abdomen ; femora finely and densely punctate. 33. similis Szépl. 


(3.) 4. Posterior margin of head simple; pronotum nearly smooth ; 
petiole distinctly longer than rest of abdomen ; ‘femora 
smooth and scarcely punctate ..................... 34. flaviceps Elliott. 


(2.) 5. Frons rugose-punctate; second flagellar joint nearly twice 
as long as first; peur nea twice as long as rest of 
abdomen yr 99. 88a HEdepdAeobue QoassebdeHooNE Ble J/UNCouerods CHiN. 


(1.) 6. Hind femora lamin, 


(8.) 7. Petiole as long as rest of abdomen; face and lower half of 
frons white; wings red and green iridescent ... 36. flavifrons Hlliott. 


(7.) 8. Petiole distinctly longer or shorter than rest of abdomen. 


(10.) 9. Petiole distinctly longer than rest of abdomen; pronotum and 
femoral teeth entirely black ........................ 87. fasciatus Szépl. 


(9.) 10. Petiole distinctly shorter than rest of abdomen. 
(12.) 11. Anterior frontal tubercle obsolete ; ON very short; both 


femoral teeth wihte ess. eereeeeccnenete » Beh flavonotatus Elliott. 
(11.) 12. Anterior frontal tubercle normal; Bcoipat Hath shang posterior 
femoral tooth black.. EA esa Tee, gut . 26. ruficollis Ender). 


1. LEUCcODON Kieff. 
Diastephanus leucodon Kieff. (4), p. 116, 2. 


2. Head subglobose; frons densely arcuate striate, vertex 
with three transcarine; occiput densely trans-striate, dull; 
posterior margin of head produced into a short collar. Basal 
flagellar joints normal. Neck elongate, densely striate ; semi- 
annular coriaceous with smooth posterior margin. Mesonotum 


and median segment coriaceous, with large, diffuse and superficial 


HYMENOPTEROUS FAMILY STEPHANIDA. 803. 


punctures. Scutellum centrally smooth and shining, marginal 
rows of punctures large and deep, lateral lobes dull. Pleura 
coarsely alutaceous. Petiole densely trans-striate, as long as 
rest of abdomen. ‘Terebra nearly as long as body, spicula red, 
sheaths red-brown, with a clear yellow ring before the black 
apex. Hind legs with cox densely trans-striate; femora 
bidentate ; tibize compressed in basal two-thirds. 

Black; head except apices of mandibles, six basal antennal 
joints, prosternum, tegule, anterior legs except femora above, 
apex of hind cox, base and apex of hind femora, basal two- 
thirds of hind tibiz, and apex of petiole clear red; space between 
anterior ocellus and posterior margin of head, anterior femora 
above and apical third of hind tibie brown; base of petiole, 
basal half of third tergite, a spot on each side of fourth and 
the hind tarsi yellowish white ; hind femoral teeth milk-white. 
Wings hyaline, stigma and nervures subhyaline. 

Length 10 mm. 

Habitat: Cameroons. Type in National Entomological 
Museum, Berlin. (Conradt.) 

Very like D. fuscidens Kieff., but differs chiefly in the 
sculpture of frons, pro- and meso-notum, and in the colour of 
the terebra and of the femoral teeth. 


2. PALLESCENS Schlett. 


Stephanus patlescens Schlett., p. 123, 2. Diastephanus 
pallescens Ender]. (2), p. 475. 


@. Frons finely and more or less distinctly transrugose ; 
occiput obconical, anteriorly rather coarsely trans - striate, 
posteriorly more irregularly rugose. Posterior margin of head 
bordered. Cheeks longer than scape. Second flagellar joint 
scarcely longer than first: third one and a half times as long as 
second. Neck elongate, very slender, finely trans-striate above ; 
semiannular moderately finely rugoso-punctate, its posterior 
margin smooth. Mesonotum coarsely and irregularly punctato- 
rugose. Scutellum finely punctate rugose. Mesopleurze smooth 
and shining above, rather dull beneath, with a few indistinct 
punctures; metapleurz and median segment somewhat coarsely 
eribrate-punctate, not separated. Petiole trans-striate, scarcely 
as long as rest of abdomen, which is feebly shining or dull. 
Terebra shorter than body, sheaths entirely black. Hind legs 
with coxe transrugose; femora finely trans-striate and dull, 
tridentate; tibize longer than femora, compressed to beyond 
middle. 

Black; face ferruginous, dull flavous towards eyes and on 
temples ; mandibles and antennz basally rufesecent. Anterior 
legs chestnut-brown; teeth of hind femora whitish. Wings 
hyaline. 

Length 11 mm.; terebra 8 mm. 

Habitat: Philippines. Type in Town Museum at Hamburg. 
Proc. Zoou. Soc.—1922, No. LIV. DA 


804 MR. E. A. ELLIOTT ON THE 


. This species resembles D. leucodontus Schlett., but the 
anterior tubercle is fully developed, second flagellar joint shorter, 
pro- and meso-notum more coarsely sculptured and hind femora 
tridentate. From D. salomonis Westw. it may be at once known 
by the conformation of the neck of prothorax and the tridentate 
hind femora. 


3. FUSCIDENS Kieff. 
Diastephanus fuscidens Wieff. (4), p. 118, 9. 


9. Head subglobose; frons and vertex irregularly rugose, 
latter with three transcarine in front; occiput trans-striate, 
dull; posterior margin of head produced into a collar. Second 
flagellar joint one and a half times as long as first; third longer 
than first and second together. Pronotum coriaceous, with 
polished posterior margin; mesonotum transrugose ; scutellum 
centrally smooth and shining, marginal rows of punctures 
superficial. Pleurz alutaceous. Median segment alutaceous, 
with large, diffuse punctures. Petiole densely trans-striate, as 
long as remainder of abdomen, Terebra as long as body, spieula 
ved, sheaths entirely black-brown. Hind legs with coxe trans- 
striate ; femora tridentate ; tibize compressed to beyond middle. 

Black; head except apices of mandibles, four basal antennal 
joints, basal half of third segment entirely, anterior legs, apex of 
hind coxee, hind femora entirely and hind tibiz basally red ; 
hind tarsi testaceous; apical joint of all tarsi black; femoral 
teeth brown. Wings hyaline, stigma and nervures subhyaline. 

Length 11 mm. 

Habitat: Cameroons (Conradt). Type in National Kntomo- 
logical Museum, Berlin. 

Very like D. lewcodon Kieff., differing chiefly in the irregularly 
rugose frons, transrugose mesonotum, and in the colour of 
terebral sheaths and hind femoral teeth. 


4. Birol Szépl. 


Fenatopus birdi Szépl., p. 5380, 2. Diastephanus 6, Enderl. 
(2), p. 475. 

2. Frons irregularly, not transversely rugose; occiput short 
and rugose; posterior margin of head bordered. First and 
second flagellar joints of equal length. Neck short, subquadrate, 
anteriorly transversely, then irregularly rugose; semiannular 
coarsely rugose. Mesonotum coarsely rugose; scutellum centrally 
smooth, otherwise diffusely punctate. Mesopleurze with large, 
dense punctures, anteriorly dull. Median segment with large 
punetures, which are confluent and form rugosities on the apical 
half. Petiole finely striate, about as long as rest of abdomen. 
Terebra slightly shorter than body, sheaths black. Hind coxe 
finely trans-striate, their femora finely and densely punctate 
tridentate, 


HYMENOPTEROUS FAMILY. STEPHANID. 805 


Black ; cheeks and temples yellowish ; mouth-parts,. base of 
antenne, anterior tibie mostly, and all tarsi yellow-red. Wings 
hyaline, stigma brown, tegulz basally yellow. 

Length 11 mm. 

Habitat: Sinbang, New Guinea. Type in Hungarian National 
Museum. 


5. BILINEATUS Elliott, 
Diastephanus bilineatus Klliott (2), p. 162, @. 


@. Head finely trans-striate; two carine between the 
posterior ocelli; all frontal tubercles distinct; posterior margin 
of head bordered. Second flagellar joint one and a half times as 
long as first; third longer than second, but shorter than first and 
second together. Neck of prothorax elongate, finely trans-striate, 
remainder smooth; mesonotum subglabrous, very diffusely punc- 
tate ; pro- and meso-pleurz very finely trans-striate ; metapleurz 
and median segment cribrate punctate, not separated. Petiole 
trans-striate, as long as rest of abdomen, which is smooth 
and shining. ‘Terebra shorter than body, sheaths rufescent, 
with a subapical band # mm. broad, before the } mm. bread 
black apex. Hind coxe and femora distinctly, their tibi less 
strongly trans-striate, the femora tridentate, tibie compressed to 
middle. 

Black; head rufeseent, mandibles except extreme apex, frons 
centrally broadly and orbits up to the level of the anterior 
tubercle flavous; point of abdomen below terebra white, 
Anterior knees pale rufescent, hind legs darker, with knees and 
metatarsi pale. Central femoral tooth white. Wings hyaline, 
iridescent ; stigma and nervures brown. 

Length 8-83 mm.; abdomen 5 mm.; petiole 23 mm.; terebra 
7mm. 

Habitat: Pusa, Bihar; G. R. Dutt, 12.v.09, Chapra, Bengal; 
Mackenzie. 

The whole of the lower part of the head is white, excepting 
the more or less narrow rutfescent lines separating the frons from 
the inner orbits. The sculpture of the head, the colour of the 
terebra and that of the femoral teeth will suffice to distinguish 
this species from all those with tridentate hind femora yet 
known. (May 1919.) 


6. graciis Kieff. 
Diastephanus gracilis Kieff. (5), p. 28, 2. 


@. Body, especially abdomen, unusually slender. Frons dull 
and densely trans-striate, arcuately round anterior ocellus ; 
vertex with five transcarine; occiput rugose, without central 
impression ; posterior margin of head simple. Second flagellar 
joint little longer than first ; third shorter than first and second 

D4* 


806 MR. E. A. ELLIOTY ON THE 


together. Pronotum elongate, shining, finely punctate, posterior 
margin rugose. Mesonotum coarsely transrugose; scutellum 
with central lobe finely punctate, with large marginal punctures, 
separated by rows of small punctures from the lateral lobes, 
which are alutaceous. Mesopleurz finely alutaceous; meta- 
pleure reticulate rugose, the impressed part alutaceous. Median 
segment finely punctate with diffuse larger punctures, and with 
a broad crenulated suleus on anterior margin.  Petiole very 
slender, finely trans-striate, much shorter than rest of abdomen, 
which is coriaceous. Terebra little shorter than body, spicula 
red-brown, sheaths black. Hind legs with cox densely trans- 
rugose; femora finely alutaceous, tridentate; metatarsus partly 
and second tarsal joint entirely white pubescent beneath. 

Black; head except apices of mandibles and occiput, four 
basal antennal joints, apex of petiole, basal half of third segment, 
apex of hind femora, the tibize and tarsi yellow or red ; posterior 
coxa and femora brown. Wings whitish, stigma dark bordered, 
nervures pale yellow. 

Length 13 mm.; terebra 12 mm. 

Habitat: Mundame, Johann Albrechtshéhe, German Hast 
Africa.. Conradt, 1906. 


7. TERTIANUS Morley. 
Diastephanus tertianus Morley, p. 108, 9. 


®. Head finely reticulate throughout; posterior tubercles 
and vertical carina small; cheeks much longer than scape; 
posterior margin of head simple. Second and third flagellar 
joints of equal length and about half as long again as first. Neck 
longitudinally sulcate, laterally obliquely carinate; semiannular 
microscopically sculptured and dull, without any smooth posterior 
margin. (Mesonotum destroyed by pin.) Scutellum dull and 
diffusely punctate. Mesosternum shining, finely alutaceous. 
Mesopleurz alutaceous and sparsely punctate; metapleure finely 
transearinate, confluent with median segment, which is finely 
reticulate and sparsely punctate. Petiole transaciculate, shorter 
than rest of abdomen, which is dull. Terebra rather shorter than 
body, sheaths black. Hind legs with the coxz narrow, dull, and 
apically transaciculate ; femora dull, tridentate; tibize longer than 
femora, constricted to middle. 

Black; head, base of antenne, prothorax, apex of petiole, base 
of second segment, basal two-thirds of third, and the legs red; 
base of hind femora and a mark inside apical third of hind tibize 
black. Wings hyaline; stigma lanceolate, centrally translucent ; 
nervures brown. 

Length 12 mm.; abdomen 7 mm.; petiole 3 mm.; terebra 
103 mm. 

Habitat: Lesapi River, Mashonaland. Nov. 1897; 


Guy 
Marshall, 


HYMENOPTEROUS FAMILY STEPHANID®. 807 


8. NOVA-GUINEENSIS Szépl. 
Penatopus nova-guineensis Szépl., p. 529, 9. Diastephanus 
nova-g. Enderl. (2), p, 476. 


2. Frons finely rugose, laterally more transversely ; vertex 
transrugose ; occiput short and rugose, with distinet central 
apical sulcus; posterior margin of then horder ed. First flagellar 
joint as long as second. Pro- and meso-notum coarsely rugose, 
neck short and broad. Scutellum only centrally smooth. Meso- 
pleuree dull, densely aciculate, with large, diffuse punctures. 
Median segment rather smooth, with large punctures. Petiole 
densely trans-striate, shorter than rest of abdomen. ‘Terebra 
little shorter than body, sheaths black. Hind legs with coxe 
coarsely transrugose; femora densely and finely alutaceo- 
punctate, tridentate. 

Black; head except vertex, base of antenne, anterior tibie and 
all tarsi yellow-red; anterior femora brown. Wings hyaline, 
nervures brown; stigma centrally yellow, tegulz black, 

Length 10 mm. 

Habitat: Stephansort, New Guinea. Type in Hungarian 
National Museum. 

Szépligetti states that in his #. nova-guineensis, birdi, similis, 
and nigripes the median nervure is prolonged outside the basal 
nervure. Hnderlein (2) gives as the definition of his subgenus 
Diastephanus: * ah “al submedian cell entirely wanting, ‘there 
being at most a tiny remnant of the anterior bounding nervure,’ 
while, in /enatopus, this nervure extends to the full extent of an 
external submedian cell. There might thus be some doubt as to 
the position of the above species; as, however, Enderlein places 


them in Diastephanus, 1 follow him. 


GS. FLAVOMACULATUS Enderl. 
Stephanus flavomaculatus Wnderl. (1), p. 205, 2. Diaste- 
phanus f. (2), pp. 474-75. 


2. Frons indistinctly arcuate rugose: all frontal tubercles 
acuminate; vertex coarsely, occiput finely transrugose, without 
central impression. Posterior margin of head slightly bordered. 
Pronotum finely rugose, laterally deeply punctate just in front 
of the polished posterior margin; neck elongate. Mesonotum 
very coarsely transrugose ; Santali punctate, with central 
smooth space. Metanotum longitudinally striate. Metapleuree 
and median segment finely reticulate rugose, separated by a fine 
raised line. Petiole rather finely trans-striate, longer than rest 
of abdomen. ‘lerebra shorter than body, reddish-yellow, sheaths 
rust-yellow, apically black. Hind coxee finely trans-striate; hind 
femora tridentate, teeth yellow. 

Black; face, cheeks, five basal antennal joints, prosternum, 
anterior legs, and hind tibie ferruginous; median line and lateral 
margins of frons, base of petiole, apex of hind femora and spots 


808 MR. KE. A. ELLIOTT ON THE 


on segments 3-5, also apex of last segment yellow. Wings 
hyaline, eres pale brown. 

Length 123 mm. 

Habitat : i ohann Albrechtshéhe, N. Cameroons. 4. Conradt, 
1896. 

There appears to be a very remarkable similarity between the 
specimens of this group of Stephanidee from Johann Albrechtshohe, 
North Cameroons, and, since several are described from single 
specimens only, and none from any sufficient number, there may 
be doubts as to the distinctness of the species. The difficulty of 
deciding this matter is greatly increased by the fact that the 
descriptions vary in the amount of the details; the relative length 
of the basal antennal joints, and especially details as to the hind 
legs and femoral teeth, are often wanting. In addition to this, 
Dr. Enderlein (1) gives a table of the African pee of 
acters 
differing from those in the detailed descriptions. vine tere~ 
bellus—median segment (a) not separated from metapleure, (6) 
separated by a raised carina; togoensis—scarcely or not at all 
separated ; flavomaculatus—neck CO obliquely striate throughout, 
(6) finely rugose; brevipetiolatus—tace (a) irregulariy raprcrell une, 
(6) finely arcuate rugose. While provisionally retaining these 
species, further information is required to settle their claims, 


10. RoTHKIRCHI Schulthess. 


Diastephanus rothkirchi Schulthess, p. 283, 9; Morley (1), 
joe dha, 2 

@. Frons irregularly and partly arcuately rugose; vertex 
with four transcarinee; occiput strongly transrugose, without 
longitudinal impression. Posterior margin of head sharply 
bordered. Scape as long as first and second flagellar joints 
together; second longer than first; third shorter than first and 
second together. Pronotum finely transrugose; semiannular 
centrally finely transrugose, laterally coarsely and irregularly 
punctate. Mesonotum coarsely rugoso-punctate; scutellum 
centrally smooth, with a few marginal ‘punetur es, lateral rows of 
punctures distinct, lateral lobes finely transrugose. Mesopleurze 
finely rugose and diffusely punctate; metapleure confluent with 
median segment, and, like it, irregularly and coarsely rugoso- 
punctate, interstices and bottom of punctures very finely trans: 
rugose. (A few lines before this, the author states ‘‘ median seg- 
ment longer than mesonotum ‘and scutellum, finely facetted, 
between the facets very finely transrugose.”) Petiole finely trans. 
striate, longer than rest of abdomen; second segment basally 
coarsely rugoso-punctate, remainder of abdomen smooth. ‘'Terebra 
as long as body, spicula red-brown, sheaths brown, paler before 
the black apex. Hind legs with coxee and femora finely trans- 
rugose, latter tridentate; teeth in three groups, the central of one 
tooth only, largest of all; the basal and apical groups ef two 
each, of which the inner is smallest ; apex of tibiz and inside of 
tarsi clothed with dense red-grey pubescence. 


HYMENOPTEROUS FAMILY STEPHANIDA, 809 


Black; head red except apices of mandibles and the occiput ; 
4-5 basal antennal joints, pronotum laterally and beneath, a basal 
band on third tergite, ventral surface of second sternite, all coxe 
and legs red; petiole basally, knees and middle metatarsi basally 
white, hind coxe and base of hind femora black. Wings hyaline, 
stigma elongate, narrow lanceolate, brownish; nervures dark 
Brown. 

Length 10 mm.; abdomen 6 mm.; petiole 3°5 mm.; wings 
3°) mm.; terebra 10°5 min. 

Habitat: Cameroons. Oberlieutenant v. Rothkirch, German 
Colonial Defence Force. 

A specimen in the British Museum (M’Langa, Nyassaland, 
S. A. Neave, 2.1. 1914) differs from the above description only 
in having a more red-brown petiole and second segment black. 

This species comes near D. szepligetit Knderl. and D. gracilis 
Kieff., but differs from both in size and sculpture. The number 
of minor denticulations between the larger hind femoral teeth 
appears to vary, but we do not possess sufficient details to decide 
whether this is an individual or specific difference. 


11. FLAvIDENTATUS Enderl. 
Diastephanus flavidentatus Ender]. (3), p. 204, 2. 


®. Head rather small, globose. Neck strongly transrugose, 
laterally more finely and closely, semiannular finely trans-striate, 
apically more closely, posterior margin laterally rugose. Meso- 
pleuree sparsely and coarsely punctate, posterior margin smooth, 
upper third finely transrugose and yellow pubescent. Petiole 
finely transrugose; second segment dull, finely punctate and 
basally indistmetly rugose. Terebra rather longer than body, 
sheaths with broad yellow band before apex. Hind femora 
bidentate ; hind tibiz and metatarsus yellow pubescent. 

Black; face, cheeks, lateral borders of frons, and a median line 
yellow ; antenne brown, basally paler. Anterior legs pale ferru- 
ginous; a streak on outside of basal fourth of front tibie, basal 
fourth of middle tibiz and basal half of middle metatarsus yellow. 
Hind legs black-brown, tarsi rufescent, femoral teeth yeilowish. 
Wings hyaline; stigma pale yellow, nervures brown. 

Length 113-14; mm.; petiole 4:1-5:2 mm.; terebra 133- 
15 mm. ; fore wing 7-83 mm. 

Habitat: Formosa, 1911; Kankau, 1912; Hoozan, 1910. 
H. Sauter. Co-types in German Ent. Museum at Dahlen and 
in Stettin Museum. 

Enderlein (/.¢.) states that the only other species in this 
subgenus having yellow femoral teeth are his D. maculifemau 
and D. ruficollis. The detailed descriptions state clearly that in 
IBY, maculifemur both teeth are white, and in D. ruficollis the 
anterior tooth is yellow, the posterior black. The present species 
differs from D. raficollis in the sheaths of terebra being yellow- 
banded, hind tibiz pubescent, and both femoral teeth yellow; from 
D. maculifemur in smaller head, longer terebra, and in the colour 
of terebral sheaths and femoral teeth. 


810 : MR. E, A. ELLIONT ON THE 


12. LEUCODONTUS Schlett. 


Stephanus lewcodontus Schlett., p. 121, 9. Diastephanus 1. 
Enderl. (2), p. 475. 

2. Frons finely, transversely coriaceo-rugose; anterior 
frontal tubercle wanting (otherwise always present); occiput 
very short, finely trans-striate, more coarsely behind the ocelli. 
Posterior margin of head simple; cheeks longer than scape. 
Second flagellar joint one and a half times as long as first; third 
longer than first and second together. Neck elongate and 
slender, obsoletely trans-striate; semiannular subnitidulous, 


with a central longitudinal trans-striate fovea. Mesonotum. 


finely and irregularly transrugoso-punctate. Scutellum entirely 
smooth. Mesopleurz smooth and shining, with a few superficial 
punctures below; metapleuree smooth and shining in front, 
rather coarsely reticulate-rugose behind. Median segment 
centrally and basally smooth, laterally with large punctures, 
which are more dense apically. Petiole very finely transrugose, 
as Jong as rest of abdomen, which is smooth and_ strongly 
nitidulous. The terebra broken off. Hind coxe slender, trans- 
striate; hind femora entirely smooth, bidentate; hind tibiz 
compressed to beyond middle. 

Black; frons rufo-testaceous, vest of head ferruginous; antennee 
basally testaceous; pronotum and legs brown; teeth of hind 
femora white, and hind tarsi white-marked. Wings entirely 
hyaline. 

Length 10 mm. 

Habitat: Sarawak. Type in Royal Nat. Hist. Museum, 
Berlin, 

The absence of the usually well-developed frontal anterior 
tubercle will distinguish this species from all others. It closely 
resembles J. pallescens Schlett. and I’. indicus Westw. It may 
be distinguished from both by the short occiput, simple posterior 
margin of head, and by the sculpture of the hind femora; from 
F. indicus Westw. it further differs in the neuraticn. 

A specimen in the British Museum, from Singapore, has 
a white-banded terebra; length of body 103 mm.; terebra 
9 mm. 


13. ELEGANS Elliott. 
Diastephanus elegans Elliott (2), p. 31, 2. 


2. Frons finely arcuate striate; vertex trans-striate ; occiput 
nearly smooth. Space between the posterior ocelli trans-striate. 
Posterior margin of head bordered. Basal flagellar joints normal. 
Pronotum very finely trans-striate and rather shining, basal 
margin smooth; mesonotum almost smooth. Central lobe of 
scutellum finely punctate. Propleure finely striate; mesopleure 
smooth and shining; metapleure and median segment cribrate 
punctate, separated by a carina. Petiole extremely finely trans- 
striate, shorter than rest of abdomen, which is smooth and 


HYMENOPLEROUS FAMILY STEPHANIDA. 811 


shining. Terebra shorter than body, sheaths white-banded. 
Hind coxe trans-striate, their femora bidentate; tibize compressed 
to middle. 

Black; head rufo-testaceous, vertex slightly darker; three 
basal joints of antenne, prosternum, tegule, and anterior legs 
rufo-testaceous, the femora rather darker, tarsi paler, middle 
metatarsus whitish; hind legs darker, with the metatarsus paler. 
Femoral teeth white. 

Length 64 mm.; abdomen 33 mm.; petiole 1; mm.; terebra 
5 mm. 

Habitat: Singapore. H. N. Ridley, 1900. Type in British 
Museum. 

A delicate insect, distinguished by the space between the 
posterior ocelli being striate, instead of, as usual, carinate, and 
by the white femoral teeth. 


14, cartnrFrrRons Ender. 
Diastephanus carinifrons Ender]. (4), p. 292, 2. 


@. Head rather small, globose; frons rugose to coarsely 
reticulate, transrugose between tubercles, from the anterior cf 
which a strong carina runs longitudinally down frons. Vertex 
strongly transrugose, behind this a median longitudinal impres- 
sion, and the occiput finely and closely trans-striate ; posterior 
margin of head bordered. Second flagellar joint one and a half 
times as long as first; third as long as first and second together. 
Neck of prothorax strongly, semiannular finely transrugose, 
posterior margin polished smooth. Mesonotum anteriorly 
irregularly transrugose, posteriorly coarsely rugoso-punctate ; 
scutellum smooth, apically diffusely punctate. Mesopleurze 
smooth, with a transrugose longitudinal impression, the apical 
third in front of the impression strongly and densely punctate. 
Metapleure and median segment deeply reticulate, scarcely 
separated. Petiole finely trans-aciculate, apical margin narrowly 
smooth and polished, longer than rest of abdomen, which is 
smooth, basally rugose. Terebra longer than body, spicula 
ferruginous, sheaths black, white-banded before apex. Hind legs 
with coxe long and slender, densely trans-striate; femora 
smooth. 

Black; head except vertex and apices of mandibles, and 3-4 
basal antennal joints ferruginous; anterior margin of frons and 
a median streak below anterior tubercle ferruginous; posterior 
margin of pronotum yellow; anterior legs red-brown; hind 
tarsi and tibie brown. Wings hyaline, nervures pale brown. 

Length 14 mm.; abdomen 94 mm.; petiole 53 mm.; terebra 
16 mm. ; fore wing 74 mm. 

Habitat : Soekaranda, Sumatra. 1 9. Dr. Dohrn. 

Not unlike D. dohrni in colour, but petiole and terebra 
proportionally longer, sheaths of latter white-banded; sculpture 
of pro- and meso-notum different. 


812 MR. E. A. BLLIOTT ON THE 


15. NiGRIPES Szépl. 


Fenatopus nigripes Szépl., p. 531, 9. Diastephanus n. 
Enderl. (2), p. 475. 


2. Frons coarsely rugose; occiput trans-striate with central 
impression ; posterior margin of head bordered. Flagellar joints 
normal. Neck short and broad, transrugose, semiannular smooth. 
Mesonotum coarsely rugose; scutellum smooth. Mesopleurz 
aciculate, with diffuse large punctures. Median segment with 
large, more or less confluent punctures, interstices aciculate. 
Petiole finely trans-striate, as long as rest of abdomen. ‘Terebra 
as long as body, sheaths white-banded before apex. Hind legs 
with coxe transrugose ; femora finely aciculate, bidentate. 

Black; head and base of antenne red; middle tibie basally 
and the metatarsus white; hind tibiz basally red-brown. Wings 
hyaline, stigma and nervures black. 

Length 20 mm. 

Habitat: Sumbava, Lesser Sunda Is. Type in Hungarian 
Nat. Mus., Budapest. 


16. parvicers Ender. 
Diastephanus parviceps Knderl. (4), p. 296, 2. 


Q. Head very small and globose. Frons finely alutaceous 
(under a 27-power lens it appears finely and densely aciculate) ; 
vertex and occiput finely and densely transaciculate, with a sub- 
obsolete impression behind. Posterior margin of head bordered. 
Second flagellar joint one and a half times as long as first; third 
as long as first and second togetber. Neck granulate aciculate, 
semiannular smooth; (mesonotum damaged); scutellum polished, 
with a few marginal punctures on central lobe, lateral lobes finely 
punctate. Mesopleure with a few coarse punctures on anterior 
half, apex finely and closely punctate. Metapleure and median 
segment coalescent with hexagonal cellular impressions, honey- 
comb-like. Petiole very finely and closely transaciculate, longer 
than rest of abdomen, which is smooth, only extreme base: of 
second segment finely punctate rugose. Terebra nearly as long 
as body, spicula ferruginous, sheaths black, white-banded raafone 
apex. Hind coxe slender, shining, finely and lightly trans- 
striate; hind femora shining smooth, with microscopic reticula- 
tions. 

Black; head (except apices of mandibles, vertex, and frons to 
anterior tubercle, which are black- brown), base of antenne, front 
tarsi, middle tibiee and tarsi ferrt uginous-yellow ; hind tibize and 
tarsi brown. Wings hyaline, nervures brown; stigma very long 
and narrow, hyaline. 

Length i mm.; abdomen 62 mm.; peticle 3? mm.; terebra 
104 mm.; fore wing 6 mm. 

Habitat: Soekaranda, Sumatra. 1 2. Dr. Dohrn. 


HYMENOPTEROUS FAMILY STEPHANIDE. 813 


17. FRONTILINEA Morley. 
Diastephanus frontilinea Morley (1), p. 109, @. 


@. Frons finely reticulate and white pubescent; vertex 
longitudinally aciculate; occiput very finely and closely trans- 
aciculate; frontal tubercles prominent and acuminate, the 
anterior rather larger. Posterior margin of head reflexed, 
Antenne with first and second flagellar joints of equal length 
and rather shorter than third. Neck multicarinate, semiannular 
smooth and shining. Mesonotum very short, shining and 
sparsely punctate. Seutellum large and flat, with the impressions 
marked oniv by two rows of three punctures in front, smooth and 
shining. Mesopleure finely alutaceous and pubescent, with a few 
scattered punctures ; metapleure smooth and finely subaciculate, 
separated by a strongly-marked sulcus from the median segment, 
which is coriaceous between very large and partly confluent 
punctures. Petiole transaciculate, longer than rest of abdomen ; 
second segment basally narrowly scabrous, remainder of abdomen 
smooth; apex of sixth segment discally acutely emarginate. 
Terebra shorter than body, sheaths white-banded before apex. 
Hind legs with coxe trans-striate; femora alutaceous, bidentate ; 
tibiz as long as femora, compressed to a little beyond middle and 
there internally excised. 

Black ; mouth-parts, inner and outer orbits testaceous ; a longi- 
tudinal line down centre of frons and the tubercles rufous; 
anterior tibie testaceous and hind tarsi red. Wings narrow, 
hyaline ; stigma lanceolate and subhyaline. 

Length 104 mm.; abdomen 54 mm.; petiole 3 mm.; terebra 
9 mm. 

Habitat: Rangpur, on border of Eastern Bengal and Assam ; 
25. vii.05. Type in Pusa collection. 


18. TRILINEATUS H]hiott. 
Diastephanus trilineatus Elliott (8), p. &1, 2. 


®. Frons and vertex very finely arcuate striate, occiput 
trans-striate; carinee between posterior ocelli subobsolete ; 
posterior margin of head bordered. Second flagellar joint half 
as long again as first; third as long as first and second together. 
Pronotum trans-striate; mesonotum diffusely punctate; scutellum 
smooth. Propleure smooth; mesopleure finely striate and 
punctate; metapleure and median segment eribrate punctate. 
Petiole trans-striate, apically smooth, as long as the smooth re- 
mainder of abdomen. ‘Terebra shorter than body, with subapical 
white band. Hind coxe trans-striate, their femora bidentate ; 
tibize compressed slightly beyond middle. 

Black; mouth-parts flavous, inner and outer orbits and frons 
centrally and two basal joints of antenne flavous; vertex and 
occiput nigro-rufescent ; legs more or less rufescent. | Wings 
hyaline. 


814 MR. E. A. ELLIOTT ON THE 


Length 9-138 mm.; abdomen 5-8 mm.; petiole 23-4 mm.; 
terebra 83-12 mm. 

Habitat: Hoabinh, Tonkin. R. V. de Salvaza; Aug. 1918. 
Two specimens in the British Museum. 

The colour of the head in this species is exactly as in 
D. bilineatus Elliott, from which it differs in the bidentate hind 
femora and concolorous teeth. 


19. cHinensis Elliott. 
Diastephanus chinensis Elliott (2), p. 78, 2. 


@. Frons very finely striate, transversely below, becoming 
longitudinal above, round the anterior tubercle and in the ocellar 
space; vertex and occiput finely longitudinally striate ; posterior 
margin of head bordered. Basal flagellar joints normal. Pro- 
notum trans-striate, semiannular more coarsely, especially laterally. 
Mesonotum coarsely punctate; central lobe of scutellum with 
diffuse large punctures, lateral lobes with the external half long1- 
tudinally striate. Propleure longitudinally striate above and 
smooth beneath ; mesopleurz alutaceous; metapleure coarsely 
punctate above, trans-striate beneath, separated by a carina from 
the cribrate-punctate median segment. Petiole finely trans- 
striate, with extreme apex smooth, as long as rest of abdomen, 
whieh is smooth and shining. Terebra slightly longer than body, 
spicula rufescent, sheaths black, with subapical white band. 
Hind coxe trans-striate; hind femora finely alutaceous and 
shining, bidentate; tibiae compressed to beyond middle, alutaceous, 
the explanate part less strongly sculptured. 

Black; head except vertex, two basal antennal joints and the 
anterior legs rufescent. Wings hyaline, nervures brown. 

Length 12 mm.; abdomen 8 mm.; petiole 4 mm.; terebra 
135 mm. 

Habitat: China; Haut Mékong,: Tong King. R. V. de 
Salvaza. Type in British Museum. 

Distinguished by the sculpture of the frons and vertex, and 
especially by the striation of the outer lobes of the scutellum, 


20. aLuTACcEUS Morley. 
Diastephanus alutacews Morley (1), p. 109, @. 


@. Body entirely alutaceous and dull throughout. Head 
finely transrugose in front; all frontal tubercles small; vertex 
wilh one carina behind basal tubercle; a few weak transcarinse 
at posterior margin of occiput, which is bordered. Antenne as 
long as head and thorax ; second flagellar joint one and a half 
times as long as first; third only as long as second; all flagellar 
joints well discreted inter se. Neck with a discal longitudinal 
suleus, apically only laterally transcarinate, semiannular basally 
explanate. Scutellum as broad as long, apically obtuse, the 


HYMENOPTEROUS FAMILY STEPHANIDA., 815 


usual impressions or rows of punctures are wanting. Meso- 
pleurze smoother than mesonotum and pubescent; metapleure 
confluent with median segment, which is explanate and slightly 
transcarinate immediately before its apex. Petiole shorter than 
the remaining segments, second basally constricted, sixth apically 
discally emarginate. Terebra shorter than body. Hind coxe as 
long as femora, which are strongly imerassate, bidentate, and 
basally angulate below ; tibiz longer than femora, compressed to 
middle and there excised. 


Text-figure 6. 


Diastephanus alutaceus. 


Rufo-testaceous; eyes, ocelli, apex of mandibles, frenum, spot 
at base of petiole, circular spot on each side of basal third of 
second segment, and a larger spot on each side of fourth black ; 
femoral teeth white; terebra rufo-testaceous, apically black. 
Wings lacteous-hyaline, with a circular pale fuscous spot in the 
discoidal cell, and another very faint one below the centrally 
infuscate stigma. 


Length 10 mm.; abdomen 6 mm.; petiole 24 mm.; terebra 
8 mm. 

Habitat: Lonely Mine, Rhodesia. H. Swale, 12.v. 1914. 
Type in British Museum. 


816 MR. E. A. ELLIOTT ON THE 


21, BREyIPETIOLATUS Enderl. 


Stephanus brevipetiolatus Knderl. (1), p. 205, 2. Dia- 
stephanus 6. Enderl. (2), p. 475. 


2. Head small; frons finely arcuate rugose; anterior 
tubercle very long and pointed, the four others indistinct. 
Vertex coarsely transrugose; occiput finely and closely punctate, 
slightly rugose; posterior margin of head bordered. First and 
second flagellar joints of equal length, third and fourth also 
equal, and twice as long as first. Pronotum smooth and dull. 
Mesonotum irregularly transrugose; scutellum centrally dull, 
not punctate, the anteriorly convergent rows of punctures deep. 
Meso- and meta-pleure and median segment finely alutaceous, 
latter not separated from metapleure. Petiole dull, only apically 
sub-transrugose, stout, clavate, and only half as long as remainder 
of abdomen. ‘Terebra as long as body, spicula yellow, sheaths 
brown. Hind coxe finely trans-striate. 

Black ; frons, vertex, and apex of mandibles black ; remainder 
of head, four basal antennal joints, and band at base of third 
seoment ferruginous ; legs ferruginous, femora basally and above 
and hind tibize centrally black. Wings hyaline, nervures pale 
brown. 

Length 8 mm. 

Habitat: Johann Albrechtshohe, North Cameroons. IL. Con- 
radt. 

The unusually short and stout petiole, together with the dull 
pronotum and scutellum, distinguish this species from all] others. 

The group of species, including D. schletterert, togoexsis and its 
var. fasciatus, flavomaculatus, and brevipetiolatus, all from Johann 
Albrechtsho6he, have much in common, and require further 
elucidation. 


22. TRILOBATUS Elliott. 
Diastephanus trilobatus Elliott (3), p. 82, 2. 


@. Frons and vertex extremely finely, occiput more coarsely 
trans-striate ; posterior margin of head very finely bordered. 
Seape slightly longer than cheeks. First and second flagellar 
joints of equal length, third about as long as first and second 
together. Pronotum transrugose, apically coarsely, basally more 
finely, with extreme base smooth; mesonotum and scutellum 
smooth, with a few coarse punctures ; mesopleurse smooth above, 
punctate below; metapleure cribrate punctate; median segment 
coarsely and irregularly punctate. Petiole shorter than rest of 
abdomen, finely trans-striate, remaining segments smooth. 
Terebra very slightly longer than body, sheaths black. Hind 
coxee basally punctate, remainder trans-striate; femora bidentate; 
tibiz compressed to beyond middle. 

Black; head beneath, inner orbits, and frons flavous; the 


HYMENOPTEROUS FAMILY STEPHANIDA. 817 


upper margin of the colour is trilobed, the central lobe extending 
to the lower tubercle, the outer lobes triangular; antennz basally 
ruto-testaceous ; posterior metatarsi white. Wings hyaline, 

Length 11 mm.; abdomen 7 mm.; petiole 3 mm.; terebra 
(ig aan 

Habitat: Hoabinh, Tonkin. R. V. de Salvaza. Aug. 1918. 
Type in British Museum. 

Differs from D. trilineatus Elliott in the shorter second 
flagellar joint, sculpture of mesonotum and mesopleure, and black 
terebral sheaths. In D. trilineatus and simillimus two descending 
rufescent streaks divide the space between the eyes into five about 
equally broad strips of colour, and the lower tubercle is entirely 
black. In D. étrilobatus the narrow central streak extends to 
the top of the tubercle and the outer streaks are broadly 
triangular. 


23. DOHENI Enderl. 
Diastephanus dohrni Enderl. (4), p. 291, 2. 


9. Head medium size; frons anteriorly regularly, posteriorly 
irregularly transrugose, an irregular longitudinal impression below 
anterior tubercle, space between tubercles arcuate rugose; vertex 
transrugose; occiput rather wavily trans-striate with central 
longitudinal impression and finely bordered. Second flagellar 
joint one and a half times as long as first: third as long as first 
and second together. Pronotum dull, indistinctly transrugose in 
front, posterior margin polished smooth. Mesonotum smooth, 
with diffuse coarse punctures and apically transrugose; scutellum 
smooth, six deep punctures on each side. Mesopleurz smooth, 
anterior third densely and finely punctate, with diffuse larger 
punctures, posterior half coarsely punctate on each side of a 
central carina. Metapleurze and median segment coalescent, 
irregularly and laterally reticulately punctate. Petiole finely 
trans-striate, apically transrugose, shorter than rest of abdomen, 
which is smooth, basally slightly rugose. Terebra as long as bedy, 
spicula red-yellow, sheaths entirely black. Hind legs with coxe 
slender, finely and densely trans-striate; femora smooth, with 
microscopic sculpture. 

Black; head except vertex and apices of mandibles, 4—5 basal 
antennal joints ferrvginous; anterior legs pale ferruginous, hind 
tibie and tarsi dark brown. Wings hyaline; nervures brown, 
stigma hyaline, apically very pointed. 

Length 18 mm.; abdomen 114 mm.; petiole 54 mm. 
18 mm.; fore wing 93 mm. 

Habitat: Soekaranda, Sumatra. 1 9. Dr. Dohrn; Jan. 1894. 

This species is not unlike D, carinifrons Enderl. in colour and 
in habitus, but differs chiefly in the structure and sculpture of 
head, shorter petivle and terebra, the latter having entirely black 
sheaths. 


; terebra 


818 MR. E. A. ELLIOTT ON THE 


24. QUADRIDENS Elliott. 
Diastephanus quadridens Elliott (3), p. 31, 2. 


@. Frons strongly arcuate rugose, vertex and occiput strongly 
trans-striate and sulcate; three carine between posterior ocelli ; 
ocellar space rugose; posterior margin of head bordered. Second 
flagellar joint one and a half times as long as first; third as long 
as first and second together. Prothorax coarsely trans-striate, 
basally more finely and extreme base smooth; mesothorax trans- 
rugose; marginal punctures of scutellum deep. Propleure finely 
striate; mesopleurze punctate above, smooth beneath ; metapleurze 
and median segment strongly cribrate punctate. Petiole finely 
trans-striate, as long as remaining smooth, shining segments. 
Terebra longer than body, sheaths black. Hind coxe and femora 
densely and finely trans-striate, latter bidentate, but the two 
prominences on basal half so unusually developed as almost to 
rank as teeth; tibise compressed to middle. 

Black; head and two basal antennal joints rufo-testaceous ; 
vertex nigrescent. Wings hyaline. 

Length 15 mm.; abdomen 10 mm.; petiole 5 mm.; terebra 
17 mm. 

Habitat: Luang Prabang, Indo-China. R. V. de Salvaza; 
5.x.1917. Type in British Museum. 

This species is characterized by the unusual development of the 
femoral subsidiary tubercles, as well as by the sculpture.of the 
prothorax, mesopleure, hind coxe, and femora. 


25. TEREBELLUS Enderl. 


Stephanus terebellus Enderl. (1), p. 204, 9. Diastephanus (2), 
p. 475. 


@. Head small; frons arcuate rugose, vertex coarsely, occiput 
finely transrugose; posterior margin of head sharply bordered. 
First and second flagellar joints of equal length, third and fourth 
also equal and each twice as long as first. Pronotum elongate, 
polished smooth, punctured only in front and round scutellum, 
which is polished smooth. Metanotum very narrow, coarsely 
longitudinally rugose. Mesopleurz polished smooth above, 
densely pubescent and diffusely punctate below; metapleure 
diffusely, posteriorly more densely punctate, separated by a slightly 
raised costa from the median segment, which is finely alutaceous, 
laterally diffusely punctate, with a fine longitudinal median line. 
Petiole finely trans-striate, about as long as rest of abdomen. 
Terebra only half as long as body, sheaths black, apically ferru- 
ginous. Hind coxee finely trans-striate. 

Black; face, cheeks, temples, 5 basal antennal joints, anterior 
femora, hind femora beneath, third and fourth sternites ferru- 
ginous. Wings hyaline; stigma and nervures pale yellowish. 
Antenne from sixth joint brown; hind metatarsus whitish. 

Length 83 mn. 

Habitat: Lolodorf, South-East Cameroons. L. Conradt. 


HYMENOPTEROUS FAMILY STEPHANIDS. 819 


This species appears to be characterized by the small head, 
smooth pro- and meso-thorax, and very short terebra. 


26. RUFICOLLIS Hnderl. 
Diastephanus ruficollis Ender}. (3), p. 205, 2 3. 


@ 6. Head small, globose. Neck strongly transrugose, 
laterally more finely; semiannular trans-striate, basally move 
coarsely and posterior margin laterally irregularly rugose. 
Mesopleurze dull, with large diffuse punctures, upper thud finely 
granulate punctate, yellow pubescent. Petiole very slender, 
finely trans-striate ; second segment dull, finely punctate, basally 
more granulate. Terebra in Q slightly shorter than body, 
sheaths entirely black. Hind femora bidentate; hind coxe 
smooth, hind tibiz and metatarsus without pubescence. 

Black; head ferruginous yellow, vertex more brown in Q, 
redder in 3 ; antenne darkish rufo-testaceous, apically brown ; 
posterior half of pronotum ferruginous; anterior coxe and legs 
pale ferruginous, all tarsi rufescent. Anterior femoral tooth 
yellow, the posterior black. Wings hyaline; stigma long and 
yellow, nervures brown. 

Length, 2, 103 mm.; petiole 3-2 mm.; terebra 103 mm. 
fore wing 6 mm. gd, 8 mm.; petiole 2°2 mm.; fore wing 
4:4—4°6 mm. 

Habitat: Formosa. H. Sauter; May-June 1912. Co-types in 
Dahlen and Stettin Museums. 

Differs from PD. flavidentatus in the entirely yellowish head, 
not pubescent hind tibie and metatarsus, black terebral sheaths 
and black postericr femoral tooth. 


27. MACULIFEMUR Ender]. 
Diastephanus maculifemur Enderl. (4), p. 294, 2. 


Q. Head rather large, globose. Frons densely and coarsely 
punctato-rugose, the rugosities run from the median line 
obliquely backwards, apically more transversely ; space between 
frontal tubercles longitudinally striate: vertex with three or 
four strong transcarine ; occiput wavily trans-striate and finely 
punctate, with narrow central impression. Cheeks buccate; 
posterior margin of head bordered. Second flagellar joint ene 
and a half times as long as first; third almost longer than first 
and second together. Pronotum densely trans-striate, more 
strongly towards apex, posterior margin polished smooth. Meso- 
notum apically strongly transrugose, dorsally with deep diffuse 
punctures and microscopic striation, a deep impression before 
scutellum, which is polished smooth, with deep marginal punctures 
and subobsolete median impression. Mesopleure polished 
smooth, diffusely punctate beneath, apically finely and densely 
punctate; metapleure and median segment separated by a 
carina between two rows of small punctures, and both with large 
hexagonal to circular punctures, honeycomb-like. Petiole very 


Proc. Zoou. Soc.—1922, No. LV. 55 


820 MR, E. A. ELLIOTT ON THE 


finely and densely transaciculate, with narrow smooth posterior 
margin, about as long as remainder of abdomen, which is smooth, 
basally finely rugoso-punctate. Terebra shorter than body, 
spicula ferruginous, sheaths entirely black. Hind coxe slender, 
shining, very finely transaciculate; hind femora smooth, shining, 
with microscopic reticulations, bidentate. 

Black; head except apices of mandibles, and three basal 
antennal joints ferruginous; posterior margin of pronotum 
yellow-brown. Basal fourth of intermediate tibiz, basal half of 
their metatarsus, a small elongate oval spot near apex of hind 
femora outside, and the femoral teeth white. Wings hyaline, 
nervures brown ; yah very long pa POOR, yellow hyaline. 

Length 13-153 mm.; aloclonaen vi- me mm.; petiole 33— 
44 mm.; terebra 10— 121 mm.; fore wing 74-84 mm. 


“Eitan: Perak, 1 @ ; Malacca, 1 on 


Text-figure 7. 


Text-figure 8. 


Diastephanus sulcatus. 


Diastephanus sulcatus. 


28. sutcatus Elliott. 

Diastephanus sulcatus Elliott (3), p. 80, 2. 

@. Face irregularly rugose, vertex ee occiput trans-striate, 
with very distinct longitudinal sulcus; three stout, curved 


carine between the posterior ocelli ; Bus: margin ‘of head 
bordered. Scape longer than cheeks. Basal flagellar joints 


HYMENOPTEROUS FAMILY STEPHANIDA. 821 


normal, Neek finely trans-striate, semiannular smooth in front, 
basally arcuate striate ; mesonotum rugose ; scutellum laterally 
strongly punctate; metanotum longitudinally carinate. Pro- 
pleure smooth; mesopleure finely trans-striate, apically punctate; 
metapleure and median segment cribrate punctate, the latter 
with an elongate triangular central basal punctate and very 
finely trans-striate and dull space, bounded by large punctures. 
Petiole trans-striate, shorter than the remaining smvoth 
segments. ‘Terebra shorter than body, spicula pale red, sheaths 
black. Hind coxe finely trans-striate, their femora glabrous, 
bidentate ; tibiz compressed to middle. 

Black ; head red, apices of frontal tubercles and the carina on 
the vertex blaek; two basal antennal joints rufo-testaceous, 
third darker red, apically black. Anterior tibize and all tarsi 
rufescent ; femoral teeth white. Wings hyaline. 

Length 13 mm.; abdomen 73 mm.; petiole 3 mm.; terebra 
11 mm. 

Habitat: Luang Prabang, Indo-China. H. V. de Salvaza ; 
29.ix.1917. Type in the British Museum. 

The sculpture of the median segment resembles that of 
D. leucodontus Schlett., which the author describes as ‘‘ basally 
and centrally smooth,” and agrees also in the colour of the head 
and of the femoral teeth. It differs in seuipture of head and 
prothorax and black terebral sheaths. 


29, SALOMONIS Westw. 


Stephanus salomonis Westw. (5), p. 128, 9. Diastephanus s. 
Morley (1), p. 107; Brues, p. 100; Elliott (2), p. 74, 2. 


@. Frons and occiput arcuately rugose, latter with slight 
longitudinal sulcus; posterior margin of head bordered. Second 
and third flagellar joints of equal length, and about one-third 
longer than first. Neck elongate, glabrous, centrally deeply im- 
pressed and laterally vallately elevated ; semiannular smooth, 
with a few apical transcarine, and a band of indistinct punctures 
before the base. Mesonotum rather densely punctate; meso- 
pleure subglabrous, with a few superficial punctures; median 
segment with large, rather dense, but not confluent pune- 
tures. Petiole rather longer than rest of abdomen, trans- 
striate; second segment basally constricted and slightly rugu- 
lose, remaining segments smooth. ‘Terebra slightly shorter 
than body, sheaths entirely black. Hind coxe trans-striate, 
hind femora alutaceous, shining, bidentate; hind tibize com- 
pressed to middle. 

Black ; face below antenne, inner orbits partly, outer orbits 
entirely pale stramineous; mandibles and anterior legs bright 
ferruginous; hind legs with coxe, base of femora and apex of 
tibia more or less black, In the type-specimen the basal half 

55* 


822 MR. E. A. ELLIOTT ON THE 


of the hind femora is black all over, but in the other above only. 
Wings hyaline. 

Length 16 mm.; abdomen 10 mm.; petiole 53 mm.; terebra 
15 mm. 

Habitat: Solomon’s Island, New Hebrides (Westwood) ; 
Solomon Islands (Woodford). 

The specimen described hy Westwood, now in the eo 
Museum, was taken during the voyage of H.M.S. ‘ Heraid’ 
1866. The abdomen is wanting, but the three-jointed tarsi prove 
it to bea female. In the same collection is a perfect specimen, 
taken by C. M. Woodford, probably about 1886, from which the 
description has been completed. 

The extraordinarily deeply-excised neck of pronotum is very 
distinctive of the species; I know of no other at all like it. 

Mr. T. C. Brues (J, c.) describes a specimen from Waiai, 
Solomon Is., which he considers to be this species. It differs in 
colour, in the legs being entirely black, except apical half of 
hind femora and basal constricted part of their tibiz, second 
flagellar joint rather longer. The peculiar formation of the neck 
is not noted, but it is said to be finely trans-rugose, mesonotum 
irregularly rugose-reticulate in front, smooth behind. Allowing 
for the per sonal equation in descriptions, and the common 
variation in the colour of the legs in insects, I think the specimen 
described is truly this species. 

Schletterer (p. 128) tentatively synonymizes this species with 
his D. pallescens from the Philippines; from it, however, 
D. salomonis differs in the above-named formation of the pro- 
thorax, the proportionate length of the basal antennal joints, and 
in having bidentate hind femora; these in D. pallescens are tri- 
dentate. 


30. CELEBENSIS Szépl. 

Frenatopus celebensis Szépl., p. 531, 9. Diastephanus Ender. 
(2), p. 475. 

Q. Frons coarsely and indistinctly transrugose ; occiput finely 
transrugose, with indistinct central impression ; posterior margin 
of head bordered. Flagellar joints normal. Neck elongate, 
smooth in front, feebly trans-striate behind; mesonotum and 
scutellum smooth; mesopleurze smooth, anteriorly rugose-punctate. 
Median segment cribrate punctate. Petiole finely trans-striate, 
nearly as long as rest of abdomen. Terebra rather shorter than 
body, sheaths black. Hind legs with coxe finely transrugose ; 
femora shining, extremely finely alutaceous, bidentate. 

Black; temples, cheeks, and inner orbits yellowish; mouth, a 
median stripe on frons, and base of antenne yellow-red; anterior 
tibiee, basal half of hind tibiz, and the tarsi brown-red. Wings 
hyaline, stigma centrally yellowish, tegule brownish. 

Length 16 mm. 


HYMENOPTEROUS FAMILY STEPHANIDZ. 823 


Habitat: Toli-Toli, North Celebes. Type in Hungarian 
National Museum, Budapest. 


31. szePLicErt Enderl. 
Diastephanus szepligetii Enderl. (2), p. 476, 9. 


©. Frons irregularly rugose; vertex with two transcarine ; 
occiput densely granulate rugose. Temples inflated. Scape 
shorter than second flagellar joint, which is scarcely twice as long 
as first; third and fourth of equal length and shorter than first 
and second together. Pronotum polished smooth; propleurz 
very finely punctate in front, centrally finely and densely acicu- 
late, smooth behind, separated from pronotum by a deep sulcus. 
Mesonotum dull, diffusely punctate, amen lorly rugose. Scutellum 
smooth, with large diffuse marginal punctures. Mesopleure 
rugose and finely punctate; metapleure coarsely reticulate 
rugose, separated by a longitudinal carina from median segment, 
which is somewhat smooth, diffusely and deeply punctate, apically 
rugose. Abdomen rough, finely and densely punctate; petiole 
shorter than rest of ‘albelonmnet, Terebra shorter than body 
(sheaths broken). Hind cox finely and densely punctate, 
posteriorly finely striate. 

Black: head ferruginous, except apices of mandibles, vertex, 
and occiput; scape, first flagellar joint, apices of femora, tibize and 
tarsi except last tarsal joint, ferruginous. Wings hyaline, stigma 
brown-yellow, nervures golden- seo as 

engule Le 5 mm. shdlannera 9 mm.; petiole 34 mm.; spicula 
13 mm.; fore wing ae mm.; expanse of wings 173 mm. 

nbn: : fom Leone, West Africa. 


32. SIMILLIMUS Elhott. 
Dastephanus simillimus Wlhott (3), p. 82, 2. 


9. Head extremely finely trans-striate; flagellar joints normal. 
Neck rather finely trans-striate, semiannular smooth; mesonotum 
punctate; mesopleuree smooth above, punctate beneath ; median 
segment and metapleure cribrate punctate. Petiole very finely 
trans. striate, slightly longer than the remaining smooth segments. 
Terebra much shorter than body, sheaths black. Hind coxe 
trans-striate; femora smooth, bidentate; tibiz compressed to 
middle. 

Black; front of head coloured exactly as in D. trilineatus 
Elliott, to which it bears a strong resemblance. Wings hyaline. 

Length 123 mm.; abdomen 74 mm.; petiole 4 mm.; terebra 
9 mm. 

Habitat: Hoabinh, Tonkin. Aug. 1918; R. V. de Salvaza. 

This species appears to differ from D. trilineatus Elliott chiefly 
in the proportional length of the second flagellar joint, in 
sculpture, and im the entirely black terebral sheaths. ‘Type in 
British Museum. 


824 MR. E. A. ELLIOT! ON THE 


33. SIMILIS Szépl. 


Fenatopus similis Szépl., p. 530, 3. Diastephanus, Enderl. 
(2), p. £75. 


3. Frons finely alutaceo-rugose; occiput short and rugose ; 
posterior margin of head bordered. First and second flagellar 
joints of equal length. Neck short, finely rugose, dull, semi- 
annular coarsely rugose, as is also the mesonotum. Scutellum 
diffusely punctured, centrally smooth. Mesopleure finely rugose, 
not punctate; median segment with large, but not confluent, 
punctures. Petiole finely trans-striate, about as long as re- 
mainder of abdomen. Hind legs with coxe finely trans-striate ; 
femora finely and densely punctate, tridentate; tarsi four-jointed. 

Black; head ferruginous, vertex nigrescent, temples and cheeks 
yellowish; anterior legs except middle coxe and the hind tarsi 
yellow-red. Wings hyaline; stigma and nervures brown. 

Length 1] mm. 

Habitat: Simbang, New Guinea. Type in Hungarian 
National Museum, Budapest. 

This species closely resembles D. biréi Szépl. from the same 
locality, and may prove to be the male of that species. 

Széepligetti states that mm this species, in D. fasciatus 3, and 
in H. wiistneti gS the hind tarsi are four-jointed. This is 
undoubtedly an error, as the said tarsi are five-jointed in all 
other known males, excepting only S. fibiator Schlett., in which 
they are three-jointed as in the typical female. 


34. FLAVICcEPS Hlliott. 
Diastephanus flaviceps Elliott (1), p. 181, 3. 


_ 6- Frons alutaceous, dull; vertex and occiput in front 
arcuate rugose, becoming transverse towards posterior margin, 
which is simple. First and second flagellar joints of equal 
length, third a little longer. Prothorax almost smooth; meso- 
notum trans-striate in front, central row of punctures and 
lateral impressions distinct ; median segment coarsely punctate. 
Petiole trans-striate, a little longer than rest of abdomen; 
which is smooth and shining. Hind coxe trans-striate; femora 
smooth, tridentate ; tibize compressed to beyond middle. 

Black; face, frons, base of antenne, and anterior legs testaceous; 
cheeks pale red, occiput dark red: apex of prothorax and the 
hind femora rufescent. Wings hyaline. 

Length 11 mm.; abdomen 74 mm.; petiole 4 mm. 

Habitat? The type in the British Museum bears a label 
“BW, Sm. coll. 79. 22.” without locality. 


35. FUSCINERVIS Cam. 
Fenatopus fuscinervis Cam. (4), p. 101, 3. 
3. Frons rugose-punctate, laterally closely trans-striate ; 


HYMENOPTEROUS FAMILY STEPHANIDA, 825 


vertex closely trans-striate, as also the occiput, which has an 
indistinct longitudinal impression. Anterior three frontal 
tubercles stout, conical; the posterior smaller and more rounded. 
Second flagellar joint not quite twice as long as first. Pronotum 
closely aciculate, basally smooth, with an apical transcarina. 
Mesonotum centrally trans-striate; scutelluam smooth. Meso- 
pleure basally and apically sparsely punctate; metapleure 
reticulate, lower half smooth, with four suboblique carine. 
Median segment strongly reticulate. . Petiole closely trans- 
striate, nearly twice as long as rest of abdomen, which is 
short ovate. Hind legs with coxe finely trans-striate ; femora 
rather dull, basally transrugose, tridentate; tibiae compressed 
to middle. 

Black; head dark red, vertex nigrescent, outer orbits yellow ; 
pronotum basally, second and third abdominal segments laterally 
testaceous ; anterior coxe, trochanters, tibie and tarsi rufo- 
testaceous ; hind tarsi red. Wings hyaline; stigma and 
nervures pale fuscous. 

Length 12-14 mm.; abdomen 53-7 mm.; petiole 33-43 mm. 

Habitat : Kuching, Sarawak. Type in British Museum. 


36, FLAVIFRONS Elliott. 

Diastephanus flavifrons Elliott (1), p. 131, 3. 

S. Frons irregularly rugose, vertex transcarinate, occiput 
trans-striate ; posterior margin of head bordered. Second flagellar 
joint nearly twice as long as first; third not quite as long as 
first and second together. Pronotum trans-striate, deeply 
impressed at apex; mesonotum centrally smooth, laterally rugose, 
lateral impressions distinct. Mesopleure finely striate; meta- 
pleurze punctate; median segment coarsely punctate. Petiole 
trans-striate, as long as rest of abdomen, which is smooth 
and shining. Hind cox trans-striate; hind femora smooth, 
bidentate. 

Black; face and lower half of frons white, the edges of the 
colour sharply defined, that of the black forming an inverted W, 
and the mandibles apically broadly black, sharply defined; frontal 
tubercles rufescent; two basal antennal joints pale red; apex of 
pronotum obscurely, second and third abdominal segments, 
anterior tibi# and tarsi rufescent. Wings hyaline, with red and 
green iridescence. 

Length 11 mm.; abdomen 8 mm.; petiole 4 mm. 

Habitat: Quop, West Sarawak. G.H. Bryant; iv.1914. Type 
in British Museum. 

This species 1s distinguished by the colour of the head; the 
black colour of the mandibles, which is unusually broad, has a 
straight margim against the white of the face, while the lower 
border of the dark upper part of frons forms a clear-cut broad 
inverted W. 


826 MR. E. A. ELLIOTT ON THE 


37. FASCIATUS Szépl. 


Fenatopus fasciatus Szépl., p. 529, ¢. Diastephanus Enderl. 
(2), p. 475. 

3. Frons finely and distinctly transrugose; occiput short and 
transrugose; posterior margin of head bordered. First flagellar 
joint subglobose, little shorter than second. Pronotum elongate, 
slender, transrugose. Mesonotum and scutellum nearly smooth. 
Mesopleure anteriorly finely rugose, indistinctly punctate. 
Median segment with large, superficial, and not very dense punc- 
tures, interstice alutaceous. Petiole densely trans-striate, longer 
than rest of abdomen. Hind coxe densely and irregularly trans- 
rugose; hind femora finely alutaceo-punctate, dull, bidentate ; 
hind tarsi four-jointed. 

Black; head except vertex, base of antenne, prosternum, 
anterior legs, hind femora centrally, hind tarsi, and apical angles 
of second segment yellow-red; tegule flavous, with black spot. 
Wings hyaline, nervures yellow-brown, stigma centrally flavous. 

Length 9 mm. 

Habitat: Sumbava. Type in Hungarian National Museum, 
Budapest. 


38. rLAvoNOTATUS Hlliott. 
Diastephanus flavonotatus Elliott (2), p. 32, 3. 


3. Frons extremely finely tvansversely coriaceo-rugose ; 
vertex with four carine: occiput short, finely trans-striate, its 
posterior margin simple. Anterior frontal tubercle alnost 
obsolete. Scape a little shorter than cheeks. Second flagellar 
joint one and a half times as long as first. Neck elongate and 
slender, the whole pronotum obsoletely trans-striate, becoming 
subnitidulous basally, with a short longitudinal fovea. Seutellum 
smooth; mesonotum nearly smooth. Pro- and meso-pleurz 
smooth, the latter diffusely punctate; metapleurze smooth in 
front, reticulate rugose behind; median segment coarsely punctate. 
Petiole very finely trans-striate, shorter than the remaining 
strongly nitidulous segments, Hind coxe slender, trans-striate ; 
their femora smooth, with two large white teeth; tibia com- 
pressed to beyond middle. 

Black; frons flavous, vertex ferruginous, occiput nigrescent. 
Antenne testaceous; pronotum subrufescent. Anterior legs 
rufo-testaceous. Hind coxe black, femora ferruginous, centrally 
paler; tibie darker, apically and the metatarsus flavous, other 
tarsal joints rufescent. Petiole black, remaining segments 
rufescent, a large circular flavous mark on each side of third 
segment near base. Wings hyaline, iridescent; nervures pale 
fuscous. 

Length 195 mm.; abdomen 64 mm.; petiole 3 mm. 

Habitat: Kuching, Sarawak. 14.v.1900. ‘Type in British 
Museum. 


HYMENOPTEROUS FAMILY STEPHANIDA. 827 


This may prove to be the male of D. leucodontus Schlett., also 
from Sarawak, with which it agrees in the obsolete frontal — 
tubercle, short oeciput, and very largely in sculpture. The round 
flavous spots on third segment are very distinctive, and I do not 
feel sure enough of the connection to unite them. 


DouUBTFUL SPECIES. 


The following species cannot be placed, in consequence of 
defective descriptions :— 


MErGISCHUS CANADENSIS Davis. 


Megischus canadensis Davis, Trans. Amer. Ent. Soc. xxiv. 
p-. 349 (1897), @. 


“ ©, 14 mm., ovipositor 18 mm. 

* Black, with cheeks beneath and posterior tarsi dirty yellow; 
four anterior legs with tibie, tarsi and apical trochanter, also 
hind ‘torent tere, pale piceous; wings hyaline, nervures and 
stigma reddish-brown: the sculpturing | is different from and 
much finer than in the preceding species (MM. floridanus, see 
below), the head, including face, is ee reticulate, very 
finely so on the occiput - : roltorness coarsely reticulate ; petiole 
irregularly transversely wrinkled; five spines of the head are 
mere transverse prominences and the anterior ocellus is in a 
broad, rather deep basin ; guides of ovipositor broken. 

“One specimen from Toronto, Canada, collected July L5th by 
Mr. 1. ©. Priddey.” 


STEPHANUS DIADEMA Fab. 

Fenus diadema Fab. MSS. Type in Mus. Kiliee. 

Stephanus diadema (Fab.), Westw. Trans. Ent. Soc. Lond. iii. 
p. 277 (1843). 

“Tam unfortunately unable to determine whether this un- 
described insect be specifially different from the last (9. bicolor) ; 
the notes which I made at Kiel simply stating that the cnus 
diadema of the Fabrician cabinet isa Stephanus with a ved head.” 

There appears to be no other reference to this species, though 
the name appears in all lists of Stephanide. 


STEPHANUS DIVERSUS Schiett. 

Megischus furcatus Brullé, Hist. Nat. Ins. Hym. iv. p. 539 
(1846); Westw. Trans. Ent. Soc. Lond. 1850-51, p. 228, ©. 
Stephanus diversus Schlett. Berl. Knut. Zeit. xxxili. p. 133 (1889), 
Qo 

‘Niger, alis fere fuscis, abdominis segmento secundo levigato, 
valvulis terebre nigris. Kem. 

“11 est noir, avec les joues roux et la base des mandibles d’un 


828 MR. E. A. ELLIOTT ON THE 


roux ferrugineux. Les ailes sont lavées de bistre et leurs 
nervures sont noires. Cette espéce ressemble beaucoup a la 
précedente (annulator=furcatus Lep. & Serv.). Elle n’en difféere, 
pour ainsi dire, que par sa taille moindre, par la couleur plus 
foneée de ses ailes et par les valves de sa tariere qui sont plus 
gréles, sans anneau blanc, et dont l’extrémité n’est pas lanceolée 
comme dans Meg. annulator- Long. du corps 0-030, de la tariere 
0:040. Hab. le Brésil; collect. de M. Serville. 

“Le nom furcatus, qui porte cette espéce, vient sans doute de 
ce que le prothorax est echancré en avant. Cette disposition 
existe dans le J. annulator, dont la partie antérieure du 
prothorax est V@un roux ferrugineux.” 

Schletterer writes :—“ Brullé has described under the name of 
annulator, a species previously named furcatus by Lepeletier and 
Serville in 1825. His fwrcatus, which is sufficiently distinguished 
from furcatus Lep. & Serv. by the entirely black terebral 
sheaths and darker wings, must be re-named : I call it diversus.” 


MEGISCHUS FLORIDANUS Davis. 

Megischus jloridanus Davis, Trans. Amer. Ent. Soc. xxiv. 
p-. 349 (1897), 2 do. 

“2 $— 20 mm., ovipositor 22 mm.; ¢ 6 mm. 

‘“* Black, with head, thorax, and tarsi ene reddish, decidedly 
soin ¢ ; guides-of the ovipositor in 2 with a broad white band 
near the apex; wings yellowish fusco-hyaline, nearer hyaline 
in d ; front, vertex, hind coxe, and petiole strongly rugose and 
more or less reticulate, distinctly so on vertex and prothorax, 
and in Q on petiole, becoming finer towards apex; front above 
antennee very rough and with five tubercles distinct, the last 
two placed between the hind ocelli; heel segment of bind tarsus 
with large, dense scopal pads. 

“Two specimens from Florida, collected by Mr. T. C. Priddey.” 


STEPHANUS FRONTALIS Klug, Westw. 

Stephanus frontalis Klug, MSS.; Westw. Trans. Ent. Soc. 
Lond. iii. p. 276 (1843) (sine deseript.). 

“Species adhue inedita, in Mus. Reg. Berolinensis conserv 
Habitat Caput Bone Spei.” 


STEPHANUS SPOLIATOR Smith. 

Megischus spoliator Smith, Journ. Proc. Linn. Soc. Lond. 
vil. p. 6 (1884), 2. Stephanus spoliator Schlett. Berl. Ent. 
Zeit. xxxiil. p. 117 (1889), 


“M. niger, mandibulis et antennarum basi rufis; thorace 
rugose; pedibus anticis et intermediis ferrugineis, tarsis posteri- 
oribus rubris ; alis hyalinis. 

‘Female. Length 7 lines. Black; the mandibles, palpi, and 
five basal joints of the antennee ferr uginous; the face rugose, the 


HYMENOPTEROUS FAMILY STEPHANIDS. 829 


front with three short acute tubercles placed in a triangle, 
behind which the head is transversely rugulose ; behind the eyes 
is a broad cream-coloured stripe, which extends to the base of 
the mandibles. The thorax rugose, with large punctures on the 
metatborax; the anterior and intermediate legs ferruginous. 
Abdomen: the first segment, which forms the petiole, trans- 
versely striated ; the rest of the abdomen smooth and shining ; 
the ovipositor more than one-third longer than the body; the 
wings hyaline, the nervures black.” 

This species is from Waigiou Island, near New Guinea. It 
appears to show affinities with D. salomonis Westw. and P. pictipes 
Roman, both from the same region, but the details are too scant 
to place it with certainty. 


MEGISCHUS TARSALIS Smith. 

Megischus tarsalis Smith, Journ. Proc. Linn. Soc. Lond. v. 
ama (USO). Ox. 

““M. niger, pedibus anticis et intermediis ferrugineis, tarsis 
posterioribus rubris, alis subhyalinis. 

“Female. Length 9 lines. Black; the head coarsely sculpturated, 
on the face transversely so; the front with a transverse ridge, 
before which are two acute tubercles touching the eyes, and a 
central, more elevated one, a littlein advance. The thorax coarsely 
punctured, the prothorax forming an elongated neck; the anterior 
and intermediate legs ferruginous; the dilated apical portion of 
the posterior tibiz and the tarsi bright ferruginous; the posterior 
coxe rugose, the femora bidentate; wings fuscous, the nervures 
dark brown. Abdomen: the ovipositor the length of the body ; 
the basal segment or petiole finely striated transversely, the 
following segments smooth and shining; the ovipositor with a 
wide fascia of white a little before the apex. Hab. Bachian. 

“This species differs from J/. coronator in the form and situation 
of the tubercles on the front of the head; the neck is much 
longer and more slender; the ovipositor is as long as, but not 
longer than the body.” 

Schletterer tentatively synonymizes this species with S. tarsatus 
Sichel, with which it corresponds in the infumation of the wings, 
length and colour of terebra, sculpture of petiole, and, to some 
extent, in the sculpture of head and thorax. Jt differs in having 
the head black, thorax coarsely punctate, and in the much 
smaller size. The details given are insufficient for certain 
identification. 


BIBLIOGRAPHY. 
Anpre (Ep.). Species des Hyménoptéres d’Europe et d’Algérie, vii. bis, 1901. 
AsHMEAD (W.H.). Classification of Ichneumon Flies, 1900. 
Biscuorr. Ergebniss d. Deutsch-Zud-Afric. Exped. 1907-08; iii. 1911. 
Buiancwarp (H.). Orbigny, Dictionnaire universelle’Vhistoire naturelle, xii. 1848. 
Botspuvau(J.A.). Voyage de l’Astrolabe, 1832. Faun. Knt. 11. 1835. 


830 MR. E. A. ELLIOTT ON THE 


Bruges (C. L.). Bulletin Museum Comp. Zoology, Harvard College, U.S. Am., lxii. 
1918. 


Brutus (A.). Histoire naturelle des Insectes—Hyménoptéres, 1846. 
Cameron (P.) (1). Biologia Centrali- Americana, ix. 1887. 
(2). Proc. Zoological Society, London, 1901. 
(3). Journal, Straits Branch, Royal Asiatic Soc. 37, 1902. 
Cones i ih : “6, BOR: 
(5). Tijdschrift voor Entomologie, Ixviii. 1905. 
(6). Nova Guinea, v. 1906. 
(7). Annals of South African Museum, v. 1906, 
(8). The Entomologist, xliv. 1911. 
(9). Proc. Linn. Soc. N.S. Wales, xxxvi. 1911-12. 
(10). Ann. Soc. Ent. Belg. lvi. 1912. 
(11). Bull. Mus. Comp. Zool. Harvard Coll., U.S. Am., Ixii. 1918. 
Cosra (Acu.) (1). Annali Museo Zool. Univ. Napoli, ui. 1866. 
(2). Rendic. Accad. Sc. Napoli, 1866. 
Cresson (Ei. T.) (1). Proceedings Eutom. Soc. Philadelphia, U.S. Am., 1866. 
(2). Trans. American Entom. Soc. Philadelphia, iv. 1872. 
(3). is a ih Ss xviii. 1889. 
Davis. Trans. Amer. Entom. Soc. Philadelphia, xxiv. 1897. 
Exitorn (E. A.) (1). Entomologist, 1. 1917. 


(2). : ii, 1919. 
(3). os li. 1920. 
(4). ap liv. 1921. 


EnpERLEIN (G.) (1). Archiv fir Naturgeschichte, Berlin, Ixyii. 1901. 

(2). Zoologische Anzeiger, xxxili. 1908. 

(3). af oo SRR OMe 

(4). Stettin, Entomologische Zeitung, Ixviil. 1906. 

(5). Entomologische Mittheilungen, ]. 1912. 
Fasricius (J.C.) (1). Entomologia Systematica, Suppl. 1798. 

(2). Systema Piezatorum, 1804. 

Forster (A.). Verhand. Naturh. Ver. d. Preuss.-Rheinlands, xii. 1855. 
Grisopo. Museo Civico di Storia Naturale di Genova; Annales, xiv. 1879. 
Junie (J.). Nouv. Méth. de classer les Hyménoptéres et les Diptéres, 1807. 
Kierrer (J. J.) (1). Bull. Soc. Hist. Nat. Metz, xi. 1904. 

(2). e A a ser. 3, ii. 1911. 

(3). Berliner Entomologische Zeitung, li. 1907. 

(4). Bull. Societé Hutomologica Italiana, xli. 1909. 

(5). Hrgeb. des Deutsch-Zud-Atric. Exped. 1907-08 ; iii. 1910. 

(6). Société Entomologique de France; Annales, Ixxx. 1911. 
Lamarcr (J. B.). Hist. naturelle des animaux sans vertebres, iv. 1817. 
Larrernue (P. A.). Hist. naturelle des Crustacés et des Insectes, xiii. 1808. 
Le Penerrer (A. L. M.) & Servirnn (A.). Encyclopédie méthodique, x. 1827. 
Monrrovuzier (P.). Société d Agriculture, Lyon, vil. 1857. 
Moruey (Ciaupr) (1). Entomologist, 1. 1917. 

(2). ap li. 1919. 
Ness as Esenpeck (C. G.). Hymenopterorum Ichneumonibus affinium mono- 
vraphiz, 1. 1834. 
Panzmr (G. W. F.). Fann. Insectorum Germanicze initia, pt. 76, 1801. 
Roman (1). Arkiv for Zoologi, Stockholm, ix. 1915. 
(ONG m a xi, 1917. 


Say (T.). Keating’s Narrative of Expedition to the Sources of the St. Peter's 
River &c., 1823, ii. Appendix, 1825. 


HYMENOPTEROUS FAMILY STEPHANIDA. 831 


Saussure (H. pz). Mission Pavie Indo-Chine, 11. 1901. 
SemEnow. Hore Soc. Entom. Rossice, xxv. 1891. 
ScunErTEReRr (A.). Berliner Entomologische Zeitschrift, xxxiii. 1889. 
Scuuurz (W. A.) (1). Spolia Hymenoptera, 1906. 

(2). Berl. Ent. Zeit. li. 1906-7. 

(8). Zoologische Annalen, Wiirzburg, iv. 1911. 
Scuuntuess (Dr. A.). Deutsch. Entomologische Zeitschrift, 1914. 
StcHen (J.) (1). Société Entomologique de France; Annales, ser. 3, vi. 1860. 


(2). ” * 50 a ser. 4, v. 1865. 
Sairu (F.) (1). Journ. Proc. Linn. Soe., Zool. 11. 1857. 

(2). 55 5 ty) veglS6L. 

(3). xs % » vil. 1864. 


(4). Trans. Ent. Soc. Lond. ser. 3, 1. 1862-64. 
SrapErmMann (H.) (1). Berliner Entomologische Zeitschrift, xxxvii. 1892. 
SzppLicgettr. Termés. Fiiz. xxv. 1902. 
Wesrwoop (J. 0.) (1). Griffith, Animal Kingdom, Class Insecta, ii. 1832. 
(2). Annals & Mag. of Natural History, vil. 1841. 
(3). Trans. Ent. Soc. Lond. iii. 1841-43. 
(4). op 3 3 ser. 2, 1. 1851. 
(5). Thesaur. Ent. Oxon. 1874. 
Zscuacu (J. J.). Museum N. G. Leskeanum, 1789. 


THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 833 


EXHIBITIONS AND NOTICES. 
May 9th, 1922. 


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


The Secretary exhibited, and made remarks upon, a medal 
struck to commemorate the 150th Anniversary of the Royal 
Academy of Belgium. 


Mr. F. Martin Duncan, F.Z.S., exhibited, and made remarks 
upon, a series of cinematograph-filis illustrating various stages 
in the life-history of the Wood-Ant (Formica rufa) and of the 
Common Wasp (Vespa germanica), and asked the Society to 
accept the series for its Zoological Film Library. 


Dr. Cas. F. Sonnrac, F.Z.S., exhibited, and made remarks 
upon, a series of specimens and lantern-slides illustrating some 
points in the Anatomy and Physiology of Whales. 


May 23rd, 1922. 


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


The Secretary read the following Report on the Additions to 
the Society’s Menagerie during the month of April, 1922 :— 

The registered additions to the Society’s Menagerie during the 
month of April were 369 in number. Of these 95 were 
acquired by presentation, 105 were deposited, 156 were purchased, 
3 were received in exchange, and 10 were born in the Menagerie. 

The following may be specially mentioned :— 

A pair of Abbott’s Duikers (Cephalophus spadix), new to the 
Collection, from Usambara, presented by H.H. Sir Horace Byatt 
on April Ist. 

1 Gambian Pouched Rat (Cricetomys gambianus), from French 
West Africa, presented by Curtis G. Lampson, Hsq., on April 3rd. 

1 King Guereza (Colobus polycomus), from West Africa, 
deposited on April 28th, 


834 THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 


2 Vociferous Sea-EHagles (Haliaétus vocifer) and 2 Tufted 
Umbres (Scopus wmbdretia), from Tanganyika Territory, pre- 
sented by H.H. Sir Horace A. Byatt, K.C.M.G., on April Ist. 

2 Tufted Umbres (Scopus wmbretta), presented by Mr. J. D. 
Loveridge, F.Z.S., on April Ist. 

2 Australian Cat-birds (Hluradus viridis), from New South 
Wales, new to the Collection, purchased on April 21st. 

6 Laughing Kingfishers (Dacelo gigantea), from South Australia, 
purchased on April 13th. 

4 Kagus (Rhinochetus jubatus), from New Caledonia, pur- 
chased on April 21st. 

2 Tawny Frogmouths (Podargus strigoides) and 5 Regent 
Birds (Selewcides chrysocephalus), purchased on April 21st. 


Mr. R. 1. Pococr, F.R.S., F.Z.S., exhibited, and made remarks 
upon, a living example of the rare Bush-Dog Speothos venaticus. 


Mr. EK. G. Boutencer, F.Z.S., and Mr. F. Martin Duncan, 
F.Z.S., exhibited, and made remarks upon, a cinematograph 
record which they had taken illustrating the life-history of the 
Axolotl (Amblystoma tigrinum). 


Mr. D. Sers-Smiru, F.Z.S., exhibited, and made remarks upon, 
a series of photographs he had taken of some recent important 
additions to the Society’s collection of Birds. 


The Rev. H. N. Hurcurtnson, M.A., F.Z.8., exhibited, and 
made remarks upon, a plaster cast of a model reconstruction of 
the marine reptile Peloneustes philarchus, a Pliosaur from the 
Oxford Clay, stating that he had received valuable assistance 
from Mr. Ed. Godwin in completing the model. 


June 13th, 1922. 
Prof. KE. W. MacBripg, D.Sc., LL.D., F.R.S., Vice-President, 


in the Chair. 


The Secretary read the following Report on the Additions to 
the Society’s Menagerie during the month of May, 1922 :— 

The registered additions to the Society’s Menagerie during the 
month of May were 489 in number. Of these 336 were acquired 
by presentation, 14 were deposited, 122 were purchased, 4 were 
received in exchange, and 13 were born in the Menagerie. 

The following may be specially mentioned :— 

1 Bush-Dog (Speothos venaticus), from Northern Brazil, pre- 
sented by H. J. Lynch, Esq., on May 23rd. 


THE SECRETARY ON THE SOCIETY’S AQUARIUM. 835 


1 Californian Sea-Lion (Ofaria californica), born in the 
Menagerie on May 27th. 

2 Huropean Beavers (Castor fiber), from the River Nidelven, 
Norway, purchased on May 23rd. 

A collection of birds from Colombia, presented by Mr. W. K. 
Pomeroy. F.Z.8., on May 6th, including 2 Colombian Red- 
rumped Hangnests (Cassicus uropygialis), new to the Collection ; 
2 Crimson-backed Tanagers (Rhamphocalus dimidiatus) ; 4 Colom- 
bian Crested Colins (Hupsychortyx leucopogon), and others. 

1 Sclater’s Curassow (Crax sclatert), 1 Crested Curassow (Crax 
alector), and 2 Purplish Guans (Penelope purpurascens), from 
Brazil, presented by Dr. Paulo de Silva Prado on May 12th. 

3 Leadbeater’s Uockatoos (Cacatua leadbeateri), hatched in the 
Menagerie on May 19th. 

1 Shovel-footed Ceratophrys (Ceratophiys cultripes), from Morro 
Velho, Brazil, new to the Collection, presented by George Chalmers, 
C.M.Z.8., on May 29th. 


The Secretary briefly described the technieal side of the 
Council’s scheme to establish an Aquarium in the Society’s 
Gardens. 


Proc. Zoou. Soc.—1922, No. LVI. 56 


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PAPERS. 


Page 


A Study of the Remarkable Tortoise, Testudo loveridgii Blgr., and the Morphogeny 
of the Chelonian Carapace. By Joan B. Procrsr, F.Z.8S. (Plates L-III.; Text- 
figures 1-21; and Table.) 


Notes on the Anatomy of Cacopus systoma, an Indian Toad of the Family Engystoma- 
fide. By D. W. Duvanesan, M.A. . (Text-figures 1-19.) ......-.......-.--2:. 


A Review of the Cetacea of the New Zealand Seas.—I. By W. R. B. Otiver, F.L.S., 
ewe Swe (Plated LTV.) 62. ...-. 


On the Dental Characters of certain Australian Rats. By Prof. F. Woop Jones. 
Me aces Se) ata eine nswnl et ate evel ne loth e canine) VellayaUn le. alielal slay ai¢ wle)nic)e'eaielaieim eae 
On the Structure of the Enamel in the Primates and some other Mammals. By 
J. THornton Carter. (Plates I—-VII.) 


Miocene Proboscidia from Baluchistan. By C. Forster Coorrr, M.A., F.ZS. 
(Plates L-IV.; Text-figures 1-12.) .... 


On Commerson’s Dolphin and other Species of Cephalorhynchus, By Sir S. F. 
Harmer, K.B.E., F.R.S., F.Z.S. (Plates I-III.) 


ee cere ee eS ee eo oe ee oo ee OOH ds OD 


i 


The Comparative Anatomy of the Tongues of the Mammalia.—VII. Cetacea, Sirenia, 
and Ungulata. By Cuaruzs F. Sonnrac, M.D., F.Z.S. (Text-figures 25-30.) .... 


The External Characters of Scartwrus and other Jerboas, compared with those of 
Zapus and Pedetes. By R. I. Pocoox, F.R.S., #.Z.S. (Text-figures 29-37.) ...... 
A Revision of the Isopod Genus Ligia (Fabricius). By Haroun Gorpon Jacxgon, 
WNIBS eee hee San Clabes ScD) acre cree ctalreinis: ote opetciia ote eitase cumrenn fete aie crate charsieccrsale 


Monograph of the Hymenopterous Family Stephanide. By Ernest A. Exxrott, 
B.Z.S., F.E.S. (TLext-figures 1-8.) 


CeCe ere re ems e st CF EH se ser. ~c eo seeder esnse ee 


483 


527 


557 


599 


609 


659 


683 


LIST OF PLATES. 


1922, Parr III. (pp. 483-835). 


Page 
J. B. Procrr: Pl. I. Testudo loveridgit Blgr. (} nat. size.) .. 483 
Pl. If. Radiograph of 2 type-specimen ...... 483, 
PI.III. Radiograph of § type-specimen ...... 483 
W. R. B. Oxiver: Pl. I. Neobalena marginata «. 02.2.2. semen 557 
Pl. Il. Cephalorhynchus hectori and Kogia 
ONGUICEDS 6 vi cin id ns ea eee 557 a 
Pls. IIL-IV. Mesoplodon bowdoini ..........+. 557 
J. Tuornron Carrer: Pls. I-VI. Structure of the Enamel in Pri- 


C. Forster Coormr: Pls. I—-LV. Miocene Proboscidia from Balu- 
chistan “isisesscs ee ee OOS 


Sir 8. F. Harmer: Pl. I. Cephalorhynchus commersonii .......... 627 

Pl, dee; commersonii aud C. heavisidet........ 627 

Pl. IIT. Species of Cephalorhynchus..........-. 627 

H. G. Jackson : Pls. I.-II. The Isopod Genus Ligia (Fabricius).. 683 
NOTICE. 


The ‘ Proceedings’ for the year are issued in four parts, paged consecutively, 
so that the complete reference is now P. Z. 8S. 1922, p.... The Distribution 
is usually as follows :— 


Part I, issued in March. 
ee lels .; June. 
vs lel By September. 
Re oh tat EA incense December, 


‘Proceedings, 1922, Part II. (pp. 205-481), was published on June 28th, 
1922. 


The Abstracts of the ‘ Proceedings,’ Nos. 229-231, are 
contained in this Part. 


The dates of Publication of ‘ Proceedings’ 1830-1858 will be found in the 
‘ Proceedings’ for 1893, page 436, 

The dates of Publication of ‘ Transactions’ 1883-1869 will be found in the 
‘ Proceedings’ for 19138, page 814. 


PROCEEDINGS 


OF THE 


GENERAL MEETINGS FOR SCIENTIFIC BUSINESS 


OF THE 


ZOOLOGICAL SOCIETY 
OF LONDON. 


y of Ce 
: sey 


Sy 
JUNS - 19237 


1922. 


S 


PART IV. 


CONTAINING Paces 837 to 1276, wiry 13 Puatrs anp 
. 145 'TExt-FIGURES. 


FEBRUARY 1923. 


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


LONDON : 
MESSRS. LONGMANS, GREHN, AND CoO., 
PATERNOSTER ROW. 


Oral [Price Twelve Shillings.) me 


LIST OF CONTENTS. 
1922, Parr IV. (pp. 837-1276). 


EXHIBITIONS AND ‘NOTICES. 


: Page 

‘nu Secretary, Report on Additions to the Society’s Menagerie during the months of 
June to September, 1922 0! ate...) wie cate clarence eve tviepe ete /2)cia/-1-gmee ferry seagate eee 1278 
Tne Secretary. Exhibition of photographs of Zebras and Oryx from Kenya ......... . 1275 
Mr. BE. T,. Newron, F.R.S., F.Z.S. Exhibition of tanned skin of a Frog .............. 1275 

Mr, CO. Tate Recan, F.R.S., F.Z.S. Exhibition of specimens of leather made from Sharks’ 
SIGIINSy eeye ere stele aie ate tartare day ete tie teloe were Seisid isis jac! lewis tore inlangye starerelaee aie See eee 1275 

Prof. G. Exaior Sutrn, M.D., F.R.S., F.Z.S. Exhibition of photographs of a fossil 
Tooth (Hesperopithecus) from Nebraska ..... i pals velsitol oteausiclater seis) o ieee eae 1275 


Dr. A. Sirrx Woopwarp, F.R.S., F.Z.S. Exhibition of drawings and photographs 


comparing teeth of a Bear, Chimpanzee, and Hesperopithecus ......+s++ee+s++++-- 1275 
Tue Secretary. Report on Additions to the Society’s Menagerie during the month of 

(OLE O Sere LS IE See IN ana entamat a dicls Wey Oe atimh asses Bis S RekD Cows A ee a cht Roe a ieee aes 1276 
Prof. E. W. MacBripz, F.8.8., F.Z.S.  Hxhibition of lantern-slides bearing upon 

Dr. Kammerer's experiments on Amphibia <0 -t)-«.0.. de esse anata idee naeiiete 1276 
Dr. A. Surrn Woopwarp, F.R.S., F.Z.S8. Exhibition of a skull and tusks of a Mammoth 

ie OITUR SLI OUL EN eae ye arate aoe eTocs SUES te: ba) Soe aco leucoee s esetn aneee 1276 


Mr. D. Suru-Smity, F.Z.8. Exhibition of the shed lining of the gizzard of a Hornbill .. 1276 


PAPERS. 

Page 

37. Notes on Hast African Birds (chiefly nesting habits and stomach contents) collected 
1915-1919" By Arruur Lovertpen, WHS) © MLZIS) 22 ona. cei s e seie ODT 

38. On the Myology and Classification of the Wombat, Koala, and Phalangers. By 
Cuarues F. Sonnrac, M.D., F.Z.S. (Dext-figures 31-43.) .................... 863 

39. Experimental Evidence that Oommensalism may be beneficial to Crustacea. By 
TOO ATR TE} Jeon, IDAStese IIIS TAR oo os soda noon ou Goan dn onele eons . 897 

40. Description of a New Lizard of the genus Chaicides, from the Gambia, living in the 
Society’s Gardens. By E.G. Bounmnomr, ZS. .-..-...-.-.-.0- cee ee 899 


41, Onthe Parasitic Nematoda collected from Mammalian Hosts which died in the Gardens 
of the Zoological Society of London during the years 1919-1921; with a descrip- 
tion of three new Genera and three new Species. By G. M. Vuvers, M.R.C.S., 
DEIR ENOL. IAs (CUieqmatstnesss NENG) sagaacaadaacacgodoodso obo cLsoacsseace | SO 


Contents continued on page 3 of Wrapper, 


MR. A, LOVERIDGE ON EAST AFRICAN BIRDS, 837 


Oils Notes on East African Birds (chiefly nesting habits and 
stomach contents) collected 1915-1919. By Arruur 
Loveripes, F.E.S., C.M.Z.S. 


[Received October 22, 1922: Read November 21, 1922. ] 


The following notes are based on a collection of about a 
thousand skins representing 51 families, 216 genera, and 402 
distinct species, besides several hundred eggs ; it does not include 
a collection made in Uganda, but refers to those from the coastal 
colonies—Kenya Colony (then B.E.A.); Tanganyika Territory 
(then G.E.A.); Portuguese East Africa; Natal. 

No object would be served in publishing the whole list of 
species, and the following field-notes selected from my diary 
relate to the nesting habits, stomach contents, Kuropean 
migrants, and other items of general interest. 

The principal localities mentioned in the following pages 
are :— 

Kenya Colony —Bissel, Fort Hall, Frere Town, Kabete, 
Kedong Valley, Kijabe, Lake Naivasha, Nairobi, Voi. 

Tanganyika Territory.—Dar-es-Salaam, Dodoma, Kongwa, 
Longido West, Makindu (Msiha River), Morogoro, Mpwapwa, 
Mt. Meru, Ngari Mtoni, Soko Nassai, Tabora, Uluguru Mts. 

Portuguese Hast Africa.—Lorenzo Marques, Lumbo (mainland 
opposite Mozambique). 

Natal.—Durban. 

_I should like to take this opportunity of thanking Dr. V.G. L. 
van Someren, Dr. Hartert, and Mr. Arthur Goodson for 
undertaking the identification of many of the specimens. To 
Dr. Hartert, | am also indebted for his kindness in correcting 
proofs and seeing these notes through the press. 

Some half-dozen new species or races contained in the collection 
have been described by Dr. van Someren, 


NECTARINIA REICHENOWLI Fisch. 

This Sun-bird is particularly associated in my mind with the 
red inflorescences of Leonitis leonurus, of which it is particularly 
fond. Clinging to the stem of one of these plants, the bird 
rapidly runs Around the circlet of red flowerlets, in quick 
suecession driving its beak into the tube-like bases of tHe flowers 
and passing on to the next till that particular plant is 
exhausted (Kijabe, 26. vi. 15). 


CHALCOMITRA SENEGALENSIS INESTIMATA Hartert. 


The species has a very wide distribution in Kast Africa, having 
been met with in strikingly different types of country. Tn 
captivity it does well on sugar slightly moistened ; to ae I added 

Proc. Zoou. Soc.— 1922, No. LVII. Dil 


838 ; MR. A. LOVERIDGE: NOTES ON 


a few ants, for without such additional insect diet they do not 
thrive for long. It was a surprise how quickly the bird took to 
this form of food, sitting on the edge of the bowl and sipping at 
the mixture, with occasional intervals in which it gave vent to 
the characteristic whistling chirp. I have seen the Scarlet- 
breasted Sun-bird hovering in the air and taking termites on the 
wing (Morogoro, 10. iv. 18). 

Nest with one egg (Morogoro, 19.11.17); nest with two eggs 
(Morogoro, 13.111. 17); building (Morogoro, 9. x11. 17). 


CHALCOMITLA VERREAUXI A. Sm. 


Verreaux’s Sun-bird was found nesting on the Bluff at Durban 
(21. xii. 14); the two eggs were of a chocolate colour. 


ANTHREPTES COLLARIS Vieill. 


The Natal Collared Sun-bird also found nesting on the Bluff 
with two eggs (Durban, 21. xii. 14). 


MoraciInuA VIDUA Sundev. 


The East African Pied Wagtail is one of the sweetest songsters 
in the country, and quite rivals a canary. Its favourite perch 
when singing appears to be the ridge of a roof; in such a situation 
I have seen three or four of these wagtails in company with a 
row of swallows (Morogoro, 14.11.17). 

A nest containing three hard-set eggs was found on the brink 
of a waterfall at Kabete—such a waterfall and surrounding scenery 
as one might find in the Vale of Neath (Kabete, 26. v. 15). 


MoraciLLA FLAVA CAMPESTRIS Pall. (cf. Ibis, 1921, p. 666). 


During the month of November 1918 large flocks of Yellow 
Wagtails were to be met with at Dar-es-Salaam. They were 
particularly plentiful on the land near the sea-front, where they 
ran at the heels of the transport oxen which were pastured there. 


Macronyx CRocEvs Vieill. 


A nest of the Yellow-breasted Pipit containing a single egg 
which disappeared the following day (Nairobi, vii. 15). 


SERINUS STRIOLATA AFFINIS Richmond. 
Nest with three eggs (Kabete, 24. v. 15). 


PASSER GRISEUS SUAHELICUS Neum. 


A nest of the Swahili Grey Sparrow containing three eggs in 
the masonry of a house. Both nest, eggs, and situation similar 
to those of the English Sparrow (Kongwa, 25. iv. 17). 


HYPOCcHERA ORIENTALIS Reichw. 


Found a freshly-killed male lying dead in the bush without 
any visible signs of violence ; one leg was missing from the knee, 


EAST AFRICAN BIRDS. 839 


but this was evidently an old injury, as it had long since healed. 
Whilst examining it, my attention was attracted by the 
demonstrations of a Shrike (Harpolestes senegalensis orientalis), 
and a short search discovered its nest with the hen bird sitting 
on the clutch. Was it possible that the Shrike had killed the 
Weaver, which had been unable to escape owing to its crippled 
condition ? (Morogoro, 29. iii. 17). 


COoLIUSPASSER ARDENS TROPICUS Reichw. 


Nests of the Red-collared Whydah with three eggs in each 
were found on 9.iv.17 and 1.vi.17, and many more between 
those dates, the species being quite common at Morogoro 
(Morogoro, 1917). 


CoLIUSPASSER LATIOAUDA Licht. 


Many nests were found on the plains at Nairobi between 
22.v.15 and 5.vi. 15, containing three eggs, which comprises the 
clutch. Two clutches of three eggs were found at the same place 
on 7.v. 19. (Nairobi). 


CoLIUSPASSER EQUES Hartl. 


Four nests with two eggs in each were found between 13 & 
23.i11.17. It would appear that two is often the full clutch. 
The nest is similar to those of other members of the genus. At 
first a framework in the shape of an oval is made between two or 
more stout grasses at a height of 3 feet from the ground, or 
thereabouts. A loosely-built domed grass net is woven on to the 
framework, and the eggs laid in the bottom of the nest without 
lining of any kind (Morogoro, 1917). 


CoLIUSPASSER JACKSONI Sharpe. 


The males of Jackson’s Whydah and the foregoing species 
dance round and round the female as she sits on a tuft of grass, 
the object of the dance being to display the extraordinary long 
black tail-feathers which are put on for the breeding season. 
The dance is really a hop, and it is quite ridiculous to see the 
birds appearing and disappearing as if shot into the air by some 
unseen spring. Round and round goes the bird, till the grass is 
trodden down in the vicinity of the central tuft to such an extent 
that one could hardly credit that it was the work of so small a 
creature. Doubtless they resort to the same spot day after day. 
The dancing rings, about two feet in diameter, are common 
enough, and become almost bare of grass. The lengthy tail 
seems to hamper the bird in flight as it leaves the grass or 
bushes; occasionally one is seen with its tail at right angles to 
its body as it sits on a branch or briar. Unwieldyas they appear, 
attempt to catch one, and you always find you just fail; at the 
eleventh hour the owner of the tail manages to get under way 
(Nairobi, 1. v. 9). 

57* 


840 MR. A. LOVERIDGE: NOTES ON 


PYROMELANA XANTHOMELANA Riipp. 

Nesting in company with P. migroventris and P. flammiceps. 
Nests containing three eggs each found on 20,111.17 and 5.iv.17 
(Morogoro, 5.iv. 17). 


PYROMELANA NIGRIVENTRIS Cass. 

Nests of the Black-bellied Bishop-bird with full clutches of 
three eggs were found from 14.11.17 to 30.11.17. So common 
were these nests in places, that as many as thirty could be found 
in about the same number of square yards. The most favoured 
locality where I met with them was on the edge of a belt of dense 
thorn-bush scrub at Morogoro. Sharp spear-grass mingled with 
other kinds grew rank to one’s shoulders, whilst scattered thorn- 
bushes obscured by the luxuriant vegetation waited to entangle 
the feet of the unwary. Several species of Yellow Weavers flew 
about in the trees from which their pendent nests were hung ; 
below were the agitated Bishop-bicds, whose scarlet-and-black 
plumage showed to best advantage against the rich green of the 
freshly-grown grass. Many other birds frequented the spot, and 
joined their voices to the tumult which greeted the intruder. 
The nests of the Bishop-birds were so similar to those of the 
Ooliuspasser just described that a fresh description is unnecessary 
(Morogoro, 14. 11.17). 

On one oceasion (20.ii.17) when visiting this spot, I found a 
dead female Black-vented Bishop-bird sitting upon three eggs 
containing live young! A few ants were crawling about the eyes 
and mouth of the dead bird, but it was so fresh that I was able to 
leave it twenty-four hours before skinning. Seventeen days 
later I examined the same nest and found two more eggs in it 
(9. 111.17), so it appeared as if the male had installed another mate 
in the nest. The death of the bird may also be explained by the 
presence of a Green Mamba (Dendraspis angusticeps), which was 
lying sunning itself on a thorn-bush not 20 feet away. It is 
possible to suppose that it had bitten the bird, which was just 
able to flutter back to its home before succumbing to the effects 


of the poison (Morogoro, 9.111. 17). 


PYROMELANA FLAMMICEPS Swans. 

The eges of the Fire-ecrowned Bishop-bird are the same hedge- 
sparrow-egg blue as the preceding species, but are readily 
distinguishable by their slightly larger size. Nests containing 
full clutches of three eggs were found between 23.ii1.17 and 
5, iv. 17 (Morogoro). 

QUELEA INTERMEDIA Reichw. 


Nest with three eggs (Nairobi Plains, 7. v. 19). 


QUELEA CARDINALIS Hartl. 
Four nests of the Cardinal Finch containing three eggs each 
were found (Nairobi, 7 & 8.v. 19). 


EAST AFRICAN BIRDS. 84] 


SPERMESTES sp., either CUCULLATA Swains. or scuTATA Heugl. 

Green-headed Mannikin. Two and four eggs respectively 
(Kabete, 24.v.15); two eggs (Soko Nassai, 22.v.16); one egg 
(Morogoro, 30.iv.17) ; two eggs (Nairobi, 7. v.19). 

I have frequently found the hen-birds sitting on their eggs in 
the deserted nests of other species of Weaver, which they do to 
save themselves the trouble of building. A nest was situated in 
a very exposed thorn-bush on an exposed hillside at Kabete. 
Two of the eges were perfectly fresh and two were perfectly bad, 
somuch so that they exploded when pricked with a pin ; the fifth 
egg was empty, with a very small hole in its side such as might 
have been made by a bird’s claw, and the contents removed by ants. 
Another of these weaver-finches was sitting on its eggs in the 
nest of a weaver (Ploceus xanthops?) which had been built in 
sedges growing in the waters of a swamp at Soko Nassai; yet 
another had adopted the retort-shaped nest of another species of 
Ploceus, which was situated in a banana-palm, about 10 feet from 
the ground, growing on the banks of the river at Morogoro. 


UR2GINTHUS BENGALUS ANGOLENSIS Linn. 


One egg of the Blue-breasted Waxbill in a nest (Lumbo, 
28. x. 18). 


URGINTHUS NIASSENSIS Reichw. Niassa-land Bengali. 

On the outward voyage to East Africa, when at Lorenzo 
Marques, | made a note in my diary that a small blue bird had 
built its nest in a thorn-bush just above that of a wasp, and 
speculated whether it had been done for protection. I forgot 
the incident entirely until indexing my notes three years later. 
_The species was probably U. 6. angolensis. ‘Two years later, at 
Morogoro, | found many nests of U. niassensis. In nineteen 
cases out of twenty these were built over the paper nests of wasps, 
so that one could not approach one’s hand through the thorns 
before the wasps began to threaten and fly round.. 1 watched a 
pair of these birds building over a wasp’s nest, and so am certain 
that it is not the wasps which build beneath the bird’s nest for 
the sake of shelter from the rain, though doubtless the arrange- 
ment is mutually beneficial. At Mombasa, I have seen the same 
thing with U. b. brunneigularis, but at Nairobi, where one of 
these charming little blue weavers is very common, I never 
recollect having seen its nest above a wasp’s. The same at 
Lumbo, where U. 6. angolensis was very abundant and its old 
nests were to be seen in almost every thorn-bush. 

I was very interested in coming across this paragraph in Belt’s 
‘The Naturalist in Nicaragua.’ ‘A yellow and brown flycatcher 
builds its nest in these bushes, and generally places it alongside 
that of a banded wasp, so that with the prickles and the wasps it 
is well guarded ” (p. 222). 

Five, four, and two eggs were found in three nests on 13. iii. 17, 
and three eggs in a nest on 30.111. 17 (Morogoro) 


842 MR. A. LOVERIDGE: NOTES ON 


LAGONOSTICTA SENEGALA, subsp. n. 


The birds exhibit a preference for the thatch of grass huts 
for a nesting-site, and make themselves perfectly at home, paying 
but little attention to the human occupants of the place ; in one 
such case the favourite perch was the towel-rail. This nest 
contained three eggs and a parasitic egg slightly larger ; the male 
bird was sitting on the lot (Morogoro, 21. vi. 17). “Another nest 
was found to contain four eggs on 30. ui. 17. 

These charming little birds, which may often be seen in 
company with the blue Ureginthus hopping about on paths at 
one’s very feet, have been rather aptly christened “‘ Animated 
Plums” by Sir Frederick Jackson, on account of the rosy or plum 
colour of the males of most of the species. Just before writing 
up these notes, | was watching a pair sitting side by side on the 
bough of a fir-like tree; they had sidled up to one another till 
they could get no closer. The hen was preening the head 
plumage of the male, and afterwards touched beaks, making as 
pretty a little group as one might wish to see. 


PYTELIA MELBA KIRKI Shelley. 


Found a nest with three pure white eggs in a very low bush. 
The nest was very similar to that of Ureginthus, being built of 
flowering grass heads loosely put together, with a tubular 
entrance in one side (Morogoro, 9. iv. Wi). The species is often 
to be found hopping about after dusk, feeding long after other 
birds have gone to roost for the night (Morogor 0, 23. vii. 17). 


AMBLYOSPIZA UNICOLOR Fisch. 


Builds a finely-woven nest attached to reed-stems over water ; 
such a nest containing three eggs was found (Morogoro, 20. iii. 17). 


PLOCEUS REICHENOWL Fisch. 


Clutches consist of two or three eggs ; several such were found 
between 20.v.15 and 20.vi.19. One of these birds fell from a 
tree just as an acquaintance of mine was passing beneath. It 
was still warm when brought to me and not a feather disarranged. 
The only signs of violence were blood oozing from the nostrils 
and the beak full of clotted blood. On skinning, I found 
extensive hemorrhage around the eyes and ears, in intestines, 
lungs, and about the heart. It appeared to me to have died 
from the poison of a venomous snake, probably the Boomslange 
(D. typus) (Nairobi, 18. ix. 19). 


PLocknus Nicricers Layard. 

Some scores of nests of the Black-headed Weaver were 
examined at Morogore between 13.i11.17 and 11.iv.17, and 
found to contain pure white eggs, white eggs with red blotches, 
pure blue eggs, and blue eges with red blotches ; the usual number 
for a clutch appeared to be two; three, however, was not 
uncommon. A bird which I teok to be this species was en- 
gaged in stripping the leaves of a Bussu Palm into threads. It 


HAST AFRICAN BIRDS. |. - 843 


accomplished this by flying to a frayed edge and taking this in its 
beak, flying slowly down with it (Morogoro, 20. xii. 16). 

One of the Black-headed Weavers surprised me to-day by 
dodging after a Praying Mantis in the manner of a flycatcher. 
Almost immediately after catching the insect, however, it dropped 
it; presumably owing to the mantis making good use of its fore- 


legs (Morogoro, 7. xii. 17). 


Pioceus sPEKEI Heugl. 

Out of alarge colony of freshly-made nests found on the plains 
and dependant from a thorn-tree, only one was found to contain 
anything, and that but a single fresh egg (Nairobi, 10. vii. 15). 


ORIOLUS BRACHYRHYNCHUS L&TIOR Sharpe. 

The note of the Black-headed Oriole is as lovely as its plumage. 
On a hot afternoon, when all is still, you may be surprised by 
suddenly hearing a liquid ery of “tokio” ringing out very clearly 
quite close to you. You look in the direction from which the 
sound came and hear it. again, but perhaps behind you. The 
bird appears to be aware that its cry is likely to draw attention 
to it, and so frequently after giving a call it slips away to some 
fresh perch, where it is concealed by the foliage. The natives 
say that it calls “‘ ndugu, ndugu,” which is Kiswahili for “ brother, 
brother.” By imitating the call, I have frequently been success- 
ful in attracting the bird to the very tree beneath which I was 
waiting (Morogoro, 7. vii. 17). 


AMYDRUS MORIO RUPPELLI Verr. 

Riippell’s Red-winged Starling and 4. fenwirostris go about in 
the same flocks, for both species fell to one shot from a gun when 
they were pillaging fruit-trees in the Uluguru Mts. 4. walleri 
was taken later near the same place. A solitary bird was shot 
on the lower slopes of the mountain near Morogoro, where they 
very rarely come; its neck was stripped bare of feathers, which 
gave it so peculiar an aspect that I quite failed to recognise it 
till shot. ‘The only explanation I could think of was that it had 
been snared at some time or had been worried by lice and 
scratched them off; the skin was, however, uninjured. At Fort 
Hall, A. m. riippelli made such a noise on the roofs and gutters of 
the houses that it was a perfect nuisance; they seemed to have 
adapted themselves to the habitations of man as much as the 
English Starling. 


SPREO SUPERBUS Riipp. 

The White-banded Glossy Starling will build in holes in cliffs 
or trees, or make a great untidy nest in thorn-bushes. <A pair 
were seen carrying food to their chirping young in a. cliff-fissure 
in the Kedong Valley (vii. 15). A nest containing young was 
found in a hollow tree at Longido West (30.1.16). Another 
nest with young was found in a thorn-tree at Dodoma (20. xii. 
18), whilst nests with bright blue eggs were found at Mpwapwa 


(30. iv. 17). 


844 MR. A. LOVERIDGE: NOTES ON 


Corvus scaPuLAtus Daud. 


Pied Crow. At Lumbo, I found a broken egg-shell in a culti- 
vated patch, and on enquiry found that the native proprietor had 
pulled the nest down from a cocoanut palm; after this I searched 
and found a good many nests, one containing three very large 
young ones and another some newly-hatched (Lumbo, x. 18). 


DicRURUS AFER LUGUBRIS Rupp. 


At Government Farm a White-shouldered Crow was giving 
vocal expression to its feelings after the manner of its kind, 
cawing hoarsely in the midst of a field of stubble. From a 
nearby mango-tree, stooping like a hawk, came a Black Drongo, 
and went straight for the croaker, who took to wing at the last 
moment. Its more active assailant landed on its back when 
about 40 feet from the ground and gave the Crow a good 
pecking, then, clearing off, it returned triumphantly to its perch. 
{ searched for, but found no signs of anest; possibly the Drongo 
had an ear for music and disapproved of the raucous cries 
(Morogoro, 20. x11. 16). 


LANIUS COLLARIS HUMERALIS Stanley. 


This Shrike, which is very common at Nairobi, breeds all the 
year round, for at any season the young birds may be seen on 
the telegraph wires clamouring for food with shivermg wings. 
Nests with eggs were found on the following dates :—One 
(13.1.15); two (22.v.15); four (1. vi. 15); three (5, vi. 19). 
Nearly all the nests were built in thorn-trees. There is no lack 
of thorns for their larders, and grasshoppers appear to be the 
most common food; huge caterpillars were also found, whilst on 
one occasion a small frog (Rappia sp.) was found impaled on the 
point of an aloe. A friend told me that he once found a small 
blue weaver (Ureginihus sp.) impaled on a thorn and still 
alive. With diabolical ingenuity the Shrike had twisted other 
thorns across in front to prevent the bird freeing itself (Nairobi, 
1. viz-15). 


LANIARIUS SUBLACTEUS Cass. 


A nest of the Lesser Pied Shrike with one egg (Lumbo, 
28. x. 18). 


HLARPOLESYTES SENEGALA ORIENTALIS Cab. 

The Greater Red-winged Bush-Shrike was found nesting on 
the 26th and 29th of March, 1917. In both instances only two 
eges were laid. The nests were very shallow, composed of a 
base of small twigs and rootlets with a lining of much finer 
roots. ‘They were both in bushes, one being at a height of 4, 
the other 5, feet from the ground (Morogoro, 29.111.17). 
Nest with young about 7 feet from the ground (Morogoro, 
4.iv. 17). 


pAST APRICAN BIRDS. 845 


MALACONOTUS POLIOCEPHALUS BLANCHOTI Steph. 

Chestnut-breasted Thick-billed Shrike. One of these beautiful 
Shrikes was seen sitting on its nest in the fork of a rubber 
sapling which had sprung from the stump of a felled rubber-tree. 
The nest was at a height of 18 feet from the ground, and was 
slightly hidden by a few withered yellow leaves, but all the 
surrounding trees in the plantation were bare of foliage. The 
egos, of which there were three in the nest, were slightly 
liver than an English Blackbird’s, and were thickly freckled 
with brown and faint purple on a white ground. One contained 
a chirping chick, already breaking from its shell, and this I left ; 
the other two were unfertile aad easily low (Morogoro, 
ay 2g MO) p 


EUROCEPHALUS RUPPELLI Reichw. 


The White-headed Shrike was found building a very beautiful 
and solid nest on the tip of a branch of mimosa-thorn quite 
12 feet from the trunk and perhaps 9 feet from the ground 
(Kongwa, 25. iv. 17). 


CRATEROPUS KIRKI Sharpe. 


A nest of this species was taken on a mimosa bough about 
10 feet from the ground. It held three eggs of Kirk’s Babbling 
Thrush and one slightly larger egg of a Cuckoo (Coreg caper) ; 
all the eggs were uniform ereonich! blue (Lumbo, 22.x.18). A 
cock-bird was found sitting on three eggs (Morogoro, 23. i11. 17). 


PYCNONOTUS BARBATUS MICRUS Oberhols. 


The Yellow-vented Bulbul breeds during the greater and lesser 
rains, if not all the year round. Many nests were found at 
Morogoro on the following dates :—-T'wo young ones in a nest 
in boughs of a flamboyant (8. xii. 17); two eggs and three eggs 
in a similar situation (14. xi1.17); young in a nest in a bunch 
of bananas in palm (16.1.18); two eggs (26.1i1.17); and one 
egg (11. 1v. 17). 

After a night of heavy rain, I witnessed a somewhat unusual 
sight. Not 15 vards from where I was standing, termites 
were flying from their nest-holes. No less than seven species of 
birds were engaged in catching them, of which five were seed 
or fruit eaters. The birds were two Weavers (Lagonosticta sene- 
gala subsp.), two Waxbills (U. niassensis), one Finch (S. acterus 
barbatus), one Sparrow (P. g. swahelicus), two Pied Wagtails 
(M. vidua), and two of these Bulbuls, whilst, to add to the colour- 
effect, a dainty Scarlet-breasted Sun-bird (C. s. incestimata) 
hovered above and captured the termites on the wing (Morogoro, 


10. iv. 18), 


PYCNONOTUS TRICOLOR PALLIDUS Roberts, 


Very abundant at Lumbo. Five nests, each containing two 
eggs, were found on 28. x. 18. 


846 MR. A. LOVERIDGE: NOTES ON 


> 


ANDROPADUS INSULARIS Hartl. 


Many nests of the Coast Olive Bulbul were also found, and 
five clutches of two eggs each taken; two would appear to be 
the normal number for a clutch. The eggs are white, scribbled 
and blotched with brown and faint purple at the larger pole, in 
some cases all over. The nest is small and shallow, composed of 
twigs, and lined with fine rootlets (Lumbo, 28. x. 18). 


PRINIA MYSTACEA TENELLA Fisch. 


Nests of the White-eyebrowed Grass-Warbler, with three eggs 
in each, were found on 22.11.17, 5.iv.17, and 21.vi.17 at 
Morogoro. 


SYLVIELTA WHYTEI Shelley. 


This Warbler constructs a most beautiful purse-like nest, 
which it hangs from the branches of a thorn-bush. Three nests 
with eggs were found at Lumbo on 28.x. 18, and a bird was 
sitting on two eggs at Morogoro on 28. x1.18. Two appears to 
be the normal clutch. 


CISTICOLA LUGUBRIS Riipp. 

A deserted nest of the Brown-headed Grass-Warbler, con- 
taining one egg, was found; from the interior roof of the nest 
depended the paper nest of a wasp, whilst a small company of 
solitary bees had their comb in the bottom (Ruaraka River, nr. 
Nairobi, 11. x. 19). 


CisTICOLA ERYTrHROPS Hartl. 
A nest of the Rufous-fronted Grass-Warbler containing three 
eggs (Nairobi, 5. vi. 15). 


CICHLADUSA GuTTATA Heugl. 

The Spotted-crested Ground-Thirush is a very sweet singer, 
generally holding forth from thickets just before sunset; it does 
not expose itself when singing. ‘The mud nest is built upon a 
braneh or, more rarely, in a fork; it is very deep. ‘The eggs 
are of a uniform blue colonr. A broken egg beneath a nest on 
12.1.175; im another nest young on 22.11.17; two eggs in a 
nest found on 4.iv.17 and two more in one on 6. iv. 17 (Moro- 
goro, 1917). 


TCHITREA CRISTATA SUAHELICA Reichw. 

I have seen this Paradise Flycatcher hovering up the trunk of 
a tree, picking the insects off the bark whilst on the wing. It 
constructs a beautiful cup-shaped nest, which it covers with 
lichen; one such was attached to some epiphytes at a height 
of 8 feet above a mountain torrent, and contained young (Uluguru 


Mts., 28. xi. 18). 


EAST AFRICAN BIRDS. S47 


RIPARIA RUFIGULA Fisch. 


Nest of the Rufous-throated Rock-Martin, containing three 
eges, attached to the wall of a cattle-shed (Kabete, 6.v, 18). 


Hirunpo suirut Leach. 

A nest of the Wire-tailed Swallow with three eggs (Moro- 
gore, 23. vii. 17); with two eggs and apparently deserted (Frere 
Town, 1. vii. 19). At Morogoro on 14.11.17, I made a note that 
thousands of swallows had been gathering on the telegraph wires 
for the past few days, apparently for migration. 


HiRUNDO PUELLA UNITATIS Scl. 

The Smaller Stripe-breasted Swallow was associated with the 
preceding species in the assemblage for migration at Morogoro 
(14.11.17). A dead bird was found upon her nest, which 
contained three eggs, one of which was plain white (Kongwa, 
20. iv. 17). 

Along the river these birds nested against the rocks, but almost 
every house in Morogoro had its nest with the typical long 
entrance passage of mud. The birdsare most persistent, and will 
continue to build in rooms, where they are not welcome and where 
their nests are knocked down again and again. The earliest date 
at which [ found a nest at Morogoro was on 25.vi.17, and the 
latest 15. vill. 17. Three eggs is the invariable clutch. 

On 23. vu.17, I found a nest with no entrance tunnels, and 
broke a small hole into it, but found no eggs. On again exam- 
ining it a few days later, there were three eggs, and the hole 
I had made had been neatly finished off with mud to form a 
window or third outlet. In one instance I opened a nest care- 
fully, and found three eggs; two were white with faint reddish 
speckles and the third pure white (23. vil. 17); these I took. On 
1.viii. 17, I again visited the nest ; the small hole [ had made had 
been repaired, and three eggs again laid of which one only was 
white. On 15. vii.17 a third visit was made, and again three eggs 
found of which one was white. A fortnight later the hole had 
been again repaired, and the bird was sitting. I did not molest 
her further, as such persistence deserved its reward, and it was 
the variation in the clutches that caused me to rob her. 

Calling on an English sergeant on one occasion, I found him 
nursing one of these swallows, which had a broken wing ; he was 
endeavouring to feed it on crumbs and meal. He had, it trans- 
pired, thrown a stone at the bird as it sat chirping on the roof- 
ridge: to his surprise and grief he hit it, and was now endea- 
vouring to restore it to health by a vegetarian diet !! (Morogoro, 
Ziieelen lrd)s 


Hirrunpo Emini Reichw. 

Kmin’s Swallow was only once met with at Morogoro (31. vil. 
17) when evidentiy on migration, three birds in poor plumage 
being seen. A paiv of these birds, after starting the foundations 


of twenty-one nests on the roof-rafters of my oftice verandah, 


848 MR. A. LOVERIDGE: NOTES ON 


appear to have at last selected a spot exactly above the most 
frequented place—i.e., between the stairs and door (Nairobi, 
8. vil. 119). 

15.vi.19. The birds sat with dabs of mud in their beaks, 
uttering their peculiarly sweet metallic notes. The nest is about 
finished. 

22.vi.19. During the week the entrance tunnel has been 
under construction, the work proceeding very slowly. 

28.vi. 19. The birds appear to have gone away. 

1. viii. 19. The weather has been very cold and dull during the 
past month, and the swallows have only twice put in an appear- 
ance, when they added a few feathers to the nest. 

7. viii. 19. Felt in the nest, and there appeared to be no lining 
worth mentioning 

14. viii. 19. Nest with lining of grass and feathers, and one 
broken egg with a streak of red in it, lying on the ground, having 
been pulled down by a ruthless native. When did the egg get 
incubated ¢ Is it possible I did not feel to the bottom of the 
nest on the 7th inst. ¢ 


CYPsELUS AFFINIS Hardw. 


The Indian Swift is extraordinarily abundant at Frere Town, 
where great numbers of them fly about, screaming, towards 
evening ; in fewer numbers they are to be seen at most hours 
of the day. They build along the cliffs or under the eaves of 
verandahs. The nests are constructed of feathers and straw, so 
cemented together with mucilage that they have the strength 
and consistency of cardboard. A large number of nests were 
examined on 1.vii.19; ten of these held young, three held 
incubated eggs whose clutches numbered four, four, and three 
respectively. One nest had a single fresh egg in it. 

A nestling was picked up at Morogoro, 29. x. 17. 


TACHORNIS PARVUS Verr. 


My notes on the curious nesting habits of the Lesser Palm-Swift 
have already been published *. On July 17th, 1917, I revisited 
the colony which were nesting in the cocoanut-palms (not 
banana-palms, which was printed in error). One nest was 
completely upside down, having been built that way; the bird 
was brooding two eggs which were nearly ready to hatch. 
Another nest had a bad egg which had a small hole in the top of 
it, probably made by a bird’s claw. Another held newly-hatched 
young; yet another was occupied by downy young. In one 
instance both birds were on the nest and appeared to be paired. 
Some nests were unfinished, possibly abandoned. 

On 4.11.18, I again visited the spot ; in one nest there were 
two fresh eggs, several hard-sat clutches in others, newly-hatched 
and half-grown young in the rest. 


* © Observations on the Nesting Habits of the Palm Swift etc.” Mem. Proc. of 
the Manchester Lit. & Phil. Soc. vol. lxi. pt. ii. (1919). 


EAST AFRICAN BIRDS. 849 


CAPRIMULGUS EUROP&US Linn. 
The Kuropean Nightjar was collected at Morogoro (24, ix. 17). 


CAPRIMULGUS FOSSEI MOSSAMBICUS Hartl. 


A single egg, perfectly fresh, was picked up and brought to me 


by a native. A good series of birds were collected (Lumbo, 
28. x. 18). 


CorAcIAS GARRULUS Linn. 

The European Roller on migration was met with on several 
occasions. Animmature female was shot at Nairobi (29. x. 15). 
Large numbers of them were seen at Longido West (ii. 16). 
Specimens were obtained from a flock at Morogoro (10, xii. 17). 


CoracrAs CAUDATUS Linn. 

The Lilac-breasted Roller was seen clinging to a blasted tree 
just below a suitable nesting-hole. On both occasions when I 
approached the tree I was made the subject of very noisy demon- 
strations, so I concluded that they were nesting in the hole 
(Bissel, 20. xii. 15). 

A pair of birds were obtained at Kongwa (23.iv. 17) in very 
fine plumage, and during that year they were extraordinarily. 
abundant at Morogoro. In the stomachs of several specimens 
were found many nauseous and warningly coloured grasshoppers 
(Zonocerus elegans Thunb.). 

Some three pairs lived in the neighbourhood of the camp at 
Lumbo. One pair frequented a large baobab-tree, which served 
as a jumping-off ground for their little excursions to rag the 
soberly-garbed crows, which they chivvied mercilessly about. 
After these little excursions, one or both of them would soar 
higher and higher above their favourite tree, and then come 
tumbling down in the most extraordinary way, more like a winged 
bird that is making an effort to recover itself than a healthy one. 
Towards October 1918, 1 saw one of the birds outside a hole near 
the top of the baobab, but thought climbing out of the ques- 
tion. At the end of the month a native brought mea Roller, 
which he told me he had caught on its nest in a hole in a tree ; 
it was sitting upon two eggs which he had broken. TI refused to 
purchase the bird, but from the information that he gave me 
I concluded that it was from this baobab that he obtained it, and, 
if so, this would explain their energy in driving off the crows. 
These birds were extremely shy of approach, whilst those met 
with at Morogoro were quite the reverse. 


MELITTOPHAGUS BULLOCKOIDES Smith. 

A colony of some fifty White-fronted Bee-eaters was found in 
an artificial pit some 20 cubie feet in size; the nesting-holes 
had been excavated in the sides of the pit (Ngari Mtoni, nr. 
Arusha, 17.iv. 16). 


850 MR. A. LOVERIDGE: NOTES ON 


Merops APIASTER Linn. 


A single pair, in very worn plumage, of the Huropean Bee- 
eater were obtained at Morogoro (17. x. 17). 


LOPHOCEROS DECKENI Cab. 


A male and immature female Von der Decken’s Hornbill were 
shot on 1. vi. 17 at Morogoro. The male had two large complete 
mantids in its stomach, whilst the female had only beetles. ‘The 
young female was strikingly different from the male. Its bill 
was black instead of orange-red and yellow ; the wings were also 
spotted with white. 


LorHOCEROS MELANOLEUCUS SUAHELICUS Neum. 


The Swahili Crowned Hornbill was common at Morogoro, the 
type-locality for the subspecies. 1 extract the following note 
from my diary under date 3. xi. 17 :—“ The last few days we have 
had a small flock of these hornbills flying past in the morning and 
evening; apparently they return to the big trees along the river 
to roost. They usually frequent a group of trees quite close to 
my quarters for a short time before continuing their flight; one 
is soon made aware of their arrival by their shrill whistling 
eries. The birds do not sit together, but on separate branches, 
from which they make sudden and absurd dives as if they were 
top-heavy and could maintain their balance no longer. These 
little excursions are apparently after insects, though their 
stomachs more often contain the remains of fruit.” And on 
4,ii.18:—‘“ One of these hornbills, which flew over the camp 
to-day, apparently had nesting (!) material in its beak.” 

Whilst out in the bush one evening, I came upon an open 
glade, thereby disturbing the most remarkable assemblage of 
typical African birds which I have ever met with in so small a 
compass. Some twenty Hoopoes (U. africana) rose, and perched 
for a few moments in the surrounding trees; a large red-billed 
Hornbill (Z. m. swahelicus) sailed off in its own peculiar dipping 
flight. Wood-Hoopoes (2. ¢. schalow?) raised their noisy outery. 
A red-headed Woodpecker (C. scriptoricauda) paused in its oceu- 
pation of hammering a tree, caught sight of me, and dived off 
with a shrill whistle, so that the glade which was so full of life a 
moment before was left to myself (Morogoro, 14. vii. 17). 


LoPHOCEROS NEUMANNI Reichw. 


A female of Neumann’s Hornbill, which was being noisily 
mobbed by a party of small birds whilst in flight, was found 
to have a praying mantis in its stomach (Morogoro, 28. xi. 17). 


ALCEDO SEMITORQUATA Swains. 

Half-collared Kingfisher. A native brought me two beauti- 
ful nestlings ; their head, back, and wings were turquoise-blue, 
their breasts fawn-colour (Morogoro, 28. xi. 17). 


EAST AFRICAN BIRDS. 851 


CoRYTHORNIS CYANOSTIGMA Rupp. 


Malachite Kingfisher. A nest of young ones was found in the 
river-bank at Morogoro (1917). 


FLALCYON LEUCOCEPHALA Vieill. 

Brown-bellied Kingfisher. Having just stepped out of my 
tent to walk across to my grass-hut one morning, I heard a 
screech and a shrill whistling cry. There was a rush of small 
birds about nie, and I was just in time to see a South African 
Lanner (Falco biarmicus) rise from its stoop with a kingfisher 
in its claws and speed away to a big tree some 20 yards off. 
I snot the Lanner promptly, and it fell stone-dead with the 
kingfisher in its claws (Morogoro, 17.1. 18). 


HALCYON CHELICUTI Stanley. 


Striped Kingfisher. Hach morning as I wake I hear a pair of 
small kingfishers start shrillmg—this coined word seems to 
describe the noise better than anything. The note is not unlike 
‘‘ neewit,” whistled through the top front teeth, but the one bird 
follows up its mate’s call, so that there is one continuous sound, 
and at first I thought it was one bird calling. As each in turn 
utters its note, it spreads its wings like a fan, then raises them 
above the level of its head, lowers and then closes them, repeating 
the performance constant!y. It is a pretty spectacle to witness, 
as the wings and tail are partially pale blue and displayed to 
advantage. As these little kingfishers are no larger than 
sparrows, and select the topmost branches of the acacia-trees for 
their performance, it was some time before I could find out the 
perpetrators of the noise (Morogoro, 26. x. 17). 


HALCYON ALBIVENTRIS ORIENTALIS Peters. 

Oriental Brown-hooded Kingfisher. A young lizard (Gerrho- 
saurus flavigularis) was in the stomach of a specimen shot at 
Dar-es-Salaam, 26. vi. 18. 


COLIUS STRIATUS AFFINIS Shelley. 


Mouse-bird. I believe Dar-es-Salaam Colies are referable to 
this race, which was found nesting with three eggs on 1. ii. 19. 
A nest and three eggs were taken at Nairobi on 7.v. 19. 


COLIUS INDICUS PALLIDUS Reichw. 


Was collected at Lumbo, where there were many flocks in 
July and August ; its cry was like that of a plover (Stephanibyz). 
After August they were only seen singly or in pairs; this should 
have put me on my guard, but it was not till I happened ona 
nest containing three eggs that it occurred to me that the birds 
had disbanded for breeding. Search being made, a dozen nests 
were found in October, but nearly all of them contained highly- 
incubated eggs or young. The eggs are round, dull stony-white 


852 MR, A. LOVERIDGE: NOTES ON 


in colour, streaked, blotched, or freckled with reddish-brown in 
great variety. The nests were all situated in thorn-trees; those 
that were putting forth green shoots were. given preference ; 
they had deep foundations of thorny sprays, upon which was built 
a cosy nest of moss, lined with fine fibres or grass. The old birds 
sit very close, and make no attempt to leave if merely looked at ; 
one’s hand must be approached unreasonably close before they 
will slip off. The clutch was invariably three (Lumbo, 28. x. 18). 


ScHIZORHIS AFRICANA LEUCOGASTRA Reichw. 

Many species of Plantain-eaters were collected, but none were 
more difficult to obtain than this species, which frequents sandy 
thorn-bush country. At Dodoma its harsh cry was quite a 
feature of the scrub-life. The ‘ha-haa” (emphasis prolonged 
on second syllable) has quite a jeering sound as the bird rises from 
some conspicuous perch it has chosen on the top of a thorn-tree 
(commanding a good view of the dense thorn patch), and, flying 
off for a short distance, leaves the perspiring hunter to push his 
way through another few hundred yards of tangled undergrowth, 
with the probability of a recurrence of the flight, which seems 
always to be accompanied by the ery (Dodoma, 5. xii. 18), 


CENTROPUS MONACHUS Riipp. 

A series of skins were obtained at Dar-es-Salaam and Lumbo. 
At the latter place the bird was very abundant, and was found 
to be feeding largely on maggots in the human feces so common 
in the bush about the camp. The throat plumage in most of the 
skins was soiled with fecal matter, and a good deal was found in 
the stomachs of the birds along with the maggots. ‘The offensive 
smell made the birds most unpleasant to skin. I watched a bird 
feeding in the early morning. ‘The natives say that the skulking 
habits of this Cuckoo amongst the bushes are an evidence of its 
sense of shame for the diet which it has adopted (Lumbo, 
21. vii. 18). 

At the back of my tent was a large mango-tree whose base 
had been wired in to form a large circular chicken-run, in which 
a hundred and fifty fowls were kept. Karly one morning one 
of these Cuckoos flew into the mango-tree, and as the shadow of 
the Cuckoo fell across the enclosure, a panic seized the fowls, 
which flew and ran em masse against the wire-netting with a 
bang ; there they crowded and trampled upon each in the greatest 
confusion. They take no notice of the crows passing overhead 
or perched in the nearby palms. Undoubtedly the Cuckoo was 
mistaken for a hawk (Lumbo, 25.ix. 18). 


CoccysTEs GLANDARIUS Linn. 

The Great Spotted Cuckoo, of Europe, was first seen about 
September 1917, when a pair flew over the camp at Morogoro. 
Later, a male was shot on 27.x.17 and another on 29.i. 18. 
A gaudily-coloured grasshopper (Zonocerus elegans) was found 
in fair numbers in its crop, 


EAST AFRICAN BIRDS. 853 


Coccystes CAFER Licht. ; 

The African Pied Crested Cuckoo was collected at Morogoro 
(10. xii. 17) and Lumbo (22. vii. 18); at the latter place an egg was 
found in the nest of a Babbling Thrush (C. kirhki), as already 
mentioned (22. x. 18). 


*CucuLus canorus Linn. 

The European Cuckoo was twice obtained at Morogoro in 
February 1918 (12th and 26th). The birds were flying across 
the railway-line from south to north about a mile outside the 
town. 


Lypius zomB# Shelley. 

Were very common at Lumbo, going about in pairs at the 
time of my stay. Perched on the bare branches of the baobabs, 
they went through the most extraordinary courting (2) antics, 
accompanied by eries no less unique (Lumbo, 11. vii. 18). 


TRICHOLEMA LACHRYMOSA Cab. 

The Black-faced Barbet was collected at Morogoro (11. vii. 17) 
and Dodoma (22.11.18). Indian corn was found in the stomach 
of one of the specimens from the first locality. 


TRACHYPHONUS EMINI Reichw. 

Common at Dodoma, where they were going in pairs, bowing 
and singing to one another. <A nest oth a ainalle young one 
was Goma | in a hollow thorn-tvee (Dodoma, 5. xi1. 18). 


CHALCOPELIA CHALCOsPILOS Wael. 
Green-spotted Dove. Two eggs in nest (Lumbo, 24. ix. 18). 


TURTUR SENEGALENSIS Linn. 
Laughing Dove. Two eggs in nest (Morogoro, |. viii. 17). 


TYMPANISTRIA TYMPANISTRIA Temm. 
Two eggs in nest (Kabete, 24. v. 15). 


P@CEPHALUS FUSCICAPILLUS Verr. 

I followed a Brown-headed Parrot to a blasted tree, in 
which was a woodpecker’s hole some 20 feet from the ground. 
Failing to get my hand in, I returned in the evening with two 
boys, who chopped the hole open after half-an-hour’s labour. 
One of the stupid fellows placed the three large white eggs in 
his handkerchief, put same in his pocket and slid down the tr ee, 
breakine all three, which were perfectly fresh (Morogoro, 25 
wale WF) 

The species was also collected at Dar-es-Salaam and Lumbo; 
at the latter place I had a young one, which was very tame and, 
though loose, made no attempt to fly away. 

Proc. Zoou. Soc.—1922, No. LVILI 58 


854 MR. A. LOVERIDGE: NOTES ON 


AGAPORNIS PERSONATUS Reichw. 

Parties of this handsome little Love-bird flew about with great 
rapidity, screaming noisily ; they were first met with at Kongwa. 
They were nesting in the holes of a baobab-tree, whose apertures 
they had apparently partly closed with a defence of thorns. Two 
eggs were found at the foot of the tree side by side, and meg- 
sured 2x? inches. How they came to be there I cannot say, for 


they contained embryos nearly ready for hatching (Kongwa, 
26.iv. 17). 


Buso AFRICANUS Temm. 


The Spotted Kagle-Owl was collected at Morogoro and Lumbo. 
At the latter place one was being mobbed by a flock of crows, 
which had driven it from the shelter of the bush and were 
pursuing it through the mangroves on the sea-shore. It alighted 
on a sand-bank, and was encompassed by a circle of crows, cawing 
with outstretched necks. The group would have made a unique 
photograph (Lumbo, 3, 1x. 18). 


PANDION HALIAHTuS Linn, 
The Osprey was found nesting at Lake Naivasha with young 
on 16. vu. 15. 


Fatco BiaRMicus Temm. 

A female of the South African Lanner was shot whilst 
devouring a chicken in the topmost branches of a baobab-tree 
(Morogoro, 11.xii.17). As already mentioned, the cock-bird 
was shot with a Kingfisher in its claws (Morogoro, 16.1. 18). 


Fanco cuvieRt Smith. 
The African Hobby was shot while eating a Yellow Wagtail 
(Motacilla campestris) (Morogoro, 31. x. 17). 


DISSODECTES DICKINSONI Sclater. 

Dickinson’s Kestrel, shot at dusk in a baobab-tree, was found 
to have beetles, locusts, and a praying mantis in its stomach 
(Morogoro, 18. vii. 17). 


CERCHNEIS TINNUNCULUS Linn. 


The Common Kestrel was very common at Morogoro and 
Dodoma; at the latter place particularly it was rarely that 
there was not one in sight, usually sitting on Huphorbia 
bushes, which were scattered about the thorn-bush. One 
which I had in captivity throve well on the remains of 
specimens which I was preserving. On one occasion, hearing 
a succession of screeches outside my tent, I ran out to see 
another Kestrel rise from the cage and fly to a tree near 
by. ‘The captive bird was clinging to the wire-netting of its 
extensive run and screeching, being answered by the bird in 


EAST AFRICAN BIRDS. 855. 


the tree for some minutes. Nearly a month later a flight of 
locusts visited the neighbourhood, and my bird exhibited such 
excitement that I released it to see what it would do. Flying 
straight to a large tree in the middle of the camp, it captured a 
locust ; for the next half-hour it remained about the tree, taking 
short flights after the insects and returning to its perch to eat 
them. ‘The following were found in the stomachs of specimens 
collected :—A lizard (Hremias spekit) and skink (Mabuia varia) 
and many locusts in one (Morogoro, 4. xii. 17); a lizard (Latastia 
longicaudata) (Morogoro, 5.11.18); a small bird (Dar-es-Salaam, 
17. xi. 18); a rat and lizard and a parasitic worm (A. recto- 
vaginata) (Dodoma, 6, xii. 18). 


PrRNis Aprvorus Linn. 

The Honey-Buzzard was twice shot at Morogoro (23.1. & 
l.iv.18); both were females, the former with grey head, the 
latter with brown. In the stomach of the former were found 
a large number of pupzeand grubs of the grey hornet (Belonogaster 
griseus). It has been said that the feathers around the eyes 
of this Buzzard protect it from stings, but anyone who has seen 
the long stings of Belonogaster might be forgiven for doubting it. 


ELANUS CHRULEUS Desf. 

The Black-shouldered Kite—a rare visitor to England—is com - 
paratively common in the Hast African bush, where its dove- 
coloured plumage makes it look something like a sea-gull when 
on the wing. Specimens were obtained at Nairobi, Dodoma, 
Morogoro, and Lumbo. In the stomach of one was a mouse skull, 
beautifully cleaned and undamaged (5. ii. 18). 


AQUILA RAPAX Temm. 

The Tawny Eagle was shot at Morogoro (9.11.18) and 
Dodoma (6. x11. 18). The former had its stomach full of putrid 
goat’s flesh, and the latter with bullock flesh and offal thrown out 
from the slaughter-house, where it was keeping company with 
vultures and kites. It had a parasitic worm (Ascaris depressus) 
in its stomach. On the road from Bissel to Longida, I rode up 
to au Hagle which was feeding on a dead mule. It allowed me 
to approach within 30 feet and, when I reined-up, looked at me 
for a moment, and then went on coolly tearing off strips of flesh 
with its beak, which it then transferred to its claws, to be held 
whilst rendered still smaller. In the Kedong Valley two young 
Eagles, just fledged, were seen sitting on the rocks at the base of 
the towering cliff, where their nest evidently was (17. vii. 15). 


LopnHoairus occrprranis Daud. 

This Crested Kagle was found to be nesting in February. The 
nest was situated in a treeat a height of 80 feet from the ground. 
The cry of the old birds when one approached reminded me of an 

58* 


856 MR. A. LOVERIDGE: NOTES ON 


English Sparrow-Hawk’s note when disturbed. A. few days later 
the birds began sitting. It was quite impossible to reach the 
nest, not merely because there was not a branch for 60 feet up, 
but because a swarm of bees nearly two feet in length were 
under the junction of the first branch and the trunk, whilst 
a second swarm hung from the big limb on which the nest 
was placed, about 30 feet from the main stem (Morogoro, 
20. ii. 17). 


MELIERAX GABAR Daud. 


Gabar Goshawk. The feet of three birds were in the crop and 
stomach; they appeared to be those of Grass-Warblers (Moro- 
goro, 2.vi.17). The melanistic form was obtained a few days 
later at the same spot (Morogoro, 5. vi. 17). 


ASTUR SPHENURUS Rupp. 


Stomach of Ruppell’s Goshawk was found to contain locusts 
(Morogoro, 12. vil. 17). 


Asrur TacutRo (?NyAnz@ Neum.). 

Probably it was this race of the African Goshawk which I obtained 
at Nairobi (20. x. 15) and Morogoro (20. xi. 17). I found the nest 
of this species on November 17th ; it was situated at the top of a 
tree whose bark was studded with thorns: the tree was on the 
banks of a stream in thick bush. On November 20th, I revisited 
the nest at daybreak aud saw the hen feeding the young; she 
was very shy and, being disturbed, did not return. In the 
evening she was on the nest, but slipped silently away as we 
approached. We had brought ropes, which assisted the native 
in his ascent; climbing was a very tedious business, as the 
thorns had to be cut away all up the trunk. The two young in 
the nest were of very different size, and one would think there 
was quite a week’s difference in their ages; the smaller bird was 
still in the down stage. 

After removing the young, I sat down in the gathering gloom 
and watched the dark outline of the nest silhouetted against the 
sky ; an indistinct “something” sailed past, and the next moment 
the mother could just be seen alighting on the edge of the 
nest. She had scarcely time to close her wings when I fired, 
and she fell forward dead into the empty nest. This involved 
another climb for the boy. After gathermg up the rope, 
hatchet, and spoils we stumbled back through the bush to camp, 
guided by the gleaming thread of water which lay in the river 
bottom. 

J skinned the bird at once —it was in fine condition—and then 
fed the young with the flesh cut into strips; they ate with avidity. 
Next morning I fixed up a very fair imitation of their home on 
the top of a bush in the python’s cage. On the 28th they began 
to feed themselves, and were found of standing up in the nest. 


EAST AFRICAN BIRDS. 857 


They held down scraps of meat in their claws, tearing it into 
still smaller scraps with their beaks. Up to this time they 
were fed on the flesh of monkey, lemur, rat, bullock, kite, hawk, 
egret, hornbill, kingfisher, voller, wheatear, wagtails, weaver, 
erow, puff adder, and chameleon. All these seemed to agree 
with them except that of the bullock and the chameleon fat, both 
of which they disgorged. 

On December Ist the larger of the young Goshawks was 
just beginning to fly about the cage when it disappeared; the 
12-foot python must have eaten it. I had kept him so gorged 
with dead fowls that I never imagined he would molest them. 
The following day, hearing my monkeys making a great outcry, 
I hastened out to find the python sliding silently towards the 
bush on which the nest had been placed. I removed the remaining 
hawk just in time, for on passing back to my tent I saw the 
python had raised itself to the nest over which he was running 
his snout. Having nowhere else to keep it, I left the bird in an 
open grass hut. 

On December 7th it took trial flights about this hut. When 
the two young ones had been in the python’s cage I had been 
much puzzled by finding their excreta about 4 feet from the 
nest. Now, I observed the remaining bird hop out of the box in 
which it was, turn round on the edge, cock up its tail and 
discharge the fluid “ whitewash” to a distance of 3 feet. This 
shows how they are able to keep the nest clean. It disgorges 
pellets of fur 13" x 3" in size. 

The young bird was not in the least like its parent, having 
large pear-shaped brown blotches on the breast like 7. biarmucus. 
The mother, of course, has’ transverse fine striations. On 
December 13th I missed the bird; the night before it had flown 
about 500 yards and stayed out all night. Someone accidentally 
discovered it in a rubber-tree, and, in trying to recover it, struck 
it with a butterfly net, which caused its death the following 
night, as a small clot of blood was found on the spine when 
dissected. 


BUTASTUR RUFIPENNIS Sundev. 


A Buzzard Eagle was fairly plentiful at Morogoroand Dodoma. 
The stomach of a specimen examined contained scarabs (Morogoro, 


Biles 18): 


CIRCARTUS PECTORALIS Smith. 


Tn the stomach of a Black-breasted Harrier Hagle was a pellet 
of rodent fur, in its gullet was a MHissing Sand Snake 
(Psammophis sibiians) 2 feet in length, its tail being doubled 
back for about 9 inches; it had been swallowed head first. 
Curiously enough, four days earlier I stalked a handsome Barred- 
breasted Eagle in the same tree in which [I shot this specimen. 
On raising my head when within range I lost sight of it, but a 


858 MR. A. LOVERIDGE: NOTES ON 


minute later it rose from the mealies two hundred yards from the 
tree, with a snake in its talons. It came so low over my head 
that I was able to recognise the snake as a Hissing Sand Snake. 
The Eagle then ascended in great spirals, taking the snake from 
its talons with its beak and gulping it down as it ascended. I 
was so interested that I forgot to shoot till the bird had finished, 
and then when I did fire, missed, so that I was unable to 


identify the Hagle, which was an unfamiliar one (Morogoro, 
A & 8.vi.17). 


ASTURINULA MONOGRAMMICA Temm. 


Variously known as the African Buzzard-Eagle, Lesser Whist- 
ling Hawk, and One-streaked Hawk. The first name from a 
popular point of view is a very absurd appellation conveying 
the idea of great size. One-streaked Hawk seems to me the best 
popular name conveying some idea of the bird. A female shot 
at Morogoro had a Side-walking Spider (Palystes sp.) in her 
stomach. A male was obtained a few months later (Morogoro, 
2. iv. & 9. vii. 17). 


Crreus MACRURUS Gmel. 


In the stomach of a Pale Harrier was found a young snake 
(Rhamphiophis oxyrhynchus) and a skink (Mabuca striata), whilst 
in tnat of a second were three species of lizards (Hremias 
spekit, Gerrhosaurus flavigularis, and Mabuia striata). Here 
the Pale Harrier seems to prefer a reptilian diet (Morogoro, 
31.1, & 28, 11.18). 


MELANOPHOYX ARDESIACA Wael. 


The Black Heron feeds along the shore at Dar-es-Salaam when 
the tide is out; it was there very difficult of approach. On one 
occasion, as I was returning home, one of these small Herons flew 
over the causeway and dropped on a muddy flat not far from 
where there was some grass. I was on the far side of the water 
and immediately made for the place. The bird | found was 
performing some very curious antics. Running swiftly forward 
for about six steps, it abruptly halted, placed its beak to the 
ground, and raising its wings, brought them forward and 
downwards, so that for a moment its head was entirely hidden 
by the outspread wings which were brought down to the 
mud. This it repeated again and again until, reaching the 
water, it turned about and practised its steps in the opposite 
direction. 

I imagined that it was a cock performing some courting display, 
but on skinning found it to bea hen. Taking advantage of each 
moment when its head was concealed, I wriggled forwards through 
the grass till within range, when I dropped it, very dead indeed, 
but a perfect specimen (Dar-es-Salaam, 1. vii.18). Six months 
later I saw these birds performing the same antics on the fore- 
shore opposite the ruined Governor’s Palace. 


EAST AFRICAN BIRDS. 859 


ARDEA MELANOCEPHALA Vig. & Child. 


The Black-headed Heron was met with at Dar-es-Salaam 
(26. vi. 18), Morogoro (29.xi.18), and Tabora (13.xi1.18). At 
Tabora they were nesting in a great tree in the midst of culti- 
vated ground, the birds winging their way home of an evening 
from their feeding grounds. One of the Morogoro specimens was 
shot stalking about a stubble field, and its stomach on examination 
was found to contain the remains of rats, apparently five in 
number. 


ARDEA CINEREA Linn. 


The Common Heron of Europe was twice collected at Dar-es- 
Salaam (5. vii. & 19. xi. 18), the circumstances being as follows :— 

Just 60 feet below the bank on which our house is built, 
and perhaps a hundred yards away, lies a one-time estuary of 
the sea; the fresh water is now shut off from the sea by a 
causeway, but a series of large pipes still allow of communication 
between the two at high tides, after which infusion the water 
is brackish. Here various waterfowl are wont to feed on the 
freshwater shrimps and small fish, including the Mud Skipper 
(Periophthalmus sp.), which is present in large numbers; while 
at daybreak each morning a pair of herons are to be seen standing 
in the shallows. 

Every morning for eight days past I tried to stalk them, 
but they were too wary, taking to wing whilst one was still a 
hundred yards off. Once disturbed the birds would circle round 
in the blue above, and then, heading over the causeway, would fly 
seawards, doubtless to reeommence feeding in one of the quiet 
bays or lagoons with which the coast abounds. However early 
I arrived, the birds were there before me, but determined to 
succeed this morning I started out at 4 o’clock. 

When I left the house it was pitch dark except for a waning 
moon whose faint beams were reflected on the shimmering sur- 
face of the dark water. Stumbling around the estuary in the 
spongy soil where crickets were shrilling away, I reached the far 
end, and was carefully withdrawing my feet from a particularly 
boggy piece of ground, when suddenly I became aware of the 
presence of a bird standing motionless in the shallow water 
scarcely 30 feet away. 

I hesitated for a moment, lest in the deceptive light the bird 
might be nearer or farther than it appeared, as I was anxious not 
to damage the skin; also the report of the gun echoing across 
the water might disturb the slumbers of our neighbours 
occupying the houses along the opposite bank. Ere I had made 
up my mind the heron rose with a hoarse squawk, and, spreading 
its great wings, disappeared into the darkness. A smaller species 
arose and settled again further out. 

Pursuing my way round the estuary I arrived at an old 
stranded buoy close to which there was a favourite fishing-ground 


860 MR. A. LOVERIDGE: NOTES ON 


of the birds. Sure enough, before I was within fifty yards of it, the 
bird again arose and departed across the water. Its mate generally 
fished near a stone some little way further along the bank, and 
there I espied it; the recognition was mutual, and it winged off 
to a promontory where I could just distinguish it standing in 
the water. 

In pursuance of my original intention, | now lay down in the 
wet grass within range of the stone and here waited for half-an- 
hour until day broke behind a row of tall palms, which later 
formed a lovely picture silhouetted against the rosy dawn. The 
grass, however, was very short owing to the activities of the 
Sanitary Corps, so that as the light increased, realising how 
conspicuous I must soon appear, I crawled away to a shallow 
irrigation ditch fringed with longer grass and within range of 
the buoy. Half-an-hour passed which the mosquitoes relieved of 
all monotony. It was quite a new point of view to see them 
arrive like so many vultures outlined against the sky, and then to 
alight on the stems of grass which were already burdened with 
dew. 

At 64.m., reveille sounded and some K.A.R. askaris from a neigh- 
bouring camp came down to wash clothes, but instead of taking to 
flight at their approach, my heron, now clearly to be seen across 
the shining water, sedately stalked further out. My patience well 
nigh exhausted, T rose on one elk bow, and, seeing how light it had 
become, was on the point of giving up when another native came 
down to the opposite bank, and as it appeared probable that he 
would pass near the bird I resolved to wait a little longer and 
crouched low again. 

A hoarse cry from the Heron caused me to look up; afar I 
beheld the big bird rising from the water. Down I ducked, 
face and hand well into the grass, not daring to move lest I 
should attract attention. A few moments of anxious suspense, 
then swish. swish, swish sounded its pinions close overhead, as it 
circled round preparatory to alighting. Then came a, gentle 
splash as it struck the water. 

Very cautiously I raised my head and looked through the 
grass stems, where, not 40 feet away, stood my victim, sus- 
piciously turning its head from right to left as it strode along. 
Being cramped and stiff, I feared to raise my gun lest I should 
muddle things, and the weary hours of waiting be wasted; so 
again I lay quiet until, in its walk, the bird should come into line 
with the barrels of the gun as it lay on the ground—bang! 

Overhead a raucous squawk and the sound of wings! What 
then. Surely I could not have missed. Hastily and awkwardly 
I rose to my feet. No! There lay the victim of my murderous 
early rising floating quietly on the water, stone dead; a shot had 
passed through its neck and a single spot of blood eed on one 
leg. It floated listlessly to and fro as I waded out to bring it in. 
Far out towards the sea a speck in the sky was all that was to be 
seen of the other bird, which, all unknown to me, had come up 


EAST AFRICAN BIRDS, 86) 


close behind where I was lying, and, at the sound of the shot, had 
risen with acry. The dead bird measured 59 inches from tip 
of beak to the tip of its longest claw (Dar-es-Salaam, 5. vii. 18). 


ScOPUS UMBRETTA BANNERMANNI C. Grant. 

Kast African Hammer-head. At Dar-es-Salaam I found a nest 
from which the young had just flown, according to the natives 
living about 30 feet away. They said that the bird had bred 
there for many years; its nest,in the fork of a tree, was scarcely 
10 feet from the ground and easily accessible. 1 was surprised 
that the thrifty Sees had not pulled clove the load of sticks 
for firewood (8. ii. 19). 


Crconta cicontA Linn. 

The White Stork was only met with once and then in thousands. 
It was at Morogoro (19.11.18), when about 4 P.M. my attention 
was drawn to a great flock of birds high in the air; others were 
coming round the western spur of the Uluguru Mtns. from a 
S.W. direction. It was a most wonderful sight, and reminded 
one of the Gannets flying about the Bass Rock, with this 
difference: that in lieu of the limitless sea for a background, one 
had the forest-clad slopes of the Uluguru reaching up into the 
clouds. And over the top and down the slopes, or circling mid 
the clouds, still the birds came, the specks grew larger and 
larger, then showed white and black until finally they were 
recognisable as storks. 

Thousands upon thousands came sailing through the air 
without visible effort. By scores they settled in the tr ees, stiff and 
erect, their white plumage against the forest foliage forming a 
not easily forgotten picture. It was very evident what had 
brought them, for they pursued the locusts (which had appeared 
the day before) on the wing, swallowing them in wholesale 
fashion. In flight they carried their red legs straight out behind, 
but when intending to alight, they were allowed to hang down 
and swing freely to and fro like a parachute coming to earth. 
There is a curious mechanism of the knee-joints, necessitating 
the leg being out straight or bent at right angles; in either 
position it locks. After death, if the leg was placed in any other 
position, it flew to one or the other of these of its own accord. 

Millions upon millions of the locusts continued steadily driving 
past like rain ; the air was vibrant with the noise of their flight 
and with the steady pinion-beats of the pursuing storks. Others 
of the insects settled on the vegetation, which was a-rustle with 
them as they hopped, or took short flights, out of one’s way. The 
measurement of the outstretched wings of one of the storks was 
6 feet 3 inches. 


ABDIMIA ABDIMIt Licht. 


The following morning the storks were still about, and circled 
over the hospital, where they were made targets of by a number 


862 MR. A. LOVERIDGE ON EAST AFRICAN BIRDS. 


of foolish officers, who blazed away at them with service rifles. 
One of those shot was brought to me, and I recognised it as the 
White-bellied Stork (4. abdimiz). When one considers what 
myriads of locusts such thousands of birds must consume—the 
stomachs of those shot being distended with the insects,—what 
folly it is to destroy them needlessly. 

Someone reported having seen a Secretary Vulture among the 
sterks, which were picking up locusts among the stubble of a 
mealie-field. Numbers of hawks, eagles, and rollers were 
harrying the locusts also. 


Crex crex Linn. 


The Corn-Crake or Land-Rail was taken at Nairobi during the 
first nine days of May 1916 and 15.iv. 19. 


CotuRNIX DELEGORGUE! Hartl. 


A. Harlequin-Quail was obtained under rather peculiar cireum- 
stances, and shows how close these birds will squat. I was 
walking across a patch of grass-grown land in Nairobi when 
something rustled at my very feet ; jumping to one side, expecting 
to see a snake, I found it to be a quail that I had trodden upon 
inadvertently (Nairobi, 2. vu. 15). 


The following records of European migrants may be of 
interest :— 


CHARADRIUS ALEXANDRINUS Linn. Kentish Plover. Dar-es- 
Salaam (13.1. 19). 


CHARADRIUS HIATICULA Linn. Ringed Plover. Nairobi (12. x. 
15); Dar-es-Salaam (14.1. 19). 


TrInNGA suBARQUATA Giild. Curlew-Sandpiper. Dar-es-Salaam 
(14.1. 19). 


TrinGA MINUTA Leisl. Little Stint. Lumbo (5. x. 18). 
CALIDRIS ARENARIA Linn. Sanderling. Dar-es-Salaam (15.1. 19). 


Toranus HypoLeucus Linn. Common Sandpiper. Lumbo 
(27. vii. & 4. 1x. 18). 


Numentus pHmopus Linn. Whimbrel. Dar-es-Salaam; often 
seen in January 1919, but not collected. 


NumeEntus Arquatus Linn. Curlew. Dar-es-Salaam (7. vi. 18); 
Lumbo (25. vii. 18). 


HimANtopus HimMANToPUS Linn. Black-winged Stilt. Tabora ; 
seen only (11. xii. 18). 


H 2MATOPUS OSTRALEGUS Linn. Oyster-catcher. Dar-es-Salaam ; 
often seen during 1918. 


MYOLOGY AND CLASSIFICATION OF THE WOMBAT, ETC. 863 


38. On the Myology and Classification of the Wombat, 
Koala, and Phalangers. By Cnaries F. Sonnrag, 
M.D., F.Z.8., Anatomist to the Society. 


[Received August 1, 1922: Read November 7, 1922. } 
(Text-figures 31-43.) 


ConTENTs, 

Page. 
Heitroducthiontees eee eee eee ee eee SOS. 
MEV OO pyaar ens eae e ante eee eee, OOS 
Classificationyeeee te ee eee en SOG 
pibliograp hye eee eee eee eae eee eo 

Introduction. 


The most remarkable features in the existing classifications of 
the Diprotodont Marsupials* are the positions assigned to the 
Wombat, Koala, and Phalangers. Some zoologists have included 
the Koala in the Phalangeride and kept the Wombat separate. 
Others have included the Koala and Wombat in the Phas- 
colarctide. And others again have formed three families— 
Phalangeridze, Phascolarctide, and Phascolomyidz. It is evident, 
therefore, that there is considerable difference of opinion as to 
the true positions of these animals. To form a lasting and 
natural classification one must examine all anatomical data, so as 
to exclude convergent and adaptive characters, and those which 
represent varying degrees of persistence of primitive conditions 
which were present in their common ancestor. The exclusion of 
these conditions, leaves us with data which are of value for 
purposes of classification. And these can be divided into major 
and minor groups. Minor characters are employed to emphasise 
the classification based on major ones. 

In the examination of animals received at the Society’s 
Prosectorium + I observed so many points in their myology 
hitherto unrecorded, or different from existing accounts, that I 
have considered it fit to set them down as a separate section in 
this paper. As the anatomy of the skeleton, teeth and viscera is 
already well known I have simply mentioned the most distinctive 
characters in the section on classification. 


Myotoey. 
Muscles of the Head and Neck. 


The Platysma in all species is muscular in the face and neck. 
It is continuous with the facial, orbital, and auricular muscles, 


* Pocock (9) and Osgood (7) have reproduced most of the systems of classification 
which have been proposed. 

+ Phascolomys mitchelli ( 2), Phascolarctos cinereus (2), Phalanger orientalis 
(adult female and female mammary fetus), Pseudochirus peregrinus (2 2 2). 
Many points in the anatomy of Trichosurus vulpecula have been described by me in 


a former paper (10). 


864 DR G. F. SONNTAG ON THE MYOLOGY 


but the fusion is best marked in Phascolarctos. Fibres run to 
the pads of vibrissee, but none are continuous with the well- 
developed muscles in the labial labrets in Phascolomys. There is 
no essential point of difference between the platysma in these 
forms and the Phalangeride. 

Masseter :— The general form is similar in Phascolomys (text: fig. 
31 A) and Phascolarctos* ; in the former the anterior border is 
thick and muscular, but it is thin and tendinous in the latter, 
In Phalanger (text-fig. 31 B) and Pseudochirus (text-fig. 31 C) the 
form differs from the above: in the former the surface is 
subdivided by sulci into three parts, but it is undivided in the 
latter. 

Pterygoids :—Macalister (5) observed that the external ptery- 
goid is small and fused with the internal pterygoid in Phascolarctos, 
but I found it separated and well developed in all forms. 

Digastric (text-figs. 32-34 T):—In Phascolarctos 1t is mono- 
gastric and fused with the mylo-hyoid, genio-hyoid, and omo- 
hyoid; and it is inserted into the mandible. I have already 
shown that it enters into a sheet with mylo-hyoid and sterno- 
hyoid (10), and I believe that the lateral fibres of the sheet 
correspond to the digastric. Young (14) described two bellies 
and a tendinous inscription. In Phascolomys the broad anterior 
bellies conceal the mylo-hyoid, and strong intermediate tendons 
unite them to the posterior bellies. In Phalanger the broad 
anterior bellies do not completely cover the mylo-hyoid, and the 
posterior bellies are fusiform. In Pseudochirus the monogastric 
muscles conceal the greater part of the mylo-hyoid. In no species 
has the digastric any connection to the hyoid bone. There is, 
therefore, a close resemblance between the digastric muscles in 
Phascolarctos and Pseudochirus. 

Mylo-hyoid (text-figs. 32-34, M-H.M):—In Phascolomys and 
the Phalangers the mylo-hyoid runs from the mandible to the 
hyoid bone. But in Phascolarctos it does not get any attach- ~ 
ment to the hyoid. In my specimen it is fused with the 
sterno-hyoid ; and there is, in reality, a sterno-mandibularis. 
Young (14) does not mention fusion of mylo-hyoid and sterno- 
hyoid, but notes that the mylo-hyoid has no attachment to the 
hyoid bone. 

Omo-hyoid (text-figs. 32-34, O-H.M) :—In Phascolomys and the 
Phalangers it is attached to the hyoid bone, but no central tendon 
is present. In Phascolarctos, however, it avoids the hyoid bone 
and passes forwards to enter the tongue, wherein it forms a 
lingualis. 

The ansa hypoglossi, which supplies the omo-hyoid and _ pre- 
tracheal muscles, is derived from the cervical plexus alone in 
Phascolarctos, Phascolomys, and Pseudochirus. In Phalanger 1 
observed the nerve receiving connections from the first three 
cervical and the hypoglossal nerves (text-fig. 31 D). 


* P. ZS, 1921, p. 548, text-fig. 538. + Ibid., p. 549, text-fig. 54, 


OF THE WOMBAT, KOALA, AND PHALANGERS. 865 


 Pretracheal Muscles:—In all the animals described in this 
paper the sterno-hyoid muscles could easily be separated from the 
subjacent sterno-thyroids. The sterno-hyoids are fused in 
Phascolurctos and have no connection to the hyoid bone, so they 


Text-figure 31. 


Seas Ss | 
MASSETER” “ss oe 
The masseter muscle in Phascolomys mitchelli (A.), Pseudochirus peregrinus (B.), 
and Phalanger orientalis (C.); D: the ansa hypoglossi in Phalanger 
orientalis. 
C2 and C8: cervical nerves; A-H: ansa hypoglossi; P.G: parotid glands 
XII: hypoglossal nerve. 


play no part in its movements. But in all other species they are 
separate and receive an insertion to the hyoid. The sterno- 
thyroids are essentially similar in all. 

Hyoglossus :—Young (14) describes it as a transverse band, 
bridging over the genio-glossi, and having no hyoid attachment 


866 DR. C.F. SONNTAG ON THE MYOLOGY 


in Phascolarctos. I did not, however, observe this band at all. 
Macalister (5) regarded it as a piece of the mylo-hyloid. In. 
Phascolomys and the Phalangers it runs from the hyoid bone to 
the tongue. 


It is, therefore, evident that the Koala has many traces of 


Text-figure 32. 


an l 
-M.M_ O-H.M. 


S-M.M CL 
Anatomy of the neck in Phascolomys mitchelli. 


CL: clavicle; CL-M.M: cleido-mastoid; C.L.T: cervical thymus; Dig.M: 
digastric; E.J.V: external jugular vein; L.C.C.A: left common carotid 
artery; M-H.M: mylo-hyoid; M.M: masseter; O-H.M: omo-hyoid; 
P.F.V: posterior facial vem; S-H.M: sterno-hyoid; S-M.G: submaxillary 
gland; S-M.M: sterno-mastoid; T.M: trapezius; XII; hypoglossal 
nerve. 


primitive lamination in its platysma, mylo-hyoid, sterno-hyoid, 
and hyoglossus muscles. And in these conditions it differs 
entirely from Phascolomys and the Phalangers. 

Sterno-mastoid, Cleido-mastoid, Cleido-occipitul (text-fig. 32, S-M. 
M, Cl-M, Cl-O):—All are present in the Phalangers, but the 


OF THE WOMBAT, KOALA, AND PHALANGERS. 867 


last is absent in Phascolarctos and Phascolomys. The insertion of 
sterno-mastoid and cleido-mastoid is into the occipital crest. 
Genio-hyoid :—In my specimen of Phascolarctos it runs from the 
symphysis of the mandible to the hyoid bone and thyroid carti- 
lage, so it is in reality a genio-hyo-thyroid. This arrangement 


Text-figure 33. 


Anatomy of the neck in Phalanger orientalis (adult). 


CL-O: cleido-occipital ; Pect.Maj: pectoralis major; T.G: thyroid gland. Other 
letters as in last figure. In a mammary foetus the cervical thymus concealed 
the lower part of the neck. Note the vertical anterior jugular veins and the 
horizontal vena transversa. 


differs from Young’s account (14). As no other thyro-hyoid 
muscles are present, they are probably replaced by a part of the 
genio-hyoid. 

The genio-hyoids and stylo-hyoids are the only muscles 


868 _ DR. Cc. F. SONNTAG ON THE MYOLOGY 


attached to the hyoid bone in my specimen of Phascolarctos, for 
the pharyngeal constrictors have no hyoid origin. 

Scaleni:—In all forms the sealenus anticus is absent, as the 
scalene group is behind the brachial plexus. The scalenus medius 
and posticus are present. 

Splenius:—In Phascolarctos it is divisible into two parts 
(capitis and colli), but it is a single sheet in Phascolomys and the 
Phalangers. 

Prevertebral Muscles:—Young (14) described these muscles 
in Phascolarctos, and pointed out that they are more or less 


Text-figure 34. 


Mi 


WLU! 
Anatomy of the neck in Pseudochirus peregrinus. 


Letters as in text-fig. 32. 


inseparable. The rectus capitis anticus major is the largest; it 
runs from the bodies of the first three dorsal vertebre and all 
cervical transverse processes to the basi-occiput. Rectus capitis 
anticus minor cannot be isolated. Longus colli consists of fibres 
running between the bodies and transverse processes of the first 


OF THE WOMBAT, KOALA, AND PHALANGERS. 869 


* three dorsal and cervical vertebre. In Phascolomys I observed 
similar fusion of these muscles. A single thick muscular sheet 
extends from the third dorsal vertebra to the basi-occiput 
(text-fig. 35). A small bundle of fibres becomes detached, and 
runs to the front of the third cervical vertebra when it fuses with 
the bundle from the opposite muscle. The main mass, in my 
opinion, represents rectus capitis anticus major. The mesial part 
represents longus colli, and the rectus capitis anticus minor is 
absent. In Phalanger there is more separation between the longus 
colli and rectus capitis anticus major, and there is no rectus 
capitis anticus minor. But in Psewdochirus there is considerable 


Text-figure 35. 


Prevertebral muscles in Phascolomys mitchelli. 


Letters in text. 


fusion. It appears, therefore, that the conditions in Phalanger 
do not adhere so much to the primitive condition as do those in 
Phascolarctos, Phascolomys, and Pseudochirus. 


Muscles of the Back, Thorax, and Abdomen. 


Trapezius :—In all forms the origin extends from the occiput 
to the seventh dorsal spine, although Young (14) gave the eighth 
spine as well in Phascolarctos. The clavicular insertion varies, 
however. In Phascolarctos and Pseudochirus most of the fibres go 
to the clavicle, but some cross it and run to the clavicular deltoid, 


Proc. Zoou. Soc.—1922, No. LIX. 59 


870 DR. C. F. SONNTAG ON THE MYOLOGY 


In Phascolomys, on the other hand, most of the fibres cross the 
clavicle and fuse with the clavicular deltoid ; and some fuse with 
the cleido-mastoid. In Phalanger the conditions are intermediate. 
The scapular insertion is, however, similar in all. And in no case 
is there a tendinous intersection where the fibres cross the 
clavicle. The fusion of the trapezius and clavicular deltoid is a 
persistence of the primitive lamination, and is best marked in 
Phascolomys. 

Rhomboideus :—In all forms it is an indivisible sheet whose 
origin extends from the occiput anteriorly to the thoracic region 
posteriorly, and it thickens from before backwards. It reaches 
the fifth dorsal spine in Phascolomys, but stops at the third 
in Phascolarctos, Phalanger, and Pseudochirus. As the rhom- 
boideus shows no division in any of these animals they all exbibit 
a persistence of the primitive condition of the muscle. 

Latissimus Dorsi:—This muscle arises from a variable number 
of posterior thoracic spines and the lumbar fascia, but there are 
neither iliac nor scapular origins. Costal slips may exist. The 
following origins were present in my specimens :— 


Phascolarctos : spines 4-10; no costal origin. 
Phalanger : spines 4-12; slip from last rib. 
Pseudochirus : spines 4-12; no costal origin. 
Phascolomys : spines 5-15; slips from lower six ribs. 


The insertion of the latissimus 1s very similar in all, and it is 
intimately connected to the teres major and dorso-epitrochlearis. 

Levator Anguli Scapule :—Young (14) states that it arises from 
the front of the lateral mass of the atlas along with the first slip 
of the serratus magnus in Phascolarctos ; but he does not mention 
whether it is fused with the omo-atlantic. It is inserted into 
the scapular spine. In Phascolomys I observed it arising in 
company with the omo-atlantie from the caudal border of the 
lateral mass (text-fig. 36 A). And it is closely connected to the 
upper part of the serratus magnus. It is attached to the root of 
the spine of the scapula. In Phalanger (text-fig. 37) it arises in 
common with the omo-atlantic, but 1s separate from the serratus 
magnus. It is inserted into the dorsal surface of the scapula in 
the anterior part of the supraspinous fossa. In Pseudochirus, on 
the other hand, itis inseparable from the serratus magnus, and its 
insertion never reaches the spine of the scapula. It is, therefore, 
evident that Phascolomys and Phascolarctos difter from the 
Phalangers in the insertion of the levator scapule. So the 
conditions of the levator scapule and serratus magnus are degrees 
of persistence of primitive lamination. 

Omo-atlantic (text-figs. 36 & 37):—In Phascolomys and Phasco- 
larctos it runs, widening as it goes, from the lateral mass of the 
atlas to the outer part of the spine of the scapula. And the 
lateral fibres cover the dorsal part of the shoulder joint. In 
Phalanger and Pseudochirus it divides into two parts—a broad 
mesial and a narrow lateral slip—~and these are attached to the 


OF THE WOMBAT, KOALA, AND PHALANGERS. 871 


outer half of the scapular spine. Cunningham (2) describes it as 
two muscles fixed to the whole length of the seapular spine 
in Cuscus. The conditions in the Phalangers cannot be due to 
their leading an active arboreal life, as other active animals 
have a single muscle. They are probably remnants of primitive 
laminations. 


Text-figure 36. 


Scapular muscles in Phascolomys mitchelli. 


I-S: infra-spinatus; L.A.S: levator anguli scapule ; L.H.T: long head of triceps; 
O-A: omo-atlantic; S.M: serratus magnus; S-S: supra-spinatus. 


Serratus Magnus (text-figs. 36-38) :—In the Marsupialia there 
is a more or less intimate connection between the levator scapule 
and the serratus magnus, thus showing that these muscles are 
part of the same sheet. And the origin of the serratus extends 
from the anterior cervical to the posterior thoracic region. In 
Phalanger the origin extends from the second cervical vertebra 
to the seventh rib. The cervical slips are all distinct. In 
Pseudochirus there are eight thoracic slips. The cervical slips are 


a9* 


872 DR. C. F. SONNTAG ON THE MYOLOGY 


also coarser than the thoracic ones, but they are closer together. 
In Phascolarctos Young (14) described six cervical and nine 
thoracic slips, and I observed the same. But Macalister (5) 
recorded seven costal slips. As regards Phascolomys, Macalister 
pointed out that the cervical and thoracic parts are separate, and 
he described them in detail. 


Text-figure 37. 


INT 
AN 


Scapular muscles in Phalanger orientalis. 
T.Ma: teres major. Other letters as in text-fig. 36. 
Intercostal Muscles :—The number of sets of intercostals varies, 
as the result of variations in the number of ribs, but the 
characters of the muscles are essentially similar in all :— 


Phascolarctos:; ten pairs of interspaces. 
Phascolomys: fourteen pairs of interspaces. 
Phalangeride ; twelve pairs of interspaces. 


OF THE WOMBAT, KOALA, AND PHALANGERS. 873 


Muscles of the Ventral Abdominal Wall :—In the Phalangers at 
my disposal the muscles of the ventral abdominal wall were all 

firmly united together and the linea alba was very obvious. ‘The 
rectus abdominis had eight intersections, and the first one was 
so wide that there is apparently a rectus abdominis and rectus 
thoracis. This condition was also present in Young’s specimen of 
Phascolarctos (14). In Phascolarctos and Phalanger the pyrami- 
dalis conceals the rectus, and gets one insertion into the sternum 


Text-figure 38. 
7 
HiT 


~ 
— 
ot 


SN 


Scapular muscles in Phascolanctos cinereus. 


Letters as in text-fig. 36. 


and last costal cartilage, but it is not so extensive in Phascolomys. 
These animals are, however, essentially similar as regards all the 


muscles of the ventral abdominal wall. 
Dorsal Abdominal Muscles :—The quadratus lumborum is well 


developed in Phascolomys and the Phalangers. 


In Phascolarctos 


it is more primitive, for it is practically absent, or, as Young (14) 


suggests, it may be replaced by intertransverse muscles connecting 


874 DR. OC. F. SONNTAG ON THE MYOLOGY 


the lumbar vertebre. The psoas parvus exceeds the magnus 1 
all forms, but the disproportion is not so great, and the parvus 1s 
more tendinous in Phascolomys. Macalister (18) describes it as a 
weak muscle in the latter. It is inserted into the pubis behind 
the corresponding marsupial bone in all forms. 


Muscles of the Anterior Huxtremity. 


The Dorso-epitrochlearis arises from the posterior border of the 
latissimus dorsi and its tendon in Phalanger and Pseudochirus, 
and it remains muscular to its insertion into the tip and inner 
border of the olecranon. In VPhascolarctos it is thin, and a 
considerable part is fascial. In Phascolomys it arises chiefly from 
the tendon of the latissimus, and it is firmly bound down by deep 
fascia. When the fore-limb is abducted the muscle becomes tight 
soonest in the latter. And the distant extremity is more 
evidently continuous with the anconeus internus in Phascolarctos. 

Pectoral Muscles :—Cunningham (2) analysed the pectoral mass, 
and described a pectoralis major composed of superficial and deep 
parts, a pectoralis minor, and a pectoralis quartus. In my 
specimens the conditions present differed in some respects from 
those described by Young, Cunningham, Macalister, and others. 

Pectoralis Major :—In Phascolomys Macalister (18) showed that 
the superficial part has clavicular, sternal, and costal origins. In 
my specimen (text-fig. 39 A) it arises from the mid-sternal line 
and fascia over the abdominal muscles in the infra-sternal fossa ; 
there is no clavicular nor costal origin. It is inserted into the 
peetoral crest of the humerus and fuses extensively with the 
clavicular deltoid. The deep part arises from the manubrium 
sterni, crosses the pectoralis minor and quartus, and is inserted 
by a narrow tendon into the outer tuberosity and upper part of 
the pectoral crest. It is separated from the first part. In 
Phalanger the superficial part does not arise from the entire 
length of the sternum, but in other respects it resembles the 
superficial part in Phascolomys. The deep part has a more 
extensive origin from the body of the sternum. In Pseudochirus 
the conditions are similar to those in Phalanger, but it is difficult 
to distinguish the superficial part from the clavicular deltoid in the 
fused muscular mass. In Phascolarctos the superficial part has an 
origin from the inner third of the clavicle, so the clavicular deltoid 
does not reach the sterno-clavicular joint as it does in Phascolomys. 
I have already described and figured muscles resembling those in 
the latter in Mandritlus leucopheus. 

Pectoralis Minor:—In Phascolomys it arises from the posterior 
three-fourths of the sternum, and it is inserted into the great 
tuberosity, coraco-humeral ligament, and coracoid process. It 
remains separate from the quartus, but their insertions are 
contiguous. In Pseudochirus the minor and quartus are in 
contact throughout the greater part of their course. But in 
Phalanger and Phascolarctos they are separate. 

Subclavius :—The insertion varies. In Phalanger it is attached 


OF THE WOMBA?, KOALA, AND PHALANGERS. 875 


to the outer two-thirds of the anterior border of the clavicle. 
In Pseudochirus the conditions are similar to the above. In 


Clavicular muscles in Phascolomys mitchelli (A.), Phascolarctos cinereus (B.), and 
Phalanger orientalis (C.). 
Cl.M: cleido-mastoid; D.D).D 2: deltoid; H.H: head of humerus; P.M; 
pectoralis major; S-M: sterno-mastoid; $.D.F: slip of deltoid to forearm: 
T.R: trapezius. 


Phascolarcios it runs to the outer sixth of the clavicle. The 
attachment is more extensive in Phascolomys, for it is fixed to the 


876 DR. GC. F. SONNTAG ON THE MYOLOGY 


outer sixth of the clavicle, the upper border of the acromion, and 
the spine of the scapula. I believe the extent of this muscle is 
partly due to varying degrees of functional activity. 

Deltoid (text-fig. 89 A-C) :—In Phascolomys the origin forms a 
continuous line along the whole length of the clavicle, the 
acromion and lateral third of the spine of the scapula. The 
clavicular fibres receive the trapezius, and the acromial fibres and 
trapezius form a powerful cephalo-humeralis. The clavicular 
deltoid also fuses with the superficial part of the pectoralis major, 
and the two are inserted into the pectoral crest. From the 
scapular fibres a long, narrow, strap-like band runs to the deep 
fascia over the radial border and dorsum of the forearm and 
carpus. In Phascolarctos the muscle is also entire, the clavicular 
origin is not so extensive, and the narrow band given off fuses 
with the supinator longus. In Phalanger and Pseudochirus the 
clavicular origin is not as extensive as in Phascolomys, the 
clavicular and acromial parts are separate, and no slip is given off 
to the forearm. 

Scapular Muscles (text-figs. 36—38):—I agree with Macalister 
(18) that the supraspinatus is larger than the infraspinatus in 
Phascolomys and the Phalangers, but I observed the reverse in 
Phascolarctos. Young (14) states that the infraspinatus isslightly 
larger than the supraspinatus. The subscapularis and teres major 
are essentially similar in all, and there is a certain amount of 
fusion between the latter and the latissimus dorsi and dorso- 
epitrochlearis. ‘The teres minor is fascial in Phascolomys, thin 
and with a tendinous attachment to the scapula in Phascolarctos, 
and thin and muscular in Phalanger and Pseudochirus. Kven 
when it is muscular there is a considerable degree of fusion with 
the infraspinatus. Meckel (6) states that the absence of a teres 
minor as a distinct muscle appears to be general amongst 
marsupials. 

The characters of the Coraco-brachialis in many forms, including 
the animals described in this paper, have already been recorded, 
and Cunningham (2) has collected. the observations. My own 
observations do not differ from his. 

Biceps :—The two heads of origin are in contact, but not fused, 
in Pseudochirus, and the two bellies (gleno-ulnar and coraco- 
radial) are fused from the middle of the arm onwards. In the 
distal portion of the arm the two parts separate again and run to 
the radial tuberosity and coronoid. In Phalanger the tendons of 
origin are fused to form a tendinous sheet. The muscle bellies 
remain fused to the distal part of the arm where the stout radial 
and slender ulnar components are given off. In Phascolomys and 
Phascolarctos the origins are conjoined, but the bellies separate 
sooner from one another in the latter. 

Brachialis Anticus:—In all forms it arises on the outer 
surface of the shaft of the humerus, and winds round it to be 
inserted into the ulna with, or deep to, the gleno-ulnar component 
of the biceps. In Phascolarctos it is fused with the biceps, but 


Of THE WOMBAT, KOALA, AND PHALANGERS. 8i7 


it is separate from it in Phascolomys and the Phalangers. 
Macalister (18) showed that the conditions in Phascolomys are 
similar to those in many Polyprotodonts and Diprotodonts. 

Triceps (text-figs. 36-38) :—In Phascolomys I observed the long 
head arising from the lateral two-thirds of the axillary border of 
the scapula, and the humeral heads are indistinguishably fused. 
Macalister (8) stated that the long head arises from more than a 
third of the bone. In Phascolarctos the long head arises from a 
third of the axillary border of the scapula, and it passes to the 
distal third of the arm before it unites with the fused humeral 
heads. In Phalanger and Pseudochirus the long head also arises 
from a third of the axillary border, but it unites high up with the 
humeral heads. é 

_ Anconei:—The anconeus internus, according to Cunningham (2), 

is generally present in the Marsupialia. In Phascolomys, 
Phalanger, and Pseudochirus it is a marked pavallel-sided band 
running from the internal condyle to the olecranon, and the 
ulnar nerve passes under cover of it. In Phascolarctos it appears 
to be more continuous with the dorso-epitrochlearis. In Phasco- 
larctos and Phascolomys it is not connected to a fibrous band 
which gives origin to the flexor carpi ulnaris ‘he anconeus 
externus is present in all forms, but it is least in Phascolomys, in 
which it has no connection with the triceps. 

Pronator Radii Teres:—Cunningham (2) pointed out that this 
muscle has no coronoid head in the Marsupialia. ‘The insertions 
vary. In my specimens the conditions are as follows :— 


Phascolarcios: to middle two-fourths of the radia! shaft. 
Phalanyer : to distal two-thirds of the radial shaft. 
Pseudochirus: to distal two-thirds of the radial shaft. 
Phascolomys : to distal half of the radial shaft. 


flexor Carpi Radialis:—This muscle is well developed. It 
exhibits a variable amount of fusion with the other members of 
the superficial flexor group, but the fusion is least in Phascolarctos. 
It is inserted into the base of the second metacarpal in Phasco- 
lomys, Phalanger, and Pseudochirus, but it runs to the third 
metacarpal in Phascolarctos. 

Palmaris Longus is present in all. In Phascolomys Macalister 
(18) described a true palmaris longus and a palmaris accessorius, 
both arising from the internal condyle. In my specimen the 
accessorius does not arise from the condyle, but it is implanted 
into the side of the tendon of palmaris longus in the distal third 
of the forearm; and it has a very slender tendon which runs to 
the palmar pad. In Phascolarctos, Phalanger, and Pseudochirus 
there is no trace of the accessorius. 

Hlexor Carpi Ulnaris:—In all the animals described in the 
present paper the insertion is into the pisiform bone. But 
Macalister (18) described it as being attached to the fifth meta- 
carpal in Phascolomys. Young (14) describes it as giving off 


878 DR. C. F. SONNTAG ON THE MYOLOGY 


prolongations to the fifth metacarpal and the unciform in Phasco- 
larctos. 

Hlexor Sublimis Digitorwm :—This muscle is frequently strongly 
connected to the flexor profundus, and it ig sometimes called the 
flexor perforatus because the profundus tendons pierce its tendons, 
which are very slender. he sublimis and profundus tendons 
separate out from the strong flexor mass in the distal part of the 
forearm. 

Pronator Quadratus:—The extent of the interosseous space 
occupied by the pronator quadratus varies considerably. In 
Phascolomys 1 observed it extending over the distal half of the 
space, but Macalister says it 1s weak and only occupies the distal 
third. In Phascolarctos it occupies the lewer fifth, but the extent 
is even less in Phalanger and Pseudochirus. Cunningham (2), 
however, described it as covering one third of the bones in Cuscus 
(Phalangisia maculata). 

Supinator Longus :—My observations on Phascolarctos difter in 
some respects from those of Macalister (5) and Young (14). It 
consists of two parts. The proximal division arises from the 
shaft of the humerus from close to the neck downwards, and it is 
thin and almost aponeurotic. The distal pait, which is more 
muscular, arises frem the lateral supracondylar ridge and by 
fibres which fuse with the extensors of the wrist. The two parts 
unite into a strong tendon which, passing under the abductor 
pollicis major, is inserted into the scapho-lunar bone. Young 
points out that it is both a radial flexor and supinator. It receives 
a slip from the deltoid. In Phalanger it is also large, but not 
divided into two parts. It springs from the prosimall two-thirds 
of the lateral supracondylar ridge. No slip runs into it from the 
deltoid. These conditions are also present in Pseudochirus. The 
conditions in Phascolomys are, however, very different. It is 
thin and aponeurotic in parts, and it receives a strong band from 
the deltoid. 

Supinator Brevis :—The extent of the radius embraced by this 
muscle varies. In Phalanger it covers less than the upper fifth, 
in Phascolarctos it covers the upper fifth, and in Phascolomys it 
envelops the upper two-thirds. 

Hatensores Carpi Radiales:—The existing records show that 
there is considerable variation in these camels in the Marsupialia. 
In Phascolomys there is a single muscle arising from the external 
condyle and lateral ridge of the humerus and running to be 
inserted into the second and third metacarpal bones. But it is 
evident that the long and short extensors are combined in the 
single muscle. In Phascolarctos both long and short extensors are 
present and are inserted into the radial borders of the second and 
third metacarpals respectively. In Phalanger both muscles are 
present, but the brevior is a large and powerful musele with 
three heads of origin as described by Cunningham (2). 

Extensor Communis Digitorum :—In all forms it arises from 
the external condyle. In most Marsupialia it sends four tendons 


OF THE WOMBAT, KOALA, AND PHALANGHRS. 879 


to the four inner digits. But, as Young (14) mentions, and I 
observed myself, tendons go to each of the five digits in Phasco- 
larctos. 

Extensor Secundus Digitorwm :—This muscle, which corresponds 
to the extensor minimi digiti of human anatomy, has been fully 
described in many Marsupialia by Cunningham (2), Young (14), 
and Macalister (5 & 18). In Phascolarctos it is in reality double. 
One muscle runs to the third digit as an extensor medius, and 
the other is inserted into the fourth and fifth digits. In Phasco- 
lomys and the Phalangers only the latter part is present. 

Extensor Carpi Ulnaris:—In Phascolomys it arises from the 
external condyle of the humerus and upper part of the shaft of 
the ulna, and is inserted into the mesial border of the shaft of the. 
fitth metacarpal bone. In Phascolarctos the ulnar origin head is 
similar to the above. But in Phalanger there is no ulnar head. 

When a superficial view of the extensor muscles is taken in all 
the animals described in this paper, it is seen that intermuscular 
septa are well marked in Phascolarctos. In Phascolomys and the 
Phalangers there is more fusion of the bellies of the muscles. 

Extensor Ossis Metacarpi Pollicis:—Vhis muscle, as has already 
been pointed out by several authors, includes the extensor primi 
internodii pollicis. In Phascolomys, according to Macalister (18), 
it arises from the dorsum of the shaft of the ulna and the inter- 
osseous membrane, crosses the extensor carpi radialis tendon, and 
is inserted into the trapezium and thumb metacarpal. In Phasce- 
larctos I observed it arising from the dorsum of the proximal 
third of the part of the ulna bordering the interosseous space, 
from the membrane, and from the greater part of the dorsum of 
the radius from the orbicular ligament distally. In Phalanger 
and Pseudochirus it also arises from both bones and the inter- 
osseous membrane, but the ulnar origin is lower and more 
extensive than in Phascolarctos, and the muscle belly gives off two 
tendons which wind over the extensor carpi radialis and supinator 
longus tendons. 

Extensor Secundi Internodii Pollicis: There is no trace of this 
muscle in Phascolarctos, and in this respect I agree with Young 
(14) and differ from Macalister (5), who stated that it is arranged 
as usual. In Phascolomys it runs from the lower third of the 
ulna to the last phalanx of the pollex. The tendon gave a short 
slip to the fascia over the metacarpo-phalangeal joint of the index. 
In Phalanger it also gives slips to the index and pollex, and 
Cunningham (2) describes two separate muscles in Phalangista 
maculata. It is therefore evident that the only extensor of the 
pollex in Phascolarctos is the additional tendon of the extensor 
communis digitorum ; and in no other Marsupial is this arrange- 
ment present. 

Intrinsic Muscles of the Manus:—The observations of 
Cunningham (2), Macalister (5, 18), Ruge and Young (14, 15) 
have given us a very complete account of the anatomy of these 
muscles. Young, in particular, has shown how they have become 


880 DR. C. F. SONNTAG ON THE MYOLOGY 


modified in accordance with habits. He has also shown that the 
Koala is very different from the others, his conclusions being as 
follows :—‘‘ The digits form two groups, of which one, including 
the thumb and index finger, is opposable to the other, ‘this latter 
comprising the three remaining digits, ‘The index digit, in point 
of fact, constitutes a second thumb; and, in comformity with its 
unusual freedom of action, its special muscles are well developed. 
Moreover, as the resuit of the arrangement of the digits into two 
groups, the middle line of the hand no longer passes through the 
third digit, but along the fowrth. The muscles of the hand are 
correspondingly modified, and hence, so far as regards their 
insertions, the arrangement is very different to what is ordinarily 
found.” 


Muscles of the Posterior Hautremities. 


Gluteus Maximus :—In Phascolarctos it has no origin from the 
iliac crest, but springs from the sacrai and caudal vertebre. The 
coarse fibres sweep over the great trochanter, thereby producing 
a rounded prominence, and are inserted into a considerable length 
of the femoral shaft. The fibres are continuous postero-mesially 
with the lateralis caude. In Phascolomys a broad, long agitator 
caude shuts off the gluteus maximus from the sacral and caudal 
vertebral spines, snl a strong intermuscular septum, which gives 
origin to some of the fibres of both muscles, intervenes between 
them. The gluteus maximus also arises, as in all Marsupialia 
except the Koala, from the iliac crests posteriorly, and from the 
lumbar fascia, The whole muscle appears flattened, and it is 
inserted into the back part of the great trochanter. Cunningham 
(2) describes how the gluteus maximus is divided into three parts 
in Phalangista maculata, but there 1s no obvious separation into 
distinct parts in my specimen of Phalanger orientalis. There it 
is a long fan-shaped sheet composed of gluteus maximus, tensor 
fascize femoris, and agitator caude. It extends from the lumbar 
fascia anteriorly backwards along the iliac crest and spines of the 
sacral and anterior four caudal vertebre. The part of the sheet 
representing the agitator caude has much coarser fibres than the 
other ere hey The insertion of the sheet does not differ in any 
essential point from that in Phalangista maculata, already 
described by Cunningham. In Pseudochirus the conditions 
resemble those in Phalanger, but the fibres of the agitator caude 
are not so coarse, or so separate from the other parts of the 
muscle. 

Gluteus Medius:—In all forms it is large, and frequently 
exceeds the maximus in size. In Phascolarctos it arises from the 
iliac crest, the gluteal surface of the ilium, and the sacral spines. 
It is inserted into the great trochanter. The fibres are in parts 
fused with the subjacent gluteus minimus, but there is no 
lamination as described by Young (14). In Phascolomys it is 
difficult to separate the medius and mimimus. In Phalanger and 
Pseudochirus the medius and minimus are separate. The Gluteus 


OF THE WOMBAT, KOALA, AND PHALANGERS. 881 


Minimus does not differ in any essential in the animals described 
in this paper except for the degree of adhesion to the medius. 
The Gluteuws Quartus is present in all. 

Pyriformis :—In all forms this is a strong triangular muscle 
arising from the side of the sacrum and running to the summit 
of the great trochanter of the femur. The Obturator Haternus is 
likewise similar in all, and is large. 

Obturator Internus:-——This muscle is absent in Phascolomys, but 
present in all other forms. The Gemelli are present in all. They 
are both enormous in Phalanger, of moderate size in Phascolarctos, 
and the inferior one is large in Phascolomys. 

The Quadratus Femoris is absent in Phascolomys. In Phasco- 
larctos it arises entirely from the great sciatic ligament. In 
Phalanger it arises from the tuber ischii. The insertion is into 
the posterior border of the great trochanter. 

Sartorius :—In the Marsupialia this muscle acts as an extensor, 
and an interval, filled with fascia, is present between its upper 
part and the quadriceps. It arises in all from the anterior 
superior iliac spine; and it is inserted into the inner side of the 
quadriceps tendon (Phascolarctos), or the inner aspect of the patella 
and knee-joint (Phalanger, Pseudochirus, Phascolomys). 

Quadriceps Hatensor :—The rectus femoris has only the straight 
head in Phascolomys, but both heads are present in Phascolarctos, 
Pseudochirus, and Phalanger. In the other elements these 
animals are all essentially similar, and the vastus externus 
component is large. The tendon is inserted in all into the tibia. 
Tt contains a cartilaginous patella in Phascolomys, Phalanger, and 
Pseudochirus, but none in Phascolarctos. Waterhouse (17) stated 
that Phascolomys also has no patella, and used this character 
among others to show that they are related. As many observers 
have recorded the presence of the patella in the Wombat, it is 
evident that this is not a character linking it to the Koala, as 
Waterhouse believed. 

Gracilis :—When the skin is removed it is seen that the muscles 
on the mesial aspect of the thigh are concealed by the gracilis to 
a variable extent. It arises from the symphysis pubis and a 
variable extent of the descending ramus, and from the mesial 
extremity of the marsupial bone. It is inserted into the 
proximal half of the mesial border of the shaft of the tibia. In 
Phascolomys, Phalanger, and Pseudochirus it goes to the proximal 
half of the tibia, but in Phascolarctos it only runs to the proximal 
third. 

Biceps Cruris:—In Phascolarctos the biceps and semimem- 
branosus have a strong, common tendon of origin from the ischial 
tuberosity and caudal vertebrae. The biceps is thin and triangular, 
and inserted into the fascia over the proximal two-thirds of the 
fibula. No femoral component enters into the biceps. Young 
(14) points out that the nature and extent of the insertion are 
associated with the semi-flexed attitude of the limb. In Phalanger 
the muscle arises from the ischial tuberosity, along with semi- 


882 DR. C. F. SONNTAG ON THE MYOLOGY 


membranosus, and from the caudal vertebre, the two parts being 
separate. No femoral head is present. The ischial part expands 
and becomes fan-shaped ; it is inserted into the fascia over the 
fibula in the proximal two-thirds of the leg. The caudal part is 
at first superficial to the ischial part, but winds round it and 
divides into two portions. One of these fuses with the ischial 
part. The other fuses with semitendinosus, and then is inserted 
into the middle of the subcutaneous mesial surface of the tibia. 
The conditions present differ in several respects from Cunningham’s 
account of the muscle in Phalangista maculata. In Pseudochirus 
the conditions are essentially similar to those in Phalanger. In 
Phascolomys it arises entirely from the ischial tuberosity along 
with semimembranosus, and it is inserted into the fascia over the 
proximal third of the fibula, and into the bone itself, but it is not 
divisible into two parts as in the Phalangeride. No femoral 
head is present. Its insertion is less than in Phascolarctos and 
the Phalangeride. 

No bicipiti accessorius is present in any of these animals. 

Semimembranosus (text-fig. 40):—In all forms it is muscular 
from origin to insertion, and there is a variable amount of union 
to the biceps ; and the course in all is from the tuber ischii to the 
mesial aspect of the leg. In Phascolarctos it is inserted into the 
quadriceps tendon, internal tuberosity of the tibia, and the fascia 
of the leg; and some of the fibres fuse with the triceps adductor 
femoris. In Phalanger the insertion is moved farther distally. 
It avoids the adductor and quadriceps, passes under the internal 
lateral ligament of the knee, and is attached to the anterior 
tuberosity of the tibia. And, as Cunningham (2) pointed out in 
Phalangista maculata, it must rotate the leg on the thigh and act 
as a powerful flexor. In Phascolomys it is inserted into the 
mesial aspect of the head of the tibia. 

Semitendinosus (text-fig. 40):—In Phalanger it is inserted into 
a narrow strip of the middle of the ventral border of the shaft of 
the tibia. But Cunningham (2) described it as going to the 
mesial surface of the bone in Phalangista maculata. In Phasco- 
larctos it is inserted into the mesial aspect of the tibia at the level 
of the prominent tubercle on the anterior (ventral) border. In 
Phascolomys it runs to the mesial border of the tibia in its middle 
third. No tendinous inscription is present in any of these 
Marsupialia. The muscle fuses with the biceps in Phalanger and 
Phascolomys, but is quite free in Phascolarctos. But Macalister 
(18) described it as free from other muscles in Phascolomys and 
Phalanger. 

Gastrocnemius :—In Phalanger, as in Phalangista maculata, the 
two parts from origin to insertion are quite separate. The inner 
head arises from the back of the internal condyle of the femur, 
and its tendon is inserted into the tuberosity of the os calcis. The 
outer head arises from the proximal sixth of the shaft of the 
fibula and from the intermuscular membrane between it and 
the peronei; its tendon ig inserted into the tuberosity of the 


OF THE WOMBAT, KOALA, AND PHALANGERS. 883 


os caleis deep to that of the mesial part of the muscle. Only a 
thin membrane connects the two parts. In Phascolarctos the two 
parts arise as in Phalanger ; at the junction of the proximal and 
middle thirds of the leg a strong, narrow aponeurosis, which 
ultimately forms the tendo Achillis, begins to appear on the outer 
part of the muscle, and the inner part is attached to it, In 


Text-figure 40. 


Adductor and hamstring muscles of Phascolarctos cinereus (above), and Phalanger 
orientalis (below). 
A.M: adductors; B: biceps; G.S.N: great sciatic nerve; I.L.L: internal lateral 


ligament of knee; Q.E: quadriceps extensor; S-M: semi-membranosus ; 
S-T : semi-tendinosus. 


Pseudochirus the two parts unite, so it resembies that in Phasco- 
larctos rather than Phalanger. In Phascolomys the inner head 
arises from the back of the internal condyle, and from a small 
piece of the popliteal surface of the femur on both mesial and 
lateral aspects. The outer head arises from the sesamoid bone 
at the back of the external condyle. Both remain separate for a 
considerable distance, the mesial part overlapping the lateral one, 


884 DR. C. F. SONNTAG ON THE MYOLOGY 


But they are inserted into a tendo Achillis. It is therefore 
evident that the degree of separation of the internus and externus 
differs in these Marsupialia. 

Soleus :—In Phascolarctos and Phalanger there is no separate 
soleus, and the observations of Cunningham (2), Macalister (5), 
and Young (14) have shown that it is really contained within the 
gastrocnemius externus, as the latter has a fibular origin. More- 
over, the plantaris lies under cover of the inner part of 
gastrocnemius externus, and not under the part which corresponds 
to soleus. In Phascolomys the soleus has a small fibular head, 
but no tibial origin. 

Plantaris :—In Phalanger and Phascolarctos it is well developed, 
and it is inserted into the plantar fascia, but Young (14) observed 
it inserting into the tendo Achillis in one Koala. In Phascolomys 
it is absent. 

Tibialis Anticus :—In all forms it arises from the outer surface 
of the shaft of the tibia, and the interosseous membrane; but its 
insertion varies. In Phascolomys, according to Macalister (18), it 
is inserted into the entocuneiform. In myspecimen two tendons 
are given off. One goes to-the entocuneiform, and the other to 
the hallux. The latter takes the place of the extensor longus 
hallucis, which is absent as a separate muscle in my specimen. In 
Phalanger and Phascolarctos the insertion is into the entocuneiform 
alone. And in these genera a separate extensor hallucis arises 
from the fibula. 

Extensor Digitorwn Longus :—Macealister (18) describes it in 
Phascolomys as arising from the fibula and front of the tibia, and 
running to the four toes. In my specimen (text-fig. 41) it arises 
from the tibia, fibula, and interosseous membrane. It soon divides 
into slender inner and stout outer portions. The former gives 
three delicate tendons to the first, second, and third toes, but the 
three stout tendons of the lateral portion go to the inner, dorsal, 
and lateral parts of the inner toe. In Phascolarctos (text-fig. 41) 
the muscle likewise consists of two parts, whereas Young (14) 
states that in being separable into two portions the Koala differs 
from such marsupials as have been examined. Three tendons 
emerge. The first is slender and divides into slips for index and 
medius. The second is of medium size and runs to annularis. 
The third is powerful and divides into tendons to annularis and 
minimus. In Phalanger three tendons are disposed as in 
Phascolarctos, but the muscle itself is not divisible into two. 
There is, therefore, a greater resemblance between these arboreal 
forms than there is between Phascolarctos and Phascolomys. 

Peronei :—The muscles composing the groups differ considerably. 
In Phascolarctos, as pointed out by Young (14), there are four 
components—longus, brevis, quarti metatarsi, and quinti meta- 
tarsi,—and all are fibular in origin. In Phalanger, as in Phalan- 
gista maculata, there are three components—longus, brevis, and 
some muscular slips representing portions of the extensor brevis 
digitorum. In Phascolomys Macalister (18) described long and 


OF THE WOMBAT, KOALA, AND PHALANGERS. 885 


short peronei and an accessory quinti tendon detached from the 
last for the first phalanx of the outer toe. In my specimen this 
last slip is absent, but I believe it is represented by one of the 
slips of the extensor longus digitorum. 

Ruge pointed out that the ewtensor brevis digitorum was 
originally derived from the peroneal group of muscles. It is 
present in all Marsupialia, but varies in the degree of development. 
In Phalanger it runs to the four toes, In Phascolarctos it is 


Text-figure 41. 


Extensors of foot in Phascolomys mitchelli (above) and Phascolarctos cinereus 
(below). 
E.L.D: extensor longus digitorum ; E.L.P: extensor longus hallucis; P: peronei; 
T.A: tibialis anticus. 


inserted into the two inner toes. And in both forms it arises 
from the fibula. In Phascolomys it arises fron the outer part of 
the tarsus, and is inserted into the inner pair of toes. 

Tibialis Posticus:—This muscle has already been fully described 
by Cunningham (2), Young (14), and Macalister (5, 18). In 
Phascolomys it is a single muscle arising from the back of the tibia 
and running to the inner part of the scaphoid. In Phascolarctos, 
Phalanger, and Pseudochirus it is double. In the Koala two 
fibular slips run to the scaphoid and entocuneiform ; but in the 
Phalangers both tibial and fibular bellies run to the scaphoid. 


Proc. Zoou. Soc.—1922, No. LX. 60 


886 DR. C. F. SONNTAG ON THE CLASSIFICATION 


The flexor longus digitorum includes flexor longus hallucis, and 
arises from tibia and fibula. Superficial and deep tendons are 
present, and the latter pierce the former, as in the manus. 

The poplitews occupies practically the whole interosseous space. 

The intrinsic muscles of the pes have already been fully 
described (2), and the mid line in Phascolarctos has been moved, 
as In the case of the manus, with resulting modifications in 
structure. 


Summary of Myological Features. 


1. In all animals described in this paper there are traces of 
adherence to the primitive lamination which characterises the 
development of mammalian muscles in general. In Phalanger, 
Pseudochirus, and Phascolomys we observe :—1. Inseparability of 
the prevertebral muscles ; 2. Undivided state of the rhomboideus ; 
3. Fusion of the trapezius with the pectoral fascia and clavicular 
deltoid; 4. Biceps flexor cruris has only one head of origin; 5. 
Separate character of the hamstrings; 6. Degrees of separation 
of levator scapule and serratus magnus. In Phascolarctos the 
following additional ones are present :—7. Platysma continuous 
with muscles of orbit, auricle, and mouth; 8. Sterno-hyoid pro- 
longed to mandible; 9. Styloid muscles a single sheet; 10. 
Digastric and mylo-hyoid fused ; 11. Hyoglossus transverse and 
not attached to hyoid; 12. Omo-hyoid passes into tongue; 13. 
Internal oblique has tendinous intersections; 14. Quadratus 
lumborum absent; 15. Supinator longus very long; 16. Pronator 
quadratus small; 17. Coraco-brachialis double; 18. No extensor 
secundi internodii pollicis; 19. Origin of gluteus maximus; 
20. No soleus; 21. Tibialis posticus double: ; 22. Characters of 
peronei; 23. Intrinsic muscles of manus anal pes. 

2. Many of the myological features are similar in all these 
animals, in many other Marsupials, and in animals belonging to 
the other Mammalian orders. 

3. Most of the characters not included in the above groups are 
adaptations to suit modes of life. 

4. In the small group of characters—omo-trachelian and 
deltoid—not included in the three preceding groups, the Koala 
and Wombat differ from the Phalangers. 


CLASSIFICATION. 

All zoologists have separated Phascolomys from the Phalan- 
geride, so the work of the systematist is limited to fixing the 
proper position for Phascolarctos. To do so it is necessary to 
examine all characters, both external and internal. The large 
assemblage of data so obtained must then be reduced by excluding 
characters of little or no value for purposes of classification. Tn 
the first place, one must remove those which are varying degrees 
of persistence of primitive conditions which were once present in 
their common ancestor. Secondly, structures which are similar as 


OF THE WOMBAT, KOALA, AND PHALANGERS. 887 


the result of convergence must be excluded as they are not 
evidence of affinity. ‘Thirdly, those features which are common 
to all the animals under consideration, to many other Marsupials, 
and to other Mammals cannot be employed in making a classifi- 
cation of the Wombat, Koala, and Phalangers. We are then left 
with a series of useful characters, and those which are sheltered 
deep down in the body should be of value, for they are less liable 
to be influenced by climate and habits than are the more super- 
ficial characters. 

The anatomical characters must be arranged in four groups :— 

A. Those in which Phascolarctos and Phascolomys differ from 
the Phalangeride. They support the systems of Weber (12) and 
Winge (13), and Weber's family of Phascolarctidz is employed 
here for the Koala and Wombat. 

B. Those in which Phascolarctos and the Phalangeride differ 
from Phascolomys. Special attention must also be paid to the 
comparison between Phascolarctos and Pseudochirus. ‘These 
support the systems of Thomas (11), Bensley (1), and Gregory (4). 

C. Characters in which Phascolarctos differs from Phascolomys 
and the Phalangeride. 

D. Characters differing in all forms. 

Groups C and D are evidence in favour of Pocock’s system. 

In estimating the value of the anatomical data one must always 
bear in mind the habits and the character of the diet of these 
animals. Phascolarctos and the Phalangeride are arboreal 
animals, but lead different kinds of life. The latter are active, 
but the former is a clumsy, sluggish creature, clinging to branches 
by its modified feet. Phascolonys, on the other hand, is an active 
animal. Its hind-limbs are fossorial and its fore-limbs are for 
active progression. A study of the myology shows that these 
modes of life are accompanied by numerous differences in the 
muscles. As regards the diet, that of Phascolarctos and Phasco- 
lomys is bulky, whereas that of the Phalangeride is not; and the 
Phascolarctidee have specialisations in the structure of their 
alimentary canal to meet the demands imposed by it. Finally, 
although the life habits differ, we find the arboreal sluggish 
Koala and the active fossorial Wombat possess numerous 
characters of great importance in their skeletons and soft parts, in 
which they differ from the active arboreal Phalangeride. We 
must always aim at discovering the part played by each structure 
in the animal economy before we can assess its true systematic 
value. Some characters are well understood, but there are others 
of whose general adaptive purpose we are totally ignorant. 


A. Characters in which the Phascolarctide differ from the 
Phalangeride. 


Yaxternal Characters :—Pocock (9) showed that the supratragus 
in the Phalangers and many other Mammals is a prominent ridge 
with a well-developed lobe; but it is low, inconspicuous, and 


60* 


888 DR, C. F. SONNTAG ON THE CLASSIFICATION 


devoid of lobate thickening in the Phascolarctide. This, in my 
opinion, is a character of importance, for it does not appear to 
depend on modes of life. Pocock has also shown how the various 
genera can be separated by the auricular characters. As regards 
the mamme the Phascolarctide have only two, but the Phalan- 
geride have four. Pocock found two only in Trichosurus. The 
tail is a well-developed organ in the Phalangeride, and rudi- 
mentary but muscular in the Phascolarctide ; probably the 
reduction is the result of life habits: ‘The Koala clings only by 
its hands and feet to the branches, so the tail would be useless ; 
and a tail would be in reality an impediment to the Wombat. 
The Phalangeride have vibrisse on the wrist and hind foot, 
but no traces of these tactile hairs are present in the Phasco- 
larctidee. Moreover, the integuments of the hind feet differ. 
In the Phalangeride the skin is striated over the well- 
marked pads, but it is granular on the reduced pads in the 
Phascolarctide. 

Myology :—In the Phascolarctide there 1s no cleido-occipital, 
the omo-trachelian is single, and the unbroken deltoid gives a slip 
to the forearm. In the Phalangeride, on the other hand, the 
cleido-occipital is present, the omo-trachelian is double, and the 
deltoid, which is in two parts, does not give a slip to the forearm. 
Although these are prominent differences, they are of very minor 
importance in classification. 

Circulatory System :-—In the Phascolarctidee the pericardium 
adheres to the diaphragm, the post-caval vein does not conceal the 
abdominal aorta, and anterior jugular veins are absent. In the 
Phalangeride the pericardium is connected to the diaphragm by 
two membranous sheets, and the post-caval vein conceals the 
posterior part of the abdominal aorta. I found anterior jugular 
veins in Z'richosurus and Phalanger, but not in Pseudochirus ; 
perhaps they exist in the latter, for my specimens had been 
preserved so long in spirit that the veins, if present, had shrunk 
to minute proportions. 

Alimentary Canal :—Cheek-pouches are absent in the Phalan- 
geride, but present in the Phascolarctide ; they are well-marked 
in Phascolarctos, but rudimentary in Phascolomys. I believe, 
however, that these diverticula are, in reality, adaptive in function. 
The stomach is simple in both families, but possesses the so-called 
gastric gland in the Phascolarctide. That structure has been 
regarded by many zoologists as of special value for showing the 
close relationship between Phascolarctos and Phascolomys. Indeed, 
Forbes (3) regarded it as the main test. But Johnstone (16) 
showed that it is no special gland ; it is an evagination and folding 
of the mucous membrane plus an extension of the normal gastric 
glands. It is in reality a means of increasing the secreting area 
vf the stomach to fulfil the demands imposed by a bulky diet. 
The Phalangers do not consume a bulky diet, so cheek-pouches 
are absent, and there is no trace of the gland patch. Unless 
there were other characters showing a close relationship between 


OF THE WOMBAT, KOALA, AND PHALANGERS. 889 


Phascolarctos and Phascolomys the gland patch alone could not 
be employed as a test of affinity. 

The liver also possesses distinguishing characters. In the 
Phalangeride the caudate lobe is present, but the main lobes are 
not complicated by secondary sulci. In the Phascolarctide, on 
the other hand, there is no caudate lobe, and many secondary 
sulci subdivide the main lobes. 

Phascolarctos predominates over Phascolomys in the size of 
the cheek-pouches, the complexity of the gland patch, the sub- 
division of the main hepatic lobes, and the length of the gall- 
bladder. 

As regards the dentition, Waterhouse (17) pointed out long ago 


Text-figure 42. 


The hard palate in Phascolomys mitchelli.* 


that “the approximation to the Rodent-like dentition which is 
exhibited by the Wombat is perceptible in the Koala, in the 
smaller development of the posterior incisors and canines of 
the upper jaw, and the total absence of those premolars which, 
in the typical Phalangers, intervene between the canine and 
_the five molars of the upper jaw, and the incisor and corre- 
sponding teeth in the lower jaw.’ Bensley (1) has shown, 
however, that the dental characters which would point to a 
resemblance between Phascolarctos and Phascolomys are probably 
due to convergent evolution, and Forbes (3) rightly considered 
that the dental characters in Phascolomys are adaptive in 


* Compare with Plate V in my paper on the Koala (10). 


890 DR. CG. F. SONNTAG ON THE CLASSIFICATION 


character. Tomes (19) showed that the dental enamel in 
Phascolomys possesses histological characters which separate it 
from most, if not all Marsupialia, but he did not specifically 
mention Phascolarctos. It appears, therefore, that in the dentition 
we have an assemblage of characters brought about by adaptation 
and convergence, so they cannot be used to show a close affinity 
between the Koala and Wombat. 

Generative Organs and Development :-—In the Phascolaretidee 
the vaginal culs-de-sac are separate, but they coalesce in some of 
the Phalangeride, at least in T'richosurus (Horbes). As regards 
the foetal membranes, Professor J. P. Hill informs me that they 
are fundamentally the same, but differing in details, in the Koala 
and Wombat. Moreover, the conditions in the Phascolarctidee 
differ entirely from those in the Phalangeride. 

Osteology :—I have compared the conditions in the material at 
my disposal with Owen’s account (8), and the essential points are 
shown in the subjoined table :— 


Phascolarctide. Phalangeride. 
1. Pterygoid processes large. 1. Processes small. 
2. Intermaxillary bones increased trans- | 2. Bones not increased transversely. 
versely. 
3. Palatal process of maxilla forms a| 38. Maxille form a small part of 
considerable part of boundary of boundary. 


incisive foramina. 
4, Palatal foramina entirely on palate | 4, Palatal foramina extend into maxill, 
bones. 
. Groove obsolete. . Groove runs from foramen rotundum 
to Gasserian fissure. 
. Margin of tentorium ossified. 
. Convolutions of mid spongy bone 
numerous and delicate. 


5 
. Not ossified. 6 
7 

. Halves of mandible united. 8. Halves separate. 
0 


5 
6 
7. Mid spongy bone simple * 
8 
9 
0 


9. Seventh cervical vertebra perforated. . Not perforated. 
. Process beneath the sixth cervical | 10. Process greatly expanded. 
process short. 
11. Anterior arch of the atlas partly | 11. Arch entirely osseous. 
cartilaginous. 


Jn many of the osteological characters not included in the 
above list there are differences in degree only in the Phalangers, 
Phascolarctos, and Phascolomys. Doran has shown that the ear 
ossicles of Phascolarctos differ entirely from those in the Phalan- 
geridse, but they have some points in common with those of 
Phascolomys. 


B. Characters in which Phascolarctos and the Phalangeride 
differ from Phascolomys. 


External Characters :—In Phascolomys flaps of skin from the 
lower lip protrude into the diastemata, but this is a new develop- 
ment, associated, as in many Rodentia, with the type of dentition. 
There is, however, no trace of these in the other animals. As 


* Owen’s account is relied on here as the skulls in my possession are not perfect as 
regards the turbinate bones, 


ee 


OF THE WOMBAT, KOALA, AND PHALANGERS, aol 


regards the manus and pes, it is necessary to consider both the 
external appearances and the internal structure. Pocock (9) and 
others have already described the external appearances, and the 
following conclusions can be drawn from their accounts :— 

1. The fore foot possesses no special modifications in Vricho- 
sUurUs. 

2. The conditions in Phascolomys, which are adapted for a 
fossorial and ambulant life, have some resemblance to those of 
Trichosurus, but differ in many ways from those in Phascolarctos 
and the Phalangeride. Among these are the length of the 
digits, the shortness and bluntness of the claws; and there is no 
division of the digits into two opposing groups. The pads are 
reduced. 

3. In Phascolarctos and the Phalangeride the pollex and index 
can be opposed to the long axis of the manus, and Young regarded 
these two digits as constituting one group, the remaining three 
digits forming a second. These conditions are slightest in 
Phalanger, intermediate in Pseudochirus, and most highly 
developed in Phascolarctos. The latter differs from all Phalangers, 
except 7’richosurus, in the appearance of the pads. In Phalanger 
and Pseudochirus the pads are striated, but they are granular in 
Phascolarctos and Phascolomys. Moreover, myological literature 
shows that Phascolarctos is peculiar in many ways. I agree with 
Pocock’s scheme, which shows that the characters of the manus in 
Phascolomys differ from those in the other animals. And although 
the others have features in common, there are sufficient characters 
to distinguish Phascolarctos from the Phalangeride. 

Bensley (1) described and figured the pes of the Diprotodonts, 
and drew important conclusions as regards the affinities of the 
animals. He pointed out that the pes in Dromicia is prototypal 
for the Phalangerid, and those of Phalanger and Pseudochirus 
conform more or less closely to it. But the conditions in 
Phascolarctos show no close relation to those in Pseudochirus. 
The general conformation of the digits is the same, but the pads 
are all reduced, the sole being covered with soft, granular skin as 
in Phascolomys. The hallux is much farther back than in 
Pseudochirus, so it is more perfectly opposable. A. slightly 
greater displacement would put its axis in line with that of the 
fourth digit, which it opposes. The foot differs in Phascolarctos 
and Phascolomys both in external appearance and internal 
structure. 

Although the pes of Phascolarctos differs trom that in Pseudo- 
chirus the dental characters are similar, but reach a higher degree 
of development in the former. Bensley (1) showed that these 
genera have quadrituberculate molars with selenoid cusps. The 
upper teeth have reduced external styles, with or without 
intermediate conules. In Phalanger and Trichosurus the quadri- 
tuberculate molars have bunoid cusps. And the upper teeth have 
neither external styles nor intermediate cusps. Moreover, as 


892 DR. C. F. SONNTAG ON THH CLASSIFICATION 


shown on page 889, there are some resemblances between the teeth 
of Phascolarctos and Phascolomys which have resulted from 
convergence. These, however, must be excluded for purpose of 
classification. 

Osteology :—The contrast in the skeletal characters are tabulated 
as follows :— 


Phascolarctos and the Phalangeridx. Phascolomys. 


po 


. Zygoma extended vertically and has | 1. Zygoma has a marked twist. 
no twist. 

. Elements of occipital bone confluent. | 2. Elements not all confluent in many 

skulls, but not in all. 

. Characters of supra-occipital. 

Occipital processes arise from ex- 

occipital. 

Bulla from alisphenoid. 

. Petrous has a large cerebellar pit. 

. Six pieces in sternum. 

. Characters of humerus. 

. No third trochanter on femur. 


Processes from petro-mastoid. 


Bulla from temporal bone. 
. Pit obsolete. 
. Four pieces in sternum. 


COTA Bo vo 


Oo NOON 


. Third trochanter present. . 


Myology :—Phascolomys difters from the Phalangeride and 
Phascolarctos in the absence of the quadratus femoris, obturator 
internus and popliteus, and in having only one head to the rectus 
femoris. : 

Circulatory System :—Phascolomys has no trace of a vena 
transversa (text-fig. 32). 

Tt is necessary to remark on the comparison between Phasco- 
laretos and Pseudochirus, because these have been placed close 
together in systems of classification. In a detailed examination 
of their anatomy one finds they differ from other Phalangers and 
Phascolomys in the character ef the teeth and straightening out 
of the inflected angles of the mandible. As regards the visceral 
anatomy and myology, there is no way in which they can be marked 
out from other forms. I can see no reason other than the 
condition of the teeth and mandible for separating Pseudochirus 
from the other Phalangers, but I consider these alone as sufficiently 
important to do so. 


When the characters mentioned in groups A and B are reduced 
by the elimination of the unimportant ones the following lists are 
obtained :-— 


A. Characters separating the Phascolarctide from the Phalan- 
geridee :-— 


1. Supratragus, tarsal vibrisse, skin of pads and tail. 
2. Gastric gland patch and liver. 

3. Foetal membranes. 

A, Position of palatal foramina. 

5. Characters of symphysis menti. 

6. Characters of the mid-turbinal bone (Owen). 

7. Characters of the ear ossicles, 


OF THE WOMBAT, KOALA, AND PHALANGERS. 893 


B. Characters separating Phascolomys from Phascolarctos and 
the Phalangeridze :-— 


. Formation of the alisphenoid bulla. 

Site of occipital processes. 

Pieces in sternum. 

Third trochanter of femur. 

Dentition. 

. Some of the characters of manus and pes. 


C. Characters in which Phascolarctos differs from the Phalangers 
and Phascolomys. 


External Characters :—Pocock (9) showed that there is a 
rhinarium in Phascolomys and the Phalangeride, but no true 
rhinarium is present in Phascolarctos; and the nostrils of the 
latter are also peculiar. In the same paper he deals with the 
characters of the marsupium, and points out that Phascolomys is 
intermediate between the Phalangeridee and Phascolarctos. More- 
over, the pouch of the latter is not closely related to that of any 
genus of the Phalangeride. 

Myology :—In the summary of myological features at the end 
of the first part of this paper I enumerated many conditions 
representing adherence to the primitive lamination of the muscles. 
But these cannot be used for purposes of classification. 


The other points are given in the following table :— 


Phascolarctos. Phascolomys and Phalangeride. 
1. Occipital crest curves backwards. 1. Not so. 
2. Alisphenoid bulle as long as ex-| 2. Not so in  Phalangers. Bulla 
occipital processes. temporal in Phascolomys. 
3. Characters of nasal bones. 
4. Anterior margin of septum nasi con- | 4. Convex. 
cave. 
5. First dorsal spine scarcely exceeds 
seventh cervical. 
6. No patella. 6. Patella thin and cartilaginous. 
7, Vagina musculina present. 7. Absent. 
8. No: azygos lobe. 8. Right lung has azygos lobe. 
9. Epiglottis entire and far from tongu ue. | 9. Epiglottisnotched and close to tongue, 
10. No faucial tonsil. 10. Faucial tonsils present. 
11, Thick ureteric venous plexus. 11. Absent, 


D. Characters differing in all forms. 


In the first part of this paper I described no fewer than thirty- 
two myological features differing in all forms. But these fall 
into primitive and adaptive groups, and are useless for purposes 
of classification, As regards visceral anatomy, there are many 
points differing in all forms. Some of these are undoubtedly 


894 DR. C. F. SONNTAG ON THE CLASSIFICATION 


adaptive, but others cannot be included in that category. The 
visceral and osteological characters are tabulated as follows : — 


1. Oral vestibule, palatal formula, tongue. 

2. Spleen, thymus, cervical lymph-glands. 

3. Division of the right auricular appendix. 

Branches of the aortic arch. 

Number of dorsal vertebre, ribs, intercostal soft parts. 
. Relative length of facial part of skull. 

. Characters of glenoid cavity. 

Size and perforations of lachrymal bone. 

. Characters of scapula. 


b 


Text-figure 43. 


The scapula in Phascolomys mitchelli (A.): Phascolarctos cinereus (B.) ; 
and Phalanger orientalis (C.). 


Conclusions. 


From an examination of these four groups of characters I have 
come to the following conclusion :— 


1. Phascolarctos cannot be included in the Phalangeride. 
Although dentition and foot structure have led some observers to 
put it in that family in close relation to Pseudochirus, there are 
many very important points in which it difters from them. If 
they are united it will be by superficial characters, and one will 


OF THE WOMBAT, KOALA, AND PHALANGERS. 895 


break up the series of fundamental characters uniting the Koala 
and Wombat. 

2. Phuscolarctos is united to Phascolomys by characters whose 
importance far outweighs the resemblances between the former 
and the Phalangeride. The major ones enumerated on page 892 
are regarded as of value by anatomists and evolutionists. 

3. The characters peculiar to Phascolarctos are not sufficiently 
important to place it ina separate family. But they are useful 
for distinguishing it from Phascolomys in the family Phasco- 
larctidee. 

4, Tagree with Weber and Winge that there should be only 
two families, one for the Phalangers, and one for the Koala and 
Wombat; and there are characters of sufficient importance to 
place each of the animals in the latter in a separate subfamily. 
My views are tabulated thus :— 


Family PHASCOLARCTIDA, 
Subfamily 1. PHASCOLARCTINA. 
2. PHASCOLOMYIN®. 


Family PHALANGERIDA. 


Subfamily 1. PHALANGERINAE, 
Phalanger, Trichosurus. 

Subfamily 2, PsEuDOCHIRINE. 
Pseudochirus. 


5. If one simply enumerates characters and strikes a balance 
he will obtain one form of classification, but if he considers each 
feature and tries to detect the part it plays in the animal 
ceconomy his results will be quite different. ‘They will lead him 
to place the Koala and Wombat in the same family. 


BIBLIOGRAPHY. 


1. Brensuey, B. A.—‘‘ On the Evolution of the Australian Marsu- 
pialia.” Trans. Linn. Soc. London (2), ix. pp. 83-217. 

2. Cunnineuam, D. J.—-Voy. H.M.S. ‘Challenger,’ Zool. v. pt. xvi. 
pp. 1-92 (1882). 

3. Forses, W. A.—‘‘On some Points in the Anatomy of the 
Koala (Phascolarctos cinereus).” P.Z.S. 1881, pp. 180-195. 

. Grecory, W. K.— The Orders of Mammals.” Bull, Amer. 

Mus. Nat. Hist. xxvii. pp. 1-524 (1910). 

. Macauister, A.—“ On the Muscular Anatomy of the Koala.” 

Ann. Mag. Nat. Hist. vol. 11. p. 329 (1872). 

Mecxet, J. F.—Anat. Comp. vol. vi. pp. 166, 167. 

. Osgooo, W.—Field Mus. Nat. Hist. Chicago, Publication 207, 

Zool. Series, vol. xiv. no. 1 (1921). 

. Owen, R.—‘*On the Osteology of the Marsupialia.” Trans. 
Zool. Soc. vol. 11. pp. 379-408 (1841). 


896° 


a: 


. Youne, A. H. 


MYOLOGY AND CLASSIFICATION OF THE WOMBAT, ETC. 


Pocock, R. I.—‘*The External Characters of the Koala 
(Phascolarctos) and some related Marsupials.” P. ZS. 
1921, pp. 591-607. 


. Sonnrac, C. F.—“ The Comparative Anatomy of the Koala 


and Vulpine Phalanger.” P. Z. 8. 1921, pp. 547-577. 


. Tomas, O.—Cat. Marsupialia and Monotremata in Brit. 


Mus. 1882. 


. Wepser, M.—‘ Die Siiugethiere. 1904. 
. Winer, H.—E Museo Lundii, viii. pt. 1 (1893). 
. Youne, A. H.—‘*The Muscular Anatomy of the Koala.” 


Journ. of Anat. & Phys. vol. xvi. pp. 217-242 (1882). 
“The Intrinsic Muscles of the Marsupial 
Hand.” Journ. of Anat. & Phys. vol. xiv. p. 158 (1880). 


. JOHNSTONE, J.—‘‘ On the Gastric Glands of the Marsupialia.” 


Journ. Linn. Soc. Lond., Zool. xxvii. pp. 1-14 (1899). 


. WATERHOUSE, G. R.—Natural History of the Mammalia, 


vol. i.: Marsupiata (1846). 


. Macauister, A.—‘‘On the Myology of the Wombat and 


Tasmanian Devil.” Ann. & Mag. Nat. Hist. (4) v. pp. 153— 
173 (1850). 


. Tomes, C.—Phil. Trans. 1849. 


COMMENSALISM BENEFICIAL TO CRUSTACEA. 897 


39. Experimental Hvidence that Commensalism may be 
beneficial to Crustacea. By Hpwarp B. Poutton, 
D.Se., F.R.S., F.Z.S., Hope Professor of Zoology and 


Fellow of Jesus College in the University of Oxford. 
[Received November 7, 1922: Read November 7, 1922. | 


The special associations of Crustacea with organisms believed to 
be disliked by their enemies—-with Sponges, Ascidians, and Sea- 
anemones—has probably been derived froma more generalized 
use of animal and vegetable growths for the purpose of 
concealment. The following experiments proved that two con- 
spicuously coloured forms associated with Hermit-crabs were 
intensely distasteful to fishes, and that one of the crabs, when 
deprived of its associate, was greedily devoured. The experi- 
ments were carried out, with the help and advice of Prof. W. 
Garstang, in the Marine Biological Laboratory at Plymouth in 
the summer of 1890. The results have often been described, but 
never published. 

(1) Pagurus bernhardus and its Sea-anemone, Sagartia para- 
sitico.—These Crustacea were commonly dredged up at Ply- 
mouth, and large specimens bore on the shells—generally whelks 
—from two as to as many as six of these large brightly coloured 
actinians. Prof. Garstang had already obtained much indirect 
evidence of the value of the association, for he had often found 
the young Hermit-crabs, too small to carry a Sea-anemone, 
in the stomach of gurnards and other fishes. They had been 
swallowed entire, their borrowed shells and all. He had, however, 
never found in the fishes any of the larger crabs living in shells 
suited for carrying Sagartia™. 

I first tested the nematocysts of the Sagartia by touching it 
with the tip of my tongue, and at once experienced a sharp smart 
which endured for many hours. Pieces of about the size and 
shape of the bits of meat on which the fish in one of the tanks 
were accustomed to be fed were then cut from a Sea-anemone 
and thrown into the tank. Misled by this, a few fishes seized 
pieces of the Sagartia; but no sconer had one been received into 
the mouth, than it was shot out again with much force, and the 
fish shook its head violently from side to side, apparently feeling 
the same smart which I had experienced myself. After these 
first trials not one of the fish would touch the pieces, and it was 
obvious that the great majority saved themselves by yielding 
to the stimulus provided by the behaviour of the others. 

2) Pagurus cuanensis and the Sponge, Suberites domuncula,.— 
This small Hermit-crab, also common at Plymouth, inhabits a 
cavity in the sponge, the tip of its tail being fixed in a small 


* These facts are recorded in “Colours of Animals,” Internat. Sci, Ser., Lond. 
1890, p. 203. 


898 COMMENALISM BENEFICIAL TO CRUSTACEA. 


gastropod shell, while the rest of its body lies in a spiral canal 
continuous with that of the shell and gradually prolonged from 
it with the growth of the two associated organisms. The small 
shell was evidently inhabited by the young crab when the asso- 
ciation began, but had become deeply buried in the sponge at the 
time when the experiments were made. The sponge was of the 
usual orange-red colour, although they are sometimes dark and 
comparatively inconspicuous, In size and shape it resembled a 
smallish potato. 

When disturbed or frightened in any way, the crab darted 
back into its sponge exactly as other Hermits dash into their 
shells. 

In order to obtain pictures of this species and P. bernhardus 
under as natural conditions as possible, the objects dredged up 
with each species were arranged in a dish, which was lowered into 
an aquarium so that its edge was in contact with the glass front 
and its contents well lhghted from above. The crabs were 
secured in the most convenient position by tying to a stone in 
the dish. Although the string was wound so as not to touch 
P. cwanensis, and was, in fact, separated from its body by a con- 
siderable thickness of sponge, the crab quite understood the 
situation, and spent the whole time the artist was at work in 
trying to wear through the string with its two large claws. 
Thus occupied and stretching out of its canal, it was an 
admirable sitter. 

Prof. Garstang had already recorded that sponges as a group 
are intensely disliked by fishes*, but I was anxious to test this 
special example and compare its palatability with that of the crab. 
Pieces like those of the Sagartia were therefore thrown into the 
aquarium, but were recognized as unpalatable even more quickly 
than the others, and seized by fewer fishes. When taken they 
were instantly rejected. ‘The Pagurws was then offered deprived 
of his sponge; there was a wild struggle, and the fortunate 
captor swallowed him in a moment. 


It was evident that both these Hermit-crabs are associated with 
organisms possessing qualities rendering them unpleasant to fishes 
—qualities advertised by conspicuous warning colours. One of 
the crabs was shown on this occasion to be very palatable to fishes, 
and the other previously by Prof. Garstang, to be much sought 
after by them. Although the experiments described were very 
few, the results were so definite and clear that there can be little 
doubt about the meaning of the association in these and other 
similar examples of commensalism in Crustacea. 

The beautiful illustrations, projected upon the screen, were 
painted in 1890 by my friend the late Mr. H. M. J. Underhill, of 
Oxford, who has reproduced the appearance of the living animals 
in their natural surroundings with wonderful skill. 


* “ Colours of Animals,” 7. ¢. 


NEW LIZARD OF THE GENUS CHALCIDES. 899 


40. Description of a New Lizard of the genus Chalcides, 
from the Gambia, living in the Society’s Gardens. 
By KE. G. Boutencer, F.Z.8. (Curator of Reptiles). 
[Received October 5, 1922: Read November 7, 1922. ] 


The Society recently received from H.H. Capt. C. H. Armitage, 
C.M.G., a generous donor to its collection, a number of reptiles 
from the Gambia. Among these was a Lizard of the genus 
Chalcides which is evidently undescribed, and for which IL 
propose the name of Chalcides armitaget, after its discoverer. 


CHALCIDES ARMITAGEI, sp. n. 


Snout more conical than wedge-shaped, but with distinctly 
projecting labial edge. Eye small. Har-openings not larger 
than the nostril, on a line with the mouth. Nostril pierced 
mostly in advance of the suture between the rostral and the 
first labial. Supranasals united. Frontal longer than broad. 
Fourth labial entering orbit. Body very elongate. Sides of 
body not distinctly angular. Scales perfectly smooth, in 24 rows 
round the middle of the body. Limbs short tridactyle. Length 
of hind-limb not quite equal to distance between eye and fore- 
limb. Fore-limb equals in length distance between posterior 
border of orbit and snout. ‘Tail shorter than head and body. 

From snout to vent 119 millim.; tail 82 mm. 

Great variation is to be found in the structure of the Lizards 
of the genus Chalcides, especially in the proportions of the limbs 
and body, and in the number of digits, where we find all stages 
of degeneracy, there being species with five, four, three, or even 
two fingers or toes. A linear arrangement based solely on the 
number of the digits would be unnatural, as I have attempted to 
show in a recent paper*. The shape of the snout, whether 
conical or wedge-shaped, is of importance in tracing lines of 
descent ; this character being also taken into consideration, the 
species here deseribed as new cannot be placed in the group 
Ch. sphenopsiformis, delislii, and sepoides, its position being 
intermediate between Ch. mionecton (with four fingers and toes, 
and 24 or 26 scales round the body), and Ch. mauritanicus (with 
two fingers and three toes, and 10 scales round the body). 


* P.Z.S. 1920, vol. i. p. 77. 


NEMATODA FROM MAMMALS WHICH DIED IN THE GARDENS. 901 


41. On the Parasitic Nematoda collected from Mammalian 
Hosts which died in the Gardens of the Zoological 
Society of London during the years 1919-1921 ; with 
a description of three new Genera and three new 
Species. By G. M. Vuvers, MOW.C'S., L.E.CE., 
F.Z.S., Beit Memorial Research Fellow, and Assistant 
in the Department of Helminthology, London School 
of Tropical Medicine. 

[Received October 24, 1922: Read November 7, 1922. | 
(Text-figures 1-10.) 


The parasites referred to in this paper were collected from 
mammals which died in the Gardens from October 1919 to 
June 1921, during which period the writer held the position of 
Honorary Parasitologist to the Society. 

In a previous Report, published in 1920 (1), I gave a list of 
the parasites which were collected during the first eight months 
of this period. Certain of these are again mentioned in this 
paper, and to avoid confusion are marked in the Systematic List 
with an asterisk (*). 

Leiperenia galebi, from the Indian Hlephant (Hlephas indicus), 
and Galoncus tridentatus, from the Clouded Leopard (felis 
nebulosa), have both been described as new species by Dr. M. 
Khalil in previous papers in 1922 (2). The methods employed 
in collection have been referred to in my Report of June 1920. 

The whole of the material has been worked out in the 
Helminthological Department of the London School of Tropical 
Medicine under the Directorship of Prof. R. T. Leiper, to whom 
I am indebted for his kind assistance and many valuable 
suggestions. 

In all, 43 species were collected from 36 hosts. Three of these 
are apparently new to science ana are described below. 

It is necessary to create three new Nematode genera :— 


(1) Troglostrongylus, for a new species of Metastrongylid worm 
found in the frontal sinus of a Leopard Cat (felis 
bengalensis). Type 7. troglostrongylus, gen. n., sp. n. 

(2) Cyhcosmrura, for the Spirurid from the stomach of the 
Tiger originally described as Spiroptera subequalis by 
Molin. Type C. subequalis (Molin, 1860), gen. n. 

(3) Papillosetaria, for a new species of Filariid worm from the 
peritoneal cavity of Zragulus stanleyanus. Type P. tra- 
gult, gen. n., sp. n. 


In Tragulus stanleyanus a second new species also occurred ; 
this I have named Setaria javensis, sp. n. 
Proc. Zoou. Soc.—1922, No. LX1I. ol 


902 MR. G. M. VEVERS ON NEMATODA FROM MAMMALS 


Systematic List of Nematoda collected. 


Ryappiasoied R. & H., 1916. 
ATRACTID & Travassos, 1919. 
* Leiperenia galebi Khalil, 1922. 
* Probsitmayria vivipara Ransom, 1907. 
Oxyurowed R. & H., 1916. 
OxyuRtipD # Cobbold, 1364. 
*Oxyuris equi Schrank, 1788. 


ASCAROIDEA R. & H., 1915. 
AscAaRID & Cobbold, 1862. 
Ascarina& Travassos, 1913. 
Ascaris transfuga Rud., 1819. 
Belascaris marginata (Rud., 1802), Gedoelst, 1911. 
Belascaris mystax (Geder, 1800), Leiper, 1907. 
Toxascaris limbata R. & H., 1911. 
Toxascaris leonina (v. Linstow, 1902), R. & H., 1911. 


HEeTEROCHEILIDAR. & H., 1915. 
Gorzin&{ Travassos, 1920. 
* Contracecum osculatum (Rud., 1802), Baylis, 1920. 


HEeTEeRAKIDAR. & H., 1914. 
SuBULURIN& Travassos, 1914. 
Subulura distans (Rud., 1809), Kh. & H., 1912. 


STRONGYLOIDEA Weinland, 1858. 
STRONGYLID2 Baird, 1853. 
SPRONGYLIN# Stossich, 1898. 
Strongylus vulgaris (Looss, 1900), R. & H., 1909. 
Strongylus edentatus (Looss, 1900), R. & H., 1909. 
Cylicostomum nassatum (var. parvum), Yorke & McFie, 
Cylicostomum goldi Boulenger, 1916. [1918. 
Cylicostomum bicoronatwm (Looss, 1900), Gedoelst, 1903. 
Usophagostomum apiostomum (Willach, 1891), R.&H., 
(1905. 
ANCYLOstomINz R. & H., 1909. (1909. 


Ancylostoma malayanwm (Alessandrini, 1905), R. & H., 
Ancylostoma ceylanicum Looss, 1911. [ 1909. 
Ancylostoma pluridentatum( Alessandrini, 1905), R.& H., 
Ancylostoma caninum (Evcolani, 1859), Hall, 1915, 
*Ancylostoma conepati Solanet, 1911. 
Galoncus perniciosus (v. Linstow, 1886), Railliet, 1918. 
Galoncus tridentatus Khalil, 1922. 
*Uncinaria criniformis (Goeze, 1782), Railliet, 1899. 
Characostomum asmilium R., H. & Joyeux, 1913. 


ee 


WHICH DIED IN THE ZOOLOGICAL GARDENS. 903 


TRICHOSTRONGYLID4& Leiper, 1912. 
TRICHOSTRONGYLINEH Leiper, 1908. 
*Hemonchus contortus (Rudolphi, 1805), Cobbold, 1898. 


METASTRONGYLID& Leiper, 1908. 
ME?rAStRONGYLINE Leiper, 1908. 
*Troglostrongylus troglostrongylus, gen. n., Sp. 0. 


Sprruromea R. & H., 1915. 
SPIRURIDZ Oerley, 1885. 
SPIRURIN® Railliet, 1915. 


Habronema chevrenxi Seurat, 1913. 
Spirocerca sanguinolenta (Rud., 1819), R. & H., 1911. 


Streptopharagus armatus Blane, 1912. 
Cylicospirura subequalis (Molin, 1860), gen. n. 


RicTruLARIiIyp # Railliet, 1916. 
Rictularia affinis Jigerskiold, 1910. 
Rictularia cahirensis Jigerskiold, 1910. 
hictularia plagiostoma (Wedl, 1861), Will.-Suhm 1873. 


GNAYTHOSTOMIDS R. Blanchard, 1895. 
Gnathostoma spinigerum Owen, 1836. 


FILAROIDEA Weinland, 1858. 
FIuARiIiID & Claus, 1885. 
Hinartn& Stiles, 1907. 
*Milaria gracilis Rudolphi, 1809. 
Filaria martis Gmelin, 1790. 
Setaria labiato-papillosa (Aless., 1838), R. & H., 1911. 
Seiaria hornbyi Boulenger, 1920. 
Setaria javensis, sp. 0. 
Papillosetaria traguli, gen. n., sp. n. 


TRICHUROIDEA Railliet, 1916. 
TRICHURID2 Railliet, 1915. 
TRICHURIN® Ransom, 1911. 
Trichuris ovis (Abildg., 1795), Smith, 1908. 
Trichuris trichiura (Lanneeus, 1771), Stiles, 1901. 


ANCYLOSTOMA PLURIDENTATUM (Aless., 1905), R. & H., 1909. 
In 1905, Alessandrini described a Hookworm from Felis mitis, 
Brazil, which he named Uncinaria pluridentatum (8). In 1909, 
Railliet and Henry placed the species in the genus Ancylostoma 
(4). 
Looss, in his Monograph on the Hookworms (1911) (5), refers 
Gl 


904 MR. G. M. VEVERS ON NEMATODA FROM MAMMALS 


to, and gives a brief description of the worm, with figures 
copied from the original paper by Alessandrini. The only 
measurements given are the size of the body. In 1916, Clayton 
Lane (6) suggested that A. pluridentatwm might not be a valid 
species; he also suggested that if Alessandrini had not made an 
ervor and the species was valid, it should be made the type of a 
new genus. 


Text-fig. 1. 


Text-fig. 2. 


oe 
rr iF 
200 p1. 20041. 


Ancylostoma pluridentatum (Aless.). 


Mouth capsule. Lateral view. Mouth capsule. Dorsal view. 


My material was collected from the small intestine of /elis 
tigris, Malay States. 

I have compared the general measurements, mouth capsule, 
and bursa with the original description and figures given 
by Alessandrini, and find that they agree in all but three 
points :— 

(1) The length of the body. Measurements given by Ales- 

sandrini: Male 6-8 mm., female 6°5-9 mm. 
Measurements of my material from Felis tigris : 
Male 8:5—9 mm., female $-10°5 mm. 


WHICH DIED IN THE ZOOLOGICAL GARDENS. 905 


(2) The appearance of the mouth capsule when viewed from 
the dorsal respect. In my specimens the mouth capsule 
appears mueh deeper than the mouth capsule in Ales- 
sandrini’s figure. 

(3) Length of the spicules: 

Alessandrini’s measurement = 960 pm. 
Material from /elis tigris = 800 p. 


These differences do not seem to indicate that the worms from 
F. tigris are of a different species from those described by 
Alessandrini from /, mitis. The variations noted might well 
be ascribed to the occurrence in a different host. 

Tt would seem that the figures of the mouth capsule, as given 
by Alessandrini, were made from somewhat flattened material ; 
I have therefore prepared new figures of the head from the dorsal 
and lateral aspects (text-figs. 1 & 2). 


Text-fig. 3. 


Text-fig. 4. 


100. 
Characostomum asmilium R., H. & Joyeux. 
Mouth capsule. Dorsal view. Dorsal ray of bursa. ¢. 


CHARACOSTOMUM AsMILIuM R., H. & Joyeux, 1913. 


This species was first described by Railliet, Henry and Joyeux 
in 1913 from Cercopithecus patas (7). 

These authors describe the worm fully, but do not give figures 
of the bursa of the male or the cephalic extremity from the 
dorsal aspect. 

Two males were found in the small intestine of a Pig-tailed 
Macaque (Macacus nemestrinus) and one female from a Slow 
Loris (Wycticebus tardigradus). 


906 MR. G. M. VEVERS ON NEMA'TODA FROM MAMMALS 


The opening of the mouth is not terminal, but turns slightly 
dorsalwards as in Ancylostoma and Necator, 

The bueeal aperture is surrounded by three concentric chitinous 
rings; these have no sete. The dorsal gutter is very well 
marked, and is rugose on the dorsal aspect. 

A figure is given illustrating these two points, as they are not 
mentioned i in the description of ‘the original material. A diagram 
of the dorsal ray of the bursa in the male is also given. 


Me@rastroNGyLin#& Leiper, 1908. 


TROGLOSTRONGYLUS TROGLOSTRONGYLUS, gen. n., Sp. Nn. 


In a previous paper (1) I made reference to a parasite which 
I found in the frontal sinus of a Leopard Cat (Welis bengalensis). 
At that time [ was uncertain as to its systematic position, and 
provisionally diagnosed it as a “‘species inquirenda” in the 
genus Synthetocutlus, with which it has certain affinities. The 
female, however, is ovoviviparous, and the male has remarkably 
long spicules with palmate expansions covered with minute 
spines, which characters not only establish it as a new species, 
but separate it from Synthetocaulus. 

It also has certain characters in common with the genus 
Hemostrongylus, ». species of which (ZH. suberenatus) was described 
by Railliet and Henry, from the bronchi of a Leopard, in 1913. 
The disposition of the rays of the bursa in this worm is different ; 
in H. subcrenatus the postero-lateral and median-lateral rays are 
fused, whereas in 7. troglostrongylus the antero-lateral and 
median-lateral rays are jomed. HH. subcrenutws, moreover, has 
no accessory piece (8). 

On these grounds, therefore, I have considered it necessary to 
create a new genus—7oglostrongylus—for this worm. 

It would appear that the only other Nematode parasite which 
has the frontal sinus as a habitat is the worm /ilaroides muste- 
larwm, which lives in the frontal sinus of the Weasel (A/ustela 
vulgaris) (9). 

The systematic position of /. mustelarwm is somewhat obscure, 
but it probably falls into the Spiruroidea, which has no affinities 
with the Strongyloidea, into which 7. troglostrongylus belongs. 
A comparison of these two forms shows certain points of simi- 
larity in structure which seem to indicate parallelism in evolution 
due perhaps to similarity in habitat. 


TROGLOSTRONGYLUS, gen. n. 


Generic diagnosis.— Metastrongyline : Head simple, two lips, 
no buccal capsule. Cuticle covered with fine longitudinal stria 
tions. The bursa of the male is small in comparison with the 
length of the body. The ventral rays are short, and each formed 


WHICH DIED IN THE ZOOLOGICAL GARDENS. 907 


of two rays fused. The antero-lateral and median-lateral rays 
are fused together to form one large vay. ‘he postero-lateral 
and externo-dorsal rays are single and approximately the same 
size. The dorsal ray is single and broad, owing to the fusion of 
all its elements. 

The spicules are long and equal, they are tesselated through- 
out and carry pectinate lamelle along their inner edges; each 
ends in a palmate expansion, the fingers of which are webbed 
with a cuticular expansion which bears minute spines. ‘The 
female is ovoviviparous. . 

Type-species, 7'roglostrongylus troglostrongylus, sp. n. 


TROGLOSTRONGYLUS TROGLOSTRONGYLUS, Sp. N. 


Host. Felis bengalensis. 

Locality. India. 

Habitat. Frontal sinus. 

Specific diagnosis.—Troglostrongylus: Thin filiform greyish- 
white worms, tapering rather abruptly at each end. 

The mouth is guarded by two inconspicuous lips, each bearing 
two minute papille. There is a slight cuticular expansion at 
the head and extending about 1 mm. along the body; on this 
cuticular expansion are faint transverse striations. The rest of 
the worm is covered with very fine longitudinal striations. 

The cesophagus in both sexes is of the simple muscular bulb 
type, measuring 500 » in length and 100 » in diameter at the 
bulb. The excretory pore and nerve ring are situated 250 p 
from the anterior end of the body. 

Male-—Measures 12 mm. in length by -4 mm. in breadth. 
The bursa is small and rounded, with a slight median notch in 
its margin at the point where the distal end of the dorsal ray 
meets it. The diameter of the bursa is*} mm. The rays of the 
bursa are somewhat asymmetrical. The ventral rays are small 
and double; the antero-lateral and median-lateral rays are fused 
into one large ray. 

The postero-lateral and externo-dorsal rays are single and 
short, in some cases not reaching the margin of the bursa. The 
elements of the dorsal ray are fused into one large, broad ray, 
which terminates in the median notch of the bursa. 

The spicules are equal and measure 2:25 min. in length; they 
are dark brown in colour and are tesselated. In addition, along 
the inner edge of each there are short transverse comb-like 
projections, which may be single or sometimes divide into two or 
even three branches. In the distal third these interlock and 
produce a union of the two spicules. Hach spicule ends in a 
palmate expansion, the cuticular web of which is covered with 
minute spines. The accessory piece is dagger-shaped and 
measures 270 p in length. 


908 MR. G. M. VEVERS ON NEMATODA FROM MAMMALS 


Female.—Measures 20-24 mm. in length by ‘7 mm. in breadth. 
The tail tapers sharply and ends in a blunt point, near the end 
of which are situated two small laterally placed papille. The 
anus is 300 yw from the tip of the tail. 

The valves is situated just behind the middle of the body. 


Text-figure 5. 


* 200 


TIT? « 


"Si 


Ip URlneeconeus troglostrongylus, gen. N., Sp. Nn. 


a. Caudal extremity of male. Lateral view. 

6. Caudal extremity of male. Ventral view. 

c. Egg. X 200. 

d. Caudal extremity of female. 

e. Head of female. Lateral view, showing excretory pore. 


The eggs contain a living embryo at birth, and hatching takes 
place almost immediately, embryos being found in. the naso- 
pharynx, lungs, cesophagus, stomach, and intestines. The eggs 
measure iia by 7Ow. EKmbryes found in the nasopharynx 
measured 240 yu. 


WHICH DIED IN THE ZOOLOGICAL GARDENS. 909 


SPrruRINa Railliet, 1915. 


CYLICOSPIRURA SUBAQUALIS (Molin, 1860), gen. n. 


Tn 1913, Seurat described a Spirurid from the stomach of Felis 
ocreata Gmeiin (10). This worm he considered to be identical 
with Spiroptera subequalis Molin (11), and placed it in the genus 
Spirocerca, as it conformed in many respects to the type-species 
of that genus (Spirocerca sanguinolenta of the Dog). 

He published a description with a drawing of the mouth 
capsule of the worm taken from a single specimen, which he 
states was very much flattened by the pressure of the cover-glass. 
My material consisted of three females and one male collected 
from the stomach of a Tiger (felis tigris), Malay States. 

The measurements of these specimens correspond in every way 
to those of the worm described by Seurat from /’. ocreata, except 
in regard to the mouth capsule, which is much wider in the 
latter, due no doubt, as Seurat suggests, to the pressure. This 
writer also states that the chitinous teeth in the mouth capsule 
are tricuspid, whereas in my specimens they are bicuspid. 

Von Drasche in 1882 revised Molin’s type material, and figures 
the cephalic and caudal extremities of the male. In his figure 
ef the mouth-parts he shows each of the chitinous teeth as 
bicuspid (12). 

LT have also compared my material with an unpublished drawing 
of the cephalic extremity made by Prof. R. T. Leiper from 
Molin’s type-specimens in the Vienna Museum, and f find it 
agrees In every respect. 

Seurat, therefore, has either made an error of observation, 
due perhaps to the flattened state of his specimens, or he has 
confused another species with S. subequalis, of Molin. 

Moreover, as the mouth capsule of this worm with its armature 
of teeth in no way conforms to the type of the genus Spirocerca 
(S. sanguinolenta), I wish to propose a new genus, Cylicospirura, 
with C. subequalis as the type-species. 


CYLICOSPIRURA, gen. n. 


Generic diagnosis.—Spirurine: Body elongated, tapering 
slightly anteriorly. Mouth circular, surrounded by six small 
papille. The mouth capsule is deeper and conical in shape, with 
the apex of the cone in apposition to the anterior end of the 
esophagus. It is provided with six triangular chitinous plates 
arranged radically, each of the internal free ends of which termi- 
nates in a bicuspid tooth which projects slightly beyond the 
entrance to the mouth capsule. 

The tail of the male is twisted spirally, and is furnished with 
a narrow symmetrical bursa, which has four pairs of preanal and 
two pairs of postanal papille. The spicules are unequal, the 


910 MR. G. M. VEVERS ON NEMATODA FROM MAMMALS 


long spicule being in the type-species more than five times the 
length of the short spicule. 

The vulva is situated in the anterior half of the body. The 
female is ovoviviparous. 

Endoparasitic in the stomach of Carnivora. 

Type-species, Cylicospirura subcequalis (Molin, 1860), gen. n. 


Text-fig. 7. 


Text-fig. 6. 


S00 yu. 


100 ps. 


Cylicospirura subequalis (Molin), gen. n. 
Mouth capsule. Caudal extremity of male. 


STREPTOPHARAGUS ARMATUS Blanc, 1912. 


In 1912, Blane gave a short description (without figures) (13) 
of this species in what he termed a preliminary note. As he does 
not seem to have continued his researches on this worm, I have 
made some drawings from material collected from a Pig-tailed 
Macaque (Macacus nemestrinus) from India. 

No mention is made in the original description of the guber- 

naculum in the male; this was probably an oversight. 

Spiroptera pigmentata v. Linstow, 1897 (14), from Cercopithecus 
albigularis Africa, undoubtedly falls into this genus, but must be 
regarded as a separate species on account of the difference in the 
length of the spicules. 


WHICH DIED IN THE ZOOLOGICAL GARDENS. 911 


Seraria Viborg, 1795. 
From the peritoneal cavity of Tragulius stanleyanus, from Java, 
two species of Setaria were recovered: of one species only four 
females were found, and of the other one male and one female. 


Both of these species are apparently new, and one falls into a 
new genus. 


Text-figure 8. 


f Kf i i 
a a 
ier 


Streptopharagus armatus Blanc. 


a. Mouth. 


6. Caudal extremity of female. 
e. Cephalic extremity of male. 
d. Caudal extremity of male, showing spicules and accessory piece. 


SETARIA JAVENSIS, Sp. n. 


Host. Tragulus stanleyanus. 

Locality. Java. 

Habitat. Peritoneal cavity. 

Material. Four female worms; no males were collected. 

Specific diagnosis —— Setaria: Body tapering gradually towards 
the anterior end of the body; head rounded and not separated 
from the rest of the body. 

The peribuceal ring is ovoid, the dorso-ventral diameter being 
greater than the lateral. 

Four submedian head papille are present, 404 from the 
anterior end of the body. 

The nerve ring is situated 200 uw from the opening of the 
mouth, 


912 MR. G. M. VEVERS ON NEMATODA FROM MAMMALS 


Female 115 mm. in length and ‘5mm. in maximum breadth. 


(Hsophagus has a total length of 7-2 mm. and a maximum breadth 
of 110 p. 
The anterior portion of the csophagus is 400 in length. 


Text-figure 9. 


\ 


100 p. 


Setaria javensis, sp. n. 


a. Cephalic extremity of female. 
6. Caudal extremity of female. 


The anus is distant 320 » from the caudal extremity, which 
tapers gradually and ends in a small knob. The vulva is situated 
at a point 500 « from the anterior end. Ovoviviparous. 

The embryos in wéero measure 2104 X 7p. 


In addition to the foregoing species, I recovered another 
Filariid worm from the abdominal cavity of the same host 
(Tragulus stanleyanus). ‘This species 1s evidently very closely 
related to the genus Setéaria, but there are two points of generic 
importance which separate it from this genus—namely, the struc- 
ture of the mouth-parts and the presence of bosses irregularly 
scattered over the cuticle of both sexes; in other respects it agrees 
with the generic diagnosis of Setaria, as given by Boulenger 


(15). 


WHICH DIED IN THE ZOOLOGICAL GARDENS. 913 


PAPILLOSELARIA, gen. n. 


Generic diagnosis.—filartine: Body cylindrical, filiform, taper- 
ing considerably at the posterior extremity in both sexes. 
The mouth is guarded by two lateral lips, which are lined by 


Text-figure 10. 


400p. 
100p:. 


6 (a 


Papillosetaria traguli, gen. n., sp. n. 
a. Caudal end of male, showing spicules and papillee. 
b. Cephalic extremity of female, showing vulva. 
c. Caudal extremity of female. 


an oval ring of chitin. The cuticle of both sexes is covered 
irregularly with bosses, except in the regions of the head and 
tail. There are four head papille—two lateral and two sub- 


914 MR. G. M. VEVERS ON NEMATODA FROM MAMMALS 


median. The cesuphagus consists of two parts—a short anterior 
portion and a longer and thicker posterior portion. 

The male is Smaller than the female, its attenuated caudal 
extremity ending in a close spiral. Preanal and postanal papillee 
are present. The spicules are unequal, the longer consisting of 
two parts, the shorter dagger-shaped with an expansion at the 
proximal end. 

The tail of the female is curved dorsally, and bears two lateral 
appendages close to the caudal extremity. The vulva is near 
the anterior end of the body. The eggs are thin-shelled. Ovo- 
viviparous, parasitic in the peritoneal cavity of mammals. 

Type-species, Papillosetaria traguli, sp. n. 


PAPILLOSETARIA TRAGULI, sp. n. 


Host. Tragulus stanleyanus. 

Habitat. Peritoneal cavity. 

Locality. Java. 

Specific diagnosis.—Papillosetaria: Body tapering at both 
extremities, more especially towards the tail. Head rounded, 
not separated from the remainder of the body. The mouth is 
guarded by two lips, iined by an oval ring of chitin which 
projects very slightly beyond the mouth. The head is surrounded 
by four papille: two lateral and two submedian. ‘The body, 
with the exception of the two extremities, is irregularly studded 
with staall cuticular bosses. 

Male.—Leneth 5:7 em.; maximum breadth 280 yp. 

The anterior portion of the csophagus measured 500 p» in 
length, the posterior region 9°1 mm. 

The nerve ring is 200 » distant from the anterior end. 

The tail is coiled in a close spiral. , 

The ano-genital opening is 120 w from the caudal extremity, 
which ends in a small rounded knob. 

There are three pairs of postanal and four pairs of preanal 
papillee. 

The spicules are unequal, the larger measuring 370 » in length 
and consisting of two portions—an anterior cylindrical portion 
and a terminal twisted portion ending in a sharp point. 

The shorter spicule is 85 » in length, and also consists of two 
portions—a proximal expanded portion and a distal dagger- 
shaped portion. 

Female.—Length 14°5 em.; maximum breadth 460 p. 

Anterior portion of the cesophagus is 400 » long, the posterior 
portion 9°6 mm. 

The anus is °5 mm. from the caudal extremity; the tail portion 
has a distinet curve dorsally. 

The tail ends in a bifid appendage ; near the extremity are two 
lateral appendages. 

The vulva is 650 from the head end. 

Ovoviviparous; embryos measure 230 « in length. 


WHICH DIED IN THE ZOOLOGICAL GARDENS. 915 


NeEMATODS PARASICtES ARRANGED ACCORDING to Hosts. 
PRIMATES. MAMMALIA. 


CERCOPITHECIDS. 
MAacacus NEMESTRINUS. Pig-tailed Macaque. India. 
tCharacostomum asmilium R., H. & Joyeux, 1913. 
Hsophagostomum apiostomum (Willach, 1891), R. & H., 
1905. 
Trichuris trichiwra (Linneeus, 1771), Stiles, 1901. 
TStreptopharagus armatus Blane, 1912. 
Macacus RHESUS. Rhesus Monkey. India. 
Bsophagostomum apiostomum (Wallach, 1891), R. & H., 
1905. 
Trichuris trichiura ( Linneus, 1771), Stiles, 1901. 
TSpirocerca sanguinolenta (Rud., 1819), R. & H., 1911. 
Paprio sPHINX. Guinea Baboon. Africa. 
Trichuris trichiura (Linneus, 1771), Stiles, 1901. 


CEBID. 
CrBUS FATUELLUS. Brown Capuchin. Guiana. 
Filaria gracilis Rudolphi, 1809. 
LaGorurix INFUMATUS. Smoky Woolly Monkey. S. America. 
Filaria gracilis Rudolphi, 1809. 
JALLITHRIX JACCHUS. Common Marmoset. Brazil. 
Subulura distans (Rudolphi, 1809), R. & H., 1912. 


PROSIML#. 
LEMURID&, 
NYcTricEBUS TARDIGRADUS. Slow Loris. Malay. 
tAncylostoma malayanum (Alessandrini, 1905), R. & H., 
1909. 
Tt Characostoma asmilium, R., H. & Joyeux, 1913, 


CARNIVORA. 
FELIDG. 

Friis neo. Lion. Africa. 
Toxascaris leonina (v. Linstow, 1902), R. & H., 1911. 

Frnis vrcris. Tiger. Mailay. 
Belascaris mystax (Leder, 1800), Leiper, 1907. 
Galoncus pernicosus (v. Linstow, 1886), Railliet, 1918. 
Ancylostoma ceylanicum Looss, 1911. 

TAncylostoma pluridentatwnr( Alessandrini, 1905), R.& H. 
1909. 

Cylicospirura subequalis (Molin, 1860), gen. n. 


) 


916 MR. G. M. VEVERS ON NEMATODA FROM MAMMALS ‘ 


Friis parpus. Leopard. India. 
Belascaris mystax (Zeder, 1800), Leiper, 1907. 
Gnathosloma spinigerum Owen, 1836. 
Ancylostoma ceylanicum Looss, 1911. 


Fetis NeBULOSA. Clouded Tiger. Assam. 


Belascaris mystax (Zeder, 1800), Leiper, 1907. 
Gnathostoma spinigerum Owen, 1836. 
Galoncus tridentatus Khalil, 1922. 
Ancylostoma ceylanicum l.ooss, 1911. 
~Habronema chevreuxi Seurat, 1913. 


FELIS VIVERRINA. Viverrine Cat. India. 


Belascaris mystaa (Zeder, 1800), Leiper, 1907. 
Ancylostoma ceylanicum Looss, 1911. 


FELIS BENGALENSIS. Leopard Cat. H. Indies. 
Belascaris mystax (Geder, 1800), Leiper, 1907. 
Gnathostoma spinigerum Owen, 1836. 

tTroglostrongylus troglostrongylus, gen. N., Sp. 0. 

Fenis syLvEstris. Wild Cat. Inverness-shire. 

Belascaris mystax (Zeder, 1800), Leiper, 1907. 


CyN&LURUS JuBATUS. Cheetah. Africa. 
Belascaris mystax (Zeder, 1800), Leiper, 1907. 


VIVERRIDA. 
PARADOXURUS HERMAPHRODITUS. Malayan Paradoxure. 
Malay. 
Gnathostoma spinigerum Owen, 1836. 
Rictularia plagiostoma (Wedl, 1861), Will.-Suhm, 1873. 
Muneos munco. Indian Mongoose. India. 
tFilaria martis Gmelin, 1790. 


CANIDS. 
VULPES VULPES. Common Fox. Britain. 
Belascaris marginata (Rud., 1802), Gedoelst, 1911. 
Rictularia affinis Jagerskiold, 1910. 
Uncinaria crintformis (Goeze, 1782), Railliet, 1899. 
Canis LAGopus. Arctic Fox. Arctic regions. 
Toxascaris limbata R. & H., 1911. 


Canis PRiM&vUs. Indian Hunting Deg. India. 
Spirocerca sanguinolenta (Rud., 1819), R. & H., 1911. 


CANIS AUREUS. Common Jackal. India. 


Toxascaris limbata R. & H., 1911. 
Ancylostoma caninum (Krcolani, 1859), Hall, 1913. 


WHICH DIED IN THE ZOOLOGICAL GARDENS, 917 


Canis suBpatus. Red Wolf. 8S. America. 
Ancylostoma caninwm (Evcolani, 1859), Hall, 1913. 


CANIS OCCIDENTALIS. Arctic Wolf. S. America. 
Toxascaris limbata R. & H., 1911. 


+CANIS AZAR. Azara’s Fox. S. America. 
Rictularia cahirensis Jigerskiold, 1910. 


LYCAON CAPENSIS. Cape Hunting Dog. 8S. Africa. 


Belascaris marginata (Rud., 1802), Gedoelst, 1911. 
Toxascaris limbata R. & H., 1911. 


MUSTELID4. 
ConEPATUS PROTEUS. Cordova Skunk. Argentine. 
Ancylostoma conepati Solanet, 1911. 


URSIDA. 
Ursus arcros. Brown Bear. Caucasus. 
Ascaris transfuga Rudolphi, 1819. 


OTARIIDS. 
OTARIA CALIFORNIANA. Californian Sea-Lion. N. Pacific. 
Contracecum osculatum (Rud., 1802), Baylis, 1920. 


PROBOSCIDEA. 
ELEPHANTIDA, 
ELepnas 1npicus. Indian Hlephant. India. 
TLeiperenia galebt Khalil, 1922. 


UNGULATA. 
EQuID:. 

Equus onaGER. Onager. African origin. Bred in Gardens. 
Strongylus vulgaris (Looss, 1900), R. & H., 1909. 
Strongylus edentatus (Looss, 1900), R. & H., 1909. 
Cylicostomum nassatum (var. parvum), Yorke & McFie, 

1918. 
Oylicostomum bicoronatum (Looss, 1900), Gedoelst, 1903. 
Cylicostomum goldi Boulenger, 1916, 
Oxyuris equi Schrank, 1788. 


Equus Grevyr. Grevy’s Zebra. Africa. 
Probstmayria vivipara Ransom, 1907. 
Cylicostomum nassatum (var. parvum), Yorke & McFie, 
1918. 
Strongylus vulgaris (Looss, 1900), R. & H., 1909. 
Prog. Zoon, Soc.—1922, No. LX1i. 62 


918 MR. G. M. VEVERS ON NEMATODA FROM MAMMALS 


Bovips&. 
CrepHAtopHus sp. Duiker. S. Africa. 


Setaria lubiato-papillosa (Alessandrini, 1838), R. & H., 
IMI 


Hipporracus Equinus. Roan Antelope. Africa. 
tSetaria hornbyi Boulenger, 1920. 
Hemonchus contortus (Rud., 1803), Cobbold, 1898. 


Ovis vienrEI. Urial Gad. India. 
Trichuris ovis (Abildg., 1795), Smith, 1908. 


AMMOTRAGUS LERVIA. Barbary Sheep. Morocco. 
Hemonchus contortus (Rud., 1803), Cobbold, 1898. 


TRAGULIDEA. 


TRAGULUS STANLEYANUS. Stanley’s Chevrotain. Java. 
TSetaria javensis, sp. N. 
tPapillosetaria tragult, gen. n., sp. n. 


[+ Signifies that the parasite has not been found in this host before. ] 


REFERENCES. 


1. Vevers, G. M. 1920.—‘ Report on Entozoa collected from 
Animals which died in the Zoological Gardens of 
London during eight months of 1919-1920.” Proe. 
Zool. Soc. Lond, 1920, p. 405. 

2. Kaauin, M. 1922—‘*A Revision of the Nematode Para- 
sites of Elephants, with a description of four new 
species.” Proc. Zool. Soc. Lond. 1922, pp. 205-279. 

Kuan, M. 1922.—‘* Galoncus tridentatus sp. nov., a new 
Ankylostome living in Fibrous Nodules in, the Intes- 
tine of a Leopard.” Ann. & Mag. Nat. Hist. ser. 9, 
WO, TX FOE NG: 

3. ALESSANDRINT. 1905.—‘‘Su di alcune Uncinarie parassite 
dell’ uomo ed altri vertebrati.” Boll. Soc. Zool. Ital. 
Roma, anno xiv. p. 23. 

4, Ratuuier & Henry. 1909.—“ Sur la Classification des Stron- 
gylide: (2) Ankylostominz.” C. R. Soc. de Biol., Paris, 
5. fév., pp. 168-171. 

5. Looss, A. 1911.—‘“The Anatomy and Life History of 
Agch ylostomum duodenale Dub.” Rec. Sch. Med., Cairo, 
part i. vol. iv. p. 173. 

6. Crayton Lane. 1916.—‘ The genus Ancylostoma in India and 
Ceylon.” Ind. Journ. “Med. Res. vol. iv. 1916-17, 

TA. 

7. ieee Henry & Joyeux. 1913.—‘‘Un nouveau Stron- 
gylide des Singes.” Bull. Soc. Path. Hxot. tome vi. 
avril 1913, pp. 264-267. 


10. 


11. 


12. 


13. 


14. 


15. 


WHICH DIED IN THE ZOOLOGICAL GARDENS, 919 


. Ratiurer & Henry. 1913.—“ Un Hemostrongyle des bronches 


dun léopard.” Bull. Soc. Path. Exot. tome vi. 1913, 
pp. 451-457. 


. von Linstow. 1873.—“ Hinige neue Nematoden nebst Bemerk- 


ungen tiber bekannte Arten.” Arch. f. Naturg., Berlin, 
39, vol. i. pp. 300-302, figs. 9-16. 

Srurat, L. G, 1913.—“ Sur deux Spiroptéres du Chat ganté 
(Felis ocreata Gmelin).” C. R. Soc. de Biol. vol. lxxiv. 
pp. 676-679. 

Morty, R. 1860.—“ Una monografia del genere Spiroptera.” 
Sitzungsb. d. k. Akad. d. Wissensch. Wien: math.- 
naturw. Cl. vol. xxxvii. pp. 911-1005. 

von Drascue, R. 1882.—‘‘ Revision der in der Nematoden- 
Sammlungen des k.k. zoologischen Hofcabinetes befind- 
lichen Original-Examplare Diesing’s und Molin’s.” 
Verhandlungen Zool.-bot. Gesellsch. in Wien (1882), 
WOle soeana jos INA. 

Buanc, G. 1912.—** Un Nematode nouveau (Streptopharagus 
armatus, n. gen., n. sp.) parasite du Macaque (JZ. cyno- 
molgus).” C. R. Soc. de Biol. vol. Ixxii. p. 456. 

von Lriystow. 1897.—‘“‘ Zur Systematik der Nematoden 
nebst Beschreibung.” Arch. mikr. Anat., Bonn, 
vol. xlix. (3) 20 Mai, pp. 608-622. 

Boutenemr, C. L. 1920.—‘“ On some Filariid Parasites of 
Cattle and other Ruminants.” Parasitology, Camb., 
vol, xu. p. 341. 


CESTODA FROM MAMMALS WHICH DIED IN THE GARDENS. 92] 


42, On the Cestode Parasites from Mammalian Hosts which 
died in the Gardens of the Zoological Society of 
London during the years 1919-1921; witha description 
of a new Species of Cyclorchida. By G. M. VEvERs, 
M.R.C.S., L.R.C.P., F.Z.S8., Beit Memorial Research 
Fellow, and Assistant in the Department of Helmin- 
thology, London School of Tropical Medicine. 


[Received November 6, 1922: Read November 21, 1922. | 
(Text-figures 11-12.) 


During the period under review eleven species of Cestoda were 
collected from thirteen mammalian hosts dying in the Gardens. 
One of these is new to science, and has been placed in the genus 
Cyclorchida Fuhr. ‘This is the first record of a member of this 
genus occurring in a mammal, the only other species (C’. omalan- 
cistrota) being from a bird (Platalew leucorodea}. Two of the 
species are larval forms; of these it is interesting to note 
Multiceps (Conurus) serialis from the lung of the Coypu (Zyopo- 
tamus coypus), bred in Britain. The only other record of the 
occurrence of this parasite in this particular host is that made by 
Pagenstecher in 1877, the Coypu in this ease being one which 
had been in captivity in the Berlin Zoological Gardens (1). 
Apart from this, the only other larval form noted was that of 
Tenia hydatigena (Cysticercus tenuicollis) from three Ungulates. 


Systematic List of Species. 


CYcLoPHYLLIDEA (Adult forms). 
MESOCESTOIDID# Fuhrmann, 1907. 
MesocrstorpEs Railliet, 1893. 
M. lineatus (Goeze, 1782), Railliet, 1893. 
ANOPLOCEPHALID& Kholodk, 1902. 
ANOPLOCEPHALIN® R. Blanchard, 1911. 
Monirezia R. Blanchard, 1911. 
M. expansa (Rudolphi, 1810), R. Blanchard, 1891. 
Liystowin& Fuhrmann, 1907. 
INERMICAPSIFER Janicki, 1910. 


I, pagenstechert Janicki, 1910. 
TI, settit Janicki, 1910. 
I, interpositus Janicki, 1910, 


999 MR. G. M. VEVERS ON CESTODA FROM MAMMALS 


HYMENOLEPIDIDE R. & H., 1909. 
Diryiipun# Stiles, 1896. 
CyctorcHipa Fuhrmann, 1907. 
C. crassivesicula, sp. n. 
T#Ni11p# Ludwig, 1886. 
Tanne Stiles, 1896. 


Heurnococcus Rudolphi, 1801. 
E. granulosus (Batsch, 1786), Rud., 1805. 


Tania Linneus, 1758, 


T. tenieformis (Batsch, 1786), Wolfhiigel, 1911. 
YT’. balaniceps Hall, 1910. 


CYCLOPHYLLID&£A (Larval forms). 
T#niip# Ludwig, 1886. 
Mutticers Goeze, 1782. 


M, (Cenurus) serialis (Gervais, 1847), Stiles & Stev., 
1905. 


Tanta Linneus, 1788. 


T’. hydatigena Pallas, 1766. 
(Syn. Cysticercus tenuicollis Rud., 1810.) 


CYCLOPHYLLIDEA. 
ME8SsOCESTOIDID#& Fuhrmann, 1907. 


Mesocestoides lineatus (Goeze, 1782), Railliet, 1893. 


The occurrence of this species in a Fox and two Badgers, all 
from Britain, 1s interesting, as it opens up the question of the 
occurrence of two species, J/. lineatus and WM. litteratus, in this 
country, and also throws some light on the question of the 
validity of the latter species. 

Hall stated in 1919 (2) that he could not express an opinion 
on the validity of J. litteratus, but that he retained the name 
although he could not see any evidence from material or deserip- 
tions for supposing that it was a separate species. In the 
examination of the material from the three hosts mentioned 
above, certain variations in the size and shape of the head, neck, 
segments, and internal.anatomy were noticed, all of which might 
be accounted for by differences in the mode of preservation. In 
the case of the material from one Badger, the host had been dead 
some time, and the worms were also dead and well extended in 
consequence; in the case of the other Badger the worms were 
collected and killed at once, in consequence of which each strobila 
was much contracted, causing distortion of the internal organs of 
each segment. 


WHICH DIED IN THE ZOOLOGICAL GARDENS. 923 


The material from the Fox was dead when preserved, and 
therefore well relaxed. 

Iam of opinion that all three batches of material collected 
from these three hosts belong to the same species (H. lineatus), 
the differences between them being of a transitional or varietal 
nature, or due to differences in the methods of preservation. 

With regard to the validity of the species J/. litteratus ; 
considering the extreme variability of J/. lineatws, coupled with 
the faet that both have been recorded from the same hosts, there 
seems to be no reason for keeping MW. litteratus as a separate 
species, especially as none of the descriptions given for it, 
including the original by Batsch (1786) (3), contain any points 
on which to differentiate it from J/. lineatus. 


ANOPLOCEPHALID# Kholodk, 1902. 
Linxstowin# Fuhrmann, 1907. 
IyermicapsiFer Janicki, 1910. 


(a) Inermicapsiter pagenstecherit Janicki, 1910. 
(b) Inernucapsifer setii Janicki, 1910. 
(c) Inermicapsifer interpositus Janicki, 1910. 


In listing these three species of /nermicapsifer from the Rock 
Rabbit (Procavia capensis) I act with great diffidence, my reasons 
being similar to those given by Douthitt in his monograph on 
the Anoplocephalide, 1915 (4). 

It was only by the application of the key to the subfamily 
Linstowine, given by Douthitt, that I was able to make any 
attempt to diagnose my material. 


HYMENOLEPIDID# Railliet & Henry, 1909. 
DipyLipin& Stiles, 1896. 
CycrorcHipA Fuhrmann, 1907, e.p. emend. 


Generic diagnosis. Dipylidiine. Rostellum armed with a 
double crown of hooks which have a very large dorsal root and a 
emall hook pertion. 

Genital pores unilateral. Genital canals pass between the 
longitudinal excretory vessels. Cirrus pouch communicating 
with the genital cloaca by a narrow canal opening upon a large 
papilla. Testicles very numerous, entirely surrounding the 
female genital glands. Uterus ventral, growing laterally between 
the excretory vessels into the cortical parenchyma. 

Adults in birds and mammals. 

Type-species, Cyclorchida omalancistrota (Wedl, 1856), Fuhr., 
1907. 


924 MR. G. M. VEVERS ON CESTODA FROM MAMMALS 


‘CYCLORCHIDA CRASSIVESICULA, sp. n. (Text-figs. 11, 12.) 


Host. Paradoxurus hermaphroditus (Pallas). 

Locality. Malay States. 

Habitat. Small intestine. 

Length of strobila 16-20 em. Maximum width 1-75 mm. 

The head is 250 u long by 280 broad. There are four suckers, 
each measuring 100 » in diameter. In front of the suckers there 
is a rostellum 100 w in length and 200 p broad at the base. The 
rostellum is armed with a double crown of ten large and ten 
small hooks which interdigitate one with the other. 
- The large hooks measure 65 p in length from the point of the 


Text-figure 11. 


Cyclorchida crassivesicula, sp.n. Scolex. 
hook to the end of the haft, and 27 » from the point of the hook 
to the end of the guard. 

The small hooks measure 43 «x from the front of the hook to 
the end of the haft, and 13 uw from the point of the hook to the 
end of the guard. . 

The hook portion is relatively very small compared with the 
basal portion formed by the haft and guard. The neck is 
500 » leng by 180 broad. The first segments are equal in 
width to the neck, and are very short, gradually increasing in 
both dimensions to the posterior end of the strobila, where they 

- measure ‘D mm, in length by 1-75 mm. in breadth. 
Tn shape the segments are trapezoidal, broader than long, the 


WHICH DIED IN THE ZOOLOGICAL GARDENS. 925 


posterior border of each segment overlapping the anterior border 
of the following segment. 
The genital pores are unilateral and protrude somewhat. 


Male reproductive organs. 


The testes are 16-20 in number, oval in shape, measuring 
70-100 p in their longest diameter. 

In the younger segments they are arranged in a circle round 
the female genitalia. In the more mature segments they are 
_pushed out laterally by the ovaries but still retain a suggestion 
of their circular arrangement. ‘There is a large seminal vesicle 
which lies in the anterior third of the mature segments. In the 
ripe segments in which the male elements are spent, this vesicle 
resolves itself into a finely-coiled tube, which is continuous with 
the vas deferens. 

The cirrus pouch is 150, long, and opens through a papilla 
into the genital atrium or cloaca. 


Text-figure 12. 


Cyclorchida crassivesicula, sp. 0. 
A. Mature segment, showing male and female genitalia. 
B. aandc. Ripe fertile segments. 
b. Unfertile sezment showing shape of uterus. 


In the mature segments the cirrus is extended; it measures 
140 4 in length and 20 p in breadth at the base, and is covered 
with small spines, which are directed backwards. 


Female reproductive organs. 


The two ovaries lie side by side in the centre of the segment ; 
they are round lobulated organs measuring 160 ~—180p in 
diameter. 

Between the ovaries a very minute shell-gland is situated. 
Immediately beneath the shell-gland lies a group of yolk-glands, 
which together form a lobulated kidney-shaped organ measuring 


926 MR. G. M. VEVERS ON CESTODA FROM MAMMALS 


100 » in length, having the hilum directed towards the shell- 
gland. The vagina, which arises from a point between the 
ovaries, is 165 p in length, 

As it approaches the genital opening it increases in width to a 
maximum of 20 «, and at a point 30 from the genital aperture 
it narrows again to a fine duct, and opens into the genital atrium 
alongside the opening of the cirrus pouch. 

The uterus lies horizontally in the segment, and in the 
unfertilised segment is a sac-like lobulated organ (text-fig. 12, 6). 
In the ripe segment the uterus distends the whole segment and, 
pushes its way to the extreme edges on both sides. The eggs are 
single, oval in shape, and double-shelled. 

The diameters of the outer shell are 1404x100,: the 
embryophore is much smaller, measuring only 40 « in diameter. 
No embryonal hooklets were seen. 

In 1908 Smith, Fox, and White (5) described a cestode from 
Paradoxurus grayi which they named Tenia paradoxuri. From 
their description 7’. paradoxuri would seem to have no affinities 
with C. crassivesicula, the rostellum in the former species being 
unarmed and the genital pores irregularly alternating. 

Up to the present the genus Cyclorchida is represented by the 
single species C. omalancistrota from the Spoonbill (Platalea 
leucorodia). 

Cyclorchida crassivesicula, sp. n., agrees with the generic 
diagnosis of Cyclorchida as given by Ransom (1909) (6) in every 
respect except in the statement that the adults occur only in 
birds ; I have therefore emended the generic diagnosis to read 
“ Adults in birds and mammals.” 

A comparison of the description of C. omalancistrota, as given 
by Lithe in 1909(7), with my material offers no point of generic 
significance upon which to base a different genus. 


Ta#niip# Ludwig, 1886. 
Taniin® Stiles, 1896. 


Echinococcus granulosus (Batsch, 1786), Rudolphi, 1805. 


The diagnosis of this species, as given by Hall in 1919 (2), is of 
such latitude as to admit forms showing a variation of almost 
100 per cent., and is evidently a careful compilation of the 
characters as described by various writers. Although there are 
many cases on record of larval forms which vary considerably 
from the typical Hydatid produced by the species FZ. granulosus, 
up to the present the occurrence of more than one species of 
adult does not seem to have been fully investigated. 

Stiles in 1906 (8) suggested that an adult form from the 
Dingo might be a new species. 

Liuthe in 1910(9) redescribed Tenia oligarthra Diesing from 
Felis concolor, Brazil, and compared it with Tenia echinococcus 
V.Sieb. From his description it would appear to be a different 


WHICH DIED IN THE ZOOLOGICAL GARDENS. 927 


species, and therefore should be included in the genus 
Echinococcus. The specimens of adult Hchinococcus collected 
from the small intestine of the Cape Hunting Dog (Lycaon 
capensis) agree with the broad diagnosis as given by Hall except 
in regard to the eggs and the hooks. The eggs in my material 
measure 50 4 x 40, whereas Hall gives 36 uw x 30 pw as the 
maximum. The shape of the hooks in my material differs con- 
siderably from the figures of Leuckart and Stiles, the notch 
between the handle and the guard being exceedingly deep. 

I have diagnosed my specimens as Hchinococcus granulosus, 
but I do so provisionally, feeling that further study and com- 
parison with forms from other hosts may prove them to be 
another species. 

Until a careful comparison is made of material from Australia, 
South America, Africa, and Europe it would be unwise to make 
any definite decision. 


List of Cestode Species arranged under Hosts. 
CARNIVORA. 
VIVERRIDE. 
VIVERRICULA MALACCENSIS. Indian Civet. India. 
Tenia tenieformis (Batsch, 1786), Wolfhiigel, 1911. 
GENETTA GENETTA. Common Genet. Spain. 
Tenia tenieformis (Batsch, 1786), Wolfhiigel, 1911. 


PARADOXURUS HERMAPHRODITUS. Malayan Paradoxure. 
Malay. 


Cyclorchida crassivesicula, sp. ni. 


CANID&. 
Vupses voters. Common Fox. Britain. 
Mesocestoides lineatus (Goeze, 1782), Railliet, 1893. 
CANIS AUREUS. Common Jackal. India. 
Tenia balaniceps Hall, 1910. 
Lycaon capensis. Cape Hunting Dog. 8. Africa. 
FEichinococcus granulosus (Batsch, 1786), Rudolphi, 1805. 


MUSTELIDA. 


MELEs MELES. Common Badger. Britain. 
Mesocestoides lineatus (Goeze, 1782), Railliet, 1893. 


RODENTIA. 
OCTODONTIDA. 


Myororamus corpus. Coypu. 8. America. 
Multiceps (Cenurus) serialis (Gervais, 1847), Stiles & 
Stev., 1905. 


928 


CESTODA FROM MAMMALS WHICH DIED IN THE GARDENS. 


HYRACES. 


PROCAVIIDA. 


PROCAVIA CAPENSIS. Cape Hyrax. 8. Africa. 
Inermicapsifer pagenstechert Janicki, 1910. 
Inermicapsifer settw Janicki, 1910. 
Inermicapsifer interpositus Janicki, 1910. 


UNGULATA. 


2. 


BovibDaé. 
Hipporragus Equinus. Roan Antelope. Africa. 


Cysticercus tenuicollis Rudolphi, 1810. 
Moniezia expansa (Rud., 1810), R. Blanchard, 1891. 


AMMOTRAGUS LERVIA. Barbary Sheep. Morocco. 
Cysticercus tenwicollis Rud., 1810. 

Ovis vienEI. Urial Gad. India. 
Cysticercus tenuicollis Rud., 1810. 


REFERENCES. 


. PagenstEcHER, H. A. 1877.—“ Zur Naturgeschichte der 
Cestoden.”  eitschr. f. wissensch. Zool. Leipzig, 
Ol soos (ID Nyy, al), jos TWO) s). yolk, 10), 

Haut, M. C. 1919.—“* The Adult Teenioid Cestodes of 
Dogs, Cats, and of Related Carnivora in N. America.” 
Proc. United States Nat. Mus. vol. lv. pp. 1-94. 

. Barscn, A. J. 1786.—Naturgeschichte der Bandwurm- 
gattung uberhaupt und ihrer Arten insbesondere, nach 
den neuern Beobachtungen im einem systematischen 
Auszuge. 

. Doursirr, H. 1915.—“Studies on the Cestode Family 
Anoplocephalide.” Illinois Biological Monograph, 
vol. i. Jan. 1915, No. 3, p. 58. : 

. Suita, Fox, & Wuire. 1908.—‘‘ Contributions to Systematic 
Helminthology.” Univ. Penn. Med. Bull. vol. xx. 
. 282. 

: Re oa B. H. 1909.—‘ The Tenioid Cestodes of North 
American Birds.” Bulletin of the United States 
Museum, No. 69. 

. Line. 1909.—* C. omalancistrota.”’ Die Susswasserfauna 
Deutschlands. Parasitische Plattwittrmer Cestodes, 
Heft 18, p. 99, fig. 102. 

. Stites, OC. W. 1906.—“ Illustrated Key to the Cestode 
Parasites of Man.” Hygienic Laboratory Bulletin, 
No.°25, p. 76. 

. Liun. 1910.—“ Cystotinien Stidamerikanischer eliden.” 
Zool. Jahrbiicher, Supplement 12, Heft 3, p. 689. 


MAMMALS FROM THE INNER HEBRIDES, 929 


43. On a further Collection of Mammals from the Inner 
Hebrides. By Ivor G.S. Montagu, F.Z.8. 


[Received October 14, 1922: Read November 21, 1922.} 
(Text-figure 1.) 


Systematic INDEX. 


Page 

Sorex araneus grantii .. by SCROS ERAS ENIEHE Has lnoeosnoaOHenRM an nants oi 
Apodemus hebridensis ae, eaelbeyss They Soe lcnagneeeweeetientioney OM 
Apodemus hebridensis ghia, subsp. N. 1.0.2.6... eeeeee eee 985 
Apodemus hebridensis tural, subsp. V.  ......... eevee eee 985 

' Apodemus hebridensis larus, subsp. N.  .........0. eee se. 936 
Microtus agrestis fiond, SUbSp. D. ......-.e eee eee eee eee eeeeee ee 940 


(Measurements in the Tables are in millimetres throughout.) 


- The recent description of Hebridean mammals by G. E. H. 
Barrett-Hamilton and M. A. ©. Hinton in P.Z.S. 1913*(1), in 
“A History of British Mammals’ (2),andin Ann. & Mag. Nat. Hist. 
1913 (3), and by M. A. ©. Hinton in Ann. & Mag. Nat. Hist. 
1914 (4), neither completely exhausted the subject nor, in most 
groups, sought to make any definitive summary. Of the three 
wild genera most abundantiy distributed in the Hebrides—Sorew, 
Apodemus, and Microtus,—only Apodemus has been the subject 
of a detailed review, which, owing to paucity of material, was 
of necessity left indeterminate as far as certain islands were 
concerned. Valuable results were accordingly to be expected 
from the examination of further mammal material from these 
localities, and for this reason it was thought desirable to extend 
the scope, originally of an entomological character, of a private 
expedition to the Inner Hebrides undertaken by some students 
at Cambridge University, to include the collection of mammals ; 
specimens, described in this paper, were thus obtained on Islay 
and Jura by I.G.S. Montagu, and on Gigha by G. EK. Hutchinson 
and G. L. R. Hancock. The results achieved by the inspection 
of this and other material in the British Museum collection 
have cleared up several of the indeterminate points in (4), have 
shown the existence of certain forms of Sorex araneus which 
have an interesting bearing on the history of the species, 
and have pointed the way to a thorough revision of the British 
forms of Mierotus. It should be noted that, again, in the case of 
no group is a definitive summary attempted ; final view of the 
relationships of such a group as that treated in this paper is, in 
the writer’s opinion, only possible when forms from all the islands 
inhabited by that group have been examined. Accordingly the 


* (1) “On a Collection of Mammals from the Inner Hebrides,” P. Z. 8. 1913. 
(2) ‘A History of British Mammals.’ Gurney & Jackson, 1912-19. 
(8) “'Three new Voles from the Inner Hebrides, Scotland,” A.M. N.H. 1913. 
(4) “Notes on British forms of Apodemus,” A.M.N.H. 1914. 


930 MR. I. G. S. MONTAGU ON A COLLECTION OF 


plan here adopted has been the tentative indication of these 
relationships without final systematic grouping. 

~ It should be noted here that the work of collection in the 
Inner Hebrides was greatly facilitated by the invariable courtesy 
of the inhabitants. 

The writer further takes this opportunity to express his thanks 

to M. A. C, Hinton, without whose generous advice the results here 
described would largely have lacked their comparative value. 


Review of Sorex araneus forms from the Inner Hebrides. 


The satisfactory description of the characters of the Hebridean 
Common Shrews involves a general survey of those of Great 
Britain and of the most nearly related Continental forms. 


Dorsal Ventral 


Form. Colour. wash. Size. pe. 
S.a. araneus. Continental ...... Dark. Pale to dark. Various. Present. 
S.a.castaneus. British Southern Light. Paletodark. Large. Present. 
S.a.castaneus. Scottish Highland Light. Pale to dark. Small. Present. 
MUTA SLEW Cesena sete cca.) ledecscee rey Me weir ek pe ale Small. Present. 
(Greney IMR Uebeecetapeeron deb eco csecce / Dane, Jae. Small*. Present. 
ligne SRE | Gooteactncobdatanseerecyooo IDeA JN)nseranti Large. Absent. 


* With large cranial characters. 


It will be seen from the above table that no character can 
be cited to distinguish the Jura and Gigha shrews from some 
specimens of the Continental form. Continental S. a. araneus 
includes forms exhibiting distinct local variation from the 
typical; a small form ranges from Denmark, through Northern 
Germany and the Netherlands, into France; the large typical 
form is found not only in Scandinavia but in Southern and 
Eastern Germany; shrews from the Harz Mountains and from 
South-Eastern Norway show a light ventral surface, the former 
thus resembling the Jura and Gigha forms in both size and 
colour. It should, moreover, be noted that the typical form of 
S. a. castaneus is to be found only in the southern parts of Great 
Britain, the shrew of the Scottish Highlands being readily dis- 
tinguishable from it by its generally smaller size and smaller 
skull. These forms all deserve systematic separation ; subspecitic 
status, however, would undoubtedly obscure their relationships 
in such a manner as to render their present recognition inadvis- 
able. It follows that the revision and satisfactory arrangement 
of European shrews, as that of any other thoroughly examined 
group, is impossible without a quadrinominal nomenclature. In 
the absence of such a nomenclature the writer is unprepared 
to make, and would certainly regard as unsatisfactory, an attempt 
to distinguish systematically the forms of Jura and Gigha from 
S. a. araneus. In a similar manner the maintenance of specific 
status for the Islay shrew obscures its relationships, and the 
writer accordingly reduces its status to a subspecific one. 


MAMMALS FROM THE INNER HEBRIDES. 931 


It may be noted as an interesting circumstance that the neigh- 
bouring Highland form of SS. a. castaneus is approached in size 
most closely by the shrew of the island most recently separated 
from the mainland, Jura, and least by that of the longest 
separated, Islay. This coincides with the view that smallness in 
S. araneus is a modern character, developed in Scotland, for 
example, since the separation of Islay from the mainland. 
Similarly, light dorsal coloration might be regarded as yet more 
modern, developed throughout Great Britain since the separation 
of Jura. That largeness in the group is primitive is confirmed, by 
fossil evidence, which can, however, obviously throw no light on 
the question of ancestral colour. The view that dorsal lightness is 
primitive in Common Shrews has been considered to be supported 
by its universal presence in young specimens. If, however, this 
latter view be correct, the Jura and Gigha shrews must have an 
unusual history ; on the one hand they may be regarded as con- 
vergent forms, strikingly parallel with Continental S. a. araneus ; 
on the other hand they must be regarded as representatives of a 
successful modern group, which, having excluded the older light 
group from the European mainland, has itself, in part, been 
excluded from the British mainland. The improbability of both 
these assumptions encourages the statement that the description 
of the Jura and Gigha shrews tends to confirm the view that 
dorsal lightness in S. @. castaneus is a secondary development 
from a dorsal clove-brown condition ; this view is perhaps sup- 
ported by the fact that, even in S. a. castaneus, the young coat is 
of a slightly lighter tint than is that of the average adult. 


SOREX ARANEUS ARANEUS Linneeus. 

Sorex araneus castaneus Barrett-Hamilton & Hinton, P. Z.S. 
1913, p. 823 (1). 

Motoria] examined and Ditnepotee Six adults (3 dg, 3 2) 
collected by Sheppard in May 1912 in Jura. Five ‘adults 


(2 3,3 2) and six young adults (3 3, 3 9) collected by Hutchin- 
son in June and July 1922 in Gigha. 


JURA. HB. T. HF. 
Average of six adults ........0.:0.cceee cece 72°5 35°3 12 

GIGHA. 
2. Adult ¢, 30.6. 22. ... 70 37 13 
oR E MS MRYOU NOP A Ny 76 40 14, 
GA OK age grou) Mk 2 70 32 11°5 
TN BNEW TG POON Tae uel 71 34, 12°5 
15. ON D2 tk ire! 77 375 12 
ie Young adult Ge OONG 2 oer 62 35* 12 
TOMER eee ANON TROD: 63 37 13 
14. Hf ep HELEN nM (el 66°5 34 12 
SNe hte Nh, Only 30016) 99. 71 39°5 13 
Aa) AM ipa ORSON G22: 68'5 38 12 
Ue OU ANT kate Prem gene a a 59 36 1g 
Average ‘of five adults .. nabieivig 72'8 383 12°6 

The term “‘young adult”’ is fohlied Ae specimens in the light summer coat, their 


feet being abundantly clothed with hair. 


* Tail damaged in trap, 


932 MR. I. G. 8S. MONTAGU ON A COLLECTION OF 


Description. Both these forms exhibit a dark, clove-brown, 
dorsal coloration and a light ventral surface with traces of pale 
wash ; they are thus indistinguishable in appearance from certain 
specimens of Continental S. a. araneus. In size (head-and-body 
and condylobasal lengths) they resemble the neighbouring High- 
land WS. a. castaneus in being small. The Gigha form, however, 
has large cranial and jaw measurements not usually associated 
with small length of skull. This may represent a secondary 
development, or may represent certain survived characters from 
the primitive general largeness. The sum of these characters 
indicate that the Jura and Gigha shrews exhibit considerably 
more affinity with S. a aranews than with S. a. castaneus, and it 
would certainly be incongruous to leave them with the latter 
group. While they exhibit sufficiently distinct combinations of 
characters to justify separate recognition, the writer does not 
propose, for reasons stated elsewhere, to obscure their relationships 
by systematising them as subspecies. 


SoREX ARANEUS GRANTII Barrett-Hamilton & Hinton. 


Sorex grantii Barrett-Hamilton & Hinton, P. Z.8. 1913, 
p- 824 (1). 

Habitat. Islay. 

New Material examined and Dimensions. Five adults (2 3,3 2), 


one young adult(@), and one aged specimen (<¢) collected by 
Montagu in June 1922. 


HB. Ns HF. 
Gh ANGIE ia EGOS “Lescubnebodadce dea dec 74, 37 12 
UV. pga eh 18, 6s90 1 Noe) eae eee 35 12 
3 WO 076009.) ot ene ne nee 31 12 
Tey BON PIs C4220 ena tee eee eM TT 37 12 
Oa ne @ Sale. Op.” |e OU naa Nan z 6 38 11°5 
15 eVounesadulfyoe 21 Gh22 myers eee 72 40 12 
TY, Neral Eig WOE GEES: cco coo sodond soe asedine ov 60 39 13 
Average ot 1922 adultsi(5)\\- ae -ceesdeees: 73 35°6 11:9 
Se notallleadulitish(23) meee eee Owls 35:9 12°45 


Description. The new specimens of this form confirm the 
description by Barrett-Hamilton and Hinton in (1). An exami- 
nation of all specimens gives the following figures :— 


pP* present on both sides in 10 specimens, 45 p.c. of those examined. 
ee, » one side ,, 5 i 23) 4 


29 99 
,, absent ,, both sides ,, 7 “ Ba 


79 99 


No. 12 is a most remarkable specimen, the discovery of which 
has an important bearing on the bionomics of the shrew. 
L. E. Adams collected data, his interpretation of which has been 
generally accepted, demonstrating that S. a. castaneus is an 


MAMMALS FROM THE INNER HEBRIDES. 933 


annual, each generation surviving about eighteen months. Thus, 
according to this hypothesis :— 


In spring and early summer of year w there are present adults of 
year «-1 and young of year « bemg produced. 

In late summer and autumn of year # there are present young 
adults of year « and adults of year w-1 dying off. 

In winter of years e-—x+1 there are only adults of year « 
generation. 


The abundant data in support of this view need not be recapitu- 
lated, suffice it to say that all winter specimens were found to be 
indistinguishable in point of age, and that no really aged specimen 
had hitherto been examined. This theory was extended to 
embrace many other Huropean shrew forms. No. 12, however, 
is undoubtedly above normal age; the teeth are almost com- 
pletely worn away, the dorsal black and the ventral silver coats 
are grizzled. No specimen like it is in the British Museum 
collection, nor has one been recorded. It should be noted that 
the proportion (1/29th) of examined S. a, grantii specimens 
that this individual represents does not encourage the view that 
there is a tendency to longevity in the subspecies, but causes 
one to regard its occurrence as being as exceptional as in 
S. a. casteneus. 

In reducing the status of this form to a subspecific one the 
writer is following the method of Barrett-Hamilton and Hinton, 
who, on p. 432 of (2), write: ** This mouse (JZ. a. macgillivrait) 
might well be called a species were it not that the subspecific 
name indicates its relationships.” From this point of view specific 
status was, in the first place, incongruous for the Islay shrew, 
and the necessity for systematising it as a subspecies of S. araneus 
has been enhanced by the description of the shrews from Jura 
and Gigha, the characters of which emphasize the relationships 
of the Islay form. It should be added that this view is accept- 
able to M. A. C. Hinton. 


SorEX MINUTUS MINUTUS Linnezeus. 


Material examined and Dimensions. One adult ( ¢ ) collected 
by Montagu in June 1922 in Jura. 


HB. ae HF. 
ip, NGI ON OR GORL aos 59 38 * ut 
Average of sixteen nabs from the ee iamades (1) 55°5 35°4 10'1 


* Tail damaged in trap. 


Description. This shrew has now been reported in the Hebrides 
from Lewis, N. Uist, Benbecula, 5S. Uist, Barra; Skye, Sanday, 
Higg; Coll, Tiree; Islay, Jura, Gigha; Great Cumbrae, Arran, 
Bute. It had not hitherto been reported from Jura. The 
present specimen, it should be noted, is noticeably large in all 
dimensions. 


Proc. Zoou. Soc.—1922, No. LXITI. 63 


934 MR. I. G. 8. MONTAGU ON A COLLECTION OF 


Sorex skull dimensions. 


Sorex araneus araneus. 


JURA. Chl. Mb. bbe) be: M. 
Average of three specimens ...... 18°50 5°15 9°10 4°85 9°34 
(2) (2) (2) 
GIGHA. 
Gy ANGIOMNG: Gy abo cop sasedompsacs vc ooo JUSTE) 5°64 9°66 5°23 9°68 
TUBS Re al Oe A Rite Sak MoU ene Babli silltey yy 5°38 6°60 5°35 9°62 
Ho gy Mowing axGlOuly B) eso55e 000 18°44 5°36 9°50 5°33 9°73 
Average of eight specimens ...... 18°55 5°39 956 . 5°30 9°74, 
: (4) (4) (3) 
Sorex araneus grantit. 
Isnay. 
SapAtd ulti Mees lS Alt 5°30 9:27 5:23 9°65 
eA amt by cote cause heh GRE eRCa 18°81 5°42 9°24 5°40 9°57 
Saami te) inf LCE U ae Miata eek ge iat ILA G) 5°65 9°50 5°33 9°82 
Te iN eee eae ele Stee ee Ne i PLO BOD 5°38 9°22 5°16 9°70 
ND Aceddl Gi ste tse oat ae eee ORO. 5°50 9°77 5:00 9-76 
Average of nineteen specimens ... 18°85 5°32 9:26 5°15 9°65 
(48) (16) (18) 
Sorex minutus minutus. 
JUBA. 
IWS ANGI So cooaeAesaeepobsessbaendesa. 0 duane yh 4°33 7:45 4°35 7-89 


Notes on Apodemus. 


Hinton remarks, on p. 119 of (4), that “in practically every 
island of the Hebrides, differentiation from the parent stoek has 
proceeded so far that the most logical course would be to describe 
the mice of each island as a distinct subspecies.” In his paper, 
however, lack of abundant material compelled him temporarily 
to group the mice of Ghia and Tiree with those of Great Cumbrae 
as A. h. cumbre, and the mice of Jura and Islay with those of 
Mull as 4. h. maclean. The distinguishing characters he observed 
in them are well confirmed by the extended series now available, 
and others are made apparent; there is left ne doubt of the 
distinct nature of each, which is accordingly here separated. 


APODEMUS HEBRIDENSIS TIRA, Subsp. n. 

Apodemus hebridensis cumbre Hinton, A. M.N. H. 1914, 
p. 128 (4). 

Habitat. Tiree. 


Material examined and Dimensions. Four adults (3,1 2) 
collected by Sheppard in July 1912. 


HB. T. HE. E. 
Type 150. Adult 9, 18.7.12.......... 100 88 22 13°5 
Average of all adults (4) ............... 102°5 84°25 23:71 13°5 


Description. In general dorsal colour this form nearly resembles 
A. h. cumbre; the flanks are slightly less bright than those of 
A. h. cumbre, and are little lighter than the back. The con- 
trast between flanks and belly is not striking; the latter is 
silvery, with slight trace of the pectoral spot. 


MAMMALS FROM THE INNER HABRIDES. 935 


The size of this form is greater than that of the other rufous 
mice, A. h.cumbre and A.h. ghia, with which it was formerly asso- 
ciated. The tails are short, and the ears are considerably 
shorter than those of other forms of A. hebridensis. 

The only complete skull available, though that of an old 
specimen, is noticeably small. The dimensions indicate a broad 
rostrum like that of the Sodor (Gigha, Jura, Islay) forms, and a 
peculiar long, narrow nasal; in other proportions the skull seems 
fairly to resemble that of the Cumbrae form. 


APODEMUS HEBRIDENSIS GHIA, subsp. n. 


Apodemus hebridensis cumbre Hinton, A. M.N. H. 1914, 
p. 128 (4). 

Habitat. Gigha. 

Material examined and Dimensions. One adult (@ ) collected 
by Sheppard in May 1912. Three adults (3) collected by Han- 
cock and Hutchinson in July 1922. 


HB. at HF. E. 

SerAtdul Gal p22en eee eee seeeeceee ss 94, 89 23 14°5 
TeV) Cf) QuzlanNin ined OVE a9 95 23 155 
ES SE TEND OO rats eel 97 88* 25 15 
Average of all adults ............... 95°75 89°7 23°4 15 


* Tail damaged in trap. 


Description. The black dorsal hairs are distributed, not so as 
to dull the general colour from rufous to brown as in the Mull, 
Islay, Jura, and typical 4. h. hebridensis forms, but in more 
definite splashes; thus the back remains rufous with black 
spangled more abundantly in a median position than on the 
flanks. There is bright contrast between the flanks and the 
silvery belly, the line of demarcation being distinct. The pectoral 
spot is slight, though more distinct than in A. h. cwmbre and 
AN e607 

The appendage measurements correspond to those of A. h. 
cumbre and the typical form. 

The skull measurements, when compared with those of 4. h. 
cumbree, indicate a relatively small zygomatic breadth, cranial 
width, and coronal cheek-teeth length. In these measurements 
the skull of A. %. ghia more closely resembles that of the Islay 
than that of the Cumbrae form. 


APODEMUS HEBRIDENSIS TURAL, subsp. n. 


Apodemus hebridensis maclean Hinton, A.M. N. H. 1914, 
p. 130 (4). 
Habitat. Islay. 
Material examined and Dimensions. Seven adults (2 ¢,5 2) 
collected by Sheppard in May and August 1912, two adults (2) 
63* 


936 MR. I. G. &. MONTAGU ON A COLLECTION oF 


in the Royal Scottish Museum, and four adults (3 gd, 1 9) and 
one young specimen (¢ ) collected by Montagu in June 1922. 


HB. Te HF. E. 

Type 2. Adult gj 17.6.22. oe gl 91 25 15 

AV N36. 02," cna amen ET 98 26 16 

WA el) 2 0.86, 22: eaten Ree 98 89 23 14, 

LS ting Cor aK ADA Rau Benen Gos 91 88 24, 16 

DPelmmdlatunercs yo Os O22 merece 62 62 24. 14 
Average of 1922 adults (4)............ $4:25 91°5 24°5 15°25 
Peeeeealleadultsi (ld) eeaeeneeeees 93°6 84:6 23:1 14:4 


Description. This form well resembles A. h. maclean in general 
colour; there is, however, less contrast between the back and 
flanks, the latter being distinctly more rufous than the lighter 
flanks of A. h. maclean. ‘The ventral surface is silvery and the 
pectoral spot even less distinct than in A. h. maclean. 

The appendage measurements of Sheppard have often been 
found to require revision, and the fact that in this case 
they do not agree with those of Montagu indicates that too 
much reliance should not be placed on them. ‘There can, 
accordingly, be little doubt that the ears of Islay mice are longer 
than those of the Mull form, with which they were formerly 
associated. The tail, also, like that of the Cumbrae, Gigha, and 
Jura mice, is noticeably longer than that of the Mull and 
typical A. h. hebridensis forms. The hind feet, far from being 
small as suggested on p. 130 of (4), are extremely large; it may 
be noted that in the young specimen the hind foot is equal to 
40 per cent. of the head and body length. 

The measurements of the recently collected skulls largely con- 
firm the differences noted by Hinton in comparing this form 
with 4. h. maclean. The skull of A. h. tural is to be distin- 
guished from the latter by its greater zygomatic and interorbital 
breadths, broader and deeper brain-case, rather larger bull, and 
broader rostrum and masseterie plate. 


APODEMUS HEBRIDENSIS LARUS, subsp. n. 


Apodemus hebridensis maclean Hinton, A. M. N. H. 1914, 
p. 130 (4). 

Habitat. Jura. 

Material examined and Dimensions. Three adults (2 5,1 2) 
collected by Sheppard in May 1912, and two adults ( @ ) collected 
by Montagu in June 1922. 


HB. eo) SER. E. 

Type 123. Adult g, 18.5.12. ........ 100 59¥ 23 14°5 
TQ Nig. Oy 26ICN22.) eee eG 75 23 15°5 

DO) Mins, NiO 26) 6822. hae Red 94. 23 15°5 


Neue 92°6 83°7 22°8 14°8 


* Tail damaged im trap. 


MAMMALS FROM THE INNER HEBRIDES. 937 


Description. This form resembles A. h. tural in general colour, 
but is dorsally darker, being the darkest form of A. hebridensis ; 
the dark hairs are distributed evenly, though more abundant 
in a median position. The ventral surface is silvery and the 
pectoral spot is faint. 

The appendage measurements resemble those of A. h. tural, 
the ears being longer than those of A. h. maclean. 

Like A. h. tural the skull measurements resemble those of 
A. h. hebridensis and differ from those of A. h. maclean in showing 
larger zygomatic, interorbital, and cranial breadths, larger cranial 
depth, and broader rostrum and masseteric plate. The skull, 
however, resembles that of A. hk. maclean and differs from that of 
A. h. tural in the smallness of its bulle; further, it shows a 
peculiarly large width of the incisive foramina. 


Apodemus skull dimensions. 


(3) (2) (2) (8) (4) (3) 
Gt. Cumbrae. Tiree. Gigha. Islay. Jura. Mull. 
No. A.h. Ah. Ah. A.h. Ah. A.h. 
of D. cumbre. tire. ghia. tural. . larus. maclean. 
a 23°16 22°70 24°36 23°53 23°84 23°73 
la. 100 100 100 100 100 100 
g 110°75 110°00 10790 109°36 109°40 109°83 
3. 56°56 56°00 53°40 54°20 54°65 53°03 
4. 16°80 17°40 17°60 17°69 17°50 17:28 
5. 51:00 50°50 48°20 48°66 49°38 48°26 
6. 35°60 35°50 35°72 35°13 34°99 34,06 
To 45°56 46°30 46°20 45°78 46°03 45°90 
8 26°60 26°45 26°37 26°39 25°81 25°93 
9. 40°23 42°10 39°45 40:05 40°85 41°80 
10. 11:91 11°70 11:90 OT 12°08 11:93 
11. 54°33 55°95 55°90 54°34 54°65 54:03 
12. 28°90 28°65 29°08 29°34 29°47 29:90 
13. 23°56 24°70 22°22 23°40 24°09 24°70 
14, 7-46 773 8°66 8:13 9°09 6°86 
15. 18°96 19°80 19°62 19°30 19°94, 18°83 
16. 10°66 10°35 1141 10°77 11:89 10°26 
17. 16°25 16°25 16:02 15°92 16°22 16°05 


Number in (_) of skulls on which averages are based. 


1. Condylobasal length: average in 8. Condyle to front surface of bulla. 


millimetres. 9. Nasal length. 
Reductions :— 10. Nasal width. 

La. Condylobasal length equals 100. 11. Palatal length. 
9,. Occipitonasal length. 12. Diastema. 
8. Zygomatic breadth. 13. Length of incisive foramina. 
4. Interorbital constriction. 14. Width of incisive foramina. 
5. Cranial width. 15. Rostral breadth. 
6. Cranial depth, in middle. 16. Width of masseteric plate (least). 


7, Postmolar length; condyle to m3. 17. Cheek-teeth, coronal length, 


938 MR. I. G. §. MONTAGU ON A COLLECTION OF 


Mus muscutus muscutus Linneus. 


Material examined and Dimensions. Three (2 3, 1 2) collected 
by Montagu in June 1922 in Islay, and one (¢) collected by 
Montagu in June 1922 in Jura. 


Isnay. HB. a HF. EK. 
NO), ANchulti 5 US, OsPR bloke cop cope von 82 82 18 13 
TSE th eh yr CAO SHE P-AUM Nosebad badeesobn 78 76% 17 13 
SP ES OR RS OA! i ayancaauntoseo 75 75 17 10 
JURA. 
18. Adult ¢, 25. 6.22. 71 71 175 13 


* Tail damaged in trap. 
Description. All the specimens, which were trapped in fields, 
are in the yellowish outdoor pelage. 


Review of the British forms of Microtus agrestis. 


Definitions of various subspecies of JZ. agrestis have appeared 
from time to time (1, 2, 3) describing forms from the Hebrides 
and from the Scottish mainland. These definitions, however, 
have been written not by the same person, and often without 
facilities for surveying the whole material from all these localities ; 
the existing descriptions accordingly lack comparative value in 
some respects, such as character of fur; for such characters are 
largely incapable of reference to other than subjective standards. 
The writer therefore gives here, in tabular form, a comparison 
of the various types of fur recognised as peculiar in British 
M. agrestis forms. 


No. TyPrican. Isnay. GIGHA. ree. Mock. 
1. Intermediate. Sleek. Shaggy. Shaggy. Shaggy. 
2. Soft. Soft. Harsh. Soft. Soft. 

3. Rufous. Rufous. Rufous. Rufous. Rufous. 

4. 12 10 14 14 14 

5. 2 2 2 2 2 

6. General. General. Abundant. General. General. 

rele 16 16 16 16 

8. Irregular yellow. Uniform dark aand b. Uniform More uniform 
grey. white. yellow. 

9. 6 6 6 6 6 

16. Grey, 2 Grey, 2 Grey, 2 White, 2 Grey, 2 

11. 8 8 8 

12. Pale, 2 Bright, 2 Pale, 2 


a, irregular yellow; 6, uniform dark grey. 


Dorsal fur characters :— 
1. Appearance. 
2. Texture. 
3. General colour. 
. Coloured hair, length grey bases. 
0 ey a pe coloured tips. 
. Black hair, distribution. 
Seas alene- thas 


4 
5 
6 
7 


MAMMALS FROM THE INNER HEBRIDES. 939 


Ventral fur characters :— 


8. General colour. 
9. Short hairs, length grey bases. 


LOR a », and colour tips. 
11. Longer hairs, length grey bases. 
TA. Oa, i » and colour tips. 


Muck, MW. a. luch; Higg, UW. a. mial; Gigha, M.a. fiona; Islay, W. a. macgil- 
livraii; Typical, WU. a. exsul; N. Uist, 8. Uist, Jura, Arran, Mull, I. a. neglectus; 
Mainland, Bute. 


An examination of the skulls from all the localities visited 
leads to the conclusion that the present systematic grouping, 
based on the degree of reduction of the 4th inner angle of m1, 
is an inaccurate one. This character is one which, seldom constant 
even locally, varies very readily, and undoubtedly its variation 
does not correspond with degree of differentiation in other 


Text-figure |. 


Das 


ne 
. 


D.6. Dé 


Diagram. Temporal shield: Type 1. D.2. Temporal shield: Intermediate type. 
D.3. Temporal shield: Type 3. D.4. Posterior part of palate: Type 2. 
D.5. Posterior part of palate: Type 1. D.6. Anterior part of jugal of 
M. a. fiona. D.7. Anterior part of normal jugal. 


respects from the primitive type. Steps are being taken to 
secure specimens from most of the hitherto unvisited islands, 
and M. A. C. Hinton and I. G. 8. Montagu are undertaking a 
definitive revision of the group. Accordingly, in spite of the 
incongruous position of many WM. agrestis forms, no attempt is 


940 MR. I. G. S. MONTAGU ON A COLLECTION OF 


made here to define them in conformity with the present estimate 
of their position. An exception is, however, made in the case of 
M. a. fiona of Gigha since the material representing it is new 
and undescribed. It is further considered advisable to indicate 
the lines along which the projected new groupings will be 
effected. 


Group 1. 
Temporal shield type 1. 
M. a. macgilliwran. Islay. 
Fur colour peculiar ; palate type 1; 4th i,a. distinct. 
M. a. ewsul. Jura. 
Fur colour typical; palate type 1; 4th i.a. distinct 
or vestigial. 


Intermediate Group. 


Temporal shield intermediate type. 
M. a. fiona. Gigha. 
Fur colour peculiar; palate type 2; 4th i.a. vestigial ; 
jugal peculiar. 
Group 3. 
Temporal shield type 3. 
Division 1. Palate type 1. 
M.a.mial. Hige. 
Fur colour peculiar: 4th i.a. distinct. 
M. a. exsul. N. Uist, S. Uist. 
Fur colour typical; 4th i.a. distinct. 
M. a. exsul. Mull. 
Fur colour typical; 4th i. a. distinct or vestigial. 
Mixed Division. Palates of both types. 
M, a. neglectus. Mainland. 
Fur colour typical; 4th i. a. distinct, vestigial, or absent. 
Division 3. Palate type 2. 
M. a. neglectus. Bute. 
Fur colour typical; 4th i.a. distinct, vestigial, or absent. 
M. a. exsul. Arran. 
Fur colour typical; 4th i.a. distinct or vestigial. 
M. a. luch. Muck. 


Fur colour peculiar; 4th i. a. distinct, vestigial, or absent. 


MICROTUS AGRESTIS FIONA, subsp. n. 


Microtus agrestis exsul Barrett-Hamilton & Hinton, P. Z. S. 
1913, p. 830 (1). 

Habitat. Gigha. 

Materral examined and Dimensions. One adult (3 ) collected by 
Sheppard in May 1912, six adults (3 ¢, 3 2) and two immature 


specimens (1 g, 1 ©) collected by Hancock and Hutchinson in 
June and July 1922. 


MAMMALS FROM THE INNER HEBRIDES. 941 


HB. T. HE. E. 

8, A\ohilit By Me oP oe 119 39 18 12 
Myppensy ala 2.7522) 121 35 20 115 
Oh By WOUER. 129 38 19 11:5 

i, 4 Op OBO. 107. 35 19 11 
By, Oa Dieeee, 113 36 18 12°5 

1D, VO ino. 117 34 19 12 

18}, Jorenmnennpars Zo 5 Ph P/G IZ oe scae soe ce 97 31 19 9 
Demon 30NG) 20 ulate: 97 32 18 9°5 
Average of all adults (7) ............... 118°7 3557 18:°7 118 


Description. This form is, externally, of peculiar appearance. 
The long dorsal fur is darkened, and rendered harsh by the 
profuse abundance in it of long black hairs. There appear to 
be present in the subspecies two types of ventral surface ; this 
diversity may prove to be a secondary sexual character. Females 
1, 12, 2 mm.) have ventrally only short grey hairs, and exactly 
resemble in this respect WM. a. macgillivraii. Males 3, 5, 9, 
13 (imm.), 127 and femele 6 have, besides the short grey hairs, 
longer hairs with tips pigmented brighter yellow than those of 
the typical form, and distributed in similar irregular patches. 
The character of No. 6 prevents one from formulating a detinite 
opinion of the nature of this diversity until further material is 
available. 

In general body size and dimensions of appendages this form 
resembles M. a. easut. 

The skull shows a temporal shield like that of M/. a. ewsul in 
its main characters, but resembling that of MW. a. macgillivrati 
in certain minor respects. The shield is, further, unique in 
showing angle a* greater than 90°; in all other forms examined 
this angle is less than 90°. The posterior part of the palate is 
grooved. The jugal is peculiar in showing in dorsal view an 
abrupt anterior curve about 3mm. from its junction with the 
rostrum. The 4th inner angle of m1 is distinguishable in all 
specimens but is never more than vestigial. 


* See Diagram 2. 


Anna) 
Heal 


i 


FZ ..(S- 19220) Nie HRA laa 


a VI. Sp. S. An 
ee i ae aan ne OKs 


Kee 


Ce ieee 


‘Fem. 


NEW INDIAN SPECIES OF AULODRILUS BRETSCHER. 


bas 
Wi 
a 


Yi 


(Nn 


yh 
ee, 


ACN 


2e4 2 


NEW INDIAN SPECIES OF AULODRILUS BRETSCHER. 


i 


At ‘ Ha 
NAS 


he 


DINU 
CAN Auli 
Rrtitelts 

y 


by 
i} i 
HH 
¥, atid 
p 


eZee S's 


1922, MEHRA, PI. Ill. 


HA 


i 
j 
HHL + ae 


ees 


NEW INDIAN SPECIES OF AULODRILUS BRETSCHER. 


NEW SPECIES OF THE GENUS AULODRILUS. 943 


44, Two New Indian Species of the little-known Genus 
Aulodrilus Bretscher of the Aquatic Oligocheta belong- 
ing to the Family Tubificide. By H. R. Mrnra, M.Sc., 
Zoological Department, Benares Hindu University. 
(From the Zoological Laboratory, Cambridge.) * 

[Received July 27, 1922: Read November 21, 1922. ] 


(Plates 1.—-11I.+; Text-figures 1-9.) 


CONTENTS. 


Page 

(1) Introduction and Previous Work on the Genus ..... ...... 948 
(2) Diagnosis of the Genus and Species ........................... 945 
(3) Aulodrilus kashi, sp. n. ERM Schad ee OAIG 
(a) Habits and Repersal hamectersy canst epee eh na OAS 
(internal Amatomiyseom iene eee eee en OAS 

(4) Aulodrilus stephensont, SP. Vd. ..0...-0.ccecee eevee veseeeee ees 963 
(@)pExternalyChanacterses noes ec sseseaceencceeee GOS 

(6) Internal Anatomy ........ .. $964 

(5) Remarks on the Systematic postion ia anes enue ... 965 
(G)eS umm aiey eee eaten Se ean tne sep eatin a he imnc cre OO Th 
(7) List of Rerorencest PR eae eA Oe te aetna Ee EOS 


(1) InrRopuctIon AND Previous WoRK ON THE GENUS. 


The material which forms the subject of this paper was 
obtained from the mud at the bottom of a big pond known as 
Rewari-talab in Benares. This pond has its source of supply in 
the waste water from the city waterworks, in which the water 
from the Ganges is mainly used for the supply of the town. 

In the same mud were also found many specimens of Limmno- 
drilus socialis Stephenson and Branchiura sowerbyi Beddard, 
common Indian Tubificids; Branchiodrilus hortensis Stephenson, 
Dero limosa Leidy, and Aulophorus furcatus Oken, gilled Naido- 
morphs. Till recently the only Tubificidee known from India 
were represented by four species, each belonging to a different 
genus. Besides Limnodrilus socialis and Branchiura sowerbyi 
there were only two other rather rare species, Bothrioneuron iris 
Beddard and Monopylephorus parvus Ditlevsen. In a paper 
just published (15) Stephenson has announced the discovery of 
Tubifex tubifex and Aulodrilus remex in that country. It appears 
that Z'ubifex, which is a common European Tubificid, is rare in 
India and probably in the Hast. The species described in the 
present paper belong to the genus Azzlodrilus, which is at present 
the least known genus of the Tubificide. This genus was estab- 
lished by Bretscher (4) in 1899 for A. limnobius in Switzerland. 


* Communicated by Dr. J. SrEPHENSON, F.Z.S, 
+ For explanation of the Plates, see p. 969, 


944 MR. H. KR. MEHRA ON TWO NEW INDIAN 


He places it as an appendix to the Tubificide, though he thinks 
that it fits neither with that family nor with the Lumbriculide, 
and suspects that it may be necessary to found a new family for 
it. Piguet (11) described a second species in 1906 and placed it 
among the Naidide as Vaidiwm pleuriseta; but later transferred 
it to Aulodrilus, which he considers to belong neither to the 
Tubificide: nor to the Naididze, as he saw retractile penes in front 
of the ventral sete of segment 7. He did not complete the study 
of the worm, and so could not be sure about its position. 
Stephenson in a paper (15) published in 1921 describes another 
species, 4. remeaw, from India as above-mentioned, 

The genus, which hitherto has been characterized mainly by 
the character of the needle setze (crotvhets), is now well estab- 
lished, as will be seen from the following description. The 
genital organs, unknown to all the previous authors, are described 
at full length in the present paper and decide the position of the 
genus in the family Tubificide. The fact that the needle sete 
possess a small outer prong (the one on the outer side of the 
curve of the shaft), which is not the usual condition in the 
Microdrili, is not of very great generic importance, because in 
some Tubiticids, e.g. in Tubifex velutinus Grube, some crotchets of 
a similar kind are present; and moreover in A. stephensoni the 
outer small prong is missing altogether, the needle sete being 
thus singly pointed and not forked. 

The hinder end of the body is narrower and thinner, and does 
not show any signs of segmentation. Sometimes a faint indica- 
tion of external segmentation may be visible, but the zone of 
budding lies some distance in front of the anus, where a large 
number of new segments are being formed. One can easily 
recognize the worm by its peculiar hinder end. This feature was 
previously described by Piguet in A. pleuriseta, and Stephenson 
has recently confirmed it in A. remex, and remarks “the most 
posterior region shows no proliferation, nor even any segmenta- 
tion, but there is a zone of proliferation and formation of 
numerous new segments some little distance in front of the anus.” 
Later on, he adds “such a budding zone as that of the present 
genus is so far as I know unique.” 

Aulodrilus connects the genera Branchiura, Kawamuria, 
Psammorcytes, and Tubifex; it also shows some resemblance to 
Monopylephorus africanus Michaelsen in the points that it 
possesses a spermiducal chamber, penial sete and spermathece in 
the 9th segment. In certain points it also bears some similarity 
to the Naididz. One point of similarity, namely the anterior 
position of the genital organs, is however merely a case of extreme 
variation within the genus Aulodrilus and hence not of any 
generic importance. This follows from the description here given 
of a second species, undoubtedly belonging to this genus, in which 
the reproductive organs lie in the usual position for the Tubificide, 
That it is a Tubificid is beyond any doubt, but it shows a greater 
simplicity of structure than any other genus of the family. 


SPECIES OF THN GENUS AULODRILUS. 945 


it is remarkable that the atrium here is situated far forwards, 
and is followed by a convoluted tube of great length which is 
enclosed in a thick covering of muscle fibres forming a structure 
somewhat similar to that of the coelomic sac in Branchiura and 
Kawamuria. I have called the tube following the atrium and 
enclosed in the ccelomic sac the “atrial duct” and restricted the 
term atrium to the anterior rounded portion into which the 
prostate opens. The use of the muscle fibres forming the ccelomic 
sac, or better the muscular sac, is to evert the terminal part of 
the atrial duct, which thus projects out of the “ spermiducal 
chamber ” as a fairly long pseudo-penis. 

The term “spermiducal chamber” used by Goodrich in the 
ease of Vermiculus is retained here too, for a similar structure, 
i.e. the ventral depression of the body-wall, into which the atrial 
ducts open and the penial setze lie. 

The prostate gland as usual in the family is voluminous and 
composed of cells, which when full of secretion become disorganized 
and lose their cellular structure, the secretion being thus massed 
together flows directly into the atrium. The prostate appears to 
arise as a part of the atrial epithelium, which bursts out of the 
atrium as a sort of hernia or outgrowth. 

The penial setal sacs are provided with masses of large pear- 
shaped gland cells, which lie in the body-cavity outside the setal 
sacs themselves though connected with their walls. 

I wish to express my warmest thanks to Mr. F. A. Potts 
and Dr. J. Stephenson of Edinburgh for valuable suggestions in 
connection with this work. 


(2) DiaGNosis oF THE GENUS AND SPECIES. 


The chief characters of Aulodrilus are the following :—Dorsal 
sete capilliform, uncinate or oar-shaped (with thin flattened 
distal extremity); ventral sete uncinate. he uncinate sete 
(crotchets) have the outer prong (the one on the outer side of the 
curve of the shaft) much smaller than the inner, or the outer 
prong may be absent. The penial sete, of very different form, 
are the modified ventral setee of the 7th or 10th segment. The 
atrium is small and subspherical; it is followed by a long con- 
voluted atrial duct, which is enclosed by a thick covering of 
muscle fibres to form the “muscular” or “coelomic sac.” The 
terminal part of the atrial duct is capable of being evaginated to 
form a long soft pseudo-penis. The sperm-sac is formed by the 
backward extension of septum 6/7 or 9/10, and the ovisac by that of 
septum 7/8 or 10/11. The clitellum occupies segments 7 and 8, or 
10 and 11. The spermathece lie in segment 6 or 9. The hearts 
are in segment 6 or 8, Supra-intestinal, sub-intestinal, and 
integumental capillary networks are absent. The formation 
of new segments takes place some distance in front of the anus. 
Asexual reproduction by fission as in the Naididze does not take 
place. 


946 MR. H. R. MEHRA ON TWO NEW INDIAN 


Aulodrilus kashi vesembles A. remex in possessing somewhat 
similar oar-shaped sete in addition to the ordinary needles ; but 
its hair sete are longer than the needles. The first nephridium 
lies in the 11th or 12th segment. It is twice as long as 4. remew. 
The genital organs are in segments 6 and 7, the clitellum is on 
segments 7 and 8, and spermathece are in the 6th segment; the 
spermiducal chamber occupies segment 7. 

In A. stephensoni there are no oar-shaped needles, in which 
point it markedly differs from the above species. The needles 
are here singly pointed and not forked—a point in which 
it differs from all the other species. The genital organs here lie 
in segments 9 and 10, the clitellum extends over segments 10 
and 11 and the spermathece are in segment 9. The penial setz 
are the modified ventral sete of the 10th segment. 


(3) AULODRILUS KASHI, sp. n. 
(a) Habits and Haternal Characters. 


The worms live invariably in tubes formed of small foreign 
particles of sand and debris cemented together with mucus 
secreted by the glands in the skin; in the undisturbed condition 
the posterior portion generally remains outside the tube in the 
water, and is waved gently to and fro somewhat like the tail of 
Tubifex, Limnodrilus, and other Tubificids, while the anterior end 
and greater portion of the body remains buried in the tubes inside 
the mud. On being disturbed, however, the worms withdraw 
their freely moving tails with fair rapidity. When placed isolated 
in water, they do not come out of the tubes very readily, but 
when after a few minutes they do come out, they move like other 
Tubiticids. 

About 40 specimens of this form were obtained between 
February and April, of which 27 were sexually mature. They 
vary from 20-28 mm. in length and hence are much larger than 
A.remex. The hinder end of the worm is much thinner, the 
diameter being 13 mm. The diameter near the anterior end is 
‘26 mm. ‘The colour is yellowish-red due to the blood seen 
through the transparent body-wall. 

The prostomium is bluntly conical, about 90 in length. The 
number of segments varies from 51-70, plus a considerable 
number of undifferentiated new segments, followed by a narrow 
terminal region of about °78 mm. in length, which does not show 
any sign of segmentation (PI. ITI. fig. 11). The number of new 
segments forming in the region of budding varies considerably, 
but it usually is large. The small narrow region behind the zone 
of proliferation and in front of the wide terminal anus Piguet 
regards as physiologically a gill. According to him it possesses a 
rich cutaneous vascular plexus, but this I have not been able to 
observe in my specimens. The anus can dilate during life, 
forming what Piguet calls a ‘‘ branchial fossa.” The anus is wide 


SPECIES OF THE GENUS AULODRILUS. 947 


and terminal. ‘The worms do not reproduce asexually by 
budding. 

Sete.—The setze of both the dorsal and ventral bundles begin 
in the second segment, and are absent in a number of developing 
segments in front of the hinder end of the body. 

There are three kinds of sete in the dorsal bundles: capilliform, 
slightly hooked and doubly forked needles of the sigmoid type, and 
oar-shaped sete with somewhat flattened or fan-shaped distal 
extremity, which sometimes may end in a fine point. The 
number of sete in a dorsal bundle varies from 8-10 of which 
2-4 may be capilliform; these are generally absent in the first 
two or three setigerous segments, and hence begin in the 4th or 
5th segment. 

A hair seta (text-fig. 1, A) is 99-104» in length, always 


Text-figure 1. 


A 
\ F 


A. Hair seta; B. Uncinate seta of a dorsal bundle; C and D. Dorsal sete flattened 
near the distal end (oar shaped) ; E. Uncinate seta of posterior ventral bundle ; 
F. Penial seta. X 540. 


longer but more slender than the needle. It is nearly straight 
with a slight sickle-shaped curve and tapers to a fine point. 
The needles of the ordinary kind (text-fig. 1, B) are, as usual, 
doubly curved and slightly hooked at the distal extremity, 
which is forked. In length this seta varies from 75-92»; it 
bears a swelling or nodulus, which is situated much nearer 
the distal extremity, the proportions of the distances from it 
to the proximal and distal ends being about 2:1. The 
prong on the outer side of the curve of the shaft is smaller 
and much narrower than the one on the inner side, appear- 
ing as fine as a line or a denticle; in some preparations it 
may be broken off, and then the seta appears as a single pointed 
needle. The dorsal sete of the third kind (text-fig. 1, C and D) 
are flattened at the distal extremity and appear oar-like. They 


948 MR. H. R. MEHRA ON TWO NEW INDIAN 


resemble the webbed setze described by Lankester in Tubifex, but 
for the prongs at the sides of the web. The distal end may be 
either rounded or bluntly pointed. Their length as measured in 
two specimens is 66-69 » in one and 78-80 » in the other; the 
nodulus here also is situated near the distal extremity, the 
proportions of the distances proximal and distal to it being 2:1 
as in the ordinary needles. The oar-shaped needles are less 
numerous and are often found in the segments in front of the 
middle of the body (in A. remex these sete begin in segment 13 
in one specimen and 7 in another, but they continue to the hinder 
end of the body). The thin flattened distal end shows com- 
paratively faintly, is marked by one or two vertical lines and has 
not the distinct outline of the rest of the shaft. 

The ventral sete (text-fig. 1, E) are, generally speaking, similar 
to the forked needles of the dorsal bundles. ‘The inner prong is 
much broader, about four times as thick or even more than the 
outer one, which looks lke a fine process and in length may be as 
long as or slightly shorter than the inner one. The shaft of the 
seta in the posterior part of the body is shorter and more strongly 
curved than in the anterior part. In the anterior part of the 
body their length is 90-100 p, in the posterior 60-70. The 
nodulus is situated here also much nearer the distal end, the 
proportions being—the distance proximal to nodulus: distance 
distal to nodulus: 3:2 in the sete in the anterior part, and 
about 2:1 in those of the posterior part of the body. 

The penial sete are always present during the sexual phase, 
and in this species are the modified ventral sete of the 7th 
segment. They lie (Pl. I. fig. 1) inside the spermiducal chamber 
and posterior to the openings of the atrial (ejaculatory) ducts. In 
one specimen the penial sete occupied a position anterior to these 
openings. Their number is 1-3, generally 2 per bundle. Their 
length is about 25 mm. The form of the seta (text-fig. 1, F) is 
very different from that of any of the dorsal or ventral sete. The 
shaft is slightly curved and is somewhat broadened in its distal 
portion; its tip comes toa fine point and looks somewhat like 
the end of a spear. ‘The distal portion has a blade-like inner 
edge and an outer narrow thick border, which is continuous with 
the thick proximal part. There is a narrow longitudinal de- 
pression on one face of the blade, which gives it to some extent 
the appearance of aspoon. The breadth at the base of the 
shaft is about 7», while at the distal expanded portion near the 
tip it is about 12 p. 


(b) Internal Anatomy. 


The cuticle lines the spermiducal chamber, which is therefore 
to be considered as a depression of the ventral body-wall. Large 
mucous cells are a prominent feature of the surface epithelium ; 
these no doubt secrete the mucus, which with the addition of 
foreign ‘particles forms the tube of the worm. ‘The cells of the 


SPECIES OF THE GENUS AULODRILUS. 949 


lateral line are small, about 15 by 6-9 m in size, have an irregular 
form and contain a conspicuous rounded nucleus of 4» diameter ; 
they are only 4-5 in number in each transverse section. ‘ 

The clitellum extends over segments 7 and 8, which contain the 
principal genital organs; it leaves free the area on the ventral 
surface occupied by the spermiducal chamber (Pl. I. fig. 4). It 
is opaque white in appearance and merges gradually into the 
epidermis in front and behind. Its cells are about 70 » in height, 
i. e., about four times as high as the ordinary epidermal cells. In 
the clitellar cells the ceil outlines are lost, and the nuclei dis- 
appear, though these may be visible in the early condition lying 
here and there near the inner border. The cells have a coarse 
granular appearance; the mucous gland cells are present in the 
clitellum also. : 


Text-fig. 2 A. Text-fig. 2 B. 


Vertical sections through the penial setal sac and the glandular masses in con- 
nection with it in two different specimens. The penial seta is rooted in the 
wall of the setal sac. XX 390. 


[For explanation of letters used in the figures, see p. 969. | 


Setal Sacs——In the sexual phase the ventral setal sac of 
segment 7, which contains the penial sete, is nearly circular in 
outline as seen in a transverse section of the body (Pl. I. 
fig. 2). Its wall consists of tall columnar epithelial cells of 
about 18 by 2:5 » size, containing a few nuclei near the outer 
periphery; outside the epithelium there is a thin layer of muscle 
fibres. Cell outlines are not recognizable in the greater portion 
of the penial setal sac, except for a small portion on its ventral 
surface, on account of the secretion, with which the cells seem 
to be filled; the nuclei are also mostly lost in this region 
(text-fig. 3), though they are easily recognized in the ventral 
portion of the wall, where the secretion is not present. The 

Proc. Zoou. Soc.—1922, No. LXIV. 64 


950 MR. H. R. MEHRA ON TWO NEW INDIAN 


setal sac is continued below as a tube to open externally, and 
is there lined by cells which have the same structure as those of 
the epidermis at the opening, out of which the penial sete project. 

A very obvious feature in the constitution of the penial setal 
sac is the presence of several conspicuous lobules of a glandular 
mass, which lie outside it in the body-cavity though connected 
with its wall (text-figs. 2 A & B and text-fig. 3 & PI. I. 
fir. 2). The lobules vary in diameter generally, and each is 
continued at its narrow inner end into the wall of the setal 
sac. The various lobules are separated from one another by a 
few narrow bands of muscle fibres. The epithelial wall of the 


Text-figure 3. 


Vertical section through the penial setal sac of a specimen in a more advanced 
stage of maturity. ‘The glandular masses here are much bigger and lobed. 
The secretion of the gland cells is seen passing into the epithelial wall of the 
setal sac. X 220. 


setal sac, as has been pointed out, does not show any cell out- 
lines for the greater part (text-fig. 3), and this is specially the 
ease where the lobules pass into it, but is filled with a coagulum, 
which is as a matter of fact the secretion. It seems to me that 
the secretion of the gland cells which constitute the various 
lobular masses passes into the epithelial cells and is thus stored 
up there ; it is also likely that the epithelial cells are capable of 
secreting too. Hach penial seta is rooted only in the innermost 
portion of the wall of the sac. The whole structure and appear- 
ance of the glandular masses and their connection with the 
epithelium of the setal sae resembles a somewhat similar condition 


SPECIES OF THE GENUS AULODRILUS. 951 


shown by the prostate and the atrium in these worms, as we shall 
see later. ‘That the epithelial cells of the setal sac, filled with 
secretion generally lose their cellular structure, except in the 
small ventral portion, which is continuous externally with the 
outer tubular part of the sac and not being connected with the 
glandular cells is hence devoid of secretion, can be easily followed 
in sections. 

Each glandular mass consists of a number of somewhat elon- 
gated pear-shaped cells closely pressed together. The cells have 
a frothy appearance on account of the secretion they are filled 
with, have mostly lost their cellular outlines, are deeply stained 
with hematoxylin or similar stains, and are similar to those of 
the prostate; only a few nuclei are present and all this tends 
to show that after secreting they lose their structure. 

The septa are thin throughout the body. The first septum 
lies between the third and fourth segments. 

Alimentary canal.—The buccal cavity is capable of extrusion ; 
the epithelial cells lining it are devoid of cilia. The pharynx 
occupies segments 2 and 3, and is covered over by a small amount 
of chlorogogen pigment; its wall is folded so that about 3-4 
diverticula which may be again divided into two arise from it; 
in one case a fairly long diverticulum arising from the anterior 
part ran independently for some distance and then opened into 
its posterior part. The pharyngeal epithelium is composed of 
high columnar cells with nuclei about the middle of their length, 
and provided with cilia of large size, which arising from the 
opposite walls meet in the somewhat narrow lumen of some of 
the diverticula. Ina few specimens having a more slender body 
the pharynx was not folded, but here the cells of the dorsal wall, 
about 45-70 » in height formed a sort of disc-like structure pro- 
jecting into the pharyngeal cavity, which consequently appeared 
of a somewhat crescentic form. The cells lining the ventral 
portion of the wall here were only 27-36 » in height and 4°5 » in 
breadth. Outside the pharyngeal epithelium lies a fine layer of 
circular muscle fibres, from which fine strands of muscles arise 
here and there, specially on the dorsal and lateral sides and 
connect it with the body-wall. 

The pharyngeal gland cells lie in groups separated by the 
muscle strands, which form a sort of loose network around them 
and largely fill up the body-cavity of the pharyngeal region. Hach 
gland cell takes a deeper hematoxylin stain than the rest of the 
tissues ; it consists of a broad outer expanded portion in which 
lies the nucleus with the nucleolus and a long inner narrow 
portion, which can be easily mistaken for a connective tissue or a 
fine muscle fibre. The cell groups surround each of the pharyn- 
geal diverticula referred to before. In sections most of the gland 
cells are cut in such a way that their long inner stalks become 
completely separated off from their outer expanded portions and 
look like fibres. Hence arises a difficulty in tracing the cells to 
their inner ends. That the fine stalks of the gland cells reach 

64* 


952 MR. H. R. MEHRA ON TWO NEW INDIAN 


the fine basal membrane surrounding the pharyngeal epithelium 
is beyond doubt, and whether they penetrate the epithelium lying 
in between the cells to open into the lumen [| have not been able 
to see. The pharyngeal gland cells extend also behind the 
pharynx and lie in masses of a somewhat irregular kind in the 
body-cavity near the first septum; I believe that these also send 
processes to the alimentary canal. The csophagus occupies 
segments 4-8 and differs from the intestine in its lesser diameter. 
The epithelium is separated from the surrounding layer of muscle 
fibres by a fairly continuous sinus, which is interrupted at a few 
places by the epithelial cells meeting the muscular layer. The 
esophagus suddenly dilates about the middle of the 8th or the 
beginning of the 9th segment to form the intestine, which in 
structure corresponds to the cesophagus; it occupies most of the 
available space in the segment and is attached to the body-wall 
by muscular strands. 

A large number of parasites belonging to the group Ciliata 
Astomata (Pl. III. fig. 12) were found in the intestine of the 
majority of the specimens. 

Blood vascular system.—The blood is yellowish-red in colour 
and devoid of corpuscles. The dorsal and ventral blood-vessels 
are the only longitudinal trunks present, and they are connected 
with each other by a pair of commissural vessels in every segment. 
The dorsal vessel is contractile and extends throughout the length 
of the body; it occupies a dorsal position only in the first six 
segments, after which it turns to lie ventrally on the left side 
near the ventral blood-vessel. 'The ventral vessel divides in the 
first segment to form a pair of slightly coiled vessels which unite 
above to form the dorsal vessel. The peri-visceral vessels are large 
and undulating; they lie in the posterior part of the segment in 
front of the septum. In the region of the pharynx they form a 
series of loops around it; im the posterior portion of the body 
they lie very close to the body-wall and make it appear vascular, 
but they do not branch to form such a cutaneous plexus as is 
present in some of the Tubificids. In the 6th segment the dorsal 
vessel communicates with the ventral vessel by a pair of short 
lateral hearts. The intestine also is surrounded by a sinus, which 
lies outside the gut epithelium and 1s interrupted at a few places 
by the epithelial cells touching the muscular layer outside; it is 
directly connected with the dorsal vessel by means of a short 
vessel once in each segment. ‘The dorsal vessel is usually en- 
larged in the 7th segment, where it turns on the left side; the 
peri-visceral vessels in the region of the sperm and ovisacs become 
enlarged in the sexual phase and lie coiled round them. The 
absence of supra-intestinal and sub-intestinal vessels as well as 
the parietal network indicates the simple condition of the vascular 
system, which is emphasized by the presence of a fairly continuous 
peri-intestinal sinus as in the Naidide. The vascular system 
therefore is much simpler than that of Tubifex, Ilyodritus, and 
Branchiura. 


SPECIES OF THE GENUS AULODRILUS. 953 


Nephridia.—Vhe nephridia are constructed on the same plan 
as in Zubifew. They are unpaired and lie on one side of the 
body near the ventral surface close to the ventral blood-vessel. 
The nephrostome is followed by a short uncoiled tubular portion 
with thin walls, which after piercing the intervening septum 
undergoes several windings, before it becomes continuous with 
the convoluted tube of the glandular mass. A small pear-shaped 
enlargement of the tube before it passes into the glandular mass 
is seen only in sections with some difticulty. The tube in the 
glandular mass is coiled two or three times to form well-marked 
loops; it then emerges as a somewhat wider tube with thin walls, 
on ventral side nearer the anterior end of the nephridium, and. 
not at the posterior end as in Yubifex. The terminal free 
portion of the tube is short and expands near the distal end to 
form a small vesicle, which opens to the exterior at the nephri- 
diopore, a short distance in front of the ventral sete. The 
first nephridium lies in the 1ith or 12th segment, after which it 
is present regularly in every segment, except in the terminal 
portion of the body. 

Nervous system.—The cerebral ganglion is deeply cleft in front, 
slightly so behind. The median lobe, which characterizes the 
brain of Twhifex and other Tubificids is here absent, but a median 
nerve is given off anteriorly to the prostomium. The peri-pharyn- 
geal cords unite in the 2nd segment to form the first ganglion 
of the ventral nerve cord. he giant nerve fibres so charac- 
teristic of the nerve-cord in the Tubificide are absent here, 

Genital organs.—I collected on several occasions between the 
middle of February and the end of April a few specimens having 
these organs at various stages of maturity. The genital organs 
are placed far more anterio} rly than in any other Tubificid, 7. e. in 
segments 6 and 7; in other Tubificide they are never present in 
frout of the 9th segment except in Tudifex (Llyodrilus) bedoti, 
where they le in segments 8 and 9. In this respect Aulodrilus 
kashi comes near the family Naidide, in which the sexual organs 
lie in segments 5 and 6. The gonads are the first organs to appear 
when the period of sexual maturity sets in. In one specimen 
only testes were present; on the other hand where the sexual 
organs were fairly advanced, the testes had entirely disappeared, 
having given rise to an enormous mass of developing sperms, 
which filled the 6th segment and the sperm-sac. The ovaries 
are always present till the latest stage of maturity and are quite 
large, as in other Tubiticids. 

The testes are ovoid masses, one on each side, attached by a 
narrow base to the posterior face of septum 5/6 near the ventral 
body-wall. 

The ovaries in a stage of advanced maturity are a pair of large 
massive organs of a somewhat ovoid or irregular form with a 
small basal stalk, by which they are attached to the posterior face 
of septum 6/7 near the ventral parietes. They remain throughout 
the sexual phase, even when the ovisac is filled with ova and 


954 MR. H. R. MEHRA ON TWO NEW INDIAN 


yolky substance—a feature which distinguishes the 'Tubificidee 
from the Naidid, in which the ovaries disappear early like the 
testes. Hach well-developed ovary measures about :088 mm. in 
height and 095 mm. in breadth; it extends as far back as the 
celomic sac which lies in the hinder part of the segment. It is 
composed of ova at various stages of maturity, somewhat loosely 
arranged and not consolidated as in the Megadrili, and a large 
mass of yolk. The yolk generally fills up a large space in the 
centre of the ovary, while the ova lie all around it forming the 
periphery ; in two cases, however, the yolk extended on the upper 
border, and the ova were lying then on the anterior, ventral and 
posterior margins. It seems probable that only the cells around 
the periphery of the ovary develop into ova, and that the central 
cells break up, lose their individuality and form the large amount 
of yolk-matter. The developing oocytes, when of sufficiently 
large size get discharged with a part of the yolk and find their 
way into the ovisac, where they complete their further growth. 

There are no special blood-vessels for the ovary, or any part of 
the genital apparatus; the peri-visceral blood-vessels and the 
dorsal vessel become much enlarged in the region of the body 
containing them. 

The vas deferens lies in the 7th segment, and opens internally 
by a large seminal funnel in the segment in front, piercing 
septum 6/7 ; it opens externally in a large ventral depression of 
the body-wall—the spermiducal chamber. Hach duct lies in two 
segments and consists of the following parts: the seminal 
funnel, the vas deferens proper, the atrium with the prostate 
and the atrial duct, which is very long, much coiled and enclosed 
in a thick coat of muscle fibres to form a spherical or ovoid 
muscular or coelomic sac. 

The seminal funnels are situated in the 6th segment—z. e. the 
segment in front of that in which the tube itself lies, near the 
ventral parietes, projecting freely and lying in front of the lower 
part of septum 6/7, below the opening of the sperm-sac (text- 
fig. 4). Each funnel about 75m in height is cup-shaped with 
everted lips; in an advanced stage it is wide and shallow reaching 
about 75» in breadth. It is lined by a single layer of columnar 
ciliated cells of 20-28 height except near the margin of the 
upper lip, where the height is somewhat less; the cells are about 
3-6 1 broad, are wider on their inner ciliated border and narrower, 
appearing somewhat fibre-like at their outer end. The epithelial 
cells contain large oval nuclei, 5-6 « in greatest diameter placed 
about middle of their length ; the nuclei are somewhat smaller in 
the lower marginal cells of the funnel. The cilia are fairly con- 
spicuous. The peritoneal layer does not appear to be present 
outside the funnel epithelium. The elongated sperm-heads of 
11-12°6 y length with the tails about four times in length lying 
behind form a regularly arranged dense mass near the funnel 
opening (text-fig. 4). The lower lip of the funnel lies near the 
yentral parietes, about 32 » distance from it. 


SPECIES OF THE GENUS AULODRILUS. 955 


The vas deferens is a short slightly curved tube leading back 
from the seminal funnel and opening behind into the anterior 
end near the ventral margin of a somewhat swollen chamber, the 
atrium. The duct is circular in transverse section, and of nearly 
the same breadth throughout, 7. e. about 25 u. The lumen is only 
4 in diameter. It is lined by cubical epithelial cells of 7°5 uw 
height and without definite cell-outlines ; the oval nuclei of 7:5 
greatest diameter lie at right angles to the height of the cells. 
Outside the epithelium there is present a thin layer of circular 
muscle fibres, which as the seminal duct joins the atrium becomes 
much thicker and is continued into the thick muscular coat of 
the atrium. A few nuclei seen in sections here and there outside 
the muscular layer indicate the presence of the peritoneal layer. 


Text-figure 4. 


5e0. 
2 


Hi 


Ke 


IN CSe Ns} 
111... ~ NV hi) Zeaoreeees 
ee ce 
e 


We 


Semi-diagrammatic vertical section of the vas deferens apparatus as 
compiled from several successive sections. X ca. 540. 


¥ 


In the early stage the vas deferens is shorter and runs nearly 
straight to open at the anterior end of the atrium ; it is also 
narrower, the breadth then being only about 14, including a 
lumen of 2°54. The epithelial cells are cubical, the muscular 
coat around.the epithelium is feebly developed or even in a still 
earlier stage absent. The epithelial cells of the vas deferens are 
altogether devoid of cilia, while in the case of T’ubifex twbifew and 
Limnodrilus claparédeianus the cilia are quite conspicuous. In 
the latter genus I had the opportunity of examining the vas 
deferens on several occasions and found the epithelium provided 
with long cilia arising from the basal granules, which are visible 
in the living worm under a high power. It is also interesting to 
note here, that the vas deferens throughout the sexual phase is 
short. 


956 MR. H. R. MEHRA ON TWO NEW INDIAN 


What corresponds to the atrium in other forms is a long 
. tubular structure consisting of three parts. The first part, that 
into which the vas deferens opens, is a dilated chamber, ovoid in 
form, to which I restrict the term atrium; and the rest of the 
tube I call the atrial duct. This is much convoluted and enclosed 
in a thick sheath of muscle fibres which form a chamber— 
the coelomic or muscular sac. The atrium has its long axis 
parallel to that of the body (Pl. I. fig. 3, text-fig. 4; and 
Pl. If. figs. 5, 6, & 7); in a fully mature form it is about 
143 » in height, 92-114 » in breadth and 192 y or about 
i mm. in length; in the posterior portion however, its height 
is only about 74p. Its wall consists of a muscular sheath of 
5-7 p thickness, which is surrounded externally by a thin layer 
of peritoneum; and of the inner epithelium, which in many of 
the specimens approaching advanced sexual maturity loses its 
cellular character, appears simply frothy and stains like the 
coagulum with which the atrial cavity is filled. Though the cells 
have lost their structural features and nuclei, it can be recognized 
that they have increased in size. The cells lining the hindermost 
part of the atriuin retain, however, their columnar outlines and 
their nuclei. The secretion of the larger, anterior portion of the 
atrium appears to be similar to that of the prostate cells. 

Prostate.—The prostate gland is characteristic of the Tubificide 
among the Microdrili. 

Connected with the atrium on its ventral side nearer its 
rounded anterior end there lies a voluminous mass of gland cells, 
which surrounds it, and a greater portion of the vas deferens in 
front; it has the form of an irregularly shaped lobate mass, some- 
times reaching as far back as the coelomic sac. The mass lies 
dorsally to the ovary, which occupies a ventral position in this 
region of the body (Pl. I. fig. 3). The cells are large and 
pear-shaped with the nuclei contained in the outer swollen 
portion; their narrow inner portions or fine ductules lie near 

the centre of the gland (Pl. II. fig. 5) and converge towards 
the point where ihe prostate is in communication with the 
atrium, while the broad outer portions of the cells lie towards 
the periphery. When the prostate is fairly well developed, 
which was the case in most of the specimens under examination, 
the ductules or the inner portions of the cells lose their entity 
and become more or less disorganized ; and in the period of more 
advanced maturity, the outer portions of the cells become more or 
less dissolved, and the cellular structure disappears, only a few 
nuclei being left here and there. As the cells lose more and 
more in structure, the secretion becomes collected in a mass 
and passes straight into the atrium, where the prostate is 
connected with it (Pl. IL. figs. 6 & 7). The prostate com- 
municates with the atrium by an opening 42 » wide; here 
both the muscular and epithelial layers of the atrial wall are 
absent, so that the prostate appears to arise as an outgrowth 
of the atrial epithelium. In the mature worm the atrial 


SPECIES OF THE GENUS AULODRILUS. 957 


epithelium is thus replaced by a mass of secretion, which has 
apparently originated in the prostate. Vejdovsky (16), who 
studied the development of the atrium and the prostate has 
stated that the latter is formed by the proliferation of the cells of 
the lining epithelium of the atrium at a point, where the 
muscular and peritoneal layers are interrupted, so that the 
prostate cells and those of the atrial epithelium are intimately 
connected with one another; this agrees very well with what 
I find in sexually mature specimens. Though I am not in a 
position to say anything at present as to the origin of the 
prostate, its connection with the atrium certainly suggests its 
origin from the latter. It is, however, clear beyond doubt that 
the prostate cells become disorganized when full of secretion, 
especially in their long inner ductule-like portions, and the 
secretion is discharged mechanically as a stream of inwardly 
moving fluid running through the centre of the glandular mass 
into the atrial cavity at the point of communication. 

Muscular or ccelomic sac.—The atrium is followed by a narrow 
duct of about the same diameter as the vas deferens, which after 
a short length of about 22 w undergoes a few irregular windings 
and may be called the proximal portion of the atrial duct (or the 
middle portion of the atrium, if it is regarded as a part of the 
atrium itself). It is continued into a longer and much wider 
terminal part, which also undergoes several windings, fairly 
regularly arranged one below tie other, till it passes vertically 
downwards to open into the spermiducal chamber. The convo- 
luted middle and distal regions of the atrium which constitute the 
atrial duct are bound up together and enclosed by a thick coat of 
muscle fibres so to form an ovoid structure. somewhat similar to 
the ceelomic sae of Branchiura and Kawamuria; the muscle fibres 
forming its walls are not compactly arranged, but somewhat 
loosely connected, so that there are left some narrow spaces here 
and there in between the strands of fibres, which have a few 
nuclei at certain places. The muscular sac extends ventrally as 
far as the spermiducal chamber; the muscle fibres forming the 
wall are continued in places as strands above and below into the 
musculature of the adjacent body-wall, where they pass through 
the layer of longitudinal muscle fibres to reach the layer of 
circular muscle fibres, and thus attach the sac to the body-wall. 
The height of the sac varies ordinarily from 182 4 to 240; ina 
specimen of smaller size it was 125°5; its breadth varies from 
115-182. The narrow proximal portion of the atrial duct is 
14-17 p in diameter, having a lumen circular in transverse section 
with a diameter of about 5; it occupies the topmost portion of 
the muscular sac, in front of which it leads out from the atrium. 
It is lined by cubical cells of size 1 by 3 ~ with conspicuous nuclei, 
but no definite outlines; outside it there is a thin layer of 
circular muscle fibres, which is continuous in front with the thick 
muscular wall of the atrium. 

The epithelial cells of the wider distal portion of the atrial 


958 MR. H. R. MEHRA ON TWO NEW INDIAN 


duct are of a different appearance altogether. They ave not of 
the same uniform height throughout, being tallest about the 
middle, where they are 8:5 4 high; they are 14°3 » broad at the 
base; the inner portion of the cell projects into the lumen and 
contains the prominent rounded nucleus of 3:5 u diameter (text- 
fig. 4; Pl. Il. figs. 6 & 7). The atrial duct in this region is 
much thicker than in the proximal part, being about 28-40 p 
in diameter as it gradually descends towards its termination : 
its lumen is 16-22 wide. Outside the epithelium there is a 
covering of muscle fibres, which is slightly thicker than that 
of the proximal part of the duct owing to the presence of 
an additional layer of longitudinal muscle fibres external to the 
circular layer. 

The paratrium, a blind tubular outgrowth of the atrium 
characteristic of Branchiura, Kawamuria, and Bothrioneuron is 
here absent. 

The spermiducal chamber is the median quadrangular depres- 
sion on the ventral surface in the 7th segment formed as 
an invagination of the body-wall. The term has been used by 
Goodrich for a similar structure in Vermiculus. The chamber is 
large, about 115 uw deep, 200-250 uw long and about 360 p broad ; 
the margin of its external opening is generally puckered. The 
openings of the atrial ducts lie inside at its anterior angles, one 
on each side, and in several specimens during life one or two 
penes were seen projecting out of it; sometimes, though rarely, 
the chamber is everted to form a papilla-like structure bearing 
the openings of the atrial ducts on its surface. The diameter of 
the terminal portion of the atrial duct near its opening is about 
32, and the epithelial cells lining it gradually become of a 
uniform size, till they have the same form as those lining the 
inner wall of the chamber, which is the inturned epidermis. The 
cuticle of the epithelial lining of the chamber is continuous with 
that of the body-wall outside. The spermiducal chamber acts 
probably as a sort of sucker during copulation. Although there 
is no direct muscular mechanism for deepening the chamber, the 
radiating muscle fibres attaching the muscular sacs to the body- 
wall by contraction can indirectly pull it to a certain extent and 
thus deepen it to produce a sucker mechanism. The chamber 
was more deepened in the specimen in which the penis was pro- 
truded ; and it appears that the contraction of the muscle fibres 
of the muscular sacs, which lie on its top, helps in deepening the 
chamber also. 

Penis.—While examining several specimens in the living con- 
dition I saw in some cases one or two long cylindrical soft penes 
projecting out of the spermiducal chamber for a long or short 
distance. After proper fixation, sections were cut of two such 
specimens and the structure of the penis was investigated. This 
organ is covered externally by epithelium which is really the 
inner wall of the wide terminal portion of the atrial duct, and can 
be distinguished as such by the peculiar character of its cells, which 


SPECIES OF THE GENUS AULODRILUS. 959 


are high and low at regular intervals, and swollen at their inner 
projecting borders, which here lie outwards; the layer of muscle 
fibres outside the epithelium in the atrial duct forms here the 
inner wall (text-fig. 5). In the centre of the penis is a tube—a 
part of the atrial duct, the wall of which is continuous with that of 
the outer tube at the terminal opening. It is obvious therefore, 
that the penis is here formed by the eversion of the terminal 
portion of the atrial duct carrying with it the more proximal 
part as the central. tube. The narrow proximal portion of the 
atrial duct does not seem to be drawn into the penis. The 


Text-figure 5. 


Part of transverse section of the body through the penis, ccelomic sac, 
and a part of the atrium and prostate. X 390. 


eversion of the atrial duct is in all probability brought about by 
the contraction of the muscle fibres forming the muscular sac, 
which in turn are connected with the muscular layer of the body- 
wall. The penis in these worms is not a permanent structure and 
hence is very different from that in many Tubificids; it should be 
regarded as a pseudo-penis. 

The sperm-sac is a median pouch formed by the extension 
backwards of septum 6/7; it opens anteriorly by a wide mouth in 
the 6th segment, and lies generally dorsal to the esophagus in 
the 7th and a part of the 8th segments. Its walls are as thin as 


960 MR. H. R. MEHRA ON TWO NEW INDIAN 


the septum from which it is formed and consist of a thin layer of 
muscle fibres covered both on the inner and outer sides by a 
peritoneal layer of thin flattened cells. It is filled with sperms 
in various stages of development, and is in appearance and’struc- 
ture similar to the sperm-sac in the Naididee. 

A large portion of the body-cavity in the 6th segment is 
separated off laterally and ventrally from the smaller peripheral 
portion, contains all the organs belonging to the segment, and is 
filled with a huge mass of developing sperms. This part is sur- 
rounded by a wall, which is continued dorsally into the body-wall, 
its side walls become continuous ventrally to form the ventral 
portion of the wall, except at the ventro-lateral corners where 
the spermathecal ducts interrupt it as they pass out to their 
openings, and in the transverse sections of the worm here the 
peripheral chamber is thus divided up into a median ventral 
and two lateral parts (Pl. IT. figs. 8&9). The wall which so 
separates the two portions of the body-cavity in the segment 
is composed of modified peritoneal layers, with a layer of 
circular muscle fibres in between. The outer peritoneal layer 
consists of branched cells of an irregular outline, the fibrillar 
branches given off from the various cells anastomose forming 
a sort of loose network, which with fairly big spaces enclosed 
in between gives the whole layer the appearance of paren- 
chyma and is about five times as thick as the rest of the wall 
(Pl. III. fig. 10). The middle layer, consisting of circular 
muscle fibres, becomes continuous with the same layer of the 
ventral body-wall at the openings of the spermathecal ducts. 
The inner layer consists of cells fairly regularly arranged and not 
of the same breadth throughout, less regular on the lateral 
portions of the wall. This layer is continued dorsally into the 
peritoneal layer of the body-wall, which also in this segment may 
be parenchymatous in character, hence sometimes the whole 
peritoneal cavity, 7.e. the space outside the wall of the central 
chamber, is filled with parenchymatous tissue formed by these 
peritoneal layers. The wall of the central chamber in the front and 
hinder parts of the segment gradually comes near the body-wall, 
and consequently the peripheral cavity becomes much more 
reduced ; posteriorly it completely joins the body-wall about the 
level of the ventral sete, in front of the seminal funnel; 
anteriorly it unites with the body-wall near septum 5/6, of which 
it appears to be a backward continuation, for in structure like 
the septum it is composed of three layers, the outer and inner 
peritoneal ones of more or less parenchymatous character with 
the middle one of muscle fibres. It seems clear that a good deal 
of support is afforded to the spermathecal ducts and proper 
protection to the developing sperms by this arrangement. 

The anterior sperm-sac described in many Tubificids is here 
absent. The ovisae is formed as a pouch by the extension back- 
wards of septum 7/8 and lies in segment 8, extending behind nearly 
to septum 8/9. It surrounds a part of the sperm-sac and extends 


SPECIES OF THE GENUS AULODRILUS. 961 


about + mm. distance behind it. It is filled with a large mass of 
yolk granules, and also contains a number of ova at various stages 
of maturity. 

The female funnel is very small. Its posterior wall (the one 
nearer the septum behind) is represented by a patch of columnar 
epithelial cells of about 25 « by 5 p size lying on the anterior face 
of septum 7/8 near the ventral parietes, below the point where this 
septum is bulged backwards to form the ovisac. The anterior 
wall is, however, smaller and somewhat less conspicuous in sections. 
The epithelial cells of the funnel contain oval nuclei of 2°5 
diameter. 

The oviduct is extremely short, about 40, in length; it runs 
backwards and downwards somewhat obliquely to pass through 
_the body-wall, through which it runs as a narrow irregular 
channel of about 25y length and 4y breadth. The female 
opening is wider at the inner margin of the epidermis and 
narrower at the outer. 

The spermathece occupy segment 6; they lie freely for the 
greater part of their length in the central chamber of the body- 
cavity and are attached to the partition wall at the ventro-lateral 
corners as they pass downwards to their openings, which lie about 
the middle of the segment, much in front of the ventral sete. 
They are of considerable length and bent, so that each consists of 
two parts, a vertically elengated terminal portion—the duct, and 
an inner sac-like portion—the ampulla. The duct is not sharply 
marked off from the ampulla, but gradually passes into it. The 
ampulla is simple, and in the later stages of maturity quite large. 
The length of the spermatheca is 234-305 p. The duct is circular 
in transverse section, about 88-133 in length and 48:5» in 
diameter; in one case it measured 175, in length. Its wall is 
17-20 » thick, and is composed of an inner lining of columnar 
epithelial cells, surrounded by a coat of muscle fibres, with a thin 
layer of peritoneum outside. The epithelium is composed of a 
single layer of tall columnar cells of 10-14 height and 3p 
diameter, containing oval cr somewhat elongated nuclei near the 
periphery, which lie with their long axis parallel to the height of 
the cells. The muscular sheath of 2 thickness consists of an 
inner thin layer of circular muscle fibres and an outer thicker one 
of longitudinal muscle fibres. The peritoneum as usual consists 
of a layer of thin flattened cells with prominent nuclei. Near 
the opening the wall of the duct is less thick than in the upper 
part; the epithelial cells here are less tall, being nearly of the 
same height and form as those of the epidermal cells with which 
they are continuous; here the muscular coat is also thinner and 
is continued into that of the body-wall. The nuclei of the epithelial 
cells in the terminal part are not elongated, but somewhat 
rounded, and lie about the middle of the cell. The ampulla is 
much larger than the duct; its length varies considerably—in 
three specimens it was 145, 185, and 295 1; the maximum breadth 
was about 120. Its wall is much thinner than that of the duct, 


962 MR. H. R. MEHRA ON TWO NEW INDIAN 


but the three layers are recognizable. The epithelium consists of 
cubical cells of not more than 6°5 p height; in the advanced stages 
of sexual maturity the cells lining the inner blind portion are 
still lower, while those near the duct are somewhat square in 
outline. The muscular portion of the wall is much thinner, and 
consists of the same layers as in the duct. The thin peritoneum 
continuous with that of the duct forms the outermost layer. 
The greater thickness of the wall of the duct is due to the 
greater height of its epithelial cells and thickness of the muscle 
layers. 

Tn one specimen the spermathecze were found to be in an early 
stage of development (text-fig.6). Here it has the form of a small 
rounded chamber with an external opening and is not differentiated 
into duct and an ampulla. Im size it measures 120p by 90p. 
Its epithelium is composed of tall cells with oval nuclei lying 
about the middle and is surrounded on each side by a mass of 
small peritoneal cells with prominent nuclei, a few of which are 


Text-fig. 7. 


= = BUNA D)e\ (Sh 


esp ey : 
“Lefofete fore feisy) 
m7. 


Stages in the formation of a spermatheca. > 840. 


continued to form a thin layer above. There is no layer of 
muscle fibres present outside the epithelium, the cells of which 
have the same form and structure as those of the epidermis, with 
which they are continuous. In another specimen the spermathecee 
are better developed, and one of them was about 160 » by 68 » in 
size; here they have the form of an upright tube with an external 
opening. It is composed of columnar cells with conspicuous 
nuclei lying fairly near the periphery. Outside the epithelium a 
thin layer of circular muscle fibres surrounded by a few peritoneal 
cells is present, but the layer of longitudinal muscle fibres is not 
yet formed. The wall is 22 thick and the lumen 13, wide. 
Herealso there is as yet no indication of the ampulla (text-fig. 7). 
The spermathece thus arise as small rounded sacs by an 
invagination of the epidermis, then they assume a tubular form, 
but the differentiation into the duct and ampulla takes place 
much later. In both these cases there were no spermatozoa in 
the spermathece. 


SPECIES OF THE GENUS AULODRILUS. 963 


(4) AULODRILUS STEPHENSON], Sp. 0. 


All the worms I obtained conform to the above description 
with the exception of one. ‘This was first examined in the living 
condition and then studied from longitudinal sections. The 
description, though far from complete, gives the main features of 
the anatomy. : 


(a) Haternal Characters. 


The length is about 17°5 mm. The prostomium is bluntly 
conical. The number of segments is 56. The spermathecal 
openings lie on the ventral surface in the 9th segment much in 
front of the ventral sete. The spermiducal chamber lies on the 
ventral surface of the 10th segment: as it is very shallow, the 
opening of the atrial duct is clearly seen in each of its antero- 
lateral corners. During the examination of the living worm this 
chamber was not made out and the openings of the atrial ducts 
were supposed to be paired male openings; but the closer 
examination of the entire mount and subsequently the sections 
revealed the presence of the shallow chamber. 

Setew.— Dorsal and ventral setee are present in all the segments 
from the 2nd segment backwards. The dorsal sete are about 
3-9 per bundle, each bundle being composed of 2-3 hair sete, 
and 1-6 sigmoid needles. The uncinate seta (text-fig. 8) is about 


Text-figure 8. 
A. 


Uncinate seta of A. stephensoni as seen in balsam preparation of the 
entire specimen. X 540. 


half the size of the hair seta and ends in a fine point at the distal 
extremity. The small narrow outer process, or the distal prong 
which is present in the sete of 4. kashi, is not seen. It may be 
that its absence is due to breakage while making preparations. 
The nodulus is situated at about one-third of the distance from 
the distal end, and the distal portion of the shaft beyond it 
projects outside the body-wall. 

The ventral sete are similar to the dorsal needles and are 7-10 
ina bundle. The penial sete are the modified sete of the 10th 
segment and are 1-2 in a bundle; they lie in the spermiducal 
chamber slightly outside the opening of theatrial duct. In shape 
and length they appear like those of A. kashi; the breadth near 
the base is about 5-7 pu. 


964 MR. H. R. MEHRA ON TWO NEW INDIAN 


(b) Internal Anatomy. 


The clitellum is twice as thick as the general epidermis and 
occupies segments 10 and 11. 

The penial setal sac is large, rounded, about 102» in diameter. 
Its wall is about 30 u thick and the lumen 42 4 in diameter. In 
connection with the sac there is a large glandular mass similar in 
structure to that in A. kasht. The first septum lies between 
segments 3 and 4. 

The pharynx extends up to the 4th segment, in which the 
esophagus begins. The intestine commences in the 9th segment ; 
it is larger till the 20th segment, after which it gradually narrows 
towards the posterior end, where again it slightly broadens, being 
stretched by a few muscle fibres, which attach it to the body-wall. 

The dorsal and ventral blood-vessels are connected by, an 
undulating commissural vessel in every segment. From the 
sections I have been able to find the position of hearts in 
segment 8. 

The testes had disappeared, but a large mass of developing 
sperms fills up the central chamber of the-9th segment and the 
sperm-sac as in dA. kashi. The structure and position of the 
seminal duct, prostate, atrium and muscular sac agree in essential 
respects with the description of these organs in A. kashe. 
The seminal funnel is cup-shaped with everted lips appearing 
somewhat like a thistle funnel; it is attached near the ventral 
parietes to the anterior face of septum 9/10 and lies in the 9th 
segment. ‘The columnar epithelial ceils lining it are 14-20» in 
height. The yas deferens (text-fig. 9) is short and 17, in 
diameter, its wall being 6 » thick ; itslumen is 5-6 p in diameter ; 
it is lined by cubical epithelial cells, which are covered by a thin 
layer of circular muscle fibres. The atrium is very much like that 
of the other species, but it is smaller, being 80 u by 45°6 p in size 
and lies here in the 10th segment. The cells lining the cavity 
have lost their cellular appearance owing to the secretion with 
which they are filled, and are surrounded by a thick muscular 
sheath. 

The prostate is large and massive; it extends behind as 
far as the muscular sac. It is similar in structure to that in 
A. kashi, and opens here also into the antero-ventral border 
of the atrium. : 

The atrial duct (text-fig. 9) is much convoluted ; it consists of 
proximal and distal portions, and is enclosed by a thick covering 
of muscle fibres which forms the muscular (ccelomic) sac. The 
atrial duct as it leaves the atrium and before it enters the 
muscular sac is 28 in diameter. Its lining epithelium consists 
of cubical cells, outside which there is a thin layer of circular 
muscle fibres. The proximal portion of the atrial duct is larger 
and has a wider lumen—about 11 —than in A. kashi. The 
distal portion of the atrial duct is 31 » in diameter and is lined 
by an epithelium, which as in the other species is alternately 


SPECIES OF THE GENUS AULODRILUS. 965 


high and low. In the taller portions near the inner margin are 
the nuclei. The muscular sac is smaller in this species; it is 
about 140 u in length and 114 p in height. 

The sperm and ovisacs are formed here by extension backwards 
of septa 9/10 and 10/11 respectively. 

The peripheral portion of the body-cavity in the 9th segment 
is separated here also by a partition as in 4. kashi. 

The ovaries are large and attached to the posterior face of 
septum 9/10 near the ventral parietes lying in the 10th segment. 

The spermathece lie in the 9th segment; in structure they are 
similar to those in the other species, except that the duct here is 
somewhat smalier. 


Text-figure 9. 


Longitudinal section through vas deferens apparatus of 
Aulodrilus stephensoni. X 220. 


The present species is principally distinguished from the former 
by the position of the genital organs, which here lie in segments 
9 and 10. The seminal funnel hes in the 9th segment, while the 
rest of the efferent apparatus occupies the 10thsegment. The sperm- 
sac and ovisac are formed by septa 9/10 and 10/11. The spermi- 
ducal chamber is very shallow and lies in segment 10. The 
uncinate sete are somewhat different from those of the other 
species, and the penial sete are the modified setz of the 10th 
segment. 


(5) REMARKS ON THE Systematic Posrrion oF THESE Worms, 


It will be apparent from the above description that Auwlodrilus 
must be referred to the family Tubificide, although in A. hashi 
the anterior position of the genital organs, which lie here in 
segments 6 and 7, suggests a position closer to the Naidide, in 
which family these organs liein segments 5and 6. In A. stephen- 
sont however, the sexual organs are present in segments 9 and 
10, though in other structural details there does not seem to be 
any marked difference between the two species. This clearly 
decides the position of this genus in the Tubificide. Another 
point, in which these worms resemble the Naididz, is the presence 


Proc. Zoou. Soc.—1922, No. LXV. 65 


966 MR. H. R, MEHRA ON TWO NEW INDIAN 
\ 
of only one sperm-sac, which in appearance and structure is quite 
similar to that in the Naidide; the anterior sperm-sac, which is 
present in many Tubificids is not present here. The main 
portion of the body-cavity in segment 6 or 9 is cut off from a 
peripheral portion, and forms the central chamber, which contains 
all the organs and is filled with the developing sperms—a peculiar 
feature of this genus. ‘That the worms do not reproduce 
asexually by budding, which is a common occurrence in the 
Naidide, separates them sharply from that family. There are, 
however, some features in which Azlodrilus shows a simplicity of 
structure comparable to that met with in the Naidide, e.g. in the 
vascular system, where the presence of hearts in the 6th segment 
in A. kashi* and the absence of supra- and sub-intestinal vessels 
and integumental network is noteworthy; the brain also is with- 
out a median lobe, and there are no giant nerve fibres in the 
ventral nerve-cord. 
The distinct Tubificid characters are as follows :— 


(1) Structure of efferent apparatus with an atrium and massive 
prostate followed by a long convoluted atrial duct enelosed in the 
muscular or coelomic sac. 

(2) Constant presence of a large ovary throughout the sexual 
phase. 

(3) The large massive prostate is connected with the atrium as 
in Tubifex and some other Tubificids. It appears to be developed 
as an outgrowth of the atrial epithelium. 

(4) Presence of a spermiducal chamber. 

(5) Position of genital organs in 4A. stephensont in segments 
9 and 10. 

(6) Penial setal sacs provided with large massive glands as in 
Tubifex (Peloscolex) velutinus. The penial setz are also very long 
like those of the above species, and very different from the ventral 
sete. 


The anterior position of the genital organs in A. kashi, I think, 
is a character only of specific rank not showing in any direct way a 
closer relationship to the Naidide, for in some genera such as 
Megascolex and Buchhelzia t there are a few species in which the 
genital organs are placed one or more segments in front of those, 
which they generally occupy in the genus. In Tubifex (Llyo- 
drilus) bedoti the genital organs lie in the 8th and 9th segments 
and hence more anteriorly placed than in other members of the 
family. Among the Tubificidee Awlodrilus seems to have relations 
on the one hand to genera T'ubifex, [lyodrilus, and Psammorcytes, 
and on the other to Branchiura and Kawamuria. In possessing 
hair sete, the long penial sete and the massive glands in 


* In A. limnobius, A. plewriseta, and A. remex also the hearts lie in the 6th 
segment. 

+ In Buchholzia appendiculata Buchholz, these organs lie in the 7th and 8th 
segments, while in B. fallax Michaelsen, they occupy the usual position in the 
family, 7.¢6., in segments 11 and 12. 


SPECIES OF THE GENUS AULODRILUS. 967 


connection with the setal sacs, and in the absence of the paratrium 
this genus resembles Zubifex and Psammorcytes. But it differs 
from these genera in the following important points :— 

(1) Position of the zone of formation of new segments some 
distance in front of the anus. 

(2) Presence of a spermiducal chamber. 

(3) Presence of a muscular or coelomic sac. 

(4) Variable position of genital organs. 

(5) Absence of spermatophores and a true penis. 

(6) Absence of supra-intestinal and parietal vessels. 


In possessing hair setee, an anterior atrium and a ceelomic sac, 
and in having no spermatophores, it bears a remarkable resem- 
blance to Branchiura and Kawamuria, the chief differences being 
the absence of a paratrium, the presence of penial setz and a 
spermiducal chamber, and the absence of a true penis, which is 
said to be present in Kawamura. 

Aulodrilus resembles Monopylephorus africanus Michaelsen, 
in possessing a spermiducal chamber, penial sets, and sperma- 
thee in the 9th segment, and in the absence of spermatophores 
and a paratrium. 

The spermiducal chamber perhaps acts as a sucker during 
copulation. As this structure is present in Monopylephorus, 
which is however, in other respects very different from the present 
genus, I think this organ is correlated with the absence of a true 
penis and therefore probably is due to convergence in these 
genera. 

(6) Summary. 


(1) The diagnostic characters of the genus and the species are 
given. 

(2) The reproductive organs hitherto unknown are described 
in detail. 

(3) The prostate is large and massive ; it opens into the atrium 
near the antero-ventral margin. The prostate cells soon after 
they are functional and filled with secretion lose their structure 
and become disorganized, while the secretion passes as an inwardly 
moving mass into the atrium at the point where the gland is 
connected with it. The atrial epithelium also at this time under- 
goes a great change on account of the secretion by which its cells 
become replaced having lost their entity. The manner of con- 
nection of the prostate with the atrium suggestis its origin as an 
outgrowth of the atrial epithelium. 

(4) The coelomic cavity in the segment which contains the 
spermathece is separated off from the peripheral portion by the 
formation of a ventro-lateral wall, which is composed of a central 
muscular layer surrounded on either side by a peculiar parenchy- 
matous tissue of peritoneal origin. The central chamber thus 
formed contains all the organs of the segment and is also filled 
with the developing sperms; the spermathecal ducts pass through 
the ventro-lateral corners of the wall on their way to the exterior. 


65* 


968 MR. H. R. MEBHRA ON TWO NEW INDIAN 


(5). The spermathece are ectodermal in origin, and arise as an 
invagination from the epidermis, 

(6) As regards its systematic position Aulodrilus belongs to the 
Tubificide, although it resembles the Naidide in some of its 
features. Among the Tubificide it is related to ZTubifex and 
Psammorcytes on the one hand, and Branchiura and Kawamuria 
on the other. The spermiducal chamber, which is present in 
Monopylephorus, also seems to be independently derived in these 
genera, and perhaps acts as a sucker during copulation. 


(7) List or REFERENCES. 


(1) Bepparp, F. H.—A Monograph of the Order Oligocheta. 
Oxford, 1895. 

(2) Bennam, W. B.—* Atrium or Prostate.” Zool, Anz. xiii. 
pp. 868-372 (1890). 

(3) Bexnam, W. B.—On some new Species of Aquatic Oligocheta 
from New Zealand. Proc. Zool. Soc. London, vol. ii. 
(1903). 

(4) BretscueR, K.—Beitrag zur Kenntnis der Oligocheten- 
fauna der Schweiz. Rev. Suisse Zool, vi. p. 388 (1899). 

(5) Dixon, G, C.—Tubifex. L.M.B.C. Memoir, 1915. 

(6) Eisen, G.—Preliminary Report on Genera and Species of 
Tubificide. Bih. K. Vet.-Ak. Handl. v. no. 16, 26 pp. 
(1879). 

(7) Eisen, G.—Oligochetological Researches. Ann. Report, 
Commissioner of Fish and Fisheries, Washington, 1885, 
pp. 879-964. 

(8) Goopricn, EK. S.—On the Structure of Vermiculus pilosus. 
Quart. Journ. Mier. Sc., n. s. vol. xxxvii. (1895). 

(9) Micuartsen, W.—Zur Kenntnis der Tubificiden. Arch, Natg. 
Jahrg. 74 Bd. 1. pp. 129-162 (1908). 

(10) Micuanisen, W.—Oligochetain‘ DasTierreich.’ Berlin, 1900. 

(11) Pieuerr, K.— Observations sur les Naididées et révision 
systematique de quelques espéces de cette famille. Rev. 
Suisse Zool. xiv. p. 218 (1906). 

(12) Picuxr, K., et Brerscumr, K.—Catalogue des Invertébrés de 
la Suisse, Fascicule 7. Genéve, 19138. 

(18) SrepHeanson, J.—Aquatic Oligocheta from Japan and China, 
Memoirs, Asiat. Soc. Beng. vol. iv. (1917). 

(14) Srepurnson, J.—‘*On a new Species of | Branchiodrilus 
and certain other Aquatic Oligocheta, with Remarks 
on Cephalization in the Naidide.” II. Branchiura 
sowerbyt. Rec. Ind. Mus. vol. vii. (1912). 

(15) SrzPHENSON, J.—‘ Oligocheta from Manipur, the Laccadive 
Islands, Mysore, and other parts of India.” Rec. Ind. 
Mus. vol. xxii. part v. no. 34 (1921). 

(16) Vespovsxy, F.—‘System und Morphologie der Oligo- 
cheten.” Prag, 1884. 


SPECIES OF THE GENUS AULODRILUS. 969 


EXPLANATION OF THE PLATES. 
Explanation of Letters used in Figures. 


a. anus; aér. atrium; ap.at.d. aperture of atrial duct; atz.d. atrial duct ; 
b.v. blood-vessel; 6.w. body-wall; ec. cuticle; c.c. central chamber; cl. clitellum ; 
cem.s. ccelomic (muscular) sac; ¢.m,f. layer of circular muscle fibres; c.s.s. cavity 
of setal sac; d.b.v. dorsal blood-vessel ; d.atr.d. distal portion of atrial duct ; e.gl.c. 
epidermal gland cell; fem,f. female tunnel; gl.c. gland cell; gl.pe.ss. glands in 
connection with penial setal sac; int. intestine; l.m.f. layer of longitudinal muscle 
fibres; /.p.w. lateral portion of partition wall; m. muscles; meg.n. meganucleus ; 
nm. nodulus of seta; o. opening of vas deferens into atrium; ces. cesophagus: 
ov. ovary; ovis. ovisac; p.atr.d. proximal portion of atrial duct; p.c. peritoneal 
cells; pe.s. penial seta; pe.s.s. penial setal sac; per.c. peripheral chamber ; 
ph. pharynx; pr. prostate; pro. prostomium; prs. prostate secretion; prolif. 
proliferating cells; p.z. partition wall; s. seta; se. secretion; sep. septum 6/7 ; 
sf. seminal funnel; sp. sperms; sp.ch. spermiducal chamber; sp.s. sperm-sac ; 
spth. spermatheca; spth.op. spermathecal opening; v.n.c. ventral nerve cord; 
as vas deferens: v.b.v. ventral blood-vessel: v.p.w. ventral portion of partition 
wall. 


[ Figs. 1-12 illustrate Aulodrilus kashi and fig. 13 Aulodrilus stephensoni. 
All the figures except 1, 3, and 13 are drawn with camera lucida. | 


Fig. 1. Ventral view of anterior portion of the worm showing spermathecal 
openings in the 6th and spermiducal chamber in the 7th segment. 
Penial setze and apertures of the atrial ducts lie in the spermiducal 
chamber. X ca. 95. 


Vig. 2. Transverse section through the penial setal sac. X 220. 

Hig. 3. Semi-diagrammatic, compiled from several successive longitudinal sections 
showing the genital organs. X ca. 120. 

Fig. 4. Transverse section of the body through the spermiducal chamber and its 
opening. X 120. 

Fig. 5. Transverse section showing the prostate opening into the atrium. The 


prostate and atrial epithelial cells have mostly lost cellular structure and 
are converted into the secretion. XX 390. 

Figs. 6 & 7. Longitudinal sections through the prostate, atrium, and celomic sac. 
The prostate cells are disorganized, having lost their structure, and the 
secretion passes in a mass into the atrium. The atrial cells have also 
lost their entity, being replaced by the secretion. > 390. 

Figs. 8 & 9. Transverse sections in the spermathecal region. The celomic cavity 
in the 6th segment is divided by a partition wall into a central chamber 
and a peripberal portion. In fig. 8 one spermatheca is opening to the 
exterior. >< 95. 

Fig. 10. A part of section in fig. 8 highly magnified. The partition wall consists 
of a circular layer of muscles in the middle with a layer of parenchy- 
matous cells on either side. > 540. 

Fig. 11. Terminal portion of the body. The part just in front of the anus shows 
no signs of formation of new segments. XX 97. 

Fig. 12. Ciliate astoma parasites. 

Fig. 13. Ventral view of a part of the body in 4. stephensoni showing spermathecal 
openings in the 9th and shallow spermiducal chamber with penial setz 
and openings of atrial ducts in the 10th segment. X ca. 95. 


THE ANATOMY OF A HAMMEREHEAD SHARK. 971 


45. A Contribution to the Anatomy of a Hammerhead Shark 
(Zygena malleus Shaw). By J. H. Luoyp, M.Sc. 
(Birm.), F.Z.S., and Epira M. SuEpparp, B.Sc. 
(Wales), F.Z.8S., Zoological Department, University 
College, Cardiff. 

[Received April 20, 1922: Read November 21, 1922. ] 
(Text-figures 1-7.) 

The specimen on which this paper is based was given to us by 
Professor W. N. Parker, of University College, Cardiff, to whom 
our best thanks are due for his generosity. The specimen was 
sent to him some years ago by the late J. J. Neale, Esq., a local 


trawler owner, but unfortunately the fishermen had eviscerated 
it immediately after it was caught. 


Text-figures 1 & la. 
2@.C. 


Head in Ventral View. 


a.c. ampullary canals. n.g. nasal groove. 
gs. gill slits. m.m. nictitatinz membrane. 
m. nostril, 
la. Dermal Denticle. 


In this short paper we intend to include notes on the structure 
of the skull, brain, cranial nerves, and membranous labyrinth. 
Owing to the long immersion in spirit to which the specimen had 
been subjected the preparation of the skeleton proved to be a 


972 MR. J. H. LLOYD AND MISS BDITH SHEPPARD ON 
matter of some difficulty, as the connective tissue had hardened 
to an extraordinary extent and the cartilage had become exceed- 
ingly brittle. We hope later to publish an account of the 
structure of the visceral arches, limb-girdles, vertebral column, 
and fins. 

A description of this fish appears to be desirable as, with the 
exception of the external characters, we have not been able to 
find any consecutive account of its anatomy. 

Both Garman (4) and Day (8) have given a species definition 
and figured the entire animal, the former including it in the 
genus Cestracion under the name of Cestracion zygena Linn. 
Gegenbaur (5) gives two figures: of the head and briefly mentions 
the species 1n his text, but our observations do not agree with his 
in all respects. The macroscopic structure of the brain has been 


briefly described and figured by Busch (2) and Miklucho-Maclay 
(12). 


SKULL. 


The craniwm bears at its anterior end a stout rostrwm which 
consists of three bars. ‘Iwo of these (d.r.) arise from the dorsal 
surface and run forward and downwards to unite with a median 
ventral bar (v.r.). The latter is expanded anteriorly to form a 
small flattened plate, which is perforated by a single small 
foramen. We were unable to find any trace in our specimen of - 
the rostral appendages (‘ Anhang der Rostrum’) figured by 
Gegenbaur (5). 

At the anterior end of the cranium is a large fontanelle (a.f.) 
extending from between the two dorsal bars of the rostrum to 
the ental bar of the rostrum. This fontanelle, which therefore 
occupies the whole of the anterior end of the cranium, is completely 
covered by a strong sheet of fibrous connective tissue. 

When seen from the dorsal surface (text-fig. 2) the most 
prominent features of the skull are the large, lateral, wing-like 
expansions of the olfactory regions (0.c.). ‘These expansions are 
dorso-ventrally compressed, and their distal extremities taper 
almost to a point. The orbits (0.) are situated immediately 
posterior to the distal portion of the capsules. 

The depression into which the 2nd to 7th cranial nerves emerge 
in a normal skull also bears the eye, and is therefore the orbit. 
In Zygena malleus the orbit is a separate structure, and we have 
therefore decided to call the depression into which the above 
mentioned cranial nerves emerge the pre-auditory depression 
(p.a.d.). This has resulted in a few other changes in the nomen- 
clature of related parts, but it appears preferable to us to do this 
rather than to retain terms which give incorrect impressions of 
the relationships of the regions to one another. 

The orbit is composed of a post-olfactory process (p.o.) which 
runs backwards and outwards from the olfactory capsule to form 
the anterior three-quarters of the circumference of the orbit on 
the dorsal surface. The orbit is completed dorsally by a bar of 


tHE ANATOMY OF A HAMMERHEAD SHARK. 973 


éartilage, the auditory process (a.p.), which arises from the antero- 
dorsal corner of the auditory capsule. ‘The auditory process 
broadens out distally, and the broad distal portion forms the 
posterior fourth of the orbit. The process throughout its length 
acts asa support to the posterior border of the hammer. The 
relations of this bar and the shape of the orbit are incorrectly 
figured by Gegenbaur (5). 

The olfactory capsules are dorso-ventrally flattened and project 
laterally from the front of the cranium. They do not come 
into apposition with one another in the median longitudinal line. 
Hach cartilaginous capsule is completely closed except for the 
small nostril, which is situated antero-ventrally and a short 
distance from the distal end. 


Text-figure 2. 


Sap’ 
: 
‘ 


c 


p3 \ ; Rane 
pad. ea ) 


Dorsal View of Skull. 


a,f. anterior fontanelle. o.c. olfactory capsule. 

a.p. auditory process. p.a.d. pre-auditory depression. 

d.r. dorsal bars of rostrum. pf. posterior fontanelle. 

nf. nerve foramen. p.o. post-olfactory process. 
o. orbit. v.r. ventral bar of rostrum. 


The cranium is slightly dome-shaped dorsally, and the auditory 
capsules are attached to it postero-laterally. Between the two 
auditory capsules, and on the dorsal surface of the cranium is a 
shallow median depression, the posterior fontanelle (p.f.). This 
fontanelle contains four apertures, two on each side, which will 
be referred to later in the description of the membranous 
labyrinth. 

Hach auditory capsule exhibits on its dorsal surface a supra- 
auditory process (sa.p.) which extends upwards from the socket, 
which is found on the side of the capsule, for the articulation of 
the hyomandibular. About half an inch mesial to each supra- 
auditory process is a small foramen for the passage of a nerve. 


974 MR. J. H. LLOYD AND MISS EDITH SHEPPARD ON 


The ventral view of the skull shows that a groove, about one 
inch in length, runs from the distal end of the olfactory capsule 
to the nostril. The end of the capsule also serves as a further 
support to the anterior of the orbit. Gegenbaur (5) has incor- 
rectly figured the extent of the cavity of the olfactory capsule 
and the position of the nostril. 

A very slender bar (s.c.) of fibrous tissue, which is perforated 
by a fine canal, runs from the auditory process to the olfactory 
capsule, and then runs along the posterior edge of the capsule. 
The structure of this bar suggests that it is a sensory canal. It 
is figured as far as it could be traced in our specimen, but we do 
not believe that it is complete. It has been figured, for about a 


Text-figure 3. 


Ventral View of Skull. 
ep.y. ethmo-palatine groove. 
f. foramina. 
n. nostril. 
ng. nasal groove. | anterior to the nerve indicates the 
s.c. sensory canal. extent of the olfactory sac. 
s.o. 1st spino-occipital foramen. | 


p. process on cranium for articulation 
of ethmopalatine process. 
I. olfactory nerve. The dotted area 


third of the length shown in our drawing, by Gegenbaur (5), and 
he has lettered it (x), but does not explain it in any way. . 

The ventral side of the cranium (text-fig. 3) is marked by a 
deep ethmopalatine groove (ep.g.) into which fits the narrowed 
middle portion of the upper jaw. On each side of this narrowed 
portion of the jaw is an ethmopalatine process. Hach of these fits 
against a corresponding process (p.) on the cranium, one being 
situated immediately anterior to each of the two outer edges of 
the ethmopalatine groove. The processes on the cranium are 
perforated on the mesial side by a single large foramen, which 
opens dorsally into the pre-auditory depression In addition in 
this regior tkere are four smaller foramina on each side. 


HE ANATOMY OF A HAMMERHEAD SHARK. 975 


Another pair of large foramina, one on each side, is seen 
perforating the inferior pre-auditory ridge (i.p.r.), which runs 
anteriorly from the auditory capsule. 

Two small apertures (s.o.) ventral to the foramen magnum 
serve for the passage of the first spino-occipital nerve. 

The posterior view of the skull shows the apertures for the 
ninth and tenth cranial nerves in the normal position; the 
tairly large foramen of the vagus nerve (x) at the sides of 
the occipital processes, and the small foramen for the glosso- 
pharyngeal nerve (1x) at the postero-lateral corner of the auditory 
capsule. 

The posterior portion of the skull seen in lateral view (text- 
fig. 4) shows a number of foramina. The most posterior of these 
is that of the vagus nerve (x), and slightly anterior to this is the 
foramen of the glossopharyngeal nerve (1X). 


Text-figure 4. 
7 SP pf : 
W. ee ue aa VAM 
S Sd. /p. 


Lateral View of Posterior portion of Cranium. 


a.s. articular facet for hyomandibular 
cartilage. 


p. articulating process on ventral side 
of cranium. Seeker inserted to 
show large foramen. 

sap. Supra-auditory process. 
h.a. passage for hyoidean artery. s.p.r. superior pre-auditory ridge. 
The posterior seeker marks the 
foramen perforating the inferior 
i.p.r. inferior pre-auditory ridge. | ILX. C pre-auditory ridge. 
—X. Cranial nerves. 


b.v. passage for blood-vessel. 


i.c. interorbital canal. 


The pre-auditory depression exhibits the following apertures :— 
‘the largest, which is situated near the posterior of the depression, 
is the foramen of the fifth, sixth, and seventh cranial nerves 
(Vv, VI, VII), excepting the ophthalmic branches of the fifth and 
seventh nerves. Ventral to this are three small apertures, the 
anterior of which serves for the passage of the hyoidean artery 
(i.a.). The posterior of the three is the interorbital canal 
(i.c.). The third aperture, which is situated between and slightly 
dorsal to the others, is the foramen of the third cranial nerve (141). 

_ The large optic foramen (11) is situated immediately anterior to 
the foramen of the chief branches of the fifth and seventh and 


976 MR. J. H. LLOYD AND MISS EDITH SHEPPARD ON 


the sixth nerves, and the single foramen of the ophthalmic 
branches of the fifth and seventh nerves (v & vit, op.) is immediately 
dorsal to it. 

In front of the ophthalmic foramen is a small aperture 
which serves for the exit of the pathetic nerve (iv), and ventral 
and slightly anterior to this is a small opening for the passage of 
a blood-vessel (6.v.). 

The pre-auditory depression is bounded on its dorsal and ventral 
edges by well-marked superior and inferior pre-auditary ridges. 
The inferior ridge (7.p.r.) is the better developed and runs forward 
from the auditory capsule to the olfactory capsule. The superior 
ridge (s.p.7.) becomes less marked anteriorly and disappears before 
reaching the olfactory capsule. 


MermpBraAnous LABYRINTH. 


As has been stated above, the posterior fontanelle of the cranium 
contains four apertures, two on each side of the median longi- 
tudinal line. ‘The anterior and smaller member of each pair 
serves to transmit the ductus endolymphaticus, whilst the larger 
posterior aperture leads into the perilymph cavity surrounding 
the posterior vertical canal. The posterior aperture apparently 
corresponds to the tympanic aperture described by Howes (11) in 
Raia, and also mentioned by Goodey (6) as occurring in Chlamy- . 
doselachus anguineus. i 

Unfortunately the superficial portion of the ductus endolym- 
phaticus was destroyed during the preparation of the skull, and 
we are therefore unable to give any account of its course outside the 
cranium. Judging, however, from the shallowness of the posterior 
fontanelle, and the fact that the skin was closely attached to the 
cranium in this region, it most probably runs straight to the 
dorsal surface of the head. 

The membranous labyrinth (text-figs.5 & 6) of the right side 
of the head has been dissected out, by removing the surrounding 
cartilage, and serves as the basis of the following description. 

In this account we have adopted the nomenclature used by 
Stewart (15) in preference to that used by Retzius (14). 

The ductus endolymphaticus (d.e.) rans downwards and bifur- 
cates at its lower end; a main branch going to the sacculus (s.) 
and a smaller branch opening into the recessus wtriculd (1.u.). 
Our observations on this point do not agree with those of Ayers 
(1), who states :—‘“‘T have not seen the bifurcate endolymphatic 
duct, described by Hasse for the species which he studied, but the 
two endolymphatic tubes observed by E. H. Weber I have found 
beautifully developed in Sphyrna zygena, and there is not the 
slightest doubt as to their occurrence.” 

We have been unable to trace Hasse’s statement in his paper (8) 
or to ascertain what species he studied. There is, however, no 
doubt whatever that the endolymphatic duct does bifurcate at its 
lower end. The second duct present is not an endclymphatic 


THE ANATOMY OF A HAMMERHEAD SHARE. 977 


duct but leads into the posterior perilymph cavity. This obser- 
vation agrees with that of Goodey (6) on the posterior duct in 
Chlam ydoselachus anguineus. 

The Utriculus, as in nearly all Hiner opranenet is divided into 
two portions, anterior and posterior, which do not communicate 
directly with one another, but indirectly through the sacculus. 

The anterior utricle (w.@.) is circular in transverse section and 
gives off the anterior canal (c¢.a.) at its dorsal end. The canal 
runs forward ana slightly outward, and expands at its lower end 
into the anterior ampulla (a.a.). The anterior utricle and the 
anterior canal together form an oval. 

The anterior utricle also gives off posteriorly, at about the 
middle of its length, the external canal (c.e.), which runs at first 
downward and outward, and secondly horizontally. At its 
anterior end it expands into the external ampulla (a.e.), which 
communicates again with the anterior utricle through a short 
canal running dorsal to the recessus utriculi, but having no direct 
connection with it. 

On the ventral side of the anterior utricle is a sac, the recessus 
utricult (7.u.), which is somewhat flattened from side to side, and 
almost triangular in lateral view. It communicates with the 
anterior utricle through a small aperture, whilst another small 
aperture, situated near the opening into the utricle, opens into the 
sacculus. 

The posterior utricle (u.p.) is the portion of the labyrinth 
situated nearest to the median longitudinal line of the cranium. 
It has direct communication with the sacculus through a short 
duct, the ductus utriculo saccularis posterior (d.u.s.p.). The dorsal 
end of the posterior utricle gives off the posterior canal (c.p.), 
which runs downward and outward and after expanding into the 
posterior ampulla (a.p.) opens into the ventral end of the utricle. 

The sacculus (s.) and lagena (J.) are laterally compressed ; the 
latter being given off as a simple pouch at the infero- posterior 
end of the sacculus. ‘The outer face of the sacculus and lagena is 
directed outward at an angle to the median plane. 


NERVE SUPPLY, 


The membranous labyrinth is supplied by branches of the 
eighth cranial nerve. 

The main nerve gives off a branch which runs beneath the 
recessus utriculi and then bends upward. It then bifureates to 
form the ramulus ampulla anterior (r.a.a.) and the ramulus 
ampulia externus (r.a.e.). A longerand finer branch, the ramulus 
ampulla posterior (r.a.p.), comes off from the main nerve and runs 
on the inner side of the sacculus to the posterior ampulla, giving 
off tiny ramuli to the sacculus and lagena en route. We were 
unable to trace the ramulus utriculi in our specimen, and it is 
probably very small. 

The membranous labyrinth resembles in many respects that of 
Cestracion philippt. 


\ 


978 MR. J. H. LLOYD AND MISS EDITH SHEPPARD ON 


Text-figures 5 & 6. 


ert! 


GA eel SU AUI ES 


Right Membranous Labyrinth. (Outer view.) 


Right Membranous Labyrinth. (Inner view.) 


a.a. ampulla anterior. l. lagena. 
a.e. ampulla externus. p.d. perilymph duct. 
a.p. ampulla posterior. r.a.a. ramulus ampulla anterior. 
c.a. canalis anterior. r.a.é. ramulus ampulla externus. 
c.e. canalis externus. r.a.p. ramulus ampulla posterior. 
c.». canalis posterior. ru. recessus utriculi. 
d.e. ductus endolymphaticus. uw.a. utriculus anterior. 

d,u.s.p. ductus utriculo  saccularis u.p. wtriculus posterior. 


posterior. VIIL. 8th cranial nerve. 


THE ANATOMY OF A HAMMERHEAD SHARK. 979 


BRAIN AND CRANIAL NERVES. 


Busch (2) and Miklucho-Maclay (12) have briefly, and on the 
whole accurately, described and figured the macroscopic structure 
of the brain. Owing to the poor condition of our specimen we 
are not able to add much to their account, but there are a few 
points on which our observations do not agree with theirs, 

The brain is antero-posteriorly shortened and, in dorsal view, 
only a portion of the olfactory lobes (0.1.), the prosencephalon ( f.b.), 
the much convoluted cerebellum (c.), the restiform bodies (7.b.), and 
the short medulla oblongata (m.) are visible (text-fig.7). The 
diencephalon and optic lobes are completely covered by the 
cerebellum. 

The prosencephalon is single and shows no trace of a median 
fissure. 

The large paired olfactory lobes are situated ventral to the 
prosencephalon ; only their anterior ends being visible in dorsal 
view. 

Owing to the intimate relation of the olfactory lobes and the 
prosencephalon Busch (2) has named them the “‘ lobes communes,” 
and a section through the forebrain appears to justify this name. 
On the other hand, in his diagram he shows the prosencephalon 
as a paired structure and the olfactory lobes as unpaired. This 
is opposed to our observations. 

The cerebellum is roughly triangular in shape, the base of the 
triangle being closely apposed to the posterior of the prosen- 
cephalon, and its apex posterior. It is primarily divided into 
three segments ; two of these being anterior, right and left, and 
the third running obliquely backward from the right. The left 
anterior segment is approximately equal in size to the posterior 
segment, but the right anterior segment is considerably smaller. 
All three segments are further divided into smaller segments, 
which in turn are sulcated. 

Ventral to the posterior end of the cerebellum are two small 
oval lobes called by Miklucho-Maclay (12) the lobé trigemini (1t.). 

The restiform bodies (r.b.) are well developed and occupy their 
normal position. The medulla oblongata (m.) is very short, and 
the fourth ventricle is correspondingly reduced. 

In ventral view the olfactory lobes completely cover the 
prosencephalon. The optic chiasma lies immediately posterior to 
the olfactory lobes. 

The infundibulum is well developed, and the saccus vasculosus 
shows on each side. Portions of the optic lobes are also visible in 
this view. 

An examination of a median longitudinal section confirms 
Miklucho-Maclay’s observation that a wedge-shaped portion of the 
cerebellum projects into the ventricle of the “ Zwischenhirn.” 

The following Notes on the cranial nerves were made chiefly 
during the preparation of the skull, and are consequently far from 
complete. 

The olfactory nerve is large and is given off laterally from the 


980 MR. J. H. LLOYD AND MISS EDITH SHEPPARD ON 


olfactory lobe. It runs attached to the posterior border of the 
laterally elongated olfactory sac, which is placed in the laterally 
expanded olfactory capsule. ‘The portion of the nerve adherent 
to the sac is called the olfactory bulb by Busch (2), whose figure 
shows it expanded rather more than is the case in our specimen, 
To verify Busch’s statement we have sectioned a portion of the 
nerve taken from the region of thesac, but we have failed to find 
any trace of nerve cells. ‘This, however, may be accounted for 
by the fact that our material was not in the best state of pre- 
servation. 


Text-figure 7. 


Dorsal View of Brain. 
ec. cerebellum. o.l. olfactory lobes. 
fb. prosencephalon. 7.6. vestiform bodies. 
1.¢. lobi trigemini. v. 4th ventricle. 
m. medulla oblongata. J, IJ, V. cranial nerves. 


The optic nerve runs on the ventral side of the hammer just 
anterior to the auditory process. 

The oculomotor nerve runs for the greater portion of its length 
posterior to the optic, but crosses it ventrally about an inch from 
the orbit and is distributed to the inferior oblique, anterior 
rectus, and inferior rectus muscles. A distinct branch of the 
nerve runs parallel to the main branch and supplies the superior 
rectus muscle. 

The pathetic nerve runs antero-dorsal to the optic and supplies 
the superior oblique muscle. 

The ophthalmic branches of the fifth and seventh nerves run 
directly to the anterior end of the dorsal surface of the snout. 
The fifth, in addition, sends off a lateral branch to a patch of 
ampullary canals just mesial of the orbit. 

The mandibular and maxillary branches of the trigeminal have 
the normal distribution. 

The abducens runs ventral to the auditory process and supplies 
the posterior rectus muscle. 


THE ANATOMY OF A HAMMERHEAD SHARK. 981 


The hyomandibular branch of the seventh nerve runs posterior 
to the mandibular branch of the fifth and gives off an external 
mandibular. 

The auditory nerve has already been described in relation to 
the membranous labyrinth. 


SUMMARY, 


The following are the important points dealt with in the 
paper :— 

1. The skull, except the jaws, has been described in detail. 

2. The membranous labyrinth has been described and figured 
for the first time. 

3. The existing accounts of the macroscopic structure of the 
brain have been revised, and notes have been included on the 
relationships of most of the cranial nerves. 


LITERATURE. 


1. Avers.—‘‘ Contribution to the Morphology of the Vertebrate 
Kar.” J. Morph. vi. 1892. 
2. Buscu.—De Selachiorum et Ganoideorum  Encephalo. 
Berolini, 1848. 
3. Day.—The Fishes of India. 1878. 
4, Garman.—‘ The Plagiostoma.’ Cambridge, Mass., Mem. 
Mus. Comp. Zool. Harvard Coll. 36, 1913. 
5. GrGensaur.—— Unters. z. vergl. Anat. d, Wirbelthiere, ii. 
1872. 
6. Goopry.—“ A Contribution to the Skeletal Anatomy of the 
Frilled Shark, Chlamydoselachus anguineus Gar.” Proc. 
Zool. Soc. 1910. 
7. Gooprico.—A Treatise on Zoology. Pt. 9, Ist Fase. 
Cyclostomes and Fishes. London, 1909. 
8. Hasse.—‘‘ Bemerkungen tber das Gehodrorgan der Fische.” 
Verhandl. der Phys. Med. Gesell. in Wurzb., N. F. i. 1868. 
9. HasweLut.—‘‘ Studies on the Klasmobranch Skeleton.” Proc. 
Linn. Soc. N. 8. Wales, vol. ix., 1884. 
10. Hemprich & HEurenserc.—Symbole Physice. Zoologica. 
Berlin, 1899. 
11. Howrs.—‘“ On the presence of a Tympanum in the Genus 
Raia.” Journ. Anat. & Physiol. vol. xvii., 1883. 
12. Mrxiucno-Macriay.—Beitrige z. verg]. Neur. d. Wirbelthiere. 
I. Das Gehirn. d. Selachier. Leipzig, 1870. 
13. Parxer.—‘‘ Notes on Carcharodon rondeletii.”. Proc. Zool. 
Soc. 1887. 
14. Rerzius.—Das Gehorogan d. Wirbelthiere. Bad. I. Stock- 
holm, 1881. 
15. Srewart.—‘‘On the Membranous Labyrinth of Certain 
Sharks.” Journ. Linn. Soc., Zool. vol. xxix., 1906. 
Proc. Zoou. Soc.— 1922, No. LX VI. 66 


alg 

ia terg) 

sapere 
alas iil ‘ 


tae 
Baier 
een 
iret 


Hi 
zi 


6 


P. Z.S.1922, KIRKPATRICK & METZELAAR, PI. I. 


COMMENSALISM BETWEEN A HERMIT CRAB & A POLYZOON. 


evan p 
if ahs Ais, 
ue 
: HN 


P. Z. S. 1922, KIRKPATRICK & METZELAAR, PI. II. 


14 


COMMENSALISM BETWEEN A HERMIT CRAB & A POLYZOON. 


ON AN INSTANCE OF COMMENSALISM. 983 


46. On an Instance of Commensalism between a Hermit 
Crab and a Poiyzoon. By R. Kirkpatrick, F.Z.8., 
and Dr. J. MarzEeLaar. 

[Received August 4, 1922: Read October 24, 1922. ] 
(Plates I., I.*) 


General.—In the year 1906, Mr. F. P. Vermeulen, Ymuiden, 
Holland, started with a fishery expedition to the West African 
shores with the object of seeing if the “‘langusts” of Cape Blanco 
and its neighbourhood would be worth exploitation. The rich 
material collected by him was given to the Zoological Society, 
“ Natura Artis Magistra,” at Amsterdam. It contains many 
Hermit Crabs, the commonest of which is Petrochirus granuli- 
manus Miers. 

The younger specimens of this Crustacean live in small shells 
of Turritella brevialis, Natica fulminea, Nassa miga, Terebra 
senegalensis, Aporrhais pes pelecani, and Dorsanum politum. These 
samples come from Cape Blanco and the large Greyhound Bay 
(‘Baie du Levrier”) close by, from depths not exceeding 
20 metres. A few specimens were also met with on the Senegal 
coast. 

Now, some of these “‘youngster” home-shells show a thin 
incrustation of a Polyzoon, a few layers thick near the orifice 
and only one or two layers over the rest of the surface (PI. I. fig. 3). 

If we compare the older young and adult pagurids with the 
former, we notice the said colony has increased very much, 
resulting in a heavy turnip- or potato-like spheroidal mass, 
about 6 cm. in diameter (fig. 1), completely involving the original 
contour of the shell. At one side there is a funnel, about 2 cm. 
wide externally and filled in by the heavy claw of the Petro- 
chirus. A section (fig. 2) shows the gasteropod shell covered 
entirely by a dense, stony crust of the Polyzoon in numerous 
layers, amounting to 56. The shell-substance does not appear 
to have been eaten away as in shells encrusted by Lepralia edax. 
The free margin of the orifice of the larger specimens is made 
up wholly of Polyzoan layers folded on themselves (fig. 2). On 
dissection, the calcareous Polyzoan mass appears to be homo- 
geneous, excepting that a few barnacles have been overwhelmed 
and suffocated. ‘The most sound colonies have a perfectly smooth 
surface of a deep violet-brown colour in spirit, and they form a 
typical feature of the fauna of the Cape Blanco coast. But for 
reasons not yet explained the colony may lose its power of 
resistance against destructive forces of the environment. Every- 
one knows the difference between “living” mollusc-shells, so 
eagerly wished for by conchologists, and the porous, brittle shells, 
the legal inhabitants of which have been replaced by boring 


* Wor explanation of the Plates, see p. 990. 


66* 


984 MR. R. KIRKPATRICK AND DR. J. METZELAAR 


animals. Exactly the same is the case with our Polyzoon. The 
simple colonies from one station have all changed into microcosms 
of sessile invertebrate life. Large specimens of Lithodomus 
easily lodge in the thick, limestone-like wall without touching 
their motor-pagurid. ‘There are the big colonies of a compound 
ascidian (Distomus sp.), purple barnacles, bushy hydroids upon the 
Polyzoan wall (PI. II., fig. 14). Although very interesting from 
an cecological point of view, we shall not describe these compound 
colonies as they do not show constant features, the original 
Paguro-Polyzoan association having lost its exclusive character. 
Meanwhile it illustrates the fact that mobility of substratum is 
favourable to diverse forms of sessile animal life. 

By the way, we may notice one rather striking fact. Although 
the weight of its limestone house with weeded roof mast become 
a nuisance to the Hermit, however powerful it may be, let us 
mention one point in favour of its security. In one of the 
hydroid tufts a spawning Sepia has glued several eggs. So 
this professional crab-eater remained unconscious of the close 
proximity of its prey, and the most critical seeptic must admit 
that this argument, taken from nature itself, is stronger than any 
aquarium experiment as to the efficiency of this pagurine mode 
of life. 

As a counterpart to this, we may add that remains of Hupa- 
gurus bernhardus are often found in the stomachs of Cod trom 
moderate depths in the North Sea. 


Description of the Polyzoon, based upon 43 adult and 3 young 
samples sent to the Natural History Museum, London. 

A glance with a good lens at once shows the Polyzoon to be a 
member of the great Membraniporidan group. The surface of 
the zoarium is covered with a membrane, which can be peeled 
off. The zocecia are arranged in straight, longitudinal rows, the 
latter every now and then bifureating. The zoccia are not 
abreast transversely, but in quincunx. A five of diamonds in 
cards, with the pips uniformly enlarged till they nearly meet, 
would illustrate the plan. 

The zoocia (Pl. I. figs. 4, 5, 6) are oval, and on an average 
0-6 mm. long and 0°3 mm. wide in the middle. The margins of the 
zocecia, in well-preserved material are mapped out by dark brown 
membranous or chitinous lines. Busk calls attention to similar 
“brown lines” in his description of Membranipora denticulata 
Busk. 

The calcareous margin is strongly granulated, and slopes 
inwards and downwards to a finely serrated edge bordering the 
oval opesia. 

At the proximal or basal end of each zocecium and immediately 
below the opesia there are two triangular, flattened, hollow 
tubercles, separate in the young stage, but joined into a single 
rectangular block in older zoecia (figs. 4, 5, 6). The brown line 
separately surrounds each triangular tubercle and also each 
rectangular block. 


ON AN INSTANCE OF COMMENSALISM. 985 


The ectocyst in the surface layer of zowcia is membranous. 
The operculum is 0°06 mm. long and 0:08 mm. wide, and with 
the rim thickened, especially laterally. In the older layers of 
zoecia the opesia is filled in, not by membrane, but by a thick 
punctate, calcareous plate, convex on its lower surface, and 
further, the zocecial walls become thickened, so that the zoarium 
becomes hard and stony. MRosette-plates multiporal; usually 
four in each lateral wall; two, occasionally several, in the 
distal wall. 

There are no avicularia and no ocecia. 

There need be no hesitation in assigning the Polyzoon to the 
genus Conopewm Gray* (genotype: Jlacroiwii Audouin fT) as 
amended by Norman and Canu & Bassler §. 

But the determination of the species has been a more difficult 
problem. One high authority, to whom the material had been 
sent, identified the species as Membrunipora tehuelcha VOrbigny. 
We were at first inclined to regard the species as a variety of 
Conopewm lacroiaii Audouin, but finally we have come to the 
conclusion that the species is new to science. We propose to 
call it CoNOPEUM COMMENSALE, sp. n. 

The distinguishing characters of the new species are: the thick 
zocecial walls, the thick calcareous opesial plate, and the dense 
stony multilaminate zoarium. Neither the well-known British 
Con. lacroixii, as described by Busk and Hincks, nor the 
Mediterranean form, as described and figured by Audouin and 
Savigny, have any of these characters. If in the British form 
of C. lacroixit occasionally one layer grows over another, there 
are no calcareous opesial in-fillings; and, moreover, ‘the tri- 
angular hollows on each side above the aperture” (Hincks, Brit. 
Mar. Pol. p. 130) have each an opening, covered by membrane, 
and do not fuse into a single rectangular block. This fusion 
does, however, take place occas ionally in the typical form, figured 
in the ‘ Description de Egypte,’ pl. x. fig. 9. The new species 
differs widely from JM. tehwelcha a Orbigny. We had the good 
fortune to find numerous specimens encrusting the alga Macro- 
cystis pyrifera from the Straits of Magellan and from Valparaiso 
and the coasts of the Tehuelchan region (the T. being a great 
tribe of Patagonian Indians). The zoecia (fig. 12) are very 
different in shape and character, being thin-walled, elongated, 
rectangular boxes with sharp edges; and the triangular knob 
and fused rectangular structures of C. commensale Kirkpatrick 
& Metzelaar are replaced by a pair of rather long, blunt 
caleareo-chitinous “spines.” J. fehuelcha, in our opinion, is little 


* Gray, List of British Animals.—Part I. Radiated Animals, pp. 108, 146 (1848). 

+ It is held by several authorities that Audouin’s “ Lacroixii” is a synonym of 
Millepora reticulum Linné. 
it Norman, Nat. Hist. East Finmark. Ann. & Mag. Nat. Hist. (7) xi. p. 586 
(1903). 

§ Canu & Bassler, North American Harly Tertiary Bryozoa. U.S. Nat. Mus 
Bull. 106, text pp. 84, 86 (1920). 


986 MR. R. KIRKPATRICK AND DR. J. METZELAAR 


other than a South American representative of Membranipora 
membranacea Li. 

The study of the triangular objects in ©. lacroixii Audouin 
and in the early stages of C. commensale Kirkpatrick & 
Metzelaar has, we think, thrown an interesting light on certain 
structures found in widely separate families and genera. The 
precise nature of the “ triangular hollows” * in CO. lacroixii is not 
definitely known, but the view that they are diminutive zoccia, 
aborted owing to exigencies of space, appears to us a reasonable 
one. Cramped zocecia are often found among the rows and layers 
of ordinary zocecia. The openings of the inter-zoccial triangular 
hollows of O. lacroixii have granulated margins, precisely com- 
parable with the edges of ordinary zoccia, and under a high 
power the whole structure is fairly similar in aspect to an 
ordinary zocecial box somewhat distorted. 

In C. commensale the triangles have a calcareous roof, and are 
bounded each by'a separate brown line. Where the triangles 
are fused into one rectangular block, there is only one line round 
the block. A fundamental and—with the exception of Loao- 
soma—universal character of the Polyzoa is that of budding 
so as to form colonies. Limitation of space must lead to a 
struggle among the buds, with partial or complete suppression of 
some. 

It would appear that in ©. lacroixti each zocecium gives off 
three distal buds, the lateral ones being almost, but not wholly, 
suppressed. 

A careful examination of many species of Cheilostomes has 
convinced us that the paired spaces, knobs, tubercles, and spines 
so commonly present are all comparable with the “ triangular 
hollows” of C. lacroixii, and that they are modified zoecial 
buds f taking on strange shapes and functions. In Membranipora 
membranacea L., for instance, triangular spaces will be found at 
or near the growing edges of the zoarium; in older cells the 
“hollows” (really inter-zocecial boxes) grow up into long spines 
or tubercles, as if, owing to the limited basal area, they took to 
building skyscrapers. 

In WM. tehuelcha dOrbigny (fig. 12) the same event has 
happened. Here the tops of the spines are often membranous, 
recalling the membranes over the openings of the triangles in 
C. lacroixti. That this interpretation is not a mere fancy, is 
shown by the resemblance of the early stages of the spines 
near the growing edges of colonies to the triangular areas in 
C. lacroixii. 

In WM. tuberculata Bose, so abundant on Gulf weed, the 


* Hincks, ‘ British Marine Polyzoa,’ p. 180. 

+ Of. the “‘origelle” of Jullien, “des bourgeons charnus, développés sur l’endo- 
cyste,’ ‘which can reproduce zocecia, avicularia, etc. (Bull. Soc. Zool. France, xi. p. 607, 
1886). Later, “ joncturies,’ which produce zoccia, are distinguished from “ origelles 
évolutives,’ which form avicularia, spines, etc. (Jullien & Calvet, Bryozoaires 
‘ Hirondelle,’ 1903, p. 18). 


ON AN INSTANCE OF COMMENSALISM.. 987 


tubercles, at first separate, may join together to form single 
blocks as in C. commensale ; a remarkable example of the former 
Species is shown in fig. 11, where the blocks have developed so 
enormously as almost to obliterate the opesia. 

In descriptions, the triangular hollows, spines, tubercles, etc. 
are usually associated with the distal or oral end of the zocecium, 
but they should perhaps rather be associated with the proximal 
or basal part of a zoeecium. On the triple-bud hypothesis it is 
certainly more rational to associate the three buds, no matter 
what their subsequent history and disguise might be ; and further, 
the knobs, spines, etc. are often clearly seated on the front of 
the basal end of a zoccium. ‘Take, for example, the little knobs 
on the zoecia of Membranipora savartii Audouin, shown on the 
base of each cell in Savigny’s figure (‘ Description de |’ Egypte,’ 
Atlas plex ties 10) *. 

In Vhairopora armata MacGillivray (fig. 13) the two club- 
shaped tubercles are situated one on each side of the oral orifice 
of a zocecium, and, it must be admitted, a little imagination is 
required to realize that even here the tubercles should perhaps 
be associated with the proximal end of the following zocecium. 
One might assume that the two lateral buds had been given off 
at right angles to the central one, instead of at a forward-opening 
acute angle. 

The avicularian cell in this species appears beautifully to 
confirm the theory of homology of tubercles and triangular 
boxes; for here, in place of tubercles, and on each side of and 
above the avicularian opening, there is a triangular box with a 
clear space in the roof (fig. 13). 

Possibly the quineuncial plan of growth so common in Cheilo- 
stome Polyzoa may be due to adaptation to limited space. 
Zocecia giving off three buds at the distal end would not be well 
able to grow abreast; accordingly the buds (and their adult 
equivalents) often alternate transversely. 

The fusion of two lateral hollow knobs into one rectangular 
hollow box in C. commensale may be connected in some way with 
reproduction, for in decalcified specimens there can usually be 
seen a brown body or developing new polypide in the zoecial 
cavities adjacent to these boxes. 


Discussion.—On two points our description of the new species 
is open to adverse criticism. Firstly, we attribute specific value 
to zoarial characters. During an early period of the study of 
Polyzoa, zoarial characters alone were considered, most of the 
encrusting and branching calcareous Cheilostomes being relegated 
to Lepralia and Hschara respectively. Later, zocecial characters 
became all-important, the zoarial ones being entirely put aside. 


At the present time the tendency is to take all characters into 


* There is rarely any trace of the knobs in the specimens of this species from 
tropical seas or from Japan. 


988 MR. R. KIRKPATRICK AND DR. J. METZELAAR 


consideration, judging each case on its own merits. A good 
example of the specific value of zoarial characters is afforded by 
Lepratia bifurcata Waters, found in deep water off Capri. Here 
we have a typical Lepralia, but one that always grows in a very 
definite, peculiar, and characteristic way, viz. as a bifureating 
colony with two little wings. No matter how closely the 
characters of the individual zocecia resemble those of some par- 
ticular encrusting Lepralia, it would be difficult to regard the 
bifurcating colony as a mere variety or variation of the en- 
crusting one. Similarly, we regard the dense, stony, multi- 
laminate Polyzoon from Cape Blanco as specifically distinct from 
the slender, thinly encrusting C. lacroiwit, the modified character— 
if we may adopt the neo- -Lamarekian line of thought—having 
perhaps resulted from epecal, e. g. well-fed life. 

Biology.— Again, the adoption of the name ‘ commensale” 
might be considered a doubtful proceeding. We think the name, 
however, to be convenient, not only from the point of view of 
easy identification, but justifiable as indicating the real relation- 
ship existing between the Polyzoon and the Crustacean. There 
is evidently no question of true symbiosis or mutualism sensu 
strictiore, if we define that as ‘legal relation between two 
different organisms,” principally, because Petrochirus granuli- 
manus can live and thrive without C. commensale. Leaving the~ 
young Crustacea aside, we find the adults associated with the 
sponge Sauberites domuncula, the compound zoantharian Corticifera 
lutea, and the Polyzoon Lepralia edax. 

But the reverse does not seem to be the case. Polyzoa growing 
on the hydroids and corals of the Vermeulen-expedition did not 
include C. commensale among them. Nor did we notice it upon 
living molluses from the same localities. Possibly the Polyzoon 
could survive for a time on an vuntenanted shell with much 
diminished vitality, but so far there is no evidence on this point, 
although we must take into consideration that the West African 
material has been collected by one who was not a zoologist. 

According to Prof. Calvet, any smooth surface is suitable to the 
Membranipore, and “associations of hermit-crabs and Polyzoa 
are common enough.” Even so, we never saw them before in 
such a definite non-accidental mode and shape. The Paguro- 
Polyzoan association described here appears to be a definite and 
not an accidental one, the Crustacean and Polyzoon being more to 
each other than casual messmates; for it is certain they derive 
special advantages from each other’s presence. 

Hermit Crabs are widely distributed all over the world, and 
are notorious for their militant and aggressive nature. Hnsconced 
in their shells they are veritable tankers, spreading dismay 
among their helpless victims. Every now and again, when the 
soldier has to leave his fortress, his softened body is exposed to 
danger. But frequently some other organism (Sponge, Coelente- 
rate or, more rarely, Polyzoon) adapts itself to the shell, and 
gradually acquires certain special characters. The encrusting 


ON AN INSTANCE OF COMMENSALISM. 989 


animal forms an extra panoply, not merely thicker, but more 
extended, especially around the shell orifice, thereby generally 
ensuring a longer tenancy to the Causa This feature of 
extended growth 4 is illustrated in fig. 2, where the opening of the 
fortress is seen to be formed, not of shell, but of infolded layers 
of the Polyzoon. Again, the Polyzoon obviously benefits; for 
it is in alliance with a vigorous and successful marauder, and 
although sedentary by nature, is continually being carried to new 
and rich pastures. 

Accordingly, the large size of the new species is not surprising. 

We return now, after this economic excursion, to the pecu- 
harity of the association described. Against critics who deny 
the non-accidental nature of it. we draw attention to one main 
point. Setting apart those favourite Pagurine mates: Suberites 
and the epeecal Zoantharians, there remains the fact that a 
“special seat” is reserved to Conopewm commensale among the 
hosts of sessile epeecal animals, ready to populate every available 
spot in the tepid tropical coast-waters. In a very short lapse of 
time a typical complex is formed. This follows from the fact 
that among the many home-shells of adult Petr. granulimanus 
only one was found naked, obviously inhabited only a short time 
before it was caught; intermediate gradations to full-grown 
adult colonies are wanting. Now, what is the reason that 
among so many rivals the tiny Polyzoon regularly wins the 
battle? We have seen that the other candidates may win the 
second round if the Hermit migrates to a spot unfit for its 
Polyzoan comrade to live in. In the struggle for life the 
champion is slain at last. Wow at last a fair chance is given to 
everybody; here you have Accident playing its part, and the 
result ....a chaotic conglomerate of sessile marine life (fig. 14). 

Since. the time of Darwin, Law has taken the place of Acci- 
dent in biological science. 

Our case is in some respects comparable to the well-known 
association of Hupagurus bernhardus and Hydractinia echinata. 
In this case the Pagurid inhabits the whole “littoral” region 
up to a depth of 270 metres, but Hydractinia echinata is a 
coastal form which does net follow the adolescent Hupagurus 
to depths exceeding those of the centre of the North Sea. 
At the lower limit of its occurrence the Hydractinia is often 
replaced by Alcyonidium gelatinosum. Now, although Hydrac- 
tinia echinata has been found in a few instances without its 
mate Hupagurus bernhardus, is there anybody who will seriously 
contest the particular relation between these two organisms ¢ 
Cf. Hupag. pubescens, which occur in the North Sea as a 
rule, associated with Swberites jficus, but exceptionally with 
Zoanthus sp. 

So far as we know, the present case has not yet been described. 
It is a remarkable fact that Chevreux and Bouvier, among the 
rich material of the ‘ Melita,’ described 17 species of Pagurids 
(Mém. Soc. Zool. France, v. 1892), but only noticed ‘ coquille 


990 ON AN INSTANCE OF COMMENSALISM. 


recouverte de bryozoaires” in the case of Petrochirus granuli- 
manus. The ‘“ bryozoaires” have probably been Con. commen- 
sale, but they were not examined. 

In the volume dealing with the results of the inquiries of the 
‘Travailleur’ and ‘ Talisman’ immense numbers of Polyzoa are 
mentioned, but no reference to the present case. 


Summary and Conclusion.—The present Polyzoon from the 
Cape Blanco region is a new species, viz. Conopeum comimensale 
Kirkpatrick & Metzelaar. It is most nearly related to C. lacroixiw 
Audouin. 


The paired “triangular spaces,” paired knobs, tubercles, or 
spines so commonly present in many species of Cheilostome 
Polyzoa are probably aborted zocecial buds, more or less sup- 
pressed or modified owing to lack of space for free growth. 

The association with the Crustacean Petrochirus granulimanus 
is a definite, not an accidental one, being a case of commensalism 
in the sense of Van Beneden. 


EXPLANATION OF THE PLATES. 
(The figures to be examined through a hand-lens.) 


PratE I. 


Fig. 1. Gasteropod shell inhabited by Petrochirus granulimanus Miers, completely 
encrusted by the Polyzoon Conopeum commensale Kirkpatrick & Metze- 
laar from Cape Blanco. Nat. size. 

2. A shell and Polyzoan crust cut in half. Nat. size. 
8. Turritella shells inhabited by Hermit Crab, and each encrusted by young 
colony of Conopeum commensale: from Senegal. Nat. size. 
4. Conopeum commensale. X 20. 
5 & 6. The same from another specimen, showing at the first separate triangular 
tubercles and the single rectangular blocks. X 20. 
. An older layer of zocecia broken into and viewed from the inferior or ‘dorsal 
aspect, showing calcareous infillings of the opesia. X 20. 
. Conopeum lacroixii, showing the two separate triangular “ spaces” 
(boxes) at the base of each zocecium. X 20. 
9. Conopeum commensale. Transparent vertical section, showing layers. X 3%. 
(Cf. fig. 2.) 

10. Conopeum lacroivii Audouin, showing separate triangular hollows: speci- 
men from Dovercourt, England. > 20. 

11. Membranipora tuber ulate Bose, encrusting two sides of alga from Algoa 
Bay ; showing remarkable blocks at the margin of colony, also ordinary 
paired tubercles. X 20. 

12. Membranipora tehuelcha VOrbigny on Macrocystis pyrifera from South 
America ; showing the two spine-like tubercles to each zocecium. 

13. Thairopora armata MacGillivray, showing zocecia with paired boss-like 
tubercles and an avicularian cell with triangular hollow boxes above 
mandible. x 20. 

N.B.—There is an avicularium with sharply-defined triangular 
mandible about 8 mm. above the middle of the lower border of the 
picture. The mandible and the triangular boxes above it have a frog’s- 
face-like aspect. In the ordinary zoccia the pairs of white boss-like 
tubercles replace the triangular “ hollows” or boxes. A lens is necessary. 


bas § 


Prater I. 
Fig. 14. Bulbous specimen secondarily overgrown with hydroid, ete. 


PIZeS 1922 NA Ee einale 


NEW SPECIES OF TRINIDAD MOTHS 


VITTY & SEABORNE, LTD., 
LONDON. 


ON NEW SPECIES OF TRINIDAD MOTHS, 991 


47. New Species of Trinidad Moths. By W.J. Kays, F.E.S.* 
[Received July 30, 1922: Read November 7, 1922. | 
(Plate I.+) 


The moths here described and figured were all (except one, 
Centronia plorator, caught by myself) taken by Sir Norman Lamont 
in the south of the island of Trinidad. Most likely, all the species 
will some day also be found on the mainland in either Venezuela 
or Guiana, as,except for numerous local races, one would not 
expect distinct island species such as are found in Oceanic 
islands, Trinidad being essentially a recently detached portion of 
Venezuela. Doubtless, there are numbers of species still unknown 
and undescribed, and very much work remains to be done before 
we can even approximately guess the number of genera and 
species that exist. 


Family Hypsip&. 


CENTRONIA PLORATOR, sp. n. (PI. I. fig. 3.) 


@. Base of eyes and base of palpi orange. Antenne filiform. 
Abdomen black with the segments below ringed with white. 
Fore wing black with a postmedian curved white band com- 
mencing very narrow on the costa but rapidly widening to about 
3mm. wide as far as vein 4, where it rapidly contracts and 
becomes suffused with black to extreme angle of wings. Veins 
showing grey on inner side of white band. Hind wing dark 
bluish black with a narrow whitish marginal band, suffused with 
black as far as vein 5, where it suddenly ceases. Below, the 
white band of both fore and hind wing completely white without 
any suffusion. Exp. 44 mm. 

Habitat. Trinidad, Siparia, 18. xi.20 (W. J. Kaye). In Coll. 
B.M.. 


Family NoropoNnT1ipD&, 


DICENTRIA NONDESCRIPTA, Sp. n. (PI. I. fig. 9.) 


Head brown with a fan-shaped tuft of raised hair behind 
antenne. Fore wing dark chocolate-brown with greenish-yellow 
shades, especially on costa beyond middle. Discoidal spot not 
conspicuous, of the same ground-colour as wing and edged on 
both inner and outer edges with greenish yellow. An indistinct 
slender pale line consisting of lunules beyond middle. Sub- 
terminal line also of pale lunules. A very ill-defined double 
submedian line, becoming lost in the ground-colour soon after 
leaving the cell, Hind wing pearly transparent with the apex 


* Communicated by Guy A. K. Marswatt, C.M.G., D.Sc., F.Z.S, 
+ For explanation of the Plate, see p. 998. 


992 MR. W. J. KAYE ON 


darker. Inner margin also dark and a pale but distinct mark 
just below anal save, Fore wing below shining blackish. Hind 
wing below shining whitish with a small dark mark in the 
middle of costa. Exp. 54 mim. 

Habitat. Trinidad, Palmiste, 25. xii. 20 (VV. Lamont). 


BorizA KALODONTA, Sp. n. (PI. I. fig. 13.) 


Fore wing greenish grey mixed with slaty purplish. A con- 
spicuous, elliptical, pale tawny spot between veins 2 and 4 near 
margin. Basal area to end or beyond cell slaty grey with three 
parallel wavy lines. A short, slender, black dash. Inner margin 
pale greenish. Outer margin broadly paler than rest of win 
with the veins showing through conspicuously. A small black 
dot between veins 3, 4, and another between veins 6, 7. Hind 
wing white with the costa narrowly dark brown, divided at 
middle with white. Thorax olive-brown. Fore wing below 
white with the veins 5, 6, and 7 heavily smoky.. Costa imme- 
diately above pale tawny. Hind wing below pure white. Exp. 
40 mm. 


Habitat. Trinidad, Palmiste, 30. x. 11 (WV. Lamont). 


PHASTIA MARICOLOR, sp. n. (PI. I. fig. 12.) 


Fore wing pale sea-green. Discoidal spot purplish white. 
Subterminal series of large irregular spots, purple on the inside 
and white exteriorly. A short purplish basal streak. Beyond 
this is a faint ill-defined band, becoming very broad on inner 
margin, of a darker shade of green to the ground-colour. Beyond 
discoidal spot is also a poorly-defined narrow band. Thorax of 
same ground-colour as fore wing, Hind wing very pale grey, 
darker towards the outer margin. Under surface of both fore 
and hind wing yellowish white, the whole of the fore wing except 
the narrow inner margin dusted with blackish. Fore legs with 
the first joint of the ‘tarsus clothed with yellowish-green hair, 
Exp. 42 mm. 

Habitat. Trinidad (NV. Lamont). 


RIFARGIA BRUNNIPENNIS. (PI. I. fig. 11.) 


Fore wing dark burnt-umber with a purplish shade. with the 
basal half the darkest. A short dark basal streak. Costa near 
apex and inner margin except at base tawny. A conspicuous, 
little, white, wedge-shaped mark between veins 2, 3 close to 
margin. Veins 4, 5, and 6 edged with tawny. Hind wing 
brownish grey, darker towards outer margin and darkest at anal 
angle. A very inconspicuous postdiseal line, becoming duplicated 
on vein 3 to inner margin. Fore wing below dull blackish. 
Hind wing below as above. ilia black at termination of veins. 
Thorax tawny. Abdomen rather darker. Exp. 46 mm. 

Habitat. Trinidad, Palmiste, 25. xii. 20 (V. Lamont). 


NEW SPECIES OF TRINIDAD MOTHS. 993 


Family NoctruipD4. 
Subfamily HrastRiana&. 


BRYOLIMNIA MONODONTA, sp. n. (PI. I. fig. 20.) 


Third joint of palpi, frons, and base of antenne white. Thorax 
light brown with a pair of white tufts behind. Fore wing with 
the base chocolate-coloured, followed by a broad pure white band 
deeply toothed on outer edge and. with a small chocolate spot on 
costa. Postdiscal area chocolate, lighter towards inner margin 
and darkening towards costa... Cilia white, with a square dark 
spot about middle and with the tornus chocolate. Hind wing 
smoky greyish black, the apical area much the darkest. Dis- 
coidal spot indistinct, blackish. Abdomen greyish black. Exp. 
26 mm. 

Habitat. Trinidad, Siparia, 16. vi. 18 (WV. Lamont). 


BRrYOCODIA CRYPTOGRAMMA, sp. n. (PI. I. fig. 19.) 


Palpi with the first joint brown, the second and third whitish 
with dark brown marks. Fore wing dull olive greenish with 
pink and brown markings. A dark mark at base and a heavy 
brown basal line. A median fascia most prominent at costa and 
much broken at and beyond middle. A postmedian, thin, much- 
curved line, most prominent on veins 1, 2 and 4, 5. A dark 
eurved apical fascia and a dark brown spot at tornus. Hind 
wing coloured as fore wing, with double submedian brown lines 
and a deeply curved postmedian single line. Underside of fore 
wing below cream-coloured with dark median and postmedian 
lines; a broad outer marginal fascia dark brown. Hind wing 
below coloured as fore wing, except that in the region of the 
anal angle the dark brown fascia is broken up with the pale 
ground-colour. Exp. 28 mm. 

Habitat. Trinidad: Caparo, Oct. 1904 (Birch); Palmiste, 
27. x1. 15 (WV. Lamont). 


PARANGITIA NIGROFULGENS, sp.n. (PI. I. fig. 5.) 


Palpi, collar, and patagia yellowish brown. Abdomen fawn- 
coloured. Fore wing dark shining coppery black. <A double 
indistinct waved black line close to base. Apex lighter than 
rest of wing and followed by indistinct double black lines. 
A curiously placed black angular mark on vein 2 close to 
edge of cell. Hind wing black with the whole of the large inner 
area dark cupreous grey. Fore wing below dull black with the 
inner marginal area paler. Underside of hind wing rather dark 
slaty grey with a darker marginal band. Exp. 24 mm, 


Habitat. Trinidad (NV. Lamont). 


994 MR. W. J. KAYE ON 


PARANGITIA MICAPENNIS, sp.n. (PI. I. fig. 2.) 


Fore wing shining brownish olive. A large apical area, ex- 
tending along costa to above end of cell and to tornus, paler with 
several short dark dashes. Discal spot uniform pale with a short 
black dash at its lower extremity externally. Opposite and close 
to outer margin a small black spet. Above these marks in 
certain lights are to be seen three short, parallel, brown dashes. 
Two short wavy lines above one another emanating from the 
outer margin. Hind wing blackish brown with a broad dark 
marginal band and a small yellow elongated mark at anal angle. 
Thorax with long raised scales. Abdomen with a raised tuft on 
third segment. Exp. 30 mm. 

Habitat. Trinidad (N. Lamont). 


Subfamily Nocruin#. 


SANYS IMPLACATA, sp. n. (PI. I. fig. 17.) 


Head, thorax, and abdomen pale brownish white. Fore wing 
narrow with the apex and inner margin rounded, pale brownish, 
darkest on the costa. A small blackish dot at origin of vein 2. 
A straight brown line across basal area and a slightly oblique 
postmedian line. A wavy double submarginal line greatly angled 
near tornus and a series of dark dots on margin. Hind wing 
similarly coloured to fore wing. <A well-marked straight line 
across centre of wing, stopping at vein 6. <A wavy, partly 
double, subterminal line much broken from vein 5 to apex. 
Exp. 30 mm. 


Habitat. Trinidad (NV. Lamont). 


PLUSIODONTA CUPRISTRIA, sp. n. (PI. I. fig. 16.) 


Fore wing dark purplish. A large burnished-copper patch at 
tornus and a central stria of same colour from the cell to the 
“tooth ” of the inner margin. A pale oblique streak from apex 
to vein 2. At apex the streak is reflexed to costa. Thorax 
purplish. Two short subcostal streaks before apical streak. A 
small shade of copper-colour in middle of outer margin. Hind 
wing white with a broad dark margin, shading off internally with 
the white ground-colour. Underside of fore wing dark brown 
with the costa narrowly pinkish ochreous. Hind wing below 
white with only a small dark area at apex. Exp. 40 mm. 

Habitat. Trinidad, Palmiste, 8.1.21 (V. Lamont). 


Subfamily Derorpin”. 


EPItoMIPTERA PURPURASCENS, sp. nu. (PI. I. figs. 4, 23.) 


Fore wing purplish with a broad, somewhat darker, central 
fascia, greenish on inner side from inner margin for half the 
length of wing. A slender wavy pencil-line within the fascia a 
little beyond middle, Outer edge of fascia very sinuous and 


NEW SPECIES OF TRINIDAD MOTHS. 995 


a slight greenish flush near tornus. Extreme apex with a short 

dark mark followed by a dark greenish-purple patch. Close to 

base a small dark dot. Hind wing purplish brown with a central 

dot followed by a dark indistinct line. Hind wing beneath much 

paler, and, in addition to the dot and dark line, there is a broad 

marginal fascia enclosing a pale wing-line. Exp, 28 mm. 
Habitat. Trinidad. 


EPrroMIPTERA MARMORATA, sp. n. (PI. IL. fig, 6.) 


Fore wing various shades of pale brown with white lines. 
Extreme base pale followed by light brown fascia edged exteriorly 
with a sharply-defined white line sharply angled near inner 
margin. Central broad fascia darker than basal band, but also 
edged with a white line which is broken and very incomplete 
in central area but well-defined near costa and inner margin. 
Apex much mottled with white and brown followed by a wavy 
short white line. Hind wing pale light brown with a dark 
central oblique shade, followed by a short much-angled white 
mark, followed again by a slender white wavy line. Exp. 25 mm. 

Habitat. Trinidad, Guaico, 18.iv.15 (NV. Lamont). 


HYPENA BRODESCENS, sp. n. (PI. I. fig. 8.) 


Head, thorax, patagia, and metathorax very dark brown. 
Abdomen blackish brown with darker dorsal tufts of hair on the 
first four segments. Fore wing olivaceous dark brown with an 
oblique basal whitish line; a median broader straight white line. 
A postmedian series of yellowish spots on the veins and a very 
fine subterminal pale dotted line. Hind wing dull blackish with 
a pale bluish submarginal line most distinct between veins | and 3, 
Exp. 36 mm. 

Habitat. Trinidad; Caparo, Nov. 1904 (#. Birch); Palmiste, 
14. v. 21 (VY. Lamont) (type), 20. xii. 21 (WV. Lamont). 


Family LIMACODID4. 


SISYROSEA GUAICA, sp. n. (PI. I. fig. 15.) 

Head and collar light brick-red. Thorax fawn-coloured; 
abdomen fawn-coloured with a reddish dorsal shade. Fore wing 
pale pinkish buff. A small black spot at the base of the cell and 
another black spot close to outer margin lying on vein 4. Cilia 
shining buff-colour. Hind wing buff, the cilia of the same 
colour. Underside of fore wing buff, with some darker shading 
towards costa. Underside of hind wing buff. Exp. 34 mm. 

Habitat. Trinidad, Guaico, 18.ix.15 (N. Lamont). 


Family LASIOCAMPID2. 


HypRIAS ALBIOCHREA, sp. n. (PI. I. fig. 14.) 


Head and thorax grey with some ochreous hairs. Fore wing 


996 MR. W. J. KAYE ON 


rather pale brown with scattered grey scaling. At base a short 
black streak above which is an irregularly-shaped greyish patch. 
This on costa is followed by two short curved lines. Some very 
indistinct, slender, wavy lines across the discal area, with the 
veins showing darker. Hind wing with the costal third greyish 
white with two or three indications of transverse lines. Basal 
two-thirds of wing tawny, pale at outer margin. Abdomen 
tawny, with some black scaling on the three basal segments. 
Exp. 42 mm. 
Habitat. Trinidad, Port of Spain, 27.i1.21 (WV. Lamont). 


Family URANIAD A. 


SYNGRIA DELICATA, sp.n. (PI. I. fig. 7.) 


Collar whitish. Thorax and abdomen pinkish fawn-colour, 
irrorated with minute black dots. Fore wing pinkish fawn- 
colour. Discoidal spot small, black, lunar-shaped, externally 
edged with pale cream-colour. A submedian, very indistinct, 
orange line running very obliquely. A very distinct postmedian 
ochreous line, palish-edged internally, also very oblique and 
sharply angled just below vein 6. Margin sharply defined 
ochreous, edged with pale colour internally. Hind wing pinkish 
fawn with fine irroration of black dots. Very indistinct sub- 
and post-median ochreous lines. Margins as in fore wing. Fore 
and hind wing below without marking gs, except the fine black 
dusting, which is less than on upper sila, Exp. 36 mm. 

Habitat. Trinidad, Palmiste, June 1915 (N. Lamont). 


Family PyRALID A, 


Subfamily Pococerins& (= Lpipaschiane). 


PococERA BRUNNAPEX, sp. n. (PI. I. fig. 1.) 


Head and thorax reddish brown. Abdomen pale brown. 
Fore wing with basal third pinkish brown with some short 
darker longitudinal lines terminated by a somewhat sinuous 
silvery-grey line. Discal area of wing silvery grey, containing a 
much waved slender line in middle. A darker shade before 
outer margin and a dark reddish-brown apex with the veins 
showing through, heavily scaled with blackish brown. Hind 
wing pearly transparent with the apex blackish. Fore wing 
below with a dark red costa, and the remainder of the wing 
blackish. Hind wing below also with costa dark red, but other- 
wise as above. Exp. 38 mm. 

Habitat. Trinidad, Guaico, 18. ix. 15 (VV. Lamont). 


Subfamily Acrorprina (= Pyraustine). 
SAMEA DELICATA, sp. n. (PI. I. fig. 22.) 


Head pale straw-coloured. Patagia with dark brown spots on 


NEW SPECIES OF TRINIDAD MOTHS, 997 


frontal edge. Fore wing pale straw-coloured with two dark 
Spots on costa close to base and some further minute points as 
far as end of cell. A submedian, a median, and a postmedian 
slender line. Beyond the last is a dark marginal band. Dis- 
coidal spot formed of a double ring, the one above the other. 
Hind wing pale straw-colour with three transverse lines, the 
inner one straight but oblique. The middle line greatly angu- 
lated in middle, and the last line greatly angulated beyond 
middle. A very fine dark subterminal line. Exp. 16 mm. 
Habitat. Trinidad (N. Lamont). 


PHOSTRIA DUPLICATA, sp.n. (PI. I. fig. 21.) 


Head and thorax dull ochreous brown. -Fore wing ochreous 
brown. Basal line curved dark brown. Discoidal spot linear 
dark brown. Postmedian line heavy dark brown, greatly set- 
back between veins 2 and 3. Fine dark points at extremities of 
veins, and colour of wing beyond postmedian line darker some- 
what than remainder. Hind wing darker than fore wing and 
blacker. A small dark spot within the cell and a curved median 
dark line, broken and waved between veins 2 and 3. Abdomen 
with the segments edged with white. Fore wing below buff with 
black discoidal spot and dotted postmedian line. Hind wing 
below buff with point in cell and faint indication of median line. 
Exp. 26 mm. 

Habitat. Trinidad (N. Lamont). 


Family ZyYG@ BNID&, 


Genus LAMONTIA. 


Head small; antenne long, slender, bipectinated. Abdomen 
small and rather slender. Fore wing long and narrow; veins 
4, 5 very close together at origin; veins 8 and 9 stalked with 10, 
vein 1] weak, 12 strong. Hind wing with veins 4,5 from same 
origin ; vein 7 from a long way from 6. Outer margin of wing 
rounded. Vein 8 approaching very close to 7 and connected with 
the usual short bar. 


LAMONTIA CALIBANA, sp.n. (PI. I. fig. 18.) 


Head, cox, pectus, and abdomen metallic greenish. An- 
tenne black, bipectinated to the tip. Collar and patagia with 
some pale mauve scales. Fore wing long and narrow, black 
at base and along the veins and costa, gradually becoming 
transparent (‘ugitive scaling) outwardly. Hind wing black, 
completely scaled, short, and much rounded. Exp. 30 mm. 

Habitat. Trinidad, Rock, 1.1.18 (N. Zamont). In Coll. 
Lamont. 

Proc. Zoou. Soc.—1922, No. LX VII. 67 


998 


oe 


OMIAAEWNHWH 


ON NEW SPECIES OF TRINIDAD MOTHS. 


EXPLANATION OF THE PLATE. 


. Pococera brunnapex, sp. n. 

. Parangitia micapennis, sp. n. 

. Centronia plorator, sp. n. 

. HKpitomiptera purpurascens, var. 
. Parangitia nigrofulgens, sp. n. 
. Lpitomiptera marmorata, sp. n. 
. Syngria delicata, sp. n. 

. Hypena brodescens, sp. n. 

. Dicentria nondescripta, sp. n. 

. Dicentria klagesi Druce. 

. Rifargia brunnipennis, sp. n. 

. Phastia maricolor, sp. n. 

. Boriza kalodonta, sp. n. 

. Hydrias albiochrea, sp. n. 

. Sisyrosea guaica, sp. n. 

. Plusiodonta cupristria, sp. n. 

. Sanys implicata, sp. 0. 

. Lamontia calibana, sp. n. 

. Bryocodia cryptogramma, sp. 0. 
. Bryolimnia monodonta, sp. n. 

. Phostria duplicata, sp. n. 

. Samea delicata, sp. n. 

. Epitomiptera purpurascens, sp. ni. 


Pyralidee. 
Noctuide. 
Hypsidee. 
Noctuidee. 
Noctuide. 
Noctuidee. 
Uraniadee. 
Noctuide. 
Notodontide. 
Notodontide. 
Notodontide. 
Notodontide. 
Notodontide. 
Lasiocampidee 
Limacodidee. 
Noctuidee. 
Noctuidee. 
Zyganide. 
Noctuide. 
Noctuide. 
Pyralide. 
Pyralidee. 
Noctuide. 


ON THE NEMATODE GENUS PHYSALOPTERA, 999 


48. The Nematode Genus Physaloptera Rud.* By R. J. 

Ortierp, M.A., Research Assistant, Dept. of Helmin- 
thology, London School of Tropical Medicine; Hon. 
Parasitologist to the Society 1921-1922. 


[Received November 4, 1922: Read November 21, 1922. ] 
(Text-figures 1-44.) 


ConTENTS. Page 
Introduction . eee ERE HEEL Le eh atte tai ol: haat ceeeee eee i OOD) 
Material and Agknouletennants: Laine ashes asad Bae ci Co Stg RN 1002 
Methods of Study... : ASR Nets oe space eee EEE LOUD 
Conall Thoragiee of Epaaaiontan: nod canababeanon ecandaspevaucay aon ccom day 
Characters of Systematic Importance ........5.....00.. cece eee sen een eee 1007 
SKVSUO MARINO TSOGNION 1 Lae cho nad seo cad sacaaiBedeobue nse aug dooeacsaontsedoctene. MmOUlI 
Species Tarren Re SRO dociaee eNdionNoBosbaaeseea, |12U (60) 
Species wrongly attributed to ee PONIES SapescP coh eeobacoidoanannuce ell Lou 
Bilblioemap hiya wes tie ka asa a RRM Nee eS REL eee eae St eT Oa: 
InTdexpotSMeciestes: sea VNU CRORE A aay Une Diane oo reek tee ot TON 
INTRODUCTION. 


Rudolphi (1819), when establishing the genus Physaloptera 
expressed some doubt as to its validity. He gives the following 
as its generic diagnosis:—‘ Corpus teres élasticum utringue 

attentuatum. Os orbiculare. Cauda maris deflexa utr ringue alata, 
vesicam inferam sistens. Penis tuberculo emissus.’ Dujardin 
(1845), after quoting Rudolphi’s doubt, suppressed the genus and 
transferred the species to the genus Spiroptera. Diesing (1851) 
re-established the genus, and defined if in such a way that it 
could not be confused with any other genus of Nematodes. He 
added two more species, one of which—Ph. mucronata—was 
found to be an Ascarid by Molin. 

Leidy (1856) gave a short and incomplete description of four 
more species, two of which—JP%h. constricta and Ph. contorta— 
can be definitely ruled out from the genus, as they probably 
beiong to the family Gnathostomide as redefined by Baylis and 
Lane (1920). 

Up to the appearance of Molin’s monograph on the genus in 
1860, 18 species had been described as belonging to it, namely 
9 by Rudolphi, 2 by Creplin, 4 by Leidy, 2 by Diesing, and 1 by 
Hemprich and Ehrenberg. Of these, 5 are not members of the 
genus and | is doubtful. 


* Communicated by Prof. R. T, Lurene, F.Z.8. 
Oa 


1000 MR. R. J. ORTLEPP ON THE 


Molin’s monograph contains the descriptions of 22 species, of 
which 9 are new and 4—PA. colubri (Rud., 1918), Dies., 1851, 
Ph. linbata Leidy, 1856, Ph. abjecta Leidy, 1856, and Ph. spirula 
Hempr. & Ehren., 1828—are listed under Species Inquirende. 
In his introduction to this work he eliminates the four species 
Ph. strongylina Rud., Ph. mucronata Dies., Ph. saginata Rud., 
and Ph. tenwicollis Rud. from the genus, stating that Ph. mu- 
cronata and Ph. tenwicollis are Ascarids, Ph. saginata a 
Spiroptera, and that Ph. strongylina belongs to a new genus. He 
places Ph. megalostoma Crep., 1829, as a synonym of Ph. alata 
Rud. 

Schneider, in his ‘ Monographie der Nematoden’ (1866), de- 
scribes 9 species of Physaloptera, of which 4 are new, namely 
Ph. digitata, Ph. swbalata, Ph. truncata, and Ph. spiralis. Prior 
to his publication, anc after the appearance of Molin’s monograph, 
the only other new Physaloptera described is Ph. nvwris-brasilt- 
ensis Diesing, 1861, from Aus brasiliensis. 

In 1883, von Drasche amplified Molin’s monograph by pub- 
lishing a revision of the Molin and Diesing types deposited in 
the Vienna Museum; his work is of considerable value, as he 
paid special attention to the structure of the lips and caudal 
extremity of the males, and also gave figures of them all. He 
confirmed Molin’s diagnosis that ’h. mucronata Dies., 1851, 1s 
an Ascarid. 

Von Linstow, in his ‘Compendium der Helminthologie’ 
(1878) and in his “ Nachtrag” (1889) to this work, records 40 
species—31 in the first work and 9 additional ones in the latter. 

The next landmark was the appearance of Stossich’s monograph 
on the genus in 1889. He here gave the description of 37 species, 
26 of which he considered valid, and the rest he listed under 
Species Inquirende. Those species referred to in this work and 
not in Molin’s are as follows:—1 species—Ph. torquata, of 
Leidy, 1886; 5 species of von Linstow—VPh, crassa, 1879, Lh. 
dentata, 1883, Ph. malleus, 1883, Ph. striata, 1883, and Ph. pyra- 
midalis, 1879; 4 species—Ph. digitata, Ph. spiralis, Ph. subalata, 
and Ph. truncata, described by Schneider in 1886; and 1 species 
of Diesing —Ph. muris-brasiliensis, 1861. Of Stossich’s 37 species, 
11 are recorded from reptiles, 11 from birds, and 15 from 
mammals. 

Tn 1911, Gedoelst published his ‘Synopsis de Parasitologie,’ in 
which he listed 5 species, viz. 2 from man, | from carnivores, 
and 2 from birds. 

Hall (1916) gives the descriptions of 7 species, viz. 1 from 
Hyracoidea, 5 from rodents, and 1, PA abbreviata, which is a 
reptilian parasite, but had been recorded from Graphiurus murinus 
by Parona (1909). Hall doubts Parona’s diagnosis, and I agree 
with him that the presence of this parasite in a rodent is very 
suggestive of an error in its identification. 

Up to the present day the number of species has been con- 
siderably increased, thanks principally to von Linstow and 


\ 


NEMATODE GENUS PHYSALOPTERA. 1001 


Seurat. Won Linstow added 19 species to those of his mentioned 
above, one—Ph. amphibia—-being the only record of the oceur- 
rence of a member of this genus in amphibians, and another— 
Ph. cawcasica-—being the first species described from man. 
Seurat has added 8 new species and two varieties of Ph. alata, 
Teeeels diteylt). 

The latest addition to the literature of this group is two papers 
by Irwin-Smith reviewing the parasites found in reptiles. In 
her first paper (1921) she gives a host list of all the reptilian 
Physaloptera, as well as a bibliographical catalogue in connection 
with them. In her second paper (1922) she brings together the 
description of 13 species, all from lizards, and states that of the 
15 species recorded from this group of reptiles, only 9, perhaps 10, 
are valid, the others being either synonyms or not members 
of this genus. 

In Stiles and Hassall’s ‘Index Catalogue of Round-worms’(1920) 
80 species and 2 varieties of Physaloptera are listed, many of 
which, however, are either synonyms or do not belong to this 
genus. The addition of 3 species by Gedoelst (1916), 1 by 
Hall and Wigdor (1918), and 1 by Travassos (1920) not listed 
in the catalogue, and 7 new species described in this paper 
brings the number recorded up to 92. These are distributed 
among the principal host groups as follows :—43 species from 
mammals, 20 species and 2 varieties from birds, 28 species from 
reptiles, 1 species from amphibians, and 1 species, Ph. bonnet, 
sp. n., from a host in Dutch Guiana of which its native name 
is “Sapakara,” but of which I have been unable to obtain the 
scientific name. 

The following list comprises all the species recorded from each 
of these host groups :— 


AMPHIBIA. 
Ph. amphibia v. Linst., 1899. 


REPTILIA. 
Ph. abbreviata Rud., 1819. | Ph, longissima, sp. 0. 
Ph. abjecta Leidy, 1856. | Ph. monodens Mol., 1860. 
Ph. affinis Gedoel., 1916. |  Ph.mucronata Dies., 1851. 
Ph. alba Stoss., 1902. | Ph. obtusissima Mol., 1860. 
Ph. aloisii-sabaudie Par., 1907. eee pallaryt Seurat, 1917. 
Ph. antarctica v. Linst., 1899. | Ph. paradoxa v. Linst., 1908. 
Ph. britanica Skrjabin, 1916. | Ph. phrynosoma, sp. n. 
Ph. chameleontis Gedoel., 1916. | 227A. quadrovaria Leiper, 1908. 
Ph. colubri (Rud., 1819), Dies., 1851. | Ph. retusa Rud., 1819. 
Ph. constricta Leidy, 1856. | Ph. simplicidens, sp. un. 
Ph. contorta Leidy, 1856. Ph. sonsinoi vy. Linst., 1895. 
Ph. dentata v. Linst., 1883. Ph. spiralis Schn., 1866. 


striata v. Linst., 1883. 
varani, Parona, 1889. 


> 


| 

| 

| 
Ph. gracilis, sp. n. [1917, | Pe. 
Ph. leptosoma (Gerv., 1848), Seurat, | Ph. 


~ 


1002 


MR. R. J. ORTLEPP ON THE 


AVKS. 

Ph. acuticauda Mol., 1860. Ph. inflata (Mol., 1860), Stoss., 1889. 
Ph. alata Rud., 1819. Ph. malleus vy. linst., 1883. 

us » Vv. chevreuxi Seurat, 1914. Ph. megalostoma Crep., 1829. 

16 »  V. nouvelt Seurat, 1915. Ph. ovata v. Linst., 1907. 

Ph. bilabiata Crep., 1829. Ph. rotundata v. Linst., 1906. 

Ph. brevicauda v. Linst., 1909. Ph. saginata Rud., 1819. 

Ph. bulbosa v. Linst., 1906. Ph. striata v. Linst., 1883. 

Ph. crassa v. Linst., 1879. Ph. strongylina Rud., 1819. 

Ph. crosi Seurat, 1914. Ph. subalata Schu., 1866. 

' Ph. fusiformis v. Linst., 1902. Ph. tenuicollis Rud., 1819. 
Ph. galinieri Seurat, 1914. Ph. truneata Schn., 1866. 
MAMMALTA. 

Ph. anomala Mol., 1860. Ph. mawvillaris Mol., 1860. 
Ph. brevispiculum v. Linst., 1906. Ph. mephites Solanet, 1909. 

Ph. brevivaginata Seurat, 1917, Ph. mordens Leiper, 1908. 

Ph. capensis, sp. n. Ph. muris-brasiliensis Dies., 1861. 
Ph. caucasica v. inst., 1902. Ph. nasilionis Gedoel., 1916. 

Ph. cesticillata, Sons., 1889. Ph. numidica Seurat, 1917. 

Ph. circularis v. Linst., 1897. Ph. papilloradiata v. Linst., 1899. 
Ph. citilli (Rud., 1819), Hall, 1916. Ph. papillotruncata Mol., 1860. 
Ph. clausa Rud., 1819. Ph. preputialis vy. Linst., 1889. 
Ph. celebs v. Linst., 1897. Ph. pyramidatis y. Linst., 1879. 
Ph. digitata Schn., 1866 Ph. rara Hall & Wigdor, 1918. 

Ph. dilatata Rud., 1819. Ph. ruwenzorti Parona, 1907. 

Ph. dispar v. Linst., 1904. Ph. semilanceolata, Mol., 1860. 

Ph. elegantissima Stoss., 1902. Ph. sciuri Parona, 1898. 

Ph. gemina v. Linst., 1899. Ph. spivula Hempr. & Ehren., 1828. 
Ph. getula Seurat, 1917. Ph. tacapeusis Seurat, 1917. 

Ph. guiarti Garin, 1913. Ph. terdentata Mol., 1860. 

Ph. ineurva v. Linst., 1908. Ph. torquata Leidy, 1886. 

Ph. inermis v. Linst., 1906. Ph. torresi (Tray., 1920). (1912. 
Ph. limbata Leidy, 1856. Ph. tumefaciens Henry & Blane, 
Ph. magnipapilla Mol., 1860. Ph. turgida Rud., 1819. 

Ph. malayensis, sp. n. 


Doubtful Host. 


Ph. bonnei, sp. n. 


MATERIAL AND ACKNOWLEDGMENTS. 


The present work originated in the examination and deter- 
mination of a number of species of this genus collected by the 
author at the Prosectorium of the Zoological Society of London, 
while holding the post of Parasitologist to the Society. Prof. 
Leiper suggested that the work should be extended, so as to 
include a redescription of as many as possible of the recorded 
species of this genus, together with a survey of the whole group. 
With this aim in view he handed over to me, for study and 
identification, his valuable collection of Physaloptera, most of 
which had been personally collected by him during his expedition 


NEMATODE GENUS PHYSALOPTERA. 1003 


to Uganda in 1906. I wish here to express my indebtedness and 
thanks to the Zoological Society of London for the honour and 
privilege of collecting these parasites in their Gardens, and to 
Professor Leiper my sincere thanks and gratitude for having 
entrusted me with his material, for his valuable advice and 
criticisms, without which the work could not have been under- 
taken, and for the use of his valuable library. 

In addition to the above material, the writer was able to 
examine representative material of all of Molin’s species, the 
types of two of Diesing’s species, and examples of six of Rudolphi’s, 
including his types of PA. clausa deposited in the Helminthological 
collection of the Zoologische Abteilung der Naturhistorischen 
Staatsmuseum, Vienna. J wish to express my indebtedness and 
thanks to Hofrat Dr. Ludwig von Lorenz-Libtirnat, Director of 
the Zoological Department, for the privilege afforded me to 
examine this material, and especially also to Dr. Carl Graf 
Attems, custodian of the Helminthological collection in the 
Museum, for the excellent facilities so generously placed at my 
disposal. 

Molin did not set apart any material of his species as types, 
neither did von Drasche when re-examining Molin’s material. In 
consequence of this, I have taken the bottle with the lowest 
number to represent the type, except in the case of Ph. obtusis- 
sima and Ph, terdentata. In the former case a paratype of the 
species was examined; in the second case the bottle with the 
lowest number did not contain Molin’s species, but a repre- 
sentative of a later-described species of von Linstow’s, namely 
Ph. preputialis; consequently a bottle without any regular 
number upon it, but with the legend Y1074 on the stopper, was 
taken as the type; this material had been determined by Molin 
and also redetermined by von Drasche. The number of the 
bottles containing what I have taken as the type materials of 
Rudolphi’s, Diesing’s, and Molin’s species are recorded under the 
description of the individual species concerned. 

The types of the two new species, Ph. phrynosoma and 
Ph. capensis, described in this paper will be deposited in the 
British Museum of Natural History. The types of the five other 
new species, Ph. gracilis, Ph. longissima, Ph. simplicidens, 
Ph. malayensis, and Ph. bonnei, are in the Helminthological 
Department of the London School of Tropical Medicine. 


MeEtrHops or STuDY. 


The fresh material obtained from the Prosectorium of the 
London Zoological Society was collected and washed in normal 
saline, and as much as possible of the structure of the worms 
determined while they were still in the living state. They were 
then killed in hot 70 per cent. alcohol, according to the method ° 
recommended by Looss, after which they were preserved in a 
70 per cent. alcohol + 5 per cent. glycerine solution. For 


1004 _ -MR. R. J. ORTLEPP ON THE 


further study of this, and the rest of the material it was necessary 
to render them transparent by some clearing medium. Langeron’s 
lactophenol was at first used, but, although this medium is 
excellent for small nematodes, it did not give satisfactory results 
with the larger members of this group, as it did not render the 
specimen clear enough to make out the details of the spicules. 
Consequently beechwood creosote was used in its stead. In using 
this latter medium, the parasites were transferred into strong 
alcohol, about 95 per cent., and left therein for a longer or 
shorter period according to the size of the worms; after a 
sufficient interval to allow for the thorough penetration of this 
alcohol, they were placed in the creosote and examined, when 
clear, under the microscope by placing them on a larger slide 
and rolling them under a coverslip when necessary. After exami- 
nation it is essential to remove the creosote from the worms, as 
they tend to become very dark in colour if returned into the 
preserving fluid without removal of the creosote. To obviate 
this, the parasites were first immersed for at least two hours in a 
1-2 per cent. acid alcohol solution (70 per cent.), after which 
they could be transferred into the preserving fluid with apparent 
safety. 

All the measurements, except that of the length of the worms, 
were taken by means of an ocular micrometer, the value of each 
of whose divisions had been previously determined. 

The details of the female genitalia were made out in most 
species by simple clearing ang rolling of the parasites; in the 
remaining species it was necessary to dissect out these uteri 
under a dissecting microscope, and to examine them separately 
when mounted on a slide: where, however, there was ample 
material of the former, the genitalia were also dissected out. 

Where there was sufficient material available, the details of 
the lips were determined by first cutting the lips off from the 
body and then examining each lip from the inner and outer 
surface when cleared and mounted on a slide. 

All the figures except one—caudal bursa of Ph. colwbri—are 
original. All those accompanied by a scale were drawn by means 
of a camera lucida; the others are free-hand drawings made 
from the specimens. 


GENERAL CHARACTERS OF PHYSALOPTERA. 


These worms are rounded elongate and taper gradually in their 
anterior half and posterior third ; they vary considerably in size, 
the largest specimens known being 10 em. long and nearly 3 mm. 
thick. The cuticle is loosely attached to the body, and in nearly 
all eases is extended over the lips to form a prepuce-like collar ; 
in a few species a similar reduplication 1s present over the caudal 
extremities of both sexes. The cuticle always shows a very fine 
transverse striation, which often requires high magnification for 
its detection. 


NEMATODE GENUS PHYSALOPTERA. 1005 


Two cervical papille are present. These are situated laterally 
a short distance behind the cephalic extremity. They are small 
and spike-like, and stand out at right angles to the body. 

The excretory pore is ventral in position, and is situated a 
very short distance posterior to the cervical papille. It gives 
exit toa thin duct, which at first passes obliquely inwards and 
forwards and then bends backwards to join an oval, elongate, 
and unicellular gland pressed against the ventral sur face of the 
cesophagus. 

Two lips are present. These are lateral in position, and have 
their inner faces flattened and their outer somewhat convex; in 
lateral view they may be semicircular, dome-shaped, or more or 
less triangular in outline. They are always simple and their 
pulp is never subdivided. Each lip carries a variable number of 
teeth on its anterior and inner border ; these consist typically of : 
(1) an outer median tooth, generally triangular in shape and of 
variable size ; (2) an inner median tooth immediately internal to 
the outer median: it may be larger, smaller, or of the same size 
as the outer tooth, and its tip may be either simple and pointed 
or broadened out and tripartite; (3) two inner lateral teeth, 
one situated in each of the submedian fields of each lip on its 
inner surface: they are smaller than the outer tooth and are 
always split ; (4) a series of small denticles arranged in a linear 
series on the inner surface of the lip, dorsad and ventrad of the 
median teeth. The external median tooth is always present, but 
any or all of the other teeth can be absent. Two papille are 
present on the external surface of each lip; these are situated 
one in each submedian field towards the angies of the lips; they 
are generally dome-shaped. A number of observers mention the 
presence of an additional external papilla in the mid-line of the 
lip. I have not detected any such papille in any of the specimens 
studied, and [am consequently led to the conclusion that a small 
shoulder-like bulging, somtimes present on the outer surface of 
the lip, has been mistaken for a papilla. 

The cesophagus 1 is a straight and cylindrical organ, mcreasing 
shghtly in thickness towards its posterior extr emity. Tt is 
always divisible into two parts, namely,a short, transparent, and 
slightly thinner anterior muscular portion showing transverse 
muscular striations, anda posterior glandular portion which is 
thicker, oqaque, and granular. Its lumen is lined with chitin 
and is triradiate. Seurat mentions the presence of a buccal 
cavity between the lips and esophagus. I was not able to detect 
any such space in my material except in one, Ph. gracilis, sp. n., 
where I think the space was due to contraction; in all my other 
specimens the esophagus immediately followed the lips. 

The nerve ring is large, and encircles the muscular esophageal 
portion in its posterior half. 

The intestine is straight, and its connection with the cso- 
phagus is slightly thicker than the rest of the organ, which has 
a uniform thickness throughout. It opens to the exterior by a 


1006 MR. R. J. ORTLEPP ON THE 


short and oblique rectum, lied internally with chitin and 
accompanied on its outer surface by a number of rectal glands. 

The tail of the female is generally short and conical; it is 
supphed with a unicellular gland; two lateral caudal pores are 
present, situated generally behind its middle. 

The male caudal extremity is ineurved ventralwards, and 
carries on either side a cuticular expansion, which passes across 
the ventral surface, some short distance in front of the anus, 
to join its fellow of the other side. It is always supported by at 
least four pairs of long pedunculated papillae, generally arranged 
circumeloacally. Its wenn al surface is nearly always ornamented 
either by longitudinal ridges, or with longitudinal rows of 
tubercles, or spikes or rounded prominences. In addition, there is 
present a variable number of ventral papille (?absent in Ph. guiarti 
Garin) which ave generally sessile ; typically these consist of three 
pre-anal and five pairs of post-anal papille, but this number may 
vary ln some species. 

In the female the vulva is always situated in the anterior half 
of the body. Itis a circular aperture, generally situated on a 
slight elevation. It leads into an unpaired ovijectoral portion, 
which in its turn is followed by a double or multiple portion of 
the female genitalia. ine ovijectoral part consists typically 
of three subdivisions, viz.: (1) a thick-walled cylindrical and 
muscular vaginal part, lined internally with chitin; (2) a 
middle swollen portion, generally having its lumen filled with 
egos: this part I have interpreted as ‘‘ege-chamber”; and (3) a 
posterior cylindrical portion of about the same thickness as the 
vagina, but generally shorter: this part I have designated in the 
descriptions by the term ‘“‘common trunk”, as it probably 
represents a fused portion of the uteri. The first two parts are 
always present, but the last is often absent. The double or 
multiple parts consist of two or more tubules connected by a 
thinner part to the common trunk or egg-chamber, and com- 
prising a uterus, a receptaculum seminis, an oviduct, and an 
ovary. ‘The uteri are large, and are filled with numerous eggs ; 
they fill practically the whole body-cavity, and pass hadkiaics 
more or less parallel to each other to within a few millimetres 
from the posterior extremity; they then pass forwards, become 
thinner, and join on to the receptacula seminis. The receptacula 
seminis are small, oval, rounded, or pyriform, with their greatest 
diameter further away from the uteri. The oviducts are fine 
ducts, connecting the oviducts to the receptacula seminis ; their 
transition into the ovaries is generally gradual, but into the 
receptacula it isabrupt. The ovaries are long and taper gradually 
to their tip: they at first pass irregulary forwards, but recurve to 
end generally in the anterior body half. 

The eggs are relatively small, oval, smooth, and very thick- 
shelled, and are embryonated before being laid. 

The male genital tubule is unpaired, and is situated ventrad 
to the digestive tract. It consists of three well-defined portions, 


NEMATODE GENUS PHYSALOPTERA. 1007 


viz.: (1) a posterior and short but elongately pyriform ejacu- 
latory duct, with its thickest part away from the genital orifice; 
(2) a long vesicula seminalis, of the same dimensions throughout 
and slightly thicker than the ejaculatory duct; and (3) a much 
thinner and tapering testis, which passes forwards into the 
anterior third of the body, after which it recurves, and passes 
backwards to end at about the middle of the body. The ejacu- 
latory duct and vesicula seminalis are straight, but the testis 
performs irregular longitudinal loops in its forward course. The 
accessory male genital apparatus consists of two spicules, of which 
the left is generally long and slender, and the right shorter and 
stouter; they may, however, be of the same length, or the right 
may be slightly larger than the left. 


CHARACTERS OF SySrEMATIC IMPORTANCE. 


Seurat (1914) was the first to draw attention to the systematic 
value of the number of the uteri, and in his writings on the 
Physaloptera invariably groups his species according to whether 
they have two or four uteri. In my investigations on this genus 
I have found also forms possessing as many as 1 uteri, ae con- 
sequently have classified them into the three groups—Didelphys, 
Tetradelphys, and Polydelphys. 

The number of the uteri and their mode of origin from the 
ovijector appear to me to be characters of primary systematic 
value, and the question arises whether they are not of generic 

significance. 

Irwin-Smith (1922) considers it ‘‘ undesirable to establish a 
new genus entirely on a character which can be determined only 
vy dissection.” J agree with her that it does not appear legitimate 
so create a genus on only one character; and consequently I have 
looked for some other characters common to each of these groups, 
but have not been able to find any. The whole genus is so 
compact, and its individual species have so many characters in 
common, that at the present state of our knowledge of the 
Physaloptera, I am compelled to retain them all in the same 
genus, however much I have felt inclined to split them up for the 
sake of convenience. 

Travassos (1920) separates from the genus Ph ysaloptera four 
new genera, limiting the genus Physaloptera to those forms 
possessing similar and subequal spicules, two uteri, four pairs of 
pedunculated papille, and having no reduplication ‘of the cuticle 
over the caudal extremity. He gives the following key to these 
five and three other nearly related genera :— 


1. Spicules similar and subequal. 
A. Two uteri. 
a. Four pairs of pedunculated papille ; no Se -like sheath 


at the posterior extremity” ............ es... Physaloptera. 
b. A prepuce-like collar present at the postene 10r ety Sani ... Chlamydonema. 
¢e. Hight pairs of pedunculated papillee.....................000-0-008 Lhubunea. 


13}, TNSI0) ABT E cabaret bce Peale ce ese nieek Ounce rates Buta See lal MN ec RA ACTH ted Me He 


9 


1008 | MR. R. J. ORTLEPP ON THE 


2. Spicules dissimilar, and their sizes very different. 


AA. Two uteri. 


aa. Four pairs of pedunculated papille ........................... Abreviata. 
6b. Nine to ten pairs of pedunculated papille. 
ec. Vulva in anterior half; ovijector very long ............_Heliconema. 
dd.) Noulva)nean to) thevanuspeeeeeasenentece sso eseeee ete se) enOLeous: 
E318}, (I OWED (RUG an denGdoxlasonaopd venue sdeane box sea can bon dod obewnolonockaposans,, \ALA/OSOHIG 


Except for the genus Twurgida, Travassos gives no generic 
diagnoses for his new genera, so that in order to form a conception 
of the three genera Chlamydonema, Abreviata, and Leptosoma, we 
are limited to the skeletal framework of his key. He also does 
not indicate which species he regards as the type of each of these 
three genera, but as he has used the term Vurgida in raising the 
species Ph. turgida to generic rank, I take it that Chlamydonema 
will have as its type-species Ph. preeputialis von Linstow, 1899, 
because Chlamydonema felineus is the name used by Hegt (1910) 
in describing a nematode parasite from the Domestic Cat, being 
apparently unaware of its physalopteran characters or its identity 
with Ph. preputialis. Abreviate is presumably Rudolphi’s 
species Ph. abbreviata raised to generic rank, and Leptosoma is 
presumably Ph. leptosoma (Gerv ais) Seurat, 1917, raised to a new 
genus. 

In view of Seurat’s investigations and my own observations on 
the genus Physaloptera, all Travassos’ genera appear to me to 
be untenable. Chlamydonema, besides the characters it has in 
common with Travassos’ restricted genus Physaloptera, has in 
addition the prepuce-like collar over the posterior extremity ; 
now, Ph. tumefaciens Henry & Blanc, 1912, also shows this 
characteristic, although not to such a marked extent as 
in Ph. preputialis, but as it has four uteri, it cannot be 
included in the genus Chlamydonema, so that this genus is not 
valid. 

The genus 7urgida has all the characters of the genus Physa- 
loptera as restricted by Travassos, except that it 1s polydelphoid, 
so that its validity 1s based on this single characteristic. As 
shown above, I do not regard the multiplication of the uteri by 
itself as of more than specific value. Ph. capensis, sp. n., 18 also 
polydelphoid, but the mode of origin of its uteri is quite different 
from that in Ph. turgida, so that if the genus Z’urgida is accepted, 
then a special genus would have to be created for Ph. capensis, a 
procedure which I consider quite unjustifiable. 

Ph. abbreviata, as vedescribed by Seurat (1914 6), has four uteri, 
so that on this ground alone the genus Abreviata must be 
suppressed, as Travassos creates it for those forms possessing two 
uter1 and having very equal and dissimilar spicules; besides, the 
tetradelphoid species Ph. tumefaciens and Ph. magnipapilla have 
only slightly unequal spicules, which character also does not 
tally with Travassos’ conception of his new genus. 

The genus /eptosoma, according to TMravacsos: key, is char- 
acterised by having four uteri and very unequal and dissimilar 


NEMATODE GENUS PHYSALOPTERA. 1009 


spicules. According to Seurat (1917a@), who has studied Ph. 
leptosoma, this species has only two uteri, so that again it 
appears that Travassos has made a new genus on wrong data. 
At any rate the name Lepiosoma is not available, as according to 
Seudder (1882) it has already been used by Leach (1819) for a 
beetle, by Risso (1826) for a crustacean, by Nardo (1827) for a 
fish, and by Boisd. (1833) for a lepidopteran, while the form 
Leptosomus was used by Vieill. (1816) for a bird, and by Schonh. 
(1826) for a coleopteran. 

With regards to the genera Vhubuncea Seurat, 1914, Heli- 
conema Trav., 1919, and Proleptus Duj., 1845, 1t does not 
appear necessary to discuss them here, as they do not affect the 
status of the genus Physaloptera. 

Seurat (1915-16) has also proposed dividing the genus into 
four groups according to the teeth. These groups he characterises 
briefly as tollows :— 


1st group of Ph. galinieri Seurat. 


The internal median tooth tripartite and very large, passing 
over the small external tooth. 


2nd group of Ph. clausa Rud. 


The external and median teeth are of the same height. 
3rd group of Ph. alata Rud. 


The external tooth is very large and triangular; the internal 
tooth (tripartite) is small. 


Ath group of Ph. abbreviate Rud. 


The external tooth is enormous and triangular; the internal 
tooth is very small and difficult to see. 


lf we adopt this division, we would be obliged to add a 
5th group of Ph. colubri (Rud.), Dies., characterised by the 
presence of a large and triangular external tooth and the entire 
absence of the internal median tooth. 

In subdividing any group of animals, our first aim is to bring 
together those forms which possess close affinities to each other, 
and not those forms which, when grouped together, render it 
easier or more convenient to us to determine them. Seraut’s 
divisions, when submitted to this test, are consequently not 
tenable, as it would imply that all the polydelphoid forms are 
closely related to such didelphoid forms as Ph. preputialis and 
Ph. maxillaris. 

Assuming that the line of evolution has proceeded from the 
didelphic to the polydelphic forms, then, I think, we have to 
recognise two ways, both starting from a form with genitalia of 
the Ph. clausa (text-fig. 1 A) type. The one group would evolve 
from this type through stages like Ph. preputialis (text-fig. 1 B), 
Ph. tumefaciens (text-fig. 1 C) to Ph. turgida (text-fig. 1 D), and 


1010 MR. R. J. ORTLEPP ON THE - 


the other along such stages as Ph. obtusissima (text-fig. 1b), 
Ph. abbreviata (text-fig. le), Ph. paradoxa (text-fig. 1d) to 
Ph, capensis (text-fig. 1 e). 


Text-figure 1. 


My 


i 


A 


d. 
J 
Ay Be ac 
Dp 
“INXS 
ce. 
GIS 


Schematic representation of the probable methods of Uterine evolution from the 
didelphoid to the polydelphoid conditions. 


A= Ph. clausa type; B= Ph. preputialis type; C= Ph. tumefaciens type ; 
D= Ph.turgida type; 6 = Ph. obtusissima type; ¢ = Ph. abbreviata type; 
d = Ph. paradoxa type; e = Ph. capensis type. 


For the specific determination of the material examined, the 
following characters were especially noted :— 


Number of uteri and their mode of origin. 

Number and arrangement of the male caudal papillee. 

Relative lengths and shape of the spicules. 

Number, shape, and size of the labial teeth. 

Cuticular markings on the ventral surface of the imale 
bursa. 

Position of the vulva. 


OU Go bo 


ee 


NEMATODE GENUS PHYSALOPTERA. 1011 


7. Presence or absence of a cuticular sheath at the posterior 
extremity. 
8. Size of the eggs. 
9. Relative length of the cesophagus. 
10. Positions of the excretory pore and cervical papille. 


The value attaching to each of these characters is indicated by 
its position in the above list. 


SYSTEMATIC SECTION. 


The remainder of the paper is divided into two parts. Part I. 
contains the descriptions and diagrams of all the species 
personally studied. Part II. gives an extract of the principal 
characteristics of all the species not personally examined. In 
most cases these extracts are based on the original descriptions 
and drawings; where this is not the case, the source is recorded 
under the species concerned. In each part the parasites are 
classified into three groups according to whether they were 
di-, tetra-, or polydelphoid, the arrangement proceeding from 
Mammals to Birds and to Reptiles. 

Ph. acuticauda, however, has been placed together with 
Ph. preputialis and Ph. malayensis after the reptilian didelphoid 
forms, because of the similarity of their female genitalia. Species 
Inquirende, and species which were described as belonging to 
the genus Physaloptera but which have since been found to 
have been misplaced, are listed at the end of Part IT. 

The hosts cited after the species in Part I. are the hosts from 
which the material examined was collected; the hosts in Part IT. 
are those from which the material collected formed the basis of 
the descriptions. A more detailed list of hosts from which the 
separate species have been recorded is furnished by Stiles and 
Hassall’s ‘ Index Catalogue of Round-worms ’ (1920). 


PART I. 
Generic Diagnosis (emended). 
Superfamily SPrrvurorDe. 
Family PHYSALOPTERID&. 


Subfamily PuysaLoPrerinz. 


Mouth with two large simple and lateral lips, each armed with 
a variable number of teeth and carrying two external papille. 
Cisophagus consists of an anterior muscular and a_ posterior 
glandular part. Cuticle generally reflected over the lips. Caudal 
extremity of male with lateral ale joining each other across the 
ventral surface in front of the anus. At least four pairs of 
pedunculated papiile supporting the ale; these generally arranged 
eircumeloacally. A variable number of ventral caudal papille, 


1012 MR. R. J. ORTLEPP ON THE 


generally three pre-anal and five pairs post-anal. Spicules un- 
equal, subequal, or equal. Vulva in anterior half of body. Two, 
four, or many uteri. Eggs oval, smooth, thick-shelled, and 
embryonated when laid. 

Parasitic normally in the digestive tract, generally the stomach, 
_ of mammals, birds, and reptiles, but very rarely in amphibians. 


Type-species, Ph. clausa Rud., 1819. 


Grove Didelphys. 
(1) PHysatoprera cuausa Rudolphi, 1819. (Text-figs. 2-4.) 
Syn. Spiroptera clausa (Rud.), Duj., 1845. 


The material examined consisted of many specimens from 
Erinaceus europeus in the Vienna Museum (bottles 4447 & 4450), 
also some specimens from the same host in the collection of 
Prof. R. T. Leiper. The three lots of material were in all 
respects identical. 

The cuticle appears smooth, but under high magnification a 
very delicate transverse striation is s2en ; it is only very slightly 
reflected over the base of the lips. 

The cervical papille are lodged in small depressions of the 
cuticle, and are situated from 115 to 165 » behind the level of 
the hind end of the muscular cesophagus; the excretory duct 
opens in the ventral mid-line at the same level as the cervical 
papille or just behind them. 

The lips are almost semicircular in lateral view, but appear 
square in dorsal or ventral view. Hach bears a medium-sized 
conical outer tooth with obtuse tip, and just internal to it a 
median tooth of equal height with three tips to its free end. 
No lateral teeth ave present. The subdorsal and subventral 
external papilla are small and dome-like. 

The csophagus immediately follows the lips; it is straight, 
and forms in the female 1/7°8 to 1/9th, and in the males 1/5:4 to 
1/56 of the total body-length. Its muscular part is distinetly 
set off from the glandular, and is also slightly thinner ; in the 
females it forms about 1/8°5, and in the males 1/7th of the whole 
organ. The nerve ring encircles it in its posterior quarter. 


Female. 

Mature females vary in length from 30 to 50 mm., with a 
maximum thickness of 1:3 to 1:6 mm. The body is attenuated 
in its anterior 1/3rd and posterior 1/5th ; it is terminated by a 
bluntly conical tail 1/45th to 1/55th of the total body-length, 
with caudal pores in its posterior half. 

The postoion of the vulva divides the body in the ratio of 
1:1:7 to 1:2; it is situated on a slight elevation, and leads 
into a bent vagina 1:6 mm. long by 95 » broad ; the egg-chamber 
is slightly shorter but thicker, measuring 1:3 mm. long by 325 yw 
broad. From its posterier end the uteri, of which there are two, 


NEMATODE GENUS PHYSALOPTERA. 1013 


directly take their origin, so that a common trunk portion is 
absent. 

The eggs are oval and thick-shelled, measuring 52 yp long by 
38 « broad, 


Male. 

The males varied in length from 15 to 28 mm., with a breadth 
of 825 to 875 pw. The body is attenuated only on its anterior 
half. 

The caudal bursa is large, forming about 1/11th of the total 
length ; it is bluntly rounded posteriorly, and is closely coiled 
ventralwards. The lateral expansions are well developed, and 


i 


Text-figure 


Physaloptera clausa Rud. 
A = Anterior extremity of body. 
B = Externo-lateral view of ip. 
C = Ventral view of lip. 


the four pairs of pedunculated papillae supporting them are equi- 
distant, two pairs being pre-anal and two pairs post-anal. There 
are 13 ventral papille, of which three are pre-anal and the rest 
post-anal. The pre-anal papille are situated in a row imme- 
diately anterior to the anus, the middle papilla being larger than 
the other two. The first two pairs of post-anal ventral papille 
are small and situated in a row just behind the anus; the remain- 
ing papille are equidistant and large, and they divide the tail 
into quarters ; In some cases the last two pairs may be slightly 
approximated to each other. 

The left spicule is nearly half as long again as the right, 
and is also much slenderer; itis 740 long by 33 » broad at 


Proc. Zoou. Soc.—1922, No. LX VITI. 68 


1014 _ MR. B. J. ORTLEPP ON THE 


its base and tapers to a fine point; the right spicule is stout, 
495 w long and 50 broad at its base, and also terminates in a 
fine point. 

Host: Hrinaceus europeus. Stomach. Hurope. 

Types in bottle 4447 in the Vienna Museum. 

My observations on this species differ in many respects from 
those of Seurat (19176) made on worms obtained from Hrinaceus 
algirus and Hrinaceus deserti. These differences apply more 
especially to the male. Seurat finds that of the three pre-anal 
ventral caudal papille the two lateral ones are far removed from 


Text-figure 3. 


Physaloptera clausa Rud. 


A = Caudal extremity of female. 
B = Terminal parts of female genitalia. 
C = Egg-shell. 


the median papilla and are also stalked; in my material these 
three papille are situated close together in a line immediately in 
front of the anus, and the lateral papille are non-pedunculate ; 
also the median papilla is large and nearly twice the size of 
the other two, whereas Seurat finds this papilla to be very 
small. 

With regard to the post-anal ventral papille, the third pair is 
stalked in Seurat’s material and sessile in mine. 

Further, the spicules of his specimens differ from those of 
mine both in length and in shape; in his material the right and 


NEMATODE GENUS PHYSALOPTERA. 1015 


left spicules are respectively 300 and 360 long, and only the 
left has a pointed tip. In my material the spicular lengths are 
as follows: right 495, left 740, i.e. the right is more 
than half as long again and the left twice as long. Both of 
them terminate in sharp points. 

These, besides other minor differences, e.g. size of egg, length 
of ovijector, position of the vulva, etc., seem to indieate that the 
specimens studied by me belong to different species; and as I 
examined what I take to be the type-material, I think Seurat’s 
specimens must be regarded as a new species, which, however, is 
very closely allied to Ph. clausa, as shown by the nature of the 
teeth and mode of origin of the two uteri. 


Text-figure 4. 


Physaloptera clausa Rud. 
A = Caudal extremity of male. B = Spicules. 


Schneider (1866) states that the inner teeth are smaller than 
the outer, and figures a male bursa having the papille arranged 
very similarly to those figured by Seurat; he does not definitely 
state that he has examined the Vienna material, although he 
quotes the same host from both Vienna and Berlin; it is probable 
that he examined material only from the latter locality, and that 
would partly account for the differences between his and my 
observations ; it is therefore probable that he also is not dealing 
with Ph. clausa, but with a species identical with or very closely 
allied to that described by Seurat. Unfortunately Schneider’s 
description is very brief, so that I am not able to make a closer 
comparison between his and Seurat’s accounts. 


68 


1016 MK. R. J. ORTLEPP ON THE 


(2) PHysALoprera PAPILLOTRUNCATA Molin, 1860. (Text-fig. 5.) 


The material examined consisted of the types of the species 
deposited in the Vienna Museum; this material had also been 
examined by von Drasche. 

The cuticle is very finely striated transversely, and in the 
specimens examined it was not reflected over the lips; a coarse 
ringing is present in some specimens, probably due to some 
shrinkage. 

The cervical papille are situated from 150 to 250 u behind the 
level of the posterior margin of the muscular cesophagus, and the 
excretory duct opens to the exterior either at the same level or 
about 75 « further back. 

The lips are hemispherical in lateral view, and each has only 
two teeth, namely a large and blunt outer tooth recurved out- 
wards, and a smaller tripartite inner tooth. Hach lip bears on 
its subdorsal and subventral outer surface a dome-like papilla. 

The cesophagus is straight, and forms in the females from 
1/5°6 to 1/7th, and in the males 1/5th of the total body-length. 
Its muscular part is slightly thinner than the following glandular 
part, and forms from 1/8°6 to 1/10th of the whole organ: it is 
encircled by the nerve ring at the junction of its fourth and last 
fifths. 


Female. 

The females examined varied in length from 32 to 68 mm., 
with a maximum breadth of from 1 to 2°? mm. The body is 
much attenuated in its anterior half, and slightly less so in its 
posterior quarter, where the body is terminated by a short and 
truncated tail 1/66th to 1/80th of the total body-length; its 
caudal pores are situated just behind its middle. 

The vulva opens on a slight elevation, and is situated rela- 
tively far back; it divides the body in the ratio of 1: 1:8 to 1:12; 
it leads into a slightly curved and muscular vagina 1:5 mm. 
long by 96, broad; this passes into the egg-chamber, which is 
1:3 mm. long by 210 broad. The trunk which follows is of the 
same length and breadth as the vagina; from its posterior end 
there arise the two uteri which at first pass forwards, but soon 
recurve and pass down the length of the body. 

The eggs are oval and thick-shelled, measuring on the average 


52 pw long by 30 « broad. 
Male. 


The males are much smaller than the females, and are 
attenuated in their anterior third, the rest of the body being 
of about the same thickness; they vary in length from 20 to 
25 mm. with a breadth of 750 to 800 p. 

The bursa is large, and forms about 1/10th of the body-length ; 
it has well-developed cuticular expansions, and is rounded at its 
tip. The ventral surface is traversed by longitudinal rows of 
protuberances. The four pairs of stalked papille are equidistant 


NEMATODE GbNUS PHYSALOPTERA. 1017 


from each other, the first, however, being more ventral in origin ; 
two pairs are pre-anal, and the other two pairs are post-anal in 
position. The three pre-anal ventral papille are situated in a 
row immediately in front of the anus. There are five pairs of 
post-anal papiile, of which pairs 1 and 2 are small and situated 
close together immediately behind the anus, one pair being 
slightly more posterior than the other; pairs 3, 4, and 5 are 
equidistant from each other; the distance between the 2nd and 
3rd pairs is about half that between the 3rd and 4th pairs, and 
the distance between the 5th pair and the tip of the tail is about 
twice that between the 4th and 5th pairs. 


Text-figure 5. 


OS mm. 


Physaloptera papillotruncata Mol. 
A = Cephalic extremity of body. 
B = Inner view of median labial teeth. 
C = Caudal extremity of male. 


The spicules are equal and straight; the right spicule is slightly 
slenderer than the left, and ends in a sharp point; the end of 
the left spicule is slightly flattened out to form a small spear- 
head; they are 360 long by 35 and 38 thick at their base 
respectively. 

Host: Myrmecophagu jubata. Stomach. Brazil. 

Types in bottle 4477 in the Vienna Museum. 

Discussion.— See Ph. maxillaris. 


(3) PHysALOPTERA SEMILANCEOLATA Molin, 1860. (Text-fig. 6.) 
The material examined was somewhat shrunken, but in 


excellent preservation. 
The cuticle is finely striated, and shows, in addition, an 


1018 - MR. R. J. ORTLEPP ON THE 


irregular course ringing, probably due to contraction ; in some 
cases it is completely reflected over the lips, and in others only 
the base of the lips are covered. 

The cervical papille are situated about 300 » behind the level 
of the hind end of the muscular cesophagus, and the excretory 
duct opens about 90 « further back. 

In the specimens examined the cwsophagus was thrown into 
loops, so that a correct estimation of its length could not be 
determined; roughly, it forms in the male about 1/5th, and in 
the female about 1/6th of the total length. 

The lips are flatly rounded, and each bears two large terminal 
teeth of the same height ; the outer tooth is markedly truncated, 
whereas the inner is foliaceous and tripartite. Each lip has on 
its outer surface a prominent subdorsal and subventral papilla. 


Female. 
The females are coiled, and mature specimens average in length 
from 21 to 37 mm. leng by ‘9 to 1°5 mm. thick. The body is of 


Text-figure 6. 


Physaloptera semilanceolata Mol. 


A = Externo-lateral view of head. 
B = Caudal extremity of male. 


a more or less uniform thickness throughout, except for the 
anterior i/10th and posterior few millimetres. The tail in all 
specimens examined was bent dorsalwards. It tapers only 
slightly towards its tip, which is very bluntly rounded; the 
caudal pores open in its posterior 1/5th. 

The vulva opens very far forwards, in some specimens its 
position being just anterior to the hind end of the esophagus 
and in others just behind it. It leads into a long and coiled 
vagina about 3 mm. long by 110m broad; its posterior end 
gradually widens to join the swollen egg-chamber, which, when 
distended with eggs, measured 2°75 mm. long by 440 pw broad. 
The common trunk is relatively short and slightly thinner than 
the vagina; it measured about 475 w long by 95 thick. The 
two uteri which arise from it may first pass forwards, and then 
recurve to pass down the length of the body. 


NEMATODE GENUS PHYSALOPTERA. 1019 


The eggs are oval and thick-shelled, and are already embryo- 
nated in utero; they are 50 p long by 32 » broad. 


Male. 

The males are also coiled, and the body is attenuated in its 
anterior third. The specimens examined measured from 25 to 
32 mm. long by 750 to 875 thick. The bursa is relatively 
narrow and pointed, and three of the four pairs of pedunculated 
papilla: are pre-anal in position. The three pre-anal ventral papille 
are equidistant from the cloaca, and the central papilla is slightly 
larger. There are five pairs of post-anal ventral papille, of which 
the first two pairs are small and situated in a row immediately 
behind the anus; the third pair is at the junction of the Ist and 
2nd sixths of the tail, the fourth at about the middle, and the 
last at the junction of the 3rd and last quarters of the tail. The 
ventral surface is covered with longitudinal rows of irregular 
tubercles. 

The spicules are slightly unequal, of the same thickness, and 
slightly curved ; both taper to fine points; the right is 574 pw long 
by 384, thick at its base, the left is 730 long by 38'p at its base. 

Host: Vasua narica. Stomach. Brazil. 

Discussion.—The small pair of papille between the 4th and 
5th pairs of ventral post-anal papille described by von Drasche 
are the openings of the caudal pores. 

Affinities.—See Ph. mawillaris. 


(4) PHYSALOPTERA MAXILLARIS Molin, 1860. (‘Text-fig. 7.) 


The material examined consisted of the types of this species 
deposited in bottle 4458 in the Vienna Museum; the material 
was in an excellent state of preservation. 

The cuticle is finely striated transversely, and is partly or 
wholly reflected over the lips. The cervical papille are situated 
half a millimetre behind the junction of the two csophageal 
parts, and the excretory pore about 50 y further back. 

The lips are rounded or slightly conical; each has two terminal 
teeth—numely, a large triangular outer tooth with obtuse tip 
shghtly bent outwards, and an inner semi-membranous tooth, 
tripartite at its free end, and of the same size as the outer tooth. 
Two large conical papille are present on the outer surface of 
each lip. 

The csophagus is straight, and thickens gradually towards its 
posterior end ; in the female it forms 1/5:2 to 1/5:4, and in the 
male 1/6th to 1/6°4 of the body-length ; its anterior tenth forms 
the muscular part, which is slightly thinner than the glandular, 
and is encircled by the nerve cord in its posterior third. 


Female. x 

The females are stout, and are attenuated only in their anterior 
third and tail region ; this latter part tapers abruptly to form 
a short and pointed tail 1/73rd of the body-length, and having 
its caudal pores situated in its middle. Mature forms vary in 


1020 MR. R. J. ORTLEPP ON THE 


length from 20 to 36 mm., with a maximum breadth of 800 u 
to 1°24 mm. 

The vulva is slightly protuberant, and its position divides the 
body into the ratio of 1: 1°6. The vagina is long, and varies in 
length from 21 to 34 mm. with a thickness of 95y. ‘The 
egg-chamber is elongately oval, and is 1°7 mm. long by 435 pu 
in diameter. The common trunk is short, and varies in length 
from 290 to 320; it is slightly thicker than the vagina; from 
its posterior end the two uteri take their origin, and may 
either pass forwards and then bend backwards, or else pass 
posteriorly directly. 

The eggs are oval and thick-shelled, and measure 43 p long by 
31 pw broad. 


Text-figure 7. 


Mmm. 


Physaloptera maxillaris Mol. 
Caudal extremity of male. 
Male. 

The males are much attenuated from behind forwards, having 
their maximum thickness just above the bursa; they vary in 
length from 23 to 32 mm., with a thickness of 550 to 800 nu. 

The bursa is elongated and somewhat pointed, and has its 
caudal expansions well developed in its middle portion. Its 
ventral surface is ornamented by two types of cuticular bosses, 
those in the ventral area, above and below the anus, being 
rounded, simulating those found in Ph. preputialis, while those 
on the lateral areas are in the form of much-broken ridges; the 
transition from the one type to the other is gradual. 


NEMATODE GENUS PHYSALOPTERA. 1021 


The four cireumcloacal lateral papille are provided with very 
long stalks, especially the Znd and 3rd pairs. The ventral 
papille in the immediate vicinity of the anus are small, and 
consist of three pre-anal papille in a row, and two pairs post-anal, 
also in a row. Further down the tail there are three additional 
pairs; these are equidistant from each other, the most anterior 
being situated just behind the level of the last pair of stalked 
papillee. 

The spicules are slightly bent, unequal, and the right is 
stouter than the left; both taper to sharp points. The left is 
1-2 mm. long by 40, thick at its base, and the right is 560 p 
long by 57 p thick at its base. 

Host: MJephites chinche. Stomach. Brazil. 

Types in bottle 4458 in the Vienna Museum. 

Some specimens of this species, collected from Mephites 
meplitica, were handed over to me for identification by Mr. R. I. 
Pocock, F.R.S.; he obtained them from animals introduced into 
Northumberland (Hngland) from Canada, but how long they had 
been in England before the parasites were collected it was not 
possible to definitely find out. J wish to express my thanks to 
Mr. Pocock for this material. 

This material differs from the types studied in that they are 
slightly larger, and the bursa of the male appears rounded and 
lobulated ; however, I believe these characters of the bursa to be 
due to shrinkage, as the worms had been killed in formalin. 

Solanet in 1909 described another species—Ph. mephites— 
from J/ephites suffocans, Buenos Aires. Unfortunately I have 
not been able to consult his description, and consequently cannot 
discuss the relation of this species to Ph. mawillaris. 

The arrangement of the ventral bursal papille, the presence of 
two uteri, general characters of the teeth, ally this species to 
Ph. semilanceolata, Ph. clausa, and Ph. papillotruncata. Ph. semi- 
lanceolata differs from it by its truncated outer tooth, more 
anterior position of the vulva, shorter left spicule, and type of 
ornamentation on the bursa. It differs from Ph. papillotruncata 
by its longer and pointed spicules, ornamentation on the bursa, 
and smaller size of the adult females. Ph. clawsa is distin- 
guished from it by the size of the spicules, markings on the 
bursa, and absence of a common trunk to the unpaired female 
genitalia. 


(5) PuysaLoprera ANOMALA Molin, 1860. (Text-fig. 8.) 


The material examined consisted of three males and one 
female; type-material in bottle 4446 in the Vienna Museum. 
The material was in excellent preservation. 

The cuticle is finely striated transversely, and is partly reflected 
over the base of the lips; the cervical papille pierce it on either 
side from 320 to 400 « behind the level of the posterior end of the 
muscular cesophagus; the excretory pore is found about 100 p 
further back. 


1022 MR. R. J. ORTLEPP ON THE 


The lips are large and conical, and each carries a large conical 
but blunt tooth; the inner tooth is large and of equal height, 
with its free end divided into three points, of which the middle 
one 1s slightly shorter than the other two. No lateral teeth are 
present. Externally each lip carries a subdorsal and subventral 
conical papilla. 

The esophagus has its anterior muscular part thinner than 
the rest, and the nerve ring encircles this part in its posterior 
quarter. ‘The esophagus forms in the male about 1/5'4, and in 
the female 1/56 of the total body-length. 


. Female. 
The solitary female was 36 mm. jong and just under 2 mm. 


Text-figure 8. 


Te 
we HT 
Wi Att 
Perea 


Physaloptera anomala Mol. 


A = Outer view of median teeth. 
B = Caudal extremity of male. 


broad in its middle; the body is much attenuated in the anterior 
half, but only slightly so posteriorly; it is terminated by a 
bluntly-pointed tail, 1/60th of the body-length, and having the 
caudal pores in its posterior half. 

The vulva is situated just behind the end of the csophagus, 
and divides the body in the ratio of 1: 4:2; it leads into a coiled 
vagina 1‘) mm. long by 95» thick; this jos with a large and 
spacious egg- -chamber 2 mm. long ‘by 560 » broad, which pos+ 
terlorly narrows suddenly to join the 400 pe long common trunk, 
which is of the same thickness as the vagina. ‘The posterior end 


NEMATODE GENUS PHYSALOPTERA. 1 023 


of the trunk divides into two branches, each of which connects 
up with one of the two uteri. 

The eggs are thick-shelled and oval, measuring 38 p long by 
26 pw broad. 


Male. 

The males are from 30 to 32 mm, long by 1 to 1:2 mm. broad; 
they are attenuated towards both extremities and the bursa is 
shghtly incurved. 

Lhe caudal bursa is large, and is covered ventrally by 
longitudinal rows of cuticular elevations. The four pairs of 
pedunculated papille are situated three pairs pre-anal and a pair 
post-anal. Of the three pre-anal ventral papille the middle one 
is nearer the anus. There are five pairs of post-anal papille, of 
which the pairs | and 2 are small, and situated in a row imme- 
diately behind the anus. Pairs 4 and 5 are iarge, and are 
situated in the middle area of the tail. The distance between 
the 3rd and 4th pairs is about four times the distance between 
the 2nd and 3rd pairs, and twice that between the 4th and Sth 
pairs. 

The spicules are short and equal, each measuring 495 long 
by 32, broad at their base; they are straight, tubular, and 
pointed. 

Host: felis onca. Stomach. Brazil. 

Types in bottle 4446 in the Vienna Museum. 


(6) PHYSALOPTERA MURIS-BRASILIENSIS Diesing, 1861. (Text- 
ey 5) 


Syn. Spiroptera bilabiata Mol., 1860 (not Spiroptera bilabiata 
(Crep., 1829), Duj., 1845). 

The material studied consisted of ten females in bottle 4525 
and two males in bottle 4524 in the Vienna Museum. ‘This 
material constituted the types of this species. 

The cuticle forms a ring round the base of the head, but leaves 
the lips naked; it is very finely striated transversely. The 
cervical papille are situated about 250 ~ behind the level of the 
hind end of the muscular cesophagus, and the exeretory pore is 
situated about 100 « further back. 

The lips are large, and each bears two large outer papille. 
The external tooth is small, and the inner is of the same size, 
foliaceous, and tripartite ; no additional teeth were observed. 

The muscular cesophagus is markedly thinner than the rest of 
the organ, and forms a little less than 1/10th of its whole length. 
Vhe nerve ring encircles it in its posterior 1/4th. The esophagus 
forms in the female about 1/8th and in the male about 1/6th of 
the total body-length. 


Female. 
The length of the female varies from 35 to 43 mm., with a 
maximum breadth of 1:1 to 1:75 mm. 


1024 MR. R. J. ORTLEPP ON THE 


The body is attenuated in its anterior third, but posteriorly has 
a more or less uniform thickness; it is terminated by a short 
and obtusely rounded tail, forming about 1/100th of the total 
body-length. 

The vulva is slightly protuberant, and is situated at about the 
junction of the first and second quarters of the body. It leads 
into a coiled vagina 2 mm. long by 80 u thick; the ege-chamber 
is about 800 » long by 300 » thick and the common trunk about 
640 » long by 112 broad; its posterior end divides into two to 
give rise to the two uteri. 

The eggs are oval and thick-shelled, measuring 45 by 26 ph 
broad, and are fully embryonated in utero. 


Male. 

The two males were respectively 22 and 28 mm. long, with a 
thickness of 870 and 950. They are attenuated in their 
anterior half and the posterior end is curved ventralwards, 


Text-figure 9. 


lam. 


200 p. 


Physaloptera muris-brasiliensis Dies. 


A = Anterior extremity of male. 
B = Caudal extremity of male. 


The bursa is somewhat heart-shaped with a drawn-out tip, the 
lateral expansions being much better developed in their anterior 
half. Of the four pairs of pedunculated papille three pairs are 
pre-anal. There are six pairs of ventral paired papille and two 
unpaired papille; of these one pair ana an unpaired papilla are 
pre-anal, the latter being nearer the anus. The first and second 
pairs of post-anal papille are small and are situated in a row 
immediately behind the anus; pairs 3, 4, and 5 are equidistant 
from each other, and are situated respectively at the 2nd, 
3rd, and 4th sixths of the tail; they are all larger than the 
first two pairs. The unpaired post-anal papilla is about midway 
between the 2nd and 3rd pairs, and is about twice as large as the 
latter. 


NEMATODE GENUS PHYSALOPTERA. 1025 


The spicules are equal, straight, and pointed, each measuring 
400 x long by 32 p broad. 

Host: Mus brasiliensis. Stomach and small intestine. Brazil. 

Type males in bottle 4524, type females in bottle 4525, both 
in the Vienna Museum. 

Von Drasche, in his account of this species, considers that the 
number and arrangement of the bursal papille may be somewhat 
different, as the material he worked with was too opaque. By 
clearing the specimens in beechwood creosote, I was able to make 
out two additional pairs of ventral post-anal papille, viz. pairs 
2 and 4, as well as the nature and lengths of the spicules. 


(7) PuysatoprerRa ALATA Rudolphi, 1819. (Text-fig. 10.) 

Syn. Spiroptera physalura Duj., 1845. 

The material examined consisted of some males and females 
from Falco gallicus and some females from Falco pennatus; both 
lots from the Vienna Museum. 

These parasites have a somewhat slender body, whose cuticle 
is very finely striated transversely; the cuticle is either com- 
pletely refiected over the lips or else leaves them quite naked. 

The cervical papille are found from 75 to 100m behind the 
level of the hind end of the muscular cesophagus. The position 
of the excretory pore appears to vary in that it was found in 
some specimens to be very slightly posterior to the cervical 
papillz, whereas in others it was as much as 200 » further back. 

The lips are somewhat spherical in lateral view, and each 
carries two terminal teeth; the inner median tooth consists of 
three large denticles, and is larger than the outer tooth; the 
outer tooth varies in size: In some specimens it is very small, 
whereas in others it is almost as large as the inner tooth; a 
series of sizes can, however, be traced in different specimens, 
showing the transitions of the smaller to the larger sized outer 
teeth. Two somewhat slender external papille are present on 
each lip. 

The cesophagus is straight, and forms in the males about 1/6th 
and in the female 1/5-3 to 1/6°6 of the body-length. Its muscular 
part is thinner than the glandular, and is encircled by the nerve 
ring in its posterior quarter; it forms about 1/9th of the whole 
organ. 

Female. 

The females from F’, gallicus were all immature except one, 
and this specimen was 21 mm. long and 540 y thick; those from 
fF. pennatus were about 30 mm. long and 700 p thick, and were 
all mature; the body is attenuated in its anterior half and also 
slightly in its posterior third; in the immature forms the tail 
forms about 1/45th of the body-length, but in the mature forms 
it is slightly longer. 

The vulva opens practically level with the general surface, 
and is situated relatively far back ; its position varies slightly, 


1026 MR. R. J. ORTLEPP ON THE 


dividing the body into the ratio of 1: 1-07 tol: 1:4. The vagina 
is straight, and may either pass forwards or backwards; it is 
muscular, about 17 mm. long and 60, thick. The egg-chamber 
may pass forwards with the vagina or be bent backwards; it is 
about half the length of the vagina, and its breadth varies 
according to whether it is distended with eggs or not. The 
common trunk is short, and is nearly 300 long; its posterior 
end divides into two to give rise to the two uteri. 

The eggs are thick-shelled and oval, measuring on the average 
41 p long and 23 » in diameter. 


Male. 

The two males are respectively 17 and 18 mm. long, and 
are *5 mm. thick in their posterior third. The body is much 
attenuated in its anterior half. 


Text-figure 1¢. 


Physaloptera alata Rud. 


A & B= Ventral view of lips, showing variations in size of the external tooth. 
C = Caudal extremity of male. 


The bursa is relatively narrow and lanceolate in shape. There 
are five pairs of pedunculated lateral papilla, of which two pairs 
are pre-anal and the rest post-anal in position ; the anterior four 
pairs are equidistant from each other, whereas the distance 
separating the 4th and 5th pairs is about three times that 
separating the other pairs; this last pair is also more ventral in 
origin than the others. There are five pairs and an unpaired 
ventral papille; one pair and the unpaired papilla are situated 
in a triangle in front of the anus, the remaining four pairs being 
post-anal. The lst and 2nd post-anal pairs are small, and are 
found on the posterior margin of the anus; the 3rd pair is 
situated just behind the level of the 4th pair of stalked papille ; 


NEMATODE GENUS PHYSALOPTERA. 1027 


the 4th pair is at the middle of the tail, and it divides the 
distance from the last pair of pedunculated papille to the tip of 
the tail in about the ratio of 1:2. 

The ventral surface of the bursa is ornamented with longi- 
tudinal rows of tubercles extending slightly posterior to the last 
pair of stalked papillee. 

The spicules are short, equal, and pointed; the right, however, 
has a slightly stouter base; they are 275 long, the left spicule 
being 25 and the right 30 thick at its base. 

Hosts: Falco gallicus. Intestine. Museum, Vienna; bottle 

4439. 
Falco pennatus. Stomach. Museum, Vienna; bottle 
4449. 

Discussion.—This material differs from Schneider’s (1866), 
von Linstow’s (1877), and Seurat’s (1914 c¢) descriptions of this 
Species in the nature of the teeth and in the arrangement of the 
male bursal papille; these two characters are in keeping with 
Ph. subalata Schn., 1866, and Ph. galiniert Seurat, 1914. As 
the specimens examined consisted of the paratypes of Ph. alata, 
it appears to me that the conception of the species has been 
erroneous. Further, it would appear that Seurat’s species Ph. 
galiniert is either a synonym or a variety of Ph. alata, because 
it agrees with Rudolphi’s species in the nature of the teeth, the 
arrangement of the male bursal papille, the very posterior 
position of the vulva, and the anterior direction of the vagina ; it 
differs from Ph. alata in its relatively shorter esophagus and in 
its slightly longer and subequal spicules. 

Rudolphi deseribed this species from three hosts in the Vienna 
Museum, viz. Falco nisus, FP. pennatus, and F. gallicus. I take 
his specimens from the first-named host to be the types of this 
species, and the material from the other two hosts to be the para- 
types. 

An extracted description of Ph. galiniert Seurat is included in 
Part IT. 


(8) PHysALoprerA RETUSA Rudolphi, 1819. (Text-figs. 11 & 12.) 

Syn. Spiroptera retusa (Rud., 1819), Duj., 1845. 

I have been able to examine two lots of material of this species, 
both having been collected from the Teguexin. The first lot 
consisted of specimens deposited in the Vienna Museum (bottle 
4497), and the second I collected from a Teeuexin which died in 
the Gardens of the London Zoological Society. Both sets of 
material were in all respects identical. 

The cuticle is very finely striated transversely and, in addition, 
shows a coarse irregular ringing. Anteriorly it is partly or 
wholly reflected over the lips. 

The cervical papille are inserted generally symmetrically on 
either side, but sometimes one may be slightly more anterior to 
the other; they are lodged a short distance behind the junction 


1028 MR. R. J. ORTLEPP ON THE 


of the two csophageal parts. The excretory pore is situated 
about 50 u further back. 

The lips are large and somewhat triangular in side view, and 
each carries on its apex a large triangular tooth. Immediately 
internal to it there isa slightly larger but membranous tooth, 
whose free extremity is tripartite. No lateral teeth are present. 
Rach lip bears on its outer surface two conspicuous papille. 

The cesophagus is long, and in the female it is about 1/7th, in 
the male slightly more or slightly less than 1/5th of the body- 
length. Its anterior region is differentiated into a short muscular 
portion, about 400 » long in fully-grown specimens and slightly 
thinner than the rest of the esophagus; the large nerve ring 
encircles it in its posterior quarter. 


Text-figure 11, 


~ 


250 fi: 


Physaloptera retusa Rud. 
A = Ventral view of cephalic extremity. 
B = External aspect of median teeth. 
Female. 

The largest female measured 45 mm. long, and 1-2 mm. broad 
in the middle; fully mature females varied in size from 20 mm. 
upwards. The body is rather robust, tapering only slightly 
towards both ends; the tail, however, is sharply set off from the 
body, and is relatively short and slender; it forms on an average 
1/107th of the total body-length, and its caudal pores are situated 
in its posterior third. 

The vulva opens on a level with the general surface, and is 
situated some distance behind the end of the cesophagus. Its 
position varies slightly in different individuals, but in general it 
is seen at the junction of the Ist and 2nd quarters of the body. 
It leads in the largest females into a relatively short and thick- 


NEMATODE GENUS PHYSALOPTERA. 1029 


walled vagina, about 1-1 mm. long and 100 » in diameter. The 
following egg-chamber is about 3/4ths of the length of the vagina 
and about twice as broad. ‘The common trunk is very short, so 
that in some cases it appears to be absent; its posterior end 
divides into two branches, each of which connects up with one of 
the two uteri. 

The receptacula seminis are slightly oval, 380 « long by 280 » 
broad; they are distinctly constricted off from the oviducts, but 
their Junction with the uteri is very gradual. 

The eggs, which on the average are 43 p long and 27 pw in 
diameter, contain fully-developed embryos im wiero; they are 
thick-shelied and oval. 

Male. 

The males are much smaller than the females, the longest 

measuring 20 mm. long and 570 « broad. Anteriorly the body 


Text-figure 12. 


La 
ie) 
250 ws, 


Physaloptera retusa Rud. 
A = Left spicule. B = Right spicule. 
C = Caudal extremity of male. 
(The scale alongside B refers also to A.) 


tapers very gradually, but posteriorly hardly any narrowing 
takes place until a very short distance in front of the bursa; 
here the thickness becomes sharply less, so that at the juncture 
of the bursa it is only about 3/4ths of the thickness further 
forwards. 

The male bursa is large, measuring about 2°3 mm, long by 
1:1 mm. broad in the largest specimens, and is sharply reflected 
ventralwards. The cuticular expansions are well developed, and 
the four pairs of pedunculated papille supporting them are 
situated equidistant from each other, two pairs being pre-anal 
and two pairs post-anal. The ventral surface of the bursa is 
covered by numerous longitudinal rows of small protuberances. 

Proc. Zoou. Soc.—1922, No. LXIX. 69 


1030 MR. R. J. ORTLEPP ON THE 


There is one pair and an unpaired median ventral papille in 
front of the anus. ‘These areso placed as to form a triangle with 
its apex directed towards the anus. Behind the anus there are 
five pairs of ventral papille, of which the first two pairs are 
situated immediately behind the anus. The third pair is placed 
at the junction of the lst and 2nd quarters of the tail, whilst the 
Ath and 5th pairs are situated further back, and are slightly 
approximated. Hquidistant from these last two pairs there is 
seen what appears to be a very small papilla, but which probably 
is the opening of the caudal glands. 

The spicules are only very shghtly unequal, the right spicule 
being the longer and more slender. Some difficulty was en- 
countered in making out their lengths in cleared specimens 
because of their paleness, and consequently they were dissected 
out by means of a dissecting microscepe. The left spicule 
broadens posteriorly to about 90 » and then gradually tapers to a 
point; it is 455 p long, and 32 wu broad at its base. The right 
spicule, which is 475 » long, and 45 p broad at its base, tapers 
gradually to a point, and in all the males examined its posterior 
extremity was recurved. 

Host: Zupinambis tequexin. Stomach. Brazil. 

In addition to the material described above, I have also 
examined a few specimens from the mouth of a Hog-nosed Snake 
(Heterodon platyrhinus, S. America) which I have referred to the 
above species. They agree with Ph. retusa in practically all 
respects except in the size of the spicules, which in this material 
are equal in length and 300 w long. Their shapes, however, 
are identical with those described above. 


(9) PaysaLoprera MONODENS Molin, 1860. (Text-figs. 13 & 14.) 


The material on which the study of this species is based was 
collected from Boa constrictor and deposited in bottle 4459 in 
the Vienna Museum. 

The cuticle is finely striated and completely reflected over 
the lips. The cervical papillae vary in their position, in some 
cases being at the level of the hind end of the muscular wso- 
phagus, and in others as much as 300, further back. The 
execretory pore is situated about 400, behind the muscular 
cesophagus. 

The lips are dome-shaped, and each bears a small subdorsal 
and subventral external papilla. The two median teeth are of 
unequal size, the inner one being much smaller; the outer tooth 
is sharply conical, the inner foliaceous and tripartite; they are 
rather small. 

The muscular cesophagus forms about 1/10th of the whole organ, 
which is about 1/5°3 of the body-length in the male and 1/8th in 
the female. The nerve ring encircles its muscular portion in its 
posterior quarter. 


NEMATODE GENUS PHYSALOPTERA. 1031 


Female. 


Mature females vary in length from 25 to 40 mm., with a 
maximum thickness of 850 to 960; its greater portion is of 
more or less uniform thickness, tapering being confined only to 
the last few millimetres of the two extremities. The tail is 
obtuse, and forms about 1/75th of the body-length. 

The vulva is protuberant, and its position divides the body into 
the ratio of 1:5. It leads into a slightly coiled vagina 1°6 mm. 
long by 65, thick. The following egg-chamber is relatively 
short and thin, being 640 long and 130 in diameter. The 
common trunk is very short, being only 160 long and as thick 
as the vagina; its posterior end divides once to give rise to the 
connections of the two uteri. 

The receptacula seminis are oblong, 240 long and 175, in 
diameter. 

The eggs are thick-shelled, oval, and embryonated in utero; 
they measure 45 by 26 pw. 


Male. 


In the males the body is attenuated only in its anterior third ; 
the longer specimens varied from 18 to 23 mm. long by 719 to 
770 p thick. 


Text-figure 13. 


aaa 


Physaloptera monodens Mol. 
A = Cephalic extremity, lateral view. 
B= Cephalic extremity, ventral view. 


The bursa is ovoid, has well-developed ale, and its ventral 
surface shows longitudinal rows of small irregularly rounded 
tubercles. The three anterior pairs of stalked papille are pre-anal 
in position. Of the three pre-anal ventral papille the central 
one is the largest, and is situated closer to anus than the other 
two. There are five pairs of post-anal ventral papille, of which 
the first two pairs are small and situated in a row immediately 
behind the anus; the 3rd pair is slightly obliquely placed at 
about the junction of the Ist and 2nd sixths of the tail; the 
Ath pair is just in front of the middle of the tail. The 
distance separating the 4th from the 5th pair is slightly less 
than that between the 8rd and 4th pairs. The caudal pores 

69* 


1032 MR. R. J. ORTLEPP ON THE 
are conspicuous, and are found between the last two pairs of 
ventral papille. 

The spicules are unequal, and the left has its end broadened out 
to form a kind of spear-head; it is 415 « long and 38 p broad at 


Text-figure 14. 


Physaloptera monodens Mol. 


A = Caudal extremity of male. 
B = Right and left spicules. 


its base. The right spicule is shorter and thinner, and tapers to 
a fine point; it is 362 w long by 32 pu broad at its base. 
Host: Boa constrictor. Stomach and intestine. South America. 
Types in bottle 4459 in the Vienna Museum. 
Vor atlinities see Ph. obtusissima. 


(10) PuysALOPTERA oprusissImA Molin, 1860. (Text-figure 15.) 


The specimens of this species which were examined were 
deposited in bottles 4463 and 4470 in the Vienna Museum ; in 
both cases the host was simply given as Colubri, N. 58. 

The cuticle is slightly reflected over the base of the lips, and is 
provided with a fine transverse striation. The cervical papille 
pierce it from 130 to 225 » behind the level of the posterior end 
of the muscular ceesophagus, and the excretory pore is situated 
about 100 further back. 

The lips are hemispherical in lateral view and slightly flattened. 
The external tooth is large and bluntly conical, and is slightly 
bent outwards; the inner tooth is membranous, triparite, and of 
the same height as the outer. The two external lip papille are 
large. 

The cesophagus is straight and short; in the female it forms 


NEMATODE GENUS PHYSALOPTERA. 1033 


about 1/9th and in the male about 1/7th of the total body- 
length. Its muscular part is slightly thinner, and the nerve 
ring is situated in its last quarter. 


Female. 

The females are rather slender, mature worms varying from 
28 to 47 mm. long by 650 to 720 « thick in their posterior third. 
The body is much attenuated in its anterior half and in its posterior 
quarter. The tail is short and bluntly conical, and forms 1/80th 
of the body-length; its caudal pores are situated behind its 
middle, and their position divide the tail in the ratio of 3: 2. 

The vulva is situated on a slight elevation at the junction of 
the Ist and 2nd quarters of the body. It leads into a straight 
vagina 15 mm. long by 65 » thick; the following egg-chamber 
is oblong and 640 long by 175 broad; the common trunk is 


Text-figure 15. 


Physaloptera obtusissima Mol. 


Caudal extremity of male. 

short, and half again as thick as the vagina ; it is 320 » long, and 
its posterior end divides to give rise to two branches, each of 
which joins up with a uterus. The proximal parts of the uteri 
may be much convoluted in some individuals; posteriorly each 
terminates in a receptaculum seminis, which is long and oval, 
measuring 400 p long by 160 y broad. 

The eggs are fully embryonated im uwlero; they are oval and 
thick-shelled, and measure 45 by 26 x. 


Male. 

The males are comparatively stouter than the females; they 
vary in length from 20 to 32 mm., with a maximum thickness of 
700 to 720 4. The body is attenuated only in its anterior third. 

The caudal bursa is elongate and somewhat lanceolate in 
shape; its ale are well developed and its ventral papille are 


1034 MR. RB. J. ORLTLEPP ON THE 


large. The stalked papille are equidistant, two pairs being pre- 
anal and two pairs post-anal. Of the three pre-anal ventral 
papillee, the central one is large, and is situated nearer the rim 
of the anus than the other two. There are four post-anal sessile 
papille, of which the first two pairs are small and arranged in 
a row immediately behind the anus; the 3rd pair is slightly 
obliquely placed just behind the level of the last pair of stalked 
papille ; the 4th pair is at about the middle, and the last pair at 
the junction of the 2nd and last thirds of the tail. 

Tbe spicules are short and slightly unequal; the right is 385 p 


long by 88, broad at its base, and tapers to a fine ; point ; the. 


left is 430 p long by 38 broad at its base; its posterior half 
broadens out to about 74 p to form a comparatively large sigteit 
head. 

The ventral surface of the tail is traversed by longitudinal 
rows of irregular tubercles of the same size and shape as those 
ee in P. monodens. 

Host: Colubri, N.58. Brazil. 

This species appears to be closely allied to Ph. monodens, with 
which species it agrees in the arrangemeut of the caudal papille 
of the male, the general shape of the spicules, markings on the 
bursa, and size of the eggs. It differs from this species, however, 
in its larger teeth, more posterior position of the vulval aperture, 
shorter cesoph agus and larger receptacula seminis. 

Von Drasche (1883 3) states that the inner teeth are very small ; 
my observations showed that they were in nearly all cases of the 
same size as the outer teeth, and could be easily seen from either 
lateral or ventral (dorsal) views of the lips. 


(11) PHysALoprera BONNET, sp.n. (Text-figs. 16 & 17.) 


The material consisted of four females and two males, collected 
from a ‘“Sapakara,” Dutch Guiana; the females varied in length 
from 41 to 50mm., and the males are respectively 26 and 33 mm. 
long. 

The cuticle is transversely striated, the strie being about 
pe4 apart. In addition there is a very distinct and coarse 
ringing.. It is only very slightly reflected over the base of the 
lips. 

The cervical papille are situated some distance behind the 
junction of the two cesophageal parts, this distance being about 
half the length of the muscular esophagus. The excretory duct 
opens to the exterior about 150 » further back. 

The lips are somewhat flattened, and each has two outer 
papille. Only two teeth are present on each lip, an outer and 
an inner median tooth. They are of the same size, the outer 
being conical and obtuse, whereas the inner is “angi Dye ows and 
tripartite. 

The muscular cesophagus is only slightly thinner than the 
glandular, and the nerve ring is situated at the junction of its 


NEMATODE GENUS PHYSALOPTERA. 1035 


3rd and last quarters. The whole organ forms in the male 
1/5°6, in the female 1/7°6 of the body-length. 


Female. 

The female is slightly attenuated towards both extremities, 
and is terminated posteriorly by a slender tail, forming 1/56th part 
of the body. Its caudal pores are situated at the junction of its 
2nd and last thirds. The maximum breadth of the largest 
female is 1-3 mm. just behind its middle. 

The vulva is situated in a depression partly overhung by an 
anterior cushion-like thickening of the body ; it divides the body 
in the ratio 1:2°5. The vagina is coiled, and is 1:2 mm. long by 
95 pw thick; the following egg-chamber is about 1 mm. long and 
twice as broad as the vagina; its posterior end divides into two, 
and each branch joins up with a uterus, a common trunk being 
absent. The anterior half of the vagina, the egg-chamber, and 
_the terminal portions of the uteri are directed forwards, the 


Text-figure 16. 
\/ 


[ 


SOO py. 


Physaloptera bonnei, sp. n. 
A = Outer view of median teeth. 
B = Terminal portion of female genitalia. 
C = Receptaculum seminis. 


uteri extending as far as the base of the cesophagus, when they 
bend sharply backwards and pass almost to the posterior end of 
the body. The receptacula seminis are pear-shaped, and are 
A80 pw long by 265 uw broad at their junction with the oviduets. 

The eggs are elongate, oval, and thick-shelled, and contain fully- 
developed embryos; on the average they are 42m long by 
25 p broad. 


Male. 


The males are much thinner than the females, the diameters 
of the larger and smaller worms being respectively 950 and 850 p. 
The bursa is large, forming about 1/14th of the body-length, and 
is nearly twice as long as broad. Ventrally it is ornamented 
with longitudinal rows of cuticular elevations. The four pairs of 
pedunculated lateral papille are equidistant, two pairs being pre- 
anal and two pairs post-anal. The three pre-anal papille are 
situated in a row just above the anus. Behind the anus there 


1036 MR. R. J. ORTLEPP ON THE 


are five pairs of sessile papille, of which the first two pairs are 
small and inserted in a line just behind the anus; the 3rd pair 
is found at the level of the last part of pedunculated papille, the 
Ath pair in the middle of the tail, and the last just anterior to 
the front limit of the posterior third of the tail. 

The spicules are of equal length, but the base of the right is 
very slightly thicker than that of the left. They are 455 » long, 
the right being 29 u broad at its base and tapering gradually to 
a fine point, the left being 32 4 thick as its base and having its 
posterior end widened out to form a spear-head 160 long by 
95 » broad. 

Host: ‘‘Sapakara.” Stomach. Dutch Guiana. 

Types in the Helminthological Department of the London 
School of Tropical Medicine. 


Text-figure 17, 


Physaloptera bonnet, sp. n. 
A = Caudal extremity of male. B = Spicules. 


This species has many characters in common with Ph. obtusis- 
sima, as shown by the general shape, ornamentation and arrange- 
ment of the papille of the male bursa, the nature and size of the 
teeth, and the general shape of the spicules. It is, however, 
distinguished from it by its longer cesophagus, slightly more 
posterior position of the vulva, the presence of a cushion-like 
swelling anterior to the vulva, the absence of a trunk portion 
behind the egg-chamber, and in that the spear-head of the left 
spicule is better set off. 

I have named this species after Dr. Bonne, of Surinam, who 
collected and presented this material to Professor Leiper. 


NEMATODE GENUS PHYSALOPTERA. 1037 


(12) PHYSALOPYERA PHRYNOSOMA, sp. n. (Text-figure 18.) 


Numerous specimens were collected from several lizards which 
had died in the Gardens of the London Zoological Society ; 
unfortunately only a few of the females are mature. 

The parasites are slender and much attenuated anteriorly ; 
their greatest diameter is in their posterior third. 

The cuticle is smooth (?), and is not reflected over the lips. 
The cervical papille are situated just in front of the level of 
the junction of the two esophageal parts, and the excretory pore 
from 50 to 100 « behind them on the ventral surface. 

The two lips are somewhat conical, and each is surmounted by 
a large conical tooth, which is inclined slightly outwards; no 
other. teeth are present. Hach lip has two papille on its outer 
surface. 

The csophagus is straight, and varies in the females from 1/6th 
to 1/7th, and in the male it is slightly less than 1/5th of the 
total body-length ; its anterior muscular part passes gradually 
into the following glandular part, there being little difference in 
the diameters of the two. The nerve ring encircles its anterior 
part just in front of the junction of its third and last quarters. 


Female. 

Fully mature females vary in length from 18 to 23 mm., with 
breadth from 610 to 760 uw in their posterior third. The genital 
aperture is non-protuberant, and is situated more or less at the 
junction of the first and second body thirds; its position varies 
slightly in different individuals, being sometimes slightly in front of 
or shightly behind this level. It leads intoa short and thick-walled 
vagina, 380 w long by 38 in diameter ; this in turn leads into a 
well-developed egg-chamber measuring 950, long by 152 
broad; the posterior end of this chamber narrows suddenly and 
passes into a short common trunk, which after a very short 
traject divides into the two uteri; the uteri pass parallel to each 
other into the posterior sixth of the body. 

The eggs are oval and thick-shelled ; they measure 50 p long 
by 36 p in diameter. 

The receptacula seminis are small and oval, measuring 133 u 
by 86 w. 

The tail is relatively long and slender, forming 1/36th of the 
total length ; its caudal pores are situated just behind its middle. 


Mate. 

The males are much slenderer than the females, their average 
length being 11 mm. by 470 » broad. The caudal bursa has well- 
developed ale, and its ventral surface is covered with longitudinal 
rows of small tubercles. Iwo pairs of the stalked papille are 

- pre-anal, and two pairs are post-anal in position; they are more 
or less equidistant from each other, but the pre-anal pair nearest 
the anus is slightly more dorsal in origin than the others. There 
are three ventral pre-anal papille, the median unpaired one being 
situated nearer the anus than the other two. IJmmediately 


1038 MR. R. J. ORTLEPP ON THE 


posterior to the anus there are two pairs of small papillee close 
together, the one pair being slightly behind the other pair ; 
a small distance behind these there is another pair somewhat 
ebliquely placed and also larger; posterior to the middle of the 
tail there are two further pairs dividing this part of the tail 
into thirds. 

The spicules are unequal, the right being slightly stouter than 
the left; both taper to fine points. The right spicule is 1804 
long and 10 p broad at its base, while the left spicule is 530 p 
long and 9 « broad at its base. 

Hosts: Phrynosoma cornutum, P. regale. Stomach. Brazil. 


Types to be deposited in British Museum of Natural History, 
London. ; 


Text-figure 18. 


en eaten 
ZOO fs. 
Physaloptera phrynosoma, sp. 0. 
A = Ventral view of cephalic extremity. B = Caudal extremity of male. 


Affinities.--Ph. abbreviata has been recorded from this host, 
but it seems doubtful whether the determination was correct ; 
this species was present in nearly every one of the specimens of 
Phrynosome examined by me, which numbered about a dozen, 
and was the only species present; it is therefore possible that 
it was this species which was mistaken for Ph. abbreviata. As 
redescribed by Seurat (19146 and 1917a), Ph. abbreviata is readily 
distinguished from Ph. phrynosoma by the fact that it has four 
uteri, whereas the latter has only two. 

The presence of two uteri and a single tooth to each lip allies 
this species to Ph. gracilis, sp.n., and to Ph. longissima, sp. 0. 5 
it is, however, easily separated from these two species by its 
much shorter and relatively stouter body. 


NEMATODE GENUS PHYSALOPTERA. 1039 


(13) PHYSALOPTERA GRACILIS, sp. n. (Text-fig. 19.) 


The material consisted of about half-a-dozen specimens, all of 
which were coiled up watch-spring like ; as no mature eggs were 
observed, it is probable that the worms, although fully developed, 
had not yet reached the reproductive stage. 

The cuticle shows a very fine transverse cuticular striation, 
and it may be partly or wholly reflected over the lips. 

The cervical papille are situated at the level of the junction of 
the two esophageal parts or just posterior to this; the excretory 
pore is situated on the ventral surface about 40 » further back. 

The lips are simple and triangular in side view, and each is 
surmounted by a large external tooth, triangular in shape, and 
having a spike-like internal tooth attached to its inner surface ; 
a row of small denticles is present on either side of it, each row 
being terminated laterally by two larger denticles. External 
lip papille were not observed. 

The cesophagus is slender, and retracted from the lips, so that 
an elongate chamber is formed between it and the lips. The 
muscular part is slightly thinner than the following glandular 
part, and forms in the male 1/12th and in the female 1/14th 
of the whole organ. The whole esophagus forms in the male 
about 1/7th and in the female 1/6°2 of the total length of the 
body. The nerve ring divides the muscular cesophagus in the 
ratio of 3: 2. 


Female. 

The females vary in length from 20 to 24 mm., and in breadth 
from 240 to 250; the body is attenuated towards both 
extremities, but this is only slightly evident posteriorly. The 
tail is short and pointed, and forms 1/97th part of the total 
length. 

The vulva is non-protuberant, and leads into a straight vagina 
410 long by 45 broad; the egg-chamber which follows is 
about as wide again as the vagina, and is 730 4 long; the common 
trunk has more or less the same dimensions as the vagina ; from 
its posterior end the two uteri take their origin, and these pass 
down the body more or less parallel to each other. The position 
of the vulva divides the body into the ratio of 1: 2. 

No mature eggs were observed. 


Male. 


The males average about 19mm. in length by 230 broad ; 
they are more slender than the females, and do not become atten- 
uated towards the posterior end. 

The bursa is small, forming 1/36th of the body-length ; it is 
nearly 24 times as long as it is broad, and its width is only very 
slightly greater than the maximum body-breadth. Its ventral 
surface is free from cuticular protuberances, except for a small 
area surrounding the anus. 

The lateral stalked papille are arranged three pairs pre-anal 


1040 MR. R. J. ORLTLEPP ON LHE 


and one pair post-anal; the lst pair is more ventral in origin 
than the others, and is also closely approximated to the 2nd 
pair; the remaining pairs are equidistant from each other. 
There are three pre-anal and four pairs of post-anal ventral 
papillee ; the Ist pair of post-anal papille are situated immediately _ 
behind the anus, the 2nd and 3rd pairs are near together just 
behind the middle of the tail, and the last pair midway between 
the 3rd pair and the tip of the tail. 

The spicules are unequal, the right being short and _ stout, 
measuring 105 long by 23 broad, the left being slender, 
measuring 273 » long by 7 pe broad at its base; the right spicule 
tapers to an obtuse point, whereas that of the left side forms a 
very fine point. 

Hest: “ hizard.’”?” Uganda. 

Types in the Helminthological Department of the London 
School of Tropical Medicine. 


Text-figure 19. 


Cla 


8 


Physaloptera gracilis, sp. n. 


A = Cephalic extremity, lateral view. B = Ventral view of lips. 
C = Inner view of lip. D = Caudal extremity of male. 


Discussion.-—The arrangement of the bursal papille in the 
male, the absence of cuticular markings on the greater part of 
ventral surface of the caudal extremity of the male, and the 
character of the lips and its teeth—each of these distinguish this 
species from all the other reptilian didelphoid forms. 

There is some similarity in the arrangement of the bursal 
papille of this species to that found in Ph. leptosoma (Gervais), 
Seurat, 1917, except that just behind the anus there are two 
pairs in this latter species instead of one. Ph. gracilis, however, 
is easily distinguished from this species by the additional 
characters of a more elongate body and of much shorter spicules. 


NEMATODE GENUS PHYSALOPTERA. 1041 


(14) PHYSALOPTERA LONGISSIMA, sp. n. (Text-fig. 20.) 


The material consisted of about two dozen specimens, some of 
which were still immature. The specimens are remarkable for 
their slenderness, the largest females having a thickness of less 
than 500. The body shows very little attenuation towards the 
extremities. 

The cuticle is finely striated transversely, and in some cases it 
forms a small swelling round the head, whereas in others it is 
reflected over the lips. 

The cervical papillz and excretory pore occupy a very anterior 
position, the former being found at the level of the nerve ring, 
and the latter half-way between the nerve ring and the level of 
the base of the muscular cesophagus. 


Text-figure 20. 


250 pe: 


Physaloptera longissima, sp. 0. 
A = Ventral view of cephalic extremity. 
B = Caudal extremity of male. 


The lips are rounded, and each carries only a single tooth, 
namely the external median ; this tooth is triangular in shape, 
and has its tip slightly recurved outwards. Hach lip carries two 
outer papillee. 

There is no difference in diameter in the two esophageal 
parts, the two merging gradually into each other; the whole 
organ attains 1/7°6 in the female and 1/6°6 in the male of 
the body-length; the muscular part, which is encircled in its 
middle by the nerve ring, forms nearly 1/8th of the whole 
organ. 


Female. 


The largest females are 30 mm. long by 438, broad, and 
possess a vulva which is situated just in front of the middle of 


1042 MR. R. J. ORTLEPP ON THE 


the body, dividing it in the ratio of 7:8; it leads into a straight 
vagina 1-3 mm. long, which is directed posteriorly ; it then turns 
forwards to join the trunk, which in its middle bends back again ; 
it is 3-5 mm., long and its anterior half is only slightly differ- 
entiated into an egg-chamber; the posterior end of the common 
trunk divides to give rise to the two uteri. The uteri pass 
posteriorly almost to the hind end of the body, then pass 
forwards before joining the oviducts and ovaries, which in their 
turn pass backwards again. 

The body is terminated by a short and rounded tail forming 
nearly 1/190th part of the body-length. 

The eggs are long, oval, and thick-shelled, measuring 59 p long, 
32 » broad, the thickness of the shell being 5 p. 


Male. 


The males average about 22 mm. long by 325 pw broad in their 
middle. 

The bursa is not sharply set off from the body, and is only 
slightly broader than it; it forms about 1/28th of the body- 
leneth ; it is little more than twice as long as it is broad, and its 
ventral surface is devoid of cuticular markings. 

The four pairs of stalked papille are all pre-anal in position, 
and arranged in two widely-separated groups of two pairs each. 
No ventral pre-anal papille were seen. Behind the anus there 
are four pairs of shortly stalked papille, which are equidistant 
from each other and from the anus. They are situated somewhat 
laterally. 

The spicules are unequal, the left being nearly twice as long as 
the right; the left spicule is slender, and tapers to a fine point; 
it is 546 long by 22 broad at its base; the right spicule is 
from 228 to 300 w long by 52 w broad at its base; near its base 1b 
has a distinet neck, after which it widens out and then tapers to 
an obtuse point. 

Host: “Snakes.” Australia. 

Types in the Helminthological Department of the London 
School of Tropical Medicine. 

Discussion.—This species 1s easily separated from all the 
didelphoid Physaloptera (1) by the arrangement of the stalked 
bursal papille, these being entirely pre-anal and arranged in two 
widely-separated groups in this species; (2) by the arrangement 
and number of the post-anal ventral papille; and (3) by the 
entire absence of cuticular markings on the ventral surface of 
the tail of the male. 

The elongate and slender body shows some similarity to 
Ph. gracilis, sp. u., from which species it is, however, quite 
distinet, as shown by the above-named three characters; it 
also differs from this species by the difference in shape of its lips 
and teeth, by the more posterior position of its vulva, and by 
its larger spicules. 


NEMATODE GENUS PHYSALOPTERA. 1043 


| (15) PHYSALOPTERA PREPUTIALIS von Linstow, 1889. (Text- 
figs. 21 & 22.) 
Syn. Chlamydonema felineus Hegt, 1910. 


Several lots of material were examined, mostly from cats. 

The body is slightly attenuated anteriorly, and is surrounded 
posteriorly by a very loose cuticle, which is partly or wholly 
reflected over the tail region. The cuticle shows a very fine 
transverse cuticular striation, and is only slightly reflected over 
the lips. 

The excretory pore and cervical papille are situated relatively 
far back, the former being more posterior in position than the 
latter. The distance from the anterior end to the cervical 


Text-figure 21. 


Physaloptera preputialis v. Linst. 


A = Lateral view of cephalic extremity. 
B = Caudal extremity of male. 


papilla in a male 15 mm. long is 875 « and the excretory pore 
990 4; in a female 17 mm. long these distances were 952 uw and 
1123 uw respectively. 

The lips are large and conical, and each carries a subdorsal 
and subventral external papilla. The external median tooth is 
large and bluntly triangular; the inner median tooth is of the 
same height as the outer, and is flattened and tripartite. 

The cesophagus is long, and varies in the female from 1/4:8 to 
1/5:4 of the body-length ; in the male it forms about 1/4°5 of the 
body-length. ‘Che muscular portion of the esophagus is slightly 
thinner than the glandular, and is encircled by the nerve ring at 
the junction of its second and last thirds. It forms about 1/8th 
of the whole cesophagus. 


NOAA MR. R. J. ORTLEPP ON THE 


Female. 


The females are generally large, and vary in length from 15 to 
48 mm. with a maximum thickness of 1 to 1‘7 mm. in their 
posterior half. 

The vulva opens a short distance in front of the middle of the 
body. It is non-protuberant, and its position divides the body 
into the ratio of 1:1°4 to 1:1:l. It leads into a thick-walled, 
straight or bent vagina 1-6 mm. long by 114 pw thick in a female 
20 mm. long; its posterior part gradually passes into the egg- 
chamber, which broadens out posteriorly so as to assume a more 
or less pyriform shape: in the same female it was 820 pu long by 
228 » thick at its posterior end. Two uteri take their origin 
from the posterior margin of the egg-chamber, a trunk portion 
being entirely unrepresented. ‘The mode of origin of the uteri is 
very characteristic, and differs from all those already described ; 
they arise from the posterior lateral margins of the egg-chamber, 
and not from its base. It thus happens that the uteri, 
although arising at the same level, are yet far removed from 
each other. 

In most individuals a dark and detachable chitinous ring 
surrounds the body in the region of the vulva. 

The tail is conical and rounded at its tip, and forms from 
1/58rd to 1/66th part of the body-length. Its caudal pores open 
in its posterior half, and their position divides the tail into the 
ratio of 3: 1. 

The eggs are oval and thick-shelled, and contain a fully- 
developed embryo; they average 49 » long by 35 w in diameter. 


Male. 


The length of the males varies from 13 to 40 mm. by °7 to 
1:3 mm. thick in their posterior third. 

The tail is elongate, and is often closely reflexed on to the 
ventral surface. its median ventral surface 1s ornamented with 
conspicuous and rounded tubercles arranged longitudinally ; 
toward the lateral margins of the tail and on to the ale 
these tubercles become gradually replaced by longitudinal ridges, 
each having a few relatively far removed breaks along its 
course, 

The caudal region, within the cuticular reduplication, appears 
pointed; this 1s due to the fact that the caudal ale, although 
present, do not open out laterally, but remain irregularly folded 
towards the ventral surface of the tail. The fours pairs of 
stalked papillae are thick and equidistant from each other, two 
pairs being pre-anal and two pairs post-anal. The three pre- 
anal ventral papille are in a transverse row, and the median one 
is much larger than the other two. Just behind the anus, one 
pair immediately following the other, there are three pairs of 
ventral papille. Two additional pairs of ventral papille are 
found further down the tail, and their positions divide the tail 
roughly into thirds. 


NEMATODE GENUS PHYSALOPTERA. 1045 


The spicules are unequal, pointed, and generally curved. The 
left spicule appears to vary in length from 1 to 1:2 mm., with a 
thickness of 32 at its base. The right spicule is slightly 
stouter than the left; its length also appears to vary from 840 to 
900 « with a thickness of 38 p» at its base. 

Hosts: felis catus domesticus. Stomach. Federated Malay 

States, British and Dutch Guiana, China and 
Ceylon. 

Felis nebulosa. India. 

Felis pardus. Stomach. Nigeria. 

Although my observations as regards the arrangement of the 
caudal papiile in the male, the position of the vulva in the 
female, and the size of the eggs differ considerably from those of 
von Listow, yet I am quite satisfied that my material belongs to 
the same species. 


Text-figure 22. 


Physaloptera preputialis vy. Linst. 


Terminal portions of female genitalia. 


The prepuce-like fold of the cuticle over the tail, the rounded 
wart-like tubercles on the ventral surface of the tail of the male, 
the nature of the teeth, the presence of a ring round the body in 
the vulval region of the female, and the fact that his material is 
described from /elis catus, easily counterbalanees the differences 
in the arrangment of the bursal papille. Von Linstow states 
that this arrangement may be found to be different, as he had 
only one male for examination, and this specimen he found 
difficult to study. The ventral papille, according to von Linstow, 
are three pre-anal, one pair immediately behind the anus, three 
papille towards the tail end, and an additional pair slightly in 
front of these three. He gives the average length of the female 
as 30 mm., with the vulva 8 mm. from the cephalic extremity 
and the eggs as 55 by 33 p. 

Proc. Zoo. Soc.—1922, No. LXX. 70 


1046 MR. R. J. ORTLEPP ON THE 


(16) PHysALOPTERA MALAYENSIS, sp.n. (Text-fig. 23.) 


In external appearance this species appears to be identical 
with Ph. preputialis v. L., with which species it was at first 
confused. This applies especially to the female, where there is a 
corresponding similarity in the genitalia. 

The cuticle is very slightly reflected over the base of the lips, 
and in most cases the lips are quite naked. The cervical papille 
are situated about 130 behind the level of the hind end of the 
muscular csophagus, and the excretory pore is found about 100 p 
further back. 

The lips are rounded, and each bears a large triangular and 
slightly recurved outer tooth, internal to which is the foliaceous 
and tripartite inner tooth of equal height. 

The csophagus forms 1/5th in the male and 1/6th in the 
female of the total length; its anterior muscular part, which is 
encircled in its posterior third by the nerve ring, is slightly 
thinner than the following part, and forms nearly 1/10th in the 
female and 1/7th in the male of the total organ. 


Female. 

Mature females vary in length from 29 to 38 mm., with 
breadth of from 1:6 to 2°1 mm. ‘The body is attenuated in its 
anterior third, but posteriorly, because of the looseness of the 
cuticle and its reflection over the tail, there appears to be only a 
very slight thinning. 

The vulva opens on a slight bulging, and is situated far back, 
just in front of the middle of the body; its position divides the 
body in the ratio of 1:1:2 to 1:15. Externally the position 
of the vulva is indicated by a dark chitinous and detachable band 
encircling the body. The vulva leads into a vagina, which pro- 
gressively thickens posteriorly to join a progressively thickening 
egg-chamber ; at its vulvular end the vagina is 100 p thick, at 
its posterior end it is 200 w thick, and the hind end of the egg- 
chamber is 330 u in diameter; the whole organ is straight, and 
measures about 3°3 mm. long. From the basal end of this organ 
the two uteri take their origin like two horns in the same 
manner as in Ph, preputialis. 

The eggs are small, thick-shelled, and only slightly ovoid ; 
they are 35 » long by 28 to 32 » broad. 

Male. 

The males are attenuated in their anterior half, and have in 
most cases the cuticle completely reflected over the tail end; 
they average in length from 18 to 21 mm., with a maximum 
breadth of 0-9 to 1:2 mm. 

In consequence of the cuticle being reflected over the tail, the 
bursal expansions are irregularly folded over the ventral surface; 
they are supported by the usual four pairs of circumcloacal 
stalked papillae. The whole of the ventral surface is traversed 
by well-marked and unbroken longitudinal ridges running more 
or less parallel to each other. The three pre-anal ventral papille 


NEMATODE GENUS PHYSALOPTERA. 1047 


are arranged in a row, and the central one is the largest. There 
are five pairs of ventral post-anal papille; pairs 1 and 2 are 
small, and arranged in a line immediately behind the anus. The 
3rd and 4th pairs are approximated to each other just behind 
the middle of the tail, and the last pair is at about the junction 
of the second and last thirds of the tail. 

The spicules are unequal, that of the left side being about 
22 times as long as that of the right side; their lengths appear 
to vary in different individuals, but this may probably be due to 
the fact that the spicules do not lie straight in the body, but 
often have a wavy course; it is thus difficult to measure them 
accurately ; in this way the left spicule was found to vary from 
1-4 to 2°5 mm. in length, with a thickness of 32 at its base; 


Text-figure 23. 


Physaloptera malayensis, sp. n. 


A = Lateral view of cephalic extremity. B = Outer view of median teeth. 
C = Caudal extremity of male. 


the right spicule is slightly stouter, and varies in length from 
580 to 957», with a breadth of 36 at its base. Aithough 
there is this marked variation in the sizes, yet 1t is worthy of 
note that the ratio of the two spicules was nearly the same in all 
eases. Unfortunately there was not sufficient material to allow 
for the dissecting out of the spicules in a series, in order to 
determine whether this variation was only apparent or real. 
Hosts: Melis chaus. Intestine. Federated Malay States. 
Felis tigris. Stomach. Federated Malay States. 


“Tiger cat.” a Federated Malay States. 
S Bush) catty ; Nigeria. 


92 
“ Hyena.” Prob. 1. striata. Stomach. Nigeria. 
AOS 


1048 MR. R. J. ORLTLEPP ON THE 


Types in the Helminthological Department of the London 
School of Tropical Medicine. 

Affinities. —This species, because of its labial armature and 
female genitalia, forms a distinct group with Ph. preputialis and 
Ph. acuticauda; to the former it is very closely related, as shown 
by the similarity of the females and the characteristic reflection 
of the cuticle over the caudal end in both species ; it differs from 
it, however, by the arrangement of its post-anal ventral papille, 
its larger spicules, and by the presence of unbroken ridges on the 
ventral surface of the bursa. 

‘The arrangement of the papille on the caudal extremity of the 
male is very similar to that found in Ph. terdentata. Apart from 
this characteristic, and that these two species are both parasitic 
in carnivores, there appears to be no close affinity between them. 
Ph. malayensis differs trom Ph. terdentata by the shape and size 
of its teeth, leneth of its spicules, presence of unbroken ridges on 
the male bursa, and in that the cuticle is reflected over the caudal 
extremity. 

(17) Paysanoprera TeRDENTATA Molin, 1860. (Text-figs. 24 
& 25.) 

Three bottles of material, Y 1074 from Velis concolor, 4511 
from Felis tigrina, and 4513 from Felis sp., labelled &h, ter- 
dentata and deposited in the Vienna Museum, were examined. 
Bottle Y 1074 contained two males, one of which had its head 
missing. Bottle 4511 contained one female. and bottle 4513 
contained one male. The male in the last-named bottle proved 
to be not Ph. terdentata but Ph. preputials v. Linst. The 


Text-figure 24, 


Physaloptera terdentata Mol. 
Outer view of lip. 


female from bottle 4511 had all the characters of the female of 
Ph. preputialis, except that the cuticle was not reflected over the 
caudal end, and consequently I am inclined to think that it also 
is Ph. preputialis. The material from Felis concoler proved to 
be Ph. terdentata, and the following description of the male is 
based on these two specimens, 

The complete male is 14 mm. long, and 400 thick in its 
posterior third. The body is attenuated anteriorly, and the 
cuticle is slightly reflected over the lips; externally it shows a 
coarse annulation, between which a fine transverse striation is 


NEMATODE GENUS PHYSALOPTERA. 1049 


visible under high magnification. The cervical papille are 
lodged about 100 behind the posterior limit of the muscular 
cesoph: agus, and the excretory pore is situated slightly, posterior 
to them. 

The lips are iarge and flatly rounded, and each carries two 
large and spherical outer papille—one in the subdorsal and one 
in the subventral line. The two terminal teeth are well-defined, 
the outer being short and stumpy, the inner large and tripartite. 

The cesophagus is straight, and consists of the usual two parts ; 
it is 2°3 mm. long or about 1/6th of the total body-length. Its 
anterior glandular part is slightly thinner than the part following, 
is 240 pu ‘long, and has the nerve ring situated in its posterior 
quarter. 

The bursa is elongate with rounded tip, and forms nearly 


Text-figure 25. 


250 ys. 


Physaloptera terdentata Mol. 
Caudal extremity of male, 


1/15th of the body-length. Its lateral ale are supported by four 
pairs of equidistant and somewhat slender stalked papillee, and are 
arranged two pairs pre-anal and two pairs post-anal in position. 
Its ventral surface has longitudinal rows of irregular cuticular 
bosses. The pre-anal ventral papille are situated close together 
in a row just above the anus, and the middle one is slightly 
larger than the other two. ‘The post-anal ventral papille consist 
of five pairs, of which pairs | and 2 are small and arranged 
in a row just posterior to the anus, pairs 3 and 4 are situated 
close behind each other in about the middle of the tail, and 
pair 5 is found at the junction of third and last quarters 
of the tail. 


The spicules are , only shghtly subequal, the left being 320 u 


1050 MR. R. J. ORTLEPP ON THE 


and the right 305 « long; they are of equal thickness, 32, and 
they possess a more or less uniform thickness in their anterior 
four-fifth, the last fifth tapering to a sharp point. 


Female. 

The female from Felis tigrina is 47 mm. long ; teeth both large 
and of equal size, the outer triangular; cesophagus 1/7°3 of body- 
length, vulva situated 18°5 mm. from anterior end, the female 
genitalia as for Ph. preputialis, and the cuticle is not reflected 
over the tail. 

Host: Melis concolor. Stomach, Brazil. 

Type males in bottle 1074 (number on stopper) in the Vienna 
Museum. 

With regard to the name of the host, the legend on the bottle 
was somewhat indistinct and appeared somewhat like /elis 
casiolaris, but as no feline of this name is known, I take it to 
mean /elis concoloris. 

I have placed this species among the didelphoid forms, because 
the female described may belong to this species, in which case 
it would come into the group characterised by Ph. preputialis 
v. Linst. 


(18) PuysaLoprera ACUTICAUDA Molin, 1860. (Text-fig. 26.) 

The material on which the study of this species is based 1s the 
type material deposited in the Vienna Museum. This material 
has also been studied by von Drasche. Ail the specimens were 
in an excellent state of preservation. 

The cuticle is very finely striated transversely, and anteriorly 
is partly reflected over the lips. In each lateral line, just behind 
the level of the posterior end of the muscular esophagus, a small 
and spike-like cervical papilla is situated, and on the ventral 
surface about 100 further back is the opening of the excretory 
organ. 

The lips are somewhat rounded in lateral view, and each 
carries only two teeth, a large and triangular outer tooth, and 
immediately internal to it a smaller membranous tooth having 
its free end tripartite. In the subdorsal and subventral lines of 
each lip there is a conical papilla. 

The esophagus, which immediately follows the ie has its 
anterior muscular part slightly thinner than the following 
glandular portion: in the male it forms about 1/5:1 of the total 
body. length and in the female about 1/7th. The glandular part 
forms about 1/7th of the cesophagus, and is surrounded by the 
nerve ring at the level of its posterior quarter. 


Female. 

The females have a length of about 30 mm., with a maximum 
breadth of just over 1 mm. ‘The body tapers in its anterior half, 
but posteriorly it narrows only in the vicinity of the tail, which 
is a conical structure about 1/56th of the total body-length, and 
has its caudal pores situated in its posterior third. 


~ 


we 


NEMATODE GENUS PHYSALOPTERA. 1051 


The vulva is situated very far forwards, opening to the exterior 
on a slight elevation in front of the hind end of the esophagus. 
The muscular vagina is very short, measuring only about 480 p 
long by 50 broad; its hind end is very slightly enlarged, and 
from its outer sides the two uteri take their origin in the same 
manner as in Ph. preputialis y. Linst.; the thin initial portions 
of the uteri are swollen in their middles, after which they again 
become thin and also much convoluted, after which they join up 
with the much thicker and egg-containing part of the uteri. 

The eges are large, oval, and thick-shelled, measuring 51 pu 
long by 42 » bread; they are fully embryonated wm utero. 


Male. 
The males are much smaller and slenderer than the females, 
measuring only 18 to 23 mm. long by 620 to 740m thick in 


: 


A = Terminal portions of female genitalia. 
B = Caudal extremity of male. 


Text-figure 26. 


Physaloptera acuticauda Mol. 


their posterior third. From about the posterior third the body 
tapers gradually towards the anterior end. 

The caudal bursa has well-developed lateral wings supported by 
four equidistant, pedunculated and paired papille, two pre- and 
two post-anal. Or the three pre-anal ventral papille the median 
one is lodged nearer the anus than the other two, and is also 
larger. There are five pairs of ventral post-anal papille, of 
which the Ist and 2nd pairs are situated in a row imme- 
diately behind the anus. The distance between the 4th and 
last pairs is about twice the distance between the 2nd and 
ord pairs. 


1052 MR. R. J, ORTLEPP ON THE 


The spicules are unequal, tubular, and end in sharp points ; 
that of the left side measures 1°89 to 2°17 mm. long by 45 yw broad 
at its base, that of the right 420 to 490 p long by 50 p broad. 

Host: Falco cachinnans. Csophagus and stomach. Brazil. 

Type material in bottle 4431 in the Vienna Museum. 

This species occupies an isolated position among the Physa- 
loptera recorded from birds; none of these in which the nature 
of female genitalia is known have the uteri arising from the 
margin of the egg-chamber in a way similar to that seen in 
Ph. acuticauda. This peculiarity of the female genitalia allies 
Ph. acuticauda to Ph. preputialis vy. Linst. and to Ph. malayensis, 
sp. n., from which species it is, however, very easily distinguished 
by the very anterior position of its vulva, different arrangement 
of the male caudal papille, and difference of the ornamentations 
on the ventral surface of the male tail, 


Group Tetradelphys. 
(19) PHysaLopreRA MORDENS Leiper, 1908. (Text-figs. 27 & 28.) 


For the study of this species, Prof. Leiper kindly placed at my 
disposal the type males (three specimens) and also two lots of 
worms collected by Drs. Turner from man in Africa. In addition, 
six tubes of worms collected by Dr. Davy from African monkeys 
were also placed at my disposal for comparison and identification ; 
all these proved to be the same species as that collected from man. 

The cuticle shows a very delicate transverse striation ; in some 
specimens an additional coarse and irregular ringing, probably 
due to a certain amount of shrinkage, is present. It is partly 
reflexed over the lips in some, whereas in others it stops short at 
the base of the lips. It is somewhat inflated, so that the cervical 
papille appear to be lodged in shallow cuticular pits. The 
cervical papille are situated slightly less than three-quarters of 
the length of the muscular esophagus behind the posterior limit 
of this esophageal part. The excretory pore opens ventrally 
from 50 to 100 » further back. 

The two lateral lips are large and rounded in lateral view, and 
are sharply set off from the body in those worms where the 
cuticle is not reflected over their bases. Hach lip carries a large 
dome-shaped external papilla on each submediun line. Four 
teeth are present on each lip, one outer and three inner; the 
outer tooth is large and triangular, and has its tip slightly 
rounded and recurved outwards. The median imner tooth is 
very small, and is attached to the base of the outer tooth; it is in 
the form of a small spike. Leiper states that its inner surface 
is modified to form a cutting-edge; to me this surface appeared 
quite flat, or at most slightly convex inwardly. The two inner 
lateral teeth are situated in the submedian lines, and are directed 
inwards ; each is split longitudinally to its base, and is lodged in 
a slight elevation of the lip. 


NEMATODE GENUS PHYSALOPTERA. ~ 1053 


The cesophagus is straight and immediately follows the lips; it 
thickens very slightly towards the posterior end. Its anterior 
1/10th, which is slightly thinner than the rest, forms its glandular 
part, and the nerve ring encircles it in its posterior third. The 
entire organ forms 1/6°4 in the male, and 1/6-2 in the female, 
of the total body-length. 


Female. 

The females are large and stout; they vary in length from 41 
to 100 mm., with a thickness of 1-8 to 2°8 mm. in their posterior 
third. The anterior two-thirds taper gradually towards the 
cephalic extremity ; the posterior 1/8th tapers to end in a short 
and conical tail 1/70th to 1/90th of the body-length ; the caudal 
pores are situated in its sont ior half. 

The vulva is situated on a slight elevation behind the hind 
end of the esophagus; its position divides the body into the 
ratio of 1:5. The general characters of the female genitalia are 
exactly similar to those described for Ph. varani, except that 
the common trunk portion behind the egg-chamber is relatively 


Text-figure 27. 


Physaloptera mordens Leiper. 


A = Ventral view of lips. B = Inner view of lip. 


larger. The four uteri are twisted about each other in a com- 
plicated way, or fill practically the whole of the body as far as its 
posterior eighth. 

The eggs are oval and thick-shelled, and vary from 45 to 49 p 
long by 32 to 34 broad. They are fully embryonated when laid. 


Male. 

The males vary in length from 29 to 34 mm., with a thickness 
of 9 to 1 mm. in their posterior quarter; the body is slightly 
attenuated anteriorly. 

The bursa is long and pointed, and is reflexed ventralwards. 
Its under surface is ornamented with small triangular spike-like 
processes arranged longitudinally, and extending as far as the 
4th pair of post anal ventral papille. The four pairs of stalked 
papullz are arranged in two groups, the anterior two pairs being 
pre-anal and the posterior two pairs post-anal in position; the 
Ist and last pairs are shorter than the other two, and their origins 
are slightly more ventral. 

The three pre-anal ventral papilla are of equal size, and are 


1054 MR. R. J. ORTLEPP ON THE 


shghtly separated from each other, the middle one being nearer 
the anus. The first two pairs of post-anal ventral papille are 
small, and are situated close together, one pair behind the other, 
in the immediate vicinity of the anus; pairs 3 and 4 are 
approximated to each other, and are situated at the junction of 
the Ist and 2nd tail thirds; the 4th pair is situated nearer the 
ventral mid-line than the 3rd pair. The last pair is found at 
the junction of the posterior tail thirds. 


Text-figure 28. 


ver 


oe yewree 


>> 
a5 
x 
>» 
y 
>> 
S 
s 


Ph ysaloptera mordens Leiper. 
Caudal extremity of male. 


The spicules are very unequal, the left being long and slender 
and about eleven times as long as the right; it is filiform, tapers 
to an acute point, and appears to vary in length from 4:6 to 
5-5 mm., with a breadth of 35. The right spicule is short and 
stout, with its apex tapering to forma long and thin point ; it is 
ATO to 500 » long by 50 to 55 » broad at its base. 

Hosts: Homo sapiens. Stomach and intestine. Africa. 

‘* Monkeys.” Stomach. Nyasaland. 

My observations differ from Leiper’s description, firstiy, in the 
number of uteri, of which he records only two; I was able to 


NEMATODE GENUS PHYSALOPTERA. 1055 


determine their number by dissection, as it is almost impossible 
to see them by simply clearing and rolling the worms. Secondly, 
in the size of the females; his longest specimens were 55mm. 
long, whereas mine reached 100 mm. And, thirdly, in the size 
of the spicules; the left spicule, according to my observations, 
appears to be longer than in his. 

The only other Physaloptera from man is Ph. caucasica 
v. Linst., 1902. The arrangement of its male bursal papillee is 
very similar to that of Ph. mordens, excevt, as Leiper has pointed 
out, that the 3rd pair of post-anal ventral papille is nearer to 
the mid-line than in Ph. mordens. These two species, however, 
appear to be distinct, as shown by the presence of only one tooth, 
smaller spicules, and larger eggs, in conjunction with the much 
smaller body in Ph. caucasica. 

The nature of the labial armature, mode of origin and number 
of the uteri, general shape and size of the male bursa, distribu- 
tion of the bursal papille, and the great inequality in the length 
of the spicules, allies this species to the reptilian species 
Ph. abbreviata, Ph. varani, and Ph. antarctica. To the mam- 
malian species Ph. numidica it is also related, as shown by the 
number and shape of its teeth and the character of its female 
genitalia. Ph. mordens is, however, easily distinguished from all 
these by its much larger size, relatively longer trunk portion in 
the femaie genitalia, and by its much larger spicules. 

The occurrence of this species in monkeys from Nyasaland is 
very interesting, and leads one to think that these Primates are its 
normal hosts, man being only an accidental one; this view is 
further supported by the fact that its only occurrence In man 
is in the natives of Hast and Central Africa. 

A parallel case in West Africa is the occurrence in monkeys 
and man of Gsophagostomum apiostomum (Willach, 1891), Rail. 
& Henry, 1905. 


(20) PHYSALOPTERA TUMEFACIENS Henry & Blanc, 1912. 
(Text-figs. 29 & 30.) 

The material examined consisted of three males and three 
females, kindly placed at my disposal by Prof. Leiper. This 
material was part of the collection of worms from which Henry 
and Blane described the species, and was presented to Prof. Leiper 
prior to the publication of their deseription. In addition, two 
males, obtained from a Macacus fascicularis which had died in 
the Gardens of the Zoological Society of London, were also 
studied. 

The cuticle is very finely striated transversely, and also shows 
a coarse cuticular ringing. Anteriorly it is reflected over the 
lips, and this may also be the case over the tail region in both 
sexes; this posterior reduplication of the cuticle may pass over 
as much as the anterior third of the bursa in the male; in the 
females it only forms a narrow collar in front of the anus. The 


1056 MR. &R. J. ORTLEPP ON THE 


cervical papillee vary in position; in one they are placed at the 
junction of the two esophageal parts, in the others up to 150 pa 
further back ; they may be symmetrical or asymmetrical in 
position. The excretory pore is found about 70 behind the 
cervical papille. 

The two lateral lips are large and conical in side view, and 
each carries two large submedian papille on their outer convex 
surface. Only two teeth are present—a large and conical outer 
tooth with obtuse tip and slightly bent outwards, and a flattened 
and broad inner tooth, of the same size as the outer, and having 
its free end divided into three cusps of the same size. 

The csophagus is straight, and forms in the female 1/5:2 to 
1/5°4, in the male 1/4°6 to 1/5:2 of the total body-length; its 
anterior 1/1lth or 1/12th forms its muscular part, which is 
slightly thinner than its posterior glandular portion. The nerve 
ring encircles it in its posterior third. 


Text-figure 29. 


\ 


/imm. 
"Z5Oph 


\ 8 G 
(IN\ 
Physaloptera tumefaciens Henry & Blanc. 
A = Lateral view of lips. B = Terminal portions of female genitalia. 
C = Receptaculum seminis. 
Female. 

The three females are 50, 38, and 36 mm. long, with a thick- 
ness of respectively 1:9, 1:5, and 1-2 mm. The anterior half 
tapers considerably, but posteriorly this tapering is limited to 
the last 4 cr 5mm, The tail is conical, and forms from 1/57th 
to 1/65th part of the body. Its caudal pores are situated in 
slight pits in the posterior half of the tail; their position divides 
the tail into the ratio of 5:2. 

The vulva opens on a slight elevation, which divides the body 
into the ratio of 1:2°9; in one of the females it is situated in 
front of the end of the esophagus; this position 1s probably due 
to shrinkage of the anterior part of the worm, as shown by the 
facts that the cuticle in this region is much corrugated and that 
the intestine immediately behind the cesophagus is thrown into 
a loop. 

The genitalia pass straight back, and are specially characterized 
by the absence of a common trunk portion behind the egg- 
chamber, and by the presence of four uteri arising all together 
from the hind end of the egg-chamber. The vagina, in the 


NEMATODE GENUS PHYSALOPTERA. 1057 


largest. female, is 1:7 mm. long by 100, thick, and the egg- 
chamber 800y long by 270 broad at the origin of the uteri. 
The receptacula seminis are pear-shaped, and are sharply 
constricted off from the oviducts, their union with the uteri 
being gradual; they are 380, long by 200, broad at their 
oviducal end. 

The eggs are oval and thick-shelled, and average 43 » long by 


27 w thick. They contain a fully-developed embryo when laid. 


Mate. 

The males vary in length from 26 to 31 mm., and are 
attenuated in their anterior two-thirds; they vary in thickness 
from 1 to 1-2 mm. 

The bursa is narrow and elongate and rounded at its tip; its 
anterior part may or may not be covered over by a posterior 


Text-figure 30. 


Ay 


$ 
‘ 


fl Lf 

| a] 
\ ‘G 
\ [ 


Physaloptera tumefaciens Henry & Blanc. 
Caudal extremity of male, 


reflection of the cuticle. Its central ventral area is ornamented 
with irregular tubercles arranged longitudinally; laterally in the 
cloacal region these tubercles are replaced by longitudinal ridges. 

The four stalked papille are short and arranged equidistant 
from each other round the cloaca. The three pre-anal ventral 
papille are situated in a row just anterior to the anus; the central 
papilla is large. There are five pairs of post-anal ventral papille, 
of which the first two pairs are small and situated close together, 
one behind the other, immediately posterior to the anus. 
Papilla 3 are situated at the anterior quarter of the tail, and 
papille 4 half-way between them and the middle of the tail. 
Papille 5 are slightly approximated to the mid-line, and are 
situated slightly behind the middle of the tail. 

The spicules are bent, thick, short, and slightly unequal. 


1058 MR. R. J. ORTLEPP ON THE 


They are of equal thickness, and both taper to fine points. The 
right varies from 475 to 520 4 long by 55 » thick, and the left 
from 740 to 870 » long, with the same thickness as the right. 

Hosts: JJacacus cynomolgus. | Sei AR aR 

Macacus fascicularis. 

My observations on this species are practically identical with 
those of Henry and Blane; the only differences are in the size 
of the egg, which appears slightly smaller in my material, and in 
the size of the spicules, mine being slightly larger. 

Ph. tumefaciens, Ph. mordens, Ph. nwmidica, and Ph. magni- 
papilla are the only 4-uterine forms described from mammals, 
Ph. mordens and Ph. numidica have teeth as in Ph. paradoxa, 
whereas Ph. magnipapilla and Ph. tumefaciens have each only 
two teeth to each lip. Ph. tumefaciens is distinguished from the 
other three species by the mode of origin of its uteri, by the 
absence of a common trunk portion in the female genitalia, and 
by the reduplication of the cuticle over the caudal extremity in 
most individuals of both sexes. 


(21) PHYSALOPrERA MAGNIPAPILLA Molin, 1860. (Text-fig. 31.) 


Some of the excellently preserved type material in the collec- 
tion of the Vienna Museum was examined. 

The body appears smooth, but under high magnification the 
cuticle is seen to be provided with very delicate transverse 
striations. It is slightly reflected over the base of the lips, and 
in some females it was reflected over the whole of the tail in a 
manner similar to that seen in Ph. preputialis. 

The cervical papille are small and spike-like, and pierce the 
cuticle about 150 ~ behind the level of the hind end of the mus- 
cular cesophagus ; from 75 to 100 » further back on the ventral 
surface the opening of the excretory duct is found. 

The lips are somewhat quadrangular, with large subdorsal and 
pri rennin al external papille, and each is pr onilad: with two teeth 
of the same height; the outer tooth has a widened tip, whereas 
the median inner tooth: is tripartite. 

The esophagus immediately follows the lips. It is straight, 
and forms in the female 1/5°5 and in the male 1/6°6 of “the 
total body-length. Its anterior muscular part is nearly 1/10th of 
the whole organ, is slightly thinner, and is encircled in its 
posterior third by the nerve rine. 


Hemate. 

The females vary from 30 to 38 mm. long by ‘9 to 1:1 mm. 
broad; they ave attenuated in their anterior fourth and posterior 
fifth, and the body is terminated by a bluntly conical tail 
1/52nd of the body-length and having its caudal pores in its 
posterior half. 

The vulva opens level with the surface, and its position divides 
the body in the ratio of 1 :2°3; it leads into a straight muscular, 
vagina nearly 2 mm. long by 80, thick, The egg-chamber is 


i 


NEMATODE GENUS PHYSALOPTERA. 1059 


slightly shorter, and about three times as broad as the vagina. 
The common trunk is about 1:1 mm. long and 754 thick; its 
posterior end divides twice, in the same way as seen in Ph. 
abbreviata, to give rise to the four uteri. 

The eggs are small, oval, and thick-shelled, measuring on an 
average 40 long by 26m broad; when laid they are already 
fully embryonated. 


Male. 

The males are much smaller than the females, and measure 
from 20 to 25 mm. long, with a maximum breadth of '7 to °85 mm. 
The body is attenuated in its anterior half, the posterior half 
being of uniform thickness. 

The bursa is pointed and recurved ventralwards. The four pairs 
of pedunculated papille are equidistant, three pairs being pre- 
anal. The three pre-anal ventral papille are small, and are 


Text-figure 31. 


Physaloptera magnipapilla Mol. 
A = Lateral view of lip. B = Caudal extremity of male. 


situated immediately in front of the anus. The first two pairs of 
post-anal ventral papille are also small and situated in a row in 
close proximity to the anus. The 3rd pair is larger and slightly 
obliquely placed ; they divide the tail roughly into the ratio of 
1:3. The 4th and 5th pairs are small and approximated 
to each other, and are situated just behind the middle of the 
tail. The ventral surface of the bursa is ornamented with 
longitudinal rows of tubercles. 

The spicules are straight, tubular, pointed, and slightly un- 
equal; the left is 450 long by 26 w. broad at its base, and the 
right is 415 « long by 20 u broad at its base. 

Host: M yrmecophage bivittata. Stomach. Brazil. 

Types in bottle 4457 in the Vienna Museum. 

My observations differ from those of von Drasche only in respect 
to the teeth; he states that the outer tooth is small, and in 
his figure shows it smaller than the inner tooth. In the worms 


1060 —, _ MR. RB. J. ORTLEPP ON THE 


examined by me the two teeth were of the same size and fairly 
large. 

The number of uteri and their mode of origin brings the 
species into the group characterised by P. abbr evict. It differs, 
however, from all the reptilian tetradelphoid forms by the 
shape of its outer tooth and by the presence of a large and 
tripartite inner tooth; this latter tooth is either absent 
Ph. colubri—or represented only by a small spike-like tooth 
at the base of the outer tooth—P. paradoxa—in the reptilian 
forms. 

The nature of the teeth and the presence of four uteri ally this 
species to Ph. tumefaciens Henry & Blanc, 1912, from which 
species it differs, however, by the difierent arrangement of its 
bursal papillee, no reduplication of the cuticle over its tail, mode 
of origin of its uteri, and the presence of a common trunk 
portion in its unpaired female genitalia. 


(22) PHYSALOPTERA QUADROVARIA Leiper, 1908. (‘Text-fig. 32.) 


The types of this species, which were kindly placed at my 
disposal for study by Professor Leiper, consisted of six mature 
females and one male; the latter unfortunately had most of its 
bursal region broken off. 

The parasites are stout, being thickest just behind their 
middle and attenuated towards both extremities. The cuticle 
shows a fine transverse striation, and is partly reflected over 
the lips. 

The two cervical papille are situated laterally some distance 
behind the junction of the two cesophageal parts. In the male 
they are 550 from the anterior end, and in the females 
slightly more than 600. The excretory duct opens ventrally 
at the same level or just behind the level of the cervical 
papille. 

The lips are large and somewhat flattened in lateral view ; 
each carries a large wedge-shaped external tooth with its tip 
slightly bent outwards, aad two much smaller lateral teeth 
slightly split. Between the median and lateral teeth, on the 
inner face of the lip, there isa row of very small denticles. 
Internal to and attached to the base of the external tooth there 
is a small and spike-like internal tooth. 

The cesophagus is relatively short, forming in the male 1/7th 
and in the female about 1/9th of the total body-length ; it is 
straight, slightly thickening posteriorly, and its anterior end is 
differentiated into a short and narrow muscular part, about 350 
long in both sexes, and having the nerve ring in its posterior 
quarter. 


Female. 
The specimens vary in length from 28 to 32 mm., with a 
breadth of -95 to 1:1 mm. The tail is relatively long and 


NEMATODE GENUS PHYSALOPTERA, 1061 


pointed, forming 1/45th of the total body-length, and has its 
eaudal pores situated in its posterior third. 

The vulva is situated on a slight elevation at the junction of 
the Ist and 2nd body fifths; it leads into a long and thick- 
shelled vagina 2°2 mm. long by 95 broad. This vagina passes 
straight backwards to join the egg-chamber, which is also large 
and straight, measuring 1-8 mm. long by 380 » broad. The hind 
end of thischamber narrows sharply to join the common trunk, 
which is 760 » long and 90 » in diameter; its posterior end divides 
into two, each of which in their turn subdivide to give rise to 
the four uteri. These two primary branches are so short as: 
to almost give the appearance that the uteri all arise at the same 
level. 


Text-figure 32. 


250 p. 


> 


Physaloptera quadrovaria Leiper. 


A = Anterior extremity of body. B = Inner view of lip, 
C = Mode of origin of uteri. 


The eggs are thick-shelled and oval, measuring 51 » long by 
36 p broad; they embryonate in utero. 


Male. 

The solitary and incomplete male is 19°5 mm. long and 750 u 
thick. Part of the left spicule—the bursa is broken off just in 
front of the anus—is seen inside the body, but nothing remains 
of that of the right side. The remaining portion of the left 
spicule is long and filiform, measuring 1-3mm.long. The spicules 
are probably very unequal. 

Host: Varanus niloticus. Intestine. Sudan. 

Discussion.—See Ph. varant. 


Proc. Zoou. Soc.-—1922, No. LX XI. 71 


1062 MR. R. J. ORTLEPP ON THE 


(23) PHYSALOPTERA PARADOXA von Linstow, 1908. (Text-figs. 
33 & 34.) 


Syn. Ph. affinis Gedoelst, 1916. 


The primary study of this species is based on materials 
collected in the Zoological Society’s Gardens from Varanus 
albigularis. All the specimens from the other hosts were kindly 
handed over to me by Professor Leiper for study and comparison. 
The material from Varanus alhigularis consisted of about a dozen 
males and females, which had unfortunately been killed in 
formalin, so that the specimens were somewhat shrunken and 
coiled. 

The shape of the body is very similar to that of P. gquadrovaria, 
but is much smaller in size, mature females varying from 18 to 


Text-figure 33. 


sae a 
é AA 2 


Physaloptera paradoxa v. Linst. 


A = Inner view of lip. B = origin of uteri. 
C = Receptaculum seminis. D= Ege-shell. 


24mm. in length with a thickness of 750 to 870 in their 
posterior third, and the males from 14 to 18mm. long by 
450 to 500 w broad in their posterior fifth. 

The cuticle is coarsely annulated and, in addition, shows a very 
fine transverse striation; it is only very slightly reflected over 
the base of the lips. The cervical papille and excretory pore 
are situated in the same level behind the junction of the mus- 
eular and glandular cesophageal parts; in the male they are 
about 550, from the anterior end and in the female about 
750 p. 

The lips are similar to those described for Ph. guadrovaria, 
except that lateral to each of the outer teeth there is another 


NEMATODE GENUS PHYSALOPTERA. 1063 


row of small denticles on the inner surface parallel to the edge 
of the lip. 

The muscular part of the cesophagus is thinner than the 
glandular part, and is encircled by the nerve ring at its base. 
The whole organ is short, and forms in the females 1/7th and in 
the males nearly 1/8th of the total body-length. 


Female. 
The vulva opens practically flush with the surface, there 
being only a slight trace of an elevation. Its position is 


post- “cesophageal, and divides the body in the ratio of 1:2°4; it 
leads into a long and thick-walled vagina about 2mm. long 
by 100 » broad ; ‘its most anterior portion is slightly twisted. 
The following egg-chamber is straight, and lies parallel to the 
intestine; it is slightly thicker than the vagina but shorter, 
measuring 1°2 mm. long by 133 w broad. The following common 
trunk is relatively short, and is thinner than the vagina ; it is 
570 pw long and 57 » thick. From its posterior end the four uteri 
take their origin at the same level, but their cavities arise by a 
double Sdmicion of the unpaired duct. The uteri at first pass 
forwards to beyond the posterior end of the cesophagus, after 
which they bend backwards and fill the body almost to the 
posterior end. 

The eggs are thick-walled, oval, and fully embryonated tn utero ; 
they measure 90 » long by 35 m broad! 

The body is terminated by a relatively long and pointed tail, 
forming 1/47th of the total length; its caudal pores are situated 
just behind its middle. 


Mate. 

The caudal bursa is large and provided with well-developed 
ale; its ventral surface is traversed by coarse cuticular eleva- 
tions arranged in longitudinal rows. The anus is a large and 
transversely oval aperture, with a thickened rim, very prominent 
when viewed from the ventral surface ; it is situated about 850 u 
from the tip of the tail. 

The four pairs of pedunculated papille are equidistant from 
each other; two pairs are pre-anal and two pairs are post-anal 
in position; of the three pre-anal ventral papille the median one 
is situated slightly closer to the anus than the other two. Behind 
the anus there are five pairs of sessile papillae: the Ist and 
2nd pairs of these are small, and situated close together just 
behind the anus, one pair behind the other; the 3rd and 4th 
pairs are closely approximated, and found: just in front of the 
middle of the tail; while the last pair is situated just posterior to 
the front margin of the posterior quarter of the tail. 

The spicules are very unequal, and appear to vary in size; but 
these variations may be due to the difficulty found in measuring 
them. The right spicule is short, and varies from 185 to 240 pu 


71* 


1064 MR. R. J. ORTLEPP ON THE 


long, with a thickness of 50, at its base; the left spicule 

is long and filiform, varying from 1:8 to 2°8 mm. long; it is 

about 15 w thick atits base ; both spicules terminate in sharp tips. 
Hosts: Varanus albigularis, Stomach. South Africa. 


Psammophis sibilans. 3 Sudan. 
“Snake.” sy Nigeria. 
‘¢ Colubrine snake.” F Sudan. 
“Snake.” a Nigeria. 


Discussion.—My observations differ slightly from those of 
Seurat (1914 5) made on parasites obtained from Varanus griseus 
and Cerastes cornutus. These differences apply to the spicules, 
which are 100 and 1:92mm. long for the right and left 
spicule respectively in Seurat’s material. whereas in my material 


Text-figure 34, 


hh 
1,\\ 
My! 


O-S 77777. 


Physaloptera paradowxa v. Linst. 


A = Caudal extremity of male. 
B = Spicules (only part of left indicated). 


the right spicule varied in length from 185 to 240 and the 
left from 1:8 to 2°8 mm. Also the eggs in his specimens are 10 p 
broader than in mine. Despite these differences I believe the 
parasite to be the same, and the difference in size of the spicules 
can be accounted for in that it is not always possible to measure 
them correctly, as they generally take up a twisted course inside 
the body. 

Von Linstow (1908), in his description of this species, draws 
attention to the absence of caudal ale on the tail of the male, 
and for this reason gave it the name “ paradoxa.” Seurat (1914) 


NEMATODE GHNUS PHYSALOPTERA. 1065 


explains this absence by supposing that von Linstow was exam- 
ining immature material, as he (Seurat) had in his possession 
immature specimens of Ph. alata in which the caudal ale were 
also absent. Unfortunately, | have not in my collection any 
Physaloptera in this condition, and consequently I accept Seurat’s 
interpretation. 

‘There is, however, another difference between von Linstow’s 
cbaemzaitens and mine; this concerns the number of the post-anal 
ventral papille. Von Linstow finds an extra pair present behind 
the second, whereas in my material there is no indication of 
these papille, neither were they present in NSeurat’s (1914) 
material. It is therefore probable that von Linstow was mis- 
taken in his observations, especially as my material and his are 
from the same host, both from South Africa, and there further 
appears no reason to doubt the identity of the two materials. 


(24) PHYSALOPTERA VARANI Parona, 1889. (Text-fig. 35.) 


The material on which the study of this species is based 
consists of three males collected from Varanus bengalensis and 
two females collected from V. indicws. I wish to express my 
indebtedness to Professor Leiper for placing this material at my 
disposal. 

The body is attenuated towards both extremities, and shows a 
coarse transverse ringing in addition toa very fine transverse 
cuticular striation. The cuticle is partly reflected over the lips. 

The cervical papille are situated laterally a short distance 
behind the junction of the two cesophageal parts; on the same 
level or just posterior to it is the opening of the excretory 
gland. 

The two lateral lips are large and tall. Each is provided with 
a large external tooth whose tip is slightly recurved ; attached 
to it on its inner surface there is a small and membranous spike- 
like tooth, and on each side of it, towards the angles of the lips, 
there is a small bifid tooth. Externally each lip carries a large 
subdorsal and subventral papilla. 

The cesophagus is long, and forms in the male 1/6th and in 
the female 1/5°5 of the total body-length. Its anterior muscular 
part is slightly thinner than the following glandular part, and 
the nerve ring encircles it about 80 w from its base. 


Female. 

The two females are respectively 35 and 17 mm. long by 1 mm. 
and 530 w broad ; the body is terminated by a pointed tail 1/80th 
of the total length, with its caudal pores just behind its middle. 

The vulva is non-protuberant, and is situated at the junction 
of the lst and 2nd quarters of the body; it leads, in the 
smaller female, into a thick-walled vagina 950 » long by 85 » in 
diameter; this passes gradually into the egg-chamber, which is 
1‘8mm. long and 170 » thick. Thecommon trunk which follows 
it is 1:-14mm. long and 50 win diameter. The first two parts pass 


1066 MR. R. J. ORTLEPP ON THE 


straight down the body, whereas the common trunk is recurved 
and passes forwards parallel to the other parts. The posterior 
end of the common trunk divides into two branches, each of 
which in their turn, after a distance of 120, subdivide to give 
rise to the connections of the four uteri. 

The eggs are oval and thick-shelled, and contain fully 
embryonated embryos before being laid. They ave 53 » long by 
32 w in diameter. 

Male. 

The males are respectively 12,21, and 24 mm. long, the largest 
being 800» thick in its posterior quarter. 

The caudal bursa is large, and nearly twice as long as it is 
broad. The papille are arranged in identically the same way as 
those described for Ph. paradoxa, as also are the tubereular 


Text-figure 35. 


Physaloptera varani Parona. 


A = Ventral view of cephalic extremity. 
B= Caudal extremity of male. 


elevations. The outline of the bursa, however, is slightly 
different, its length and breadth being respectively relatively 
shorter and broader than in Ph. paradox. 

The spicules are very unequal and bent, and both end in sharp 
points. The right is broad, with almost parallel edges except 
for its posterior tenth; it is 342 long by 25 broad at its 
base. The left spicule is long and filiform, 2:lmm. long by 
14 » broad at its base. 

Hosts: Varanus bengalensis. Stomach. Ceylon. 

Varanus indicus. af India. 

Discussion.—Seurat (1917 a) considers this species to be iden- 
tical with Ph. paradoxa v. Linst. and Ph. quadrovaria Leiper. 


NEMATODE GENUS PHYSALOPTERA. 1067 


After a study and comparison of these three species, I hold that, 
although they are nearly related, they must be regarded as distinct 
species. Ph. varani differs from Ph. paradoxa by its longer 
cesophagus, longer trunk, mode of origin of the four uteri, and 
the absence of a denticular ridge on the inner side of each lip. 
Tt differs from Ph. quadrovaria by its longer esophagus, shorter 
tail of the Female, less evolved type of origin of the uteri, and 
also by the absence of denticles on the inner surface of the 
lips. 

Ph. varant appears to be very closely related to Ph. abbreviata, 
as redescribed by Seurat (19146 and 1917 @), with which species 
it agrees In the female genitalia. It appears, however, to differ 
from Ph. abbreviata in its larger size, absence of denticles on the 
inner surface of the lips, relatively shorter bursa, and larger 
spicules. I have unfortunately not been able to examine and 
compare any examples of Ph. abbreviata. 

Ph. pallaryi Seurat, 1917, seems to be a near relative of 
Ph. varani, from which species it can, however, be very easily 
distinguished by its much shorter left spicule, the conformation 
of the circumcloacal area, and by the position of the vulva in 
front of the posterior limit of the cesophagus. 

Parona’s (1889) description differs from my findings in that 
he mentions and figures the presence of four pairs of post-anal 
ventral papille, the last three pairs of which are equidistant 
from each other, the 1st pair being situated immediately behind 
the anus. I think an error has crept in his observations: namely, 
that he has missed one pair of small papillee just behind the Ist, 
and has misjudged the distance separating the 2nd and 3rd 
pairs in his figure. I am led to this view because I think his 
specimens could not have been well preserved, as he shows the 
caudal ale having a lobuiated border, a state of affairs which 
Tt have noticed to take place in contracted specimens. I do not 
think there can be any doubt as to the identity of the materials, 
although Parona’s description is so incomplete that it can apply 
to quite a number of Physaloptera. 


(25) PHYSALOPTERA ANTARCTICA v. Linstow, 1899. (Text-figs. 
36 & 37.) 


Syn. Ph. alba Stoss., 1902. 


Female. 


Mature and fully-grown females are about 45 mm. long and 
950» broad. The body is of more or less uniform thickness, 
only tapering slightly towards the anterior end. The body is 
terminated bya short tail about 440m long. The cuticle is 
roughly ringed, and between these rings a very fine cuticular 
striation is observed under high-power magnification ; the cuticle 
anteriorly is partly drawn over the lips. ‘The cervical papille are 
lodged in shallow pits in the cuticle, and are situated about 650 p 


1068 MR. R. J. ORTLEPP ON THE 


from the anterior end; the excretory pore is seen on the ventral 
surface about 95 ~ further back ; it leads into the exeretory duct, 
which passes into a gland lying against the ventral surface of the 
esophagus. The two lateral lips are each surmounted by a large 
triangular tooth, slightly recurved at its tip; applied to the inner 
surface of each isa small spike-like tooth. On either side of the 
median tooth, in the lateral angles of each lip, there is seen 
another tooth, much smaller than the terminal tooth and split 
almost to its base; just below these and on the outer surface of 
the lip there is a conspicuous papilla. The lips are immediately 
followed by the cesophagus, which is straight and thickens slightly 
towards its posterior end. It is about 4*7mm. long or 1/9°5 of 
the body-length, and is divided into a short and narrower 


Text-figure 36. 


JOO ps. 


| 
| 
| =) 
| 
| 


Physaloptera antarctica v. Linst. 
Latero-ventral view of cephalic extremity. 


anterior part about 475 » long and a longer posterior glandular 
part. The first part is surrounded near its base by the large 
nerve ring. 

The chief characteristics of the female genitalia is the presence 
of four uteri. The vulva is a circular aperture, flush with the 
surface and situated at about the junction of the Ist and 2nd 
quarters of the body. It leads into a short and slightly coiled 
vagina, very muscular and about 90 » in diameter. It is followed 
by the egg-chamber, about 259 w in diameter; the hind end of 
this chamber constricts suddenly, so that the common trunk has 
about the same diameter as the vagina. The common trunk 
divides posteriorly into two branches, and these after a short 
distance subdivide again. The four uteri thus formed may first 


NEMATODE GENUS PHYSALOPTERA. 1069 


pass forwards to beyond the junction of the cesophagus and 
intestine and then pass backwards, or they may pass backwards 
directly. The uteri are more or less parallel, and their hinder 
ends pass gradually into the receptacula seminis, about 380 wu 
long by 150, broad. The receptacula seminis pass abruptly 
into the oviducts, 38 in diameter, which soon join the ovaries ; 
the latter pass forwards, taking more or less a convoluted course. 


Male. 


The males are slenderer than the females, fully-grown forms 
being about 32mm. long and 750, broad. The body is only 


Text-figure 37, 


Physaloptera antarectica v. Linst. 
A = Caudal extremity of male. B = Tip of left spicule. 
C = Right spicule. 


slightly attenuated anteriorly. The cuticle is of the same char- 
acter as the female, and the cervical papille occupy relatively the 
same position, being situated a short distance beyond the junction 
of the muscular and glandular portions of the esophagus. The 
excretory pore is situated ventrally about 130, further back 
than the cervical papilla. - The lips are as described for the 
‘female. 

The first part of the cesophagus is thinner than that following, 
measures about 500 long, and is surrounded at its base by the 
nerve ring. The total length of the esophagus is 3°55 mm. or 
1/9th of the body-length. 


1070 MR. R. J. ORTLEPP ON THE 


The bursa is large, being about 24mm. long by 1} mm. broad 
across the cloaca. The lateral expansions are well developed, 
and the central portion of the ventral surface is covered by 
Jongitudinal rows of spike-like tubercles. Theanus is a triradiate 
aperture, situated in the centre of a more or less circular cushion 
hke a swelling, about 420 m in diameter, and itself covered with 
small tubercles. On either side of it there are four elongate 
papille, implanted equidistant from each other. Anterior to the 
cushion there are three ventral papille of equal size and situated 
in arow; immediately posterior to the cushion there are two small 
pairs of ventral papille. There are three other pairs of ventral 
papille further down the tail, of which the middle pair is nearer 
to the anterior than to the posterior pair. These last three pairs 
of papille are each provided with a short stalk. 

_ The spicules are unequal, that of the left side being longer 
than that of the right side. Theleft spicule measures 1504 » long 
by 19 broad at its base; at its tip it widens out to form a kind 
of spear-head 64 w long by 18 » broad ‘The right spicule is stout, 
and measures 410 long by 33 w broad at its base. In its pos- 
terior two-fifths it becomes slightly widened out, after which it 
gradually tapers to a point. 
Hosts: Varanus varius. 
Python spilotes. 

My material has been identified as Ph. antarctica v. Linst. 
because of the general agreement of the measurements of my 
material with those of von Linstow’s; the only important differ- 
ence is the relative length of the tail of the female ; von Linstow 
gives it as 1/54th of the body-length, whereas in my material 
it forms only about half this length. 

Only two species have so far been described from Australian 
lizards—namely, the species described by von Linstow and Ph. 
alba described by Stossich (1902). Irwin-Smith (1922) considers 
these species to be the same, and Lam inclined toagree with her. 
Unfortunately, [have not been able to examine material from 
any of the hosts from which these two species were obtained, but 
from the brief descriptions and figures given by von Linstow and 
Stossich it appears probable that they are identical. 

Ph. antarctica, because of the nature of its labial teeth and 
mode of origin of the four uteri, is closely related to Ph. abbre- 
viata, Ph. varani, and Ph. pallaryi; it is, however, very easily 
distinguished from all these by the configuration of its male 
bursa and by the difference in size and shape of the spicules. 
I have given an extract of both von Linstow’s and Stossich’s 
descriptions in Part II. 


\ Stomach. Australia. 


(26) PuysaLoprera cotuBrir (Rud., 1819), Diesing, 1851. 
(Text-fig. 38.) 

The types of this species, deposited in the Vienna Museum, 
consisted of two immature females and the fragments of another 
worm; the specimens were slightly shrunken. 


NEMATODE GENUS PHYSALOPTERA. 1071 


The cuticle is finely striated transversely, and is partly reflected 
over the lips; the cervical papille are situated about 100 behind 
the level of the hind end of the muscular cesophagus, and the 
excretory pore about 75 w further back. 

The lips are rounded, and each is surmounted by a single large 
terminal tooth, no other teeth being present. Externally each 
lip carries a small subdorsal and subventral papilla. 

The cesophagus forms nearly 1/4°6 of the body-length; its 
anterior muscular part is thinner than the rest, and it is encircled 
by the nerve ring in its posterior third; it forms just less than 
1/10th of the whole organ. 

Female. 

The two specimens measure respectively 7 and 8mm. long by 
350 and 360 w broad ; the body is attenuated in its anterior half, 
and it is terminated behind by a relatively Jong and blunt ly 
rounded tail, which is turned up dorsally in both specimens; it 


Text-figure 38. 


Physaloptera colubri (Rud.), Dies. 
. A = Lateral view of lip. 
B = Caudal extremity of male. (After v. Drasche.) 


is about 1/30th of the body-length, and its caudal pores are 
situated in its posterior third. The vulva is only very slightly 
protuberant, and its position divides the body intu the ratio of 
1:2°5; it leads intoa vagina bent on itself and measuring 17 mm. 
long by DD p thick; the egg-chamber is 320 » long by “145 pin 
diameter, and is also bent on itself; the following common trunk 
is of about the same length as the egg-chamber, and is about 
50 « broad; its posterior end divides into two, each of which, 
after a distance of 240, again divides into two to give rise to 
the four uteri. 

No eggs were observed, the specimens being immature. 

Male. 

The following description of the male bursa is based on von 


Drasche’s description and drawing, no males being available for 
personal study. 


1072 MR. R. J. ORTLEPP ON THE 


The caudal expansions are well developed, and the four pairs of 
stalked papille are situated in two groups, two pairs being pre- 
anal and two pairs post-anal. The three pre-anal ventral 
papille are of the same size, the middle one being nearer the 
anus than the other two. 

There are four pairs of post-anal ventral papille, of which the 
lst is small and situated immediately behind the anus; the 2nd, 
3rd, and 4th pairs are equidistant, the 2nd being situated at 
the level of the last pair of stalked papille, and the last near the 
tip of the tail. 

Host: Coronella austriaca. Intestine. Australia. 

Types in bottle 4451 in the Vienna Museum. 

Tt is probable that when the male of this species is re-examined, 
it will be found that there are five pairs of post-anal ventral 
papille, and that the extra papilla will be situated near the 
Ist pair just behind the anus. 


(27) PHYSALOPTERA SIMPLICIDENS, sp. n. (Text-fig. 39.) 


The material examined consisted of three females, two of which 
were immature, one small male, and one larger male which had its 
bursa broken off. 

The cuticle is finely striated, and is slightly reflexed over the 
lips; the cervical papillae are situated in small concavities just 
behind the junction of the muscular and glandular parts of the 
cesophagus. The excretory pore is situated on the ventral 
surface about 60 » further back. 

The lips are somewhat flatly conical in side view, and each 
bears a large triangular tooth, slightly recurved outwards ; no 
other teeth were observed. Hach lip carries two papille, one 
subdorsal and one subventral, on its outer surface. 

The cesophagus is short, and in the female forms 1/13th and in 
the male 1/8th of the body-length ; its anterior muscular part 
forms about 1/9th of its length, and is distinctly separated from 
the following glandular part ; it is encircled by the nerve ring in 
its posterior quarter. 


Female 

The females are about 44 mm. long and 950 broad, and are 
attenuated towards both extremities; this is only slightly 
evident posteriorly where the body is terminated by a short and 
conical tail, forming 1/105th of the body-length. The vulva 
divides the body into the ratio 1:3°3; it leads into a muscular 
vagina 1-3 mm. long by 95 » in diameter; this in turn passes 
gradually into the egg-chamber, which is 1-7 mm. long by 325 pw 
broad; the following common trunk is short, and divides into two 
branches, each of which after a distance of 200m divides again 
to give rise to the four uteri. 

The eggs are oval and very thick-shelled, measuring on an 
average 95 w long by 41 pw broad, with a shell 9 w thick; they are 
fully embryonated in utero. 


NEMATODE GENUS PHYSALOPTERA. 1073 


Male. 

The males are respectively 14 mm. and 21 mm. long; they are 
attenuated only towards the anterior end; the maximum breadth 
of the larger male is just in front of the bursa, where it is 725 
broad. 

The bursa is large, and is nearly twice as long as it is broad ; it 
forms about 1/10-7 of the body-length. The arrangement of 
the papille is similar to that found in Ph. antarctica v. Linst., 
except that the lateral stalked papille are arranged in two groups, 
the Ist and 2nd pairs forming a pre-anal group, and the 3rd and 
Ath pairs forming a post-anal group. 


Text-figure 39. 


05mm, 


Ss 250 py. 


Physaloptera simplicidens, sp. n. 
A = Ventral view of anterior extremity of body. 
B = Caudal extremity of male. 


The spicules are slightly unequal, the left being slightly longer 
and slenderer than the right; their lengths and breadths are 
respectively 510, by 38 w and 410, by 50; both end in fine 
points, the right, however, being slightly swollen just behind its 
middle. 

Host: ‘Sleeping Lizard.” Australia. 

Types in the Helminthological Department of the London 
School of Tropical Medicine. 

Affinities.—Vhe presence of four uteri and a single tooth allies 
this species to Ph. colubri (Rud.), from which species it is, how- 
ever, distinguished by its much larger size, its relatively much 
shorter cesophagus, and by the difference in number and arrange- 
ment of the post-anal ventrai papille. 


1074 MR. R. J. ORITLEPP ON THE 


Group Polydelphys. 
(28) PHYSALOPTERA TURGIDA Rudolphi, 1819. (Text-fig. 40.) 


Syn. Turgida twrgida (Rud., 1819), Travassos, 1920. 
Spiroptera turgida (Rud., 1819), Duj., 1845. 


The material studied consisted of some excellently preserved 
specimens in the collection of the Vienna Museum, and of three 
tubes of material in the collection of Prof. Leiper. 

The cuticle is slightly inflated, and is partly reflected over the 
lips; it is transversely striated by very fine strize; the cervical 
papille pierce the cuticle just behind the level of the muscular 
esophagus, and from 200 to 300 » further back is the opening of 
the excretory gland. 

The two lateral lips are semi-conical in lateral view; each is 
surmounted by a large triangular outer tooth having its tip 
shightly bent outwards; immediately internal to it is a large 
membranous tooth of equai height and tripartite; no lateral teeth 
are present. Hach lip carries two external papille, one subdorsal 
and one subventral. 

The cesophagus is long and straight, widening gradually 
posteriorly ; in the females it varies from 1/4°8 to 1/5°5 of the 
total length, and in the male from 1/3°9 to 1/59; its muscular 
portion is thinner than the rest, and forms in the female 1/10th 
and in the male 1/8th of the whole organ. ‘The nerve ring 
encircles it in its posterior third. 


Female. 


Adult and fully mature females are large and stout, measuring 
as much as 95 mm. long by 3 mm. broad in their posterior inal 
The body is attenuated in its anterior half, but posteriorly the 
thickness is more or less uniform until the vicinity of the anus, 
where the body tapers suddenly to form a pointed tail 1/45th 
to 1/56th of the total body-length, and whose caudal pores are 
situated in its posterior third. 

The position of the vulva varies considerably, but in all cases 
it was situated behind the end of the esophagus; it divides the 
body in the ratio of 1:2 to 1:4°4. The vagina is thick-walled, 
short, and passes straight back; it is just less than 1 nm. long by 
95 w thick. It passes directly into a larger ege-chamber of about 
the same length and about 230 » broad; from the posterior end 
of this chamber a number of uteri take their origin ; beeause of 
the complicated manner in which they were entwined, it was not 
always possible to determine the exact number, but in two speci- 
mens it was definitely seen that the one had 11 and the other 
had 14 uteri. The uteri pass down the body, coiled round each 
other, almost to the posterior end, when they recurve and join up 
with the receptacula seminis, which in their turn are united to 
the ovaries by means of short and thin oviducts. The oviducts 
pass forwards again to about the middle of the body. 


NEMATODE GENUS PHYSALOPTERA. 1075 


The eges are oval and thick-shelled and relatively narrow, 


measuring 48 w by 29°5 pw. 
Male. 


The males are smaller than the females, apparently mature 
specimens measuring from 21 to 32 mm. long by *950 to 1°25 mm. 
thick; the body is nauch more attenuated anteriorly than in the 
females, this thinning beginning from about the posterior third. 

The bursa is large, and has well-developed lateral expansions ; 
the four pairs of pedunculated papille are equidistant from each 
other, two pairs being pre- and two pairs being post-anal in 
position. Of the three pre-anal ventral papillee, the middle one 
is situated nearer the anus than the other two. Behind the anus 
there are five pairs of ventral papille; of these the Ist and 


‘Text-figure 40. 


O-S 177797. 


Physaloptera turgida Rud. 
A = Lateral view of cephalic extremity. 
B = Caudal extremity of male, 


2nd pairs are situated immediately behind the anus in a 
row, the 3rd pair is slightly oblique in position, and the 4th 
and 5th are situated close opener. The ratio of the distances 
separating pairs 2 and 3, 3 aad 4 to 5, and 5 to the tip of the tail 
1S sul. 974 2 ile 

The spicules are of equal length, short, straight, and tubular ; 
each is about 410 » long by 32 » broad at its base. 

The ventral surface of the bursa is covered with longitudinal 


protuberances. 
Hosts: Didelphys cancrivora. Stomach, Brazil. 
» *¢ Manicon.” sd West Indies. 
Didelphys virginiana. si Michigan. 


Affinities—See Ph. dilatata. 


1076 _ MR. R. J. ORTLEPP ON THE 


(29) PHysaLoprera DinATaTa Rudolphi, 1819. (Text-fig. 41.) 

Syn. Spiroptera dilatata (Rud., 1819), Duj., 1845. 

Two males and two females were examined from bottle 4454 in 
the Vienna Museum. ‘The females were somewhat shrunken, but 
otherwise the material was in an excellent condition. 

The cuticle is finely striated transversely, and is loosely attached 
to the body ; it is reflected over the base of the lips. 

The cervical papille are situated about 290 » behind the level 
of the posterior limit of the muscular cesophagus, and the excre- 
tory pore about another 200 w further back. Sal ga 

The two lps are large and semi-rectangular, with projecting 
dorsal and ventral corners. Hach is surmounted by a large 
conical external tooth, and a tripartite inner tooth of the same 


Text-figure 41. 


Physaloptera dilatata Rud. 
A = Terminal portions of female genitaha. 
B = Caudal extremity of male. 


height but semi-membranous. No additional teeth are present. 
Externally each lip has a subdorsal and a subventral dome-lke 
papilla. 

The cesophagus is remarkable for its difference in relative 
length in the two sexes; in the two male specimens examined it 
formed respectively 1/4:°3 and 1/4°8 of the total body-length ; in 
the females it is relatively only half this length, forming in 
the one 1/8°6 and in the other 1/7:5 of the total length. Its 
muscular part is thinner than the following glandular, and 
forms about 1/10th of the whole organ; it is encircled by the 
nerve ring at the junction of its third and last quarters, 


Female. 
The females are large and stout, the two specimens measuring 


NEMATODE GENUS PHYSALOPYERA. 1077 


respectively 55 and 90 mm. long, with a maximum breadth of 1:9 
and 2°9 mm. respectively; the body is attenuated only in its 
anterior half, the posterior half being of the same thickness, 
except for the tail region, where the body tapers abruptly to form 
a conical tail 1/85th of the total length. 

The vulva opens very far forwards, being situated in front of 
the middle of the cesophagus; its position in one specimen 
divided the body in the ratio of 1:22. It leads into a com- 
paratively straight vagina 2; mm. long by 115 uw broad. The 
following egg-chamber is club-shaped, 2 mm. long by about 840 jw 
broad at its widest part ; many uteri take their origin from the 
outer edge of the club; 15 were present in the one specimen, and 
in the other at least 12 could be made out. 

The eggs are oval and thick-shelled, averaging 39 long and 
27 w broad. 


Male. 


The males are smaller than the females, measuring respec- 
tively 47 and 55 mm. long by 1:45 and 1:6 mm. broad. The 
body has its maximum breadth just in front of the bursa, and 
from here the body attenuates gradually towards the anterior 
end. 

The bursa is broad and sharply recurved ventralwards ; it forms 
about 1/16th of the body-length, and its ventral surface is 
covered with longitudinal rows of tubercles. The four pairs of 
pedunculated papille are equidistant, three pairs being pre-anal 
and the 2nd and 3rd pairs are much longer than the other pairs ; 
of the pre-anal ventral papille, the middle one is slightly larger 
and'is situated nearer the anus. There are five pairs of post-anal 
ventral papille, of which the Ist and 2nd pairs are small and 
situated in a row immediately behind the anus; the 2nd, 3rd, 
and 4th pairs are equidistant from each other, while the distance 
between the 4th and last pair is only about a third the distance 
separating the two preceding pairs. 

The spicules appear to be short and equal, but unfortunately 
the tip of the right spicule was broken off in both specimens, so 
that it was not possible to determine its correct length; the left 
spicule was straight and pointed, and measured 655 » long by 
57 pw broad at its base ; the base of the right spicule had the same 
thickness as that of the left side. 

Host: Lagothrix humboldtii. Stomach. Brazil. 

Affinities.—This species appears to be closely related to 
Ph. turgida, with which species it agrees in the form of the lips 
and teeth, general size, and in the anatomy of the female 
genitalia. It is, however, readily distinguished from this species 
by the position of the vulva, the difference in the relative length 
of the esophagus in the two sexes, and by the position of the 4th 
and 5th post-anal ventral papille on the male bursa. 


Proc. Zoon. Soc.—1922, No. LX XIT, 


NI 
bo 


1078 MR. R. J. ORTLEPP ON THE 


(30) PHYSALOPTERA CAPENSIS, Sp.n. (Text-figs. 42 & 43.) 


The material on which this study is based consisted of one 
female, two males, and portions of two males. 

‘The cuticle is very finely striated transversely, and is only very 
slightly reflected over the base of the lips. 

The lips are large, and each has on its outer lateral border two 
dome-like papille. The median external tooth is large and is 
situated on the summit of the lip; its tip is bluntly rounded ; 
immediately internal to it there is another tooth of the same 
height, but having its tip slightly notched ; towards the angles 
of each lip there is an additional tooth, fairly large, which is 
generally bifid and may sometimes be irregularly trifid. On 


Text-figure 42. 


val 


H 


in 


A = Ventral view of anterior extremity of body, 


An iesleall 


Physaloptera capensis, sp. n. 


—. 
a 


B = Outer lateral view of lip. 
C = Terminal portions of female genitalia. 


either side of the median tooth there 1s another bifid tooth, very 
small and somewhat difficult to make out; it is situated about 
midway between the median and lateral teeth. 

The cervical papille and exeretory pore are situated at the 
same level, about 120 » behind the junction of the two cesopha- 
geal parts. 

The muscular part of the cesophagus forms about 1/11th of the 
total length of the esophagus; it is slightly narrower than the 
following part, and is surrounded in its middle by the nerve ring. 
The whole cesophagus forms just less than 1/5th of the total body- 
length in the male and 1/10th in the female. 


NEMATODE GENUS PHYSALOPYERA. 1079 


Female. 

The solitary female is robust, having somewhat the appearance 
of a short ascarid. It is 56 mm. long, and 1:7 broad in its middle; 
it is attenuated only very slightly anteriorly. 

The vulva is situated flush with the surface and 15:85 mm. 
from the anterior end ; it leads into a long vagina 3°62 mm. long 
and 152 broad; this vagina, except for a loop at its end, passes 
straight down the body to join the ege-chamber ; this is also very 
long but not very broad, measuring 4°86 mm. long by 348 y in 
diameter. The end of the chamber joins the common trunk, 
which is 2:19 mm. long and about as thick as the vagina. The 
rest of the genitalia are remarkable for the number and mode of 
origin of the uteri; there are 14 uteri arising from six branches, 
which unite to form the common trunk; four of these branches 


Text-figure 43. 


/mm. 


Physaloptera capensis, sp. n. 
Caudal extremity of male. 


give rise each to two uteri, whereas each of the remaining two 
branches give rise to three uteri. 

The tail is pointed and 950 uw long ; the caudal pores open 380 
from its tip. 

The eggs are oval and thick-shelled, measuring 55 » long by 
32 w broad, and shell 4 p thick. 


Male. 

The males are much slenderer than the females, measuring 
24 mm. long by 900 broad just behind their middle; they are 
slightly attenuated anteriorly. 

The caudal bursa is large, and twice as long as it is broad ; it 
forms about 1/11th of the body-length; the caudal alz are well 
developed, and are supported by the four pairs of pedunculated 
papille, of which two pairs are pre-anal and two pairs post-anal 

(es 


1080 MR. R. J. ORTLEPP ON THE 


in position. On the ventral surface there are three pre-anal 
papille immediately in front of the anus, and five pairs of post- 
anal papille. The Ist and 2nd of these last are found close 
together near the anus; the 3rd and 4th are closely approximated 
to each other, and are inserted in the middle of the tail; the last 
pair is situated midway between them and the caudal extremity ; 
the whole ventral surface of the bursa is covered by longitudinal 
rows of tubercles. 

The spicules are unequal, that of the left side being over six 
times as long as that of the right. The right spicule is fairly 
stout, measuring 57 at its base and gradually tapering to a 
point; it is 420 long. The left spicule is also pointed, but much 
slenderer; it is 2°7 mm. long by 27 » broad at its base. 

Host: Xerus setosus. Stomach. South Africa. 

The types to be deposited in the British Museum of Natural 
History, London. 

The mode of origin of the uteri, in conjunction with their 
number, give this species a unique position in the genus. 

The general character of the vagina, egg-chamber, common 
trunk, and eggs are typical for the genus, so that the presence 
of numerous uteri is not a sign of very close afhnity to the 
polydelphoid species Ph. turgida and Ph. dilatata. 


(31) PaysaLoprera sp. (Text-fig. 44.) 

The study of this species is based on a single male 15 mm. long 
and °65 mm. broad. It was collected from the stomach of a 
Bandicoot which had died in the Gardens of the Zoological 
Society of London. 

From the middle forwards the body becomes attenuated, but - 
behind the middle the body is of a uniform thickness. The 
cuticle is finely striated transversely, and is reflexed over the lips 
so as to completely enclose them. ‘Ihe cervical papille are found 
457 «from the head end, 7. e. at the junction of the two cwsophageal 
parts. The excretory pore is situated just behind them. 

The mouth is bounded by two large and somewhat conical lips, 
each surmounted at its apex by a large and triangular outer tooth 
and immediately internal to it by a median tripartite tooth, of 
the same height as the outer tooth, but less chitinised. Two 
external papille are present on each lip. 

The esophagus is 3°77 mm. long; its muscular part is thinner 
and 400 long, and the nerve ring encircles it just behind its 
middle. 

The caudal bursa is twice as long as it is broad, and it forms 
1/9th of the body-length. Its chief characteristic is the irregular 
arrangement of its papille. Of these there are four stalked 
papille on the right side and six on the left. Those of the right 
side are arranged in a pre-anal group of three, far removed from 
the anus, and a post-anal papilla; those of the left side are 
arranged in a pre-anal group of five and a single post-anal papilla. 
The three pre-anal ventral papille are far removed from each 


NEMATODE GENUS PHYSALOPTERA. 1081] 


other, the centre one being large and situated nearest to the 
anus. There are five pairs of post-anal ventral papille, situated 
more or less equidistant from each other; the 2nd, 3rd, and 4th 
pairs of these are obliquely situated across the tail, whereas the 
Ist and last pairs ave arranged transversely. 

The spicules are unequal, the left being longer and slendercr 
than the right. The right spicule is 370 by 50, thick at its 
base; it tapers to a rounded tip. The left spicule is 523» long 
by 20 u broad at its base, and tapers to a fine point. 


Text-figure 44. 


OS mm, 


Physaloptera sp. 


A = Latero-ventral view of cephalic extremity. 
B = Caudal extremity of male. 


The ventral surface of the bursa is ornamented with longi- 
tudinal rows of somewhat rounded tubercles. ie 

Host: Perameles nasuta. Stomach. Australia. 

I have not attempted to classify this species, because only a 
single specimen was available for study, and the arrangement of 
its bursal papille differed so much from the typical that JI 
consider the specimen to be abnormal. 


PARY If. 
Grove Didelphys. 


(32) PHyYSALOPTERA RARA Hall & Wigdor, 1918. 

Male unknown ; length of female 24 mm., maximum thickness 
1°34 mm. 

Anterior extremity somewhat attenuated. Cuticle strongly 


annulated, and first annulation behind the head forms a sort of 
collar, into which the head is partly sunk. 


1082 MR. &. J. ORTLEPP ON THE 


Two lateral lips large, each carrying three prominent inner 
median teeth and a somewhat smaller external tooth. Hach lip 
with three external papille. 

Length of esophagus 1:16 mm. ; its muscular part 526 p» long. 
Nerve ring near base of muscular cesophagus. 

Vulva 3°63 mm. from anterior end. Vagina 880, long. 
Ege-chamber and common trunk together 2°16 mm. long. 
Two uteri. Tail blunt and 420 p» long. 

Host: Canis familiaris. Duodenum. Michigan. 

Hall and Wigdor suggest that this may be an accidental 
parasite of the ‘Dog. From the remarkable shortness of the 
cesophagus and the relative length of its muscular part, I suggest 
that this worm is abnormal, which could easily be accounted for 
if it was parasitic in a wrong host. 


(33) PHYSALOPTERA TACAPENSIS Seurat, 1917. 


Length of male 15°6 mm., female 22°3 mm. 

Cervical papille subsymmetrical, at same level as excretory 
pore immediately behind the muscular esophagus. External 
tooth strongly chitinised and truncated ; internal tooth of same 
height. Cisophagus 1/4 in the male and 1/4°6 in female of body- 
length. Female tail short and conical; male tail conical. 

Vulva immediately in front of end of esophagus. Vagina 
2°3 mm. long, egg-chamber 1:02 mm. long ; common trunk absent. 
Two uteri. Receptaculum seminis very long. Eggs with very 
thick shell (7 «); 56 x 30 p. 

Male caudal bursa with longtitudinal cuticular ornamentations. 
Character and arrangement of bursal papille exactly as those in 
the species described by Seurat (1917 6) as Ph. clausa. 

Right spicule 240 long, thick, and thimble-shaped; left 
spc ule 360 u long, slender er: ‘with conical tip. 

Host: Ctenodacts ylus gundi. Stomach. Sud-Tunisien. 

From Seurat’s description this species appears to be so closely 
related to his Ph. elawsa that it appears hardly warrantable to 
separate the two. The similarity of the teeth, shape of spicules, 
and arrangement of male caudal papille appear to me to be of 
far greater weight than the slight differences of the relative 
positions of the vulva, relative lengths of the esophagus, lengths 
of the spicules, positions of the cervical papille and excretory 
Ose and the larger egg, all of which differences may very well 
be due to the fact that the one e parasite is about twice the size of 
the other. In a given species of Physaloptera taken from the 
same host, the differences in size between one adult and another 
is often very marked, and consequently there is always a a certain 
amount of difference between the individual specimens. 


(34) PHYSALOPTERA GETULA Seurat, 1917. 


Length of male 16:3 mm., female 18°5 mm. x 970 » thick. 
Body stout, attenuated anteriorly ; cervical papille sub- 


+ 


NEMATODE GENUS PHYSALOPTERA. 1083 


symmetrical, far posterior to end of muscular csophagus and 
just in front of excretory pore. Hxternal tooth conical, slightly 
shorter than the internal tooth. 

Vulva may be found behind cesophagus, but generally it opens 
in front of its posterior termination, even so far forwards as its 
middle. Two uteri arise directly from base of egg-chamber. 
Receptaculum seminis short and ovoid, not constricted off from 
the uterus. 

Kgeg? ‘Tail short, digitiform, with rounded tip. 

Body of male smaller and slenderer than that of female. Bursa 
ornamented with spines. Three large papille just in front of 
anus; 3rd pair of post-anal papille at the level of the 5th pair of 
external papille. 

Spicules unequal, right larger and shorter with bent tip, left 
slenderer and straight. 350 and 480 » long respectively. 

Host: Mus ratius. Stomach. Morocco. 


(35) PHYSALOPTERA BREVIVAGINATA Seurat, 1917. 


Only immature female known ; 9°8 mm. long x 444 » thick. 

Hxternal tooth strongly chitinised with broad base (surbaissée) 
and bent outwards. Internal tooth tripartite, and higher than 
the external tooth; its middle part is bieuspid. Two papille to 
each lip. Cisophagus 1/4th, tail 1/35th of body-length ; tail has 
a rounded tip, and the caudal pores are in its posterior third. 

Vulva immediately in front of middle of body, 2°3 mm. from 
anterior extremity. Vagina very short, 300 ~ long; egg-chamber 
600 » long, and its posterior end passes directly into the two 
uteri. 

Host: Vespertilio kukli. Stomach, Bou-Saada, 


(36) PHYSALOPTERA ALATA, var, CHEVREUXI Seurat, 1914 

Has same character as Ph. alata; differs by its smaller size and 
very posterior position of the cervical papillae and excretory pore, 

Male 7 mm. longx515y thick. Muscular cesophagus 260 p. 
Cisophagus 1/4th of body-length. Cervical papille 370 and 360 px 
from anterior end ; excretory pore 384 p. 

Female (immature). Length 8 mm.; tail long (310) and 


conical. Vulva a short distance behind end of cesophagus. 
Host: Hawk. Bone. 


(37) PHYSALOPTERA ALATA, var. NOUVELI Seurat, 1915. 

Large nematodes with robust body and regularly attenuated 
anteriorly. Cervical papille subsymmetrical, situated more or 
less at the level of the hind end of the muscular esophagus. Two 
lateral lips, each with a large triangular external and pointed 
tooth, and a very small internal tooth with three tips. Ciso- 
phagus short, 1/6th of body-length. 

Male. Length 22-285 mm.x 900, thick. Tail conical and 
slender, 1:165mm. long. Circumcloacal area covered with small 


1084 MR. R. J. ORTLEPP ON THE 


cuticular tubercles. (Arrangement of the papilleas in Ph. alata.) 
Three pre-anal sessile papille arranged ina triangle. Four pairs 
of post-anal ventral papille; the lst and 2nd pairs of these are 
slightly stalked, and are found on the posterior lip of the cloaca ; 
the last pair is situated just behind the middle of the tail. Five 
pairs of lateral stalked papille grouped about the cloaca, the last 
inserted at the level of the 3rd pair of post-anal ventral papille. 
Right spicule robust and large, 550 » long; left spicule slenderer, 
feebly chitinised, and longer. 

Female 33 mm. long x 1:055mm. thick. Tail long, conical, and 
pointed. Caudal pores in its anterior third. Cisophagus 5-61 mm. 
long. Vulva in front of termination of cesophagus, 3°8 mm. from 
anterior end. Unpaired female genitalia very long; vagina 
2°15 mm, long and directed backwards ; egg-chamber 3°125 mm. 
long; trunk 500 » long, and divides posteriorly to give rise to 
two uteri. Receptacula seminis pyriform, 600 x 325). Egg 
elliptical, thick-shelled, and embryonated ; they measure 50 x 25 p. 


Sloss oe BUT SIDE: | Cisophagus. Bou-Saada. 
ceuprter NsUs. 


(38) PHYSALOPTERA SUBALATA Schneider, 1866. 


Length of male 32 mm. 

Tips of lips very prominent. Outer tooth longer than inner 
tooth ; edges of smaller teeth quite parallel, and pointed at the 
tip. Four pairs of stalked circumcloacal papille. Three sessile 
pre-anal papille, the middle one nearer the anus; three pairs 
behind the anus, of which the last two pairs are close together, 
one pair behind the other, and the last pair nearer the mid-line ; 
the other pair is situated at about the junction of the 1st and 2nd 
tail thirds. 

Host: Falco sp. Stomach. Brazil. 

With regard to the nature of the teeth, Schneider’s description 
differs from his figure; he states that the outer tooth is larger 
than the inner, and in his figure shows the outer tooth very much 
smaller. I take his figure to represent the correct nature of the 
teeth. 

Seurat (1914 c) tentatively referred to this species some Physa- 
loptera which he obtained from the stomach cf a Buzzard, San 
Martino, Corsica. Briefly he describes this material as follows :— 
Large and robust body. ‘Two lips, each with a small and cuneiform 
outer tooth and a large tripartite inner tooth. Cisophagus 
1/4°5 in male, 1/6th in female of body-length. Vulva in anterior 
third of body, 3 mm. behind esophagus. Vagina | mm., egg- 
chamber 750 » long ; trunk divides to give origin to two uteri. 

Bursa with four stalked cireumcloacal papille. Three pre-anal 
ventral papille form a triangle; two pairs immediately behind 
anus, 3rd pair in middle of tail. 

Spicules unequal ; left 840 long, slender, and pointed; right 
less than 400 yw long, and thicker. 


NEMATODE GUNUS PHYSALOPTERA. 1085 


(39) PHYSALOPTERA CROSI Seurat, 1914. 


Male 22 mm. long by 660 » thick, female 12-22 mm. long. 

Two lips, large, each with two external papille. Hxternal tooth 
large and triangular; inner median tooth very small and tri- 
partite. Cervical papille situated immediately behind level of 
nerve ring, 

Cisophagus 1/6th in male, 1/4th in female of body-length. 

Male bursa plus well-developed ale, and ornamented with 
tubercles. Arrangement of its papille as in Ph. galiniert and 
Ph. subalata, except that an unpaired sessile papilla is present 
between the last pair of ventral papille. Spicules equal, 300 wu 
long. 

Tail of female short, with caudal glands opening in front of its 
middle. Vulva in anterior third, | mm. behind csophagus. 
Vagina shorter than egg-chamber, which is 1200 » long x 300 u 
broad; trunk 400, long, and divides to give rise to the con- 
nections of the two uteri. Eggs 55 x 25 uy, embryonated. 

Host: Accipiter nisus. Stomach. Mascara. 


(40) PHYSALOPTERA GALINIERI Seurat, 1914. 


Length, male 21 mm., breadth 780 1; length, female 17-34 mm. 
by 1:05 mm. thick. 

Two lateral lips, each with a large and very conspicuous 
tripartite inner tooth. 

(Esophagus 1/6th in male and 1/7th in female of body-length. 
Cloaca 925 1 fromend of tailin male. Five pairs of stalked papille, 
of which the anterior 4 pairs are arranged circumcloacally, and 
the last some distance further back. Eleven ventral papille, with 
short stalks and approaching mid-ventral line; of these, 3 are 
pre-anal, 4 immediately behind the anus in a transverse row, 2 at 
the level of the posterior pair of circumcloacal stalked papille, and 
2 midway between the posterior stalked papilla and tip of tail. 

Spicules short, 360 and 380 , long. 

Vulva a short distance in front of middle of body, far removed 
from hind end of cesophagus. Ovijector passes forwards ; vagina 
2 mm. long; egg-chamber oval, 600 » long; trunk 200 long. 
Two uteri, one of which passes forwards and then recurves 
backwards, whereas the other passes straight back. Recep- 
taculum seminis sharply constricted off from oviduct and uterus, 
rounded; diameter 240. Hggs 65 x 35 p. 

Host: Aigle. (Esophagus and stomach. Ain-Oussera. 


(41) PHYSALOPTERA LEPTOSOMA (Gervais, 1848), Seurat, 1917. 
Syn. Strongylus leptosomus Gervais, 1848. 
Ph. chameeleontis Gedoelst, 1916. 
Length of male 7-8-20 mm., of female 12°5-45 mm. 
Two lips; external tooth cuneiform, slightly knobbed at its tip; 
internal tooth spike-like; lateral teeth bicuspid (these teeth more 


1086 MR. R. J. ORTLEPP ON THE 


or less as in Ph. paradoxa, except that inner median tooth is 
larger); inner denticulate border reduced and interrupted in 
places. 

Muscular esophagus very short. CHsophagus 1/6th in male, 
1/10th in female. Cervical papille asymmetrical, just behind 
muscular cesophagus. 

Vulva at level of end of cesophagus o1 just posterior : vagina 
2°2mm. long, egg-chamber 925 y long, trunk 500m long. Two 
uteri. Receptaculum seminis 400 long. Hggs 52x 35 p. 

Tail of female elongate and digitiform, with its caudal pores in 
its posterior third. 

Circumcloacal area of male with small tubercles arranged 
longitudinally ; in adult males these are spined. 

Male bursa lanceolate; 4 pairs circumcloacal stalked papille. 
Thirteen ventral papilla ; of these, 3 in front of anus, the lateral 2 
being shortly stalked; 2 pairs immediately behind anus; 2 pairs 
just in front of middle of tail, the anterior pair being stalked ; 
2 papille at posterior third of tail. Spicules very unequal ; right 
short and thick, 156 x 35; left slender and twelve times as long 
as the right, 1°92 mm. 

Hosts: Uromastix acanthinurus and U. a. var. nigriventris. 

Stomach and esophagus. Algeria. 
Varanus griseus. Stomach and intestine. Algeria. 


(42) PHYSALOPrERA CHAM@LEONTIS Gedoelst, 1916. 

Syn. Ph. leptosoma (Gervais), Seurat, 1917. 

Length of male 13-4 mm., of female 22 mm. x 815 p» thick. 

Two hemispherical lips, each with a triangular external tooth. 
No internal teeth. 

(Esophagus of male 1-9 mm., of female 3°7 mm. long; bursa 
1°44 mm. long; spicules unequal, right 370 p, left 2°1 mm. long. 
Arrangement of bursal papille as in Ph. affinis. Vulva divides 
body into ratio of 1:4. Two uteri. Eggs 56-63 x 40-44 p. 

Host: Chameleon gracilis. Stomach. Belgian Congo. 

IL agree with Irwin-Smith (1922) that this species is the 
same as Ph, leptosoma (Gervais), Seurat,1917. The close similarity 
between Gedoelst’s and Seurat’s descriptions is very striking. 
The slight differences in the lengths of the spicules, length of 
cesophagus, and position of the vulva can very well be accounted 
for as individual variations. 


Group Tetradelphys. 
(43) PHYSALOPTERA NUMIDICA Seurat, 1917. 
Male unknown. Length of female 31-9 mm., thickness 950 p. 
Two lips, each with a very large external tooth, a very small 
inner median tooth, and two bicuspid lateral teeth. Cervical 
papille subsymmetrical and situated far behind the muscular 
cesophagus, at the level of the excretory pore. Tail short and 


NEMATODE GENUS PHYSALOPTERA. 1087 


conical ; caudal pores in its posterior third. Muscular cesophagus 
short; cesophagus 1/8th of body. 

Vulva in anterior quarter of body, 3°8 mm. behind cesophagus. 
Vagina 2°16 mm. long, straight; egg-chamber long; trunk long, 
and divides dichotomously to give rise to four uteri. Recepta- 
culum seminis oval. 

Host: Dipodillus campestris. Stomach. Bou-Saada. 


(44) PHYSALOPTERA ABBREVIATA Rudolphi, 1819. 


Syn. Spiropiera abbreviata (Rudolphi), Duj., 1845. 

Body robust. Two lateral lips large. External tooth large, 
wedge-shaped, and truncated at its tip. Three inner teeth as in 
Ph. paradoxa. Muscular esophagus thick; nerve ring at its 
posterior end. CH#sophagus long, 1/3°6-1/5-4 im female, 1/4—1/5°5 
in male of body-length. 

Female. Length 7-38-20 mm., attenuated towards both ex- 
tremities. Tail conical, with caudal pores just behind its middle. 
Position of vulva variable, opening either in front of or behind end 
of esophagus; divides body into ratio of 1:2-1:6. Unpaired 
female genitalia, very long (3°5 mm.), consist of vagina, egg- 
chamber, and relatively long trunk. Four uteri arise by dicho- 
tomous division of trunk. Receptaculum seminis 150x120 p, 
sharply constricted off from both uterus and oviduct. Eggs 
50-60 x 37 uw 

Male 75-11 mm. long. ‘Tail elongate. Caudal ale large. 
Cuticular projections small and arranged in rows. Cloacal lips 
non-salient. Arrangement of papille as in Ph. paradowa. 
Spicules very unequal : left long and filiform, 2 mm. long; right 
short and stout, 180x454. Both pointed. 

This description is based on Seurat’s (1914 6, 1917 a) rede- 
scriptions of the species from worms obtained from the stomach 
of Lacerta ocellata. Bou-Saada. 


(45) PHYSALOPTERA PALLARYI Seurat, 1917. 


Length of male 8°1-10°8 mm., length of female 12°4 (larve) to 
25 mm. 

Two lips, each with its anterior border slightly trilobed. Two 
external papille very small. Hxternal tooth triangular, pointed, 
and recurved. Internal denticulate border formed of twenty 
sharp denticles. Cervical papillee at junction of two cesophageal 
parts; excretory pore at same level or just posterior. Tail of 
female very short and rounded; caudal pores in its posterior 
fifth. 

(Hsophagus long, 1/5th in the male, 1/6th in the female of body- 
length. 

Vulva 260-1200 » in front of end of esophagus. Vagina short, 
390 » long; egg-chamber 1:2 mm. long; trunk 650 » long, and 
divides dichotomously to give rise to four uterl. Hggs 65 x 32 p. 


1088 MR. R. J. ORLTLEPP ON THE 


Male tail massive, with rounded extremity. Anus opens on a 
large cushion, 100, in diameter and covered with large warts. 
First two pairs of stalked papille at the same level; 3rd pair at 
the level of the cloaca; Ist pair post-anal ventral papillae just 
behind cloaca, last pair 85 from caudal extremity. Spicules 
unequal: right pointed and thick, 150 » long; left slender, 260 wu 
long. 

Host: Agama bibront. Stomach and intestine. Marocco. 


(46) PHYsALOPTERA AFFINIS Gedoelst, 1916. 


Length of male 11-13°2 mm., of female 18°7—20°7 mm. x °8 mm. 

Two lateral lps. External tooth large and triangular; no 
median internal tooth. CHsophagus of ¢ 1/46, of % 1/5°6—-1/6°6 
of body-length. Bursa 1:4 mm. long; 2 pre-anal and 2 post-anal 
stalked papille ; 3 pre-anal ventral papille arranged in a triangle ; 
2 pairs papillee close behind anus; 2 pairs close together in middle 
of tail; 1 pair at junction of last tail quarters. 

Spicules unequal, 2°2 mm. and 270, long; club-shaped plus 
a conical tip. 

Vulva just behind esophagus or as much as 2 mm. further back. 
Four uteri. Hggs 56x40 p. 

Host: Psammophis sibilans. Intestine. Belgian Congo. 

From Gedoelst’s description this species appears to be the same 
as Ph. paradowa v. Linst. J have examined Physaloptera from 
the same host, and these have been found to belong to von Linstow’s 
species, and consequently Gedoelst’s species must be relegated to a 
synonym of Ph. paradoxa. 


Group Polydelphys. 

(47) PHYSALOPTERA TORRESI (‘Travassos, 1920). 

Syn. Zurgida torres: Trav., 1920. 

Length of male 35 to 40 mm., of female 70 to 90 mm.; 
thickness of male 1 to 1°5 mm., of female 2 to 2°5 mm. 

Two subtriangular lips, each with four external papille and 
one apical and median papilla. ‘Three teeth present on the tip 
of each lip, the median being larger and more internal in position 
than the other two; i¢ is 17m in length. Lips 85, high. 
(Hsophagus claviform, 6 to 8 mm. in length, and its muscular 
part is 400 to 430 w long 

Vulva situated about midway down esophagus. Ege-chamber 
gives rise directly to 9 to 11 uteri. Receptaculum seminis pyri- 
form, 870» long and 380, thick. Hggs 42 to 49 u long by 28 yu 
in diameter. Anus 780 to 870 4 from the posterior extremity. 

Male caudal bursa with well-deveioped ale. Four pairs of 
lateral circumcloacal pedunculated papille. Three sessile papille 
in front of the anus and two pairs of sessile papille in a row 
immediately behind the anus; three further pairs equidistant 
from each other in the anterior half of the tail. Spicules 


NEMATODE GENUS PHYSALOPTERA. 1089 


subequal and weakly chitinised, 430 to 520 u long. Anus about 
2 to 4 mm. from the caudal extremity. 

Host: Agouwti paca. Stomach. Bahia, Brazil. 

Discussion.—This species appears to be very closely related to 
Ph. dilatata Rud., 1819, the only notable difference being the 
position of 3rd, 4th, and 5th pairs of ventral post-anal papillee in 
the male. In Px. torresi these ave equidistant from each other, 
and are situated in the anterior half of the tail, the distance 
between the 2nd and 38rd pairs being small; in Ph. dilatata the 
distances between the 2nd and 3rd, 3rd and 4th pairs are about 
equal and long, while that between the 4th and 5th pairs is only 
about half this distance. 

Travassos describes and figures five external papille to each lip, 
but I very much doubt whether his observation is correct, as in 
all the species of this genus examined by me I have never seen 
more than two; he also mentious the presence of five similar 
papille in his description of Ph. turgida, but, having re-examined 
my specimens of this species, I still maintain that only two 
external papillz are present. 

There seem to be some inaccuracies or misprints in Travassos’ 
description of this species. He writes ‘ ovejector com vagina 
claviforme longa de cerca de 46 mm. de comprimento.” I think 
this should be 4°6 mm., asit is hardly likely that this organ would 
be about half the length of the worm. Further, with regard to 
the ventral pre-anal papille of the male, the description reads “3 
pares situados imediatamente acima do anus.” As he figures NG 
usual three papille in front of the anus, 1 think this sentence is 
intended to convey the fact that three papille, not three pairs, are 
situated in this position. Because of this error, the number of 
paired papille is given as 12 instead of 10 pairs in addition to an 
unpaired papilla. 

With reference to the position of the anus in the male, the 
description reads that it is ‘a cerca de 2,a 4 mm. da estremidade 
posterior.” In his French translation this sentence is rendered 
“a environ 2-4 mm. de Vextrémité postérieure.” I am accepting 
the original to be what T'ravassos meant, although from the 
construction of the two sentences it would appear that the 
translation probably describes the true state of affairs. 

It is not perhaps out of place here to draw the attention of those 
consulting Travassos’ original description of this species to the 
fact that the French translation does not always give a correct 
rendering of the Portuguese text. 


Forms with undetermined number of uteri. 
(48) PHYSALOPTERA CAUCASICA v. Linstow, 1902. 


Male 14:22 mm. long x ‘71 mm. broad; female 27 mm. long by 
1:14mm. thick. i 

Cuticle unringed. Lips with four submedian papille, and large 
cone-shaped external teeth. Cisophagus 1/5th of body- length. 


1090 MR. R. J. ORTLEPP ON THE 


Male bursa broad anteriorly and narrowed behind; small 
tubercles on its ventral face. The spicules are unequal, right 
short and broad, left long and filiform; measure -62mm. and 
1°76 mm. long respectively. One pre e-anal pair of papille (the 
median probably missed) and five post-anal pairs of ventral 
papule; the arrangement of these papille is similar to those 
found in Ph. orden, except that the relation of the 3rd and 4th 
pairs to the ventral mid-line is the reverse to that found in 
Ph. mordens. Your pairs of lateral stalked papille. 

Female tail rounded, 1/53rd of body-length. Vulva in the 
region of union of the Ist and 2nd body sixths. Hgg 57 x 39 p. 

Host: Homo sapiens. Intestine. Caucasus. 


(49) PHYSALOPTERA DIGITATA Schneider, 1866. 


Length of male 20 mm., of female 27 mm. 

Outer tooth small, inner tooth larger than outer, with its tips 
rounded. Vulva 4mm. from head endl Of ohe inner bursal 
papille, 3, 4, and 5 are more or less in a line on the ventral 
surface. Distance between 4 and 5 smaller than between 3 
and 4, Distance from 5 to tip of tail more or less equal to the 
distance between 3 and 5; 1 and 2 immediately behind anus. 
Three pre-anal papille in a triangle. Four pairs of stalked 
papille; the distance between the most anterior two pairs 
greater than that separating the others. 

Host: Felis concolor. Stomach. Brazil. 

Ph. terdentata Mol., 1860, was described from the same host, 
but it appears to be different from Ph digitata Schn., as shown by 
the difference in arrangement of its post-anal sessile papille. 
The nature of their teeth is, however, very similar. 


(50) PHYSALOPTERA BREVISPICULUM v. Linstow, 1906. 


Male 11:1 mm. long x ‘95mm. thick; female 11-4mm. long 
x1:06 mm. thick. Hach lip with three small peaks (teeth 2) 
on its summit. A preputium-like thickening over the tail is 
present. 

Cisophagus 1/6°5, tail in male 1/9°4, in female 1/22°3 of body- 
length. Spicules very short, feebly curved, -79--81 mm, long. 
Eggs 36 x31 p. 

Host: Felis rubiginosa. Stomach. Ceylon. 


(51) PHYSALOPTERA GEMINA v. Linstow, 1899. 

Length, male 11-4, female 19 mm. x °53 mm. thick. 

Two large conical external teeth; small lateral teeth ; more 
smaller ones internally. 

(sophagus 1/5-9, tail of male 1/19th, of female 1/35th of 
body-length. 

Bursa ppainted | 4 pairs lateral staiked papille arranged in two 
groups, 2 pairs pre-anal and 2 pairs post-anal. Hleven ventral 
papille, of which 3 are in a pre- -anal row, 2 in a row just behind 


NEMATODE GENUS PHYSALOPTERA. 1091 


the anus, 4 stalked and in a transverse row just in front of middle 
of tail, 2 stalked and at the junction of last tail quarters, 

Hegs 52 x 32 pw 

Host: Felis catus domesticus. Stomach and intestine. Hgypt. 


(52) PHYSALOPTERA CESTICILLATA Sonsino, 1889. 

Length 14-3 mm. and more, slightly attenuated anteriorly, 
obtuse “hetndinal, Two retractile lips, with teeth on its inner front, 
and with at least one well-developed lateral papilla. A well- 
defined collar over the lips. 

Male. Bursa of variable size, elongate and bent inwards. 
Four pairs of lateral pedunculate papillzs, and six pairs of ventral 
sessile papillae plus two unpaired papille, one of which is pre- 
anal, and the other between the last pair of ventral post-anal 
papille. Except for the posterior unpaired papille, the arrange- 
ment of the papile is similar to that found in Ph. papillotruncata. 
Ventral surface of bursa with longitudinal rows of spikes. 

Female immature. ‘Tail mucronate and anus 500 » from its tip. 

Host: Canis cerdo. Stomach. Egypt. 


(53) PHyYSALOPTERA TORQUATA Leidy, 1856. 

Males unknown. Females *25—-5 inches long by °02 inches thick 
(3-6 lines x *25 of a line). 

Body cylindrical, attenuated anteriorly; a conspicuous and 
narrow collar round the head; lips half conical, each with a pair 
of lateral papille and a group of four teeth on its apex, of which 
one is external to the others. Tail short, conical, obtuse, recurved 
from the anal aperture. 

Host: Meles labradorica. Stomach. | America. ] 

Stossich (1889) lists this species under Sp. Inq., but the nature 
of the lips and the collar round the head, together with the 
nature of the teeth, strongly suggest that this species is a true 
Physaloptera. 


(54) PaysaLoprera meputres Solanet, 1905, 

Host: Mephites suffocans. Buenos Aires. 

The description of this species, published in the ‘ Revista del 
Centro d. Hstados d. Agronomia y Veterinaria,’ Buenos Aires, 
was not available for reference.) 


(55) PHYSALOPTERA ELEGANTISSIMA Stossich, 1902, 

Length of female 50-60mm. Male unknown. 

Body large, cylindrical, and attenuated anteriorly. <A large 
more or less imbutiform cuticular expansion embracing most of 
the cephalic region except the lips. ‘'wo lateral lips, large, and 
each has three large papille—one central and two lateral. 
External tooth large and conical, supplemented by two subequal 
denticles. Tail conical and rounded. Kggs smooth and elliptical 
with very thick shell. 

Host: Ratelus capensis, Stomach, Hritrea. 


1092 MR. R. J. ORTLEPP ON THE 


(56) PHYSALOPTERA GUIARTI Garin, 1913. 


Length 35 mm., breadth 2mm. 

Body eylindrical, elongate, and yellowish in colour; trans- 
versely striated. Anterior extremity with chitinous expansions. 

Teeth absent. ; 

Male caudal extremity bent, with margin of ale lobulated and 
corrugated ; anal aperture in middle of bursa. Four pairs of 
circumcloacal papille. No internal papille. 

Host: Leptonychotes weddelli. Stomach. S. Pole. 

From Garin’s. brief description and crude drawing, this 
parasite is a Physaloptera, but the absence of teeth and internal 
papille on the tail of the male is very remarkable, and suggests 
the possibility of their having been overlooked. 


(57) PuysALoprEera ca@LeBs v. Linstow, 1897. 


Male 7 mm. long x51 mm. thick. Female unknown. 

Gisophagus 1/46, tail 1/15-4 of body-length. 

Four pairs of lateral stalled papille. ‘Three pre-anal ventral 
papillze arranged ina triangle. Five pairs of post-anal ventral 
papillae, of which the last two pairs are stalked; pairs 1 and 2 
just behind last pair of lateral papille; pairs 3, 4, and 5 in 
posterior third of tail. 

Host: Centetes ecaudatus. Madagascar. 


? 


(58) PHYSALOPLERA DISPAR v. Linstow, 1904. 


Length of male 16°6 mm. by °53 mm. broad; length of female 
25 mm. by :95 mm. thick. 

Two rounded lips surrounded by a cuticular collar. External 
tooth rectangular, with two small papille on either side; further 
back there is a larger papilla on either side. Cisophagus in 
male 1/4°9, in female 1/6°2 of body-length. 

Tail in both sexes rounded, and forms 1/19th in the male and 
1/26th in the female of the body-length. 

Caudal bursa of male with four pairs of elongately stalked 
papille; five sessile papille round the anus, one being pre-anal 
and four being post-anal ; four further pairs of post-anal papille, 
of which the most anterior pair is just behind the level of the 
last pair of stalked papille, the 2nd and 3rd pairs close together 
at about the middle of the tail, and the last towards the tail tip. 
Spicules very unequal; right broad and °35 mm. long, the left 
slenderer and °62 mm. long. 

Vulva divides the body into ratio of 3:8; eggs thick-shelled 
and embryonated, 57 x 39 p. 

Host: Hrinaceus albiventris. Stomach. Nigeria. 

Seurat (1917 6) has identified this species with his specimens 
of ‘‘ Ph. causa,” If von Linstow’s description of the lips and 
arrangement of the bursal genital papille is correct, then this 
species cannot be regarded as the same as Seurat’s specimens. 
Consequently, I think that until von Linstow’s specimens have 


NEMATODE GENUS PAYSALOPTERA. 1093 


been re-examined, his and Seurat’s specimens must be treated as’ 
separate species. 


| (59) PaysALOPTERA INCURVA v. Linstow, 1908. 


Length of male 20 mm. by ‘75 mm. broad; female 47 mm. long 
by 1:58 mm. thick. 

Two lips, dorsal and ventral; each with three teeth. 

(Hsophagus of male 1/7°6, of female 1/8th, tail of male 1/22nd, 
of female 1/235th of body- length, 

Male bursa with four stalked papillee—an unpaired pre-anal 
ventral papilla and three pairs of equidistant post-anal papille. 

Spicules unequal, right -36 mm. long and broad, left °57 mm.. 
long and narrow. 

Vulva divides body into ratio of 84:59. Hges 57 x 39m. 

Host: Erinaceus frontalis. Stomach and intestine. Kalahari. 

It is doubtful whether von Linstow’s observations in connection 
with the lips are correct. As all the other characters are 
typically those of the genus, it is probable that the lips are 
lateral in position and not dorsal and ventral. 


(60) PHYSALOPTERA CIRCULARIS v. Linstow, 1897. 


Length of male 15:2 mm.; length of female 24 mm. by 1:9 mm. 
thick. 

Two lips, each with two rounded swellings internally, and 
three external papille. 

(Hsophagus of male 1/4°3, tail of male 1/15-4, tail of female 
1/61st of body-length. 

Four pairs of circumcloacal stalked papille ; one pair of sessile 
papille in front of, and an unpaired papilla just behind anus ;. 
three pairs of post-anal ventral papillae, of which pairs 1 and 
2 are close together at about the junction of the first two. 
quarters of the ail, and the 3rd at about the middle of the tail. 

Eggs with very thick shells, 49 x 33 p. 

Host: Mus rattus. Stomach Madagascar. 

It is quite possible that von Linstow “missed an unpaired pre- 
anal sessile papilla and four small sessile papillee just behind the 
anus. If this should later be found to be the case, then there 
would be reasonable ground for supposing this species to be 
either the same as, or very closely related to Ph. muris- 
brasiliensis Dies. 


(61) PHYSALOPTERA RUWENZORII Parona, 1907. 


Length of male 9-93 mm., of female 14-17 mm. 

Body attenuated for a short space anteriorly. Large lips with 
small papille. Teeth large and conical. Esophagus 1/6th of 
Yody-length. 

Male caudal bursa large, with large circular expansion anteriorly 
and conical terminal portion; cuticular ornamentations absent ; 


Proc. Zoou. Soc.—1922, No. LX XIII, 73 


1094 MR. R. J. ORTLEPP ON THE 


four pairs of lateral and circumcloacal stalked papille; a large 
pre-anal unpaired ventral papilla; behind anus two pairs of 
ventral papille and a large unpaired papilla in front of them. 
Spicules unequal, pointed and slightly bent. 

Tail of female long and attenuated. Vulva divides body into 
ratio of 1:2; it is slightly raised and papiilate. Eggs oval, 
thick-shelled, containing partially developed embryos. 

Hosts: Arvicanthis abyssinicus. Intestine. Fort Portal, 

Africa. 
Epimys ugande (Mus ugande). Intestine. Fort 
Portal, Africa. 

(Compiled partly from Parona and Hall’s (1916) descrip- 

tions.) 


(62) PHySALOPTERA ScIURI Parona, 1898. 


Length of male 16 mm., of female 17-32 mm.; breadth of 
male ‘5 mm., of female ‘5-1 mm. 

Two large and semicircular lips, with foliaceous external 
tooth. 

Caudal bursa ef male large and lanceolate; 4 pairs of lateral 
stalked papille situated circumcloacally; a large unpaired papilla 
close behind the anus; 2 pairs of ventral papille situated 
marginally and at about the middle of the tail; a pair of small 
ventral papilla near the tip of the tail. The bursa is ornamented 
with numerous tubercles. The anus is large and circular, and 
has a scalloped and raised edge. 

Tail of female obtuse. Vulva non-protuberant, and divides 
the body more or less into the ratio of 1:2. Single uterine tube. 
Eggs small and spherical, with a diameter of 16-22 p. 

Host:  Sciurus melanogaster. Stomach and intestine. 
Mentawei. 

As Hall (1916) has pointed out, Parona’s observations with 
regard to the uteri is probably incorrect; in all other Physalo- 
ptera in which the number of uteri has been determined they 
are never less than two. 


(63) PHYSALOPTERA INERMIS v. Linstow, 1906, 


Length of male 21:5 mm. by 1:26 mm. thick; length of 
femaie 51 mm. by 1°62 mm. thick. 

(Esophagus of male 1/3°8, of female 1/8°5, tail of male 1/21st, 
of female 1/425th of body-length. 

Bursa with four stalked papille on either side and four pairs 
of equidistant ventral and post anal papille; the most anterior of 
these are situated at about the middle of the tail. Longitudinal 
striations on ale. 

Spicules 2°37 mm. long. 

Vulva divides body into ratio of 11:17. Eggs 47 x 26 p. 

Host: Sciwrus prevosti. Stomach. Koenigsberg. 


NEMATODE GENUS PHYSALOPTERA. 1095 


(64) PHysaLoprera citinui (Rud., 1819), Hall, 1916. 


Syn. Spiroptera citilli Rud., 1819. 


Male unknown ; female 7-13 mm. long by 400 p thick. 

“ Body curved. Anterior extremity very slightly attenuated. 
Head rounded, continuous with the body, and with the cuticle 
inflated. Mouth with two large lips. Six head papille, of 
which there are two conical papille situated on each side near 
the angle of contact of the two lips, and one spherical papilla at 
the base of each lip. Cuticle densely striated transversely. 
Posterior extremity of female obtuse. Vulva in anterior portion 
of body.” 

Host: Citellus citellus. Stomach. 

Hall (1916) transferred this species to Physaloptera, partly on 
the opinion expressed by Molin and partly. because the description 
could apply to a member of this genus. I follow both these 
authorities, and consequently list it as a valid member of the 
genus Physaloptera. 


(65) PHYSALOPTERA PYRAMIDALIS v. Linstow, 1879. 


Length of male 18 mm. by | mm. thick; length of female 
50 mm. x 1:5 mm. thick. 

Two pyramid-like lips, dorsal and ventral in position. Outer 
tooth pointed, inner tooth of same size and tripartite. Ciso- 
phagus 1/57, tail 1/28th in male and 1/55th in female of body- 


length. Two pre-anal and two post-anal stalked papille in 


caudal extremity of male; in addition 4 pairs of unstalked 
papille, of which one pair is pre-anal and 3 pairs post-anal. 

Vulva at junction of Ist and 2nd body fifths. Kgegs very 
thick-shelled, 42 « 33 pw. ; 

Host: Cholepus didactylus. ‘An den Sehnen der Zehen.” 
Surinam. 

This species appears to differ from Ph. papillotruncata by its 
pointed outer tooth and by the more anterior position of the 
vulva; nevertheless I am inclined to think that this species is 
identical with or a variety of Ph. papillotruncata. 

It is possible that von Linstow is mistaken in the position of 
the lips, and that they are really lateral. 


(66) PHYSALOPTERA LIMBATA Leidy, 1856. 


Length -5 inch, breadth -02 inch (6 lines x :25 of a line). 

Mouth two-lpped and surrounded by an elevated linear 
margin. Caudal extremity of male alated and having four pairs 
of divergent and stalked papille. 

Host: Scalops canadensis. Stomach. Philadelphia. 

Molin (1860) lists this species under Sp. Inq.; however, in 
view of the fact that there are two lips surrounded by what may 
be interpreted as a cuticular collar, and that the male bursa is 
alated and has four (pairs) of stalked papille, I think there is 
good ground for supposing that this species is a true Physaloptera, 

13* 


1096 MR. R. J. ORLTLEPP ON THE 


(67) PHYSALOPTERA NASILIONIS Gedoelst, 1916. 


Larve: length 6 mm. .Two lateral lips. External tooth 
triangular. CHsophagus 2°23-2°33 mm. long. Nerve ring in 
posterior quarter of muscular esophagus. Tail conical and 
blunt, 215 p long. 

Host: asilio br ead ynchus. ‘issu conjonctif.” Belgian 
Congo. 


(68) PHYSALOPTERA BILABIATA (Creplin, 1829), Dujardin, 1845. 


Two large lips, prominent and obtuse. Body much attenuated 
anteriorly. Caudal extremity of male reflexed, alate, elliptical, 
with four stalked papille. Tail of female conical and obtuse. 

Length, ¢ 0°025, 2 ultra 0:025; breadth 0-001. 

Host: Lanius minor. Intestiue. (Extracted from Stossich’s 
Monograph.) 


(69) PHYSALOPTERA MEGALOsTOMA Creplin, 1829. 


Syn. Spiroptera megalostoma Duj., 1845. 
Ph. alata Stoss., 1889. 


Head naked, with large and simple mouth. Caudal extremity 
of male with ale up to its tip; female straight, with rounded 
extremity. 

Host: Falco nisus. Proventriculus. Greifsw. 

This species is considered by Molin (1860) and Stossich (1889) 
to be the same as Ph. alata Rud. 


(70) Puysatoprera 1nFriarA (Mol., 1860), Stossich, 1889. 
Syn. Spiroptera inflata Mol., 1860. 
Length of female 7 mm.; male unknown. 
- Body striated and attenuated posteriorly. Cuticle of head 
inflated; no internal teeth on lips. Caudal extremity conical, 
with an obtuse apex. Vulva in anterior part of body. 
Host: Falco unicinctus. Stomach and esophagus. Brazil. 


(71) PHYSALOPTERA CRASSA v. Linstow, 1879. 


Male 14 mm. long by 1°5 mm. thick; female 23 mm. long by 
2 mm. thick. 

Lips pyramid-shaped, each with two outer papille and two 
median teeth, of which the inner is tripartite. Cisophagus 1/4-7 
of body-length. Cervical papille -6 mm. from anterior end, 
situated dome: ally and ventrally. 

Tail of male rounded, 1/11:7 of body-length. Bursa with six 
pairs of papille, of which the anterior five are stalked and 
arranged in a longitudinal row close to the anus; the 6th and 
unstalked papille are found on the inner side of the 5th pair. 
Cirri unequal, -66 and *36 mm, long. 

Tail of female rounded, 1/22nd of body-length. Eges 49 p by 
26. 
Host: Alauda arvensis. Intestine. Stuttgart. 


NEMATODE GENUS PHYSALOPTERA, 1097 


(72) PHYSALOPTERA TRUNCATA Schneider, 1866. 


Male 25.mm. and female 33 mm. long. 

Outer tooth knobbed, ‘mit beilartiger Schneide.” Inner 
teeth heart-shaped. 

Four pairs of lateral and stalked papille on male tail; the 
distance between the 2nd and 3rd somewhat greater than that 
between the Ist and 2nd, and between the 3rd and 4th. Three 
pre-anal papille in a row, the central one of which is small. The 
ist and 2nd pairs of post-anal ventral papille small and just 
behind anus ; three larger pairs further down. The ratio of the 
distances between the 2nd and 3rd, the 3rd and 4th, the 4th and 
Sth, the 5th and the tip of the tail is about-as 1:1:2:2. 

Host: Phasianus gallus. Stomach. Brazil. 


(73) PHYSALOPTERA FUSIFORMIS v. Linstow, 1902. 


Male 18 mm. long by 1°38'mm. thick ; female 30 mm. long by 
1°97 mm. thick. 

Two rounded lips, each with three small papille on outer 
edge. Cisophagus of male 1/3°8, of female 1/5°3 of body-length ; 
tail of male 1/23rd, of female 1/20th of body-length. Bursa 
small and heart-shaped; four pairs of lateral stalked papille; one 
pair of sessile papille in front of anus and separated from each 
other; a pair of sessile papille just behind anus; an unpaired 
papilla in middle of tail, and three more pairs behind it and 
equidistant from each other. 

Vulva in anterior half, and divides body into ratio of 13: 56. 
Eges 81 x 26 p. 

Host: Micropogon sp. Breslau Museum. 


(74) PuYsALOPTERA DENTAYA v. Linstow, 1883. 


Male 7 mm. and female 12 mm. long and -96 mm. thick. 

Two large lips, each with a large terminal outer tooth and 
spike-like inner tooth. Cervical papille -48 mm. from head end. 
(Hsophagus 1/4:2, tail of male 1/11th, of female 1/25th of body- 
length. 

Bursa lanceolate, ale ornamented with conical tubercles 
arranged longitudinally ; four pairs lateral stalked papille, all 
are pre-anal; one unpaired papilla in front of anus and three 
pairs behind it; pairs 1 and 2 close together in middle of tail, 
often fused foection:) pair 3 at junction of last two tail quarters. 

Eggs 52 x 43 p. 

Hosts: Agama sanguinolenta. Intestine. Turkestan. 

Vipera berus. Q 93 
Phrynocephalus auritus. Mouth and intestine. _,, 


(75) PHYSALOPTERA BRITANICA Skrjabin, 1916. 


Length of immature female 42 mm., breadth 6mm. Male 
unknown. 


1998 MR. R. Je ORTLEPP ON THE 


(isophagus 5°44 mm. long. Cervical papille “6 mm. from 
anterior end. ‘Tail conically pointed, -46 mm. long. Narrow 
alee on both sides of body. 

Vulva 4:25 mm. from head end. 

Two large lateral lips, each with large external tooth; on 
inner face a series of smaller triangular and pointed denticles on 
either side of median tooth ; 10-11 of these on either side, and 
the 5th and 6th of them, from the external tooth end, are large. 

Host: Agama sp. Intestine. British Hast Africa. 


(76) PHyYSALOPTERA SONSINOI v. Linstow, 1895. 


Length of male 6°5 mm., of female 6°7 mm. and -48 mm. 
thick. 

A conical tooth on top of each lip. 

(sophagus of male 1/4°6, of female 1/2-75, tail 1/9°25 in male, 
and 1/18th in female of body-length. 

Male caudal bursa elongate; 4 pairs of circumcloacal stalked 
papille; 3 pre-anal sessile papille in a triangle; 2 large and 
oval just behind anus, and slightly further back 2 smaller and 
rounded papilla; in middle of tail 4 papille in a transverse row, 
and further back 2 pairs, of which the anterior pair is large, and 
each papilla of which contains 2 pulps. 

Right spicule 1:89 mm. long; left spicule -32 mm. long and 
much broader. 

Vulva divides body into ratio of 5:31. Hggs 57 x 39m. 

Host: Agama mutabilis. Stomach. Pisa. 

Since the spicules are so unequal, it is possible that von Linstow 
has mistaken the right for the left. 


(77) PHYSALOPTERA ALOISII-SABAUDIA Parona, 1907. 


Male 6 mm., female 11 mm. long. 

Body attenuated in anterior third. Two large lips with large 
tooth, and with a series of denticles and one papilla on each lip. 
(Hsophagus 1/4th of body-length. 

Male bursa lanceolate ; without tubercles on its surface. Anus 
surrounded by papillee; four lateral papille longly pedunculate ; 
2nd and 38rd post-anal pair slightly stalked. Spicules large, long, 
and unequal. 

Female with short tail; eggs oval, small, thick-shelled, and 
embryonated. 

Host: Agama atricollis. Intestine. Africa. 


(78) PHYSALOPTERA SPIRALIS Schneider, 1866. 


Length of male 14-22 mm.; length of female 29-67 mm. 

Outer tooth pointed, inner tooth absent. Inner face of lips 
provided with denticles. A pair of these behind the tooth in 
the middle of the lip; on either side of it an additional pair 


towards the edge of the lips; dorsally and ventrally a row of 
about five. 


NEMATODE GENUS PHYSALOP'TERA, 1099 


Tail of male straight. Four pairs of lateral stalked papille ; a 
pair of sessile papille in front of anus and 3 pairs post-anal: 
of these latter the last pair is situated at about the junction of 
the last tail thirds, and the other two pairs are situated close 
together just in front of the middle of the tail. 

in the largest females the vulva is 10 mm. from the anterior 
end. Posterior part of body spirally coiled and tail slightly bent 
dorsalwards. 

Host: Amphisbena sp. Stomach. Brazil. 


(79) PHYSALOPTERA ANTARCTICA v. Linstow, 1899. 


Length of male 27 mm. by °79 mm. thick; female 42 mm. long 
by 1:18 mm. broad. 

Two lateral lips, somewhat rounded; each bears a conical 
outer tooth and two small inner side teeth. 

(Hsophagus of male 1/8°3, of female 1/8°6 of body-length ; tail 
of male 1/23rd, of female 1/54th of body-length. Male cloacal 
aperture large and rounded, and on either side of it four papillee 
with long stalks. Thirteen ventral papille, of which 3 are pre- 
anal, 4 just behind the anus in two rows, 2 on either side behind 
each other in middle of tail, and further back another one on 
either side. 

Tail of female conical, with ronnded tip; eggs very thick- 
shelled, 54-44 x 42-31 p. 

Hosts: Acanthophis antarctica. S. Australia. 

Cyclodus occypitalis. Cisophagus. 8S. Australia. 

I have identified this species with my specimens from Varanus 

varius and Python spilotes. 


(80) PHYSALOPTERA ALBA Stossich, 1902. 


Male 25-28 mm. long; female 42 mm. long. 

Weakly-developed lips, each with two large submedian papille 
and with very weak teeth. 

Male bursa lanceolate, with minute ornamentations on its 
ventral surface. Four pairs of lateral papille with long peduncles. 
Five pairs of ventral papille arranged in two groups, viz. three 
pairs towards the end of the tail and two pairs just behind 
the anus, of which the outer pair has double pulps. 

Tail of female conical. Vulva at the anterior third of the 
body. Eggs embryonated, elliptical. with very thick sheli. 

I have identified this species with Ph. antarctica v. Linst., 1899. 

Host: Cyclodus boddaertii. Proventriculus. Nuova Olanda. 


(81) PHyYsALOPTERA AMPHIBIA V. Linstow, 1899. 


Male 12 mm., female 18 mm. long by 1:05 mm. broad. 

Two half-rounded lips. External tooth large and cone-shaped, 
internal tooth small and tripartite. Cisophagus 1/4:4 in the 
male and 1/83 in the female of the body-length. 


‘1100 .. MR, R. J. ORTLEPP ON THE 


‘Tail of male 1/15°5, of female 1/82nd of body-length. Bursa 
‘with four pairs of stalked lateral papillz arranged in two groups 
separated from each other; Ist group of two papillee i is pre-anal, 
2nd group is post-anal. Four pairs of ventral sessile papillee, of 
which one pair is pre-anal and the other three are in the 
posterior half of the tail; the central pair of these last three 
pairs is slightly nearer the front than the hind pair. 

Kggs 55 x 29 p. 

Host: Kana macrodon. CHsophagus and ventriculus. Luzon. 


SPECIES INQUIRENDA, 


(82) Puysatoprera ABsecta Leidy, 1856. 


Length of female °83 inches, breadth -04 inches (10 lines x °5 
of a line). Male unknown. 

‘‘ Body most narrowed anteriorly, incurved. Mouth distinctly 
bilabiate ; lips lateral, prominent. Caudal extremity incurved, 
‘obtusely conical.” 

Host: Psammophis flagelliformis. Stomach. [U.S.A.] 


(83) PHYSALOPTERA BREVICAUDA v. Linstow, 1909. 


Two lips widened outwardly, their pulp narrower at its base ; 
a papille in their middle, and right and left of it a cone-shaped 
tooth. 

_ Male 27mm. long x °83 mm. thick; female 42 mm. long 
x ‘95mm. thick. ‘l'ail of male rounded, 1/69th of body-length. 

Bursa somewhat heart-shaped and ale ornamented with longi- 
tudinal strie. Six pairs of stalked papille—4 pre-anal and 2 
post-anal; the post-anal pairs are situated in a transverse 
row just behind the anus. Spicules equal, 1-4 mm. leng, and 
rounded at their tips. 

Vulva just behind middle of body ; it divides body into ratio 
23:21. Hggs 89 x 26 p. 

Host: Francolinus adspersus. Intestine. S.W. Africa. 

The shape of the lips, arrangement of the male caudal papillae, 
length of the right spicule, position of the vulva, and small size of 
the eggs, seem to indicate that this species is not a Physaloptera. 
From the description I am not able to place it in any known 
genus, and consequently list it under Sp. Ing. 


(84) PHYSALOPTERA MALLEUS v. Linstow, 1883. 


Length of male 20mm. by ‘76mm. thick; length of female 
42 mm. by 1:5. mm. thick. 
Two very large and rounded lips. Cisophagus 1/6°5, tail of 
male 1/33:3, of female 1/269th of body-length. Bursa oval, with 
_rounded prominences arranged in transverse rows. Six pairs of 
stalked papille, of which 4 pairs are pre-anal, and 2 pairs are 


NEMATODE GENUS PHYSALOPTERA. 1101 


post-anal in position ; these last situated just in front of middle 
of tail. Spicules unequal, the larger having a double barb at its 
tip. Eggs double-shelled, the ater being the thicker ; they are 
46 x 29 pu, 

Host: Corvus cornix. Turkestan. 

This is probably not a Physaloptera, as evidenced by the 
rounded lips without teeth, the arrangement of the bursal 
papillz, and the presence of a hook on the spicule. 


(85) PHYSALOPTERA PAPILLORADIATA v. Linstow, 1899, 


Length of male 32 mm., of female 56 mm. by 790 uw thick. 

Two rounded lips. 

CHsophagus 1/12th, tail of male 1/7th, of female 1/127th of 
body-length. © 

Bursa with 4 pairs pre-anal and 2 pairs post-anal stalked 
papille; 7 pairs of post-anal ventral papille, of which 1 pair is 
just posterior to the anus, and the remaining 6 pairs are aggre- 
gated at the posterior end of the tail. 

Eggs 26 x 12 p. 

Host: Canis lupus. Pectoral cavity. Persia. 

As figured by von Linstow, the arrangement of the caudal 
papill is very similar to that of Ph. rotuwndata v. Linst., except 
that in rotundata there are 5 papille at the base of the tail, 
Whereas in papilloradiata there are 12. I have placed this 
species under Sp. Inq. because of this similarity, and because 
Seurat has found Ph. rotundata to be a Habronema. 


(86) PHYSALOPTERA SPIRULA Hempr. & Ehrenb., 1828. 


Male 6mm. long; female unknown. Head continuous with 
‘body. Mouth papillate. Caudal extremity of body provided with 
closed bursal alee. 
Hosts: Procavia syriaca. 
Procavia capensis. 
(Extracted from Hall, 1916.) 


} Large intestine. Syriaand Arabia. 


SPECIES WRONGLY ATTRIBUTED TO THE GENUS. 


(87) PHysaLoprerRA RorUNDATA v. Linstow, 1906. 


Length of male 15°6 mm. by 1-1 mm. thick; length of female 
20°3 mm. by 1:5 mm. thick. Both ends of body rounded. 
Two large lips, each bearing a prominent papilla on each side. 
(Hsophagus 1/8°6 in male, 1/9th in female, tail in male 1/40th, 
-in female 1/20th of body-length. 
Bursa narrow, with 6 pairs of lateral stalked papille, of which 
4 pairs are pre-anal and 2 pairs post-anal. Two pairs of ventra: 
papille at tip of tail, and just in front of them an unpaired 
papilla. Spicules unequal ; right 510 p, left Bos long, soe 
with rounded tips. 


1102 , MR. R. J. ORTLEPP ON THE 


Vulva divides body into ratio of 6:15. Hegs 39 x 26 p. 
Host: Otis houbara. Intestine. Koenigsberg. 
Seurat (1914 d) has identified this species as a Habronema. 


(88) PHYSALOPTERA BULBOSA v. Linstow, 1906. 


Host: Pavo spicifer. Stomach. 

I have recently found specimens of this parasite in the proven- 
triculus of Pavo cristatus from India. From my observations on 
this material I find that this parasite is not a Physaloptera, but 
belongs to the genus Cyrnea Seurat, 1914, A detailed description 
of it will appear in a later publication. 


(89) PHyYSALOPTERA conToRTA Leidy, 1856. 


‘Body capillary, most narrowed anteriorly with the posterior 
4/5th spirally contorted ; white, with the intestine brown. Lips 
prominent, constricted from the body, trilobate. ‘Tail short 
conical, acute, in the male with narrow ale each furnished with, 
five funnel-shaped pores. Length of female from 6 lines to an 
inch ; breadth to one quarter of a line; male from one-half to 
three-quarters of the size.” 

Hosts: Hmys serrata, Emys reticulata, Cistudo carolina, and 
Kinosternum pennsylvanicum. Stomach. [U.8.A.] 

Baylis and Lane (1920) suggest that this species is probably 
identical with or closely related to Spiroxys contorta (Rud., 1819), 
Schn., 1866. I have been able to examine some specimens of 
Spiroxys contorta obtained from Emys orbicularis, and from my 
observations Baylis and Lane’s view appears justifiable. Accord- 
ing to Leidy’s description, these parasites must be excluded from 
the genus Physaloptera because of their trilobed lips. Molin 
(1860) has already doubted the Physalopteran affinities of this 
species. 


(90) PHYSALOPTERA consTRicTA Leidy, 1856. 


Length, female 13 inches, male half as long. 

Lips large, lateral, constricted from body, and trilobate. Tail 
of male with long and narrow turgid ale; that of the female 
incurved, conical, and acute. 

Host: Lropidonotus sipedon. Stomach. [U.S.A.j| 

The nature of the lips definitely excludes this parasite from 
the genus Physaloptera, no members of it having trilobed lips. 
Baylis and Lane (1920) suggest that this parasite may be 
a Tanqua. I agree with them in so far that it falls mto the 
family Gnathostomide as redefined by them, but as to its closer 
relationship to any of the genera of this family, it does not appear 
justifiable to draw any conclusions; Leidy’s brief description of 
the parasite is equally applicable to the genus Spirowys as to the 
genus J'anqua. 

Molin (1860) also expressed doubt as to this species belonging 
to the genus Physaloptera. 


NEMATODE GENUS PHYSALOPTERA. 1108 


(91) PuysALoprerA MUCRONATA Diesing, 1851. 


Host: Alligator mississippiensis. Georgia. 
This parasite has been found by Molin to be an Ascarid, and 
von Drasche has identified it with Ascaris lanceotata Mol., 1860. 


(92) PHyYsALopTERA ovata v. Linstow, 1907. 


Length of male 13°83mm.; length of female 16°8mm. by 
1-22 mm. thick. 

Mouth surrounded by 6 cones. Césophagus of male 1/4°4, of 
female 1/3-5, tail of male 1/14th, of female 1/531d of body ‘length. 

Bursa oval, with 5 pairs of se bane and lateral stalked 
papille, the ond being the longest; 2 pairs are pre-anal. 
Spicules equal 290 long. 

The vulva divides the body into the ratio of 13:31. Hggs 
narrow, 42x 13m. 

Host: Astur melanoleucus. Ventriculus. Cameroon (Kamerun). 

The nature of the lips rules this species out of the genus 
Physaloptera ; this is supported by the nature of the bursal 
papille and by the extremely narrow egg. 

The description is too incomplete to definitely say to which 
genus it should be transferred, but it appears to have some 
resemblance to the genus: Oyrnea Seurat, 1914. 


(93) PHYSALOPTERA SAGINATA Rud., 1819. 
Syn. Spiroptera saginata (Rud.), Duj., 1845. 


I examined examples of this species from Corvus cajanus and 
Caprimulgus guianensis while at Vienna, and am consequently 
able to confirm Molin’s (1860) view that this species is a Spiro- 
ptera. 


(94) PHYSALOPTERA STRIATA v. Linstow, 1883. 


Length of male 18mm., breadth -48mm.; length of female 
25mm., breadth :66 mm. 

Two dorso-ventral lips, small and rectangular, having a wavy 
anterior border and divided into three parts internally ; 4 small 
interlabia. Tail of male spirally rolled, 1/37:°5 of body-length ; 
tail of female rounded, 1/83rd of body-length. 

Caudal alee of male with longitudinal striations. Four pairs of 
pre-anal stalked papille; 1 unpaired papilla in front of anus; 
2 pairs post-anal and dividing tail roughly into thirds. 

Vulva divides body into ratio of 3:2. Hggs 46 x 25 p. 

Hosts: Tropidonotus hydrus. Turkestan. 

Ciconia alba. Stomach. Turkestan. 
_ Seurat has obtained a Cyrnea from different species of Ciconia, 
which he identifies with Ph striata. This species is undoubtedly 
not a Physaloptera, and from von Linstow’s figure and description 
of the lips I think Seurat is correct in transferring it to his 
genus. 


1104 “MR. Re J. ORTLEPP ON THE 


(95) PHYSALOPTERA sTRONGYLINA Rud., 1819. 
Syn. Spiroptera affinis Duj., 1845. 
Length of male 7:87 mm., of female 15-25 mm. to 20°25 mm. 
Body white and head appears to have papille; tail of male 
with lateral ale, so that it resembles a Strongylid tail. Spicules 
fairly long and thick. 
_ Host: Cuculus seniculus. Stomach and intestine. Brazil. 
Molin ruled this species out of the genus Physaloptera, stating 
that it belonged to a new genus. 
The description is too incomplete to warrant it being referred 
to any now known Nematode genus. 


(96) PHYSALOPTERA TENUICOLLIS Rud., 1819. 

Syn. Spiroptera tenwicollis (Rud.), Duj., 1845. 

Male unknown ; length of female 41 mm., breadth 1:15 mm. 
-Mouth rounded; anterior end sharply attenuated for a 
distance of 2°25 mm. 

Host: Malco haliaétus. Intestine. 

Molin (1860) has identified this species as an Ascaris. 


BiBLIOGRAPHY. 
Bayuts, H. A., & 1920. A Revision of the Nematode Family Gnathostomide. 
Lang, C. Proc. Zool. Soc. London. 
CREPLIN, I. C. H. 1829. Nove observationes de entozois. Berolini. — 
Dresine, K. M. 1851. Systema Helminthum. Vindobone. 
e ss 1861. Revision der Nematoden. Sitzungsb.d. k. Akad. d. 
Wissensch., Wien, math.-naturw. Cl. (1860), vol. xlii. (28). 
DRASCHE, R. von. 1883. Revision der in der Nematoden-Sammlung des k.-k. 


zoologischen Hofcabinetes befindlichen Original-Exem- 
plare Diesing’s und Molin’s. Verhandl.d.k.k. zool.-bot. 
Gesellsch. in Wien (1882), vol. xxxii. 


Dusarptin, F. 1845. Histoire naturelle des helminthes ou vers intestinaux. 
Paris. 
Fantuam, H. B., 
Stevens, J. W. W., - 1916. The Animal Parasites of Man. London. 
& THEOBALD, I. W. 


Garin, C. 1913. Recherches physiologiques sur la fixation et le mode 
de nutrition de quelques Nématodes parasites du tube 
digestif de ’homme et des animaux. Ann. Univ. Lyon, 

Fasc. 34. Jiyon. 


GEDorEtst, L. 1911. Synopsis de parasitologie de Vhomme et des animaux 
domestiques. ILierre et Bruxelles. 
ee 1916. Notes sur la faune parasitaire du Congo belge. Rev. 
Zool. Afr. vol. v. fase. 1. Bruxelles. 
Hatt, M.C. 1916. Nematode Parasites of Mammals of the orders 


Rodentia, Lagomorpha, and Hyracoidea. Proc. U.S. 
Nat. Mus., vol. 1. Washington. 


Hatz, M.C., & 1918. A Physaloptera from the Dog, with a note on the 
Wiaepor, M. Nematode Parasites of the Dog in North America. 
Journ. Amer. Vet. Med. Assoc. vol. liii. n.s. vol. vi. 

No. 6. 


Heet,J.NooRDHOEK. 1910. Chlamydonema felineum, Nov. Gen., Nov. Spec., eine 
neue parasitisch lebende Nematode. Tijdsch. der Nederl. 
Dierk. Vereen., 2nd ser. pt. xii. Leiden. 


Henry, A., & 


Branc, G. 


Trwin-Smita, V. 


Lerpy, J. 


Lerrer, R. T. 


” 


23 


Linstow, O. von. 


29 


o> 


39 


” 


39 


BB) 


3 


9 


39 


39 


32 


39 


NEMATODE GENUS PHYSALOPTERA. 1105 


1912. Le physaloptére du Macacus cynomolgus, L. Bull. 


Soc. Path. Exot. vol. vy. Paris. 


1921. Note on Nematodes of the genus Physaloptera, 
with special reference to. those parasitic in Reptiles. 
Part i1.. Proc. Linn. Soc. N.S.W. vol. xlvi.pt.4. Sydney. 

1922. Ibid. Part ii. A Review of the Physaloptera of 
Lizards. Proc. Linn. Soc. N.S.W. vol. xlvii.pt.2. Sydney. 

1904. Researches in Helminthology and Parasitology ; with 
a bibliography of his contributions to science, arranged 
and edited by Joseph Leidy, jr. Smithson. Misc. Coll. 
vol. xlvi. No. 1477. Washington. 

1908 a. Physaloptera mordens : a new intestinal parasite of 
Man. Trans. Soc. Trop. Med. & Hygiene, vol. i. London. 

19085. An account of some Helminthes contained in 
Dr. Wenyon’s collection from the Sudan. Rep. Welle. 
Res. Lab. Khartoum, vol. ii. London. 

1911 a. Notes of Recent and some New Records of Hel- 
minthes in Man of which there are few records. Journ. 
Lond. Sch. Trop. Med. yol.i. pt. 1. London. 

19116. On the frequent occurrence of Physaloptera 
mordens as an intestinal parasite of Man in Tropical 
Africa. Journ. Trop. Med. vol. xiv. London. 

1911 ¢. Some new parasitic Nematodes from Tropical 
Africa. Proc. Zool. Soc. London. 

1913. Observations on certain Helminthes of Man. Trans. 
Soc. Trop. Med. & Hygiene. London. 

1877. Helminthologica.. Arch. f. Naturg. 43 J., vol. i. 
Berlin. 

1878. Compendium der Helminthologie. Hannover. 

1879. Helminthologische Untersuchungen. Jahresch. d. 
Ver. f. vaterl. Naturk.in Wiirttemb.vol. xxxv. Stuttgart. 

1883. Nematoden, Trematoden und Acanthocephalen, 
gesammelt von Prof. Fedtschenko in Turkestan. Arch. 
f. Naturg. 49 J., vol. i. (2). Berlin. 

1889 a. Helminthologisches. Arch. f. Naturg.54J., vol. i. 
(3). Berlin. 

18896. Compendium der Helminthologie. Nachtrag. Die 
Litteratur der Jahre 1878-1889. Hannover. 

1895. Untersuchungen an Nematoden. Arch. f. mikr. 
Anat. vol. xliv. Bonn. 

1897. Nemathelminthen grésstentheils in Madagascar 
gesammelt. Arch. f. Naturg. 63 J., vol. i. (1). Berlin. 
1899. Nematoden aus der berliner zoologischen Sammlung. 
Mitt. a. d. zool. Sammi. d. Mus. f. Naturk. in Berlin, 

vol. i. Berlin. 

1902 a. Beobachtungen an neuen und bekannten Nemathel- 
minthen. Arch. f. mikr. Anat. vol. lx. Bonn. 

1902 6. Zwei neue Parasiten des Menschen. Centralbl. f- 
Bakt., Pavasitk. 1 abt. vol. xxxi. Jena. 

1904 a. Neue Helminthen aus Westafrika. - Centrabl. f: 
Bakt., Parasitk. 1 abt. vol. xxxvi. Jena. 

19046. Entozoa des zoologischen Museums der Kaiserlichen 
Akademie der Wissenschaften zu St. Petersburg. Ann. 
Mus. Zool. Acad. Imp. d. Se. d. St. P. vol. viii. 

1906 a. Nematoden des zoologischen Museums in Kénigs- 
berg. Arch. f. Naturg. 72 J., vol. i. Berlin. 

19066. Helminthes from the collection of the Colombo 
Museum.  Spolia Zeylanica, vol. iii. pt. 2. Ceylon. 

1907. Nematoden aus dem Ko6niglichen zoologischen 
Museum in Berlin. Mitt. zool. Mus. vol. iii. Berlin, 


1106 


Linstow, O. von. 


99 ” 


Morn, RB. 


Parona, C. 


29 33 


Rupowpuy, C. A. 


ScHNEIDER, A. 
ScuppeEr, S. H. 


Srurat, hl. G. 


3) 29 


SxKRJABIN, K. f. 


Sriuzs, C. W., & 
Hassaut, A. 


MR. R. J. ORTLEPP ON THE 


1908. Helminthes. Nematoden und Acanthocephalen. 
(In L. Schultze, Zool. u. Anthrop. Ergeb. e Forschungs- 
reise in Siid-Afrika, Bd. i. Lfg. 4.) Denkr. Med. Ges. 
vol. xi. Jena. 

1909 a. Parasitische Nematoden. In: Die Siisswasser- 
fauna Deutschlands (Brauer), Heft 15. Jena. 

1909 6. Neue Helminthen aus Deutsch-Stidwest-Afrika. 
Centralb. f. Bakt., Parasitk. abt. 1, vol. 1. Jena. 

1860. Una monografia del genere Physaloptera. 
Sitzungsb. d. k. Akad. d. Wissensch., Wien, Cl. vol. xxxix. 
(5). Vienna. : 

1889. Sopra alcuni elminti di vertebrati birmani raccolti 
da Leonardo Fea. Ann. Mus. civ. di Storia nat. di 
Genova, vol. xxvii. (ser. 2, vol. vii.). Genoa. 

1898. Elminti raccolti dal Dott. Elio Modigliani alle 
isole Mentawei, Engano e Sumatra. Ann. Mus. civ. di 
Storia nat. di Genova (1898-99), vol. xxxix. (ser. 2, 
vol. xix.). Genoa. 

1907. Nuove specie di nematodi africani. (Nota preven- 
tiva.) (Spedizione al Ruwenzori diS. A. R. Luigi Amedeo 
di Savoia, duca degli Abruzzi, 24.) Boll. mus. di zool. ed 
anat. comp. d. r. Univ. di Torino (586), vol. xxii. 

1819. Entozoorum synopsis, cul accedunt mantissa duplex 
et indices locupletissimi. Berolini. 

1866. Monographie der Nematoden. Berlin. 

1882. Nomenclator Zoologicus. Bull. U.S. Nat. Mus. 
No. 19. Washington. 

1914 a. Sur un nouveau Nématode parasite des Reptiles. 
C. R. Soc. Biol. vol. Ixxvi. Paris. 

1914.6. Sur deux Physaloptéres tétrahystériens des Reptiles. 
C. BR. Soe. Biol. vol. Ixxvii. Paris. 

1914c. Sur les Physaloptéres des Rapaces. Bull. Soc. Hist. 
Nat. d. ?Afr. d. Nord. 6thyear, No.9. Alger. 

1914d. Sur un nouveau parasite du Perenoptére. Bull. 
Soc. Hist. Nat.d.VAfr.d. Nord. 6th year, No.6. Alger. 

1915. Un nouveau Physaloptére des Rapaces. Bull. Soc. 
Hist. Nat. d. ’Afr.d. Nord. 7th year, No.9. Alger. 

1915-16. Contributions 4 ]’étude des formes larvaires des 
Nématodes parasites hetéroxénes. Bull. Scientif. d. 1. 
France e. d. 1. Belgique. 

1916. Nematode Parasites. Novitates Zoologice, vol. xxiii. 
No.1. London. 

1917 a. Physaloptéres des reptiles du Nord-Africain. C. R. 
Soc. Biol. vol. }xxx. Paris. 

1917 5. Physaloptéres des Mammiféres du Nord-Africain. 
C. R. Soe. Biol. vol. xxx. Paris. 

1918. Contribution a l’étude de la Faune Parasitaire de la 
Tunisie. Nématodes. Arch. d. l’Inst. Past. d. Tunis, 
vol. x. pt. iv. 

1900. A description of the Entozoa collected by Dr. Willey 
during his sojourn in the Western Pacific. Willey’s 
Zool. Results based on material from New Britain, New 
Guinea, &e. Pt. v. 

1905. Notes on a collection of Parasites belonging to the 
Museum of University College, Dundee. Proc. Camb. 
Phil. Soe. vol. xiii. pt. 2. 

1916. Parasitic Trematodes and Nematodes collected in 
British East Africa. 

1920. Index-Catologue of Medical and Veterinary Zoology. 
Subjects: Round-worms. Hyg. Lab.-Bull. No. 114. 
Washington, 


Sonstno, P. 


Stosstcu, M. 


ZooLroGicaL REcoRDSs. 


GVOVOIMEKMENIGY, | Seo sande. cadon 
abjecta scat ae 
acuticauda ............... 


EYATUTUIS La5 We aah soe anS 


alata v. chevreuxi ...... 
alata v. nouvelli ...... 
eulogy: by dagen Geraa atonal 
aloisii-sabaudix......... 
amphibia era Sa 
HAAN), so5 doasoh bss vende: 
AVTIANLOWNUCE) sas eho bnoneuboe 
[OWA ANBL coco ese cde nodane 
DOMAIN e Nien eee 4 
jbrevicaudal s4.y.2 20). 
brevispiculum ......... 


brevivaginata 


NEMATODE GENUS PHYSALOPTERA. 


JOVAAWIVIOR) oo caaconesoseacce 
[DEMNNOSD, cooseodco oss nasnos 


GAVDSINENG, casdansoncaeens o60 


eaucasica ... 


Geshicullavarcces en ceeeee 
chameleontis............ 


circularis ... 


Gib lie ren Geasecane 
GURUS, vncrdodnoaed snaasoses 
ODES iacasdobuconvanates 


colubri 
constricta ... 
contorta ...:.. 


(CUMS) Goo cogeod oe0 pUaueo COG 


erosi 


dentata rt ih 
QUICHE, coc sdcosuccsooeon- 


GhIENIPIEBY Ge boo oo, eolsosooe 


GUISTOEWE coo ecoone 
elegantissima 


AUSIRORMUNSH eeeeeeee es 
(SPNTMNIMEM ceovassss 605 coo ane 


gemina 


FRAME, coe sb0 060000 


eracilis 


ATER noo coo sdonuacsoconeee 
THOVGBUAVE) sageno canses sa0586 


jnermis 


TOW OLEH Ape cite AU nesduee eee 


leptosoma ... 


1899. Studi e notizie elmintologiche. 
Se. Nat., Mem. vol. x. 


Pisa. 


1107 


Atti Soe. tuse. d. 


1889. Il genere Physaloptera, Rudolphi: Lavoro mono- 


grafico. 


Bull. Soc. Adriat. d. Sc. Nat. in Trieste, vol. ii. 


1902. Sopra alveuni nematodi della collezione elmintologica 
del Prof. Dott. Corrado Parona. 
Comp. d. Genova, No. 116. 


1919. Material Helminthologico da Ilha da Trinidade. 


Archiv. do Mus. Nacional, vol. xxii. 


Boll. Mus. Zool. e Anat. 


1920. Contribuicdes para o conhecimento da fauna helminto- 
lojica brazileira. X. Sobre as especies do genero Turgida. 
Mem. do Inst. Osw. Cruz, vol. xii. pt. 1. Rio de Janeiro, 

1900-1920. Pt.: Vermes. 


INDEX TO SPECIES, 


Number Pages 
44, 1087 
82 1100 
18 1050 
46 1088 

7 1025 
36 1083 
37 1083 
80 1099 
77 1098 
81 1099 

5 1021 

25 & 79 1067 
68 1096 
11 1034 
83 1100 
50 1090 
35 1083 

5 1097 
88 1102 
30 1078 
48 1089 
52 1091 
42 1086 
60 1093 
64 1095 

il 1012 
5 1092 
26 1070 
90 1102 
89 1102 
71 1096 
39 1085 
74 1097 
49 1090 
29 1076 
58 1092 
55 1091 
73 1097 
40 1085 
51 1090 
34 1082 
13 1039 
56 1092 
59 1093 
63 1094 
70 1096 
41. 1085 


TMD, Seance sedascco 
HOMESITE, cancoo son cobane 
magnipapilla ............ 
mialayensisiessees aoe 
OEMS ,56460 S56 400 ova ¢on 
THERUILEYBIS)  coeecooesootede 
megalostoma ........ 
TANGO NUNES 556 060.500 cam oso000 
WAOMOIENS “5.4 annees cae 
TTDOROINS son ang scocosoagess 
TONDKCROMEM, sosssecsqsnen oe 
muris-brasiliensis 


nasilionis BaUee eS 
MUTCD roe cos ceases concen 
obtusissima 

ovata Se ea Sous iis 
allan. yee eee 


papilloradiata 


papillotruncata ......... 
DDTENCIOy. 50 yocideonoson ave 


phrynosoma 


preputialis..........:.... 


pyramidalis 
quadrovaria 


FURIE eon ene Maguearoneanate 
TROLADUXO NIE) sso doo soontacdo 
APWNKIIVAOVE|, cogone con conden 
SPYCAITENES G45 sho wocloosoogtes 
S(OMDUA, sedasouss aboadoar 


semilanceolata 


simplicidens ........ 
SONSINOW epepeseeeeeeee 


spiralis 
spirula 


Shiulabaese ee ere eee 
strongylina........... 
SOL ORUIIEH ED ccgoc ncoone soooee 
VEXCEY NEVES Guocoocosgouoen 
(MOUNDS -oas op seo booes 
WHEOKEYM ENE) Goo incoss cos eco 
WOPRONWENIE) Sacasocod saeco beh 
PORES TE URSA NEY i sate Mian 
LHCDDIVORRED) Son cavuso soecon oon 


tumefaciens 
turgida 


AEN We Wscs ouoanncereeannns 


Number 


66 


ey ee 
sii ao 
Naa nh 


whe 


THE OLIGOCHATA OF THE SPITSBERGEN EXPEDITION. 1109 


49. The Oligocheta of the Oxford Teas ones ee 
Expedition. By J. SrspHmnson, M.B., D.Sc., Lecturer 
in Zoology, Hdinburgh University. 


[Received October 9, 1922: Read November 21, 1922. ] 


(Vext-figures 1-6.) 
(Results of the Oxford University Expedition to Spitsbergen, 


No. 20.) 

ConTENTS. 
Page 
Introduction ......... budannldcnibids Song uetannubene dan eecane scenery utente a0 E102) 
Zoozeographical Raleivions, 5 ul cee ee Oso VATE 
On the “ Peptonephridia ” of ihe Enchy sdoe Baas emacs aaa aem ame el tl ies 
The Genus Henlea.......... scoon ULI 
On the Relations of the Cen Tannbr Piles hail nonin UB o0.058 1118 
IPermiodks Orr excell MATT Gos ccoccuebbconaessouee copocecooucnasoeosseooseoee | MINIS 
JEENPASTICIPEN BOI Rac opauaeacu cod doasee aon ode ead on areeane adenanc one Sexe sracentr heats niu ILS) 
{SIV SUIGLETE NIG a3 casei cp Bes ack Seo mad aon Guede decree do eee Ee OSE CHE Eee Hae Sesame tanya LIL 
HE) aura el WAU OD Te Sait at Oe cie ete cud etched atti clot Ac aes gmadveaar Macnee: MOLES, 
Genus WNais ....... aceon Sate oe a geicumn a dct ak a iedanaee HOLE 
Nais josine Veja. Poet ROP GaD RoenSS aN cian tn seta sae THEMES ts ed Lt 
ABT AUTrY WEIN GHETYEINR Alt TeD areas eerste care evr atccee a a namie re rd entree eeLeL TIO) 
Genus Henlea ......... Daoeceidon neces) 
Henlea (Henleanelia) heleotrophus, sp. NG. 'a eoomee podeoe, | BLY 
Henlea (Henlea) brucei, sp. e Ua Sioa ae tten oe eaen ee eal 
tenean (Hentecnella) tsps Cl) eressss-c eee eee eo 
Henlea (Hepatogaster) sp. O SAREE ie URN ee TLE 
Genus Lumbricillus ......... smaetlazael bs caer 2 ek ee INOS 
Lumbricillus egialites, sp. Mow Bonooaehageenenscaccacoeunoccats, Lao) 
Enumbricillus necrophagus, Sp. W....-..2.2+00..2--+0-----.. L180 
Genus Mnchytreus ...... NASIR ete See eee ad ie on UOTE 
Enchytreus albidus Henle. ilar oy Sates een Ua ne rue TUL 
Enehytreus crymodes, sp.N...... Sree ea neeto cho ceotoasde el ulleys) 
JD PORTERS, Ss) ‘doc aco sod ssesdneso soo soosbosdssbdvcs coscbende: LUGE) 
Ci@aws AMASPCOMURCUS  secGbccdecscocensieks covers voecceddbecacoone , HIBS 
ICSAC ERGEDS. So cosnsnoasddcodes ‘osodbo evo seoonosonacsccecce | LUBE 
IRONS 110) LOMMORMATOD | Gogdao sce Cagosonos cebbUbESeubhaboser ecasbuecedocuccaeo LIE 


INTRODUCTION. 


The Oligocheta obtained by the Oxford University Spitsbergen 
Expedition were contained in fifteen tubes—tour from Bear Island 
and the remainder from Spitsbergen. The specimens from Bear 
Island belonged to two species—AHnchytreus albidus Henle and 
Nais josine Vejd.,—both well known. Of those from Spitsbergen, 
a number were sexually immature and therefore indeterminable 
(Mesenchytreus sp., two species of LHenlea, Hnchytreus sp.); the 
remainder, most of which occur in more than one of the tubes, 
comprise two species of Henlea, one of Hnchytreus, and two of 
Lumbricillus, ali apparently new. With the exception of the 
Vais from Bear Island, all the worms thus belong to the family 
Enchytreide. 

Proc. Zoou. Soc.—1922, No. LX XIV. 74 


1110 DR, J. STEPHENSON ON THE OLIGOCHETLA OF THE 


Our previous knowledge of the Olgocheta of Spitsbergen and 
Bear Island is due almost entirely to Ude (17), who, in addition 
to collections from Bear Island (then examined for the first time), 
had also at his disposal a small collection from Spitsbergen, sent 
to him by Michaelsen and tentatively named, the names having 
already been entered in the ‘Tierreich ’ volume on Oligocheeta (11). 

The following species are all that have been recor ded: — 

From Spitsbergen :— 
Lembricillus pagenstechert (Ratz.) (originally determined 
and entered in the Tierreich volume as ZL. nervosus 
(Hisen) ). 
Fridericia leydigt (Veja.). 
Mesenchytreus sp. indeterminable. 
From Bear Island :— 
Marionina ebudensis (Clap.) 
Lumbricillus fossarwm (Tauber). 
Lumbricillus henkingi Ude. 
Enchytreus albidus Henle. 


The only one of these collected by the Oxford Expedition is 
Enchytreus albidus. All, as will be seen, belong to the 
Enchytreide. 

The Kochytreide have always furnished the great majority of 
Oligocheta from high latitudes, and there omits a considerable 
number of records “of species of the family from Nova Zembla, 
Northern Siberia, North Russia and Norway, Greenland, Avetic 
Canada, and Alaska—especially species of the genera owen and 
Mesenchytreus. Welch has in recent years (20, 21, 22) drawn 
attention to previous records of Enchytreeidze found actually on 
ice and snow, and has himself examined and described a number 
of species from high latitudes, and from snowfields and glaciers. 
Some of these worms show a resistance to cold that can only 
be described as marvellous. J/esenchytreus gelidus was found 
“abundantly on the snowfields”; it “also occurred on the snow 
on the mountain slope in a dense forest of fir and hemlock,” and 
‘‘on the snow below the ice front and outside of the lateral 
moraines of the Nisqually glacier; ... when placed on hard- 
packed snow during their active period, they are able to bore 
down through it at will; .. the snow on which they are found 
is not permanent through the entire season, but melts with the 
coming of summer, and ‘it therefore appears ‘that a part of their 
life-history must be spent on or in the ground.” Mesenchytreus 
solifugus var. rainierensis was “abundant on the higher snow- 
fields and glaciers of Mt. Rainier in early summer ; it was found 
on snowfields which seldom thaw during the sqrernitier. and they 
evidently pass the entire existence, generation after generation, 
in the snow and ice.” Beddard (1) mentions that an Enehytreid 
has been found frozen in a block of ice,and recovered. Mr. Elton 
writes that he has subjected the worms which I have named 
Enchytreus crymodes to an experiment in which they were frozen 
solid and that they survived. 


OXFORD UNIVERSITY SPITSBERGEN EXPEDITION. WIT 


Tt was thus to be expected that Enchytreids should constitute 
the bulk of the collection. ‘here is no mention of snow in the 
data velating to the worms; and since they thus pass at least a 
part of their life on or in ground free from snow, the conditions 
under which they live ave those of Welch’s Hnchytreus gelidus 
rather than of MW. soli/ugus var. rainierensis. 


ZOOGHOGRAPHICAL RELA TIONS. 


The Oligocheta may be divided, with reference to their habitat, 
into limnic, littoral, and terrestrial forms; and the Oligocheta 
of Bear Island and Spitsbergen may be classified under these 
headings as follows: 


LIMNIc. Litrorat. TERRESTRIAL. 
Nais josine. Lumnbricillus necrophaqus. Hlenlea brucei. 
Henlea heleotrophus. Lumbricillus egialites. Enchytreus albidus. 
Henlea spp. (2). Lumbricillus fossarum. Fridevicia leydigi. 
Enehytreus erymodes. Lumbricillus henkingi. Mesenchytreus sp. 
Enehytreus sp. Lumbricillus pagenstecheri. 
Mesenchytreus sp. Marionina edudensis. 


Enechytreus albidus. 
Hunchytreus erymodes. 


There is no record of the habitat of some of the previously 
recorded species: of Lumbricillus jossarum, which, however, is 
elsewhere found on the shore, and is therefore so entered here; 
of Hnchytreus albidus vecorded from Bear Island by Ude, which 
is also entered as littoral, since all his specimens from Bear Island 
of which the habitat is recorded were littoral, and the worm is 
elsewhere common on the shore; and of Mridericia leydigi, which 
is entered as terrestrial, since most of the Fridericias are so. 

All the smaller Oligochzeta, and therefore all the forms which 
are represented in the present and former collections from Spits- 
bergen and Bear Island, are easily transported, and hence are of 
only slight value for considerations of zoogeography. Limnic 
forms have a great diversity of means of dispersal; they may 
spread directly throughout a river system—though this, cf course, 
has no bearing on the present enqairy; their cocoons are easily 
transported in the mud which adheres to the feet of wading 
birds; some forms are known to encyst, and hence may be trans- 
ported by birds even in the adult state. As a consequence of 
this easy distribution, the same genera—sometimes even the 
same species-——are found in widely distant places; the case is 
similar to that of the Rotifera and Protozoa, of which the same 
genera and species are found in ponds and streams all over the 
world. ‘To illustrate by means of an example, there appears to 
be but one genus, Branchiodrilus, of limnie Oligocheta which is 
peculiar to India, while a number of species are found both in 
india and England, or India and Europe (species of ais, Cheto- 
gaster, Dero, Aulophorus, Pristina, etc.). 

74% 


TAL WS DR. J. STEPHENSON ON THE OLIGOCH ETA OF THD 


Littoral forms, living on the shore and exposed at times to 
immersion in salt water, have additional means of dispersal. Being, 
unlike Oligocheetes in eeneral, immune to salt water, they can be 
transported in masses of seaweed ; er more commonly their 
cocoons are so transported, entangled in masses of weed or other 
detritus. Not only can they take possession of a whole coast, and 
spread along the shore-line, but they may in this way travel over 
sea for long distances. Pontodrilus, for example, which occurs 
along the coasts of India, has a circummundane distribution. 

Even terrestrial Oligocheta are by no means always dependent 

on their own unaided activities for enlarging thei: distribution ; 
there are a considerable number which are especially liable to be 
transported accidentally—by man or other agencies,—and which 
noe an especial capacity for establishing themselves where they 
ey cte to be deposited. The smaller the worms the easier is 
their dispersal; certain small species of earthworms of the genus 
Dichogaster, which has its proper home in Africa, have spread all 
round the warmer regions of the globe. 

Few, however, possibly none, of the worms of Spitsbergen and 
Bear Island are exclusively terrestrial. Of those given as 
terrestrial in the above table, Hnchytreus albidus is often found 
on the shore; it is perhaps the most widely distiibuted of all 
Enchytreids, and oecurs all over Europe, in North America, and 
in South America as far as Patagonia and Tierra del Fuego, as 
well as within the Arctic circle 11 Greenland and Nova Zembla 
The species of Mesenchytreus found at Cape Boheman among 
plants in dry tundra was also obtained among mosses on the 
banks of a freshwater pond, more or less under water, and is 
therefore limnic as well as terrestrial. The Henlea found among 
Dr yes etc. on a hill near the coast of Freshwater Bay was also 
found in the Bruce City region, possibly in a limnic or littoral 
habitat (details of habitat are wanting for this tube). ridericia 
leydigi, previously obtained from Spitsbergen, 1s probably the 
most strictly terrestrial of all the above species, occurring’ else- 
where in earth rich in humus and under moss. 

‘There can thus be no question of anything peculiar in the 
Oligochete fanna of Spitsbergen and Bear Island. All the 
geneva are easily transported, and all (except perhaps the Naidid 
genus Vais) have previously been found within the Arctic circle 
—Henlea in Nova Zembla, N. Siberia, N. Russia, N. Norway, 
and in the Canadian Arctic; Lnchytreus in N. Russia, Green- 
land, and the Canadian Arctic; Leumbricillus in Nova Zerbla, 
N. Siberia, the Canadian Arctic, and Greenland; Mridericia in 
Nova Aembla, N. Russia, and N. Siberia. All these genera also 
oceur in Alaska. 

As to whether particular species of Spitsbergen Oligocheta 
have been evolved locally, or imported, it is as yet impossible to 
say, until neighbouring lands have been more thoroughly 
explored. ‘(here is some shght ground for supposing that some, 
at least, may have been evolved on the island. On the whole, no 


AFORD UNIVERSIVY SPITSBERGEN EXPEDITION, Wis} 


doubt, it is the commoner species of the neighbouring lands that 
are already known to us: and on the whole, no doubt, it is the 
commoner species that, from their frequency, have most chance 
of being transported. If it had been throughout a case of simple 
transportation, therefore, we might have expected to find that 
a larger number of the species of worms were known from else- 
where also. 

It seems hazardous to suppose that any of the present Oligo- 
chete fauna could be descended from ancestors existing 11 
Spitsbergen before the glacial period. It is true that we now know 
species which can live their lives through, and reproduce them- 
selves, on ice and snew. The food of such worms, according to 
Welch, writing of the specimens from Mt. Rainier, seemed in one 
case (Mesenchytreus gelidus) to be microscopic alge having the 
appearance of Plewrococcws, and the snow over which the worms 
were crawling had a red colour due to a minute unicellular plant ; 
in the case of the other worm (J. solifugus var. rainierensis) the 
food could not be determined. 

But the food-supply of glacier and snow worms must, one would 
say, be precarious: and it seems dangerous to assume its unfailing 
presence throughout the long duration of the glacial period. 

On the other hand, there are apparently no facts which seem 
to contradict “the most reasonable hypothesis of the origin of 
the present fauna—namely, an ‘accidental’ peopling since the 
period of maximum glaciation, during which time Spitsbergen 


to) 
has been as widely separated from adjacent lands as it is now.” 


On tHE ‘‘ PEPTONEPHRIDIA”’ OF THE HNCHYTRAIDA. 


Certain structures connected with the first portion of the 
alimentary canal in a number of Enchytreids are known as 
‘“peptonephridia,” or sometimes as “ salivary glands.” 

(1) Lhe best known of these structures are a pair of tubules 
which arise from just behind the pharynx, and extend backwards 
for some little distance free in the body-cavity. They are well 
developed, for example, in Hnchytreus albidus, a worm which is 
abundantly represented in the present collection. Here the 
tubules open into a dorsal diverticulum of the alimentery canal 
just behind the pharyngeal mass: they sometimes extend back- 
wards into segm. v., while sometimes they do not transgress the 
hinder limit of segm. iv.; their general course is sinuous or 
contorted, and in addition the outline of the walls is marked by 
numerous smaller sinuosities or crenations. The diameter of the 
tubules varies—it may be about 50 py, or may be less; that of the 
limen may be 42» or less; the diameter is least near the origin 
of the tube from the alimentary canal, where the whole structure 
is only 16, thick, and the lumen is very small. The walls are 
mostly thin, about 4» thick, but in places are much thicker— 
as much as 15 .; numerous nuclei are seen in the walls—in a 
transverse section of one of the tubes there may be about six, 
z.e, the lumen is intercellular. 


1114 DR. J. STEPHENSON ON THE OLIGOCHMTA OF THE 


2) In the genus Henlea there are often found other organs, 
situated in the middle line, one dorsally and one ventrally on 
the cesophagus, In segm. vi., sometimes extending forwards into 
v., and backwards behind vi. for, it may be, several segments. 
The association with the wall of the alimentary tube is here much 
more intimate than in the case of the former structures; these 
are closely apposed to the wall of the cesophagus for the whole or 
the greater part of their extent; they may even be contained for 
a Honcidemble part of their Oren within the wall, between the 
epithelial and muscular coats UWielen in H. moderata (19), which 
T can confirm in 4. brucei and A. heleotrophus). They frequently 
give branches. 

While the former structures have little about them that can 
remind the observer of uephridia, these have much more the 
appearance of the peculiar type of nephridium found in the 
Enchytreide—a spongy-looking, searcely staining tissue, with 
nuclei but no visible cell-limits; sometimes there seems to be no 
lumen in the interior—nothing more than the interstices of the 
spongy cell-substance, sometimes there are irregular spaces and 
tubular cavities, eal sometimes a single swell: defined central 
lumen. They have, however, no ciliated funnel or any kind of 
internal opening. Cejka, in the worms which he ealled Hepato- 
gasier, states (3) that there are indications of the double origin of 
these apparently single and median structures; in this I agree 
(compare the transverse section of a ventral organ of H. brucei, 
fig. 1; the description of the ventral organ in Henlea sp. (1), post., 
and of the dorsal organ in H. heleotrophus, where possibly there 
is no connection between the masses on the right and left sides) ; 
while Welch finds that in H. ochracea (22) the organs are actually 
paired, one pair dorsaily and one pair ventrally. It is, perhaps, 
this type of gland which is intended by Friend in his reference 
to the salivary glands of #. alba (7) and A. inwsitata (8), and to 
the “special g a ands” of H, bisetosa (8). 

(5) There is a third type of organ, which is apparently widely 
spread in the family. Deser iptions of these str uctures are given 
for all the newly described species below, and they also occur in 
the Mesenchytreus; LI have described and illustrated similar 
structures In a dadion worms Enchylreus barkudensis (14, and 
see also pl. vi. fig. 1 in 15) and in Mridericia carmichaeli (15). 
Briefly, these organs consist of a pair of rounded, or ovoid and 
bulb-like, masses of ceils, solid, situated behind the pharyngeal 
mass on the dorsal side of the alimentary tube, and connected 
with the tube by a stalk, which is at any rate partly solid, but is 
usually hollow in its proximal portion—i.e., the cavity of the 
alimentary tube may penetrate the basal portion of the stalk. 
Ude probably had this type of organ before him when he described 
the rudimentary salivary glands of Bryodrilus ehlersi (16), two 
small pear-shaped bodies, in which no lumen could be recognized 
immediately behind the pharynx; and, indeed, since I first met 
with these structures some years ago, I believe I have myself 


OXFORD UNIVERSITY SPITSBERGEN EXPEDITION. WLS 


conceived them as a rudimentary kind of organs of the first 
type. 

Whether all these structures should go by the same name— 
whetker they are all modifications of one original structure— 
seems doubtful. Those of the second type ave quite possibly 
nephridia. It will be remembered that the Enchytreids have, as 
a rule, no nephridia in the first six segments (Bretscher (2) has 
established a separate genus for a worm in which the first 
nephridium occurs in segm. ili., the funnel being in ii.); and the 
occurrence of dorsal and ventral ‘ peptonephridia” in segm. v1., 
extending thence, it may be, forwards or backwards, or both 
forwards and backwards, their obviously similar constitution, and 
their fundamentally paired nature would seem to justify the 
assumption that they are really the nephridia of seem. vi. which 
have become associated with the esophagus. If, however, we look 
on both dorsal and ventral organs as fundamentally paired (as we 
seem justified in doing, v. aé.), we should then have two pairs of 
nephridia belonging to segm. vi. In ZH. brucei, in addition to 
dorsal and ventral “ peptonephridia ” in segm. vi., there are also in 
the same segment—indeed in the same section—a pair of ordinary 
nephridia (the first nephridium being here in segm. vi. with the 
funnel in v.), i.e., éiree pairs in the one segment. 

Whether the organs discharge an excretory substance into the 
cesophagus, or a digestive juice—or, indeed, whether they discharge 
anything at all,—is doubtful. Cejka says that the openings are 
always hard to find in sections; from which phraseology it would 
seem doubtful whether they actually have definite mouths ; Welch 
does not mention any openings, nor have I found any in my pre- 
parations. On the whole, they are perhaps more likely to be 
excretory, since they have an intimate relation to the blood- 
spaces (in H. leptodera they run in the esophageal sinus and are 
directly bathed by the blood (Cejka, 4); in Hepatogaster also they 
are bathed by the blood in the sinus, which may penetrate 
between the individual cells; I have found the same conditions, 
v. inf.)- 

But the nephridial nature of the first group of organs, which 
have for long passed under the name of ‘“ peptonephridia ”—the 
sinuous or coiled tubes which extend back, from the pharynx, 
fr doubtful. These do not show the 
peculiar structure of the Enchytreid nephridium ; they have xno 
internal opening; indeed, the only feature that is brought forward 
to substantiate their nephiidial nature is the intracellular lumen. 
But in “Hnchytreus albidus (v. ant.) the lumen is certainly not 
intracellular. And im any case there is no fundamental differ- 
ence between intercellular and intracellular tubes: whether the 
Jumen of any tube is intracellular or not depends on the size of 
the lumen and the size of the cells of which the tube is composed. 
Suppose we start with a tube of fair size, in the cross-section of 
which several cells are seen to bound the lumagas uf the size of the 
tube is diminished, fewer cells will serve to surround the cavity, 


1116 DR. Je STEPHENSON ON THE OLIGOCHETA OF THE 


and fewer will be seen in any transverse section. Still diminishing 
the width of the lumen, two cells, and finally one single cell, will 
go all round the lumen—z. ¢., only one cell will be seen in any given 
transverse section of the tube, and the lumen will be described as 
intracellular. ‘There is thus really no reason for assuming the 
nephridial nature of these tubules of the Huchytreide, and in my 
opinion they should not be called nephridia. 

The third group of structures—the bulb-like organs—are even 
more unlike nephridia; nor, except for their asennad connection 

with the tubular organs first described, could it ever be supposed 
that they were nephridial. 1t is possible that they are, as has 
been suggested, vestigial remains of the tubules; they are con- 
nected to the same dorsal diverticulum of the alimentary canal as 
these. In Mnchytreus albidus, however, I find, along with the 
typical tubules, and just in front of and lying against the narrow 
proximal part of these, rather indefinite groups of cells somewhat 
similar to the bulb-like organs. To establish the homology of the 
bulb-like organs with the tubules it would be necessary to ascer- 
tain definitely whether they do or do not occur along with these, 
and whether or not intermediate forms exist. 

The second group of organs, then—the dorsal and ventral 
‘“peptonephridia” of the genus Henlea,—may provisionally be 
taken to be nephridia. But there is no evidence whatever of 
their digestive function, and I propose for them the name 
esophageal nephridia. 

The first group—the sinuous tubules—are quite possibly not 
nephridia at all. Since the alternative name salivary glands has 
long been in use for them, J would reject the name peptonephridia 
in favour of this. 

The third group—the solid bulb-like organs—do not seem to be 
nephridial, and their function, if they have one, is quite unknown. 
I propose a purely descriptive term, the postpharyngeal bulbs, as 
most suitable for them. 


Toe Genus Henlea. 


Henlea is a large genus, and contains species showing a wide 
variety of structure. Characteristic of a number of species are 
the cesophageal sacs in or about segm. vill.; and the question 
arises whether the presence or absence of these sacs should not be 
made the basis of a separation into two genera, or at least sub- 
genera. 

This course has already been proposed by Friend (7, 8)—forms 
with esophageal diverticula to be called Henlea, those without to 
receive the name Henleanella. In the second of the papers just 
referred to, Friend remarks that it would be advisable to make a 
third group for those species in which the alimentary tube dilates 
gradually to form the intestine. In a subsequent paper (9). 
Friend proposes a different criterion for the third group, viz., the 
origin of the dorsal vessel in the region of the Glitelium (in most, 


OXFORD UNIVERSILY SPITSBERGEN EXPEDIVLION. 1117 


species the dorsal vessel arises considerably in front of the 
clitellum) ; this third group to be called Henleana. ‘These divisions 
are apparently conceived as subgenera (certainly so in the first 
two papers). 

Friend's suggestion as regards the cesophageal diverticula seems 
particularly useful. But it is illogical and inconvenient to have 
2 double or triple principle for the simultaneous division of a 
group; the subgroups will not be mutually exclusive. What, for 
example, would become of a form with both cesophageal diver- 
ticula and a posterior origin of the dorsal vessel—-at the same 
time a Henlea and a Henleana; or, for that matter, of one with a 
posterior origin of the dorsal vessel and no diveiticula—both 
a Henleana and a Henleanella ? 

In 1910 Cejka (8) established a genus Hepatogaster for two 
species which aie obviously nearly related to Henlea, the chief dis- 
tinctive characters being the gradual dilatation of the cesophagus 
to form the intestine, the presence of a mass of tubules closely 
surrounding the alimentary tube in segm. vill., and of character- 
istic longitudinal canals in the epithelium of the hinder end of 
the digestive tract. The value of the first distinction is doubtful, 
e.g. in Henlea heleotrophus I am not certain whether or not the 
intestine can be said to arise as a sudden dilatation of the 
alimentary tube; at first I thought not, but later I altered this 
in my notes by adding the words “ this might be called a sudden 
widening, and so would agree with the diagnosis of Henlea.” 
The second of Cejka’s points of distinction occurs also in Henlea 
tubulifera Welch and H. moderata Welch; while, as Welch 
remarks (23), the value of the third distinction as a generic 
character remains to be demonstrated. Welch retains the genus, 
though with some doubt. Delphy (5) also considers it as distinct. 

The cesophageal diverticula of Zenlea, when present, may have 
walls which are fairly smooth and even, or which are variously 
folded; and I regard the ‘‘tubuliferous” condition of H. tubu- 
lifera, etc., and of Hepatogaster, as simply a higher degree of this 
folding, followed by fusion between the adjacent folds and 
between the two sacs above and below the alimentary tube. An 
intermediate condition is that of H. nasuta as described by 
Michaelsen (10, cf. especially pl. fig. 1), where the folding of the 
walls is very considerable, and there is only a very small central 
free space, but the two sacs are not fused above and below the 
cesophagus. The difference between Henlea and Hepatogaster 
seems, in fact, to be lessthan that between Henleanella without 
sacs and Henlea with them. 

For the present, then, I would retain the genus Heilea, but I 
would recognize three subgenera :— 

(1) Henleanella, with no cesophageal diverticula. 

(2) Henlea, with cesophageal sacs. 

(3) Hepatogaster, with a mass of tubules surrounding the 
alimentary tube and taking the place of the sacs. 

I would disregard the sudden or gradual widening of the tube 


1118 DR. J. STEPHENSON ON THE OLIGOCH ATA OF THE 


in these subgeneric distinctions ; but it is possible that a posterior 
origin of the dorsal vessel will be a sufficient ground for removing 
out of the genus altogether the forms in which it occurs. 


On THE RELATIONS OF THE GENERA Lumbricillus AXD 
Enchytreus. 


IT have previously (13) pointed out the close relationship 
between the above two genera, and described several species with 
intermediate characters ; Mnchytreus dubius, which I then placed 
in the genus Hnchytreus, 1 should now assign to Lumbrieilius. 
Welch also has some remarks that bear on the subject 
(19, 23). ; 

The distinguishing characters of the two genera may be said to 
be (1) the setee—-the distal portion of the shaft being straight in 
Hnchytreus, curved, so that the sete have the usual double 
(sigmoid) curve, in Lumbricillus; (2) the copulatory glands— 
usually absent in Hnehytreus, usually present in Lumbricillus ; 
(3) the testes, divided in Lumbr icillus, undivided in Hnchytreeus 5 
and (4) the penial bulb—single, compact, encapsuled, and 
penetrated by the vas deferens in Luwmbricillus, represented by 
discrete groups of cells around the end of the duct in Hnchytreus. 
In addition, red blood would be in favour of Lumbricillus, colour- 
less blood move in favour of Hnchytreus. 

Of the species described in the present paper, Lwmbricillus 
egialites has the straight sete of WHnchytreus, and so has 
L. necrophagus; while neh yireeus erymodes has a penial bulb of 
the lumbricilline type, copulatory glands, and hemoglobin in the 
blood. These species thus bear out what I have pr eviously written 
regarding transitional forms, and render a sharp separation 
between the genera impossible. Still less is there such a sharp 
distinction between the two as would justify their position as the 
representatives of two subfamilies, the Enchytreine and Lum- 
bricilline, eatablished by Hisen (6). 


PERIODS OF SEXUAL Marurity. 


Some of the worms in the collection were sexually mature, 
others immature. But it is not possible to draw any general 
conclusions from the data regarding tke season of maturity in 
high latitudes. Anchytreeus albidus was mature on Bear Island 
in June and #. crymodes in early August in Spitsbergen; but an 
umature Hnchytreus was also coll ected on August 6th. Henlea 
heleotrophus and H. brweet were mature in early August— 
Hf. heleotrophus also as ay as July 1l0th; but other species of 
Hlenlea weve still immature in early August. The two species of 
Lumbricillus were mature towards the end of July and beginning 
of August; the JMesenchytreus, however, was immature at any 
rate up to the end of July. 


OXFORD UNIVERSITY SPITSBERGEN EXPEDITION. 1119 


PARASITIZATION. 


Finally, I may be allowed to draw attention to the excessive 
parasitization of most of these worms, which goes much beyond 
anything that I have previously seen in members of this group. 
Not only were there numerous parasites, of more than one group 
of Protozoa, in the alimentary canal, but many of the worms had 
cysts in the genital region, body-wall, or ccelom, in which spores 
were forming. It seems probable that the frequent structural 
changes in the alimentary canal, as well as other changes in the 
contents of the ccelom, may be due to this parasitization. 


SYSTEMATIC. 
Fam. NAIDIDZ. 
Genus Nats Mull. em. Vejd. 
Nats sosinz Vejd. 


Bear Island; in littoral region of a tarn near Walrus Bay, in 
the water; 15.vi.1921. A single specimen, non-sexual. 


Fam. HNCHYTR IDA. 
Genus Hernia Mich. 


HENLEA (HENLEANELLA) HELEOTROPHUS, Sp. 1.* 


Spitsbergen; among mosses on the bank of a fresh-water pond, 
more or less under water, Bruce City. August Ist-8th, 1921. 

Spitsbergen, Bruce City region (no data regarding habitat) ; 
10.viii.1921. One specimen from each of these tubes. 


Length 8-10 mm.; diameter °4mm. Segments 40, with a few 
more not yet differentiated at the hinder end. Colour brownish 
in the first tube, black in the second (fixed with Bouin). Prosto- 
mium bluntly triangular, the triangle being about equilateral. 
Head-pore between prostomium and first segment. No dorsal 
pores. 

Clitellum on segments xii. and xiii., well marked. 

Sete enchytreine in form (7. e., straight in their distal portion), 
the tip mederately sharply pointed. In frort of the elitellum the 
ventral bundles have apparently four setz (not well seen), behind 
the clitellum 4, 5, 6,and once 7; the lateral bundles have usually 
five sete, sometimes 4 or 6, in front of the clitellum, and behind 
the clitellum 4, 5, 6, once 7, and onee 8. 

The ‘‘ taste organ” on the floor of the pharynx is similar to 
that described for the next species, except that its base of attach- 
ment appears to be a little narrower. 


* ‘EXedroo¢os, bred in the marsh. 


LILAO DR. J. SEEPHENSON ON THE OLIGOCH ETA OF THE 


The septal glands are bulky, the last, in segm. vi., causing 
septum 6/7 to bulge back very considerably. 

Dorsal and ventral cesophageal nephridia are present, but less 
conspicuous than in the next species. The dorsal nephridium is 
double, consisting of two masses, one on each side of the middle 
line, attached behind to the dorsal side of the dorgal vessel near 
the angle which the vessel makes with the septum (6/7) and 
laterally to this on the anterior face of the septal gland; the 
masses have perhaps a small connection with each other across 
the middle line. ach is very irregular in shape, and the attach- 
ment to the dorsal vessel and septal gland is relatively narrow ; 
portions of the mass consist of long strings of cells extending 
forwards and upwards in the segment. 

The ventral nephridium is contained wholly in the esophageal 
wall, and is rather indistinct—indeed, in one of my two series of 
longitudinal sections it is scarcely to be made out; it is not 
obviously paired, and extends behind into segm. vii. 

A pair of postpharyngeal bulbs are present—small solid groups 
of cells behind the pharyngeal mass, continuous with a dorsal 
upfolding of the cesophageal epithelium. ‘These structures may be 
compared with those described in the account of the next species ; 
but here they are very much less definite, the knobs of cells less 
distinetly limited, the lumen of the upward fold of cesophageal 
epithelium much less marked, indeed hardly present, and the 
whole smaller. 

Chloragogen cells begin in segm. vii. ; in this segment, however, 
they are only scanty. 

The cesophagus expands to form the intestine in segm. vill. In 
this segment there is a slight invagination backwards of the 
cesophagus into the broader intestine behind; there are no 
diverticula. In each of the segments behind vii. the canal is 
swollen out into an ovoid shape, and this is the case in the 
genital segments also. There are no chylus-cells. 

The ccelomic corpuscles are large, granular, and elliptical, oval, 
or fusiform in shape, of an average length of 29; small nuclei, 


f=) 
with a central nucleolar dot, are visible in some, though not dis- 


tinctly discoverable in all. 

The dorsal vessel originates in the anterior part of segm. ix. as 
a sudden swelling just behind the septum; it contains numerous 
corpuscles. The blood was probably colourless. 

The nephridia have a relatively large anteseptal portion in 
which the central lumen undergoes some windings; the post- 
septal portion is ovoid, and not large; in one specimen the duct 
appears to be prolonged back from the hinder end of the body of 
the organ, and lies on the body-wall; in a second, the duct can be 
seen in sections to come off from the under (or under and outer) 
side of the postseptal portion considerably in front of its hinder 
end. 

The sexual organs are more completely developed in the second 


specimen (taken on the 10th of August), which will therefore be 
described. 


OXFORD UNIVERSITY SPITSBERGEN EXPEDITION. OAT: 


The testes are small, and not lobed. Sperm-morule and the 
early stages of spermatozoa are free in segm. x1., but not in large 
numbers. 

The male funnel is small, subspherical in shape, 108» in 
diameter, with spermatozoa clustering round its mouth. The first 
portion of the vas deferens is of a Sduanetcr of 15 », but this is. 
only for a very short distance on both sides of the septum, and 
the main part of the duct forms a very narrow tube 6m in 
diameter, packed tightly in a coil just behind the septum and 
entirely in front of the level of the male pore. The penial body 
is of the lumbricillid type, subspherical in shape, 120 in diameter, 
compact, with a definite outline and contained in a definite 
muscular capsule. 

The ovary is small; large ova are contained in segm. X11. 

The spermathecal duct is straight for the most part, of some 
length, 28 in diameter, directed downwards and forwards; 
towards its ental end it bends inwards; there are a few pear- 
shaped lobules of cells around its ectal end, but these may be 
peritoneal aggregates and not epithelial eland- -cells. The ampulla 
is only shghtly dilated, measuring 56 in diameter; the two 
ampulle are continuous ecross the middle line—there is not even 
a narrowing of the lumen to demarcate one from the other. 
There is no connection between the lumen of the esophagus and 
that of the ampulle ; the structures do not anywhere come into 
contact. The length of the conjoined ampulle is 160 « (computed 
—it extends through 20 longitudinal sections each of a thickness 
of 8). 

There are no copulatory (ventral) glands. 

The first specimen is altogether smaller than the second, and was 
collected a little earlier. The diameter of the male funnei is 75 p; 
though here, too, spermatozoa are clustering round the mouth. 
The penial body is also about 75 in diameter. The sperma- 
thecal ampulla is no wider than the duct, though the epithelium 
is much lower and the lumen therefore more capacious; the inner 
ends of the ampulle are not in contact either with the esophagus 
or with each other; there is an interval of about 16 » between 
the ampulle of the two sides. 


HENLEA (HENLEA) BRUCEI, sp. n. 


Spitsbergen; dry tundra, among Lryas, etc., on hill near coast, 
Freshwater Bay, Prince Charles Foreland; 10.vii.1921. A single 
specimen. 

Spitsberg gen; Bruce City region (no data concerning habitat) : 
10.vin.1921. A single specimen. 

Spitsbergen; among plants growing on shingle of raised, beach, 
Bruce City; 11.vi.1921. Two specimens. 


Length 6-8 mm.; diameter 31-46 mm. Segments 40-43. 
Colour brown (probably in part at least due to. preservative). 
Prostomium short and rounded. Head-pore on dorsal surface of 
prostomium. No dorsal pores. 


Lu? DR. J. STEPHENSON ON THE OLIGOCH ZITA OF THE 


Sete enchytrweine in form; there may, however, be a hint of a 
distal curve. In some bundles the inner sete are smaller and 
slenderer than those at the ends of the bundle (Wridericia 
arrangement). In front of the clitellum the ventral bundles 
contain 4 or 5 sete, occasionally 3; the lateral bundles have 
3 or 4. Behind the cliteilum the ventral bundles have 4 or 5 
oecasionally 3 or 6 sete; the lateral 4, 5, or 6, occasionally 3, or 
in the last segments even two. 

The clitellum roughly covers segms. xii. and xill.; but in a 
specimen with fully developed sexual organs one-third of xi. was 
included, and a small portion of the hinder end of xiii. (about a 
quarter) was free. 

A taste-organ is present on the floor of the pharynx, in front 
of the level of the dorsal pharyngeal mass and of furrow 2/3. It 
is best seen in longitudinal sections, according to which it isa 
solid transverse ridge, bent somewhat forwards, with a narrow 
free edge. Its base is slightly narrowed (antero-posteriorly, as 
seen in longitudinal sections), and its total height is about five 
times the antero-posterior thickness of its base of attachment. It 
is composed of elongated cells, and being solid is distinguishable 
from the other ridges of the ventral pharyngeal wall, nich are 
merely folds of the epithelium. In one specimen the ‘pharynx i is 
being everted; it is the ventral wall alone that comes forward 
here, and the taste-organ is at the tip of the everted part, so 
that it comes just onto a level with the mouth-opening, or 
in one or two sections rather further, projecting a little from 
the mouth. Some deeply staining matter is apparently being 
taken in. 

The septal glands are bulky; the last is in relation with 
septum 6/7, covering both sides of it and thus occupying part of 
segm. Vll. 

There are dorsal and ventral esophageal nephridia. The dorsal 
organ ends behind in segm. vi., and gives a branch dorsalwards 
just in front of its ending: in front it gets into segm. v. in one 
series of sections, and ends some Aisin above the cesophagus 
and not in contact with it. In another specimen, cut into longi- 
tudinal sections, it is confined to vi., and appears aS a pr ojecting 
mass dorsally on the cesophagus, spongy in texture, hardly 
staining, irregular in shape, and in intimate connection with the 
dorsal vessel ; indeed, a proper wall of the dorsal vessel is wanting, 
or cannot be distinguished, and the blood appears to be contained 
within a sinus in the nephridial tissue. 

The ventral nephridium ends behind by branching into two at 
a level just in front of that of the end of the dorsal organ; each 
branch extends laterally for some distance, and then terminates 
by turning somewhat forwards. These branches form the chief 
part of the organ; the median portion, on the ventral cesophageal 
wall, is soon lost, am does not seem to get forwards into segm. v. 
This nephridiam also is in close relation with blood-spaces—in 
this case those of the alimentary wall (text-fig. 1). 


—— 


OXFORD UNIVERSITY SPITSBERGEN EXPEDITION. DE 


In another specimen both organs appear to be confined to 
seem. V1. 

Besides these organs, which have, from their texture, more or 
less the appearance of nephridia, there is behind the pharynx a 
narrow diverticulum of the alimentary tube, wedge-shaped or 
conical as seen in longitudinal sections, pointed above, directed 
dorsalwards and backwards, and at its tip connected with a pair 
of solid knobs, the postpharyngeal bulbs. Each knob is a cellular 
mass, ovoid or somewhat club-shaped, with the long diameter 
continuing the direction of the diverticulum. ‘he length of the 
diverticulum is 120 u, that of the bulbs 36 », and the thickness of 
the latter 20 p. 


Text-figure 1. 


|- = ---0es. 


Henlea (Henlea) brueci; transverse section through cesophagus and hinder end of 
ventral wsophageal nephridium. BU., blood in sinus-like cavities of nephridium ; 
chl., chloragogen cells; es., esophagus; v.n.c., ventral nerve cord; w.neph., 
ventral cesophageal nephridium. 


The cesophagus is narrow as far as seem. viil., where it swells 
out into the intestine. The canal is thenceforwards swollen seg- 
mentally in an ovoid manner, and is not constricted in the genital 
segments. There are no chylus-cells. 

Chloragogen cells begin, but only scantily, in the posterior part 
of segm. V1. 

In segm. vill. is a pair of esophageal diverticula. Each is sac- 
like, with folded walls; each originates from the alimentary tube 
at the hinder end of the segment, and extends forwards to its 
anterior hmit. There appears to be some variaticn in the exact 
manner of origin of the sacs; in two series of sections, both longi- 
tudinal, the sacs have independent openings into the cesophagus, 


1124 DR. J. STEPHENSON ON THE OLIGOCHATA OF THE 


one on each side; in a third, a transverse series, the two sacs join 
dorsally above the alimentary tube before they enter it. 

The dorsal vessel originates in segm. vill. between the two 
cegophageal sacs. The blood was colourless (no red staining with 
eosin—absence of hemoglobin). 

The celomic corpuscles are few in number, large, disc-like, 
oval, about 80 in long diameter; the body is granular, and a 
nucleus is present, 34 in diameter, with a more deeply staining 
dot-like nucleolus. In one specimen the corpuscles were distinctly 
smaller than in the others—only 20 » long. 

The preseptal portion of the nephridia is often pyramidal in 
shape, narrow at the free end and broader where it joins the 
septum; in any case it is of some length (40-60). The 
postseptal portion is broad and ovoid or pear-shaped, with the 
wider portion behind. The duct is narrow, cylindrical, and long 
—as long as or longer than the postseptal portion; it is curved, 
arched, or even bent into a loop with the convexity backwards. 

The testes are small, compact, either slightly lobed or not, but 
not divided (that is, not split up intoa number of elongated pear- 
shaped divisions attached iv a cluster at their narrow ends, as is 
the case in the genus Lumbricillus). A few sperm-morule he 
loose in segm. X?. 

The funnels are small, with a distinct rim of nucleated cells, 
and resemble what is known as a ‘ thistle-funnel.” Hach is 
rather longer than broad; in different specimens they measured 
92 by 56 p, 100 by 72 p, and 120 by 100 p. 

The first portion of the vas deferens, which lies in front of the 
septum, is 16 « in diameter; it very soon pierces the septum, and 
after maintaining its former size for a short distance, contracts to 
a very narrow tube, 5-8 » in thickness, which hes in a close coil 
ventrally in the anterior part of segm. xii. Just behind the septum 
and behind the position of the funnel. 

The penial body i is of the lumbricillid type, spherical, compact, 
of definite form, enclosed in a muscular capsule, about 120 in 
diameter. In one specimen it appeared to be rather shorter in 
vertical diameter, 120 p high by 160 p long. 

The ovaries are small, and consist of club-shaped lobes. Ova he 
in segm. Xi. 

The spermathecal duct is cylindrical, about 30, in diameter, 
with thick walls and small lumen; there are no gland-cells either 
on its course or round its ectal end. The ampulla is cylindrical 
or ovoid, thin-walled, 40 » in cross-section. ‘There is a direct 
connection between the ampulle of the two sides across the 
middle line dorsal to the csophagus—a fairly wide passage 24 pu 
in diameter, with a lumen of 16 y; ane connection boreen the 
two ampulle is contiguous to the esophageal wall, but there is no 
communication with the cavity of the cesophagus, a double 
layer of cells, the alimentary epithelium and the spermathecal 
epithelium, alw: ays intervening. 

in anothei specimen, although the ampulle were more swollen 


7 


OXFORD UNIVERSITY SPITSBERGEN EXPEDITION. 5} 


than in that from which the above description was taken and 
contained spermatozoa, 1 could not make out any connection 
between the two chambers: here again there was no com- 
munication between the cavity of the spermathecal apparatus 
and that of the esophagus. 

There are no copulatory g glands. 

The present species 1s not very unlike the.;common H. nasuta, 
which has been recorded from N. Russia and N. Siberia. It 
differs, however (I take as the standard of comparison Welch’s 
description, 22), in being smaller, in having a blunt prostomium, 
in the shortness of the ventral cesophageal nephridium, in the 
male funnel being here more nearly spherical, in having no 
glands at the ectal end of the spermathecal duct, in hovine a 
broader communication between the two ampulle, and in having 
no communication between spermathece and cesophageal lumen. 

It also approaches H. urbanensis Welch, found in Illinois (19). 
The present species is smaller than that one, has fewer sete in 
the ventral bundles, the ventral esophageal nephridium is smaller, 
the dorsai vessel originates in vili., not in ix., the male funnel 
is much shorter, there are no gland-cells round the ectal end 
of the spermathecal apparatus, and no communication with the 
alimentary canal. 

In many species of Henlea the inner ends of the two sperma- 
thece are bent downwards, meet, and join to form a common 
passage which enters the dorsal surface of the cesophagus. In 
both the species here described the condition is rather different ; 
instead of meeting at an angle, the spermathece of the two sides 
are continued directly into each other across the middle line, and 
there is no communication with the cesophagus. This absence of 
communication is just possibly due to the specimens not being 
completely mature—the cesophageal connection being probably 
the last portion of the apparatus to form,—though from the 
condition of the other organs this hardly seems to be the case. 


HENLEA (HENLEANELLA) sp. 


Spitsbergen ; among mosses on the bank of a fresh-water pond, 
more or less under water, Bruce City; from August Ist to 8th. 
Five specimens, none fully sexual. 


Unfortunately the specimens are rather too immature to 
describe. They belong to that section of the genus which has no 
cesophageal sacs. 

The dorsal and ventral cesophageal nephridia attracted my 
attention. The dorsal is seen in segm., vi. with a central lumen 
and lateral branches of some size; on this portion is continued 
baek a solid cellular cord, regular in form, cylindrical and well 
defined in the middle line on the esophagus as far as segm. x. ; 
the cord, although it has no lumen, seems to take the place of 
the dorsal vessel, which is not separately visible. The ventral 


Proc. Zoou. Soc.—1922, No. LX XV. 15 


1126 DR. J. STEPHENSON ON THE OLIGOCHATA OF THE 


cesophageal nephridium is confined to segm. vi.; it consists 
mainly of two masses, one on each side of the middle line; each 
mass has folded walls and a considerable lumen. 


HENLEA (HEPATOGASTER) Sp. 


Spitsbergen; among mosses on the bank of a fresh-water pond, 
more or less under water, Bruce City; end of July and beginning 
of August 1921. A single specimen. 


This specimen was taken near the last, but in a different moss~ 
zone. The cesophageal diverticula take the form of a number of 
communicating tubules, as in the worms called Hepatogaster by 
Cejka (8) and in Henlea tubulifera and H. moderaia as described 
by Welch (18, 19). JI could not discover whether there were or 
were not dorsal and ventral cesophageal nephridia, as my sections 
of the worm were not very successful. The dorsal vessel begins 
in segm. 1x. 


Genus LUMBRICILLUS Orst. 


LUMBRICILLUS EGIALITES, sp. n.* 


Spitsbergen; in seaweed at high-tide mark on shore, at head of 
Klaas Billen Bay in Icefjord, by Bruce City; last week in August 
1921. Numerous specimens. 


Length up to 17 mm.; maximum diameter °65 mm. Colour 
brownish (fixed in Bouin). Segms. 52-53. Prostomium rounded, 
hemispherical. Head-pore not made out with certainty, probably 
present. 

A cuticle is not usually separately discernible; it is sometimes 
seen in the anterior segments, when it is not more than 1 mu in 
thickness. 

Fairly numerous deeply staining (with hematoxylin) mucous 
cells are seen in transverse rows in the surface epithelium—not, 
however, camel eminences on the surface. 

Clitellum 3X1.—xill, (=23), not very well marked. 

Setze enchytreine (distal portion straight), blunt. In the 
Hee bundles in front of the clitellum 4 or 5 setze; behind the 
clitellum 3, or at the hinder end 2. In the lateral bundles in 
front of the clitellum 3; behind, 3 in most segments, 2 at the 
hinder end. 

The septal glands are in segms. iv., v., and vi.; septum 6/7 is 
bulged backwards by the last gland. 

There are no salivary glands. There are, however, a pair of 
postpharyngeal bulbs—club-shaped structures in connection with 
the dorsal wall of the alimentary tube at the junction of pharynx 
and esophagus. Their total length is 240; they are constituted 
by a solid mass of cells in the form of a knob, and a stalk which 


* °AtytaXirys, of the shore. 


OXFORD UNIVERSITY SPITSBERGEN BXPEDITION, WU 7 


is attached to a small diverticulum of the alimentary tube; this 
might be expressed by saying that the proximal portion of the 
stalk is hollew, its cavity communicating with the alimentary 
lumen. 

In the esophagus are large numbers of parasitic Ciliata. The 
cesophageal epithelium is being shed en masse into the lumen of 
the canal; in some places also the cilia are being detached and 
shed. 

The intestine dilates gradually after passing through the 
genital segments, about segm. X11. 

Chloragogen cells begin in segm. Vi. 

The coelomic corpuscles have a length of 16-24 « ; in shape they 
are elongated, fusiform, pear-shaped, or oval. They are not very 
definite structures, and often seem to be disintegrating and of 
irregular outline. Some, broader than most, are nucleated. 

The dorsal vessel begins in segm. xiv. The blood from its 
staining reactions (it stains a pinkish-brown with eosin) probably 
contained hemoglobin. 

Text-figure 2. 


~ 


hf 
I 
Cerebral ganglion of Lumbricillus egialites. 


The anteseptal portion of the nephridia consists of the funnel 
only. The postseptal portion is spindle-shaped. The stout duct 
is the hinder end of the organ bent downwards and inwards, 
sometimes forwards as well; sections of the duct are rather 
characteristic—always oval in outline and showing radial stria- 
tions in the wall. 

The cerebral ganglion (text-fig. 2) has a slightly concave 
anterior border; the posterior border is shghtly indented in the 
middle line and rounded on each side of the median indentation. 
The ganglion is about as long as broad, and is broader in front 
than behind. 

The testes have the usual constitution in the genus. Hach 
consists of a number of elongated lobes (“‘ divided” testes, using 
Welch’s term), which extend into segms. x. and xi.; each lobe is 
contained in a membranous sac. which in some eases may be fairly 


75% 


1128 DR. J. STEPHENSON ON THE OLIGOCHATA OF THE 


substantial, with a number of much elongated nuclei in its wall. 
The substance of the lobes consists proximally of a mass of cells 
only; further from their attachment they dissolve into sperm- 
morul, still contained within the saes. There is a large vacancy 
in septum 10/11; as already said, the lobes of the testes extend 
into both segments. 

‘he funnel is short and stout——about twice, or at any rate not 
more than three times, as long as wide. In a specimen in which 
the funnels were isolated by dilaceration they were not more 
than 14 times as long as wide. The vas deferens is long and 
much coiled, but in the dilacerated specimen not so tightly as 
in the next species; it is confined to segm. xii.; in diameter it 
measures 10-12 jr. 

The penial body is compact, subspherical in shape, in diameter 
130-160 p, of the usual lumbricilline type, with a strong muscular 
capsule, It causes a slight projection on the surface. 


Text-figure 3. 


Spermatheca of Lumbricillus egialites isolated by dilaceration. Amp., ampulla; 
coag., coagulum in ampulla; d., duct; gl., mass of ¢land-cells. 


The ovaries are in segm. xil.; loose ova are present in xil. and 
also in xiii. There is no ovisac. The female funnels are merely 
backward bulgings of the septum. The oviducts are not as yet 
patent, 

The spermathecal ampulla is smoothly ovoid, 90-100; in 
diameter, and communicates with the esophagus by a rather 
narrow neck. The duct is straight and as long as the ampulla. 
A large mass of gland-cells surr ounds the duct, and takes up all 
the space Iearneen the parietes and the ampulla—even surround- 
ing the basal half of the ampulla; the ae margin of this mass 
of “eland- cells is slightly lobed (text-fig. 3 

Copulatory glands (‘ ventral slands ’ ’) are present in segments 


OXFORD UNIVERSITY SPIDSBERGEN EXPEDITION. 1129 


xiv. and xv. They are small, and the wings do not meet above 
the cord, all the dorsal surface of the cord being free. The 
lateral extent of the wings is not great; each measures about 
twice the transverse diameter of the cord, and does not reach 
outwards more than halfway towards the ventral setal bundles. 
They are not composed of several layers (as in LZ. nervosus, 
according to Ude). 

Besides the parasites in the cesophagus, others were found in 
the testicular region. Here there are a number of sacs, each 
containing what § appear to be stages in the formation of spores. 

It is extremely difficult to ho which of the worms described 
by the older writers, and designated by names that are still in 
use, correspond to forms met with at the present day. The 
older descriptions are so very scanty, according to present-day 
requirements, that they frequently fit several of the species 
now recognized. In course of time, other descriptions have been 
published under the older name, supposedly referring to the same 
worm; particulars have been taken from these descriptions and 
incorporated in the diagnoses. The result is a composite picture, 
which may or may not represent the worm which the original 
deseriber had under his eyes. Another worm may come up, 
which may ft the original description equally well—and may 
therefore be identical with the original species; but it will have 
to be deseribed as new, since it is not characterized by the accre- 
tions which have gathered round the original diagnosis. 

Thus the original description of Lumbricillus pagenstecherr 
Ratzel) is very scanty, but it has been added to by Vejdovsky 
and Ude. The present worm seems to differ from the original 
description (12) in having the sete fewer in number (2-5 as 
against 6-10, 7-8 being the commonest numbers in the original), 
and straight (those of the original, while described as straight, 
had, nevertheless, a slight curve at their sharp end—‘“ mit leichter 
Biegung an der scharfen Spitze”), with blunt instead of sharp 
points. From the diagnosis in the ‘ Tierreich’ (11), which embodies 
what I have called later accretions, it differs in having small 
instead of large copulatory glands, which leave the whole of the 
dorsal surface of the cord uncovered; and in having the whole of 
the spermathecal duct closely covered with gland-cells, instead 
of loosely; the shape of the cerebral ganglion also differs con- 
siderably. 

From L. henkingi, described by Ude from Bear Island, the 
present form differs in not having S-shaped sete, in the origin of 
the dorsal vessel in xiv. (instead ‘of in xiii. ), in havi ne copulatory 
glands in xiv. and xv. (instead of in xiii. and xiv.), and in the fact 
that these glands are here very much smaller (in Z. henkingi the 
wings of the glands are four times as extensive as the ereatest 
diameter of the cor d, and they reach outwards somewhat beyond 
the setal bundles). 

The present species is not very unlike JZ. franciscanus Hisen, 
from California (varieties of the species also from Pribilov Islands 


eles DR. J. STEPHENSON ON THE OLIGOCH ATA OF THE 


and Alaska). It differs apparently in the shape of the sete, in 
the place of origin of the dorsal vessel (in LZ. franciscanus it arises, 
according to the short diagnosis, in segm. xiv. or xv., but in the 
detailed description it “has already arisen in xvi. How much 
further posteriorly it extends I do not know, as I did not section 
aN URED in the rather greater extent of the clitellum, and in 
the more restricted distribution of the copulatory g elands (clitellum 
over segms. Xl1.—xil1., glands in xiv.—xvi. in LZ. fr anciscanus). 


ET 


UMBRICILLUS NECROPHAGUS, sp. n.* 


ae 


Spitsbergen ; in dead seal, at high-tide mark on shore, by Bruce 
City at head 6 Klaas Billen Bay in Icefjord ; 25.vii.1921. A 
nuinber of specimens. 


Length up to 15 mm.; maximum diameter ‘4-55 mm., but 
many are thinner. Colour paie yellowish. Segments 51-53. 
Prostomium blunt, rounded. No mucous cells in epidermis. 

The surface is covered with a moderately thick cuticle, con- 
siderably thicker and more noticeable than that of the last species. 

Clitellun 3xi.-xih. (=23); in sections the extent appears to be 
rather different—it includes a little of xi. and gets slightly on to 
xiv. also. 

The sete are enchytreine in form (1 think I have occasionally 
detected a hint of a distal curvature) and blunt, rounded, or even 
squarish at their tips. The ventral bundles contain in front of 
the clitellum mostly 4 (occasionally 5 or 3) sete and behind the 
celitellam 3. The lateral bundles have usually 3 (occasionally 4 or 
2) in front of, and 3 or 2 behind, the clitellum. 

~The septal glands are bulky, the last, im segm. vi., causing a 
backward bulging of the septum. 

The postphary yngeal bulbs arise from a transversely placed 
eroove-like evagination of the alimentary tube behind the pharyn- 
geal mass. Each is a slender elongated eord of cells, rather 
swollen at its distal end, its general direction nearly vertical, and 
solid except in its proximal third or less, where there is a narrow 
lumen. 

Chloragogen cells begin in segm. v. 

The cesophageal epithelium i is being thrown off into the cavity 
of the tube; the cells lie loose in the lumen, and there is a copious 
granular substance between them and the remainder of the 
esophageal wall. Similarly, the intestinal epithelium is dis- 
integrating or has disappeared over large areas; the intestinal 
tube is full of a faintly staining granular substance, or of what is 
apparently a disintegrating cell-mass; sometimes the epithelium 
is separating in sheets, and the lumen is half empty. The 
intestinal tube is much swollen out, and the celom is almost 
obliterated in parts, the nephridia being pressed against the 


xs Ld . 
“ Nexpogayos, devouring corpses. 


b) 


OXFORD UNIVERSITY SPITSBERGEN EXPEDITION. WRI 


body-wall. Extreme degeneration may exist throughout the 
whole of the alimentary tract behind the septal glands. 

This condition might be due to the presence of parasites—there 
are a number of such, elongated, cylindrical, with a nucleus of 
corresponding shape, in the cesophagus (probably Ciliates, though 
no cilia could be made out on them), as well as a number of small 
cysts in the tissues. Still, the usual Ciliate parasites of the 
cesophagus of Hnchytreids do not asa rule cause such degenera- 
tion. It might perhaps be due to the parasites of the tissues, 
through their metabolic products, which may have a specific 
action on the alimentary epithelium. Or one might attribute it 
to the food (assuming that the worms were taking in material 
from the body of the dead seal in which they were found), but for 
the fact that the last species also showed degeneration of the 
alimentary epithelium, though not to such an extreme degree as 
these worms. 

Text-figure 4, 


Cerebral ganglion of Lumbricillus necrophagus. 


A lightly staining, almost perfectly homogeneous, very slightly 
granular coagulum, containing many nuclei, almost fills up the 
ecelom in the anterior part of the body. There are no definite 
lymph-corpuscles of the type of those found in the former species ; 
but the nuclei in the coagulum just described seem to have 
belonged to corpuscles which have degenerated. These changes 
seem to go along with those in the intestine, and are doubtless 
due to the same cause. 

The dorsal vessel begins in segm. xiv. The blood contained 
hemoglobin. 

The anteseptal part of the nephridia is quite small and 
shortly cylindrical. The postseptal portion is elongated and is 
continued behind into the duct, which has the same structure 
as in the last species. The whole is flattened against the body- 
wall. 

The cerebral ganglion is rectangular in general outline (text- 
fig. 4), the sides being approximately parallel and the length 


1132 DR. J. STEPHENSON ON THE OLIGOCHAZTA OF THE 


about one-third greater than the breadth, he posterior border 
is very slightly indented, and the posterior angles are well 
rounded, 

- The testis consists of numerous lobes or divisions, each contained 
in a definite sac; in each sac the distal portion of the contents 
consists of morule and developing spermatozoa. The lobes are 
contained in segms. x. and xi.; they originate from the parietes on 
both sides of the attachment of the septum. Numbers of sperma- 
tozoa are free in segm. xii. at and behind the level of the penial 
bodies and as far back as the hinder end of the segment. 
Numerous cysts containing developmental stages of parasites 
occur amongst the testis-lobes. 

In two specimens from which the male deferent apparatus was 
isolated by the method of dilaceration, the funnels appeared to be 
only a little longer than broad—not more than half as long again 
as broad. In one specimen, in which they were well seen, they 
were rather triangular in shape, the posterior end (where the vas 


Text-figure 5. 


Spermathece and intervening part of csophagus of Lubricillus necrophagus, 


isolated by dilaceration. Amp.,ampulla; coag., coagulum in ampulla; d., duct; 
gl. mass of gland-cells; @s., cesophagus. 


deferens is given off) being narrow. In sections also the funnels 
appeared short and stumpy, the proportions varying, but 
apparently most often about half as long again as broad. 

The vas deferens is narrow, 8 », and is confined to segm. xii.; it 
is tightly coiled, and is seen in sections as a number of windings 
squeezed close against the body-wall. The penial body is of the 
lumbricillid type, ovoid, somewhat flattened vertically, -19 mm. 
long and 14 mm. high. 

The large ovary, in segm. xii., is also branched into lobes. 

The spermathecal ampulla is shortly and stoutly pear-shaped, 
drawn out above into its esophageal connection; there appears 
to be a communication between the cavities of the spermatheca 
and of the esophagus, The duct is well marked off, longer than 
the ampulla, and surrounded by a large lobed mass of gland-cells 
which fills up the space between the parietes and the base of the 
ampulla (text-fig. 5), 

Copulatory glands occur in segms. xiv. and xv. They are small, 


OXFORD UNIVERSITY SPITSBERGEN EXPEDITION. TL S3 


not covering the dorsal surface of the cord; their shape is slightly 
different from those of the last species; the lateral expansions are 
level with the dorsal surface of the cord, so that there is a straight 
line all along; in the former species the wings rise above the 
level of the cord. 

I was at first extremely inclined to unite these worms with the 
last species; and I am still not quite certain that they are speci- 
fically distinct. The chief difference is that there are no mucous 
cells in the present specimens; they were stained in exactly the 
same way as the former, yet the mucous cells are extremely 
obvious at the first glance in the former preparations, while they 
are entirely absent in these. The thicker cuticle also seems to 
be a distinction, as well as the shape of the cerebral ganglion and, 
possibly, though doubtfully, the different shape of the copulatory 
glands. The appearance of the two worms is also very different ; 
L. egialites is dark brown (perhaps in part due to the fixative— 
Bouin), LZ. necrophagus pale and semi-transparent (fixative not 
stated); ZL. wyialites appears much stouter than L. necrophagus, 
though actual measurements hardly bear this out. But the two 
are closely related ; whether the differences can be put down to 
the different manner of life in the two cases seems doubtful, since 
one can hardly suppose that the occurrence of these specimens 
in a dead seal was more than accidental. 


Genus Encuytreus Henle em. Mich. 
ENCHYTREUS ALBIDUS Henle. 


Bear Island; Walrus Bay (S.H. of island); by shaking plants 
of dry tandra; 15.vi.1921. Four specimens, some of them 
(? all) mature. 

Same locality; among plants on shaly slope; 22.vi.1921. 
Two specimens, both mature. 

Same locality; among plants on top of hill (dry tundra), 
22.vi.1921. Two specimens, both mature. 


HNCHYTREUS CRYMODES, sp. n. * 


Spitsbergen ; among mosses on the bank of a fresh-water pond, 
more or less under water, Bruce City; end of July and beginning 
of August 1921. Several specimens. 

Spitsbergen; moss by salt marsh, probably reached by ex- 
tremely high tides, near Bruce City; 8.vii.1921. Numerous 
specimens. 


Mr. Elton adds a note to the above data:—‘'These” (the 
second of the two batches) ‘‘ were observed to be living on live 
moss leaves, and seemed to occur among these, but not in the 


decaying parts of the moss. They survived being frozen solid, in 
an experiment which I did.” 


* 


Keupwdons, icy cold, frozen. 


1134 DR. J. STEPHENSON ON THE OLIGOCH ETA OF THE 


Length 8-9 mm.; diameter:4 mm. Colour brownish, Seg- 
ments 34-38. Prostomium bluntly triangular, the triangle being 
about equilateral. Head-pore between | prostomium and first 
seginent; no dorsal pores. 

‘Clitellum not well marked (or scarcely formed), x1i.-xiil. (= 2). 

Sete enchytreine in form, tip fairly sharp. In front of the 
clitellum both ventral and lateral bundles have three sete, 
occasionally four: behind the clitellum they have two only. 

The hindmost septal glands are in segm. vi. 

There are no salivary glands of the type of those of Z. aliidus, 
but, as in the species of Lumbricillus, there are a pair of post- 
pharyngeal bulbs. These are small, knob-like, ovoid cell-masses 
unmediately behind the pharynx, connected with the first part of 
the cesophageal tube by solid conical cell-aggregates. The knobs 
are separated from each other in the middle line by a muscular 
strand. In other preparations of these same worms the struc- 
tures are not so clearly seen—either there is actually some 
variation or the plane of section is not so favourable. 

Chioragogen cells begin scantily in segm. v. 

The cesophagus remains narrow till past the genital segments ; 
it begins to widen in xiii. and the intestine is en Ly established i in 
xiv. here are numerous sporozoan parasites in the epithelium 
of the cesophagus. There are no chylus-cells in the alimentary 
tract. 

The dorsal vessel originates in segm. xiii. The blood stains 
slightly with eosin ; probably there was a small amount of 
hemoglobin in it originally. 

Uclomic corpuscles are present as irregular discs, staining 
little, fusiform or eval in shape, 32 in average length (the 
limits being 30-40), nucleated; but they are relatively few 
in number. 

The nephridia (text-fig. 6) have a small narrow anteseptal 
portion; the postseptal portion is swollen out, shortly ovoid or 
subspherical. The duct is long, relatively thick, and leaves the 
outer side of the postseptal mass, passing backwards for some 
distance and then downwards—-even somewhat forwards in some 
sections. 

The testes are bulky in the present specimens; they are slightly 
lobed in their distal portions. Some of the sexual cells are 
becoming metamorphosed into sperm-morule and spermatozoa, 
and these are bound together and to one of the lobes of the testis 
by an apparently imeomplcre sac. These sacs appear to be 
rupturing, and so allowing their contents to escape. In one 
specimen there is a mass of “genital cells in segm. ix. 

The funnels are long and narrow, as much as 6-8 times as long 
as broad. They may push right through the septum into seem. x., 
in which the anterior half of the rel may thus be gomtecned - 
or the anterior end may be reversed and push back septum 11/12 
so as to forma sac, in which the anterior part of the funnel is 
contained. The mouth of the funnel has a distinct lip. 


OXFORD UNIVERSITY SPITSBERGEN EXPEDITION. U5) 


The vas deferens is narrow, 7-12 in diameter, and forms 
numerous close coils in segm. xii., to which it is confined. It 
pierces and penetrates the penial body. 

The penial body is of moderate size, 12 mm. in diameter, com- 
pact, of the lumbricilline type, spherical, with a definite muscular 
capsule. 

The ovary, in segm. xii., is bulky. Septum 12/13 is bulged back 
by ova contained in segm. xii. in such a way as to form an 
incipient ovisac. 

The spermathece have each a relatively small and thin-walled 
ampulla, subspherical in form, 80-120 in diameter. The duct 
is thick-walled with high epithelium, 70-80 in diameter, 
eylindrical, longer than the ampulla and nearly as thick. There 
are a number of pyriform groups of celis round the ectal end of 
the duct. The ampulla probably communicates with the cavity 
of the esophagus at full sexual maturity, though such a patent 
communication was not visible in my specimens; in one, however, 


Text-figure 6. 


Vv. S. 
Nephridia of Enchytreus crymodes. 


Sep., septum ; v.s., ventral setal bundle. 


the epithelium of the two organs (ampulla and cesophagus) was 
continuous. The two ampulle may be separated from one another 
by a part of the esophagus; but in another case they were con- 
tiguous—indeed, their walls were fused together. In both these 
eases the cavity of the ampulle contained spermatozoa. 

Copulatory glands are present, though small, in segms. xii. to 
xvi. and perhaps xvii. They do not cover the dorsal surface of 
the cord at all. There isa small rounded papilla over the position 
of each gland where the giand-cells come to the surface. 


ENCHYTRAUS sp. 

Spitsbergen; among moss in marshy (fresh-water) area near 
Bruce City; 8. vill. 1921. A number of specimens. 

I regret that this worm is rather too immature to justify my 
giving ita name. I append a few particulars, which may enable 
it to be identified when next it is met with. 


1136 DR. J. STEPHENSON ON THE OLIGOCHETA OF THE 


Length 3-3-5 mm.; diameter "17 mm. Segments 22, or 22 
with some incomplete segments behind. Colour dark, Sete 
enchytreine in form, two per bundle throughout the body. No 
salivary glands. Ccelomic corpuscles fairly numerous, oval, 
nucleated, 12-14 im length. The dorsal vessel originates in 
segm, xii, The nephridia have a large anteseptal portion, nearly 
as large as the postseptal; the duct undergoes several windings 
in the anteseptal portion; the organ is somewh at constricted at 
the septum; and the postseptal “portion is elongated, the duct 
being continued from the hinder end. 

The testes were in an early stage, of large size, taking up a 
very large part of segm. xi.; scarcely any sperm-morule had 
formed, the testes were lobed, but not “divided.” 'The funnels 
were very small, and twice as long as broad; but it is more than 
likely that these proportions would not be maintained thr oughout 
the further development of the organ. There was a small, ovoid, 
and well-defined penial body. 

The ovaries were lobed in the same manner as the testes. The 
spermathecal ampulla was short, subspherical, and the duct eylin- 
drical, entirely destitute of surrounding gland-cells. There were 
no copulatory glands. 

The numbers of the sete (2 per bundle throughout the body), 
the large anteseptal part of the nephridia, and the presence of a 
lumbricilline penial bulb constitute a group of characters which, 
along with the small size of the worms, may permit of recognition 
in the future; but I disapprove of the practice of giving names 
to incompletely described or immature material. 


Genus MESENCHYTREUS Hisen. 
MESENCHYTREUS sp. 


Spitsbergen ; among plants of dry tundra, Cape Boheman in 
Icefjord in W. Spitsbergen; 12.vii.1921. A single specimen. 

Spitsbergen; among mosses on the bank of a fresh-water pond, 
more or less under water, Bruce City; end of July and beginning 
of August 1921. Two specimens. 


Like the last, the present specimens are too immature to 
name. 

Length 7-10 mm.: maximum diameter -6-"8 mm. Seems. 46 
—54. Colour dark brown. The worms are solid-looking, like the 
young of earthworms. A large head-pore near the tip of the 
proboscis. Sets: lumbricine in shape, stout and relatively short, 
with sigmoid curve; in the ventral bundles 4, 5, or 6 throughout 
the body ; in the lateral 2 or 3 throughout, or sometimes behind 
the clitellum 4 or 5, and occasionally, in front of the clitellum, 
one only. 

Ceelomic corpuscles rather small, 12-20 u, granular, with deeply 
staining granules or network, and a small nucleus visible only 
with difficulty. No salivary glands; postpharyngeal bulbs 


OXFORD UNIVERSITY SPITSBERGEN EXPEDITION. HUB 


present. (CHsophagus dilates fairly suddenly in segm. vii. Very 
numerous parasitic Ciliates in cesophagus; parasites also in 
cesophageal wall; a cyst outside, in the eelom. Much of the 
alimentary epithelium is degenerating or has been shed. Dorsal 
vessel originates in xiv.; the blood stains with eosin. Nephridia 
of the type found in the genus; the duct goes backwards at first, 
then forwards, ending finally a very little way behind the inter- 
segmental! groove, indeed almost in it. 

Clitellum only slightly marked, extending over xii. and xiii. 
with the greater part of xi.also. Testes lobed, but not * divided.” 
Sperm-sac present, extending into xii. Funnel as yet merely a 
plate of cells on the septum; vas deferens only as a solid cord 
of cells, cylindrical and somewhat wavy, from funnel to male 
aperture. Penial bulb as a number of loosely arranged cells 
around the terminal portion of the vas deferens. Ovisac ex- 
tending back to xvii. Spermathecal ampulla not distinguishable; 
duct narrow, no gland-cells round its ending, cylindrical, of some 
length, coming to the surface in the tract of the lateral line cells. 

The distinction of species of MWesenchytreus depends largely on 
the characters of the vas deferens and spermathecal ampulla— 
just those portions of the sexual apparatus which are most 
undeveloped in the present specimens. 


REFERENCES TO LITERATURE. 


(1) Bepparp, F. E.—A Textbook of Zoogeography. Cam- 
bridge, 1895, 
(2) Brerscumr, K.—Uber ein neues Enchytriiidengenus. Zool. 
Anzeiger, xxix. 1906. 
(8) CesKa, B.—Die Oligochaeten der russischen in den Jahren 
1900-1903 unternommenen Nordpolar Expedition. I. 
Uber eine neue Gattung der Enchytraeiden (Hepato- 
gaster). Mem. Acad. Sci. St. Petersb. (8) xxix. no. 2, 
1910. 
(4) Ceska, B.—/b. II. Uber neve Bryodrilus und Henlea- 
Aactenia 10.) (8) xxi, os O- NOM2 
(5) Detpuy, J.—Etudes sur lorganisation et le développement 
des Lombriciens limicoles thalassophiles. Paris, 1921. 
(6) Eisex, G.—Enchytreids, in Harriman Alaska Expedition, 
vol. xu. New York, 1904. 
(7) Frrenp, H.—A Key to British Henleas. Zoologist, (4) xvii. 
1913. 
(8) Frrenp, H.—British Enchytreids. VI. New Species and 
Revised List. Journ. R. Mier. Soc. 1914. 
(9) Frienp, H.—Studies in Enchytreid Worms. Henlea 
fragilis Friend. Ann. Apphed Biol. 11. 1915. 
(10) Micnaritsen, W.—Synopsis der Enchytraeiden. Abh. Ver. 
Hamburg, xi. Heft 1., 1889. 
(11) Micwartsen, W.—Oligocheta, in Das Tierreich, vol. x. 
Berlin, 1900. 


1138 THE OLIGOCH ETA OF THE SPITSBERGEN EXPEDITION. 


(12) Ravzer, ¥.—Beitriige zur anatomischen und systematischen 
Kenntnis der Oligochaeten. Zeitschr, f. wiss. Zool. 
xvill. 1869. 

(13) SrepHenson, J.—On some littoral Oligocheta of the Clyde. 
Trans. R. Soc. Edin. xlvii. 1911. 

(14) Srapuenson, J.—Oligocheta, in Fauna of the Chilka Lake. 
Mem. Ind. Mus. v. 1915. 

(15) SrepHEeNnson, J.—On some Indian Oligocheta, mainly from 
Southern India and Ceylon. Mem. Ind. Mus. vi. 1915. 

(16) Uns, H.—Beitrige zur Kenntnis der Hnchytreiden und 
Lumbriciden. Zeitschr. f. wiss. Zool. lxi. 1896. 

(17) Ups, H.—Die arktischen Enchytraiden und Lumbriciden, 
sowie die geographische Verbreitung dieser Familien. 
in Fauna Arctica, vol. ii. Jena, 1902. 

(18) Wetcu, P.S.—A new Henlea (Enchytreeide) from Northern 
Michigan. Trans. Amer. Micr. Soc. xxxii. 1914. 

(19) Weice, P. S.—Studies on the Enchytreide of North 
America. Bull. Illinois State Lab. Nat. Hist. x. 1914. 

(20) Wetcs, P. 8.—Snowfield and Glacier Oligocheta from Mt. 
Rainier, Washington. Trans. Amer. Micr. Soc, xxxv. 
II). 

(21) Weicn, P. 8.—-Alaskan Glacier Worms (Oligocheta). 
Bionomical Leaflets, McGill Univ., Montreal, no. 2, 
Wile 

(22) Wetcu, P. 8.—Report on the Canadian Arctic Expedition 
1913-1918. Vol. ix. part A: Oligocheta—Enchy- 
treeidee, 1919. 

(23) Weucu, P.S.—The Genera of the Hnchytreide (Oligocheta), 
Trans. Amer. Micr. Soc. xxxix. 1920. 


THE ANATOMY OF SCAPHULA. 1139 


50. On the Anatomy of Scaphula, Benson, with a description 
of a new Species. By EKENDRANATH GHosH, M.Sc., 
M.D., F.Z.S., F.R.M.S., Professor of Biology, Medical 
College, Calcutta. 

[Received June 19, 1922: Read November 21, 1922. ] 


(Text-figures 1-7.) 


The Lamellibranch genus Scaphula, Benson is represented by 
three species from the rivers of India and Burma. The anatomy 
of all the known species is unknown to science, with the exception 
of the gills of an undetermined species from Siam (Ridewood, 
Philosophical Transactions, Ser. B, vol. 195). The present paper 
aims at describing the, characteristic soft parts of the genus, 
based upon the animals of S. celox, Benson, and of a new species 
from Siam. ‘The specimens were received from the Zoological 
Survey of India, through the kindness of Dr. N. Annandale. 


Text-figure 1. 


Zp 


Animal of S. celox, side view (left). 10. 


The body is elongated, being two to three times as long as 
it is deep, and deeper posteriorly than anteriorly. The median 
dorsal margin is a third to a quarter less than the body- 
length, The anterior adductor muscle is much smaller than the 

osterior. Both the anterior and posterior margins slope back- 
wards. The anterior adductor muscle is narrow and elongated 
at its insertion, and is placed rather close to the anterior margin. 
The length of the area of insertion is parallel to the latter. The 
posterior adductor muscle is large at its insertion and may be 
oval or triangular in shape. It is placed a little distance from 
the posterior margin. The foot is elongated and compressed 
laterally ; it presents a shallow fissure along the mid-ventral line. 
The foot is perhaps capable of slight expansion round the fissure. 
A minute byssus cavity 1s present in the ventral fissure towards 
the posterior end of the foot. The byssus is not well developed, 
being in the form of fine silky threadsin the Siamese species. An 
obtusely-pointed process is present at the postero-ventral aspect 


1140 DR. E. GHOSH ON THE 


of the foot. The anterior retractor pedis muscle is narrow and 
elongated, and consists of two heads of insertion into the valve. 
The posterior retractor pedis muscle is narrow, and elongated at 
its insertion into the valves just in front of the posterior adductor 
muscle. It extends diagonally from the posterior end of the 
foot. 

The visceral mass 1s greatly widened out laterally into a some- 
what conical hump with an oblique ridge on the surface. The 
gills are elongated, tapering posteriorly, and are about half the 
length of the body and thrice as long as broad. The gill-axis is 
slender, rod-like, and is placed more horizontally than diagonally 
and ends in a small free posterior portion. The gill-filaments are 
placed obliquely to the gill-axis, and the refiected filaments are 
free dorsally. The labial palps are elongated, semilunar in shape, 
and are placed at an obtuse angle with the gill-axis. They are 


one-third to one-fourth the length of the body, and twice as long 


5 C . 
as broad. The cesophagus is curved, with the concavity directed 


backwards. The stomach is greatly elongated. The cecum of 


Text-figure 2. 


Animal of 8S. celox, longitudinal section. 
») t=) 


the crystalline style is small, narrow, and is fused with the 
beginning of the intestine, the latter being placed on the right 
side of the caecum. The intestine consists of a short anterior and 
a long posterior limb, and is placed towards the right side of 
the body. The rectum is placed dorsally to the heart, and ends 
in a small free portion projecting downwards and backwards 
from the posterior surface of the posterior adductor muscle. 
The pericardium is divided into two lateral portions. The two 
ventricles are widely separated from each other by a narrow 
tube, which gives rise to anterior and posterior aorte. The 
aur icles are comparatively large; they do not communicate with 
each other. The digestive glands are extensive. The genital 
organs surround the lower portion of the esophagus and occupy 
the ventral region of the visceral mass. Osphraclia and Thiele’s 
abdominal sense-organs are both absent. 

Structure of the Gills of S. celox.—The gill-filaments are 
flattened antero-posteriorly as usual. There are no interlamellar 
extensions (described in Arca pexata by Kellogg (Bull. U.S. Fish 


ANATOMY OF SCAPHULA. 1141 


Comm. 10,1890) and in Arca americana by Ridewood, l. c. p. 199), 
but interlamellar septa (present in both the above species) are 
present, extending to about one-fourth or one-fifth the height of 
the filament. The ciliary discs seem to be present only at the 
free endsand at the junctions of the direct and reflected filaments. 
They are much more numerous in all the other known genera and 
species of the Arcide, being arranged in many horizontal rows. 
The arrangement of the cilia could not be followed distinctly. 
The frontal cilia do not seem to extend much on the anterior and 
posterior faces of the filaments. The lateral ciliary rows are 
placed a little distance behind the frontal row. The chitinous 
lining is uniformly thin. 

The gills of the species of Scaphula from Siam could not be 
studied. Ridewood notes in species from Siam (the present 
species?) one horizontal row of ciliary dise to the descending 
filament and one to three such rows in the ascending one. 


Text-figure 3. 


Ventral view of the foot and visceral mass of S. celox. 


Remarks.—The genus Scaphula Benson belongs to the family 
Arcide. It was recognized as such by Gray (Ann. &. Mag. Nat. 
Hist. xix.), and by Adams and Adams (Gen. Recent Moll.). 
Fischer (Man. Conch.) makes it a subgenus of Arca,and places it 
in the middle of his list. Dall (Trans. Wagner Free Institute of 
Science, 111, pt. 4) makes it a group in the subgenus Barbatia. 
Pelseneer (Lankester’s Treatise on Zoology, v.) raises it to a 
genus, and places it near the genus Arca. Lamy (Journ. 
Conchyliol. lv.), in his revision of the genus Arca, considers it a 
subgenus, and places it near the subgenera Arca, Barbatia, Acar, 
Fossularca, and Parallelepipedum. 

The body of the animal is characterized by an antero-posterior 
elongation, perhaps correlated with the inequality in the size of 
the adductor muscles. ‘The elongation of the body has also affected 
the visceral mass and the foot toa relatively less extent. But 


Proc. Zoou. Soc.—1922, No. LX XVI. 76 


1142 DR. BE. GHOSH ON THE 


the shape, size, and the position of the anterior adductor muscle 
may have something to do with the atrophy * of the extreme 
anterior region of the body. The poor development of the byssus 
apparatus and the absence of sense-organs are probably examples 
of degeneration due to its fluviatile life. 

The animal of Scaphula agrees more with the animals of 
Byssarca Swainson, Barbatia Gray, and Acar Gray than those 
of the other genera of the Arcide in its great antero-posterior 
elongation and in the inequality in size of the adductor muscles. 
It further resembles the first two, especially in the widening of 
the body from side to side, and in the great disparity of the 
adductor muscles in size. But the foot and the visceral mass 
are strongly affected in the present genus by the elongation of 
the middle region of the body, whereas in Byssarca and Barbatia 
these organs are comparatively less elongated, the elongation 


Text-figure 4. 


/3 


Heart (from the dorsal aspect) of S. celox. 


being mainly exhibited in the posterior retractor pedis muscles. 
In Acar the body seems to be equally elongated in the middle 
region (the above-mentioned parts being affected) and in the 
posterior region beyond the posterior adductor, the mantle only 
being affected. The elongation of the posterior region of the 
body is thus greatest in Acar, least in Dyssarca, and attains an 
intermediate condition in Scaphula. With regard to the heart 
of Scaphula, we find an intermediate condition of the ventricles 
between Byssarca (with a wide separation of the ventricles) 
and Barbatia (in which they are only slightly separated), but 
marked difference from Acar, which has a single, but distinctly, 
bilobed ventricle. The position of the heart in relation to the 
rectum might have been derived from a condition seen in the 
above genera. Lastly, the projecting portion of the rectum seems 


* This is not referred to elsewhere.—ED. 


ANATOMY OF SCAPHULA. 11435 


to bring the anus directly in the line of the outward current of 
water and the exhalant aperture, and this is perhaps correlated 
with the depth of the posterior region below and behind the 
posterior adductor. In Acar the great depth of the mantle is 
attended by a great elongation of the projecting tube of the 


Text-figure 5. 


A gill-filament of S. celox (outer and inner demibranch). 


rectum. In conclusion it might be noted that the present genus 
might have originated amongst and be related to Byssarca and 
Barbatia, although modified and perhaps degenerate owing to its 
fluviatile life. 


SCAPHULA MINUTA, Sp. n. 


Specimens of this species were collected by Dr. N. Annandale, 
who kindly gave me the following information:—The species 
occurs all over the Tale Sap or Inland Sea of Singgora on the 


Text-figure 6. 


Shell of S. minata, outer surface (left valve). 


east coast of peninsular Siam. It was found attached to weeds 
and stones in both fresh and brackish water. The specimens I 
have examined were found adhering to water-plants by several 
byssus threads. 

76* 


1144 THE ANATOMY OF SCAPHULA., 


Shell.—Minute, elongated, thin, closed and deep posteriorly. 
Anterior margin slightly convex, and sloping backwards from 
above and continuous with the ventral margin. Ventral margin 
slightly convex in front, straight or very slightly convex behind 
and somewhat sloping downwards. Posterior margin convex and 
sloping downwards and backwards. Ligamentous area small, 
narrow, triangular in front of and broadly club-shaped behind 
the umbones. Ligament lozenge-shaped and occupying the 
anterior and a small area of the posterior portion of the 
ligamentous area, and half the length of the entire ligamentous 
area. Hinge teeth numerous. Central teeth obsolete. Four or 
five lateral teeth, oblique in direction. Three pesterior lateral 
teeth elongated and very oblique. Surface with a thin, light- 
brown epidermis. Diagonal ridge very prominent. Surface in 


Text-figure 7. 


Animal of S. minauta, side view (left). 


front of the ridge convex, and with fine radiating and horizontal 
lines intersecting oneanother. Surface behind the ridge concave, 
and with four stout radiating ridges in addition to minute, closely 
placed similar lines all intersected by vertical ones. Length 
4-5 mm., height 2 mm., trans. diam. 2 mm. 

The present species differs from the three other known species 
of Scaphula, Benson in its minute size, in the slight tumidity of the 
valves (the proportion of the vertical diameter to the transverse 
diameter being 1:2), and in the comparative shortening of the 
body in the antero-posterior diameter. 

The type-specimens are kept in the Indian Museum. 


Haplanation of Text-figures 1-7. 


1. Anterior adductor muscle. 2. Posterior adductor muscle. 3. Posterior 
retractor pedis muscle. 4. Free end of the rectum. 5. Stomach. 6. Outline of 
the liver. 7. Labial palp. 8. Foot. 9. Visceral mass. 10. Byssus threads. 
11. Ventral fissure of the foot. 12. Ventricle. 13. Auricle. 14, Rectum. 
15. Ciliary discs. 16. Filament of inner demibranch. 


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FATAL MEMBRANES OF CHIROMYS MADAGASCARIENSIS. 


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FETAL MEMBRANES OF CHIROMYS MADAGASCARIENSIS, 


P. Z. S., 1922, HILL AND BURNE, PI. III. 


FGTAL MEMBRANES OF CHIROMYS MADAGASCARIENSIS. 


Py Zo Soy 1922, JENNIE ANID IBUIRINTES, IPL IN, 


FQTAL MEMBRANES OF CHIROMYS MADAGASCARIENSIS. 


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P. Z. S., 1922, HILL AND BURNE, PI. V. 


sept. aliw. allcav. 12 ume.c. sepe. 


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FETAL MEMBRANES OF CHIROMYS MADAGASCARIENSIS. 


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FQ:TAL MEMBRANES OF CHIROMYS MADAGASCARIENSIS. 


F@TAL MEMBRANES OF CHIROMYS MADAGASCARIENSIS. 1145 


51. The Foetal Membranes and Placentation of Chiromys 
madagascariensis. By Professor J. P. Hint, F.R.S., and 
R.H. Burne, M.A. (With an Appendix on the External 
Characters of the Foetus, by R. !. Pocock, F.R.S.) 


[Received October 24, 1922: Read October 24, 1922. ] 
(Plates L—VI.* ; Text-figures 1-6.) 


The structure of the foetal membranes and placenta of the 
Lemurs was first made known during the years 1871-1877 by 
the researches of A. Milne Edwards (1, 2, 3) and Sir William 
Turner (5, 6). Contrary to the prevailing belief, the placenta 
was shown to be of the non-deciduate diffuse type, quite unlike 
that of the Apes and Man, and more nearly resembling that of 
certain Ungulates, particularly the Pigs. 

‘The species examined at that time belonged exclusively to the 
subfamilies Lemurine and Indrisine, of Madagascar, but in 1884 
a brief description of the foetal membranes of the remarkable 
Madagascar Lemur, Chiromys, was published by Milne Edwards 
(4), and twenty-five years later Hubrécht (10) reproduced a 
photograph of the surface of the chorion of a fetal specimen of 
the same in the collection of the British Museum and now in 
our possession (our specimen B). 

In 1894 our knowledge, hitherto confined to the Madagascar 
species of the order, was extended to the Lemurs of the Old 
World by the preliminary description by Hubrecht (8) of pregnant 
uteri of Wycticebus tardigradus and Tarsius spectrum. He 
showed that in the first-named species the placenta conformed, 
with some minor though characteristic differences, to that of the 
Madagascar Lemurs, but that the placenta of Tarsius was quite 
different, and resembled that of the Apesand Man in being dis- 
coidal and deciduate. In a later paper (9) Hubrecht gave a 
more detailed account of the development of the placenta of 
Tarsius, but no further description of that of Wyecticebus has 
appeared. In 1902, however, our knowledge of the placentation 
of the Lorisiformes was further extended by the publication of a 
lengthy paper by Strahl (14) on the placenta of the African genus 
Galage, wherein it is shown that Galago, in the details of its 
placentation, agrees more closely with WVycticebus than with the 
Madagascar Lemurs, as was to be expected. Strahl in a subse- 
quent paper (15) and more recently Jenkinson (16) have added to 
our knowledge more especially of the histology of the placenta of 
these latter forms. 

The feetal membranes and placentation have now been described 
in more or less detail in representatives of all the existing 
families and subfamilies of the two groups of the true Lemurs— 


* For explanaticn of the Plates, see p. 1169, 


1146 PROF. J. P. HILL AND MR. R. H. BURNE ON THE 


the Lemuriformes and the Lorisiformes*, and asthe outcome, we 
know that all the true Lemurs are characterized by the possession 
of the same non-deciduate, diffuse, epithelio-chorial type of 
placenta,- which differs in the two groups and from genus to 
genus only in comparatively unimportant though apparently 
quite characteristic details, the general nature of which is 
indicated in the following paragraphs (c/. also 17). 

Development is of the central type, and the entire mucosa, of 
the bicornuate uterus is involved in placental formation; for, 
though development begins in one of the uterine horns, and the 
foetus, invested in its membranes, later occupies both that and 
the body of the uterus, the chorionic sac sends a prolongation, at 
any rate during the later stages of pregnancy (Milne-Kdwards, 
3; Turner, 5; Anthony, 12), also into the unoccupied or non- 
feetal horn, forming what we shall speak of as the chorionic 
appendage. The uterine epithelium, immediately below which 
are richly developed maternal capillaries, persists throughout the 
gestation period, and is regarded both by Turner (5) and Strahl 
(14) as being actively secretory. The uterine glands also persist 
in an active functional condition. ‘They mostly open in groups 
in localised bare areas in the Lemuriformes (Milne-Hdwards, 8 ; 
Turner, 5; Strahl, 15), and in Galago among the Lorisiformes 
(Strahl, 14), or their openings are more evenly distributed at 
the bottom of the uterine crypts, as in Vycticebus (Hubrecht, 8). 
The mucosa is always more or less markedly folded, the villous 
outgrowths of the chorion fitting into the depressions or crypts 
between the folds. 

The chorion is early established as a complete bilaminar 
membrane enclosing the embryonal formation, and becomes 
secondarily vascularised by the umbilical (allantoic) vessels as 
the result of the early fusion of the outer wall of the allantois 
with its inner surface. The chorion in this way is converted 
into an allanto-chorion over the greater part of its extent. The 
allantois is voluminous and multilobulate, covering the greater 
part of the inner surface of the chorion ; and in later stages the 
amnion is fused with its inner wall, the extra- embryonal cceelom 
being obliterated. ‘The yolk-sac is early separated off as a com- 
plete vesicle, appearing in later stages as a quite small flattened 
sac (Milne Tadhrawdls, 8&4; Strahl, 14). 

The chorion, except over certain small restricted areas and at 
its vaginal extremity, is produced into villous processes, which 
amongst the Lemuriformes take the form of large laminar or 
leaf-like folds, secondarily branched, or amongst the Lorisiformes, 
of nodular processes. They are covered by a simple layer of 
trophoblast (tropho-ectoderm), below which is a richly developed 
plexus of allantoic capillaries. These villous outgrowths fit into 
complementary depressions of the folded surface “of the mucosa, 


* See our reference list, in which the species studied have been noted under the 
names of the investigators. 


FQITAL MEMBRANES OF CHiROMYS MADAGASCARIENSIS. 1147 


the placental connection being constituted by the close inter- 
digitation of the vascular villous processes of the foetal chorion 
with the vascular folds of the uterine mucosa or maternal decidua. 
There is no organic continuity between the two, and at birth the 
villous processes are simply withdrawn from their mucosal pockets, 
there being apparently no loss of maternal tissue—7.e., the 
placentation is non-deciduate. 

The secretion of the uterime glands is of high importance for 
the nutrition of the feetus, on in the Lorisiformes there are 
present pockets or recesses of the chorion (chorionic recesses, 
Hubrecht, 8; chorionic vesicles, Strahl, 14) specially adapted 
for its collection and absorption, their openings in Galago lying 
opposite the depressed bare areas on which the uterine glands 
open (Strahl, 14). Corresponding bare areas of the chorion, in 
contiguity with uterine bare areas, would seem to subserve the 
same function in the Lemuriformes, though chorionic vesicles 
of simple structure are, we find, present in Chiromys and seem 
also to occur in Lemur mongoz (Strahl, 15). 

Notwithstanding the extent of our knowledge of the placen- 
tation of the Lemurs, and the fact that in essentials the foetal 
membranes and placenta of Chironys are similar to those of other 
Madagascar forms, we offer no apology for presenting yet another 
contribution to this subject, since the aberrant nature and great 
rarity of the species make it desirable that these structures should 
be deseribed and figured more fully than has been done in the 
papers (4) and (10) referred to above. 

The following account is based on the examination of two uteri 
in a late and almost identical stage of pregnancy. Both fetuses 
appear to be near full term, and are practically fully haired with 
the eyelids open. The external characters of the foetus (Speci- 
men B, Pl. IIT. fig. 6) are described by Mr. R. I. Pocock in the 
Appendix to this paper. 

Specimen A* was obtained from a female purchased several 
years ago by the Royal College of Surgeons from Mr. J. 
Hamlyn. ‘he hinder part of the body was removed, and the 
uterus preserved im set in alcohol. 

Specimen B was originally in the collection of the British 
Museum. In 1921 it was received in exchange by one of us 
(H.) through the kind offices of Mr. Oldfield Thomas. According 
to the label accompanying the specimen, it was purchased of 
Mr. Stevens on October Ist, 1864. The uterus which accom- 
panied the specimen was strongly contracted, from which we 
judge that it had been opened in the fresh state and, after 
extraction of the fcetus enclosed in its membranes, had been 
with the latter at once plunged into spirit. Apart from the 
partial separation and disappearance in places of the super- 
ficial epithelial layers, the fcetal membranes and uterus prove 
to be remarkably well preserved. This specimen formed the 


* R. Coll. Surg. Museum, Physiol. Series, Nos. 3582.5, 3582.51, 3582.52. 


1148 PRUI. J. P. HILL AND MR. RB. H. BURNE ON THE 


subject of Hubrecht’s brief reference and figure (10, p. 115 & 
fig. 151). 

For comparison with our preparations of Chiromys, we have 
had available some sections of the placenta of Wycticebus from 
the collection of the late Dr. R. Assheton, kindly placed at 
the disposal of one of us by Mrs. Assheton, and also preparations 
made from the imbedded material of Lepilemur, which formed the 
subject of the contribution of the late Dr. J. W. Jenkinson (16), 
for the use of which we are much indebted to Prof. E. 8. Goodrich, 
E.R:S: : 


Uterus. 


The uterus (specimen A, Pl. I. fig. 1) is much dilated, and forms 
a pear-shaped mass about the size and figure of a small clenched 
fist, continuous with the vagina posteriorly by a short narrow 
cervical segment. Its convex anterior end projects forward 
further on the left than on the right, and is indented to the 
right of the mid-line by a shallow vertical furrow, which marks 
the division between the two unequally dilated horns. Internally 
this division is emphasised by the projection of a semi-lunar fold 
(Pl. I. fig. 1, fid.) from the line of the external furrow for some 
2 or 3 cm. into the uterine cavity. 

From the side of each hern, a short distance posterior to its 
fundus, projects a fold of the broad ligament carrying the con- 
torted Fallopian tube, and by its reflexion forming a deep 
peritoneal pouch, within which lies the ovary attached by a long 
ovarian ligament to the uterus. About 1 cm. ventral to the 
termination of the Fallopian tube is the commencement of the 
round ligament whence the ligament passes to the groin in a fold 
projecting freely from the surface of the broad ligament. 

In specimen B, owing to strong contraction during preser- 
vation, it is probable that the shape of the uterus is to some 
extent unnatural. It is transversely extended, being nearly 
twice as broad (at the level of the Fallopian tubes) as the dis- 
tance between its cervix and fundus. The left horn is con- 
siderably more dilated than the right, and forms a rounded 
bulbous extension of the corpus uteri, with the entry of the 
Fallopian tube placed some distance behind its anterior margin. 
The right horn is much smaller with a conical apex to which is 
attached the Fallopian tube. ‘The union of the two horns is not 
indicated externally by a furrow as in specimen A, but upon the 
antero-ventral surface the position of the internal fold that 
separates them is marked by a slight ridge. 

The uterus (specimen A) when opened along its posterior 
(dorsal) aspect was found to be completely filled both as regards 
cornua, corpus, and cervix by the feetal membranes (PI. I. fig. 1), 
the well-marked chorionic appendage occupying the right cornu, 
and a prolongation of the main sac extending into the cervical 
canal (PI. J. fig. 1, ».). Within them lay the foetus, far advanced 


FETAL MEMBRANES OF CHIROMYS MADAGASCARIENSIS. 1149 


in development, occupying the left horn and corpus uteri, with 
its back turned to the right and presenting with the head. 

This position of the factus is that generally found in Lemurs 
towards the end of gestation, though sometimes the fetus may 
be in the right horn and corpus uteri (Anthony, 12, p. 247; 
Turner, 5, p. Ls) and sometimes may offer a breech presen- 
tation (Turner, 5, p. 573 & 7, p. 278) or may lie diagonally 
(Turner, 5, p. 574). 

In specimen B, so far as we can judge, the elongate embryonal 
formation lay disposed transversely in the uterus, with the head- 
end of the foetus in the corpus uteri, directed to the right, its 
thicker remainder occupying the more extensive left portion of 
the corpus uteri as well as the left cornu, whilst the chorionic 
appendage (Pls. II1., 1V. figs. 5 & 7) projected into the right cornu. 
The mid-region of the back of the fetus would thus he closely 
apposed to the internal os uteri, in a position somewhat similar 
to that occupied by one of Turnev’s specimens of Lemur rufipes 
(5, p. 574). 

The wall of the uterus in specimen A is comparatively thin. 
Its mucous lining (except within the cervix and around the 
internal os where it is smooth) is thrown into irregular anasto- 
mosing folds with intervening depressions, within which are 
received the villous folds which project from the chorion. The 
general correspondence between the mucosal folds and depressions 
on the one hand and the laminar villous folds and the clefts 
between them on the other is clearly apparent in Pl. I. fig. 1, and 
though both sets of folds are somewhat shrunken, there was clear 
evidence of close interdigitation between the two. 

In specimen A, the cervix was occupied, as mentioned above, 
by a conical smooth prolongation of the chorion, but in speci- 
men B the feetal membranes did not extend bey ond the os 
internum, and the cervix was empty and produced internally 
into longitudinal folds. The os externum in B projected frecly 
into the vagina and possessed a lobed margin. In A, it was in 
addition guarded by two semi-lunar folds, projecting from the 
wall of the vagina, as in the unimpregnated state*. The 
cervical canal in neither case showed any indication of being 
closed, as in the human subject, by a plug of secretion. 

In specimen B, the mucosa reaches its maximum thickness in 
the right cornu, and its folds are here more markedly developed 
than elsewhere in the uterus, in correspondence with the strong 
development of the laminar villi on the chorionic appendage. 
Round the internal os, as in A, the mucosa is relatively smooth. 

A low-power view of a section through the body of the uterus 
(specimen B) is reproduced in Pl. V. fig. 12. The muscularis 
(musc.) though thin relatively to the size of the uterus, is well 
developed, and in sections in the appropriate plane is seen to con- 
sist of an outer slightly thicker layer of longitudinal smooth muscle 


* R. Coll. Surg. Museum, Physiol. Series, No. 2815 A. 


1150 PROF. J. P. HILL AND MR. R. H. BURNE ON THE 


and an inner thinner layer of circular fibres; between the two 
layers there are situated the main branches of the uterine vessels. 
In the muscularis there are present numerous endothelially lined 
cleft-like lymphatic channels. 

In the intervals between its much subdivided folds the mucosa 
appears as a relatively thin layer, well supplied with vessels. 
Over much of its surface the uterine epithelium has separated 
in our material, but in places it is quite well preserved. It 
consists of a single layer of, for the most part, relatively large 
and plump cubical cells with active looking oval or spherical 
nuclei (text-fig. 1), though here and there the cells ave narrow 
columnar or even flattened and plate-like. It varies in thickness 
from ’016 to:008 mm. Whilst its free surface tends to be smooth 
and regular, its deep surface, in the absence of a basement mem- 
brane, 18 wavy and irregular, the basal ends of its cells being 
directly apphed to the walls of the subjacent capillaries which, 
as Milne Kdwards (3) and Turner (5) have shown, form a richly 
developed subepithelial plexus. Both Turner (5) and Strahl (15) 
have maintained that the uterine epithelium is actively secretory, 


Text-figure 1. 


Section uterine epithelium and underlying capillaries. 


but of that we have no positive evidence in our material. LLym- 
phoid exudation no doubt plays a considerable role in the nutrition 
of the fotus, and it is quite likely that the uterine epithelium is 
concerned in its transference to the trophoblast cells. 

The uterine glands (Pl. V. fig. 12, wé.gl.) are mainly confined 
to the basal part of the mucosa, though not infrequently they are 
found extending far out in the folds. They show a distinct 
tendency to be arranged in groups separated by regions in which 
they are sparse or absent, but we have not been able to determine 
if they open together, on restricted bare areas as described for 
other Madagascar Lemurs by Milne Edwards (8), Turner (5), and 
Strahl (15). We have not been able to distinguish such areas in 
surface examination of the mucosa, and in the ‘sections the glands 
are seen to open between the bases of the folds by way of duct- 
like involutions of the uterine epithelium. 

The glands are most numerous in the thick mucosa of the 
right horn, and are for the most part of relatively small diameter 
(048-12 mm.), though some are much thicker (up to °35 mm.). 

The glands are lined by a single layer of low columnar cells 
with spherical deeply staining nuclei in their basal halves, and 
are actively secretory, the secretion appearing in the gland 


F@TAL MEMBRANES OF CHIROMYS MADAGASCARIENSIS. 1151 


lumina as small homogeneous spherules and as a deeply staining 
coagulum. Where the glands are enlarged and not actively 
secreting, the lining cells are cubical. In Lepilemur the glands 
are very similar to those of Chiromys, but we have observed dis- 
tinct evidence of branching. Jenkinson (16) states that they 
‘‘open at the base of depressions”; we find that in this species 
the localised areas on which the glands open may or may not be 
depressed, and that they are characterised by the marked 
thickening and folding of the uterine epithelium. These areas 
are situated opposite specially thickened patches of the chorionic 
trophoblast, to which we shall later make reference. In Vyctt- 
cebus the glands are larger than in Chiromys, but are otherwise 
similar. Hubrecht (8) states that they open on the bottoms of 
the crypts in which the villi are situated. In Galago they open, 
according to Strahl (14), partly on depressed bare areas in 
relation with chorionic vesicles, partly (in the main horn) inde- 
pendently (p.193). Strahl is of opinion that in Galago extravasated 
blood in the mucosa is taken up by the uterine gland cells, and 
is utilised for the nutrition of the fetus. In none of the material 
we have examined (Chiromys, Lepilemur, Nycticebus) have we 
seen any trace of extravasated bleod in the corium of the 
mucosa, though the later is richly supplied with vessels. 


Fata MremMBRANES. 
1. Allanto-chorion. 


The allanto-chorion, or chorion as we may term it for brevity 
(PI. I. fig. 1), forms an exact cast of the interior of the uterus, 
extending from the apex of one horn to the apex of the other, 
filling the entire corpus uteri and prolonged (in specimen A) by 
a blunt conical process into the cervix as far as the os externum. 
Its surface is covered almost all over by characteristic villous 
outgrowths in the form of more or less broad swollen pleats 
marked superficially by delicate sinuous convolutions, presenting 
an appearance as nearly as possible similar to the figure given 
by Strahl (15, pl. 36. figs. 4 & 5) of the chorion of a 30-mm. 
foetus of Propithecus. The surface of the folds thus appears to be 
less elaborated than in the Lemurs described by Turner (5, 

. 575 & 7%, p. 279) or in the later stages of Propithecus 
(Strahl, 15, p. 273). ‘The primary pleats are set for the most 
part transversely to the length of the chorionic sac, becoming 
more regularly disposed in each horn in parallel circular wreath- 
like bands. 

In specimen B (Pls. I11., IV. figs. 4, 5, & 7-10), the laminar 
arrangement is in parts masked to some extent by the depth of the 
secondary foldings which break up the primary pleats and produce 
a more lobulated surface than in the specimen A (PI. IV. fig. 9). 
This lobulation must not be confused with the appearance pre- 
sented by the characteristic polygonal villi of the genus Vycticebus 


1152 PROF. J. P. HILL AND MR. R. H. BURNE ON THE 


(Hubrecht, 8, p. 93), nor with that of the somewhat similar 
villi of Galago (Strahl, 14 & 15, pl. 38. fig. 17); it is simply a 
slight elaboration, such as that described by Strahl in the later 
stages of Propithecus, of the laminar arrangement typical of all 
the Madagascar species. 

In opening the uterus of specimen A, it was observed that the 
folds and corrugations upon the surface of the chorion fit closely 
into the corresponding depressions in the uterine mucosa. The 
connection between the two is, however, relatively slight, and by 
gentle traction the chorionic sac can be detached from the uterus 
without rupture of tissue. The degree of cohesion between the 
two differs in different parts, depending on the size and compli- 
eation of the folds and the depth to which they interdigitate 
with the corresponding irregularities of the uterine mucosa, 
being greatest in the non-feetal (right) horn, where the chorionic 
folds are most pronounced (PI. J. fig. 1) and least towards the 
cervix, where they are fewer and less prominent. 

Although, as mentioned above, the folds cover most parts of 
the chorion, there are areas from which they are almost or 
entirely absent (Pl. I. fig. 1). Such bare places are very ill- 
defined, and although at first sight they suggest the bare patches 
described. by Milne Hdwards (8) and Turner (5), and figured by 
Strahl for Propithecus (15, pl. 37. fig. 14), they can hardly bear 
a strict comparison with them. In the first place, they are in 
every way so much less definite and in the second place, they 
have apparently no uterine counterpart in the form of bare areas 
of the mucosa. 

The position of these relatively bare areas is no more definite 
than their boundaries ; some can be observed towards the base of 
the left (fetal) horn and in the region of the corpus uteri, more 
particularly towards the cervix where the chorionic foids become 
smaller and less numerous than elsewhere and ultimately vanish, 
leaving the conical process of the chorion that occupies the cervix 
perfectly smooth. In this complete absence of folds at the pos- 
terior extremity of the chorionic sac in specimen A, the placenta 
conforms to the type ‘‘ placenta en cloche,” as originally described 
in Propithecus by Milne Edwards (1). 

The chorionic sac of specimen B was figured by Hubrecht 
(10, pl. GG, fig. 151). It had been opened dorsally from 2 point 
opposite the lumbar region of the feetus to the head. In all 
essential features it resembles that of specimen A. - The laminar 
folds are fairly evenly distributed upon the surface of the sac 
opposite the right side of the foetus (Pl. III. fig. 4); they are 
closely crowded together, and are arranged for the most part trans- 
versely to its long axis. Towards the hinder end they become (as 
in specimen A.) less continuous and less closely packed than further 
forward, and show four areas apparently bare, but in reality 
covered with fine ridges and grooves, giving an appearance of fine 
striation (Pls. III., TV. figs. 4 & 10). Opposite the left side of the 
foetus (Pls. IIL, IV. figs.5 & 7) is the process or appendage, as 


FETAL MEMBRANES OF CHIROMYS MADAGASCARIENSIS. 1153 


we term it, of the chorionic sac that occupied the right horn of the 
uterus. Upon its surface the chorionic folds reach their maximum 
(as in specimen A), and display strongly marked secondary folds 
and convolutions (PL. IV. figs. 7, 8, & 9), Elsewhere upon this 
face of the chorion, and more par ticularly opposite the head of the 
foetus, the folds are relatively weak, and form low ridges separated 
by wide smooth areas; similar bare places also occur opposite 
the postero-dorsal aspect of the foetus. 

In comparison with the condition of the chorion in other 
Madagascar Lemurs, the strong lamination of the appendage 
that occupies the right (non-feetal) horn of the uterus is worthy 
of notice, This prolongation of the chorionic sac may be absent 
even at near full term (Anthony, 12, p. 247, Propithecus), and 
apparently, as a rule, enters the non-feetal horn only as pregnancy 
advances (Turner, 5, pp. 578, 587; Anthony, 12, p. 247), and 
is normally less strongly laminated than other parts of the sac. 


Allantois. 


The allantois, as in other Lemurs, is a voluminous and 
characteristically lobulated organ whose cavity, though extensive 
in the circumferential plane, is flattened radially between the 
chorion with which its outer wall is fused and the amnion which 
is fused with its inner wall. Between the entodermal lining of 
the allantoic sac and the trophoblast of the chorion is situated a 
layer of delicate fibrillar connective tissue, formed from the 
allanto-chorionic mesenchyme and prolonged to form the axes of 
the villous folds. In it run the numerous branches and factors 
of the allantoic arteries and vein on their way to and from the 
subtrophoblastic capillary plexus, presently to be described. 

The allantois consists of a main lobe, into which the allantoic 
canal opens and which accordingly represents the primitive 
allantoic vesicle, and of a number of accessory lobes of irregular 
shape, which have grown out from the primary sac and approach 
one another across the rump and back of the fetus. In 
specimen A the relations of the main sac and its various 
lobes to the underlying fceetus were ascertained by carefully 
dissecting away the chorion. Two views of the preparation * 
so made are shown in Pl. IL. figs. 2 and 3 (¢f. also text- 
figs. 2 & 3), representing the left and right sides respectively of 
the feetus. 

The main lobe (partially opened in text- figs. 2&3 and Pl. IT. 
figs. 2 & 3) lies upon the left side of the posterior trunk region of 
the foetus, and extends across the ventral surface some little way 
beyond the mid-line on to the right side, covering the feet and 
tail. I¢ communicates with ane bllaneoie game which runs 
down in the umbilical cord, by a small slit-like aperture, situated 
opposite the gap between the left knee and elbow. 


* KR. Coll. Surgeons Museum, Physiol. Series, No. 3582.52 


Dam 


1154 PROF. J. P. HILL AND MR. R. H. BURNE ON THE 


From the dorsal margin of the main cavity three accessory lobes 
stretch across the left aspect of the foetus (text-fig. 2 and PI. II. 
fig. 2): (@) a posterior lobe that runs caudally nearly to the mid- 
line of the amniotic sac; (6) a dorsal lobe that covers the flank 
and sacral regions of the fetus and extends for a short distance 
beyond the dorsal mid-line; (¢) a cephalic lobe, of sausage-shape, 
that passes directly towards the head of the feetus across the 
shoulder as far as the root of the left ear. At the base of this 
lobe 1s a small ventrally directed subsidiary pocket. 


Text-figures 2 & 3. 


Spec. A, from left and right sides respectively, of foetus, after removal of chorion, 
to show the main and accessory lobes of allantois. 


The cephalic margin of the main cavity gives rise to but one 
lobule (d), an extensive finger-shaped sac folded back upon itself 
in the form, roughly, of a Y. The sac runs at first cephalad 
across the left eye of the foetus, and then is reflected round the 
cephalic pole of the amniotic sac, giving off at its furthest point 
a blunt conical process (the tail of the Y) that extends into the 
similarly shaped prolongation of the chorionic sac that occupies 
the cervix uteri. After thus curving round the cephalic pole of 
the amnion, the lobule d passes caudad along the dorsal surface 


F@rtAL MEMBRANES OF CHIROMYS MADAGASCARIENSIS, 1155 


of the fetus, and leaving the amnion extends into and fills the 
chovionic appendage that lies within the unoccupied right horn 
of the uterus. Turner (5, p. 577) states that in L. rufipes the 
allantois does not extend into “the short diverticulum of the 
chorion which occupies the non-gravid horn.” In specimen B 
we find a similar condition. The terminal process of the lobule 
extended to the mouth of the chorionic appendage and fitted 
closely within its lip, but did not project further into its cavity. 

From the ventro-caudal margin of the main cavity arises a 
lobe (¢) that curves round the ventral surface of the caudal end 
of the foetus, covering the root of the tail and the right hind limb 
and reaching to within a short distance of the free margins of 
lobes a and b. 

Upon the right side of the fetus (Pl. IT. fig. 3 and text-fig. 3) 
the main cavity gives off from its dorsal border two lobes (f and g) 
that pass towards the dorsal mid-line across the lumbar and 
thoracic regions of the featus. The caudal of these extends rather 
more than half-way towards the mid-line of the back ; the other 
(g) is a larger, sausage-shaped sac, that dilates gradually from a 
comparatively narrow origin to a bluntly rounded extremity and 
terminates upon the back of the foetus in contact with lobe d, at 
the point at which the latter leaves the amnion to enter the 
appendage of the chorion. 

Apart from the narrowness of the commencement of this lobe, 
there is no indication in the allantois of the pedunculated sub- 
sidiary lobes mentioned by Milne Hdwards (8) as occurring in 
Propithecus. 

The umbilical cord was exposed in specimen B (Tl. IIT. fig. 6). 
it measured 4:3 cm. in length, and at its emergence from the 
body of the feetus was subcylindrical. At about the middle of its 
length it became more flattened, appearing as a ribbon-like band, 
some 6 mm. in width, bordered on each side by a thinner 
selvage. It joined the inner wall of the main allantoic lobe over 
the right latero-dorsal surface of the foetus, running, attached to 
the wall, for a distance of about 1:5 cm. before finally dis- 
appearing. In specimen A, the cord passed to the left side of 
the feetus to reach the main lobe. 

The cord (Pl. V. figs. 13, wmb.c., & 14) consists of a connective- 
tissue matrix, in which are situated the allantoic canal and the 
umbilical vessels. It is invested by the amnion, the epithelial 
layer of which is quite thin, resembling in places a cuticular mem- 
brane, and below this is a thin layer of connective tissue, rich in 
cells and not everywhere clearly marked off from the connective 
tissue of the cord. Theallantoic canal (all.can.), situated between 
the vessels, possesses a wide lumen, lined by a two-layered epithe- 
lium of the transitional type. There are three umbilical vessels, 
two arteries, and one vein. The two arteries (Pl. V. fig 14, 
umb.art.) contain blood, and their walls are histologically similar, 
each possessing a compact zone of smooth muscle just outside the 


WES PROF, J. P. HILL AND MR. R. H. BURNE ON THE 


lining endothelium. The vein (fig. 14, wmb.v.) is empty, and its 
wall lack ks the compact zone of musele and appears contracted 
Ae its lining folded. 

Kpithelial pearls occur below the amniotic investment of the 
cord in the region of its junction with the allantoie sac, and are 
probably derived from the amniotic epithelium, since they are 
also occasionally found below the amnion covering the inner wall 
of the allantois adjacent to the cord-junction. 

From the junctional region, branches of the umbilical vessels 
pass off to run in the inner wall of the main allantoic sac. Some 
of these after a short course reach the allanto-chorionic con- 
nective tissue directly in specimen B, by way of three septal 
pillars (figs. 13 & 14, sept.) which extend cbliquely across the 
lumen of the main sac between its inner and outer walls, as 
Strahl appears to have observed in Galago (14, p. 182). In 
specimen A, these septa appear to be absent. Other branches 
after a straight or slightly branched course reach the allanto- 
chorion by passing round the margin of the main sac. They are 
seen emerging in Pl. IT. figs. 2 & 3and text-figs. 2 & 3 in the angles 
of the clefts between the main and accessory lobes. Yet other 
branches run straight on from the inner wall of the main sae to 
supply those regions of the chorion that are not underlain by 
prolongations of the allantois. In the chorionie connective 
tissue, the vessels subdivide and run in all directions to supply 
the subtro ophoblastic capillary plexus of the laminar villi. 

Both Milne Edwards (8) and Anthony (12) state that the 
allantoic walls in the Lemurs examined by them are devoid of 
blood-vessels. That, of course, is an erroneous statement as 
applied to the allantois as a whole, but it is partially true of the 
accessory lobes, since the inner walls of these (remarkably delicate 
and thin, as Milne Edwards remarks) are quite devoid of vessels. 
Thus it is an unexpected and somewhat remarkable fact that 
these accessory allantoic lobes in the Lemurs are not really con- 
cerned with the vascularization of the chorion, and the question 
arises as to their possible significance. One of the functions of 
the allantois is to serve as a receptacle for the urinary fluid ex- 
ereted by the mesonephroi, and it is possible that the formation 
of these lobes js to be correlated with the presence in the fetus 
of actively functional mesonephroi. In this connection it is 
interesting to note that Hubrecht (11) records that in the 6-mm. 
embryo of Wycticebus, the glomeruli of the mesonephros are 
remarkably large. 

Whatever may have been the inciting cause of the growth of 
the allantois, there can be little doubt that its lobulated form 1S, 
as Milne Edwards (3, p. 283) suggested, a direct result of the 
position early assumed by the umbilical vessels on the walls of 
the main lobe, the first formed part of the organ. If that be 
so, then the actual form assumed by the All-mniiewte 1S @ secon- 
dary matter and likely to be inconstant im its details, even in the 


F@TAL MEMBRANES OF CHIROMYS MADAGASCARIENSIS. l1157 


individuals of the sameage. From Milne Edwards's observations 
we know that in younger specimens it is simpler than in older. 

It should be noted that the main sac and its lobes do not com- 
pletely clothe the inner surface of the chorion (text-figs. 2 & 3), 
there being left between the lobes considerable areas where the 
amnion is fused directly with the inner surface of that membrane. 

The entodermal lining of the allantois is formed bya well-marked 
cubical epithelium, quite different from that lining the allantoic 
canal. Its inner surface is not everywhere smooth, but is pro- 
duced into low folds, more numerous in the main sac than in the 
accessory lobes. Its mesodermal wall is relatively thin, and 
consists of fibrillar connective tissue, dense just outside the ento- 
dermal lining, but delicate and loose elsewhere both in the inner 
and outer walls. In parts it is reticular in character. It is 
rich in cells both fixed and migratory. 


3. Yolk-sac. 


We have not found any definite traces of the yolk-stailk and 
yolk-sac in our specimens, unless an irregular epithelial strand in 
process of cornification, which is situated below the amnion of the 
umbilical cord in the region of junction and bounded on its inner 
side by a mass of gelatinous connective tissue, represents the 
degenerate remains of the stalk. Milne Edwards (38) figures 
the yolk-sac in relatively late foetuses of Propithecus diadema 
and Indris brevicaudatus (v. especially his pl. 120, where in 
a late but not yet haired fetus of the latter species it is 
shown as a quite small, flattened, and stalked vesicle, situated 
towards the distal extremity of the allantoic stalk—i. e., approxi- 
mately in the position of our cornifying strand). Strahl (14) 
simply mentions that in a 36-mm. fetus of Galago the yolk-sac 
is “gut nachweisbar.” Jenkinson (16) states that in Lepilemur 
“‘no trace of the yolk-sac was found.” 


4, Amnion. 


The amnion forms an oval sac fitting loosely around the fcetus. 
It invests the umbilical cord throughout its length, and at the 
distal extremity of that spreads out to become loosely fused with 
the inner wall of the allantoic sac and its lobes. In the intervals 
between the latter it is fused directly with the connective tissue 
of the chorion. The most extensive areas of the amnion that are 
in this way in direct apposition and fusion with the chorion lie 
towards the head-end of the fcetus (text-figs. 2 & 3 and Pl. HI. 
figs.2& 3). There is an extensive area covering the whole of the 
right side of the head and shoulders (text-fig. 3), and narrower 
patches over the left side of the head and shoulders (text-fig. 2) 
and along the back between the apices of the allantoic lobes. As 
in the higher Primates, the expansion of the amniotic sac and its 
adherence to the inner wall of the allantois have brought about 
the complete obliteration of the extra-embryonal ccelom. 


Proc. Zoou. Soc.—1922, No. LX XVII. Ci 


1158 PROF, J. P. HILL AND MR. R. H. BURNE ON THE 


Minute Structure of the Allantoic Chorion and its Villa. 


The general appearance of the chorion and its villi in section 
is illustrated in Pls. V., VI. figs. 13, 14, and 15, 

The allanto-chorion consists essentially of the outer wall of the 
allantois fused with the chorion proper. It thus comprises the 
following layers:—(1) the allantoic entoderm; (2) the composite 
allanto-chorionic connective tissue which is continued out to form 
the cores of the complexly folded laminar villi, and in which are 
situated the branches of the umbilical vessels; (3) the superficial 
covering of trophoblast (tropho-ectoderm, chorionic ectoderm). 

The trophoblast (fig. 16, é#.) throughout its extent is singularly 
uniform in character, varying only in thickness, and that with 
no apparent regularity, though frequently it is thinned in places 
over the apices of the villous branches. It attains a maximum 
thickness of -024 mm.anda minimum of ‘005 mm. or even slightly 
less. Where it is thickest, it consists of a well-defined single 
layer of cubical cells, with their outer surfaces projecting con- 
vexly. The nucleiare large and deeply staining. The cytoplasm 
of the outer zone of the cell-body is homogeneous or very finely 
granular and strongly eosinophil; that of the deeper zone below 
the nucleus is reticular in character, and stains lightly. Thereis 
no basement membrane, and the basal ends of the cells rest, just 
like those of the uterine epithelium, directly on the endothelial 
walls of the underlying capillaries (fig. 16, cap.). These are 
relatively large and very abundant, forming a richly developed 
subepithelial plexus, first figured by Milne Edwards from injected 
specimens of Mea diadema and Indris brevicaudatus (3, 
pl. 121, figs. 1, 2, & 4). In Galago also, according to Strahl (14), 
the capillary network of the villi lies in direct contact with the 
epithelial covering. 

The mesodermal core of the villi (Pl. VI. fig. 16) 1s formed by a 
variety of fibrillar connective tissue, differentiated into two zones : 
(a) a looser less deeply staining central zone (c.t.), containing 
numerous leucocytes and lymphocytes and in which are situated 
the smaller branches of the umbilical vessels; and (6) a highly 
characteristic peripheral zone in the form of a perfectly continuous 
and more deeply staining layer of uniform thickness which lies 
immediately below, and evidently forms a support for the subtro- 
phoblastic capillary plexus. This peripheral zone (Pl. VI. fig. 16, 
c.t'.) consists of a coarse fibrous reticulum, the stronger fibres of 
which run out at right angles to the surface, so that under a low 
power the zone presents a striate appearance, suggestive at first 
sight of a columnar epithelium, more especially as the nuclei of 
its cells of origin are mainly arranged in an irregular row, defining 
the deep surface of the zone. Round the nuclei there is some- 
times visible a small amount of cytoplasm, which appears to be 
directly continuous with the fibrous network. This zone would 
seem to be distinctive of the villi of Chiromys. It is not present 
either in Lepilemur or Nycticebus. In both these forms the 


F@IAL MEMBRANES OF CHIROMYS MADAGASCARIENSIS. 1159 


trophoblast is very similar to that of Chiromys, but in neither of 
them is the subtrophoblast capillary plexus anything like so well 
developed, the capillaries being smaller and much less numerous 
than in Chiromys. 

We have, unfortunately, no preparations of Chiromys which 
show the relation of the villi to the folds of the uterine mucosa, 
and are thus unable to state definitely whether the interdigita- 
tion of the two was of the same intimate nature as occurs in 
Lepilemur (Jenkinson, 16) and Propithecus coronatus (Strahl, 15), 
or was of a looser character, approximating to the conditions we 
find in Nycticebus. In the latter, Assheton states that ‘“ the 
feetal villi appear to hang in grape-like bunches into the mouths 
of much wider depressions” (18, p. 269); and the sections cer- 
tainly suggest that considerable areas of the lateral surfaces of 
the villi Bde not come into contact with the uterine folds, and 
that intimate apposition between the trophoblast and the uterine 
epithelium is largely limited to the flattened surfaces of the ends 
of the villi. In this connection it is worthy of note that gland- 
secretion 1s present in fair abundance round the proximal 
portions of the vill. In Galago, Strahl (14) states that over the 
tips (except in the region of the terminal pits) and lateral surfaces 
of the villi the trophoblast and uterine epithelium are firmly 
attacbed to each other, but in the intervillous intervals the 
connection is less intimate. 

In Lepilemur, the villi are somewhat similar in character to 
those of Chiromys, but are readily distinguishable therefrom, 
being much less coarse and more finely branched. The mucosal 
folds are likewise very thin, as is the case also, according to 
Strahl (15), in the 60-65-mm. feetal stage of Propithecus, whilst 
their relations to the villous folds are exceedingly intimate. In 
our sections of Lepilemur, the uterine epithelium, practically 
everywhere in the villous regions, has become separated from 
the underlying connective tissue of the mueosal folds, and 1s seen 
asa relatively thin layer in close and intimate apposition with 
the thicker, more cubical trophoblast. In Propithecus, Strahl 
(15) also describes the two epithelia as being in close contact. In 
Chiromys, we incline to believe that the relations between uterine 
epithelium and trophoblast were not throughout quite so intimate 
as in the two Lemurs just referred to. 


CHORIONIC VESICLES. 


When the deep surface of the allanto-chorion is examined 
after reflection of the inner wall of the allantois, numbers of 
small, flattened, ovalish bodies of a reddish-brown colour may be 
seen embedded in its substance (Pl. VI. fig. 11). These are the 
chorionic recesses first described and figured by Hubrecht (8) in 
Nycticebus, and later described by Strahl! (14) for Galago, where 
he designated them “ chorionic vesicles.” 

These bodies were encountered in both our specimens, but 


77% 


1160 PROF. J. P. ULL AND MR. R. H. BURNE ON THE 


they were larger and more easily seen in specimen B than in 
specimen A. They varied considerably in size, the largest 
measuring 5°5 x 4°5 mm. in diameter, the smallest 15 mm. They 
numbered) about sixty, and were chiefly distributed upon the 
parts of the chorion that overlay the head region of the fcetus, 
being more numerous on the right side than on the left. 

In specimen A, although a few of these bodies were as large 
as in specimen B, the majority were no larger than a pin’s head. 
Most of them were located upon that part of the chorion which 
occupied the left uterine horn, being particularly numerous upon 
those areas where the chorionic laminz were most richly deve- 
loped. Few, if any, could be located with certainty on the 
chorion within the right horn and the body of the uterus. 

In the sections (Pi. V. figs. 13 & 14, ch.ves.), the vesicles are 
seen to be situated either in the allanto-chorionic connective 
tissue, close to the allantoic lumen, or actually in that of the 
laminar villi. The vesicle is of simple structure, and unlike those 
of Nycticebus aud Galago, is devoid of villous folds ox ingrowths 
projecting into its cavity. Its opening is small and slit- like (PEW 
fig. 13, op.). Lining the cavity is a single-layered epithelium, 
representing an involuted portion of the trophoblast, but differing 
from thatinits characters. It consists of low cubical to flattened 
cells, ‘013 to°008 mm. in thickness. The cytoplasm of the cubical 
cells is reticular in character, and encloses one or more large 
vacuoles. The nucleiare mostly basal in position, and frequently 
appear shrunken and irregular. ‘The lining rests on a very thin, 
compact layer of fibrillar connective tissue. It is noteworthy that 
the vesicle-wall appears to be entirely devoid of capillaries. The 
cavity is occupied more or less completely by a coarsely granular 
coagulum, no doubt representing the secretion of the uterine 
glands, and in it there occur, here and there, degenerating cells, 
probab ly derived from the lining epithelium. 

The chorionic vesicles of Nycticebus and Galago described by 
Hubrecht and Strahl are more highly differentiated than those 
of Chiromys, and would seem to be specially adapted for the 
collection and absorption of the secretion of the uterine glands 
(Strahl, 14). In our sections of Wycticebus, we find the vesicle 
invested externally in a fairly dense fibrous connective-tissue 
coat, in which there ave present numbers of larger and smaller 
blood-vessels. The lining of the sac is formed by a flattened to 
cubical epithelium, in contact with the under surface of which 
there are here and there fair-sized capillaries. The cavity of the 
vesicle is largely occupied by vascular villi as deseribed by 
Hubrecht, clothed by an epithelium similar to that lining the 
cavity, and between the villi is a granular material Simul to 
that found in the uterine glands. 

In Galago, Strahl (14) described the occurrence of similar 
vesicles, but found that their openings, unlike those of WVyete- 
cebus, overlie depressed areas of the mucosa en which are the 
openings of uterine glands. 


FC@LPAL MEMBRANES OF CHIROMYS MADAGASCARIENSIS. 1161 


In Chiromys, the vesicles are evidently not of the same 
functional importance as in NVycticebus and Galago, and it may 
be suggested that they serve not so much as absorptive organs, 
but rather as reservoirs for the surplus of the uterine- gland 
secretion which is probably 1 in major part absorbed directly by 
the trophoblast of the iaminar villi. 

The general occurrence of chorionic vesicles in the Lemuri- 
formes has not so far been established. ‘The only reference to 
their presence that we know of is an inconclusive statement by 
Strahl.(15) that in Lemur mongoz, he observed at one place an 
indication of a chorionic vesicle devoid of internal villi, overlying 
a groove in the mucosa which might be a uterine gland area. 
But what apparently does characterise the chorion of certain of 
the Madagascar Lemurs, e. g. Propithecus, is the occurrence 
of circumscribed chorionic bare patches, opposite which there 
occur on the mucosa depressed bare areas on which the uterine 
glands open. Such complementary bare areas (and move espe- 
cially the uterine) have been described by Milne Edwards and 
Turner (with excellent figures of the uterine areas, 3, p. 280, 
and 5, figs. 8 & 9), and more recently by Strahl (15, figs. 14, 15, 
16,& 18@ &b), but none of these observers has given any 
account of the structure of the chorionic areas, though Turner 
(5, p. 582) expressly states that ‘‘the smooth, non-villous sur- 
faces of the chorion opposite the smooth areas on the mucosa are 
engaged in the absorption of the secretion of the glands.” 

Im’ the course of examining our preparations of the late 
Dr. Jenkinson’s material of Lepilemur, we observed what we 
took to be these particular areas; and on referring again to 
Dr. Jenkinson’s paper, we realised that his description (16, p. 180) 
of the simpler conditions in what he terms ‘“‘the non-placental 
[really non-villous] regions,” illustrated in his figs. 3-7 and 15, 
actually refers to the areas which we had identified as the ‘heme 
areas of previous investigators. As Jenkinson describes, 
these non-villous areas are characterised by the transformation 
of the trophoblast covering them, into a thick layer composed of 
elongated narrow columnar cells, measuring up to -063 mm. in 
height and just about six times as thick as the normal tropho- 
blast covering the villi (cf. Jenkinson’s figs. 3, 4, & 5). The 
cytoplasm of the cells is lightly staining and coarsely reticular in 
char acter, and in some of the cells slivers is a large, clear vacuole 
situated in the basal part of the cell, below the nucleus. The 
nuclei are oval and deeply staining, and are situated near the 
middle of the cell-bodies, though sometimes they approach their 
free surface. Occasional binucleate cells were observed. Specially 
characteristic of this epithelium is the production of the outer 
ends of the cells into bluntly tapering or knob-shaped processes, 
in which, occasionally, a more or less shrivelled nucleus may be 
seen. They project into a granular material, “apparently a 
secretion of the uterine epithelium,” according to Jenkinson, and 
are no doubt absorptive in function. In the basal part of the 


1162 PROF. J. P. HILL AND MR. R. H. BURNE ON THE 


layer, there occur at intervals curious spherical cells, and imme- 
diately below it, capillaries are present in fair abundance. 

The uterine areas, related to the just-described characteristic 
areas, are more or less folded, and may or may not be depressed. 
They ave characterised by the marked thickening and ridging of 
the uterine epithelium, and by the presence ‘of the openings 
of the uterine glands. The uterine epithelium appears as a 
deeply staining layer, thrown into low ridge-like folds (Jenkinson, 
fig. 15) and about five times as thick as the ordinary epithelium 
covering the mucosal folds. It is composed for the most part 
of narrow columnar or elub-shaped cells, with deeply staining 
eytoplasm and chromatin-vich nuclei. Jenkinson records the 
presence of goblet ceils amongst the ordinary epithelial cells, but 
we have failed to detect them, and considers the epithelium 1S 
secretory. However that may be, and it is probable, coarsely 
granular material similar to that found in the uterine glands 
occurs in the interspaces between the thickened trophoblast 
and the ridged uterine epithelium ; and there can be no doubt, 
we think, that these non-villous chorionic areas have the same 
functional significance as the chorionic vesicles of the Lorisi- 
formes. Indeed, in view of the close adherence of the uterine 
epithelium to the trophoblast of the chorionic villi, characteristic 
alike of Propithecws (Strahl) and Lepilemur (Jenkinson), special 
absorptive areas in relation to the openings of the uterine glands 
would seem to be necessary if the secretion of the latter is 
to be adequately utilised. 

It may be noted that these areas attain a relatively consider- 
able size, as may be judged from Strahl’s figures of Propithecus 
(15, taf. 37). The largest area we have examined in Lepilemur 


measures over 3 mm. in diameter. 


CoNCLUSION. 


in the contribution by one of us(H.) to the discussion on the 
zoological position and affinities of ZVarsius (17), the view is 
expressed (p. 477) that the placentation of the Lemuroids 
is essentially primitive, “presenting us with a simple little 
specialised type of placenta from which the more advanced and 
presumably more efficient arrangements in the other Primates 
may quite easily have originated as the result of adaptive 
specialis sation in the course of evolution.” More detailed 
acquaintance with the Lemuroid placenta has only served to 
strengthen us in that conclusion, and has, moreover, entirely 
failed to reveal the slightest evidence in support of the view 
expressed by Hubrecht (10, p. 145) and also by Assheton 
(18, p. 268) that the Lemuroid placenta is a secondarily simplified 
and derivative type. 

These two authorities base their belief very largely on 
theoretical considerations concerning placental evolution, and 
consider that the diffuse placenta of the Lemurs is different from 
that of the Ungulata, Manide, Cetacea, ete. 


F@TAL MEMBRANES OF CHIROMYS MADAGASCARIENSIS. 1163 


In his speculations on the phylogeny of the placenta, Hubrecht 
(10) starts out by denying a Reptilian origin for the Mammalia, 
and postulates a primary phagocytic and absorptive activity on 
the part of the trophoblast. From the very start of uterine 
gestation, the trophoblast proliferating rapidly, invaded and 
destroyed the maternal decidual tissue, and as the result, there 
was formed a localised placenta of the ‘deciduate” type in 
which the maternal blood circulated either in endothelially lined 
eapillaries enclosed by trophoblast (Carnivora) or actually in 
lacunar spaces in the trophoblast itself (Insectivora, Rodents, 
ete.). That being the primary and original type of placentation, 
it follows that Ail the “diffuse” varieties of placenta in which 
such phagocytic activity on the part of the trophoblast ‘ was 
insignificant or absent (Lemurs, certain Hdentates, and many 
Ungulates)” have been derived by simplification from the more 
soe ite “‘deciduate” type. Hubrecht admitted, however, that 
‘we cannot for the present indicate the intermediate steps by 
which the simplification of a placenta of the intectetons 
or Primate type. down to that of the present Lemurs was 
brought about ...” (10, p. 115); but he saw “no reason why 
this ‘simplification should not have arisen more than once” 
(p. 145)—-7.e., the diffuse placentation of Lemurs ‘‘ was not 
necessarily obtained along the same hereditary line of develop- 
ment” as that of Ungulates. Indeed, he thinks there is sufficient 
evidence to justify the drawing of a sharp line of distinction 
between the placentation in “ghese two groups. The late 
Richard Assheton, in his critique of Hubrecht’s monamental 
paper (18), says (18, Pave): oll SEY agree with Hubrecht’s 
opinion that the diffuse placentation of the Lemurs is different 
from that of the true plicate * forms of Ungulates, Cetacea, 
some Edentates, ete.” Further he writes (p. 265): ‘‘ Whether 
the Carnivora or the extreme cumulative or extreme plicate 
[placenta] is the most primitive, it is very difficult to say” 
nevertheless he goes on to suggest that the Lemurine placenta, 
which he terms ‘ pseudo- plicate,” may possibly have been 
derived by reduction from a cumulate type such as is found 
in the Carnivora, “by way of such conditions as Hylobates, 
Semnopithecus, Cercocebus, by the gradual supersession of the 
glandular activity of the maternal uterus over the phagocytic 
activity of the foetal trophoblast, and the filling of the blood 
spaces, into which the fetal villi originally hung, with uterine 
secretions instead of extravasated maternal blood ” (p- 269). 
But why the Lemurs after having evolved a cumulate (deciduate) 
placenta should have been under the necessity of substituting for 
it the plicate (non-deciduate) type 1s not explained. 


* Assheton suggested a grouping of placental forms based on the behaviour o 
the trophoblast into (a) Cumulate, characterised by radial, more or less loca 
thickening of the trophoblast which is actively phagocytic (= ‘‘ Deciduate’’): 
(6) Plicate, characterised by tangential extension and folding of the trophoblas’ 
with little or no phagocytic activity (= ‘‘ Nondeciduate’’). 


1164 PROF. J. P, HILL AND MR. R. H. BURNE ON THE 


We must confess we are quite unable to accept the views of 
these two authorities, either in respect of the secondary nature of 
the diffuse placenta of the Lemurs or in regard to its dis- 
similarity to the diffuse placenta of other Mammals. It appears 
to us they have attempted to read the story of placental 
evolution the wrong way round. 

In our opinion, any attempt to trace the evolution of mam- 

malian placental arrangements must be based on a consideration 
of the probable conditions which obtained at the time viviparity 
replaced oviparity. Knowing the arrangement of the foetal 
membranes in the existing ton otrenies, we can picture the 
conditions with reasonable certainty. We snould postulate, 
then, that the common ancestral stock, from which the Di- 
delphia and Monodelphia diverged, possessed an arrangement 
of the feetal membranes similar to that of the existing Mono- 
tremes and of Phascolarctos amongst the Marsupials-—z. e., the 
outer wall of the embryonal formation consisted as to one-half 
or thereabouts of unsplit blastocyst wall or omphalopleure, in 
part vascularised by the vitelline vessels of the vascular area 
of the yolk-sac, and as to the remainder, of allanto-chorion, 
vascularised by the umbilical vessels of the vesicular allantois. | 
The uterine mucosa possessed a persistent covering epithelium, 
functional uterine glands, and a rich vascular supply. 

These being the structural elements involved, the primitive 
placenta was constituted simply by the close apposition of the 
omphalopleure and the allanto-chorion with the vascular lining 
of the uterus. Here we have the simplest possible type of 
placenta, partly allantoic, partly vitelline and diffuse in the 
sense that the whole of the outer surface of the embryonal 
formation is involved in the carrying on of the exchanges 
between the maternal and fcetal blood-streams. The trophoblast 
covering the entire outer surface is as yet a simple layer, absorp- 
tive but not actively phagocytic. 

Out of the primitive placenta here outlined, we see no difficulty 
in deriving on the one hand, the relatively simple placental 
arrangements characteristic of the Marsupialia and on the 
other, the varied types of placenta characteristic of the Mono- 
delphia, the non-deciduate diffuse type representing only a 
relatively slight elaboration of the primitive form, and the 
varieties of the ‘‘deciduate” type resulting from the assumption 
by the trophoblast of active phagocytic properties, the precise 
characters assumed by the placenta being in all cases determined 
by a variety of conditions, intrinsic and environmental or uterine. 
We therefore remain firm in our belief that the placentation of 
the Lemurs is genuinely simple and little specialised. 

Finally, as concerns the reputed distinction between the 
Lemuroid diffuse placenta and that of other Mammals, we 
readily grant there are differences in detail, but we quite fail 
to see that there is any fundamental difference, or ever enough 
difference to justify the calling of the former ‘“ pseudo-plicate” 


FETAL MEMBRANES OF CHIROMYS MADAGASCARIENSIS. 1165 


and the latter “plieate.” Both are plicate in Assheton s sense, 
and are characterised on the feetal side by the presence of a 
relatively passive trophoblast, markedly folded and of great 
extent, by the vascularisation of the chorion through a large 
vesicular allantois and by the reduction of the yolk-sae and its 
vessels; and on the maternal side, by the persistence of the 
uterine epithelium (at least in the majority) and of the uterine 
glands in an active functional condition. In our view, the 
resemblances far outweigh the differences in detail. 

We are accordingly in entire agreement with Jenkinson (16) 
in believing (1) that the resemblance between the non-deciduate 
placentation of the Lemuroids and that of other Mammals, e. g. 
the Ungulata, are due to their derivation “from a common 
ancestral ‘indeciduate’ type”; and (2) that the deciduate 
heemochorial type of placenta characteristic of the other Pri- 
mates has been evolved from the non-deciduate Lemuroid type, 
quite independently of that of the Rodents. Insectivora, ete.—— 
another illustration of the principle enunciated by H. F. Osborn 
that “the same results appear independently in descendants of 
the same ancestors.” 


We are greatly indebted to Mr. F. Pittock for invaluable help 
in the preparation of Pls. IJI.-VI.and to Mr.S. Steward for the 
photographs of Specimen A, Pls. I. & IT., and to Mr. Terzi for 
retouching the same. 


APPENDIX. 


On the External Characters of a Foetal Chiromys. 


Byala) POCOCK giao. 


At the request of the authors, I undertook to examine and 
describe the external features of the fetal Chiromys (Specimen B, 
Pl. IIT. fig. 6), the placenta of which is described in the pre- 
ceding pages by Prof. Hill and Mr. Burne. 

The approximate dimensions in millimetres of the specimen, a 
male apparently at full time, are as follows :—Dorsal contour 
length (snout to root of tail) 165; head and body from crown to 
root of tail 106; head from nose to occiput 37; ear from inter- 
tragal notch 21; tail 103; upper arm 25 ; fore-arm 21; paim of 
hand 12; thigh 31; lower leg 31; sole of foot 18. The general 
proportions do not differ greatly from those of the adult, the tail 
and arm being relatively as long, but the head is relatively 
larger, the leg shorter, and the digits of the hand shorter than 
in the adult. 

The coat consists of comparatively short and sparse adpressed 
hairs, brown and grey in colour; on the hands and feet and tail 


Wy 


1166 MR. R. I. POCOCK ON THE 


the tint is blacker ; the muzzle and cheeks are nearly naked. In 
the adult the coat is thick and woolly and black in colour, with 
long black and white bristles intermixed. The head is relatively 
larger, and has a more vaulted cranium than in the adult. 

The rhinarium is well developed and like that of the adult. 
So also are the facial vibrisse, the genal and superciliary tufts 


Text-figure 4. 


Nata C oy SS 


A. Head of fcetal Chiromys. B. Base of ear of same. C. Base of ear of adult. 
D. Rhinarium of fetal Chiromys. 


being specially well marked. There is asingleinterramal vibrissa. 
The eyes are much smaller than in the adult, oval and pointed at 
each end, like the eyes of a diurnal mammal. 


EXTERNAL CHARACTERS OF A FQ@XTAL CHIROMYS. 1167 


The ear is covered with short hair externally, and is flat and 
not hollowed, but the ridges and depressions characteristic of 
the adult are all indicated. In the adult the ear is naked 
internally and deeply hollowed; it is also relatively much larger 
than in the feetus, its length being equal to the distance between 
its intertragal notch and the end of the rhinarium, whereas in 
the young its length is only equal to the distance from the end 
of the rhinarium to a point just behind the eye. 

The tongue differs in at least two particulars from that of the 
adult. The distal margin of the sublingua is irregularly pointed 
and serrulated, and the median inferior ridge ends in a soft 
straight point, not ina hard down-curved hook. In the points 


Text-figure 5. 


Hand and foot of foetal Chiromys. 


in which the sublingua differs from that of the adult, it 
approaches the less specialised condition seen in the typical 
Lemurs. 

The fore limbs closely resemble those of the adult, but the 
digits are relatively much shorter. The fourth digit, for 
example, is only a little longer than the palm, whereas in the 
adult it is nearly twice the length. The pollex also is relatively 
much nearer to the second digit than it is in the adult. 

The hind limb closely resembles that of the adult in ail im- 
portant respects. It is, however, relatively shorter in all its parts. 


1168 PROF. J. P. HILL AND MR. R. H. BURNE ON THE 


The external genitalia are remarkable for the length and 
~ thickness of the penis, the distal end of which is curved forwards, 
with its posterior, or inferior, surface strongly convex. - Proxi- 
mally it arises in front between the two well-marked inguinal 
mamme, and its integument here is mesially grooved. Imme- 
diately behind its posterior point of insertion there is a little 
bilobed thickening of the skin, representing the tip of the 


Text-figure 6. 


a a 
Nee Ji ye 
| ay! 


p: 


A 


A. Lateral view of the hind quarters to show the prominence of the penis. 
B. Ventral view of the hind quarters, with penis turned aside to the left. 
m. YaammMma; p. penis; s. scrotum; w. umbilical cord. 


undeveloped scrotum, and above this on each side a shallow 
groove runs upwards and forwards to the level of the mamme, 
apparently defining the inguinal position of the testes. The 
penis, the inner surfaces of the thighs, and the abdominal region 
are much more scantily hairy than the upper and outer surfaces 
of the body and limbs. 


List of References. 

1. Mitne Epwarps, A.—‘‘ Observations sur quelques points de 
’Kmbryologie des Lémuriens.” CC. R. Acad. Sci., t. 73. 
14 Aout, 1871, p. 422. 

2. MitnE Epwarps, A.—/bid. Ann. Sci. Nat. ser. 5, t. xv. 
1872. (Propithecus, Lepilemur, Hapalemur, Chirogaleus.) 

3. Mitne Epwarps, A.—-“‘Du Placenta et des Hnveloppes 
foetales ” (des Lémuriens). Chap. ix.: ‘“* Histoire Natu- 
relle des Mammiféres de Madagascar.”. T.1. & Atlas, in 
Grandidier’s ‘ Histoire de Madagascar,’ vol. vi. Paris, 
1875. (Propithecus diadema, P. edwards, P. ver- 
reauxur, Avahis laniger, Indris brevicaudatus. ) 

4. Minne Epwarps, A.—“Sur la disposition des enveloppes 
foetales de l’Aye-Aye.” C.R. Acad. Sci. Paris, t. 99. 
1884, p. 265. (Chiromys.) 

5. Turner, W-.—‘‘ On the Placentation of the Lemurs.” Phil. 
Trans., vol. 166.1877, p. 569. (Propithecus diadema, 
Lemur rufipes, Indris brevicaudatus.) 


FETAL MEMBRANES OF CHIROMYS MADAGASCARIENSIS. 1169 


6. Turner, W.—‘‘ The Placentation of the Lemurs.” Journ. 
Anat. & Physiol., vol. xii. 1878, p. 147. 

7. Turner, W.—‘‘ An additional contribution to the Placenta- 
tion of the Lemurs.” Proc. R. Soe., vol. xliv. 1888, 
p- 277. . (Lemur xanthomystac.) 

8. Husprecet, A. A. W.—‘‘Spolia Nemoris.” Quart. Journ. 
Micro. Sci., vol. xxxvi. 1894, p. 90. (Wyctrcebus tardt- 
gradus, Tarsius spectrum.) 

9. Huprecut, A. A. W.—“‘ Ueber die Entwicklung der Placenta 
von Tarsius” ete. Proc.4th Internat. Congress, Zoology. 
Cambridge, 1899. 

10. Husprecut, A. A. W.—“ Karly Ontogenetic Phenomena in 
Mammals” ete. Quart. Journ. Micro. Sci., vol. li. 1909, 

Gs: 

11. aaa: a A. A. W.—Normentafel zur Entwicklungs- 
geschichte des Plumplori (Vyeticebus tardigradus). 
G. Fischer, Jena, 1907. 

12. Anruony, R.—‘“ Note sur un foetus de Propitheque et ses 
membranes.” Ann. Sci. Nat., ser. 9, t. vil. 1908, 
p. 248. (Propithecus verreauxit. ) 

18. Broca, P.—“ Le Placenta des Lémuriens.” Bull. Soc. 
Anthrop. Paris, 2 ser., t. xii. 1877, pp. 267-70. (Pro- 
pithecus diadema.) 

14. Srrant, H.—‘ Der Uterus gravidus von Galago agisymbanus.” 
Abhandl. Senckenbere, Gesellsch., Bd. xxvi. 1902, 
p. 155. 

15. Srrani, H.—“ Beitrige zur vergleich. Anatomie der Pla- 
centa.” Abhandl. Senckenberg. Gesellsch., Bd. xxvii. 
1904. (Propithecus coronatus, Lemur albimanus, L. 
MOongoz.) 

16. Jenkinson, J. W.—“ The Placenta of a Lemur.” Quart. 
Journ. Micro. Sci., vol. lxi. 1916, p. 171. (Lepilemur sp. 
(2 ruficwudatus.)) 

17. Hinz, J. P.—“‘ The Affinities of Zarsiws from the Kmbryo- 
logical Aspect.” P.Z.S. 1919, p. 476. 

18. Assunton, R.—Professor Hubyecht’s paper on the Harly 
Ontogenetic Phenomena in Mammals, ete. Quart. 


Journ. Micro. Sci., vol. liv. 1910. 


EXPLANATION OF THE PLATES. 
List of Reference Letters. 


all.c. allantoic cavity; all.can. allantoic canal; all.ch. allanto-chorion; adl.ent. 
allantoic entoderm ; all.qw. inner wall of allantois; ammn. amnion; cap. allantoic 
capillaries; ch.a. chorionic appendage; ch.ves. chorionic vesicle; c.¢. connective 
tissue of villus; c.¢’. peripheral zone of same; jid. fold between uterine cornua ; 
muse. muscularis; op. opening of chorionic vesicle; p. smooth prolongation of 
chorion into cervix ; sept. septal pillar between mner and outer walls of allantois ; 
tr. trophoblast; wmb.c. umbilical cord; wmb.art. umbilical (allantoic) artery ; 
umb.v. umbilical vein; wt.gl. Sine glands ; vill. chorionic villus. 


Fig. 1. Uterus, Chiromys, Spec. A., opened from the dorsal side, showing the 
chorionic sac in situ. the folded character of the mucosa, the chorionic 
appendage (ch.a.) projecting into the right cornu, and the non-villous 
prolongation (p.) into the cervix are well seen. féd. fold between uterine 
cornua. About nat. size. 


1170 


Fig, 


Fig. 


2. 


F@TAL MEMBRANES OF CHIROMYS MADAGASCARIENSIS. 


Foetal Chiromys, Spec. A. The chorion has been removed, exposing the 
main sac of the allantois and its accessory lobes on the left side of fetus. 
The head of the fcetus is towards the lower side of the plate. The main 
sac has been opened up, exposing the opening of the allantoic canal, into 
which a white rod has been passed. (Cf text-fig. 2 and text, p. 1153.) 
Shehtly reduced. 


. Feetal Chiromys, Spec. A. ‘The same preparation as in fig. 2, but from the 


right side of fetus. An extensive area of the amnion on the right side 
of the head of the foetus has been removed. (Cf. text-fig. 3 and text, 
p. 1155.) Slightly reduced. 


. View of the chorionic sac from the right side of the foetus, Spec. B, show- 


ing the arrangement and character of the laminar villi. The head of 
the fcetus is towards the left side of the plate. Slightly reduced. 


. Similar view of the left side of the chorionic sac, Spec. B, to show especially 


the chorionic appendage which projected as in Spec. A into the right 
cornu. Slightly reduced. 


. Fetus, Spec. B, from the right side, after removal from the foetal mem- 


branes. The severed umbilical cord is seen overlying the right fore-arm. 
Slightly reduced. 


. Chorionic appendage, Spec. B, extended so as to show the folded laminar 


character of the villi of its upper surface. Slightly reduced. 


. Portion of the same, more highly magnified. X about 6. 
. Portion of the exposed surface of the chorionic appendage, Spec. B, showing 


the convoluted appearance of the villi. X about 6. 


. Enlarged view of the portion of chorion, Spec. B, overlying left side of 


hinder region of foetus, showing the folded villi and the non-villous 
ridged areas. > about 6. 


. (PI. V1). View of the inner surface of allanto-chorion after reflection of 


the inner wall of allantois (all.w.), showing the chorionic vesicles 
(ch.ves.). 


. Section through body of uterus, Spec. B. X about 12. 
. Trans. section through the allanto-chorion in the region of junction of 


umbilical cord with the inner wall of allantois, showing the cord (umb.c.), 
the allantoic cavity (all.cav.) crossed by two septa (sept.), the allanto- 
chorion (all.ch.) with its villi and two chorionic vesicles (ch.ves.), in the 
left one of which the opening (op.) is visible. X about 8°5. 


. Section similar to the preceding, more highly magnified to show the struc- 


‘ture of the umbilical cord, allantois, and chorion. X about 12. 


. Section through the allanto-chorion and one of its villi, showing the 


allantoic entoderm (all.ent.), the connective tissue of the allanto-chorion 
(all.ch.) prolonged to form the axis of the villus (c.t.), in which are seen 
in section the smaller branches of the allantoic vessels. XX about 48. 


. Section through a villous branch, to show the trophoblast (¢7.), the under- 


lying capillary plexus (cap.), and the connective-tissue core (c.t.). Note 
the characteristic zone of the latter, below the capillartes (¢.t’.). > about 
360. 


EXTERNAL CHARACTERS OF THE BEAVER. Wall 


52. On the External Characters of the Beaver (Castoride) 
and of some Squirrels (Sciuride). By R. I. Pocock, 
WMelBoSicy LWA 

[ Received October 20, 1922: Read November 21, 1922.] 


(Text-figures 38-60.) 


ConTENTS. 

Page 

Introduction.. Bae Re ROC eco ORE GAN Ut Gite 
The Muzzle, Monte and maaal Vane Rs espoauTe ALes 1173 
A Me) OR Weel, curse Be a RSG Galt GUE Rae NU AMOR eae Sc\NLIUI/Y/ 
The Feet Uelnecsiet Ja nce S 
a. Feet of the Tr Spiral eae iSduinrelen eySeat Sands 1184 

b. Feet of the Tropical African Squirrels ............. 1188 

e. Feet of the European and American Squirrels...... 1190 

d. Feet of the Chipping Squirrels ...................... 11938 

e. Feet of the Bristly Ground Squirrels................ 1195 

f. Feet of the Sousliks and Marmots ................... 1198 

g. Feet of the Flying Squirrels (Petaur sae) Joy 1200 

h: Feet of the Beaver (Castor) ...............00ce0--e--. 1205 
hepAnusiandykixternall Genitalia. eee eee eeeee cae L208 

Introduction. 


The Squirrels, Marmots, and other admittedly related genera 
forming the Sciuridz were formerly associated with the Beavers, 
or Castoride, in a primary division of the Simplicidentate 
Rodents, the Sciuromorpha*. It has become the fashion in 
recent years to drop this group. Nevertheless, the two families 
are still suggestively juxtaposed by systematic writers, and no 
one seems to doubt the validity of the claim of their relationship 
based upon certain cranial characters connected more particularly 
with the structure of the zygomatic arch. Miller, for example, 
in his key to the families of Rodents of Western Europe, places 
the Castoride and Sciuride under a heading distinguished from 
the other families by having the anterior portion me the zygo- 
matic arch formed chiefly by the jugal bone and the infraorbital 
foramen small; and the opening words of his diagnosis of the 
family Castoride are ‘‘ characters essentially as in the Sciuridee.” 
Alston, Thomas, Flower, Tullberg, Winge, Max Weber, and 
others all agree in placing the Castoride and Sciuride in the 
same section of Rodents. 

As regards the division of the Sciuride into subfamilies there 
has been, and is still, some difference of opinion. By most 
authors the Marmots and Sousliks were separated as a subfamily 

—Arctomyine (Marmotine)—from the Squirrels (Séiwrine) ; but 


* Also containing the Aplodontide, about which I have no first-hand knowledge. 
This family was, however, removed from the Sciuromorpha by Thomas, P.Z.S 
1896, p. 1015. 


Why oe MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS 


Forsyth Major and Miller withheld that rank from them. On 
the other hand, the Flying Squirrels, which were usually dis- 
tinguished as the subfamily Péeromyine (Petawristine), were 
granted by Miller the full family rank as Petauristide, equiva- 
lent to the rest of the Squirrels or Sciuride. A fourth sub- 
family—Vannosciwrine—admitted as valid by some authors, was 
shown by Thomas in 1915* to be composed of heterogeneous 
elements, the evidence of affinity between the genera derived 
from skulls being entirely untrustworthy. 

The question of the generic classification of the Sciuride need 
not be discussed at length. All that it is necessary to say is 
that in 1915 Thomas clearly showed, by his study of the os penis 


Text-figure 38. 


A. Muzzle of Sciwrus vulgaris, with rhinarium exposed and partially 
concealed by hood. 

B. The same of S. saléwensis, with rhinarium exposed and entirely hooded. 

C. Ditto of Geosciurus capensis. 


or baculum, that previous attempts to affiliate the genera and 
subgenera by their skulls and teeth and other characters had 
in many instances yielded erroneous results, such as association 
of Callosciurus with Sciwrus and of Nannosciurus with Myo- 
sciurus. No living student of the family would claim that we 
know the relationship of all the groups of genera to each other. 
The vagueness and diversity of epinions on this point, coupled 
with the hope of finding unrecorded features of systematic value, 
induced me to take up the study of the external characters 
presented by the ears, feet, and other organs which had been for 


* Ann. & Mag. Nat. Hist. (8) xv. p. 386. 


OF THE BEAVER AND OF SOME SQUIRRELS. es 


the most part neglected by previous workers, partly owing to 
scarcity of materials, either fresh or preserved in alcohol, suitable 
for the purpose. The result of my observations, extending over 
many years and based upon specimens that died in the Zoological 
Gardens, form the subject-matter of the present paper. Examples 
of a good many genera have passed through my hands in that 
way ; “put in some cases I have been compelled to rely upon dried 
skins contained in the collections of the Zoological Society and 
of the Natural History Museum *. 


The Muzzle, Mouth, and Facial Vibrisse. 


The muzzle in all the genera of Sciuride and Petauristide 
is very uniform in structure. A peculiarity of the rhinarium is 
the mechanism by which the nostrils are closed. The hairy skin 
of the muzzle just behind it on the dorsal side is capable of being 
moved downwards and forwards, carrying the supranarial por- 
tion of the rhinarium with it, the two constituting, as it were, 
a kind of hood or cap over the nostrils. When completely 
lowered, the inferior edge of this hood forms a transverse or 
crescentically curved line defined from the upper lip by a narrow 
slit of the same shape. The line along which the upper half of 
the rhinarium is folded down is usually, at all events, marked 
by a transverse groove just above the upper edge of the nostrils. 
The rhinarium itself is naked and has a well-marked median 
groove between the nostrils, which are of the usual shape, with 
the typical narial slit behind and externally, and the expanded 
orifice in front and internally. They are moderately widely 
separated and look straight forwards. The rhinarium varies 
somewhat in shape. For instance, in Sevwrus vulgaris it is 
comparatively deep, and has a nearly straight or convexly 
rounded upper edge. In Sciurus saltwensis it appears to be lower 
with the upper edge not nearly so raised. It also appears to be 
broader and altogether larger relatively in Marmota than in 
Cynomys. But although these observations were made upon 
fresh specimens, the apparent height of the rhinarium varies so 
much in accordance with the expansion or contraction of the 
hood, that I have found it impossible to make any useful syste- 
matic generalisations with regard to its shape. (Text-figs. 38 & 
39, A-C.) . 

The two halves of the upper hp are completely divided up to 
the inferior median point of the rhinarium. 

The rhinarium in Castor is very different from that of the 
Squirrels. The nostrils are widely separated, look outwards and 
upwards and are valvular, closing to form a crescentic slit. 
The area around them is naked, and the wide naked internariat 
space has an oblique crest of hairs passing over the nostrils on 


* Thave to thank Mr. Oldfield Thomas not only for permission to examine the 
collections under his charge, but also for much kind help in the identification of some 
of the species of this difficult group about which I was in doubt. 


Proc. Zoou. Soc.—-1922, No. LX XVIII. 78 


1174 MR. R. i. POCOCK ON THE EXTERNAL CHARACTERS 


each side, and there is an upstanding crest on each side of the 
front of the infranarial portion of the rhinarium. The rhina- 
rium is not mesially grooved and the upper lip is not cleft, 
although the short hairs on each side of the middle line are 
inclined inwards and downwards in a different direction from 
those external to them and form adefinite crest or fringe. (Text- 
fig. 40, A, B.) 

The mouth in Sciuride and Petauristide is provided with two 
palatal lobes of hairy integument jutting inwards from the 


Text-figure 39. 


Gsintty 
WM Aten 


tt 


ont 


SN 


y 
CA tee sey RRQ 
GOSS 

Ubi 


A, Muzzle of Cynomys ludovicianus, with rhinarium exposed. 

B. » 9» Ratufa indica, with rhinarium partially hooded. 

C. Mouth and muzzle of Marmota marmota, showing the cheek-lobes 
projecting into mouth, and rhinarium exposed. 

D. Mouth and muzzle of Castor canadensis, half side view, to show mat of 
bristles (7.) inside the cheek. 


cheek. These are capable of meeting over the palate behind 
the incisor teeth, but they are never fused in the middle line. 
Two similar but smooth and altogether smaller lingual lobes jut 
inwards on each side of the tongue. (Text-fig. 39, C.) 

Definite cheek pouches are developed only in three genera— 
Tumias (Huiamias), Citellus,and Cynomys. I have not been able 
to examine them in Cvtellus, but in Tamias and Cynonys they 


Sip 


OF THE BEAVER AND OF SOME SQUIRRELS. 1175 


open into the mouth approximately on a level with the anterior 
end of the molar teeth. In Vamias they are of large size, and, 
when unstretched, reach back as far as the ear. In Cynomys 
they are much smaller and do not reach beyond the eye. Judging 
from the descriptions of the pouch of Citellus as “large,” I 
presume it resembles that of Zamias and Hutamias. Marmota 
is usually described as being without check pouches, and I failed 
to find a trace of them in the typical EHuropean species J. 
marmota. Nevertheless, in his diagnosis of the genus, Miller 


Text-figure 40. 


La ey WW ~ Wee 
(2 Re NUR 
JOG 


A. Muzzle and rhinarium of Castor fiber. 
B. Rhinarium of the same from the side. 
C. Ear of Castor fiber, with position of orifice dotted in. 


says ‘“‘ cheek pouches rudimentary or absent.” Elliot * describes 
them as small in the American species, assigned to Marmota, 
and Cory cites their presence as a character of the subfamily 
Marmotine. (Text-fig. 41, C, D.) Fresh information is clearly 
wanted on this point. 

In its palatal and lingual lobes the mouth of Castor resembles 
that of the Sciuride, but it differs in having a large pad on each 
side of the cheek, lying farther within the mouth than the 
normal lobes. Hach of these pads is provided with stiff, short 


* Field Columbian Museum, Zool. ii. p. 104, 1901. 
«7 ©The Mammals of Illinois and Wisconsin,’ p. 137, 1912. 


78* 


1176 MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS 


hairs, forming a kind of mat, and projecting downwards and 
backwards. The hairs of these pads meet in the middle line 
over the tongue in front of the molar teeth, and close the orifice 
of the throat. (Text-fig. 39, D.) 

The tufts of the facial vibrisse in all the genera of true 
Squirrels, in the Sousliks and Marmots are normal in number 
and position, the mystacials, supercilaries, and genals being long 
and comparatively numerous and the interramals, though less 
evident, being always detectable. The only variations to record 


Text-figure 41. 


A. Head of Callosciurus prevosti, showing the tufts of vibrissee charac- 
teristic of the Sciuridee. f 

B. 4, 4, Geosciurus capensis, with additional tuft of superciliary 
vibrissz. 

C. 4, 4, Cynomys ludovicianus, the curved dotted line showing the 
size and position of the small cheek-pouch. 

D. ., 5, Hutanias quadrivittatus, with the external wall of the large 
cheek-pouch cut open. 


are the absence of the interramal tuft in the Flying Squirrels 
(Petauristide) and the presence in the Bristly Ground Squirrels 
(Yerws and its allies) of a supplementary superciliary tuft of 
long vibrissz over the posterior angle of the eye. In this 
particular these Ground Squirrels are unique, so far as I know, 
amongst mammals. (Text-fig. 41.) 

The Beavers have a few short, stiff mystacial and superciliary 
vibriss, but the genal and interramal tufts are absent. In the 


— 


OF THE BEAVER AND OF SOME SQUIRRELS. 1177 


absence of the genal tufts the Castoride differ from the Scinride 
and Petauristide. 

The systematic results yielded by the characters discussed may 
be tabulated as follows :— 


a, Rhinarium unhooded, not wholly naked, nostrils wide apart, 
lateral, valvular; upper lip uncleft; mouth with a bristly 
pad inside the cheek ; ; no genal vibrisse ... mn . Castoride. 
. Rhinarium hooded, 7. e. capable of being covered by : a flap. of 
skin from the upper surface of the nose, naked, nostrils close 
together, looking forwards, not valv ular ; ; upper lip deeply 
cleft to rhinarium ; ; mouth without Pasty pad inside cheek ; 
eee vibrisse present ....... ..... NSewuride & Petauristide. 
. Two tufts of superciliary vibrissze, a an anterior and a posterior. 
Aerus and its allies. 
b’. Only the normal anterior tuft of vibrissze present. 
c. Interramal tuft of vibrisse present ......... Sciwride (including Xerzs). 
e’ Interramal tuft of vibrissm absent ..................00.000...... Petauristide. 


The Har. 


In the arboreal Squirrels the ear exhibits no marked struc- 
tural variations. That of Sciwrws vulgaris may be described as 
typical. It is oval in shape and stands away from the head from 
a point in front about on a level with the supratvagus, and behind 
from a point below the antitragus. The anterior edge is folded 
over from the summit or tip of the ear, and the overfolded rim 
inereases in thickness down to the supratragus. Beneath the 
supratragus the edge runs obliquely downwards and backwards 
into the cavity of the ear, and ceases on its inner wall about 
midway between the supratragus and the orifice of the ear. The 
orifice les at the bottom of the cavity just below the level of the 
intertragal notch, which is defined in front by a small hemi- 
spherical tragus and behind by a large angular antitragus. The 
latter fits over the tragus when the ear is closed, and posteriorly 
is continuous with a large flap resulting from the overfolding of 
the posterior rim from a point a little above the line of the 
supratragus. The supratragus is a ridge with sharply defined 
inferior and poorly defined superior edge. It forms the upper 
border of the cavity of the ear, separating it from the feebly 
hollowed upper portion, which in height exceeds the height of 
the cavity. (Text-fig. 42, A.) 

Judging from dried skins and material preserved in spir it, the 
ears of the American species 5. hudsonicus, carolinensis, niger 
(rufiventer), griseus, saltuensis, hypopyrrhus and of several forms 
related to the last, resemble in all essentials those of S. vulgaris. 
So also do the ears of Hutamias, and of the Petauristide of the 
genera Glaucomys, Hoglaucomys and Petawrista, with the excep- 
tion that in those Flying Squirrels the pinna is still more 

extended above the supratragus and the anterior edge is not 
beac to the summit. (‘ext-fig. ae eR EE) 

The ears of the Tropical Asiatic and African arboreal Squirrels 
differ from those of Sciurus vulgaris and of the American species 
above referred to in being shorter as compared with their antero- 
posterior width, in being less tubular at the base, so that the 


1178 MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS 


auditory orifice opens approximately on a level with the inter- 
tragal notch; the tragus and antitragus, especially the latter, 
are, on the whole, less well developed, and the summit of the ear 
does not project so high above the level of the head, although 
in Ratuja and Hunambulus it is in this respect intermediate 
between the ears of Sciwrus vulgaris and its allies and those of 
Callosciurus and Lunisciwrus. 

In Ratufa (text-fig. 42, C) the ear rises from the head at a point 
about on a level with the anterior end of the supratragus in noes 
and below the upper end of the antitragal thickening behind; 1 
upper edge is rounded, not pointed, and the edge of the ae 
margin is overfolded ‘from the summit. Inferiorly the over- 


Text-figure 42 


A. Har of Sciurus vulgaris. D. Ear of Funambulus palnarum. 
B. ,, ,, Hutamias quadrivittatus. E. ,, ,, Hoglaucomys fimbriatus. 
C. ,, ,, Ratufa indica. KF. ,, ., Glaucomys volans. 


0., position of orifice. 


(These figures are intentisnally drawn of approximately the same size.) 


folded edge is continued downwards and backwards into the 
cavity of the ear, ceasing near its middle. Beneath its termi- 
nation there is a shallow pit concealed within the antitragal 
thickening, which is low and not produced. There is no tragus, 
and the auditory orifice is just exposed within the intertragal ; 
noteh. In Callosciurus prevosti, C. notatus, and Tomeutes 
vittatus the ear is less prominent than in Ratufa, and its points 
of attachment to the head in front and behind are somewhat 
higher. In C. prevosti also its upper portion is narrower’ and 
more pointed. In C. vittatws and C. notatus the posterior edge 


OF THE BEAVER AND OF SOME SQUIRRELS. Wakes) 


is shightly emarginate above. In these three species the tragus is 

small or absent. (Vext-fig. 45, A, B.) In Funambulus trisiriatus 
the ear does not appear to differ in any essential points either 
in structure or relative size from that of C. nofatus; but in 
ff. palmarum it is less hairy and relatively considerably larger 
and has the posterior edge markedly emarginate above*. (Text- 
fig. 42, D.) 

In the African Squirrels, Munisciurus leucostigna and £. cepapt, 
the ears are small as in Callosciurus, and do not appear to differ 
from them, although the eavity is perhaps more subdivided by 
hollows and low ridges; but in Heliosciurus punctatus the anti- 
tragus is better developed, being provided with a very definite 
process; the tragus is a curved, soft ridge, and the orifice of 
the ear, normally concealed by the antitragus, is narrowed and 
pointed above instead of being subcircular as in Funisciurus. 
(Text-fig. 43, C, D.) 

The ears of the Citellus, Cynomys, and Marmota are mostly 
smaller than in the typical Squirrels; but within the limits of 
the genus Citellus the variation in size is considerable. In C. 
(Otospermophilus) beecheyi they are indeed relatively larger than 
in Callosciurus or Funisciurus and project above the level of 
the head; but in other species, 7.e. some of those referred to 
the subgenus Jctidomys, they have been described by Allen as 
‘“‘ generally small, sometimes rudimentary.” Nevertheless, even 
in C. beecheyi they differ structurally from the ears of all the 
typical Squirrels examined. The cavity of the ear, bounded 
above by the supratragus, is to a considerable extent filled in 
front by a thickened development of the inferior continuation 
of the overturned anterior margin of the pinna; and this 
thickening is covered closely with longish hair covering the 
cavity and closing it when the ear is folded. There is no trace 
of tragus or antitragus, the lower rim of the cavity forming an 
evenly curved line. For the rest, the area above the supratragus 
is well developed and has a very distinctly emarginate posterior 
border; the tip is somewhat pointed, and the overfolding of the 
edge of the anterior rim does not quite reach the summit. In front 
the ear rises from the head at a point a little above the anterior 
end of the supratragus, and behind the posterior border fuses 
with the integument of the head just below the posterior end of 
that ridge. (Text-fig. 44, D.) 

In Marmota marmota the ear is a little more reduced in size 
than in C. beecheyi, and the area above the supratragus is wider 
in proportion to its height; the tip is pointed, but the posterior 
border is less markedly emarginate. The points of origin of the 
free portion in front and behind are approximately at the same 


* Judge from dried skins the ears of Tomeutes seem to be like those of 
Callosciurus; but in the remarkable Bornean genus Rheithrosciurus these organs 
differ from those of the other Oriental genera in their great length. In this 
respect they surpass the ears of S. vulgaris; but details of their structure could not 
be determined on the made-up skins in the Natural History Museum. 


1180 MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS 


levels as in C. beecheyi; the cavity is similarly blocked in front 
by the hairy thickening of the injutting portion of the inferior 
continuation of the overfolded portion of the anterior rim, and 
there is no trace of tragus, antitragus, or intertragal notch. 
(Text-fig. 44, H.) 

In Cynomys the ear is very much smaller than in J/armota 
owing to the reduction in size of the area above the supratragus. 
The summit is rounded, and has its edge overfolded in continua- 
tion of the overfolding of the anterior edge. The hairy thick- 
ening projecting backwards into the cavity of the ear above the 
orifice is well developed, there is no tragus or intertragal notch, 
and at most a trace of antitragal thickening. (Text-fig. 44, F.) 

In the Ground Squirrels of the genus Xerws and its allies, 
which are as terrestrial in their habits as the Sousliks and 


Text-figure 43. 


A. Har of Callosciurus prevosti. D. Kar of Heliosciurus punctatus. 
B. ,, 5, Tomeutes vittatus. HK. ,, ,, Atlantoxerus getulus. 
C. 4, 5, Funisciurus cepapi. BR. 4, ,, Geosciurus capensis. 


at., antitragal thickening. 


Marmots, the ears are greatly reduced in size, In some cases, 
indeed, e.g. in Geosciwrus capensis, equalling apparently in that 
respect those of some of the small-eared species of Citellus. But 
they are modified for terrestrial life on quite a different plan 
from that exhibited by the Sousliks and Marmots. 

in Huxerus erythropus, for instance, a comparatively large- 
eared form, in which the summit of the ear reaches about as 
high as the level of the head, the antero-superior and posterior 
margins meet approximately at a right angle. The anterior edge 
is overfolded, and is continued beneath the supratragus, where 
it bends backwards and downwards, ceasing just above the 
tragus, or sometimes running thence backwards and upwards as 


OF THE BEAVER AND OF SOME SQUIRRELS. 1181 


a rod-like ridge to the posterior end of the supratragus. The 
tragus is a hard, rounded eminence, and the intertragal notch is 
very deep. The antitragus is a thick bulging swelling set high 
above the tragus just beneath the posterior end of the supra- 
tragus, which fuses with it. The anterior edge rises from the 
head at a point about on a level with the anterior end of the 
supratragus and the posterior edge, which is nearly straight or 
shghtly sinuous, from a point about on a level with the lower 
portion of the antitragal prominence. When the ear is closed 
the upper portion folds down on the supratragus, the antitragus 
is folded on itself and pulled forwards over the tragus, and with 
the supratragus blocks the upper part of the cavity, the inter- 
tragal notch at the same time being closed by the juxtaposition 


Text-figure 44, 


Nea 
= 
S S 
— a 
= E- 
S = 
Eg Zz 
a ‘y 
AY GY, f ry) Qn 
Giri ont “Wwe Za 


D * Fi 
A, B, C. Ear of Euxerus erythropus, in three stages of folding. 


D. Ear of Citellus (Otospermophilus) beecheyi, from a dried skin. 
KE. ,, ,, Marmota marmota. 


KF. ,, ,, Cynomys ludovicianus. 


of its anterior and posterior walls. Thus the mechanism for 


closing the ear is quite different from that of the Marmots and 
Sousliks, where the supratragus is depressed against the hairy 
thickening invading the fore part of the ear-cavity, the thicken- 
ing itself being pressed down over the orifice. Judging from a 
dried skin, the ear of Xerws rutilus is relatively a little smaller 
than in Humerus erythropus owing to the reduction in size of the 
portion above the supratragus. (Text fig. 44, A--C.) 

In Aélantoxerus getulus the ear seems to be relatively a little 
smaller than in Z#. erythropus, and projects from the head less 


1182 MR. R. I, POCOCk ON THE EXTERNAL CHARACTERS 


prominently, but resembles it in other ways sufficiently closely 
to call for no detailed description. The most important difference 
is the absence of the tragal thickening in front of the inter- 
tragal notch, so that the orifice of the ear is exposed. Owing to 
the reduction in size of the upper free portion, the supratragus 
is nearer to the summit and the latter is somewhat more 
pointed. (Text-fig. 43, H.) 

The ear of Geosciurus capensis exhibits the final stage in 
degeneration in this group *; but it is clearly derivable from the 
type seen in Atlantoxerus by the reduction of the upper portion 
above the supratragus to a younded thickened rim incapable of 
being folded downwards and backwards to any appreciable 
extent on to the supratragus, which is represented at most by a 
short ridge running forwards and upwards from the antitragus. 
The latter, as in the other species, is a rounded thickened bulge, 
less well defined than in the northern species, set high up and 
overlapping the posterior portion of the cavity. The anterior 
rim is overfolded, and its inferior continuation into the cavity is 
represented at most by a small excrescence opposite the anti- 
tragus. There is no tragus, as in Adtlantoxerus, the orifice being 
exposed at the bottom of the deep intertragal notch. The ear 
is closed by the forward movement of the thicker posterior border 
of the cavity against the anterior, there being no folding of the 
antitragus or upper rim. (Text-fig. 43, F.) 

The structure of the ears in these Bristly Ground Squirrels 1s 
not opposed to the view of their affinity with the African species 
referred to Munsciwrus, but the differences supply no evidence 
of close affinity. 

In Castor the ear is relatively small, but is as free from the 
integument of the head as in the typical Sciuride. Its outer 
surface is flat, and the anterior edge is not overfolded; it is 
uniformly covered with hair except round the cavity, where it is 
naked. The cavity isa small, irregularly triangular space bounded 
above by the supratragus, which is overlapped in front by the 
anterior edge which runs downwards and backwards to the 
orifice lying at the bottom of the cavity and concealed by the 
lower edge of the cavity corresponding to the intertragal notch. 
The posterior border of the cavity forms a convex bulge probably 
representing the antitragus, but there is no trace of tragus. 
(Text-fig. 40, C.) 

The characters of the principal types of ears above described 
may be summarised as follows :— 

a. Har simple, its anterior edge above the supratragus not over- 

IRONGUEYAN so ee doe ab .. Castor. 

a’, Kar simple or complex, its anterior ‘edge ‘above the supratragus 

overtolded. 
6. Cavity ot the ear, when folded, mainly closed by hairy, 


thickened continuation of the overturned anterior rim beneath 
the supratragus, no tragus or antitracus ... Citellus, Cynomys, Marmota. 


* Described by W. L. Sai as eee ae ee Sone absent, the ear- 
opening being a narrow diagonal slit half an inch in length. 


OF THE BEAVER AND OF SOME SQUIRRELS. 1183 


’, Cavity of the ear, when folded, mainly closed by antitragus, 
the inferior continuation of the overfolded anterior border 
not forming a hairy thickening within the cavity. 
e. Antitragal thickening set high up near the middle of the 
posterior wall of the cavity ; pinna reduced. 
d. Margin of ear a thickened rim not standing away from the 
Wendt iy Nea see ne GreOsenuurus: 
. Margin of ear standing a aw way from head dorsally .  Xer us, Kuxerus, 
: Atlantowerus. 
e'. Antitragal thickening occupying normal position and form- 
ing the postero-inferior wall of the cavity; pinna standing 
well away from head dorsally .................. Typical Squirrels, Tamias, 
Petauristide. 


Although Castor is isolated at the head of this table, it can 
hardly be claimed that its ears differ more from those of typical 
Squirrels than do the ears of Geosciurws. Section c! is roughly 
divisible into two groups, exemplified respectively by Sciwrws and 
its near allies, Tamias and the Petauristide, which have long 
or comparatively long ears, aud by the tropical African and 
Asiatic genera Funisciurus, Heliosciurus, Callosciurus, and 
Ratufa, where the ears are shorter and less tubular at thie base. 
But these groups seem to intergrade. What the ears of Rhei- 
throsciurus are like apart from Jength has yet to be ascertained. 


The Feet. 


The feet of the typical arboreal Squirrels of the family 
Seiuride have many characters in common. In the fore foot 
the first digit is reduced to an apparently functionless excrescence 
springing from or imbedded in the side of the inner carpal pad, 
and carrying a nail. Of the remaining digits, the fourth is the 
longest, the difference between the fourth and the fifth or second 
being considerable. The fourth and the third are more closely 
united at the base than they are to the fifth or second respec- 
tively, and the middle line of the artiodactyle foot passes between 
them. The digital pads and claws are well developed. The 
plantar pad is always represented by three well-defined elements. 
The undersides of the digits and of the sole are typically naked 
—hairy in some northern forms in winter,—and there are two 
well- developed carpal pads, of which the inner is the larger of 
the two. The tuft of carpal vibrisse is always present and 
large. 

The digits of the hind foot resemble those of the fore foot 
tolerably “closely in structure and relative lengths, the fourth 
being the longest; but the first digit 1s well developed and 
resembles the others, except that it is “much the shortest of the 
series. The well-developed plantar pad consists of four lobes. 
The metatarsal area is for the most part naked, the heel alone 
generally being hairy, and is supplied with two pads, an inner, 
which is the larger, and an outer, which, in addition to being 
smaller, is set more forwards. Sometimes one or both of these 
pads is suppressed, in the case, for instance, of Squirrels, which 
have hairy feet in the winter, e.g. S. vulgaris. 


1184 MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS 


a. Feet of the Tropical Asiatic Squirrels. 


In Callosciurus prevosti the claws of the fore foot are short, 
with an abruptly curved, sharp point; the three main elements 
of the plantar pad are large and elongated, and small accessory 
tubercular pads are sometimes associated with them. The 
outer carpal pad is irregularly cordate, with a notch in its 
posterior border; it is much sherter than the inner carpal pad, 


Text-figure 45. 


7 


efi 


td 


A. Right fore foot of Callosciurus notatus, nat. size. 


BE Chas >» » » Lomeutes vittatus. 

GPs See ri nna cleans As 2 

D. 4, fore ,, ,, Callosciurus prevosti. 
13 ay) Jan oe A 5 


which extends forwards along the side of the foot, so that its 
anterior end overlaps on the pollical side the posterior end of 
the internal lobe of the plantar pad, and its pollical margin, 
evenly convex, conceals from view from the lower side the nail 
of the pollex, which is planted some distance behind the anterior 
end of the pad. (Text-fig. 45, D.) 


OF THE BEAVER AND OF SOME SQUIRRELS. 1185 


The hind foot has claws like those of the fore foot, and the 
four lobes of the plantar pad are similarly large and long, 
especially the internal; and both the internal and the external 
lobes are provided with a small supplementary lobe, making the 
entire plantar pad appear to be composed of six !obes, four large 
and two small. The metatarsal pads are also large. The internal 
starting close behind the corresponding lobe of the plantar pad 
and extending backwards to within a short distance of the hairy 
heel, occupies about one-third of the naked metatarsal area. 
The external metatarsal pad, about equal in area to the median 
lobe of the plantar pad, extends obliquely inwards and backwards 
from a point close behind the plantar pad, its posterior end over- 
lapping the anterior end of the internal metatarsal pad. (Text- 

So aed 

The feet of Callosciurus notatus and of Tomeutes vittatus are 
practically alike and agree in all essential respects with those of 
C. prevosti, and judging from dried skins Callosciurus jfinlaysoni, 
C. castaneoventris, and Tomeutes hippurus have feet of essentially 
the same type. (Text-fig. 45, A, B, C.) 

In Ratufa indica the fore foot has the pads exceedingly large. 
The three plantar pads are in contact with each other and with 
the two carpal pads. The latter meet posteriorly, and the five 
pads form a continuous lobate cushion surrounding a depressed 
area of soft but coarsely granular integument. The inner 
carpal, with which the digital pad of the pollex is indistinguish- 
ably fused, is as large as the three lobes of the plantar pad taken 
together. It forms a large projection on the inner side of the 
foot behind the second digit, and constitutes a grasping organ with 
the tips of the four digits, which close obliquely upon it. (Text- 
fig. 46, A.) In the hind foot development of the pads reaches its 
anes, as is the case in the fore foot. The four lobes of the 
plantar pad are wide, long, and in contact, and form a continuous 
cushion, with undulating distal margin extending from the inner 
to the outer edge of the foot. The outer metatarsal! pad, moreover, 
is elongated and in contact at its distal end with the outer lobe 
of the plantar pad; and the inner metatarsal pad is similarly 
confluent with the inner lobe of the plantar pad, the two together 
forming a long cushion about half the width of the metatarsal 
area, and reaching back to the hairy heel. The soft integument 
of the sole surrounded by the plantar and metatarsal pads and 
the outer half of the sole as far back as the proximal end of the 
inner metatarsal pad is granular, the granules being coarser 
distally behind the plantar pad than proximally towards the 
heel, which is covered with hair up to the proximal end of the 
inner metatarsal pad. (Text-fig. 46, B.) 

RF. bicolor and R. macrourus have feet like those of RF. indica, 
and the claws of the genus Ratufa are short, abruptly curved, 
and sharply pointed like those of Callosciurus. The feet of 
Ratufa, indeed, may almost be described as an exaggeration 
of the type seen in Callosciurus by the expansion and fusion 
of the pads. 


1186 MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS 


The feet of the Palm Squirrels (Funambulus) are relatively 
longer and thinner than those of Ratufa and Callosciurus, and 
have the pads less well developed. Otherwise they are tolerably 


Text-figure 46. 


} ] re a 
es ao 


NCR « 

Bie = 
Neu! V Sf: 
\ 4 u / 
2 Wy ee ana 
N i 
ANY iy s N 

\ A S \ 

Wie ZW 


A. Right fore foot of Ratufa indica. +}. 
B. ” hind 33 29 33 32 
C. » fore ,, ,, Funambulus palmarum, much enlarged. 


D. 32 hind 23 39 29 3? 


similar to those of Callosciurws, the inner, or pollical, carpal pad 
being similarly elongated, so that its distal extremity consider- 
ably overlaps the inner lobe of the plantar pad. In the hind 


OF THE BEAVER AND OF SOME SQUIRRELS. 1187 


foot the external and internal lobes of the plantar pad are 
often subdivided, but not always, and never to the same 
extent as in Callosciurus; the inner metatarsal pad is always 
long and rod-like, but the outer in all the specimens examined 
is small, spherical, and approximately on a level with the 


Text-figure 47. 


A. Right fore foot of Rheithrosciurus macrotis. 
B. 
C. 
D. 


(These figures drawn without measurements from dried skins.) 


22 hind 3° 39 EE 33 
» fore ,, ,, Protoxerus stangeri. 


” hind ,, ,, ” ” 


distal end of the inner without overlapping it proximally. 
The heel is less hairy than in Callosciwrus and Ratufa, being 
sometimes naked in the middle line, sometimes scantily hairy. 


(Text-fig. 46, C, D.) ; 


1188 MR. R. Ie POCOCK ON THE EXTERNAL CHARACTERS 


Judging from dried skins*, the feet of Rheithrosciurus are 
relatively ‘longer and thinner dine those of COallosciurus, and the 
claws are longer. In the fore foot the three plantar lobes are 
large, the medion being well defined from the laterals. Of the 
carpal pads, the outer is the smaller and narrow, and the inner 
has an evenly convex margin and is not elongated and produced 
distally so as to overlap the plantar lobe of the same side. In 
the hind foot the four lobes of the plantar pad are not so large 
or so closely packed as in Callosciurus, and the outer and inner 
are at most indistinctly subdivided; the metatarsal area is naked 
to the heel, the outer metatarsal is considerably shorter than in 
other Oriental genera examined, and the inner metatarsal is small 
and subspherical as in /unambulus and many other genera of 
Squirrels. (Text-fig. 47, A, B.) 

In the structure of the feet Rheithrosciurus is perhaps more 
like some of the American subgenera of Sciwrws than it is like 
other Oriental genera. 

The distinguishing characters of the feet of the Oriental 
Squirrels above described may be tabulated as follows :— 

a. Pads enormous, internal carpal very prominent on the 
pollical side; metatarsal pads confluent with plantar pad, 
the internal nearly half the width of the metatarsus ...... Ratufa. 
a’. Pads less developed, internal carpal less or not noticeably 
prominent on pollical side; metatarsals not confluent with 
plantar pad. 
b. Internal carpal long, narrowed distally, and overlapping 
plantar pad on inner side; internal metatarsal long. 
c. Feet shorter and stouter, plantar pad of hind foot 
larger, six-lobed, metatarsals thicker, external pro- 
duced proximally beyond distal end of internal. Callosciurus, Tomeutes. 
ce’. Feet longer and thinner, pads smaller; external meta- 
tarsal spherical set beyond distal end of narrowed 
internal metatarsal.. ... Lunambulus. 
b’ Internal carpal pad apically, rounded, “only 2 a little ‘longer 


than external and not reaching plantar pad ; internal 
metatarsal relatively short..............ccsecsscsecsseeesseereeee Ltheithroseiurus. 


b. Feet of the Tropical African Squirrels. 


The two African species, /unisciurus leucostigma and I, cepapi, 
have feet of a more generalised type than those of the Oriental 
species of Ratufa and Calloscvurus examined, the plantar, carpal, 
and metatarsal pads being less well developed. The plantar pad 
of the fore foot is three-lobed, of the hind foot four-lobed. The 
inner carpal pad is not so elongated distally, and the outer meta- 
tarsal] is comparatively small, subcircular, or elliptical, and is set 
on a level with the anterior end of the long rod-like inner meta- 
tarsal or a little in front of it. Except that the feet are of 
stouter build, they differ but little from those of Hunambulus 
palmarum, although in the latter the inner carpal pad is produced 
distally nearly as much as in Callosciurus. (Text-fig. 48, C-F.) 


* It must be remembered, however, that the proportions of the feet vary with 
the preservation. [n fresh specimens they appear to be plumper and shorter than 
those contracted by alcoho!, and the latter are, as a rule, less shrunk than the feet 
on dried skins. 


OF THE BEAVER AND OF SOME SQUIRRELS 1189 


Judging from dried skins, the feet of F. isabella, the type of 
the genus Munisciurus, resemble those of &. leucostigma and 
cepapt. 

Protoxerus stangeri also has feet differing in no important 


Text-figure 48. 


A. Right fore foot of Heliosciurus punctatus, nat, size. 
B 


” hind) 53 » ” ” ” 
©) “fore! 4; 5, Munisevwirus cepapi; Te is 
D. >} ) hind ass be) bh) 99 39 32 
KE. ,, fore ,, ,, Funisciurus leucostigma, ;, 5, 
BK. 39 hind 3) 3 32 be) 39 3” 


point from those of Funisciwrus, although the outer metatarsal 
pad is much longer, being at least half as long as the inner. 
(Vext-fig. 47, C, D.) 

Proc. Zoou. Soc.—1922, No. LX XIX. Y) 


1190 MR. R, I. POCOCK ON THE EXTERNAL CHARACTERS 


In Heliosciurus punctatus the feet are relatively larger than 
in Funisciurus leucostigma, the plantar lobes and carpal pads of 
the fore foot are smaller, more angular, and separated ; in the 
hind foot the plantar lobes are also smaller and more spaced, 
the inner metatarsal is much shorter, being oval and well- 
defined distally but abbreviated and ill-defined proximally ; the 
inner metatarsal is like it but smaller *. (Text-fig. 48, A, B.) 

The species belonging to the three African genera above 
discussed may be distinguished as follows by their feet :— 


a. Inner metatarsal pad very short, lobes of plantar pads and 
carpal pads relatively small and separated ............5........... Hl. punctatus. 


a’. Inner metatarsal pad a long, stout ridge; lobes of plantar pads 
and carpal pads larger and closer together. 
a?. Outer metatarsal pad elongate, more than half the leneth of 
Pheer eee Aiea vot ce eunraanuaeee ts RuN ale cra \pMNNGl LECLERC Cape 
62, Outer metatarsal pad small, oval or spherical ...... FE’. leucostigma, cepapi. 


In a general sense the hind feet of Munisciwrus do not differ 
in any very important respect from those of FHunambulus; but 
all the African species examined differ from all the Oriental 
species in not having the same elongation of the inner carpal 
pad. In the length of the outer metatarsal P. stangeri approaches 
Ratufa and Callosciurus. 

Funisciurus leucostigma and cepapi, on the one hand, and Proto- 
werus stangeri, on the other, were regarded by Forsyth Major as 
representing respectively two subgenera of Xerus. By their 
feet, however, they are typical arboreal Squirrels, differing in 
no essential particulars, so far as those organs are concerned, 
from such tropical American species as Sciwrus (Hchinosciurus) 
hypopyrrhus. 


c. Feet of the European and American Squirrels. 


In the tropical American Squirrels, referred to a large number 
of subgenera of Sciurws, the soles of the feet are naked as in the 
tropical Asiatic and African forms, the heel being generally 
hairy although sometimes naked down the middle line, as in 
an example of Scvwrus saltuensis bonde that died early in October. 
The lobes of the plantar pads are well developed, the two carpal 
pads are subequal in size, the inner not projecting appreciably 
beyond the outer as it does in the tropical Asiatic forms. The 
two metatarsal pads are retained, the inner being thick, elongated, 
and curved, the outer smaller, ovate, and situated in front of the 
anterior end of the inner. The claws are longer than in the 
tropical African and Asiatic forms, The feet of S. (Echino- 
seiurus) hyvopyrrhus resemble those described above, and their 
soles are naked in mid-winter. (Text-fig. 49, 1.) 

The feet of the Squirrels of the northern parts of America are, 
generally speaking, similar to those of the warmer latitudes, with 
this difference, that the soles undergo a seasonal change in the 


* For a tabulation of the genera of African Sciuride, based mainly on skulls and 
teeth, see Thomas’s paper (Ann. & Mag. Nat. Hist. (8) iii. pp. 467-475, 1909). 


OF THE BEAVER AND OF SOME SQUIRRELS. 1191 


matter of hair growth, accompanied in some cases by the suppres- 
sion of one or both of the metatarsal pads. 

In the North American Grey Squirrel GS. (Weosciurus) caro- 
linensis) the outer metatarsal pad is sometimes absent. When 


Text-figure 49. 


ee 


KER 


LEER 


A. Right hind foot of Sciurus carolinensis, with external metatarsal pad 
absent, in summer pelage (from dried skin). 

B. Right fore foot of the same, in winter pelage. 

C. ,, hind foot of the same, in winter pelage, the external metatarsal 
retained. 

D. Right hind foot of S. hypopyrrhus, or closely related species, in mid- 
winter pelage. 


retained it is a hemispherical tubercle set far forwards in front 
of the inner, which is irregularly ovate and shorter than in 


79% 


1192 MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS 


the tropical species. An example examined in September 
had the sole of the hind foot naked except the heel, which was 
hairy, whereas one that died in the middle of December had the 
sole covered with hair as far down as the outer metatarsal pad. 


Text-figure 50. 


A. Right fore foot of Sciurus vulgaris, mid-winter. 


IB} oii SND DHTIYOL! as y 3 » ” 
Oe RORD 5 SLAP ROH COROUARS op Sp 
IDsi) ays arin 


39.39 33 22 


(C, D from dried skin.) 


The fore foot in the two specimens was quite naked beneath 
at both dates. Again, in a skin of a specimen of the North 
American Fox Squirrel S. (Parasciurus) wiger rufiventer, which 
died at the beginning of December, all trace of the outer meta- 


OF THE BEAVER AND OF SOME SQUIRRELS. es 


tarsal’ pad has disappeared, although the inner is somewhat 
larger and more elongated than in S. carolinensis, and the whole 
of the under side of the foot, including the digits, is covered 
thickly with hair, apart from this pad, the well-developed four- 
lobed plantar pad, and the digital pads. The latter are pointed 
at their proximal ends. The fore foot is similarly hairy, there 
being a distinct patch on the median area in front of the carpal 
pads and defined by the three elongated contiguous lobes of the 
plantar pad. I am not aware whether the soles of the feet m 
this species are naked in the summer or not; but analogy justifies 
the belief in this being so. (Text-figs. 49, A-C; 50, C, D.) 

Tn Sciurus vulgaris * and its subspecies the lower sides of the 
feet in summer are as naked as in tropical species, except that 
perhaps a larger area of the heel is hairy. The inner metatarsal 
pad is comparatively short and set some distance behind the 
plantar pad ; the outer is quite small and some distance in front 
of the anterior end of the inner. But in winter the soles and 
lower sides of the digits are overgrown with hair which is more 
abundant on the hind than the front foot. In the hind foot it 
obliterates the metatarsal pads and passes between the lobes 
of the plantar pads. In both feet at this season the digital pads 
are reduced to long, narrow, posteriorly-poimted lobes. (Text- 
fig. 50, A, B.) 

In the North American Red Squirrel, S. (Tanviasciurus) 
hudsonicus, | can find no trace of either of the metatarsal pads. 
According to C. B. Cory? the soles of the feet are “furred in 
the winter.” From this statement I should infer that they are 
naked in the summer, were it not that a skin in the Zoological 
Society’s collection in summer pelage, with untufted ears and 
black lateral stripe, has the metatarsus covered with hair almost 
down to the plantar pads, whereas the area round the plantar 
pads and the under sides of the digits are naked. The fore foot 
also is as naked as in tropical species of Squirrels. On the 
other hand, in the skin of a specimen that died on January 7th 
in winter pelage, with tufted ears and no lateral stripe, both 
fore and hind feet are covered with hairs beneath, as in the 
specimen of the Fox Squirrel described, only the digital, plantar, 
and carpal pads being naked. But whether the whole of the 
metatarsal area in the American Red Squirrel is mesially naked 
back to the heel in summer as in the European Squirrel, 
S. vulgaris, I do not know. (Text-fig. 51, A, B.) 


d. Feet of the Chipping Squirrels (Tamias). 


The feet of the Chipmunk, Hutanuias quadrivittatus, do not 
differ materially from those of the typical Squirrels, except that 
the third and fourth digits are very nearly equal, the fourth at 

* The feet, in summer and winter pelage, of the British representative 
(S. leucourus) of the species are figured and described by Barrett-Hamilton and 


Hinton (‘A History of British Mammals,’ pt. xxi. p. 699, pl. xxxvii.). 
+ ‘The Mammals of Illinois and Wisconsin,’ 1912, p. 128. 


1194 MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS 


most exceeding the third very slightly. Nevertheless, the tip 
of the fourth extends beyond that of the second both in the fore 
and the hind foot. The feet are narrow and long, and the lobes 
of the plantar pads are not in contact, though well developed. 
The inner lobe of the carpal pad has a rounded distal border 


Text-figure 51. 


Ay. 


a 
yy 
N \ NN \i } 


S 


= 


S 
Ss 
——— - right i ae «Ces 
SEZ fod \ Nee = 
a 5 é 
a 
= wwe 
Ai : 
f ; 


= 


ae 
—- 


= 


EZ 


A. Right fore foot of S. hudsonicus, in summer pelage (from dried skin). 
B. ,, hind ,, ,, same specimen. 
Cc. ,, fore ,, ,, Hutamias quadrivittatus. 
ID agen) LONE Sy 5g 8 i 
(A, B a little enlarged; C, D much enlarged.) 


and is not produced beyond the margin of the smaller outer pad, 
and the small pollex projects a little in front of it, which is not 
the case in the typical Squirrels, where the pollex rises from 
the outer surface of the inner carpal. The under sides of the 
digits and the area round the plantar lobes are naked in both 
fore and hind foot; but in the hind foot the whole of the meta- 


OF THE BEAVER AND OF SOME SQUIRRELS. 1195 


tarsal area is covered with hair down to the plantar pad, and 
there is no trace of metatarsal pads as in S. (Zamiasciurus) 
hudsonicus. (Text-fig. 51, C, D.) 

This description of the feet of Hutamias, based upon a single 
individual, must not be taken as applying to all the species of 
Tamias. In some dried skins of this genus in the Natural 
‘History Museum, the digits appear to resemble those of Citedlus 
rather than of Sciwrws and its allies in relative lengths ; and the 
metatarsal area is not so extensively haired as in the specimen 
of Hutamias described. 


e. Feet of the Brisily Ground Squirrels (Xerus, etc.). 


The feet of the African Ground Squirrels, Aélantoxerus getulus, 
Xerus rutilus, Huxerus erythropus, and Geosciurus capensis differ 
from those of the arboreal species hitherto described in having 
the third digit the longest and the fourth only a little longer 
than the second, so that the feet approach the perissodactyle 
type; the pollex is not suppressed to quite the same extent, but 
stands away as a distinct excrescence from the antero-external 
edge of the adjoining inner carpal pad, and this is not appre- 
ciably produced distally beyond the outer carpal; the metatarsal 
area is naked to the heel, and its pads are either absent or poorly 
developed. 

In Atlantoxerus getulus the feet are long and slender and the 
digits more nearly approach those of the arboreal Squirrels in 
length, the third only slightly exceeding the fourth, which, in 
the fore foot, is decidedly longer than the second. The lobes of 
the plantar pads are large, well developed, the hallucal lobe on the 
hind foot being nearly as large as the others-individually; the 
carpal pads also are comparatively large, and the metatarsal pads 
are at all events sometimes distinguishable, although small and 
oval. (Text-fig. 52, A, B.) 

In Luexerus erythropus the fore foot is broader and shorter, 
with respect both to the digits and sole, than in Atlantoxerus ; 
the plantar and carpal pads are relatively smaller and the claws 
are longer. The pollex is small but subcylindrical with a small 
pad remote from the carpal. In the hind foot, too, the claws are 
fonger than in Atlantoxerus, the lobes of the plantar pad are 
considerably smaller, especially the outer and inner, the latter 
being a small subcireular tubercle at the base of the hallux, only 
about one-fourth the size of the median lobe; and there is no 
trace of metatarsal pads*. (Text-fig. 53, A, B.) 

In Geosciurus capensis the feet are more robust and more 
fossorial in character than in £. erythropus. In the fore foot the 
three lobes of the plantar pad are well developed but shorter 


* X. rutilus is known to me only from a dried skin. On this the plantar lobes 
appear to be somewhat larger relatively than in H. erythropus; but there 1s no 
trace of metatarsal pads, and the hallucal lobe of the plantar pad is small and 
isolated. 


1196 MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS 


than in Atlantoxerus and in contact, and the pad of the pollex 
projeets as a large lobe jutting out from the side of the 
adjoining carpal pad. In the hind foot the three main lobes of 
the plantar pad are also well developed, larger than in Huwerus 
erythropus but shorter than in Atlantoxerus getulus; but 


Text-figure 52. 


SY Lm) 
Yy 


SN | 
\ NN 
Ny 


SA 


A. Right fore foot of Atlantoxerus getulus, much enlarged. 
Bo.) whine msamies 
Cc. ,, fore ,, ,, Marmota marmota. X %. 
ID Nae. hind sae asatn es 


the hallucal lobe is quite small, subcircular and isolated as in 
Euaerus erythropus; and as in the latter there is no trace of 
metatarsal pads, and the claws of all the digits are long and 
strong. 


OF THE BEAVER AND OF SOME SQUIRRELS. ILS), 


The feet of the Xerine Squirrels above described differ as 
follows :— ; 
a. Feet thinner; inner lobe of plantar pad of hind foot large, the 
metatarsal pads retained but poorly defined.............. .. Atlantoxerus. 


a’. Feet stouter; inner lobe of plantar pad of hind foot very 
small, metatarsal pads absent. 


b. Plantar pads smaller, pollex a relatively small subeylindrical 


OOS ccd ead andtoe noosa haa BoB ono DEaia Hon SED OBA ceo osoeee dosbue | OKA UAT 
6’. Plantar pads larger, pollex a conspicuous subcircular ex- 
GHATECOMNAD sco ucoacoocooass eee Geosciurus. 


The feet of Xerus, typified by ruéilws, fali under heading a’ of 
this table. 


Text-figure 53. 


. Right fore foot of Euxerus erythropus. X +. 


3 


» fore ,, ,. Geosciurus capensis. 
29) hind 99° £29 


A 
Be ae hinde shin As 
C 
D 


2 +>} 


1198 MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS 


J. Feet of the Sousliks and Marmots. 


A second group of Ground Squirrels represented by the 
Sousliks (Citellus) and the Marmots (Cynomys, Marmota) have 
feet like those of Xerus and its allies in that the third digit is 
the longest, and the second and fourth digits not so unequal, as in | 
the arboreal Squirrels, and the claws longer, stouter, and jess 
curved. The three genera in question, however, have very 
different feet. 

The fore foot of Citellus (Otospermophilus) beecheyi*, judging 
from a dried skin, is shorter and broader than that of the 
typical squirrels, and is naked below. The fourth digit is 
decidedly but not much longer than the second. The three lobes 
of the plantar pad and the two carpal pads are well developed, and 
the pollex is represented by a conspicuous nail attached to the 
outer side of the inner carpal pad in front. In the hind foot the 
second, third, and fourth digits are nearly equal, the third only 
slightly surpassing the others, which are subequal; the fifth 
reaches beyond the middle of the fourth; the plantar pad is 
represented by four well-developed contiguous lobes, the inner 
and outer of which are set rather far back, the greater part of 
each lying behind the other two. There is no trace of metatarsal 
pads, the whole area being covered with hair frem the heel to the 
plantar pad. (Text-fig. 54, A, C.) 

Judging from Miller’s description of the European species 
Citellus citellus, the feet of that species agree tolerably closely 
with those of C. beecheyi. He says, however, that in the fore 
foot the second and fourth digits are subequal, and that the fifth 
extends only a little beyond the base of the fourth. This does 
not apply to C. beecheyi. Similarly, he states that in the hind 
foot of C. citellus the fifth digit extends just beyond the base of 
the fourth, and the hallux not quite to the base of the second 
digit. This is not true of C. beecheyt. But the apparent differ- 
ences between the two species in the particulars mentioned are 
interesting, if true, because in the relative length of the digits the 
typical Kuropean species seems to approach the genus Cynomys 7. 

In Cynomys the fore foot is long and robust, almost as large 
indeed as the hind foot. The digits are thick and the claws long. 
The second digit is a little longer than the fourth, and the fifth 
reaches a little beyond the base of the fourth. The inner and 
outer lobes of the plantar pad are small, and together hardly 
equal the size of the wide median lobe. The outer carpal pad is 
large and the inner is rather prominent internally, and gives 
support to the claw of the pollex, which is as long as that of the 
second digit. The entire sole is naked. In the hind foot the 
second, third, and fourth digits are subequal, the third only very 
slightly surpassing the others, as in Citellus. The fifth is set 
almost as far back as the hallux and is only a little longer, 


* A Californian species. 
+ For a possible explanation of these discrepancies, see note, p. 1208. 


OF THE BEAVER AND OF SOME SQUIRRELS. ILS) 


surpassing by a little the base of the fourth. The claws on these 
digits are shorter than on those of the fore foot. There are four 
well-developed separated lobes constituting the plantar pad, and 
their arrangement is quadrate, the outer and inner lying behind 


Text-figure 54. 


A. Right fore foot of Citellus (Otospermophilus) beecheyi, from 
dried skin. 

Bisons » 99 95 Cynomys ludovicianus. Xt. 

Cc. ,, hind ,, ,, Citellus (Otospermophilus) beecheyi, from 
dried skin. 

IB baa. 9» 99 «9, Cynomys ludovicianus. X i. 


the others—an arrangement foreshadowed in Ci%tellus. There 
are no metatarsal pads, the sole being naked except for a band 
of hair passing from the inner side of the heel forwards along the 


1200 MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS 


middle of the metatarsal surface nearly or quite up to the level 
of the hallucal lobe. (Text-fig. 54, B, D.) 

The feet of Marmota marmota are relatively more robust and 
somewhat shorter in the digits than those of Cynomys. In the 
fore foot the three lobes of the plantar pad are relatively larger, 
occupying practically the entire width of the foot. The carpal 
pads also are larger, but the inner does not project laterally in 
the same way and has no trace of the pollex. Similarly, in the 
hind foot the plantar lobes are massive and do not exhibit that 
quadrate arrangement seen in Cynomys and Citellus. Behind 
the outer lobe of the plantar pad there isa hemispherical external 
metatarsal, and behind the inner plantar a short internal meta- 
tarsal with ill-defined posterior border. (Text-fig. 52, C, D.) 

A peculiarity of the feet of this species of dar mota is the 
complete absence of the pollex. Muller noted this in twenty 
skins, Tullberg’s figure showed the same thing, and I have not 
sueceeded in finding a trace of this digit in some half-a-dozen 
examples I have seen. On the other hand, Miller records the 
presence of the pollex with a flattened nail in JZ. bobak, the other 
European species. The presence of a rudimentary thumb 1s 
cited by Blanford in his generic diagnosis of MZ. himalayana, 
hodgsoni, and caudata, and American writers testify to its 
retention in Jf. monax and other N. American species. 

By the structure of the feet the Marmotine genera may be 
arranged as follows :— 

a. Feet very powerful; lobes of plantar pads and carpal pads 
large, metatarsal pads retained. 
(A, LoD laxe RIO NASTHAC on! covsboseaooceocouasvancsun andbreoboecoussnoanccens, LULMPIOOIRE. 
6’. Pollex retained.. Broce poadonwaseeeaudeodnannoued noe ARF RIOHO/ DS" WON. 


a'. Feet at most Podenarele vomentale iobas of plantar pads 
and carpal pads smaller, no metatarsal pads. 


. Feet comparatively stout, inner ora pad peiechine, a 


long claw on pollex............... ee Cynomys: 
ce’. Feet comparatively dlanélaes inner compel ead normal; 
Oller. HMI MAB oo caopecocossesdocobaHoncsenabesbodounvecusocen | COBHURIB. 


g. Leet of the Flying Squirrels (Petauristide). 

Apart from the cartilaginous rod supporting the patagium and 
jointed to the wrist just behind the outer carpal pad, the fore 
feet of the Petauristide closely resemble those of the typical 
arboreal squirrels. In Petawrista albiventer (magnificus)+ the 
claws are long and strongly curved. The digits also are long and 
markedly unequal in length, the fourth being considerably longer 
than the third and fifth, which are subequal and surpass the 
second. On the specimen examined there is practically no 
external trace of pollex and none of the nail. The three lobes 
of the plantar pad and the carpal pads are large and striate all 
over. (Text-fig. 55, A.) 


* Type, I. monax. 
+ For the specific names here used for Petawrista (olim Pteromys), see Wroughton, 
Journ. Bombay Nat. Hist. Soc. xx. p. 1012, 1911. 


OF THE BEAVER AND OF SOME SQUIRRELS. 1201 


On the hind foot the claws are similar, but the digits are 
shorter, the second, third, fourth, and fifth appearing to be nearly 


Text-figure 55. 


A. Right fore foot of Petaurista albiventer, from dried skin. 
B. ” hind 999 Dy) a 3) » ry) 0 
C. 4, fore ,, ,, Hoglaucomys fimbriatus, showing the - 
patagial rod. X i. 
ee hindes. &) sanie: 


o 


equal, although in reality the fourth is a little longer than the 
third, which is about equal to the fifth and slightly longer than 


1202 MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS 


the second. ‘The four lobes of the plantar pad are large, the 
outer and inner being larger than the others. The inner meta- 
tarsal pad is well developed, and extends as in Ratufa along the 
inner side of the metatarsal area, being in contact with the 
plantar pad distally. A large area of the heel is covered with 
hair, and on the outer side of the foot the hair overgrows the 
sole up to a line continuous with the proximal extremity of the 


Text-figure 56. 


(0 a ZS 
GC 


Y} 
SE 


\ 


A. Right fore foot of Glaucomys volans, summer pelage. 


B 39 hind 392 39 39 29 ” 3) 

N J . . 
Coes nhOne uueninnre % stramineus, winter pelage, 
D 93 hind bb) 29 re) yg 29 39 


large, long, outer plantar lobe, so that the area of naked skin 
on the admedian side of the metatarsal pad is narrow. (Text- 
fig. 55, B.) 

Blanford described the hind foot of this species, under the 
name Pteromys magnificus, as similar to that of P. inornatus, 
saying of the latter: ‘‘ The large metatarsal pad joins the inner 
toe pad. ... No supplementary pad.” This agrees with my | 


OF THE BEAVER AND OF SOME SQUIRRELS. 1203 


observations; but from the size and length of the outer. lobe of 
the plantar pad I suspect the outer metatarsal pad, referred to 
by Blanford as the supplementary pad, is retained and fused with 
it. According to Blanford the feet of P. ywnnanensis are like 
those of albiventer but larger, whereas in P. philippensis (oral) 
there is a large piriform inner and a small outer metatarsal pad, 
and P. caniceps seems to resemble philippensis so far as the inner 
pad is concerned, although Blanford makes no mention of the 
presence of an outer pad. 

In Eoglaucomys fimbriaius the fore foot does not differ 
materially from that of Petawrista albiventer except that there is 
a distinct pollical excrescence with a nail, as in the typical 
Sciuride, and the plantar and carpal pads carry coarser ridges 
not distributed all over them but limited to the highest portion 
of the pads. In some cases there is a little isolated tubercle on 
the outer side of the external lobe; in other cases this is confluent 
with the lobe, increasing its width. The hind foot is also tolerably 
similar to that of P. albiventer; the sculpturing of the pads, 
however, differs as in the fore foot and the inner metatarsal pad, 
about twice as long as wide, is set at the proximal end of the 
naked metatarsal area remote from the hallucal lobe of the 
plantar pad. In this species also, as noticed by Blanford, there 
is a small circular outer metatarsal pad behind and close to the 
outer lobe of the plantar pad. If this were enlarged and con- 
fluent with the plantar pad, the outer lobe of the latter would 
resemble tolerably closely that of Petawrista albiventer. (Text- 
fig. 55, C, D.) 

None of the other Indian species assigned by Blanford to 
Sciuropterus* retains the outer metatarsal pad. S. (Hylopetes) 
alboniger has the inner metatarsal pad oval; in S. (Petinomys) 
fuscocapillus it is very elongate; in Selomys pearsonit it is 
oval. 

In the North American species, Glaucomys volans, the fore foot 
structurally resembles tolerably closely that of the Himalayan 
Hoglaucomys fimbriatus. In a spirit specimen of unrecorded date 
the underside of the digits and of the entire sole of the foot is 
naked, whereas in a specimen of G. stramineus, which died in 
December, these surfaces are covered with hair, the pads, which 
appear to be smaller, alone being naked. The hind foot, how- 
ever, differs in two correlated characters from that of Hoglaucomys 
jimbriatus: there is no trace of metatarsal pads, and the metatarsal 
area is covered thickly with hair from the heel almost to the 
plantar pad. In the example of G. volans, with the naked sole 
to the fore foot, the undersides of the digits of the hind foot and 
the area round the plantar pad are likewise naked, whereas in 
the specimen of G. stramineus, with hairy soled fore foot, the 
lower sides of the digits and the area round the plantar pad are 


* For the generic and subgeneric names here used for these species, see Thomas, 
Ann. & Mag. Nat. Hist. (8) 1. pp. 1-8, 1908. 


1204 MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS 


hairy, like the rest of the sole, and the claws are almost concealed. 
It is probable that the difference between these two closely-related 
forms in the matter of the hairiness of the soles of the feet is a 
seasonal difference comparable to that of Sciwrus vulgaris and of 
S. (Tamiasciurus) hudsonicus. (Text-fig. 56.) 

Judging from Miller’s description, the feet of S. russicus, the 
type of the genus Sciwropterus, closely resemble those of the 
North American species. It might have been taken indeed 
from the specimen referred to above as S. (@.) stramineus, she 
soles of the fore foot being thinly haired between the pads, 
whereas those of the hind foot are covered with a “‘ dense growth 


Text-figure 57. 


/ 

A. Claw of Funisciurus leucostigna; B. Funiseiurus cepapi; C. Ratufa 
indica; D. Hutamias quadrivittatus; E. Sciwrus lindernicus ; F. Sciurus 
vulgaris; G. Petaurista albiventer; HH. Citellus beecheyi; I. Xerus 
rutilus. 


of long woolly hairs especially conspicuous along the inner * side, 
and adding much to apparent width of foot.” Miller had only 
three specimens of this species for examination, and in the 
absence of information as to the dates of their capture, it is not 
possible to guess whether there is a seasonal change in the 
hairiness of the soles of the feet or not. 

To draw attention to the facts here recorded in the hope that 
they may be amplified and, if necessary, corrected by other 


* For “inner” here read “ outer,’ the error below referred to under Castor of con- 
fusion between the inner and outer sides of the feet being repeated. The inner side 
of the foot as generally understood is the side on which the hallux, or first digit, 
rises, whereas the fringe of hairs on the hind foot of the Petauristide is always on 
the side of the fifth digit, z.e., the outer. 


OF THE BEAVER AND OF SOME SQUIRRELS. 1205 


workers with more and better material, I subjoin the following 
table :— 


a. Metatarsal area hairy, without pads .....................  Sciwropterus, Glaucomys. 


a’. Metatarsal area partially naked with the inner pad 
at least retained. 
6. Inner metatarsal pad confluent with plantar pad ; 
pollex practically suppressed, without nail ...... Petaurista albiventer. 


b’. Inner metatarsal pad distinct from plantar pad; 
pollex as in typical Sciuride. 


ce. Outer metatarsal pad retained ........................ Hoglaucomys fimbriatus. 
ce’, Outer metatarsal pad lost. 
d. Inner metatarsal pad elongate... ..........c00.ee0e Petinomys fuscocapillus. 
ad’, Inner metatarsal pad oval 2.00. 02.2... ces esecee eee Belomys pearsoni, Hylo- 


petes alboniger. 


h. Feet of the Beaver (Castor). 


Apart from minor differences, the fore foot of Castor fiber 
differs from that of the Seiuride and Petauristide in two 
important points, the presence of a well-developed pollex and the 
indistinctness of the lobes of the plantar pad. The digits are 
short, with roughened but not well-defined digital pads; the 
third is the longest, a little exceeding the fourth, the second and 
fifth being shorter, and the first, or pollex, the shortest of all. 
All are armed with long, wide, blunt, slightly curved claws, that 
of the pollex being a little narrower, sharper, and more curved 
than the rest. The plantar area is granularly reticulated, and 
the lobes of the plantar pads are obscurely represented by mobile 
folds of thickened granular skin, hardly alike in any two 
specimens or even on the two feet of an individual. Only some- 
times are the thickenings arranged so as to indicate the three 
main plantar lobes present in all the Sciuride. Behind the 
plantar area there are two well-developed elongated carpal pads. 
The inner at the base of the pollex is the smaller, and is hard, 
smooth, and grooved except at its distal end. The outer pad is 
much longer and wider, and extends considerably further up the 
wrist proximally. The comparatively wide space between these 

ads is occupied by thickened granular skin, which on the 
admedian side of the inner carpal pad, and a little in front of it, 
forms sometimes a lobate thickening, which may represent the 
inner or pollical element of the plantar pad, which should be 
present with a well-developed pollex, but which is absent in all 
the Sciuride, where the pollex is little more than a vestigial 
remnant. There isarim of naked skin behind these carpal pads ; 
but on the inner side of the wrist the hairsare directed obliquely 
towards the middle line, in a crest overlapping the posterior 
portions of the pads. There are no carpal vibrisse, which are 
present in all Sciuride. (‘Text-fig. 58, A.) 

The hind foot greatly surpasses the fore foot in size, far more 
so than in any member of the Sciuride. This is mainly due to 


Proc. Zoou. Soc.—1922, No. LXXX. 80 


1206 MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS 


the immense length of the digits, especially of the first and fifth, 
which, relatively to the other digits, are much longer than in the 
Squirrels, and give a wide span to the foot. The third and 
fourth digits are subequal, the fifth is not very much shorter 
than the fourth and is about equal to the second, and the 
elongated hallux extends beyond the middle of the second. They 
are all connected by wide webbing, which extends to the apex of 


Text-figure 58. 


A. Right fore foot of Castor fiber. X %. 

Baie i auc gE aie » ” 

C. Apex of second and third digits of the same, showing the sup- 
plementary claw (s) beneath the main claw on the second. 


the digital pads or nearly so. These pads are represented by 
areas of granular skin, and on the first and second digits they 
form movable lobes, whereas on the third, fourth, and fifth they 
are less prominent and not separately movable. <A well-known 
peculiarity is the presence of a secondary “ false claw” beneath 
the normal claw of the second digit, constituting with 16 a fur- 


OF THE BEAVER AND OF SOME SQUIRRELS. 1207 


comb, and the claws of the first and second are sharper and more 
curved than those of the others. The greater part of the lower 
side of the digits and of the webs is na ked, but the proximal ends 
of the digits and of the webs is thickly granular like the rest of 
the sole back to the heel. The greater part of the metatarsal 
area is covered with a big pad extending on the outer side almost 
as far as the base of the fifth digit, and on the inner side about 
halfway between the heel and the hallux, where its edge turns 
obliquely across the sole to meet the outer edge in a widely- 
rounded curve. This pad is widest across its middle, gradually 
and slightly narrowed behind and greatly narrowed in front. 
Between it and the base of the hallux, and separated from it by 
a narrower or wider furrow, lies a second much smaller pad, the 
outer edge of which forms the margin of the metatarsal area in 
front of the inner portion of the larger pad. The distal third of 
the smaller pad is at the base of the hallux. In front of these 
pads the skin is folded and grooved, but shows no definite trace 
of plantar pads. It is difficult to homologize the two pads above 
described with the pads on the sole of the hind foot in the 
Sciuride. Tentatively | adopt the hypothesis that the smaller 
inner pad represents the inner or hallucal element of the plantar 
pad, which is often large in the Sciuride, and that the larger pad 
represents the two normal metatarsal pads fused together and 
blended with the rest of the integument. The disappearance of 
the main elements of the plantar pads in Castor is in keeping 
with their partial or complete suppression in other fully web- 
footed aquatic forms like the Coypu and some others; and the 
retention of the pads on the metatarsus is to be explained by 
the beaver’s habit of walking on its heels when carrying objects 
on land. Whether the above suggested explanation be true or 
not, it is evident that in the Sipuchare of its fore and hind 
feet the Beaver differs greatly from the Squirrels. (Text- 
fig. 58, B, C.) 

My description of the feet was taken from the Huropean 
species, Castor jiber. The American species has similar 
feet. 

Miller’s description of the feet of the Huropean species differs 
materially from mine. The fore foot, according to him, has five 
palmar tubercles, “the three at bases of median digits large, 
ill-defined, that at base of thumb not so large as the others but 
better dsiimed: its outer surface tending to ns@onne smooth and 
horny, the posterior pad largest, covering about one-third of 
entire palmar surface.” It seems ‘clear from this that the inner 
of the three above-mentioned tubercles at the base of the pollex 
is the horny grooved pad I identify as the inner carpal pad. If 
Miller’s interpretation is correct, this is the pollical or inner 
element of the plantar pad. His “ posterior pad” must be the 
large pad I describe as the outer carpal pad. 

In his deseription of the hind foot it is evident Miller has made 


80% 


1208 MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS. 


the mistake of confusing the first with the fifth digits, for he 
speaks of the double claw as being on the fourth, whereas it is 
well known to be on the second digit *. This makes his deserip- 
tion of the proportionate lengths of the digits (and the sizes of 
their claws) different from mine. Of the pads he says :——“* Three 
elongate ill-defined pads at base of median digits, a larger and 
better-defined tubercle behind base of hallux [= fifth digit]; 
entire sole between hallucal tubercle and heel thickened and 
pad-like.’ 
The characters of the principal types of feet in the Sciuromorpha 

may be tabulated as follows :— 
a. Fore foot with well-developed pollex, lobes of plantar pads 

‘yeduced to folds of flexible integument, carpal pads very large, 

no carpal vibrisse; hind feet with all the digits very long, 

‘widely separabie and united by webs up to the digital pads ; 


lobes of plantar pad mostly obsolete, ae ra of meta- 
tarsal area covered by one large JORG ssn neooeoesboobonopsodendsascosee  Cuuorude- 
a’. Fore foot with pollex reduced to a mere excrescence or absent 
lobes of plantar pads well developed, and as prominent : 
carpal pads, carpal vibrissee always present; hind foot pl 
shorter digits, free from wels; plantar pad well developed; 
metatarsus typically with two pads, a longer inner and a 
smaller outer, both sometimes suppressed 
b. A carpal rod Suppor Ue patagium, digits as in arboreal 
SLGHIPDD ICE U A NSA A NWBNA Rail ana Gian Gagniysadcoapdeneseaguadmoneaa |) Ce CAA ZPUSU ln, 
b’. No carpal rod. 


ce. Fourth digit the ey much longer than the 


second.. faa vesserseeses, Sciuride (arloreal). 
ec’. Third sie ite loneest: “font aot Hal longer than 
RASOMG! | shossa500.000 900 ond 996000 nad ogneND Hea poN DDB DDN HOnUOD NCaCmradaa (erasure). 


The Anus and External Genitalia. 


In the. Sciuride and Petauristide the anus and external 
genitalia are normally separated. In the male the prepuce fT is 
prominent and situated some distance in advance of the anus, 
being mostly long and pendulous and even retroverted, when at 
rest, in the arboreal Squirrels, the Bristly-Ground Squirrels, and 
the Flying Squirrels, but shorter in the Marmots. Between it 
and the anus there is in the Squirrels a very large scrotum in 
the breeding season, but I have not been nai to verify this in 
the case of the Marmots. In the female the vulva is a semi- 
circular or pointed process, situated a little below the anus, and 
the clitoris is concealed within the labia. The anus has the two 
normal glands in the Squirrels, whereas in the Marmots there is 
an additional median gland. (Text-fig. 59, A, B, C.) 

The anus and external genitalia in Castoride differ profoundly 
from those of the Sciuride and Petauristide. 

Externally the male and female are alike, owing to the 
orifices of the alimentary canal and of the genito-urinary organs 


* This error no doubt arose from the fashion now in vogue of twisting the hind 
feet in made-up skins so that the hallux lies on the outer side of the foot so placed. 
+ The structure of the penis and baculum I must leave for another paper. 


OF THE BEAVER AND OF SOME SQUIRRELS. 1209 


discharging into a common chamber or cloaca, the aperture of 
which, closed by a sphincter muscle, is placed at the base of the 
thick hairy portion of the tail. In the male, moreover, there 
is no scrotum, the testes being inguinal in position, that is to say, 
beneath the integument covering the pubic portion of the pelvis. 


(Text-fig. 59, D.) 


Text-figure 59. 


(aa) VOM RY 


PAonngaaAniy, 
sex ROOTES 
Thaitokon: 
ta) 


LONE A y 

ES ae 

4 EAT yy 
C 1D), 


A. Anal and genital region of male of Callosciwrus prevosti, from below. 
B. The same of Geosciurus capensis, from the side. 

C. 4, 5 > temale Cynomys ludovicianus. 

D. 4, 5) +>», male or female Castor fiber. 

a, anus; sc, scrotum: p, prepuce; v, vulva; eZ, orifice of cloaca in Castor. 


When the cloaca of the male is opened, four orifices, two 
median and one on each side, are seen to open into it. The 
posterior and larger median orifice is the anus, the anterior and 
smaller median orifice is the common aperture of the penis and 
the castorium glands. The paired orifices, one on each side and 


1210 MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS 


remote from the anus, are the apertures of. the anal glands. 
Hach of the apertures of the anal glands leads into a deep 
narrow pouch, at the bottom of which are three papillee, each 
tipped with a long hair. This pouch can be evaginated, so that: 
the gland comes to the surface, and the strong-smelling secretion 


Text-figure 60. 


A. Diagrammatic dissection of the anus and genitalia of male Castor canadensis. 
3. The same of the female. 
C. The cloaca of the male. opened. 

Miles, a Nee heap se naall einer 
H. One of the anal glands, showing the tip with its papille and sete within 

the pouch. 

t, testis; s, pubic suture; p, penis; c, castorium glands with orifices on 
each side of preputial orifice in male and above vaginal orifice in female ; 
v, vaginal orifice; ag, anal gland; 0, orifice of anal gland; a, anus; si, 
slit through which secretion of castorium glands is discharged into cloaca 


in female. 


is discharged at the tips of the papille. Normally the penis is 
completely retracted, but it is capable of being protruded through 
the orifice above mentioned into the cloaca and thence to the 
exterior. Through this orifice the secretion of the castorium 


OF THE BEAVER AND OF SOME SQUIRRELS. 1211 


glands is also discharged. These are represented by a pair of 
immense sacs, with ‘puckered walls, one. lying on each side. 
They are filled with fluid; and the aperture of each sac opens 
close to the median aperture of the penis, the three apertures 
lying close together just within the common orifice above 
mentioned. (Text-fig. 60, A, C, E.) 

The arrangement is very similar in the female, in which the 
anal and castorium glands appear to be as well developed as in 
the male. ‘The chief difference is that the orifice of the vagina 
opens separately into the cloaca. Just in front of it there is a 
transverse slit in the wail of the cloaca, and when this is opened 
the apertures of the castorium glands may be seen on each side 
of it, just in front of that of the vagina. (Text-fig. 60, B, D.) 

To sum up, the Castoride differ from the Sciuride, so far as 
the organs just described are concerned, in the following parti- 
culars :——(1) In both sexes the anus and genitalia open into a 
common chamber or cloaca with a single external orifice. 
(2) The anal glands are of great size and open apart from the 
anus. (3) There is a pair of large castorium glands opening 
close to the generative orifice. (4) In the male there is no 
scrotum; the penis is straight and is protruded close to the 
anus. 

In view of these well-known and profound differences, it is 
singular that in the current text-books the Castoride are 
separated from the Sciuride merely by such comparatively 
trivial differences as are presented by the skull and teeth, the 
webbed hind feet and broad tail being added as of secondary 
value, and no mention whatever being made of the genitalia 
and anus. The difference between the Beavers and Squirrels 
in the anus and external genitalia may be summarized as 
follows * :— 


a. Sexes externally alike owing to the anus and genito-urinary orifices 

opening into a cloaca with a single aperture at the root of the 

tail, no hairy prepuce or vulva, the concealed orifice of the 

penis just in front of the anus; testes inguinal; a voluminous 

“castorium” gland opening close to the orifice of the penis and 

vulva; anal glands very large, with long tubular orifice......... Castoride. 
b. Sexes externally unlike, the anus and external genitalia not 

opening into a cloaca; female with hairy vulva close beneath 

anus; male with long perineal region, typically with large 

scrotum for testes in breeding season; penis long, opening by 

prominent hairy prepuce on posterior part of abdomen; no 

“castorium ” gland; anal glands small, with small orifice. 

Sciuride, Petauristide. 


From the above-given analyses of the external characters of 
the Castoride, Sciuride, and Petauristide it will be seen that, so 
far as the organs considered are concerned, there is not a particle 
of evidence of kinship between the Castoride and the other two 
families. 


* JT have not in this paper dealt with the great differences between the Beavers 
and Squirrels in the tail, because they are well known. 


TD) EXTERNAL CHARACTERS OF THE BEAVER. 


If the structural differences between the Flying Squirrels and 
the less specialised true Squirrels entitle these groups to be 
regarded as distinct families, the differences between the Beavers 
and the Squirrels must be granted much higher rank. The 
Sciuride and Petauristide may be associated as the Sciuroidea, 
and the Beavers kept apart as the Castoroidea. The Petauristidee 
are a compact group represented by several genera and subgenera; 
but, so far as my limited observations on the external characters 
are concerned, I have found no reasons for thinking them 
susceptible of grouping under headings of subfamily rank. The 
genera of Sciuride are much more varied; but their affinities are 
at present too little understood to warrant the subdivision of the 
family. 


ON THE CHONDROCRANIUM OF THE LARVAL HERRING. 1213 


53. On the Morphology of the Chondrocranium of the Larval 
Herring (Clupea harengus). By F. R. WELLS. 
[Reeeived October 18, 1922: Read November 21, 1922.] 


(Text-figures 1—11.) 


ContTENTS. 

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1. [IyTRODUCTION. 


This research was started at the suggestion of Prof. MacBride, 
the work being carried out in his laboratory in the Imperial 
College of Science, our attention being drawn to the necessity of 
studying the development ot the chondrocranium, in particular, 
of the more’ primitive Teleostei by work being done by 
Mr. Norman on the Hel, whieh, like the Herring, has no upper 
jaw in the early larval stages. 

The paper refers to the larvee of the Clupeoid, Clupea harengus : 
all observations and reconstructions were made from these only, 
although the internal morphology of the Sprat and Pilchard was 
found to be indistinguishably similar. The paper describes only 
the chondrocranium. For convenience, the larve have been 
divided into stages as follows :—5 mm., 10mm., 20 mm., 30 mm., 
and metamorphosing stages (40 mm. and 50 mm.). 

Previous work on the skulls of Teleostomi includes the fol- 
lowing :—Salmon (Parker, 1872; Stohr, 1882; Gaupp, 1903; 
Boker, 1913), Trout (Winslow, 1897), Gasterostews (Swinnerton, 
1920), Cyclopterus (Uhlmann, 1921), Syngnathus (MeMurrich, 
1884), Amiuwrus (Kindred, 1919), Selachian (Parker, 1879); 
Sewertzoff, 1899), Chondrostei (Parker, 1885; Sewertzoff, 1902), 
Lepidosteus (Parker, 1883; Veit, 1911), and a recent paper on 
Amia (Pehrson, 1922), in which is pointed out the need for the 
study of the morphogenetic development of the cranium of the 


* Communicated by Prof. E. W. MacBrips, F.R.S. 


1214 MR. F, R. WELLS ON THE MORPHOLOGY OF THE 


Teleostei, more especially of the Clupeide and Siluride. No 
work has been done on the cranial development of the 
former”. 

Material was obtained from Plymouth, and was fixed on 
the trawler as it was caught; the larger stages were caught 
in Dublin Bay in 1921; material was also obtained from 
Lowestoft. 

I wish to thank Prof. MacBride, Dr. Hogben, and Mr. H. 
Graham Cannon for valuable suggestions and assistance with 
both the work and the manuscript. 


2. MErHop. 


The specimens were fixed in Bouin’s fluid, in Chromic-Urea- 
Bouin, or in Corrosive Sublimate. The first was found to be most 
successful. A number of series of transverse sections were cut, at 
84; they were then stained in Ehrlich Hematoxylin, in Borax 
Carmine and Picronigrosine, in Mallory’s triple stain, or in 
Thionin and Orange G. The last was found to be the most 
successful. 

The usual wax model method of reconstruction was not used. 
The reconstructions were made direct as dorsal or lateral 
views on specially squared paper. The exact method is as 
follows :— 

An eyepiece micrometer, divided into squares, was used. The 
virtual size of these squares, with the various powers used, was 
measured by means of a blood-corpuscle measuring slide divided 
into hundredths of a blood millimetre. With the microscope 
used (Leitz, eyepiece 3, objective 3, draw-tube 140) one square 
was found to measure 15 mm. The reconstructions wen made 
at a magnification of 120, The paper was therefore “squared ” 
with lines 18 mm. apart vertically (or longitudinally) and 1 mm. 
apart (= 120 x 843,) horizontally (or transversely). The direction 
of the lines on the micrometer was then set according to the 
direction from which the view was taken, and the reconstruction 
mapped out directly on to the squared paper. This method is 
really a simplification of the method, which was tried at first, of 
drawing each, or alternate, section on squared paper and then 
reconstructing dorsal or lateral views by measuring these 
drawings. The new method merely eliminates the intermediate 
process. It was found to be very considerably quicker, and 
certainly as accurate as the wax model method. The same 
assumption had to be made as is made in the model method— 
that is, the assumption that one of the cartilages was straight. 
In the reconstructions made the ethmoid plate was taken for 


* Tt is interesting to note that none of the above-mentioned types have larvee, as 
do the Clupeoids, Flat-fish, &c. Theterm “larva” is here taken to refer to the free- 
swimming stage preceding metamorphosis as opposed to the newly-hatched yolk- 
sac bearing embryo. Lebour and others refer to this free-living stage as the post- 
larval stage. 


CHONDROCRANIUM OF THE LARVAL HERRING. 1215 


this centre; corrections to allow for this can quite easily be 
made from longitudinal sections of a similar larva. To make 
comparison more easy and more accurate, all reconstructions 
were made to the same scale, using the same micrometer and 
microscope. 


3. EXTERNAL CHARACTERS. 


The specimens examined varied in length from 5 to over 
50 mm.; the 10 mm. stage was taken in the greatest detail, and 
it corresponds to an age of less than a month. The larve were 
of the Herring, distinguished from those of the Pilchard and Sprat, 
as 1s pointed out by Fage (1920), by the number of myotomes, 
having 56 to 58, 51 to 52, and 46 to 48 respectively. 

The tail is characteristically short, the anus being very close to 
the end of the body. In a specimen 10°5 mm. long we have the 
following measurements :—head 1:5 mm., trunk 7°5 mm., tail 
15 mm. (Lebour, 1921). The head is nearly as broad as it is 
long (including the eyes), the lower jaw protruding forwards well 
beyond the upper. 

The only visible fins at first are the pectoral and caudal, the 
pelvic fin not appearing until a length of over 20 mm. is reached, 
and the dorsal fin is not complete until the animal is 28 to 
29 mm. long (Lebour, 1921). 

None of the specimens of 10 mm. upwards examined had the 
yolk-sac still present, although it may persist until 12 mm. 

These and other points have been illustrated in text-fig. 1. 

Metamorphosis takes place at about 40 mm. 


4, DISCUSSION. 


This research, as stated in the Introduction, was carried out to 
fill a very noticeable gap in the work done on the development 
of the Teleostean chondrocranium. ‘The paper does not pretend 
to be a complete account, and it has very little detail in it; the 
results obtained were so unexpected and abnormal that I have 
made no attempt to explain or to theorise. Time was unfortu- 
nately too short for me to make it a comparative account ; it is, 
in fact, merely a simple description of the state of affairs 
found. 

The first striking point is this: there is in the early stages no 
upper jaw at all—it does not appear until the larva attains a 
length of about 13 mm. 

Another point is that at the earliest stage examined (i. e. 
5 mm.) the hyomandibular mass of cartilage is fused both to the 
cranium and to Meckel’s cartilage. It breaks away from the first 
at about 15 mm., and the second at 25 mm. The upper jaw 
grows back, and fuses with it secondarily. The quadrate is 
formed from this mass by a constriction forming in it. 


1216 MR. F. R. WELLS ON THE MORPHOLOGY OF THE 


It is interesting that the larve showed points of interest not 
only in the chondrocranium. The development of the pituitary 


Text-figure 1. 


go. 


Meckel's Ft 
Cavtlacg \ 


Larval Clupea harengus. 15 mm. Stage. 


body does not tally exactly with the previous accounts. This 
matter was briefly examined, and will be published separately. 


CHONDROCRANIUM OF THE LARVAL HERRING. ey 


5. Tut CHONDROCRANIUM. 
a. The 5 mm. Stage. 


This description is based on the study of transverse sections of 
a specimen nearly 5-5 mm. long. Sections, for confirmation and 
comparison, of slightly longer specimens were also cut (text- 
fig. 2). 

In front, the chondrocranium is a thin plate of cartilage, in the 
form of a deep, nearly straight-sided trough, about half as deep 


Text-figure 2. 


Eth ie Ye Ss 
ee HM 
a 5mm 


Eth WE: 


HM 
MK 


C (Omm 


Lateral views of early stages of Larval Herring. 120. From reconstructions. 


as it is broad. A little further back the sides turn up, and 
separate from the median part forming large upturned pro- 
cesses, which lie on each side between the brain and the eyes. 
The median part becomes a somewhat narrower nearly flat plate, 
which soon divides into two narrow processes enclosing a large 
foramen in which the pituitary lies. These enclosing processes 
are the Trabecuize Cranii. 


1218 MR. F. R. WELLS ON THE MORPHOLOGY OF THE 


At the posterior end of this foramen the processes widen 
laterally and turn up sharply. The median part forms a tube 
enclosing the notochord. 

From near the top of the lateral upturned process there 
grows down a cartilage, laterally thin, which soon elongates to 
form a large flat plate, slating out at the top. This is the hyo- 
mandibular. Close below its fusion with the cranium there is 
a foramen, of the hyoid branch of the VIIth (facial) nerve. 

At the bottom of this plate and slightly in front of its fusion 
with the cranium there is a narrow elongated slit. From the 
posterior end of this hyomandibular there grows downwards and 
forwards a thin cartilage. This runs along the lower side of the 
hyomandibular, and then forwards and upwards. The halves (it 
is, of course, like the hyomandibular, paired) meet some way in 
front of the anterior end of the ethmoid plate; they are not 
fused, but articulate. 

(The naming of the cartilages is based on study of later 
stages.) 

There is at this stage no trace of an upper jaw (i.e. the 
palatal, pterygoid, and quadrate cartilages), the only roof to the 
mouth being the ethmoid plate. 

A specimen of 6°5 mm. was examined, and showed only a few 
slight differences; the large upturned processes are distinctly 
smaller, and the narrow elongated slit in the hyomandibular mass 
is closing up. The fusion of this mass with Meckel’s cartilage is 
stili placed as in the younger stage. 

In a 75 mm, specimen it was found that the large upturned 
processes had diminished considerably in size, being very little 
larger, comparatively, than in the 10 mm. stage. The narrow 
elongated slit in the hyomandibular has entirely closed up, and 
the fusion of the posterior process from Meckel’s cartilage to the 
hyomandibular has moved forwards, being now situated directly 
below the anterior end of the fusion of the latter to the cranium. 

Text-fig. 2C showsa reconstruction of a 7-8 mm.specimen ; the 
upturned processes have disappeared entirely, and the fusion of 
Meckel with the hyomandibular has moved right forwards to the 
anterior end of the latter, and its fusion with the cranium is as 
before. 


b. The 10 mm. Stage. 


This description of the chondrocranium is based on the study 
of transverse sections in all nine series, on whole mounts of 
specimens of Clupea harengus varying in length from 9 to 
11 mm., and on reconstructions made from a 10 mm. specimen 
(text-figs. 3 & 4). 

In front the ethmoid plate is still thin, but it has flattened 
out considerably, being at first dorsally convex and, further 
back, concave. In front this ethmoid plate is very broad, 
measuring nearly -5 mm. across; it runs back for nearly one- 
third of the length of the head before dividing to give off the 


free 
* o> 


CHONDROCRANIUM OF THE LARVAL HERRING. IY) 


Trabecule Cranii. The pituitary foramen is now about -45 mm. 
long and half as wide. Just anterior to the posterior end of this 
foramen these processes (the Trabecule Cranii) widen laterally 
as before, and turn up sharply to send forwards the long, slender 
post-orbital processes. (These might also be called the supra- 
orbital processes, but the former name is chosen as it emphasises 
the fact that they grow forwards from behind, and that they lie 
between the eyes and the brain and not over the eyes.) Small 
protrusions, corresponding to the large upturned processes of the 
5mm, stage, turn back from the ethmoid plate as if to meet 
these post-orbital processes. The orbit is thus, at this stage, 
incomplete, as it is in the 10mm. Syngnathus, whereas in the 
8mm. Amiurus and the second week Salmo it is complete. 

In the flat plate between the pituitary foramen and the post- 
orbital processes are the foramina of the VIlth nerve. 

The anterior end of the notochord lies directly behind the 
pituitary foramen, and it is enclosed in cartilage for the first 


Text-figure 3. 


Lateral view of chondrocranium. 10 mm. Stage. X 5d. 


‘Dmm. of its length. On each side of this are the auditory 
capsules, enclosed at their posterior end by cartilage. The brain 
and nerve-cord are in no place completely roofed over, the 
nearest approach to this being two thin, high processes lying 
laterally to and nearly level with the top of the nerve-cord : 
these arise from the cartilage enclosing the notochord, near the 
posterior of the head. In them is the foramen of the Xth nerve 
(vagus). The 10mm. Syngnathus, 8 mm. Amiurus, and second 
week Salmo, all have the posterior end, at least, of the brain 
roofed over. 

At this stage there is still no trace of an upper jaw. Meckel’s 
cartilage protrudes conspicuously beyond the ethmoid plate; the 
two halves are narrow but deep. The jaw widens rapidly until 
it is nearly as wide as the ethmoid plate, and then narrows slow] 
until it meets the hyomandibular. The two are still definitely 
fused, Meckel’s cartilage extending as a long forward growth 
from the anterior end of the hyomandibular. 


1220 MR. F. R. WELLS ON THE MORPHOLOGY OF THE 
The hyomandibular is a great mass of cartilage, sloping back- 
wards and outwards at the top and measuring about °5 mm. 


long, -15 mm. deep, and nearly -1 mm. broad. It meets the 
cranium just behind the post-orbital processes and between the 


Text-figure 4. 


MK. 


< 
Bia” 


, Eth, q 


Dorsal view of chondrocranium. 10mm. Stage. X 70. 


foramina of the VIIth nerve and the pituitary. The hyomandi- 
bular touches the cranium for about half its length, being 


articulated in front and fused behind. 


CHONDROCRANIUM OF THE LARVAL HERRING. OPM 


The foramen of the hyoid branch of the VIIth (facial) nerve is 
situated as in the 5mm. stage. 

At the posterior end of the hyomandibular, at the bottom, 
there articulates with it the ceratohyal, by means of a conspicu- 
ous knob, probably the appearing stylohyoid, as in Syngnathus. 


Text-figure 5. 


\ 


ray 1 Ne 
ye ei 
py ‘ ies. 
d a 4 
: i 
i : ! 
: / 
AN , / 
iy D 
Ey ' 
é { 
' 


w. 4 \\ 


jig 


Dorsal view of branchial arches. 10 mm. Stage. X 60. 


The ceratohyal runs forward as far as the anterior end of the 
ethmoid plate ; arising from the anterior end of the ceratohyal 
are the branchial arches. 

The accompanying figure is of the branchial arches of a larva 
at this stage; the figure is self-explanatory. 


c. The 20 mm. Stage. 


This description is based on reconstruction of a 20 mm. 
specimen of Clupea harengus (text-fig. 6) 

At its extreme anterior end the ethmoid plate is, as in the 
10 mm. stage, a flat plate; a little further back a median dorsal 
ridge appears. This ridge runs back as far as the two lateral 
protrusions, which are considerably developed. 

The ethmoid plate now becomes comparatively thinner; the 
pituitary foramen is also comparatively smaller. The post- 
orbital processes are thicker, and run forwards beyond the lateral 


Proc. Zoou. Soc. —1922, No. LX XXI, 81 


1222 MR. F. R. WELLS ON THE MORPHOLOGY OF THE 


protrusions and come down close to the median ridge. About 
half-way along these post-orbital processes a bridge is formed 
between them, which gives off two median thin processes, one 
forwards and one backwards. This bridge roofs over part of the 
anterior end of the brain (epiphysial bar). 
Arising from the anterior end of the upturned sides of the 
ethmoid plate, from which the post-orbital processes arise, are 
paired backward growing pieces of cartilage; these meet processes 
growing up from the sides of the posterior end of the ethmoid 
plate, and enclose the anterior end of the auditory capsules. 
From the top of these a bridge grows to cover part of the 
posterior end of the brain. The posterior end of the auditory 
capsules is enclosed as in the 10 mm. stage. The cartilage 
enclosing the notochord is as before. ‘ 
Articulated with the sides of the anterior end of the ethmoid 
plate are small paired cartilages—the early beginnings of the upper 
jaws (the palatal part of it). This appears first at about 13 or 


Text-figure 6. 


Lateral view of chondrocranium. 20mm. Stage. X 40. 


14mm. Theyrun back as faras the anterior end of the pituitary 
foramen. They have no connection whatever with either the 
hyomandibular or with Meckel’s cartilage. This is very similar 
to the 10 mm. Amiwrus, in which the palatal arises separately 
close to the ethmoid plate and the pterygo-quadrate grows 
forwards from the hyomandibular to meet it. At this stage, 
however, there is no trace of any forward growth from the 
hyomandibular. The palatal arises separately similarly in 
Syngnathus (8 mm.). 

The hyomandibular is articulated with the cranium, and in no 
place is it fused with it; it 1s a considerably larger mass of 
cartilage, comparatively, than in the 10 mm. stage. The part 
articulating with the ceratohyal has grown back and away from 
the main mass, forming even more conspicuously a stylohyoid 
cartilage. Anterior to the base of this process a long pointed 
piece of cartilage grows forwards. 

The hyomandibular is considerably elongated, and there is a 


CHONDROCRANIUM OF THE LARVAL HERRING. 1223 


slight restriction where it meets Meckel’s cartilage. At the 
lower part of this restriction the two cartilages are divided to 
indicate a coming division between them. 

Meckel’s cartilage is very much as in the smaller 10 mm. stage, 
wide at the base and tapering to a thin, horizontally flattened 
cartilage at its anterior end. 

The two foramina of the VIIth, facial, nerve (7.e. in the 
hyomandibular and in the ethmoid plate) are unchanged. 


d. Zhe 30 mm. Stage. 


The following description is based on the study of transverse 
sections of a 30 mm. specimen of Clupea harengus and on a 
reconstruction of it (text-fig. 7). 

The median ridge in this stage runs to the extreme anterior 
end of the ethmoid plate, the top buiging laterally considerably ; 
near the posterior end of this ridge it divides to enclose the 
anterior end of the brain. Just below the top bulge there is the 
foramen of the Ist nerve. 


Text-figure 7. 


Lateral view of chondrocranium. 380mm. Stage. X 20. 


The ethmoid plate runs back from this at first as in the 
previous stage, being nearly flat. It then tapers to a long 
median point, on each side of the base of which are slight knobs, 
the remains of the Trabecule Cranii, which are now absent. A 
ventral view of this is shown in text-fig. 8. 

From the cartilage enclosing the anterior end of the brain the 
paired post-orbital processes extend back as before. The bridge 
over the brain and between these (the epiphysial bar) is 
unchanged. From the posterior end of this bar there runs a 
process, along the top of the brain, to meet the cover to the 
posterior end of the brain. 

Theauditory capsules are now completely enclosed by cartilage. 
The processes which in the 20 mm. stage grew back from close 
behind the post-orbital precesses are now broken, to give long 
sloping slits in the upturned sides of the ethmoid plate. Part of 
the posterior end of the hind brain and the anterior end of the 
nerve-chord are entirely roofed over by cartilage. 


81* 


1224 MR. F. R. WELLS ON THE MORPHOLOGY OF THE 


The palatal cartilage is comparatively larger in cross-section, 
and from its posterior end there runs a thin strip of cartilage 
to meet the hyomandibular mass at its anterior end. It is 
important to notice that this fusion is entirely secondary, and, 
as pointed out above, the upper jaw grows back from the 
front. 

The hyomandibular mass of cartilage has grown considerably, 
being now nearly half as long as the whole chondrocranium. 
The articulation with the cranium is as before, the stylohyoid 
and the forward growing piece of cartilage, just in front of it, are 
even more conspicuous. Just in front of the base of this forward 
growing part the hyomandibular mass forms a narrow neck, and 


Text-figure 8. 


Ro. 


od VA 


Ventral views of ethmoid plate in the 10 mm. and 30 mm. Stages. X 80. 


spreads out again further forward. From its anterior end on top 
it sends forward the thin strip to meet the uppet jaw, and 
immediately below this there is a small protrusion which articu- 
lates with Meckel’s cartilage. At the articulation there is a 
large upward growth from Meckel’s cartilage, otherwise it is 
unchanged. 

The two foramina of the VIIth nerve (facial) are unchanged. 


e. The 40mm. Stage. 


This account is based on the study of transverse sections of a 
40mm. specimen of the Herring and on a reconstruction of it 
(text-fig. 9). 

The whole chondrocranium has considerably increased in height 
without a very marked change in length. 

The anterior end of the ethmoid plate and the median ridge 
show little change, except in the formation of the anterior eye- 
muscle canal; the rostrum is more pointed. The epiphysial bar 
is unchanged; the post-orbital processes have become compara- 
tively more massive, and from the ventral side of their anterior 


CHONDROCRANIUM OF THE LARVAL HERRING. 1225 


end a pair of small backward-growing protrusions have appeared. 
The median process running along the top of the brain is very 
much thinner, especially near the posterior end. 

The long sloping shts in the upturned sides of the posterior 
part of the ethmoid plate—mentioned as present in the 30mm. 
stage—are now closed at the upper and anterior end, leaving a 
large foramen. The roof over the posterior end of the brain is 
now complete. 

The anterior end of the palato-pterygoid is practically 
unchanged, the posterior end being somewhat enlarged to form 
an upturned knob. There is a slight restriction where it meets 
the hyomandibular mass. 


Text-figure 9. 


Lateral view of chondrocranium. 40 mm. Stage. X 20. 


The articulation of the hyomandibular with the cranium is no 
longer complete, being broken in the middle. The hyomandibular 
mass has changed very considerably—changes which are even more 
marked in the 50mm. stage,—and it is perhaps best to describe 
them in the next section. 

Considerable ossification is commencing. 


f. The 50 mm. Stage. 


[See text-figure 10.] 

The nasal septum appears at this stage at the base of the 
rostrum ; apart from this the anterior part of the ethmoid plate 
remains unchanged. The post-orbital processes are again more 
massive, and the process running medianly from the epiphysial 
bar to the posterior end of the cranium is, as it originally was, 
incomplete. 

The retro-articular process on Meckel’s cartilage is now con- 
spicuously developed. 

No other differences have appeared except in those cartilages 
concerned in the suspension of the jaws. 

Text-fig. 11 shows the suspensory apparatus of Clupea 
harengus and of Amia calva (from Torsten Pehrson, 1922) for 


1226 MR. F. R. WELLS ON THE MORPHOLOGY OF THE 


the purpose of naming the various cartilages present. The 
explanation offered in the figure does not explain the slight 
constriction in the middle of the quadrate. 

It remains, however, without doubt that the quadrate, the 
symplectic, stylohyoid, hyomandibular, and Meckel’s cartilages, 
all arise from the same single mass of cartilage, which mass was 
originally fused to the cranial part of the skull, whereas the 
palato-pterygoid cartilage arises entirely separated. 

Text-fig. 11 shows the development of the suspensory arrange- 
ment of the jaws compared with that of Amia calva, and also 


Text-figure 10. 


Lateral view of chondrocranium. 50mm. Stage. X 17. 


stages in Salmo and Syngnathus. ‘This figure tends to fill in the 
gap left by the absence of any comparison in the text. 

To assist further comparison, the following table, showing the 
rate of growth of the larva, is given :— 


de lsh fo) dubavere nun Naas nesean dehnee 4mm. 
DRWeOKS Un em a are ya teenie 4 AO) seamen, 
amromulie ecrecer ener eter 18 mm. 
Dombi inane earn, 35 mm. 
RAMMOTTNENG saancussgasasodoudoss 43 mm. 
AL TINOVONWOS So coeces MEU Io etd 55 im. 

approx. 


(H. A. Meyer.) 


6. SUMMARY. 


1. The hyomandibular, quadrate, symplectic, and stylohyoid are, 
in the 5 mm. stage, all one mass of cartilage. 

2. This mass is fused, at this stage, to the cranial cartilage and 
to Meckel’s cartilage. 

3. The palato-pterygoid forms at about 13 mm., and grows back 
to meet the quadreate. 


CHONDROCRANIUM OF THE LARVAL HERRING. 224 


Text-figure 11. 


CLUPEA Amia. 


3Omm #0 72 rem 


End wstele 


‘Ss YNGNATHUS 
20 mm »ZO 


eS 
HM. HyoMandibular Qu Quasvale n Qe p 


SHR Shylo Hyos Mk Meckels Carblago 


Sym 


Shu 


Sym Symplecic. PP Palate Ptenaod 
Simm. 


7 In 79 Bi 
Mg “sy 
Diagram comparing development of suspensory apparatus of jaws 


of Clupea, Amia, Salmo, and Syngnathus. Reduced one-half 
from dimensions printed on block. 


1228 MR. F. R, WELLS ON THE MORPHOLOGY OF THE 


A, 


5. 


» oo 


Until the 20 mm. stage is reached, the brain is in no place 
roofed over. 

The Trabecule Cranii disappear between 20mm. and 30 mm., 
leaving a median process, which does not join the posterior 
part of the skull. 


. The quadrate appears by a constriction in the hyomandibular 


mass. 

The symplectic appears as a thin pointed process from beneath 
this mass. 

Articulations of this mass with the cranium and with Meckel’s 
cartilage appear at 10 mm. and 26 mm. respectively. 

Considerable ossification is taking place at 30 mm. 


7. List or ABREVIATIONS USED IN THE [LLUSTRATIONS. 


IBA eaten inens Branchial arches. 

Oils (haar aainy Ceratohyal. 

A cee estes Epiphysial bar. 

Di HOLS 5 Sa Geb Ado Ethmoid plate. 

TMLEREGS * sinosc Foramen of I(etc.) nerve. 
de PNY ee ca Hyomandibular. 

ee ieee ence Meckel’s cartilage. 

INAS cree aee Nasal septum. 

INGO Mei ees Notochord. 

1 ee SW aacoancs Pectoral fin. 

PGs Pectoral girdle. 

EO? Wiener Post-orbital process. 

IP AEe erence Palato-pterygoid cartilage. 
Qi eee Quadrate. 

Ties Senecons ... Rostrum. 

Ria Pee sae Retro-articular process. 
Shively) Ss 03ho Stylo-hyoid. 

SVs ueeeueuins Symplectic. 

Oe Od case seas Trabecule Cranti. 


8. BIBLIOGRAPHY. 


. Baxrour, F. M.—Monograph on the Development of Elasmo- 


branch Fishes. 1878. 

Boxer, H.—Der Schiidel von Salmo salar etc. Anat. Hefte, 
1. p. 49, 1913. 

Brive, T. W.—Cranial Osteology of Amia calva. Journ. 
Anat. & Phys. 1877. 


. Brooks, H, J.—On the Osteology of the Haddock. Proc. 


Roy. Dublin Soe. 1883. 


. Faau, L.—Report of the Danish Oceanographical Expedition : 


Clupeide. 1920. 


. Huxtey, T. H.—On the Theory of the Vertebrate Skull, 


Proc. Roy. Soc. 1868. 


. SEwERTZOFF, A. N. 


CHONDROCRANIUM OF THE LARVAL HERRING. 1229 


. Lrzour, M. V.—Post-Larval Stages of Pilchard, Sprat, and 


Herring. Journ. Mar. Biol. Assoc. 1921. 


- McMouraicu, J. P.—On the Osteology and Development of 


Syngnathus. Q.J.M.S8. 1883. 


. Parker, W. K.—On the Structure and Development of the 


Skull of the Salmon. Proc. Roy. Soc. 1872. 


. Potnarp, H.B. On the Suspension of Jaws in Fish. Anat. 


Anz. 1895. 


. Scuiere, W.—Die Entwickelung der Knopfknochen bei dem 


Lachs und der Forelle. Anat. Hefte, 1904. 
Die Entwickelung der Occipitalregion 


der niederen Vertebraten. Bull. Soc. Imp. Nat. Moscow, 
1895. 


. SEwEeRTzorF, A. N.—Die Entwickelung des Selachierschadels, 


Jena, 1899. 


. Sour, P.—Zur Entwicklungsgeschichte des Kopfskeletts der 


Teleostier. Leipzig, 1882. 


. SWINNERTON, H.—Morphology of Teleostean Head Skeleton, 


Q. J. M.S. 1902. 


. TorstEN PreHrson.—Some points in the Cranial Development 


of Teleostomian Fishes. Acta Zool. 1922. 


. Unumann.—Studien zur Kenntniss des Schadels von Cyclo- 


pterus. Zeitschr. f. Naturwiss. 1921. 


. Veit, O.— Die Entwickelung des Primordialcranium von 


Lepidosteus osseus. Anat. Hefte, 44, pp. 93-225, 1911. 


. Winstow, G. M.—The Chondrocranium in the Ichthyopsida. 


Bull. Essex Inst. 1896. 


ON THE COLOURS OF WATER-MITES. 1231 


54. On the Colours of Water-Mites. By C. 8. Eiron, 
New College, Oxford *. 
[Received August 31, 1922: Read November 7, 1922. | 


ConTENTS. 

Page 
Henlintrodueti omg -ceeen od puheas meLOST 
2. The Mites of euler Daioh, Ona SEE are 1232 
3. The Evidence for Warning Coloration............ 1233 
4. The Evidence for Millerian Mimicry ............ 1235 
Se DISCUSSIOMA Wind tvcee nate oaadlanecsonsumetegeenctarne: mL ZOO 
GMS uM ay ape stems nee eancielcciee ec acioaerecsuiero eran ee Mm SOO, 


1. InrRopDuUCTION. 


There are in the British Isles over 250 species of Hydrachnide 
or Water-mites, belonging to about 40 genera [7]. The life- 
history is interesting, as the larva is usually parasitic upon some 
other fresh-water animal (e. g., Hydrachna globosa or Dytiscus, 
Nepa, ete.), although the adult is free-living. By such means 
dispersal is often effected. One result of this rather elaborate 
and risky life-history is that comparatively few adults are 
usually found. We should expect, a priori, to find some special 
method of defence among these mites, since they are not able to 
increase rapidly in numbers, in order to counteract the depre- 
dations of enemies. 

The colours of Water-mites are very varied. As this paper 
deals mainly with the scarlet species, the genera may be roughly 
divided into four groupsin order to give some idea of the colours 
which occur : 

1. All the species of the genus bright red, e. g., Hylais, Diplo- 

dontus, Hydryphantes, etc. 

2. Most of the species of the genus bright red, but a few non- 

red, e. g., Hydrarachna. 

3. A few of the species of the genus bright red, but most of 

them non-red, e. g., Limnesia, Arrhenurus, Piona. 

4, All the species of the genus non-red, e. g., dtav, Hygrobates. 

This is a large group and the colours are extremely varied. 

(These groups are not intended to express the natural relations 

of the genera.) 


The ground-colour of the mites is due to pigment present in 
the skin. Markings of various sorts are produced by the internal 
organs showing through the skin. 

It is the object of this paper to prove that warning coloration 
and probably Miillerian mimicry exist among some at least of 
the Hydrachnide. 

T am much indebted to Prof. EH. B. Poulton for help and 
encouragement during the preparation of this paper, and to 
Mr. C. D. Soar for help in identifying the mites, and for other 
valuable information mentioned later cn. 


* Communicated by Professor KH. B, Pounton, F.R.S., F.Z.8 


12323 MR. CG. 8. ELTON ON THE 


2. ‘Tue Mires or Rattway Drrew, Oxrorp. 


If there are no particular laws governing the occurrence of 
colours in mites we should expect to find chance mixtures of 
colours among the species of any one habitat. In the following 
example such was clearly not the case. During May 1921 the 
fauna of a small ditch near Oxford (herve referred to as ‘* Railway 
Ditch”) was examined by me. This ditch is connected by 
several channels with the Thames, but is usually almost stagnant. 
Only one part of it was studied, an area about 30 yards long and 
4 feet wide Since it is important to know what animals and 
plants are associated with the mites, a list of the species observed 
during the course of the work is given below. This list is not 
by any means exhaustive, but gives some idea of the type of 
habitat in which the mites were living. 


Prants: Ranunculus aguatilis Age., Hottonia palustris L., Calli- 
triche aquatica Smn., Potamogeton natans L., P. perfoliatus L., 
P. densus L. (Lemna minor L., L. trisulea L.). 

Hiruptnea: Two large species. 

Moxuusca: Limnrcew stagnalis L., L. peregra Mill., Planorbis 
corneus L., P. complanatus L., P. spirorbis Miull., Aneylus 
lacustris L., Bithynia tentaculata L., Paludina vivipara L. 

INSECTA : 

Coleoptera: Dytiscus marginalis L., Typhrydus ovatus L., 
Hydroporus palustris L., Agabus bipustulatus L., Haliplus 
sp. 

Hemiptera: Vepa cinerea L., Hydrometra stagnorum L., 
Notonecta glauca L., Corixa sp., Velia sp., Gerris sp. 

Neuroptera: Various larve. 

Crustacea: Cyclops serratulus Fischer, C. prasinus F. 

Chydorus sp. 
Asellus aquaticus L. 

Fiso: Gasterosteus aculeatus L. 

AMPHIBIA: Frog tadpoles. 

The mites and their colours are shown in Table I, 


SPECIES. CoLotr. 
Hydrarachna globosa De Geer. Scarlet. 
HA. schneidert Koen. Scarlet. 
H, distincta Koen. Searlet. 


H. fuseata Soar. Dark purple with reddish spot on back. 


Hylais hamata Koen. | Scarlet. 
Diplodontus despiciens Mull. | Scarlet. 
Hydryphantes ruber De Geer. Scarlet. 


Piona longipalpis Krend. Scarlet with black spots on back. 


Limnesia fulgida Koch. | Scarlet with black marks on back. 


COLOURS OF WATER-MITES. Wasa 


The ditch was full of breeding 3-spined Sticklebacks (Gastero- 
steus aculeatus). The males were guarding their nests and 
driving off all intruders fiercely. The mites were not abundant, 
only about 20 specimens being taken, after a good many days of 
watching and collecting. This fact made it impossible to do as 
many experiments as one could have wished. When a scarlet 
mite did appear, it was extremely conspicuous and sometimes 
swam near a male fish without being touched. 


3. Tur Evipence ror WARNING COLORATION. 


Experiments were made in May 1921 in order to see whether 
mites are eaten by Sticklebacks. 

Expt. 1. A large male 3-spined Stickleback from Railway 
Ditch was placed in a vessel] of water, and starved for three days. 
It was then offered a Daphnia pulex, which it devoured. 
Directly afterwards a Limnesia fulgida was put in. The fish 
made towards it from an inch or two away, but stopped half or 
quarter of an inch from it and swerved off. It repeated 
this after a minute or two. After that the mite was disregarded 
altogether. When given another and brighter L. fulgida the 
fish went up to it once in the same way, and afterwards ignored 
it. It was given another Daphnia which it devoured eagerly. 

Haupt. 2. The last experiment was repeated on the same fish, 
Hylais hamata being used instead of Limnesia. The mite was 
completely ignored after several inspections. 

Hupts. 3, 4,5. The same result was obtained when Diplo- 
dontus despiciens, Piona longipalpis, Hydrarachna distincta, 
respectively, were used. 

These experiments show that the fish avoided scarlet mites 
even when it was fairly hungry, that the latter were presumably 
distasteful to it, and that the fish remembered this fact from its 
former experience. 

Expt. 6. A Stickleback was placed in a vessel as before. 
A Daphnia was eaten eagerly. An Lylais hamata was put in 
and avoided by the fish. Another Daphnia was eaten. <A 
Hydrarachna fuscata was now introduced. The LZ. hamata 
accidentally bumped into the fish, which whipped reund and 
snapped up the mite, but immediately spat it out again. The 
fish then approached H. fuscata but did not touch it. A Daphnia 
was then put in and eaten. 

This experiment shows that 1. hamata was actually distasteful 
to the Stickleback, and that 1. fuscata was avoided. 

Mr. Soar tells me that he has never seen any mite eaten except 
by other mites. He once observed a Dytiscus larva which refused 
to touch a Diplodontus despiciens. Piersig [8| quotes an obser- 
vation of Dugés to the effect that Vepa refused to touch water- 
mites, but does not state what species. Thus, scarlet mites are 
avoided by two of the fiercest enemies of fresh-water animals, 
Dytiscus and Sticklebacks, both of which were present in 


1234 MR. C. S. ELTON ON THE 


Railway Ditch. [Mites will often attack each other. Piona 
longipalpis was seen to attack Limnesia fulgida and Hydrarachna 
fuscata, and Mr. Soar states that L. fulgidw will often attack 
Hylais. | 

Hydrarachna fuscata, which is not scarlet, has the peculiar 
habit of crumpling up its legs and feigning dead when touched. 
Mr. Soar states that he does not know of any other Water-mite 
which does this, although mites sometimes are found resting in 
a crumpled up position. Before we can say what the precise 
use of this habit is, more experiments are required. But 
the following occurrence shows that it may be an advantage 
sometimes. 

Hapt.7. A Hydrarachna schneideri and a H. fuscata trom 
Railway Ditch were put ina jar with a specimen of Agabus bypus- 
tulatus from a pond in Oxford Botanic Gardens in which no mites 
had ever been seen, although regular collections had been made 
for nine months. The beetle attacked and ate 1. schneideri. 
Meanwhile H. fuscata feigned dead during the disturbance and 
remained unharmed. 

This experiment does not prove anything with regard to the 
distasteful properties of 1. schneideri, since the beetle was almost 
certainly unused to scarlet mites. 

Expt. 8. In the autumn of 1921, a young 3-spined Stickleback 
from a flood-pond near Railway Ditch was starved for three 
days and then given a Tubifex, which it gobbled up. A Limnesia 
fulgida from Railway Ditch was putin with it. The fish dashed 
at the mite and caught hold of one leg, but immediately let go 
and left it. After this the fish went up to within a quarter of an 
inch of the mite several times, but did not touch it. ‘The mite 
was left in with the fish for seven days, but still remained 
uneaten, although the latter had now been starved for ten 
days. 

Expt. 9. In the end of February 1922 a young 3-spined 
Stickleback was taken in Railway Ditch and was starved for 
four days. (No mites were to be found in the ditch at this 
time.) <A Simocephalus vetulus was put in and eaten eagerly. 
A nymph of Hydrarachna paludosa 'Thon. from a salt stream at 
Marcham was now putin. ‘This species is coloured very brilliant 
scarlet. The fish went up to it and took the mite into its meuth, 
but immediately spat it out. It went up to the mite repeatedly 
after this without eating it, and followed it round for some 
time, clearly torn between its ravenous hunger and the un- 
pleasantness of the mite. The latter was now removed. 

Two days later, the Stickleback (having now been starved for 
six days) was given two Simocephalus, which it devoured. 
FH, paludosa was then introduced. This time the fish snapped 
up and spat out the mite many times, but sometimes avoided it, 
and never swallowed it. 

This experiment shows among other things the effect of 
hunger on the reactions of the fish. 


COLOURS OF WATER-MITES. 1235 


All Water-mites possess large skin-glands, provided with 
so-called ‘‘ sensory” hairs, whose function has been, so far, 
uncertain. As Piersig (3) points out, it has long ago been 
suggested that these glands secrete some unpleasant fluid; but 
he states that there is not enough evidence to decide whether 
the mites are distasteful or not. That they are so to Stickle- 
backs is certain, in view of the experiments here described. The 
idea that it is the skin-glands which make the unpleasant taste, 
is supported by the fact that the fish does not injure a mite 
when it takes it into its mouth, and therefore cannot actually 
taste its flesh; also by the fact that the mite is spat out so 
promptly. The anatomy of these glands and the nature of the 
supposed secretion require to be worked out in detail. The 
hardness of the body probably belps to render some mites 
distasteful, as indicated by the fact that a Stickleback when not 
very hungry will sometimes reject a young Asellus, after taking 
it into its mouth. But this could not apply to such soft-skinned 
mites as Mylais and Diplodontus. 

These experiments make it clear that Sticklebacks recognize 
the scarlet mites by their appearance. It is extremely likely 
that it is the colour which is remembered, although no accurate 
work has been done on Gasterostews, in order to prove this. 
But White (8) has recently shown that the American Stickle- 
back (Hucalia inconstans) and Mud Minnow (Umbra limbi) are 
well able to distinguish differences in colour as distinet from 
differences in the intensity of light. She proved by experiment 
that the fish were capable of associating different colours with 
their food, though they were unable to distinguish between fine 
shades of the same colour. The same may well be the case for 
the 3-spined Stickleback. 

There is another point which is of some importance. It may 
be asked what advantage is gained by mites, seeing that they 
appear to be uninjured by fish, even after being snapped up and 
spat out several times. The answer to this is that: (1) Mites 
must be actually destroyed by some animals before thei dis- 
tastefulness can be realized, e. g. Agabus; (2) the fluid which 
seems to be secreted by the skin-glands would ultimately become 
exhausted ; (3) being continually attacked must interfere with 
the life processes of the mite. The degree of effectiveness of 
warning colours depends among other things on the state of 
hunger of the enemies. 


4, Tue Evipence ror MULuERIAN Mimicry. 


There is no evidence at present of Batesian mimicry, 7. e. of 
edible mimics, among Water-mites. It is unlikely that it exists, 
in view of the facts that all Water-mites have the large skin- 
glands and that there are very few cases on record of mites being 
edible. However, the possibility should not be ignored. There 
is on the other hand, strong evidence of Miillerian mimicry 


1236 MR. C. 8S. ELTON ON THE 


among mites. It seems remarkable that eight species of scarlet 
mites belonging to six genera occurred in Railway Ditch. Two 
of these species are members of genera in which the other species 
are mostly non-red (Limnesia and Piona). ‘The only species which 
is not bright red, occurring in the ditch, has a special defensive 
habit, not found among other mites. The absence of any other 
non-red species is remarkable, since there is no dispersal difficulty 
to prevent them from immigrating from the Thames. The shape, 
size, and mode of swimming of all these species are much the 
same, except that Hylais trails its legs behind. 

If we exclude the possibility of coincidence, the explanation 
might be that the red mites possess some advantage which is not 
shared by the non-red ones. Now, there may be two important 
factors affecting the evolution of a group of animals in the 
direction of having a common colour :— 


(1) In the words of Prof. Poulton, “the feasibility of certain 
colours and patterns depending upon their effect on the 
vertebrate eye and thus giving the enemies as easy an 
education as possible” (4), That is to say, scarlet may 
have a more striking effect on the retina of the Stickle- 
back’s eye than any other colour, and therefore be more 
effective as a warning. 

(2) Reinforcing this would be the co-operative advantage of 
Miillerian mimicry as usually understood. 


The early stages in the evolution of a set of Miillerian mimics 
would be easier to conceive of if there was any evidence that the 
first factor is an important one, and its effect would be greater in 
the case of animals which, like these scarlet mites, are of one 
uniform colour and do not possess elaborate patterns. (The 
latter, where they occur, cannot be explained without the aid of 
the Miullerian theory.) Therefore, if it can be shown that 
Sticklebacks remember scarlet mites better than those with other 
colours, the case in favour of Miullerian mimicry will be corre- 
spondingly strengthened. The following experiments show that 
this may be so :— 


Hept. 10. The same fish was used as in Expt. 9, after being 
starved for four days. As was seen in that experiment, a scarlet 
Hydrarachna paludosa was refused after one trial. A Simocephalus 
was then put in and eaten at once. A specimen of Hygrobates 
longipalpis Herm. was introduced. This mite is coloured black 
and yellow, and is quite conspicuous. ‘The fish snapped it up and 
immediately spat it out. This it repeated fourteen times at 
intervals. Another Simocephalus was put in and eaten. The 
mite was again taken in and rejected seven times. Another 
Simocephalus was eaten. The fish now approached the mite 
four times without touching it. After that the mite was more 
or less ignored. A Limnesia undulata Mull. was now put in. 
This species is yellow with black or brown spots. The fish took 


COLOURS OF WATER-MITES. WE 


it in its mouth and spat it out twice, and afterwards avoided it. 
Hydrarachna paludosa (brilliant scarlet) was now introduced, 
and the fish avoided it after one trial. ‘There were also some 
Cypris virens present, coloured a uniform brownish green. ‘This 
Ostracod was too hard for the fish to crush and too big to 
swallow; yet the Stickleback kept rushing at the Cypris ier 
ever they moved, trying to swallow them. This it continued 
to do throughout several days. 

This experiment suggests that scarlet was remembered by the 
Stickleback better than! yellow and black, although the hunger 
of the fish has to be taken into account. But both were more 
effective than the dull uniform colour of the Cypris. 

Hapt. 11. TVhe.same fish was used as in Expt. 1. It was 
experienced as regards scarlet mites. A Daphnia was eaten 
eagerly. An <dAcercus lutescens Herm. was put in. This is a 
small mite, coloured pale pink, dark brown, and yellowish white. 
The ish snapped up the mite, but immediately spat it out again. 
This was repeated three times. After this the mite was avoided. 
A short time afterwards the fish noticed it again, and the whole 
procedure was repeated (the same number of times), A Daphnia 
was eaten. Hylais hamata (scarlet) was introduced, but avoided 
after inspection. Both mites were removed. Next morning the 
experiment was repeated. The fish saw Acercws and snapped 
it up, and then spat it out. Next time the mite was avoided. 
The mite was then removed. Thirty-five minutes later Acercus 
was put in again. The fish went right up to the mite, but did 
not touch it. After this the mite was ignored. Daphnia was 
put in and eaten. Hylais was put in and avoided. This experi- 
ment suggests that the fish was able to remember the scarlet 
much better than the other colours. It is unlikely that the 
difference in impressions made on the fish was due to the scarlet 
mite having a more unpleasant taste than the other, since both 
were spat out equally promptly. It is probable, then, that 
scarlet is the most efficient warning colour fer use in Stickleback- 
haunted habitats. Perhaps it is only the best for such habitats 
at one time of year, 7. e. in the breeding-season of the fish. 
Tt observed yellow-and-biack mites (unidentified) in a Hampshire 
pond where 3-spined Sticklebacks occurred on August 4th, 1922. 
This shows that such mites can exist in some Stickleback ponds. 
Not much is known about the food reactions of these fish, except 
that they are usually very fierce and voracious. It is necessary 
to be very cautious in applying the knowledge about food habits 
of Sticklebacks in one place to those of another. J.T. Saunders 
(5) records a pond in which the adult 3-spined Sticklebacks ate 
nothing but one kind of diatom, while the young ones in the same 
pond and all the fish of neighbouring ponds were carnivorous. 
He also states that the fish soon learn to distinguish by sight 
between different animal foods, and that they are very Benen 
to changes in their environment—e. 7., the diatom-eaters became 
carnivorous in captivity. The latter fact might throw some doubt 


Proc. Zoou. Soc.—1922, No. LX XXII. 82 


1238 MR. C. S. ELTON ON THE 


on the validity of the experiments with mites ete. described 
here; but the uniformity of the results makes it pretty certain 
that the fish would reject Water-mites in nature as well as in 
captivity. 

The fact that the fish in Expts. 10 and 11 did not learn well 
on other colours than scarlet does not weaken the case in favour 
of warning coloration, since the non-red mites used were not in 
the presence of their natural enemies. <Acercus lutescens came 
from a pond in which there were no fish. AHygrobates longipalpis 
and Limnesia undulata were from the Thames. Now, although 
Sticklebacks are abundant in most rivers, Maxwell (2) says: 
“Tt has been observed that the Stickleback is very scarce in the 
Thames.” 

The chief importance of these non-red mites here is that it has 
been shown that they might be at a disadvantage in Stickleback- 
haunted habitats (where scarlet ones are successful), and that 
non-red mites actually tend not to come in such places. 

Some other evidence may be added to this, though very little 
is known about the associations of Water- mites. 

There is a flood-pond near Railway Ditch which was full of 
3-spined Sticklebacks in May 1921 (see Hxpt.8). The very large 
and conspicuous scarlet Hylais exiendens Mill. was taken here 
swimming about among the fish, ‘The only other mite was 
Piona conglobata Koch, which is very minute and is incon- 
spicuously coloured. Hyphrydrus ovatus was unusually abundant 
in Railway Ditch in May 1921. This beetle is coloured bright 
orange below. Jt secretes a liquid when handled, which smells 
like honey and may make it distasteful. It is possible that the 
Miillerian mimicry included this beetle. Shelford (6) records 
Limnochares aquatica, which is a large scarlet mite and is unable 
to swim, as occurring abundantly in a pioneer association of 
Chara, in which were also many species of fish. Mr. Soar tells 
me that all the crawling Water-mites are red, which supports the 
idea that they are warningly coloured. 


5. Discussron. 


Scarlet is rather rare among animals, except in the tropics. 
Higgins (1) has suggested that it is due to the fact that red rays 
are more valuable (there being more energy per unit wave-length 
than in other colours of light), and are therefore usually absorbed, 
not reflected. It is more likely that the rareness of scarlet is due 
to its conspicuousness When seen against natural browns and 
greens (as in our own road-traffie warning signs). In any case, 
we generally find that animals are scarlet only when there is 
some very important use for the colour. For the reason given 
above, it makes an excellent warning colour. It is possible that 
red can be seen from further away in water than other colours. 
This is known to be the case in air. 

Any protection gained from warning coloration and Millerian 


COLOURS OF WATER-MITES. 1239 


mumicry would be, of course, only relative. For instance Piersig 
quotes Forbes, who found that fish did not entirely refuse to eat 
Water-mites, and gives as his own observation that Ranatra 
seized and sucked mites without hesitation (3), but does not state 
whether these were the natural enemies of the mites. Before 
any generalization can be made about the colours of Water-mites, 
more information is required about their associations, together 
with experiments in which their natural enemies are used. 
Mr. Soar states that many mites vary in colour individually a 
great deal (e.g. Piona rufa Koch). It is quite likely that the 
colours of other mites than those described here may have some 
different significance, 

In conclusion it may be pointed out that many Land-mites are 


bright scarlet, and no experiments appear to have been done on 
them. 


6. SUMMARY. 


1. Ecological observations are described which make probable 
the existence of warning coloration and Miuillerian mimicry 
among certain species of Hydrachnide. 

2. Experiments are described which show that these are 
distasteful to and recognized by Sticklebacks (Gasterosteus 
aculeatus). 

3. There is evidence that scarlet is the colour best remembered 
by these fish. This makes the evolution of such a set of 
Mullerian mimics more easy to understand. 

4. Other evidence is given in support of these conclusions. 


Works Quoted. 


i. Hiceins, H. H. 1873. Notes by a Field-naturalist in the 
Western Tropics. Liverpool; p. 46. 
2. Maxwewi, H. British Fresh-water Fishes. London; p. 83. 
3. Prersicg, R. 1900. ‘‘ Deutschlands Hydrachniden.” Zoologica, 
vol. xxil. p. 32. 
4. Pounron, H. B. 1887. Proc. Zool. Soc. p. 238. 
DOMUNDERS: ely LOSS) Proc. Camb: (inks Soc volo xvii 
Dd. 236. 
§. ee. V. E. 1918. Animal Communities in Temperate 
America, Chicago; p. 144. 
. Soar, C. D. 1921. Journ. Photomicrographic Soc. vol. x. 
No. 2. 
8. Waitt, G. M. 1919. J. Exp. Zool. vol. xxvii. p. 443, 


3 


(VA 
tua 
ty) 


hy 


iM 


Sap ai eey 
{ Mi i f ‘i ; Gin 
it? Cite 


SAN 
Watt 
an 


P. Z. S. 1922, STENSIO, PI. 1. 


alsph. 
! 


c.ophth. lat. 


c.ophth. lat. 


a.car.int. 


v.cerebr.ant. 


a.carint. 


DICTYONOSTEUS ARCTICUS and POROLEPIS ? sp. 


NOTES ON CERTAIN CROSSOPTERYGIANS. 1241 


55. Notes on certain Crossopterygians. 
By Enix A:son Srensto, Upsala.* 
[Received September 18, 1922: Read November 21, 1922. | 


(Plate I.¢; Text-figures 1-6.) 


INTRODUCTION. 


In the summer of 1921, just as my paper on the Devonian 
Celacanthids from Wildungen (Stensi6, 1922 a) had been sent 
to press, there arrived at the Upsala Library an interesting 
description of Musthenopteron, published by Bryant already in 
1919, which accordingly could not be dealt with in my paper just 
referred to. A few months later there also appeared a very 
important paper by Watson on the organisation of the Ceela- 
canthids (Watson, 1921). As many of the facts put forward by 
Bryant and Watson in these papers throw new light on many 
questions, I have found it of interest to give here some of the 
views at which I have arrived, partly on the basis of these facts, 
partly by examining my own material of Crossopterygians. I shall 
begin with a detailed description of what is known so far of the 
neurocranium in /ictyonostews, then turning to Husthenopteron, 
Porolepis, and the Ceelacanthids. 


DICYYONOSTEUS ARCTICUS Stensio. 
(Pl. I. figs. 1-4.) 


The material of this fish to be described below has already 
been dealt with by me in a preliminary description in 1918 
(Stensi0, 1918 b), and has further been referred to in my memoirs 
on the Triassic fishes from Spitzbergen and in my paper on the 
Celacanthids from Wildungen (Stensio, 1921; 19226, 19224, 
pp. 196-198, 204-206). It consists only of an anterior part of 
the primordial neurocranium with the dermal bones partly 
preserved. The part is large, measuring 21 em. in length and 
indicating that the head of the fish to which it belonged must 
have attained a very considerable size. 


Primordial Neurocraniwm. 

The occipital region is totally lacking, as is also the labyrinth 
region with the exception of a very short anterior part. The 
remaining two regions, the orbitotemporal and ethmoidal regions 
respectively, are as a whole rather completely preserved, though 
partly crushed and broken. 

The whole orbitotemporal region and the anterior part of the 


* Communicated by Prof. D. M.S. Watson, F.R.S., F.Z.S. 
+ For explanation of the Plate, see p. 1272. 


1242 DR. ERIK A. STENSIO: NOTES 


labyrinth region that is preserved are occupied by an unpaired 
large bone, the sphenoid (Stensio, 1921, p. 56), while in the 
ethmoidal region there is found a paired bone, the ‘exethmoideo- 
preéthmoid (cf. Stensi6, 1922 a, pp. 184-185). Both the sphenoid 
and the exethmoideo-preéthmoid consist chiefly of cancellous 
bone, and exhibit no centres of ossification nor any radiate 
structure. 

We begin our detailed deser:ption with the sphenoid. This 
bone (Sph, text-fig. 1; Pl. I. figs. 1-4) is high at the posterior 
end, but becomes rapidly lower forward, so that its height at the 
anterior end is only about a third of that at the posterior one. 


Text-figure 1. 


v.cerebrant 


acar:int be Ss} Vii 


Dictyonosteus arcticus Stensio. 
Anterior half of the neurocranium in lateral view. The lateral part of the ethmoidal 


region removed, as are also the lateral parts of the dermal bones of the cranial 
roof. Cartilage dotted. Somewhat more than a third of the natural size. 


E.pethm,, exethmoideo-preéthmoid ; Fr, frontal; Psph, parasphenoid; Sph, 
sphenoid ; a.car.int, canal for the arteria carotis interna; alsph, alisphenoid wulst ; 
bp, basipterygoid process; cx, orbito-nasal canal for vessels to or from or both to 
and from the nasal fossa; c.ophth.lat, canal through the alisphenoid wulst for the 
n. ophthalmicus lateralis; s7, sulcus for the vena jugularis (dorsal end of the sulcus 
only seen here); v.cerebr.ant, canal for the vena cerebralis anterior; I, olfactory 
canal; IJ, opticus canal; III, oculomotorius canal; IV, trochlearis canal; V, tri- 
geminus exit; V,, canal for the v. ophthalmicus profundus; VII, the probable 
place of the facialis exit. 


The width of the bone taken as a whole is rather small, except 
at the postero-ventral part where the basipterygoid process (bp) 
is situated. The posterior end of the bone seems to have been 
separated from the ossification, or the ossification following next 
behind in the labyrinth region, by a synchondrosis, and that this 
also has been the case anteriorly towards the exethmoideo- 
preéthmoid is certain. The former end is in side view angulated 
in the way shown by text-fig. 1, the latter, on the other hand, 
convexly arched. The cranial cavity extends forward through 
the whole length of the bone. 


———— 


ON CERTAIN CROSSOPTERYGIANS. 1243 


We can distinguish on the sphenoid an unpaired pars basalis, 
which forms the floor of the cranial cavity, and a paired pars 
lateralis, issuing upwards from the pars basalis along either 
of its dorso-lateral edges and extending upwards to the dermal 
eranial roof, forming the lateral wall of the cranial cavity. 

The pars basalis corresponds in its whole extension to about 
half the height of the sphenoid, its absolute height thus being 
much longer at the posterior end than at the anterior one. From 
the anterior end it first diminishes in width backwards, so that 
some distance anterior to and below the opticus exit (II) it 
forms only a rather thin wall between the orbits (text-fig. 2). 
Behind this place it continually grows wider to the posterior 


Text-figure 2. 


Psch 


Dictyonosteus arcticus Stensio. 


Transversal section through the neurocranium a short distance anterior to the 
opticus exit. Cartilage dotted. Somewhat more than a third of the natural 
size. 


Fy, frontal; Psph, parasphenoid; Sph, sphenoid ; ev, cranial cavity. 


end, which is rather broad owing to the large basipterygoid 
process (6p) on either side. The posterior surface is rather 
concave (Pl. I. fig. 3), probably in the same way as Watson 
(1921, p. 335) has found it to be in Megalichthys and in the 
Carboniferous and Permian Celacanthids. In my description 
in 1918, when the specimen had not been cleaned from the 
matrix *, [ came to the incorrect conclusion that this concavity 
was the fossa hypophyseos and the upper boundary of it the 
dorsum selle (¢f. Stensié, 19186, p. 120; pl. iv. fig. 3; pl. v. 
fies plevas): 

The ventral surface of the pars basalis is straight and plane, 
and has been situated almost horizontally; the dorsal one, on the 
other hand, slopes rather much forwards, and is concave from one 
lateral side to the other—7.e., it is somewhat grooved in a 
longitudinal direction. At about the middle of its length the 


* The cleaning of the specimen from the matrix has been very difficult, and has 
taken a long time, as the matrix was a very hard iroustone. 


1244 DR. ERIK A. STENSIO: NOTES 


latter surface suddenly deepens to a postero-ventrally directed 
fossa hypophyseos, which extends nearly to the postero-ventral 
corner of the pars basalis (fAyp, Pl. I. fig. 2), where it is 
separated from the synchondyrosis behind the sphenoid only by a 
very thin layer of bone. So far as is known, the fossa hypo- 
physeos does not in any way open on the base of the primordiai 
neurocranium or in the mouth. 

On either lateral surface of the pars basalis there is found, as 
already mentioned, farthest posteriorly the large basipterygoid 
process (bp). ‘This process partly arises along the ventral, partiy 
along the posterior edge of the lateral surface, and extends rather 
high upwards, ending dorsally with a truncated end. The 
process has a dorsal surface represented by a truncated dorsal 
end, a postero-ventral surface which has been attached to carti- 
lave, a ventro-lateral surface against which the metapterygoid 
probably articulated, aud an antero-dorsal surface which, together 
with the lateral surface of the pars basalis just amiteuion be it, 
forms a rather shallow and indistinct fossa directed with its 
opening antero-laterally and somewhat dorsally. 

In this fossa there opens a wide canal, which perforates the 
pars basalis in a transverse direction below the fossa hypo- 
physeos to the ee fossa on the opposite side. This 
canal (v.pié, Pl. I. figs. 2, 4) communicates with the fossa hypo- 
physeos by a short but wide posteriorly directed branch, and, 
though it runs ventrally of the posterior part of the fossa 
hypophyseos, must undoubtedly be the canalis transversus, and 
have been pierced by the pituitary vein (Allis, 1914; 1919 a; 
cf. also Stensid, 1922 >). From its external opening in the fossa 
described there a very distinct sulcus (sj, text-fig. 1; Pl. I. fig. 4) 
leads upwards to the dorsal end of the basipterygoid process, a. 
suleus which must have been developed for the jugular vein 
(of. Allis, 1919@; Stensiéd, 19226, loc. cit.). Accordingly there 
is no doubt that this vein in its continuation backwards must 
have run dorsally of the basipterygoid process. 

Immediately in front of the basipterygoid process of either 
side there is found close to the ventral edge of the lateral surface 
of the pars basalis a large oval Ronen (a.car.int, text-fig. 1; 
BE fos ali 2 senda) apparently leading into a vertical canal 
ascending to the cranial cavity. This canal, which must be the 
canal for the arteria carotis interna, ought to have entered the 
cranial cavity not far anterior of the fossa hypophyseos, between 
this and the opticus exit, and thus quite in the normal way 
(of Allis’ 189:72 1909 ET 19T a." Os (192 aio OIA ALO evan 
Allen, 1905; Danforth, 1912; Stensio, 19225). 

The partes laterales of the sphenoid (text-figs. 1, 2; Pl. I. 
figs. 1-4) do not diverge much from one another upwards, thus 
occupying an almost vertical position. Anteriorly they are 
rather thin, but they grow slowly thicker backwards. Along 
their dorsal margins, too, they are partly thin, and so developed 
as to show that the roof of the primordial neurocranium in the 


ON CERTAIN CROSSOPTERYGIANS. 1245 


orbitotemporal region, apart from possible fontanelles, must have 
been cartilaginous (cf. text-fig. 2). 

Hach of the partes laterales has a somewhat concave posterior 
edge, which faces postero-dorsally. This edge also is so developed 
that in part it cannot have been in contact with cartilage, but 
must have been free, forming the anterior boundary of a great 
foramen (v, text-fig. 1). This foramen must, as comparisons 
with Diplocercides make clear, be the trigeminus foramen, which 
thus has been situated in the dorsal part of the synchondrosis 
between the sphenoid and the ossifications in the labyrinth 
region (ef. Stensid, 1922 a). 

“Along the posterior edge each pars lateralis has on its lateral 
surface a vather broad but well-marked “wulst,” which extends 
from the upper edge of the pars lateralis downwards and 
somewhat backwards to the upper end of the basipterygoid 
process. This “wulst” (alsph, text-fig. 1; Pl. I. figs. ie 4), 
which IT have called the alisphenoid Swulst (Stensio, 1922 a, 
pp. 180, 196-197 ; 1922 6), is perforated close to the dorsal ond! 
by a horizontally running canal (c.ophth.lat, text-fig. 1; Pl. I. 
figs. 1-4), which has its posterior opening on tlie posterior and 
its anterior opening on the anterior side of the wulst. At the 
anterior edge of the ventral part of the same wulst, some distance 
above the basi pterygoid ipROwESS, there is found the outer opening 
of a canal (V,, text-fig. 1; Pl. I. figs. 1-4), leading backward, 
inward, and sonneine me dially to ale cranial cavity. 

The division of the eranial cavity enclosed in the sphenoid is 
os aed narrow throughout its extension oar NARS | 1 74 
Eten s 3c also Stensid, LOS TON ple eve nies: 32h tS) While 
its height decreases gently for ward, its width, on the contrary, 
seems rather to grow somewhat larger towards the anterior end. 
Owing to the considerable height of the pars basalis of the 
Speman, the division of the cranial cavity im question, with the 
exception of the large fossa hypophyseos, is situated totally in the 
dorsal half of the neurocranium, and it is evident that the 
division next behind in the anterior half of the labyrinth region 
also had a stmilar high position. 

From the shape ‘of the division of the cranial cavity just 
described, we can conclude with great probability that the 
telencephalon was long and rather high, and that the ventral 
parts of the diencephalon were str ongly developed Thus the 
anterior divisions of the brain of the fish must have resembled 
the corresponding ones in the recent Ceratodus, at least with 
regard to their general shepe (cf. Bing & Burckhardt, 1905). 

The trigeminus branches proper, except the r. ophthalmicus 
profundus and the lateralis branches accompanying these (or 
perhaps their roots), have left the cranial cavity through the 
foramen already referred to, which must be supposed to have 
existed in the upper part of the synchondrosis between the 
sphenoid and the bone or bones following next behind in the 
labyrinth region. The position and probable size of this foramen 


1246 DR. ERIK A. STENSIO : NOYES 


are seen in text-fig. 1(V). After their exit through this foramen, 
the trigeminus branches proper, accompanied by the n. buccalis 
lateralis, have evidently passed forward to the orbit laterally of 
the alisphenoid wulst and dorsally of the basipterygoid process ; 
while the n. ophthalmicus lateralis immediately took «a more 
dorsal course, entering the canal (c.ophth.lat.) in the dorsal part 
of the alisphenoid wulst and, soon after having left this, reaching 
the roof of the orbit. As the canal (c.ophth.lat.) has a rather 
small calibre, it seems to me most probable that the rv. ophthal- 
micus superficialis did not pass through it, but ran independently 
of the n. ophthalmicus lateralis as it does in Sturgeons and 
certain Siluroids, and as it certainly did also in Saurichthyids 
(cf. Stensio, 192 2 b; Workman, 1900, pp. 404-407; Herrick, 
1901, pp. 201-203). The rv. ophthalmicus profundus trigemini 
or a somewhat equivalent nerve must apparently have been 
formed already within the cranial cavity, from where it pierced 
the pars lateralis of the sphenoid independently of the other 
trigeminus branches (or their roots) through the Canale described 
above (eof. Stensid, 1921, pp. 58-59, 93, 169- IAOY 3) US 2 2 ia, 10 NS) 3 
1922 bd). 

The exit of the facialis nerve must, so far as one can judge, 
have been situated somewhat postero-ventrally of the trigeminus 
exit, so that the r. palatinus facialis in its way downwards and 
forwards must have run first posteriorly and then ventrally to 
the basipterygoid process before it reached the orbit (ef. Allis, 
1919 @; Stensio, 1922 a). 

Of the exit of the n. abducens nothing is known with certainty, 
but the nerve may perhaps have left ihe eranial cavity through 
the trigeminus foramen (cf. Stensid, 1922 0). 

The Olfactory nerve has from the anterior end of the cranial 
cavity, as we shall see, entered an antero-ventro-laterally running 
canal in the ethmoidal region. 

The opticus has left the cranial cavity far forward. It per- 
forated the pars lateralis of its side somewhat anterior of the 
middle of the length in a downward and forward direction, and 
has Beached the orbit strikingly low (JI, text-fig. 1; Pl. f. 
fies. 1, 2, 4) immediately dorso- ~caudally of the thinnest part of 
the aan ‘pasalis of the sphenoid. 

Somewhat behind the opticus exit, but only slightly higher 
than this, is found an oval foramen, which undoubtedly trans- 
mitted the n. oculomotorius (IiI, text-fig. 1; Pl. 1. figs. 1, 2, 4). 

High up, rather near the cranial roof and approximately 
midway between the opticus and oculomotorius exits, is found 
the external opening of a fine canal (IV, text-fig. 1; Pl. I. 
figs. 1, 2, 4), which must undoubtedly have transmitted the 
n. trochlearis. 

A rather wide canal (v.cerebr.ant, text-fig. 1; Pl.I. figs. 1, 2), 
which is situated high up anterior of the opticus and trochlearis 
exits, must probably have been traversed by the vena cerebralis 
anterior (ef. Stensid, 1921, pp. 168-169) on its way to the orbit. 


ON CERTAIN CROSSOPTERYGIANS. 1247 


This canal was considered by me in 1918 as the trochlearis canal 
(Stensid, 1918 6), a view which must be incorrect in the light of 
the facts now known. 

At the anterior end of the sphenoid, in the suture between 
this and the eaethmoideo. preéthmoid, a rather wide canal (cn, 
text-fig. 1; Pl. I. figs. 1 is found which leads forwards to the 
olfact eee canal, and w Tae undoubtedly has contained vessels 
to and from the nasal pit (cf. Stensi3, 1922 6). Perhaps it may 
also have been in some relation to the points of vrigin of the 
musculi obliqui of the eye. 

The arteria carotis externa must, so far as one can understand, 
already have been sent out from the arteria carotis communis 
behind the basipterygoid process, and has probably run upward 
approximately as far as to the facialis exit, then turning forward 
and continuing dorsally of the basipterygoid process to ‘the orbit. 
The arteria carotis interna, after its origin behind the basi- 
pterygoid process, must have run forward ventially to this 
process and laterally to the parasphenoid to its ascending canal 
described above in the sphenoid. The position of the ventral 
opening of the ascending canal in relation to the nerve exits 1s 
seen from text-fig. ] and P!. I. figs. 1, 2,4. The vena jugularis, 
after arriving at the dorsal end of the basipterygoid process, 
must have passed backward dorsally of this process, as already 
pointed out, then continuing backwards between the trigeminus 
and facialis exits. 

There is evidently no trigemino-facialis chamber and no 
myodome in the fish. According to the course of the nerves and 
vessels, the basipterygoid process with its antero-ventral parts 
undoubtedly corresponds to certain posterior parts of the basi- 
pterygoid process in Birgeria, with its dorsal parts on the other 
hand, corresponding to the basipterygoid process of the Ceela- 

canthids (ef. Stensid, 1922 a, pp. 205-206; 19226; ef. also 
pp. 1259-1266 below in the present paper). 

The ethmoidal region rapidly decreases in height forwards, 
becoming very low at the anterior end. It is mainly occupied 
by the paired exethmoideo-preéthmoid, only certain smaller 
anterior and dorsal parts being cartilaginous. At the postero- 
medial part the excetlnmnondleee “preéthmoid is separated from 
the sphenoid by a short synchondrosis, as already pointed out 
above. In the posterior two-thirds of the length its. plane 
ventral surface is covered by the parasphenoid. Tts lateral 
surface is accessible for investigation only in the anterior parts, 
being otherwise concealed by remains of very fractured dermal 
bones. 

Asalready mentioned, the exethmoideo-preéthmoid is traversed 
by the rather wide canal for the n. olfactorius (1, text-fig. | ; 
Pl. I. figs. 1, 2), which runs forwards, downwards, and laterally. 
The large anterior opening of this canal is incompletely sub- 
divided into a postero-dorsal division situated at the anterior end 
of the lateral surface of the region and an antero-medial division 


1248 DR. ERIK A. STENSIO: NOTES 


situated on the ventral surface of the region, the former of 
which represents the external, and the latter the internal, nasal 
aperture. In the present state of preservation, dermal bones 
form the direct boundary of the external nasal aperture on the 
dorsal side and of the internal nasal aperture anteriorly, but it is 
conceivable that in the fresh specimen these walls of the two 
nasal apertures consisted, at least in part, of cartilage. 

The r. maxillaris trigemini and the n. Tiasaraallis bucealis ought 
as usual to have had their course along the ventral edge of the 
lateral surface of the ethmoidal region, but owing to the im- 
perfect state of preservation of these parts of the region it has 
not been possible to discover whether a distinct sulcus was 
developed for them as in several other fishes (Stensio, 1921, 

. 94; 19226). For the same reason it has not been possible to 
Bataan anything with regard to the articulation facet for the 
anterior end of the palatoquadrate. 

From the facts put forward here it is easily seen that the 
exethmoideo-preéthmoid is so developed that it represents both 
the exethmoid and preéthmoid (cf. Stensid, 1921, pp. 93- 94: 
1922 a, pp. 184-186). 


Dermal Bones of the Primordial Neurocranium. 


The parasphenoid (Psph, text-figs. 1, 2; Pl. I. figs. 1, 2,4; 
Stensio, 19186, pl. v. fig. 1) is imperfectly preserved at both 
ends. The preserved parts of it cover the whole ventral surface 
of the sphenoid and the ventral surface of the exethmoideo- 
preéthmoids in the posterior two-thirds of their length. It thus 
eannot have extended much farther forward, and it is also 
conceivable that only a rather small part is lacking at its 
posterior end. There is no processus ascendens. The centre 
of ossification is clearly distinguishable, and situated midway 
between the posterior and anterior ends of the bone. At this 
place and for some distance forward and behind it the bone is 
much broader than the ventral surface of the sphenoid, so that 
it projects here with a free lateral part on either side of this 
(text-fig. 2). 

Just at the centre of ossification a rather fine median, unpaired 
canal opens on the ventral surface of the parasphenoid. The 
eanal perforates the parasphenoid obliquely upwards and back- 
wards, then continuing backwards between this and the ventral 
surface of primordial neurocranium towards a place between 
the ventral openings of the internal carotid canals. The 
canal ought, so far as one can judge from ny conditions in 
Ania (Allis, 1897, ‘pp: 620262102 Coll kaxcavat ten Olle ple sexemyite 
figs. 62, 63), to thane been tt averse by the saneenon branches of 
the rami palati ni of the facialis nerves, which ought to have 
accompanied the internal carotid arteries when these entered 
between the sphenoid and the ventral surface of the primordial 
neurocranium. Possibly the canal in question for the anterior 


ON CERTAIN CROSSOPTERYGIANS. 1249) 


branches of the rami palatini also transmitted branches from the 
internal carotids. 

If we imagine that a processus ascendens homologous with that 
in the Teleosts (ef. Stensid, 19226) was developed on the para- 
sphenoid of Dietyonosteus, this process ought to have issued from 
the stretch of the bone situated between the hind end and the 
anterior border of the internal carotid eanal and the arteria 
carotis interna, and the n. palatinus facialis would thus have 
come to run medially of it between it and the ventro-lateral parts 
of the basipterygoid process. 

Of the vomer nothing is preserved in the specimen. 

The arrangement and shape of the different bones in the part 
preserved of ‘the dermal cranial roof is seen in text- fig. 3. The 
bones were described by me in 1918, and especially later in 
connection with my account of the Celacanthids from the 
Triassic of Spitzbergen, and I have here nothing of interest to 
add to these descriptions (Stensio, 1918 4, pp. 116-118; 1921, 
pp. 1383-134). 

In the present state of preservation of the fish the external 
nasal aperture is bounded directly on the dorsal side by the 
two anterior nasal elements and a rostral, while the external 
boundary of the internal nasal aperture is probably formed by 
the premaxillary and the rostral elements behind this. 


Sensory Canals of the Head, 


The infraorbital canal anastomoses on the snout with its fellow 
of the opposite side in a well-developed ethmoidal commissure, 
which is situated in the rostral elements. ‘The supraorbital canal 
traverses the dermosphenotic, the lateral part of the frontal, and 
the three nasal elements, from the anterior one of which it enters 
the rostral plate situated next to the median line, there anasto- 
mosing with the infraorbital canal. The exact course of the 
supraorbital canal is clearly seen in text-fig. 3. 

Of the number of sensory canal organs and tubuli nothing is 
known so far. 

As we shall see from the subsequent description, the sensory 
canals on the snout in Porolepsis run exactly like those 
in Lictyonosteus. 


EUSTHENOPTERON FORDI Whiteaves. 


The material of this fish at my disposal consists of twe small 
specimens collected by Professor P. D. Quensel, Stockholm, 
during his journey to Canada in 1913, and which were kindly 
presented to the Paleontological Institution at Upsala. 

The two specimens are well preserved with regard to the 
dermal bones, but exhibit on the other hand hardly anything of 
the primordial neurocranium. My account of this below is 
therefore based exclusively on the description published by 
Bryant 1919 onthe material in the Buffalo Museum. 


L2EOD DR. ERIK A. SYENSIO: NOTES 


Primordial Neurocranium. 


The general shape of the primordial neurocranium is almost as 
in Diplocercides (Stensio, 1922a). The degree of ossification 1s, 
however, evidently much less than in this. 

“~The basioccipital and the paired lateral occipital described by 
Bryant are both rather thin, and it seems very probable, under 


‘Text-figure 3. 


Dictyonosteus arcticus Stensio. 


The anterior part of the cranial roof. Sensory canals with dotted lines 
and shading. From P. 355. 1/2. 

Ant,, anterior antorbital element; Antz, posterior antorbital element; Dsph, 
dermosphenotic (dermal postfrontal); Hr, frontal; 7, interrostral; Wa,—Nas, 
nasal elements; Pma, premaxillary; Pt, anterior postrostral element; Ptr, 
posterior postrostal element; R, rostrals; So, supraorbitals ; 2a, external nasal 
aperture ; orb, orbita; y, accessory bone-plate. 


the presumption that Bryant’s text-fig. 56 is correct, that they 
did not directly bound either the chorda, which certainly 
extended into the basis cranii for some distance, or the cranial 


ON CERTAIN CROSSOPTERYGIANS. 1251] 


cavity, but that the medial parts of the lateral and ventral walls 
of the occipital region consisted of cartilage. Hxpressed in 
another way: the ossification in the occipital region probably 
occupied only the external parts of the region, while the inner 
ones were cartilaginous. The basioccipital has a rather con- 
siderable extension forward, and actually lies with more than 
its anterior half in the labyrinth region. There cannot, how- 
ever, be any doubt of its homology. The lateral occipital is 
strikingly high, extending from the basioccipital below nearly 
to the dermal cranial roof dorsally (Bryant, 1919, text-fig. 3), 
thus partly occupying the position of the supraoccipital, which 
according to Bryant is totally lacking. ‘The lateral occipital is 
said by Bryant to be perforated by a fine canal, but as he does 
not deseribe the course of this canal, one cannot decide with full 
certainty whether it is a canal for a dorsal branch from the 
n. line lateralis or the vagus canal, although the latter alternative 
seems to be the more probable one. If however, the canal 
should reveal itself to have transmitted a dorsal branch of the 
n. linez lateralis, the vagus nerve must have perforated the 
lateral wall of the primordial neurocranium immediately anterior 
of the lateral occipital. 

In either lateral wall of the labyrinth region of the fish, Bryant, 
1919, p. 18, describes five ossifications. One of these ossifications 
he calls the pro-otic, another the opisthotic. The three remaining 
ones he considers 1t impossible to identify, and accordingly he 
does not name them. 

The description which Bryant gives of the pro-otic is not 
sufficient to make fully clear the shape and actual extension of 
the bone. As I understand his account, the pro-otie is a rather 
low bone (cf. his text-fig. 5 @ and pl. ix. fig. 3), which does not by 
far extend to the cranial roof dorsally. It is perforated by a 
canal, one of the openings of which is situated approximately at 
the middle of the length of the bone just at the boundary 
between its ventral and lateral surfaces or perhaps on the 
ventral surface, though close to the lateral edge (cf. Bryant, 
P99, texte. 4; pl. ix. fig. 3; pl. xiv. figs. 4/5). The canal 
is considered by Bryant to have transmitted the n. facialis (he 
evidently means the truncus hyoideo-mandibularis of this nerve), 
but he gives no statements either as to its course nor as to 
the position of its other opening, and the facts so far known do 
not seem to support this view. As far as can be judged from 
the conditions in recent fishes, the canal seems most probably 
to have been developed for the carotis externa on its way 
upwards from the cranial basis to approximately the height of 
the trigeminus and facialis exits (cf. Allis, 1897, pp. 497-500; 
1908, pp. 219-221; 1909, pp. 51-53, 185-187; 1911 a, pp. 284— 
287; 1914; 1919@; Allen, 1905, pp. 51-56; Danforth, 1912, 
pp. 485-442 ; Stensid, 1922). The truncus hyoideo-mandibularis 
ought not to have run so strongly downwards that it can be 
thought to have pierced the canal. 


1252 DR. ERIK A. SYENSIO: NOTES 


The exact position of the pro-otic of Bryant’s description in 
relation to the sphenoid is not clear, but it seems to me very 
probable that its anterior end in the fresh specimens ought to 
have been situated much more ventrally than it has been drawn 
in his text-fig. 5, and at the same time also closer to the sphenoid 
than in this text-fig. The bone actually seems to me to corre- 
spond only to the basal parts of the corpus of the DECC CGS 
opisthotic in post-Devonian Ceelacanthids (¢f. Stensi6, 1921, pp. 55, 
120-121; 19224, pp. 199-201). 

The *‘small quadrangular bone,” which according to Bryant is 
attached by suture to the outer and hinder margins of the pro-otic 
and which extends nearly vertically upward to the cranial roof 
(Bryant, 1919, p. 18), must evidently correspond to some dorsal 
part of the corpus of the prootico-opisthotic in the post- Devonian 
Coelacanthids. The description and the figures given by Bryant 
have failed to convince me that the bone is independent, and 
IT cannot help suspecting that in reality it may be simply a 
broken dorsal part of the pro-otic. 

The bone which Bryant interprets as the opisthotic, much 
resembles, as he points out (p. 18), the opisthotic in certain 
Stegocephalians, and undoubtedly represents an opisthotie to 
some extent. 

The two remaining bones of the region, which are both 
small, are imperfectly known, and it is not possible to decide 
whether they are broken parts of the other bones or not. 

In the anterior half of the bottom of the labyrinth region there 
is In the present state of preservation a large opening bounded by 
the basioccipital behind, by the sphenoid anteriorly, and by the 
pro-otics laterally. This oppeueinls which occurs also in Ceela- 
eanthids (Stensi6, 1921, pp. 58, 121; 1922 a, pp. 173, 202), appa- 
rently must have existed in Dictyonosteus too. It is situated 
entively behind the hypophysis, was probably traversed by the 
anterior end of the chorda, and corresponds fairly well in its 
position to the fenestra basicranialis posterior in the embryos of 
veptiles (Gaupp, 1905, pp. 663, 757-760, figs. 382, 383; Allis, 
1919¢; Stensis, 1922 a, p. 202 \ In the fresh specimens it was 
probabl: y at least to a gr eat extent filled by cartilage. 

That it cannot be the fossa hypophyseos, as Bryant pre- 
sumably means (pp. 18-19), is fully evident from the conditions 
in Dionne and the Ccelacanthids. 

The orbitotem poral region and the antero-ventral end of 
the labyrinth region are “occupied by a large unpaired bone, 
the sphenoid, which is on the saltlle very stall to that in 
Dictyonosteus (cf. pp. 2-9 above). Thus its general shape is 
as in the latter fish, and it has a paired basipterygoid 
process and a paired alisphenoid wulst situated in the same 
wey as in this fish (cf. Bryant, 1919, text-fig. 5 and pl. ix. figs. 

3), and the division of the cranial cavity enclosed in it as 
oe as seveval of the canals piercing it are in the main as in 
this. On the other hand, if Bryant’s account is correct, it 


ON CERTAIN CROSSOPTERYGIANS. 1253 


extends, contrary to that in Dictyonosieus, far forward in the 
ethmoidal region, thus probably including in its anterior parts 
the homologues of the exethmoideo-preéthmoids (¢f. pp. 9-10 
above). “Bryant’s account of the anterior parts of the bone 
is, however, very incomplete, and as in addition the figures 
he publishes are not very distinct and lack letters of reference, 
it is impossible to understand how the conditions actually are 
there. 

In accordance with the conditions in Dietyonosteus and 
Diplocercides the n. trigeminus must have pierced the carti- 
laginous part of the cranial wall just behind the dorsal halt of 
the sphenoid. The exit of the n. facialis has probably been 
situated somewhat more postero-ventrally, though also in the 
cartilaginous part of the cranial wall behind the sphenoid (cf. 
Stensid, 1922 a). 

If my opinion, just given, with regard to the position of the 
trigeminus and facialis is correct, 1t follows from this that the 
small cartilage bone that is seen in Bryant’s text-tig. 5 immediately 
beneath tbe dermal cranial roof some distance behind the 
sphenoid would be homologous with a dorsal part of the process 
lettered f by me in the post-Devonian Ceclacanthids (Stensid, 
1921), and the corresponding antero-dorsal part on either side of 
the compound occipito-labyrinth ossification in Déiplocercides. 
This small cartilage bone is, however, so far as I can find, not 
mentioned by Bryant in his description. 

The upper part of the alisphenoid wulst is pierced by a 
horizental canal for the n. ophthalmicus lateralis, exactly as in 
Dictyonosteus. ‘Vhe anterior opening of this canal is seen in 
Bry2nt’s text-fig. 5 and also, though less distinct, in his pl. ix. 
fig. 3. The presence of this canal is undoubtedly a strong 
support for my view, given above, as to the position of the exits 
of the n. trigeminus and the n. facialis. 

The canal which is stated by Bryant to open in a recess on the 
external surface of the sphenoid just anterior of the basipterygoid 
process, must evidently have transmitted the pituitary vein. 
According to Bryant it leads forwards to a ‘‘ chamber” on the 
posterior basal part of the sphenoid—a ‘‘ chamber ” which is said 
to be open behind and which, with some doubt, is taken to be the 
anterior end of a myodome. That the “chamber,” which in 
another place is termed by him a ‘‘ pocket,” cannot be a part of 
a myodome is evident, and it is quite certain that no posterior 
myodome was developed, but that the conditions were as in 
Dictyonostews in this respect. If the statements given by Bryant 
were correct, the ‘‘ chamber” in question would therefore be the 
ventral part of the fossa hypophyseos. As, however, as far as can 
be judged, it is very wide, I feel inclined to think that Bryant may 
have made a similar mistake as I did in 1918 when describing the 
corresponding part of the sphenoid m Dictyonosteus (ef. p. 1242 
above). I thus rather fear that in the “chamber” we are 


Proc. Zoou. Soc.—1922, No. LXXXITI. 3 


1254 DR. BRIK A. STENSIO : NOTES 


concerned simply with the concavity filled with stone in the 
posterior end of the pars basalis of the sphenoid, and that the 
canal for the pituitary vein actually does not open there, but 
runs somewhat in front of it, ventrally of the fossa hypo- 
physeos, exactly as it does in Diciyonosteus. 

‘The foramen seen in Bryant’s figures some distance in front of 
the upper end of the basipterygoid process at about the middle 
of the height of the sphenoid corresponds approximately to the 
external opening of the oculomotorius canal in Diciyonosteus, 
but a certain interpretation of it is not possible, as we do not 
know the course of the canal leading medially trom it, nor 
several of the other canals of the region. It actually seems to 
have a position somewhat too far anteriorly to be the external 
opening of the canal for the ramus ophthalmicus profundus 
trigemini and, on the other hand, too far back to be the external 
opening of the opticus canal. 

The foramen which in Bryant's figures is seen in the anterior 
part of the sphenoid may probably be the external opening of the 
opticus canal. This interpretation is specially supported by the 
fact that the eye has been situated far forward. 

No other foramina or canals that can be thought to have 
transmitted the n. trochlearis, the r. ophthalmicus profundus 
trigemini, the arteria carotis interna, and the v. cerebralis 
anterior are described by Bryant, nor are any canals of this sort 
clearly discernible in the figures he gives of the different 
specimens, though they must of course exist. 

As is evident from the facts given here, the vena jugularis, the 
r. palatinus facialis, and the internal carotid artery must in 
relation to the basipterygoid process have had the same course as 
in Dictyonosteus. 

Of the primordial skeleton of the ethmoidal region of the 


fish there is nothing to be added to what has already been 
written above (p. 1253). 


Dermal Bones of the Primordial Newrocranium. 


The parasphenoid is, according to Bryant’s description, 
developed in the same way as that in Dzctyonosteus and 
Saws ipterus (cf. Hastman, 1917, pl. vil. fig. 5), but is much 
more narrow than in these two forms. In my specimens there 
are only fragments of it preserved. It is noteworthy that it 
seems to be very firmly connected with the sphenoid. The 
paired vomer seems to show the normal conditions. Nothing 
of it is seen in my specimens. 

If, after these short remarks on the bones of the ventral 
surface of the primordial neurocranium, we turn to those of 
the dorsal one (text-fig. 4), it is clear at once that Bryant has 
correctly recognized the extrascapular bones, the parietal, the 
supratemporal and the intertemporal, and it ought also to be 
added here that the sutures between the extrascapular bones 


ON CERTAIN CROSSOPTERYGIANS. 1255 


mutually and between these and the bones in front are more 
correct in Bryant’s text-fig. 2@ than in the restoration pub- 
lished by me in 1921 (Stensid, 1921, text-fig. 57)*. 

Between the anterior end of the intertemporal and the postero- 
lateral corner of the frontal there is in my specimens a very 
distinct small dermosphenotic (Dsph, text-fig. 4), and I also find 
from pl. v. fig. 2 and pl. iii. fig. 2 in Bryant’s paper that the 


Text-figure 4. 


Busthenopteron fordii Whiteaves. 


Dermal cranial roof. After a specimen belonging to the Paleontological Institution, 
Upsala. The sensory canals marked with dotted contours and shading. 1/1. 


Ant, antorbital; Dsph, dermosphenotic; #Hwxt, lateral extrascapular; Hat.m, 
median extrascapular; Fr, frontal; J¢, mtertemporal; Za, lacrymal; Ma, maxil- 
lary; Wa, nasal; Pa, parietal; Pmx, premaxillary; Pér, postrostral; R, rostrals 
and interrostrals; So, supraorbital ; Stemp, supratemporal ; ifc, infraorbital sensory 
canal; Je, cephalic portion of the main lateral line; mp, groove indicating the 
position of the middle head line of pit organs; na, external nasal aperture; 07d, 
orbital entrance; pp, groove indicating the position of the posterior head line of pit 
‘organs; soc, supraorbital sensory canal. 


same bone was an independent element in the specimens investi- 
gated by him. Its exact position and relation to the sensory 
canals is shown in text-fig. 4 and in my restoration published in 
1$21 (Stensid, 1921, text-fig. 57), and will be further dealt with 
below in the account of the sensory canal system. 

The frontal in Bryant’s restorations thus has a too large 
extension postero-laterally. The pineal foramen between the 
two frontals is not distinct in my specimens, though I can 


* This was due to the fact that my restoration was made after a specimen 
showing the inner surfaces of the bones. 
Sam 


1256 DR, ERIK A. STENSIO : NOTES 


conclude with the guidance of Bryant’s figures where it must 
have been situated. 

The bone which is held by Bryant to be the postfrontal is 
clearly seen to consist of two mdependent bones in both my 
specimens (text-fig. 4), and I also find there that the posterior 
one of these two bones, which is undoubtedly a compound 
supraorbital (so), extends so far backwards as to meet the 
anterior end of tne dermosphenotic. The anterior bone (Anz), 
on the other hand, is situated mainly anterior of the orbit, 
forming the anterior boundary of the entrance to this, and is 
certainly to be interpreted as an antorbital bone. In the main, 
the supraorbital and the antorbital are correctly drawn in my 
restoration published in 1921 (Stensio, 1921, text-fig. 57; the 
suture line between the nasal and antorbital has fallen away on 
the left side of this figure). 

The postrostral elements of the fish have fused to a rather large 
unpaired plate (Per, text-fig. 4), as described by Bryant (called 
by him interfrontal), and my restoration in 1921 (text—fig. 57), 
is thus incorrect in this respect. 

The paired nasal bone is well seen in one of my specimens. It 
is situated along the lateral edge of the postrostral plate, 
extending forwards to the anterior end of this plate, backwards 
on the other hand somewhat farther (Va, text-fig.4), Its length 
thus exceeds that of the postrostral plate. The breadth is rather 
small, and constitutes only about one-half of thelength. Through- 
out its length the bone is pierced by the supraorbital sensory 
canal. My restoration in 1921 thus showed correctly its 
approximate shape and relations. It is also noteworthy in this 
connection that the left nasal bone of the specimen actually seems 
to consist of two or three independent elements, the conditions 
thus much resembling those in Dictyonosteus. 

That the shape and extension of the nasal bone in Bryant's 
vestoration (his text-fig. 2a) must be incorrect is thus evident. 
That this is the case is also seen from pl. in. fig. 2 in Bryant’s 
paper in question, where the bone is rather well displayed in 
ventral aspect and shows a longitudinal rounded ridge caused by 
the sensory canal in it. 

Anterior to the postorbital plate and the nasals, I find in the 
smallest of my specimens (the one that has the nasals well 
displayed) fragments of a transverse series of bone-plates, which 
are the rostrals and probably also the interrostrals, and finally 
anterior to these the rather narrow premaxillaries. The number 
of the rostrals and interrostrals cannot be ascertained, and it is 
also impossible to make out anything as to the exact shape either 
of any of these bones or of the premaxillaries. A rostral plate 
is also observed by Bryant (p. 9). That he has not seen more 
of these plates is easy to explain, as they are very difficult to 
observe and in larger specimens have perhaps fused more or 
less intimately with the premaxillaries or other neighbouring 
bones. ' 


ON CERTAIN CROSSOPTERYGIANS. VO57 


Dermal Bones of the Cheek. 


The dermal bones of the cheek posterior of the orbital entrance 
seem in the main to be correctly described by Bryant. In my 
specimens I can clearly observe the postorbital, the jugal, the 
squamosal, and the preopercular element. The jugal ought, 
perhaps, to have a larger extension forward beneath the orbital 
entrance than Bryant has given it in his restoration (his text- 


fig. 2). 
Viseeral Skeleton. 


The visceral skeleton is badly preserved in my specimens, but 
T find, however, that Bryant is correct in distinguishing a dermo- 
palatine (termed palatine by him), an ectopterygoid, and an 
entopterygoid. 


The Sensory Canals of the Head. 


The sensory canals proper form closed canals in the bones, and 
open outwards with tubuli in the normal way. As the bones are 
usually thickened on the inner side along the sensory canals, the 
course of these canals can be rather well traced without prepara- 
tion in those cases where the bones display their inner surfaces. 
This is partly finely shown by certain of Bryant's figures, e. g. 
pl. ii. fig. 2; pl. v. figs. 2,3. At least most of the structures 
described by Bryant as sensory canals proper are grooves indi- 
cating the position of lines of pit organs, as we shall see from the 
subsequent account. 

The supraorbital canal (soc, text-fig. 4) is very well preserved 
in my specimen nearly throughout its length. Its posterior end 
is situated in the dermosphenotic, where if anastomoses with the 
infraorbital canal. From there it runs forward through the 
lateral part of the frontal, and then pierces the nasal in a longi- 
tudinal direction. From the nasal it must have entered one of 
the lateral rostral plates, in which it probably anastomosed with 
the infraorbital canal of its side. The sensory organs in it have, 
as far as one can judge, been comparatively few, as the foramina 
for nerve-branches through its lower wall are comparatively few 
(cf. eI 1889, pl. xli.; Cole, 1898, pl. xxii.; Herrick, 1899, 
ill, Seva, 5 USI pl. Xiv., etc.). 

The infraorbital canal [(ife, text-fig. 4): defined as in my 
memoir on the Triassic fishes from Rey ee ; Stensid, 1921, 
p- 103] has its posterior end at the boundary between the 
supratemporal and intertemporal. In its course forward it 
pierces first the intertemporal, then enters the dermosphenotic, 
in which it anastomoses with the posterior end of the supraorbital 
canal, as mentioned in the description of thisregion. Just at this 
anastomosis it turns laterally, and continues laterally and down- 
wards on the cheek, where it pierces the postorbital, the jugal, 
and certainly also the lacryma]. It probably was in communi- 
cation with its fellow of the opposite side through an ethmoidal 


1258 DR. ERIK A. STENSIO: NOTES 


commissure, and it is also conceivable that its anterior parts 
anastomosed with the anterior end of the supraorbital canal as 
pointed out above, but it is not distinctly preserved anterior of 
the orbit in my specimens. 

In the jugal there issues from the infraorbital canal a jugal 
canal (ja, text-fig. 4), which runs backwards and downwards to 
the squamosal, in which it continues to the postero-ventral 
corner. In its further course this canal must have entered the 
quadratojugal, if the restoration of this bone given by Bryant is 
correct. As the most postero-ventral part of ‘the canal seems to 
be homologous with a ventral part of the preopercular canal (ef. 
Stensid, 1921, pp. 76-77), the quadratojugal ought to comprise a 
preopercular component (cf. Pearson, 1922, pp. 56-58). 

A true preopercular canal is not developed. The mandibular 
ennal is not clearly seen in any specimens. 

The cephalic portion of the main lateral line (dc, text-fig. 4) 
has its anterior end at the boundary between the supratemporal 
and intertemporal, where it is directly continuous with the 
posterior end of the infraorbital canal. It runs backward 
through the supratemporal and the lateral one of the three extra- 
scapular bones. Through a_ well-developed supratemporal 
commissure (s.com., text- fig. 4), which pierces the three extra- 
scapular bones transversely, it is in communication with that of 
the opposite side. 

On the posterior part: of the dorsal surface of the frontal there 
is found a transverse fine groove (mp, text-fig. 4), which continues 
laterally also over an adjacent part of the supratemporal. 
Another groove (pp, text-fig. 4), which, however, is very short, 
issues from the medial end of the former in a_postero-lateral 
direction. The two grooves which are described by Bryant as 
a sensory canal proper (p. 12) and which, as it seems, have been 
‘interpreted in this way also by Watson (192), p. 334), are 
undoubtedly developed for lines of pit organs. The longer of 
them probably lodges the middle head line of these organs, and 
the short obliquely running one the posterior head line (¢. 
Stensid, 1921, pp. 218, 263; 19224, pp. 224, 235; 19226: ef. also 
Allis, 1899, pp. 502-509; 1900, p. 445; 1903, p. 187; 1905, 
pp. 406, 410, 418, 440, 452, 465, 469, 474, 484: Herrick, 1901, 
oe 222— 293, pl. xiv.: Pander, 1860, taf. i11. figs. 1, 3, 4, 12, 13, 
23; taf. iv. fig. 2: Jaekel, 1911, figs. 84, 85). I find no certain 
traces of any anterior head line. 

Of the other structures described as sensory canals by Bryant, 
the posterior one of the two on the mandibula is clearly shown in 
my specimen to be a pit line, and the same ought also to be the 
case with the other ones shown in his restorations (Bryant's 
text-figs. 2c, 3 6). 

As is evident from this description, the sensory canals proper 
of Husthenopteron are developed exactly as described by Goodrich 
in Osteolepis (Goodrich, 1919) and by me in Dictyonosteus and 


ON CERTAIN CROSSOPTERYGIANS, 1259 


the Ceelacanthids (Stensid, 1921). The pit lines seem, so far 
as is known, to agree rather well with those in other Crosso- 
pterygians. 

POROLEPIS ? sp. 


Among the material collected by the Norwegian expeditions 
in the Lower Devonian strata of Northern Spitzbergen, and for 
the investigation of which | am indebted to Professor J. Kiaer, of 
Kristiania, there is an anterior part of the cranial roof, which in 
this connection deserves some remarks. The part Tha belonged 
to a very large fish, probably ae to the same genus as 
the scalés known under the name Porolepis posnaniensis Kade 
(cf. Woodward, 1891, p. 9; Stensid, 1918 a, pp. 66-73). 

On the upper surface of the part present (Pl. I. fig. 5) no 
sutures at all can be observed. If, however, the outer layers of 
the bones are removed, the radiation from their ossificatory 
centres appears, though not so distinctly as in Dictyonosteus ox 
Husthenopteron owing ‘to the less favourable state of preservation 
of the bone tissue. By means of this method, I have been able 
to observe that the division of the dermal roof covering the 
ethmoidal region consists of a large number of plates, probably 
even larger than in Dict; tyonosteus. The exact arrangement and 
extension of these plates as well as their number can, however, 
not be made out with the help of the fragmentary material at my 
disposal. One can only see clearly that the nasal is represented 
by two or three plates. ‘he supraorbital sensory canal (soc, Pl. I. 
fig. 5) runs exactly as in Dictyonostews. I+ communicates 
anteriorly with the anterior parts of the infraorbital canal 
(tfc) of its side, which in its turn anastomoses with that of the 
opposite side through a well-developed ethmoidal commissure 
(ethm.com.), the conditions thus being also in these respects as in 
Dictyonosteus. 

THE C@LACANTHIDS. 


The Ceelacanthids are now rather completely known (Stensié, 
1921, 1922 @, 19226; Watson, 1921), and J have here only 
certain remarks to make on them. 

Tt is easily seen that the sphenoid of Diplocercides agrees well 
with that of the Rhipidistids in most features, and there can 
be no doubt that the soft anatomy of the region occupied by 
this bone in Diplocercides in the main must also have shown 
great agreement with that of the corresponding region in the 
Rhipidistids. 

In Diplocercides (text-fig. 5) we can clearly see that the 
trigeminus branches proper or their roots, with the exception of 
the r. ophthalmicus profundus, which has its exit more forward, 
have pierced the cranial wall just postero-dorsally of the Mace 
pterygoid process and at the hinder border of the atisphenoid 
wulst, together with the r. ophthalmicus lateralis and n. buccalis 
lateralis (or their roots). From this place all these nerves have 


1260 DE. ERIK A. STENSIO: NOTES 


evidently run forward dorsally of the basipterygoid process. 
One of them, the n. ophthalmicus lateralis, has pierced the dorsal 
part of the alisphenoid wulst through an anteriorly running canal, 
perhaps accompanied by the r. ophthalmicus superficialis tri- 
gemini, while the other ones certainly had their passage more 
ventrally on the lateral side of the alisphenoid wulst. From the 

conditions in Pictyonostews one can conclude with rather great 
probability that in Diplocercides the vena jugularis also ought to 
have passed dorsally of the basipterygoid process. 


Text-figure 5 


Stemp.ear Patt alsph sje Fe 
i ea WAL (— ophth. tak, 
crocc . wT) ; mm ; vial 
hs Bs Ue een ale -cminrephie als 
BS —~—_ \ ae 0° 4 ‘ eli 
h 4 eon 
sissies ler) olf 
x--- Fe Bee co eteleey 
ot \ 
ose : 
i BS aoa 
oi ' ~-Epethm 
1b.@3 O ‘ %% “Pgh 


W, Wow Wye fen a.carint 
Diplocercides kayseri (V. Koenen). 


Neurocranium from the right side. Cartilage dotted, cartilage bones shaded. 2/1, 
From Stensié, 1922 (1922 a). 


FE, pethm, exethmoideo-preéthmoid ; Fr, frontal part of the dermal cranial roof ; 
Ol, occipito-labyrinth ossification; Pa.it, parieto-intertemparal : Psph, para- 
sphenoid; Sphk, sphenoid; Stemp.ext, supratemporo-extrascapular; a.car.int, 
ventral opening of the unpaired caual for the internal carotid ; alsph, alisphenoid 
wulst; ¢.z, canal for vessels to or from the nasal fossa (probably a vein canal) ; 
er.ep, eplotic crista; e7.oc, occipital crista; ¢, basipterygoid process; fev, 
anterior division of the myodome; for, canal probably for a dorsally bounded branch 
from the n. linez Jatevalis; dat.for, foramen probably for the anterior branches 
of the r. maxillaris trigemini and n. buccalis lateralis; olf, exit of the olfactory 
nerve in the nasal fossa; ophth.latz, anterior opening of the canal throngh the 
alisphenoid wulst for the n. ophthalmicus lateralis; ophth.lat3, posterior opening 
of an anterior canal for the n. ophthalmicus lateralis; II, opticus canal; III, oculo- 
motorius canal; IV, trochleavis canal; Vye, canal for ther. ophthalmicus profundus 
trigemini; VII, facialis canal; VI, probably the canal for the r. palatinus facialis 
or perhaps the canal for the abducens; IX ?, presumed position of the elosso- 
pharyngeus exit; X, canal for the n. vagus, n. line lateralis, and v. encephalica 
posterior. 


du Awelia we have a form which with regard to the position of 
the basipterygoid process agrees rather well with Diplocerciaes 
(Stensid, 1921, text-figs. 39, 40). Moreover, we have there a 
long well-developed homologue to the alisphenoid wulst of 
Diplocercides, and on the lateral surface of this homologue, which 


—— 


ON CERTAIN GROSSOPTERYGIANS. 1261 


forms a postero-ventral process from the fronto-dermosphenotie, 
there runs an antero-caudal sulcus, which certainly corresponds 
to the canal for the n. ophthalmicus lateralis in Diplocercides. 
These facts and the conditions in general in Awelia show that 
the trigeminus branches (with the exception of the r. ophthal- 
micus ‘profundus), the lateralis nerves accompanying them, and 
the vena jugularis ought all to have run dorsally of the basi- 
pterygoid process as in Diplocercides and the Rhipidistids. 

The basipterygoid process in Wimania and Maer ones though 
sibuated very high up, issues, however, just at the basis of the 
obvious homologue of the alisphenoid wulst and just below the 
level of the floor of the cranial cavity. It actually has the same 
position as in Diplocercides and Awelia, but the cranial basis 
below it has grown much deeper in these. There is thus no 
reason to believe that in Wimania and Macropomea the trigeminus 
branches, the lateralis branches accompanying these, and the 

. jugularis would have run in another way in relation to the 
basipterygoid process than in Alexia, Diplocercides, and the 
Rhbipidistids. 

Consequently the view put forward by Allis in a recent paper 
(1922, pp. 149-152) that the vena jugularis, the trigeminus 
branches, and the lateralis nerves accompanying these passed 
ventrally of the process termed the basipterygoid process by me 
seems untenable, as far as we can judge at present. It is 
absolutely certain that the basipterygoid process of my descriptions 
of the Celacanthids cannot be a postorbital process as Allis 
coneludes. A single glance at the conditions in Diplocercides and 
Aelia is sufficient to show the impossibility of this view. 

In his argument against my view of the basipterygoid process 
in the Celacanthids, Allis also says that as the hyoman- 
dibula in these fishes is evidently of the teleostome type, the 
jugular vein must have passed internal to it (Allis, 1922, p. 151; 
1915), and that this ee | cannot have run so high up 
as I have maintained, 7.e. it cannot have run dorsally of the 
basipterygoid process Te evidently bases his account (cf. Allis, 
1922, pp. 150, 151) in this point on Reis’s description, having 
probably overlooked my statements (Stensi6, 1921, pp. 114, i18— 
119) that the dorsal part corresponding to the hyomandibula of 
the iigoid arch in the Ceelacanthids is reduced, and that the 
hyomandibula of Reis’s description is simply a pari of some of 
the ossifications in the. primordial neurocranium. The epihyal, 
which in the fossil state of the Celacanthids forms the dorsal 
part of the hyoid arch, has been attached to the primordial 
neurocranium ‘only with ligaments. How far dorsally these 
hgaments extended it is impossible to say, but it seems to me 
most probable that they were rather short, and that the vena 
jugularis passed totally dorsally of them and thus dor sally of the 
hyoid arch, as it does in Selachians. Thus in this respect, 
too, the conditions do not support the view put forward by 
Allis. 


1262 DR. ERIK A. STENSIO: NOTES 


An account of the exact homologues of the basipterygoid 
process of the Celacanthids is given by me in connection with 
the description of the myodome of Sawrichthys (Stensid, 1922 6), 
and on the same occasion I also had the opportunity of dealing 
with the myodome of the Coelacanthids. As it will perhaps be 
a long time before I can get enough money to print my work on 
the Sauriechthyids, I have found it necessary to give here a 
extenso the considerations that I made there on the Ceelacanthids. 

“Tn the Celacanthids (Stensi6, 1921. pp. 53-62, 91-86, 120— 
122; 1922, pp. 177-184, 205-206) there has taken place, in 
connection with the considerable growth of the eyes, a compression 
and thinning from the sides of certain postero-ventral parts of 
the orbito-temporal region and an adjacent smaller part of the 


Text-figure 6. 


ke my fon S Gcar.ink 


Diplocercides kayseri (V. Koenen). 


Median sagittal section through the neurocranium. Cartilaginous parts restored. 
Cartilage dotted, cartilage bone shaded. 2/1. 


Fr, frontal part of the dermal cranial roof; OZ, occipito-labyrinth ossification ; 
Pa.it, pavieto-intertemporal; Psph, parasphenoid; Sph, sphenoid; a.car.int, 
unpaired canal for the internal carotid arteries; bo, basioccipital portion of the 
occipito-labyrinth ossification; ch, chorda space; cv, cranial cavity; fen, anterior 
division of the myodome; f-ryp, fossa hypophyseos; my, posterior division of the 
myodome; s, canal between the fossa hypophyseos and the myodome; so, supra- 
occipital portion of the occipito-labyrinth ossification. 


labyrinth region. These thinned parts have a vatker considerable 
fenestration close to the base of the primordial neurocranium, 
the orbits thus being in communication with each other at this 
place. The fenestration, the ventral boundary of which is formed 
by the parasphenoid, is situated ventrally of the fossa hypo- 
physeos, with which it has presumably communicated, and just 
posterior to the unpaired canal for the arteriz carotides interne, 
that ascends vertically through the basis cranil, entering the 
cranial cavity immediately anterior to the fossa hypophyseos. 


ON CERTAIN CROSSOPTERYGIANS. 1263 


The fenestration has thus approximately the same position as 
the canalis transversus of the Hlasmobranchil (canal for the 
pituitary vein), and must certainly be considered as such a canal, 
though it has become very much widened and in addition 
shortened in a transversal direction. From its posterior surface, 
which is somewhat wider than the anterior one, a short fossa 
penetrates in a backward direction between the base of the 
primordial neurocranium and the parasphenoid. To define it 
more closely, this fossa is so situated in the post-Devonian 
Ceelacanthids, that it is bounded dorso-caudally and caudally by 
the corpus of the basisphenoid and on either lateral side by the 
ventral process of the basisphenoid, while the ventral wall is 
formed by the parasphenoid. In the Devonian form Diplocer- 
cides khaysert (Stensid, 1922 a, pp. 178-179, 205-206) the fossa 
is sitnated in quite the same place as in the post-Devonian 
forms, but as there the basisphenoid is contained in and forms 
a postero-ventral part of the large sphenoid, it is conpeque ny, 
bounded there dorsally, caudally, and laterally by the pars basi- 
sphenoidea sphenoidei.”’ 

“‘ In fishes in general, as we know, the recti muscles of the eye 
have their place of origin postero-ventrally of the opticus and 
oculomotorius foramina or else postero-ventrally of a line that 
ean be drawn through these two foramina. In the Ccelacanthids 
the place where one must suppose the musculi recti to have 
originated is occupied by the widened transversal canal, and we 
can, therefore, scarcely help thinking that an expansion of this 
canal has been caused by the recti muscles. The recti externi 
have probably, as usual, invaded the transversal canal and then 
pushed postero-ventrally into the posterior wall of this canal, 
forming the short fossa described in this wall. The other three 
musculi recti have perhaps partly had their origin on some 
vertical membrane, which may have filled the anterior part ot 
the transversal canal; most probably, however, they ought to 
have originated for the most part from the dorsal surface of the 
parasphenoid on the sides of or in the transversal canal itself. 
Whatever the conditions may have been with regard to the 
origin of certain of the recti muscles, it seems, however, 
impossible not to suppose that in the expanded transversal canal 
and the fossa opening into this from behind we are concerned 
with a sort of myodome.” 

‘« Watson has recently (1921, p. 335) doubted this explanation 
of mine, which I put for ware in an incomplete form in the 
former part of this werk. He then laid stress especially on the 
fact that the space that would be regarded as a myodome could 
not under any circumstances be homologous with the myodome of 
the Paleoniscids, which of course is situated dorsally of the 
basipterygoid process and laterally of what he calls ‘ the body of 
the basisphenoid.’ It is of course true that the relation of the 
myodome to the hasipterygoid process is quite different in 
Paleoniscids and Cceelacanthids, but, as I believe I can show, 


1264 DR. ERIK A. STENSIO: NOTES 


this difference is due to the upward displacement of the 
articulation between the primordial neurocranium and the 
palato-quadrate in the Ceelacanthids (Stensié, 1922, pp. 205— 
206) from its original position at or near the base of the 
neurocranium.,” 

«For if we turn to Dictyonostews we find there a large basi- 
pterygoid process which certainly extends some distance upwards 
along the lateral surface of the neurocranium but which issues, 
however, also in part from the cranial base. This basipteryg coid 
paces really forms, if one will so express it, a very powerful 

‘wulst’ along the posterior edge of the sphenoid (cf. Stensio, 
Weal Te) 5G) where it begins immediately above the para- 
sphenoid—é. é. at the basis cranii—and continues upwards and 
somewhat backwards to about a third of the height of the 
neurocranium at this place, ending there with a deomenten upper 
end. On account of the oblique position of the process the 
anterior surface of this is directed antero-dorsally and the 
posterior one postero-ventrally. The process has in ¢ addition a 
dorsal surface and has certainly also had another one against 
which the metapterygoid probably articulated. This latter 
surface, which probably faced ventro-laterally and perhaps some- 
what posteriorly, is, however, not well preserved.” 

“The antero-dorsal surface of the basipterygoid process and 
the lateral wall of the part of the sphenoid just in front of the 
basipterygoid process in Dictyonosteus form together a sort of 
shallow antero-laterally and somewhat dorsally directed fossa at 
the inner end of which the well-developed canalis transversus 
opens. The vena jugularis has passed the opening of this canal 
and received the vena pituitaria from it, after which it has 
continued upwards and backwards in a very distinct sulcus on the 
boundary between the external] vertical surface of the sphenoid 
and the antero-dorsal surface of the basipterygoid process, until 
it reached the dorsal surface of the latter process, where 16 
presumably turned more straight backwards, situated all the 
time close to the lateral surface of the sphenoid. The arteria 
earotis interna must in its forward course have been situated 
on the outside of the basipterygoid process close to its base. At 
the anterior end of the basipterygoid process it must have turned 
in a medial direction, entering a sphenoid by a large foramen. 
The parasphenoid lacks the processus ascendens, but if this had 
been developed the arteria carotis interna would have passed 
between it and the basipterygoid process to its canal in the 
sphenoid just referred to.” 

‘Fiom the upper end of the basypterygoid process a ridge, 
which if not strong is still distinctly marked, passed upward and 
forward to the cranial roof; and just in front of this ridge, which 
T have called the alisphenoid ‘ wulst,’ we find at the top not far 
below the cranial roof the anterior opening of a canal, which, as 
far as one can judge (cf. Stensid, 1921, pp. 60, 93), must have 


ON CERTAIN CROSSOPTERYGIANS. 1265 


transmitted the n. ophthalmicus lateralis (ef also Stensid, 1921, 
p- 170) and lower, a short distance above the dorsal end of the 
basipterygoid process, the anterior opening of another canal, 
which seems to have transmitted their r. ophthalmicus profundus 
or some similar nerve.” 

“The main part of the trigeminus nerve, together with the 
lateralis nerves accompanying the trigeminus branches, has 
traversed the cranial wall rather high postero-dorsally of the 
basipterygoid process behind the alisphenoid wulst, and must 
then have run forward and a little downward dorsally of this 
process. The facialis exit must have been situated still somewhat 
farther backward, but at the same time certainly also more 
ventrally than the trigeminus exit. ‘he r. palatinus facialis 
has been thence given off in a forward and downward direction, 
and it probably came down behind the basipterygoid process, 
afterwards turning ina forward direction on the outside of the 
basal part of this process. In this part of its course the r. pala- 
tinus facialis ought to have been situated. at least for some 
distance, rather close to the arteria carotis interna. The 
n. abducens probably accompanied the trigeminus branches, and 
thus passed dorsally of the basipterygoid process, Finally, it 
may aiso be pointed out that the large fossa hyphphyseos reaches 
with its postero-ventral part into the part of the sphenoid 
situated between the basipterygoid processes.” 

‘From the facts stated, it is obvious that the formation in 
Dictyonostews that has hitherto been called the basipterygoid 
process must really in its antero-ventral part be homologous with 
a posterior larger part of the basipterygoid process in Birgeria, 
at the same time, however, also including semething more than 
this. It would have about its entire correspondence in Birgeria 
if we supposed there the myodome half of one side with the 
exception of the prepituitary part to be filled out with bone and 
this bone-mass forming a connected whole with the basipterygoid 
process forward and downward, and also with the basal parts of 
sphenoid on the medial side. In a similar way it would have 
its equivalent in Ama if in this fish we supposed the lateral 
pacts of the myodome of either side, apart from the prepituitary 
part, to be filled with bone or cartilage, but in this case of 
course it would correspond exactly only to the anterior part 
of such a bone or cartilage mass. In sharks it corresponds some- 
what but not quite exactly to a posterior part of what Allis 
(1914 @) ealls the ‘subocular shelf ’—together with an adjoining 
part of the capsula auditiva postero-ventrally of the vena jugu- 
lavis. The correspondence would in the latter case be almost 
complete if only these cartilaginous parts in the sharks were 
developed somewhat more in such a way as to form a lateral 
wall, to some extent more complete, for the fossa at the posterior 
eud of the orbit in which the canalis transversus opens ont, and 
in which the trigeminus ganglionic complex is also usually 


1266 DR. ERIK A. SPENSIO: NOTES 


situated (fossa trigemino-pituitaria, Allis, 1914a@). The basi- 
pterygoid process is Dictyonosteus thus comprises not only the 
homologue to the actual basipterygoid process, but also certain 
other parts of the cranial wall.” 

‘¢ Dorsally of the upper end of the basipterygoid process—if we 
may, for the sake of brevity, retain the term basipterygoid process 
for it—Dictyonosteus lacks every equivalent to the outer wall of 
the trigemino-facialis chamber in Birgeria, Amia, Lepidosteus, 
and Teleosts, and also to the postorbital process and the sphenotie. 
As also no processus ascendens parasphenoidei that might have 
covered this part of the cranial wall is present, the trigemino- 
facialis chamber is consequently absolutely lacking. If a myo- 
dome were developed in Dictyonostews this myodome would 
naturally be without lateral prespinal parts.” 

‘In Husthenopteron (Bryant, 1919, text-fig. 5; pl. ix. fig. 3)a 
basipterygoid process is found of essentially the same type as the 
one just described in Dictyonosteus. It only reached somewhat 
higher up than in the latter form. In a fossa between it and 
the vertical interorbital wall is situated the opening of the 
eanalis transversus in the same way as in Dictyonosteus. By 
its greater extension upwards the basipterygoid precess im 
Husthenopteron shows a certain tendency to develop in the same 
direction as in the Ceelacanthids. Fer, if we imagine its ventral 
part reduced or weakly developed, the remaining dorsal part 
would evidently come to correspond fairly closely to the basi- 
pterygoid process in the Ceelacanthids, as I have already pointed 
out in another work (Stensid, 1922 a, pp. 205-206). This means 
of course that the basipterygoid process of the Ccelacanthids 
would be homologous with at least a part of the septnm that 
separates the myodome from either trigemino-facialis chamber in 
Teleosts.” 

“This view of the basipterygoid processin the Ceelacanthids 
is also supported by the course of the nerves and the vessels. In 
Diplocercides kayseri, where the conditions are best known, we 
thus find the following conditions (Stensid, 1922q@). All the 
trigeminus branches except the r. ophthalmicus profundus must 
have passed forward dorsally of the basipterygoid process, as has 
the vena jugularis on its way backward after having received 
the v. pituitaria, which traversed the anterior part of the 
myodome from side to side. The vr. palatinus facialis must in 
all probability on its way forwards and downwards have run 
just postero-ventrally of the so-called basipterygoid process, and 
probably reached down to the level cf the cranial hase just at 
the myodome. The arteria carotis interna must have had its 
course far ventrally of the so-called basipterygoid process, and 
has, as we have seen, entered the primordial neurocranium at 
the anterior end of the myodome, where it turned directly 
upwards in an unpaired canal which opened into the cranial 
cavity immediately behind the exits of the optic nerves. With 
regard to the n. abducens, it is true that as yet nothing certain 


ON CERTAIN CROSSOPTERYGIANS, 1267 


is known about it, but presumably it accompanied the trigeminus 
branches.” * 

“The myodome in the Cceelacanthids as described here has 
pro-otic, pituitary, and prepituitary parts (cf Allis, 1919). The 
pro-otic part of it 1s represented by the fossa situated between 
the basisphenoid (pars Paste pienotilca of the sphenoid in Diplo- 
cercides) and the parasphenoid, The pituitary part is situated 
beneath the fossa hypophyseos, and is separated from this fossa 
by a roof of bone pierced only by a small opening for the 
hypophysis. The prepituitary part is very short, and is only 
homologous with the median portion of the prepituitary part of 
the myodome in the Teleosts. Homologues to the lateral portions 
of the prepituitary part of the ay odome in the Teleosts are 
sory lacking, even as potential.” 

‘As at least in the fossil state of preservation the myodome 
in the Celacanthids is bounded directly by the parasphenoid on 
the ventral side, it might be supposed to consist of both a dorsal 
and a ventral compartment, like the myodome in the Teleosts. 
If this were actually the case, the ventral compartment must, 
however, at least in certain species (Amelia, Stensio, 1921, p. 95), 
have been very short and limited only to the anterior end of the 
myodome, as the parasphenoid in these species (Awelia) beneath 
the middle and posterior parts of the myodome undoubtedly 
seems to comprise parts ossified in the ventral wall of the 

rimordial neuroeranium, and the parts so ossified there form 
fae floor of the myodome. oae 
“The myodome in the Ceelacanthids can easily be thought to 
have arisen from the conditions in the Rhipidistids in the 
following way. If in the Rhipidistids the postero-ventral parts 
of the orbitotemporal region and the adjacent part of the 
labyrinth region became much compressed from the sides and 
thinned by an increase in the size of the eye-balls, the canalis 
transversus would of course be very much shortened. If, then, 
the musculi recti externi invaded the opening of this shortened 
eanal, they would rather soon meet with their origins in the 
median line, ¢ causing a considerable widening of the whole canal. 


* Tn my paper on the Devonian Celacanthids from Wildungen (Siensid, 1922 a 
p. 183), I was of the opinion that the n. abducens in Diplocercides ought to have 
run postero-ventrally of the so-called basipterygoid process of the fish. The opinion 
maintained in the present work seems, however, more in agreement with the general 
conditions than my earlier one. 

+ “From the conditions described here in Dictyonosteus it is evident that the 
objections made by Allis, 1922 (pp. 149-152 ) against my interpretation of certain 
parts of the neurocranium in he CGcelacanthids must be untenable. There can be 
no doubt that the dorsal part of the basipterygoid process of Dictyonosteus is homo- 
logous with the corresponding part of the process of Husthenopteron and the whole 
process of the primitive Ceelacanthids (Diplocercides), and that accordingly also the 
jugular vein in this form has run dorsally of the process as it has done in Dietyo- 
nosteus. Among the post-Devonian Ceelacanthids we have in 4.vrelia a form which 
with regard to the j position of the basipterygoid process shows rather intermediate 
conditions ‘yelinreern the primitive Celacanthids (Diplocercides) and Wimania, in 
which latter the process issues very high above the cranial base.” 


1268 DR. ERIE A. SLENSIO: NOTES 


Being forced to shift their origins still farther backwards by the 
continued growth of the eye-bulbs, the recti externi muscles may 
be imagined to have pushed backwards into the posterior wall of 
the transversal canal, forming a fossa there opening in front 
into the transverse canal. Then, perhaps chiefly owing to the 
influence of the other recti muscles, a widening of the transverse 
canal forward may be supposed to have taken place at the 
expense of the bone part between this canal and the vertically 
ascending canal for the internal carotid arteries. Finally, the 
transverse canal, by the influence of the recti muscles, grew 
wider downwards as well, the floor of the canal belonging to the 
primordial neurocranium at the same time becoming much 
reduced, and finally perhaps forming only a membrane on the 
dorsal surface of the parasphenoid. The recti inferiores, pos- 
teriores, and superiores can after this have been able more or less 
easily to break through the membrane with their origins in 
seeking a firm attachment on the parasphenoid, and if: this had 
happened a ventral myodomic compartment would have arisen. 
That a myodome arisen in the way sketched here from the con- 
ditions in the Rhipidistids must lack homologues of the lateral 
portions of the prepituitary parts of the myodome in birgeria, 
Ama, and Teleosts,as the myodome in the Celacanthids actually 
does, is evident from what has been put, forward above concerning 
the Rhipidistids.” 

‘‘ In this connection finally it ought alsoto be pointed out that 
the occurrence of a myodome in the Ccelacanthids shows with full 
cer tainty that a myodome can easily develop, and that the myo- 
dome in several groups of fishes has been developed independently.” 

‘The myodome in the Ceelacanthids, if it has no ventral 
compartment, would, as is easily understood, correspond fairly 
well to the myodome in S. ornatus if the latter were extended 
forward by reduction of the larger posterior part of the bone- 
part bsph.” 

If we summarize here the results obtained by the investigation 
of the Ceelacanthids with regard to the basipterygoid process, we 
find the following. The basipterygoid process of the primitive 
ancestors of the Ceelacanthids must have been rather high, 

xtending upwards along the lateral side of the primordial 
neurocranium as in Dictyonosteus, and the metapterygoid probably 
articulated against the upper parts of this process. Through 
reduction of the ventral parts of the process the conditions 
which occur in Diplocercides seem to have arisen, and from these 
the evolution towards the post-Devonian Ceelacanthids ought to 
nage proceeded in such a way that the parts of the cranial basis 

below the remains of the basipterygoid process grew very deep, 
while at the same time the height of the parts ‘dorsally of this 
process decreased. 

in Diplocercides the sphenoid, like that of the Rhipidistids, 
comprises paired orbitosphenoid and alisphenoid components and 
an unpaired basisphenoid component. In the post-Devonian 


ON CERTAIN CROSSOPLERYGIANS. 1269 


forms, on the other hand, there are found merely posterior parts 
of the basisphenoid and alisphenoid components, while the 
anterior parts of these components, together with the orbito- 
sphenoid component, have been reduced and replaced by 
cartilage. 

In JMacropoma the part present of the alisphenoid component 
of either side is continuous with the hasisphenoid component, thus 
having there its original relation to the latter component (Watson, 
1921, p. 323). In Avelia and Wimainia, on the other hand, the 
alisphenoid component of each side has coalesced with and forms 
a descending process from the fronto-dermosphenotic of its side. 
These different conditions of the alisphenoid component in dif- 
ferent forms are of very great interest, as they show that a part 
of a substitution bone may rather easily become independent and 
coalesce with other neighbouring bones, even dermal ones (cf. 


Stensio, 1921, pp. 153-161, 181-182; Allis, 1899, 1909). 


BIBLIOGRAPHY. 


ALLEN, W. F. 1905. The Blood-vascular System of the Loricati, 
the mail-cheeked fishes. Wash. Acad. of Sci. Proc. 
vol. vii. 

Aus, H. Po. 1889. The Anatomy and Development of the 
Lateral Line System in Ama calva. Journ. of Morph. 
vol. i1. 

——. 1897. The Cranial Muscles and Cranial and First Spinal 
Nerves in Ama calva. Journ of Morph. vol. xii. 

——. 1899. On Certain Homologues of the Squamosal, Inter- 
ealar, Exoccipital, and Extrascapular Bones of Amia 
calva. Anat. Anz., Bd. xvi. 

——. 1900. The Lateral Sensory Canals of Polypterus bichir. 

. Anat. Anz., Bd. xvii. 

——. 1902. The Lateral Sensory Canals, the Eye-muscles, and 
the Peripheral Distribution of certain of the Cranial 
Nerves of Mustelus levis. Quart. Journ. Mier. Sci. 
vol. xlv. 

——. 1903. The Skull, and the Cranial and First Spinal 
Muscles and Nerves in Scomber scomber. Journ. of 
Morph. vol. xvii. 

——. 1905. The Latero-sensory Canals and Related Bones in 
Fishes. Intern. Monatsschrift f. Anat. u. Physiol., 
Bd. xxi. 

——. 1908. The Pseudobranchial and Carotid Arteries in 
Ameiurus. Anat. Anz., Bd. xxxii. 

——. 1909. The Cranial Anatomy of the Mail-cheeked Fishes. 
Zoologica, vol. xxil. 

——. 19lla. “The Pseudobranchial and Carotid Neen ice in 
Polyodon spathula. Anat. Anz., Bd. xxxix. 

—-, 191106. The Pseudobranchial and Carotid Arteries in 
Chlamydoselachus anguineus. Anat. Anz., Bd. xxxix. 


Proc. Zoou. Soc.— 1922, No. LX XXIV. 84 


1270 DR. ERIK A. STENSIO: NOTES 


Auuis, #.Pa. 1912a, The Pseudobranchial and Carotid Arteries 
in Hsox, Salmo, and Gadus, ete. Anat. Anz., Bd. xl. 

——, 19126. The Branchial, Pseudobranchial, and Caxotid 
Arteries in Heptanchus. Anat. Anz., Bd. xli. 

—---. 1914. The Pituitary Fossa and Trigemino-facialis Cham- 
ber in Selachians. Anat. Anz., Bd. xlvi. 

—-. 1915. The Homologies of the Hyomandibula_of the 
Gnathostome Fishes. Journ. Morph. vol. xxvi. 

——. 1918. On the Origin of the Hyomandibula of the 
Teleostomi. Anat. Record, vol. xv. 

——.,. 1919a. The. Myodome and Trigemimo-facialis Chamber 
of Fishes. Journ. of Morph. vol. xxxu. 

——. 19196. The Homologies of the Squamosal Bone of Fishes. 
Anat. Reeord, vol. xvii. 

——, 1922. The Myodome and the Trigemino-facialis Chamber. 
in the Ceelacanthidee, Rhizodontide, and Paleoniscide, 
Journ. of Anat. vol. lvi. 

Bine, R., & Burcxuarpy, R. 1905. Das Centralnervensystem 

von Ceratodus forsteri. Jena, Med.-Nat. Gesellsch. 
Denkschriften, Bd. iv. 

Bryant, W. C. 1919. On the Structure of Husthenopteron. 
Buffalo Soc. Nat. Sci. Bull. vol. xui. 

Coin, F. J. 1898. Observation on the Structure and Morphology 
of the Cranial Nerves and Lateral Sense-organs of 
Fishes: with special reference to the genus Gadus. 
Trans. Linn. Soc. London, 2nd Ser., Zool. vol. vii. 

Danrorra, C. H. 1912. The Heart and Arteries of Polyodon. 
Journ. Morph. vol. xxiu. 

Easrman, Cu. R. 1917. Fossil Fishes in the Collection of the 
United States National Museum. U.S. Nat. Mus. Proce. 
vol. Jil. 

Gauer, E. 1905. Die Entwicklung des Kopfskelettes. In 
Hertwig’s Handbuch der Entw.-Lehre, Bd. mn: 2, 
1901-1906. 

GoopricH, E. 8. 1919. Restoration of the Head of Osteolepis. 
Journ. Linn. Soc. London, Zool. vol. xxxiv. 

Herrick, C.J. 1899. The Cranial and First Spinal Nerves of 
Menidia, ete. Journ. Comp. Neurol. vol. ix. 

——. 1901. The Cranial Nerves and Cutaneous Sense-organs 
of the North American .Siluroid Fishes. JZb¢dem, 
vol, Xi. 

JAEKEL, O. 1911. Die Wirbeltiere. Berlin. 

Panppr, C. H. 1860. Uber die Saurodipterinen, Dendrodonten, 
Glyptolepiden wnd Cheirolepiden des devonischen 
Systems. St. Petersburg, 1860. 

Peurson,'T. 1922. Some Poimts in the Cranial Development 
of Teleostomian Fishes. Stockholm, Acta Zoologica, 
Texel, rob, 

Rets, O. M. 1888. Die Celacanthinen mit besonderer Beriick- 
sichtigung der im Weissen Jura Bayerns vorkommenden 
Arter. Paleontographica, Bd. xxxv. 


ON CERTAIN CROSSOPTERYGIANS. Oval 


Srensro, EK. A:son. 1918a. Zur Kenntnis des Devons und des 
Kulms an der Klaas Billenbay, Spitzbergen. Upsala, 
Geol. Inst. Bull. vol. xvi. 

——. 19186. Notes on a Crossopterygian Fish from the Upper 
Devonian of Spitzbergen. Upsala, Geol. Inst. Bull. 
vol. xvi. 

——. 1921. Triassic Fishes from Spitzbergen. Part I. Vienna. 

——. 1922a@. Uber zwei Ceelocanthiden aus dem Oberdevon 
von Wildungen. Paleontologische Zeitschr., Bd. iv. 

——. 19220. Triassic Fishes from Spitzbergen. Part II. (In 
manuscript. ) 

Swinnerton, H. H. 1902. A Contribution to the Morphology 
of the Teleostean Head Skeleton, based upon a study of 
the Three-spined Stickleback (Gasterosteus aculeatus). 
Quart. Journ. Mier. Sei., New Ser. vol. xlv. 

Warson, D. M.S. 1921. On the Celacanth Fish. Ann. & Mag. 
Nat. Hist., ser. 9, vol. viii. 

Woopwarp, A. 8. 1891. The Devonian Fish-fauna of Spitz- 
bergen. Ann. & Mag. Nat. Hist., ser. 6, vol. vili. 

Workman, 1.8. 1900. The Ophthalmic and Eye Muscle-nerves 
of the Catfish. Journ. of Comp. Neurol. vol. x. 


EXPLANATION OF THE PLATE. 


Fig. 1. Dictyonosteus arcticus Stensié. Anterior half of the neurocranium in 
lateral view. ‘The lateral parts of the ethmoidal region and of the cranial 
roof removed. 1/2. 

Fig. 2. Dictyonosteus arcticus Stensid. Same parts as in the preceding figure, but 
with the basipterygoid process removed to show the canal for the pituitary 
vein and the fossa hypophyseos. 1/2. 

Fig. 3. Dictyonosteus arcticus Stensid. Sphenoid from behind. 1/2. 

Fig. 4. Dictyonosteus arcticws Stensio. Antero-lateral view of the posterior half 
of the sphenoid. N.B. The very distinct sulcus for the jugular vein (s/), 
and the opening of the canal for the pituitary vein (v.pit). 1/2. 

Fig. 5. Porolepis ?sp. Anterior part of the dermal cranial roof, showing the 
sensory canals. 


The type-specimen of figs. 1-4 belongs to the Paleontological Institution, Upsala 
the type-specimen of fig. 5 to the Paleontological Museum, Christiania. 


Index letters. 


H.pethm, exethmoideo-preéthmoid; Fr, frontal; Psph,.parasphenoid; Sph, 
sphenoid ; «@.car.int, canal for the arteria cavotis interna; alsph, alisphenoid 
wulst; bp, basipterygoid process ; en, orbito-nasal canal for vessels to or from, or 
both to and from the nasal pit; c.ophth.lat, through the dorsal part of the 
alisphenoid wulst for the n. ophthalmicus lateralis; cv, cranial cavity; f-hyp, 
fossa hypophyseos ; tfc, infraorbital sensory canal ; s7, sulcus for the vena jugularis ; 
soc, supraorbital sensory canal; v cerchr.ant, canal for the anterior cerebral vein ; 
v.pit, canal for the pituitary vein; I, olfactory canal; II, opticus canal; III, 
oculomotorius canal ; IV, trochlearis canal; V, trigeminus exit ; Vj, canal for the 
r. ophthalmicus profundus,. 


THE SECRETARY ON ADDITIONS TO TIE MENAGERIE. 1273 


EXHIBITIONS AND NOTICES. 
October 24th, 1922. 


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


The Secrerary read the following Report on the Additions to 
the Society’s Menagerie during the months of June, July, 
August, and September, 1922 ;— 


JUNE. 


The registered additions to the Society's Menagerie during the 
month of June were 157 in number. Of these 75 were acquired 
by presentation, 34 were deposited, 26 were purchased, and 22 
were born in the Menagerie. 

The following may be specially mentioned :— 

1 Striped Hyena (Hyena hyena), from Kaduna, Nigeria, 
presented by Lt. L. 8. Clinton on June 20th. 

3 Cape Hyraxes, born im the Menagerie on June 19th. 

1 Greater Amethyst Sun-bird (Chalcomitra amethystina) and 
1 Malachite Sun-bird (Nectarinia famosa, from South Africa, 
presented by Mis. George Robey on June 6th. 

1 Apure Tovi Parrakeet (Brotogerys jugularis apurensis), from 
Apure, Venezuela, new to the Collection, presented by Mons. 
J. Delacour, F.Z.S., on June 12th. 

6 Pink-winged Rose-Finches(fhodospiza obsoleta), from Persia, 
presented by Alfred Hzra, O.B.E., F.Z.S., on June 12th. 

1 White-crested Hornbill (Ortholophus leucolophus), from 
Coomassie, West Africa, new to the Collection, presented by 
Major Harold W. Sidley on June 12th. 

A collection of Indian Snakes, including 2 Hamadryads 
(Waia bungarus), presented by Alfred Ezra, O.B.E., F.Z.8., on 
June 12th. 

A Tiger-Fish (Carapus fasciatus), from Brazil, new to the 
Collection, purchased on June 15th. 


JULY. 


The registered additions to the Society’s Menagerie during the 
month of July were 229 in number. Of these 120 were 
acquired by presentation, 17 were deposited, 31 were purchased, 
' 1] was received in exchange, and 60 were born in the Menagerie. 

The following may be specially mentioned :— 

1 Tiger (felis tigris), § , from Bhopal, Central India, presented 
by H.H. Raja Sir Birandra Singh, K.C.1.E., on July 13th. 

1 Kiang (Zquus kiang), 2, born in the Menagerie on July 9th. 

2 Guinea Baboons (Papio papio), 1 African Civet (Civettictis 
cwetta), 2 African Poreupines (Hystria africe-australis), 2 
Harnessed Antelopes (Zragelaphus scriptus), 1 Saddle-billed Stork 
(Hphippiorhynchus senegalensis) from Gambia, and 3 Ostriches 


1274 PHE SECRETARY ON ADDITIONS TO THE MENAGERIE, 


(Struthio camedus) from Senegal: presented by H.H. Capt. C. H. 
Armitage, C.M.G., D.S.O., on July 18th. 

1 Chestnut-capped Buarremon (Buarremon bruwneinucha), 
from Southern Mexico, new to the Collection, purchased on 
July 25th. 

| White-bellied Amethyst Starling (Pholidauges leucogaster), 
2 Broad-tailed Babblers (Crateropus platycercus), presented by 
Dr. E. Hopkinson, C.M.G., D.S.0., on July 17th. 

A Grooved Tortoise (Z'estudo calcarata), bred in the Menagerie 
on July 16th. 

A West-African Chameleon (Chameleon gracilis), new to the 
Collection, presented by H.H. Capt. C. H. Armitage, C.M.G., 
D.8.0., on July 18th. 


AUGUST. 


The registered additions to the Society’s Menagerie during the 
month of August were 187 in number. Of these 102 were 
acquired by presentation, 42 were deposited, 25 were purchased, 
3 were received in exchange, and 15 were born in the Menagerie. 

The following may be specially mentioned :— 

1 Brazilian Tapir (Vapirus terrestris), from Brazil, deposited 
on August 29th. 

16 Prairie-Marmots (Oynomys ludovicianus), from North 
America, purchased on August 22nd. 

2 Pocket-Gophers (Thomomys bulbivorus), from San Francisco, 
new to the Collection, deposited on August 15th. 

3 Greater Amethyst Sun-birds (Chalcomitra amethystina), from 
Durban, presented by A. M. Campbell, F.Z.8S., on August 13th. 

1 Verreaux’s Amethyst Starling (Pholidauges leucogaster 
verreauxi), from Durban, presented by Harold Millar, U.M.Z.5., 
on August 13th. 

A young Wart-Hog (Phacochwrus africanus) and a coilection 
of Reptiles, including 2 Green Tree-Snakes (Ohlorophis irregularis) 
and a Skink (Chalcides, sp. nova), both new to the Collection, 
from Gambia, presented by H.H. Capt. C. H. Armitage, C.M.G., 
D.S8.0., F.Z.8., on August 15th. 

1 Mamba (Dendraspis angusticeps), new to the Collection, from 
South Africa, purchased on August 11th. 

13 Buerger’s Tree-Frogs (Rhacophorus buergeri), new to the 
Collection, from Japan, presented by Mr. Tomisaburo Nagai. 

2 Sucker-Fish {Plecostomus commersoni), new to the Collection, 
from South America, purchased on August 31st. 


SEPTEMBER. 

The registered additions to the Society’s Menagerie during the 
month of September were 214 in number. Of these 87 were 
acquired by presentation, 17 were deposited, 62 were purchased, 
19 were received in exchange, and 29 were born in the Menagerie. 

The following may be specially mentioned :— 

2 Musk-Oxen (Ovibos moschatus), $ 2, from Greenland, 
purchased on September 19th. 


THE SECRETARY ON ADDITIONS TO THE MENAGERIE. WAT 


1 Tigress (Felis tigris), from Perak, presented by W. H. Jones, 
ae on September 18th. 

2 Cheetahs (Cynelurus jubatus), from Nairobi, deposited on 
Sei tom ber 25th. 

1 Nagor Antelope (Redunca nagor), 1 Gambian Oribi (Ourebsa 
nigricaudata), 3 Petas Cercopitheques (Cercopithecus putas), 
3 Nilotic Monitors (Varanus niloticus), 1 West-African Chame- 
leon (Chameleon gracilis), 1 Hissing Sand-Snake (Psammophis 
sibilans), and 1 Senegal River Turtle (Cyclanorbis senegalensis), 
from Gambia, presented by H.E. Capt. C. H. Armitage, C.M.G., 
D.S.O., EZS.. on September 19th. 

1 Spectacled Owl (Pulsatrix perspicillata), from South America, 
purchased on September 18th. 

1 Rock-Buntine (Lringillaria tahapizi), from South Africa, 
new to the Collection, presented by the Hon. Mrs. Charles Winn 
on September 26th. 

A large collection of N.-American Snakes, including 2 Copper 
Heads (7; Ineisirodon coniortriz), « Confluent Rattlesnake (Crotalus 
confluentus), and a Horned Rattlesnake (Crotalus horridus), 
presented by Dr. Howard A. Kelly on September 11th. 

2 Puff-Adders and a Reinhardt’s Snake (Prosymnu meleagris), 
the latter new to the Collection, from Mongalla, Sudan, presented 
by the Rev. A. Shaw, C.M.G., on September 2nd. 


The Sucrevary exhibited, and made remarks upon, two photo- 
graphs of Zebras and Oryx from Kenya. 
Mr. EK. T. Newton, F.B.S., F.Z.5., exhibited, and made remarks 
upon, the tanned skin of a Frog. 
b) co} 


Mr. ©. Tate Reean, F.R.S., F.Z.S., exhibited, and made 
remarks upon, some specimens of leather made from Sharks’ 
skins. 


November 7th, 1922. 
- Prof. E. W. MacBripz, F.R.S., Vice-President, 


in the Chair. 


Prof. G. Exiior Smits, M.D., F.R.S., F.Z.S., exhibited, and 
made remarks upon, a series of photographs of a fossil tooth 
(Hesperopithecus) trom Nebraska. 


Dr. A. Smira Woopwarp, F.R.S., F.Z.8., exhibited, and made 
remarks upon, drawings and photographs comparing teeth of 
a Bear, Chimpanzee, and Hesperopithecus. 


1276 THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 


November 21st, 1922. 


Sir S. F. Harmer, K.B.H., F.R.S., Vice-President, 
in the Chair. 


The Srcrerary read the following Report on the Additions to 
the Society's Menagerie during the month of October, 1922 :— 


The registered additions to the Seciety’s Menagerie during the 
month of October were 263 in number. Of these 172 were 
acquired by presentation, 40 were depesited, 21 were purchased, 
8 were received in exchange, and 22 were born in the Menagerie. 

The following may be specially mentioned :— 

2 American Bisons (Bison americanus), 5 2, from Canada, 
presented by the Canadian Government on October 23rd. 

1 Pygmy African Elephant (Zovodon cyclotis), 2 , from French 
Gaboon, deposited on October 16th. 

1 Black-throated Saltator (Saltator artricollis), from Bahia, 
Brazil, new to the Collection, purchased on October 26th. 

7 Cream-coloured Coursers (Cursorius gallicus), from Egypt, 
new to the Collection, purchased on October 4th. 

2 Black Storks (Ciconia wigra), from Poland, presented by 
H.8.H. Prince Charles Radziwill on October 25th. 

1 Kea Parrot (Vestor notabilis), from New Zealand, presented 
by Lady Ursula Abbey on October 4th. 

49 Indian Fruit-Bats (Pleropus medius) and a collection of 
Indian Reptiles, inciuding 2 Cobras, a Russell’s Viper, and a 
Banded Krait, presented by A. Ezra on October 7th. 

A collection of rare tropical fish including several species new 
to the Collection, presented by J. R. Shattock on October 8th. 


Prot, E. W. MacBripg, F.R.8., F.Z.S8., exhibited, and made 
remarks upon, a series of lantern-slides bearing upon Dr. Kam- 
merer’s experiments on Amphibia. 


Dr. A. Smita Woopwarb, F.R.S., F.Z.S., exhibited, and made 
remarks upon, askull and tusks of a Mammoth from Siberia, and 
drew attention to the remarkably perfect and fresh condition of 
the specimen. 


Mr. D. Sersa-Smuiry, F.Z.8., exhibited, and made remarks 
upon, the shed lining of the gizzard of a Hornbill. 


No. 223. 


ABSTRACT OF THE PROCEEDINGS 


OF THE 


ZOOLOGICAL SOCIETY OF LONDON.* 


February 7th, 1922. 


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


The Srecrerary read a Report upon the Additions to the 
Society's Menagerie during the months of November and 
December, 1921. 


Mr. A. H. Evans, F.Z.8., exhibited, and made remarks upon, 
a series of Cuckoos’ eggs taken near Cambridge. 


Mr. C. W. Hosiery, C.M.G., C.M.Z.S., gave a résumé of his 
paper on “The Fauna of Hast Africa and its Future,” and drew 
special attention to the urgent need for immediate action being 
taken, if the rapidly disappearing herds of Big Game are to be 
preserved from total extinction. 


Lord Ciirrorp oF CuupieIcH, F.Z.S., exhibited, and made 
remarks upon, a series of photographs of Vototheriuwm mutchelli. 


Miss L. E. Currsman, F.E.S., gave an account of the position 
and function of the Siphon in the Amphibious Molluse Ampul- 
laria vermiformis. 


Mr. G. C. Rosson, F.Z.S., exhibited, and made remarks upon, 
a series of models demonstrating the respiratory mechanism of 
Ampullaria verniformis. 


* This Abstract is published by the Society at its offices, Zoological Gardens, 
Regent’s Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Staxpence, or, if desired, sent post-free for 
the sum of Sta Shillings per annum, payable in advance. 


14 


In the absence of Lt.-Col. J. SrepHmnson, D.Sc., F.Z.8., his 
communication ‘ Contribution to the Morphology, Classification, 
and Zoogeography of Indian Oligocheeta : 


IV. On the diffuse Production of Sexual Collen in a Species 
of Chetogaster (Fam. Naidide). 


V. On Drawida japonica (Michlsn.), a Contribution to the 
Anatomy of the Moniligastridee. 

VI. On the Relationships of the Genera of Moniligastride ; 
with some Considerations on the Origin of Terrestrial Oligo- 
cheeta,” 

was taken as read. 


The next Meeting of the Society for Scientific Business will 
be held on Tuesday, February 21st, at 5.80 p.m., when the 
following communications will be made :— 


R. Broom, D.Sc., F.R.S., C.M.Z.S. 
On the Temporal Arches of the Reptilia. 
H. Burevap, Ph.D. 
Animal Communities in the Southern North Sea. 


Cuas. F. Sonnrac, M.D., F.Z.S. 
(1) On the Vagus and Sympathetic Nerves of the Hdentata. 


(2) On the Vagus and Sympathetic Nerves of Myrax 
capensis. 


C. Tate Reean, F.R.S., F.Z.S. 
The Cichlid Fishes of Lake Victoria. 


The following Papers have been received :— 
F. Batrour Browne, M.A., F.Z.S. 
The Life-history of the Water-Beetle Pelobius tardus Herbst. 


15 
EKENDRANATH GuosH, M.D., M.Sc., F.R.M.S. 


On the Animal of Scaphula Benson, with the Description of 
a new Species. 


F. W. Uricu, Huex Scorr, M.A., D.Sc., and J. WaTErRston, 
IDISGoq 18 Acts). 


Note on the Bat-Parasite Cyclopodia greeffi, and on a new 
Species of Hymenopterous (Chalcid) Parasite bred from it. 


S. K. Monreomery, B.A., B.Sc. 


Direct Development in a Dromiid Crab. 


The Publication Committee desire to remind Fellows 
that it is only by courtesy that they are allowed to 
bring guests to a Scientific Meeting. For the present 
no Fellow can be permitted to bring more than one 
guest to a Meeting. 


The Publication Committee desire to call the attention of 
those who propose to offer Papers to the Society, to the great 
increase in the cost of paper and printing. This will render it 
necessary for the present that papers should be condensed, and 
be limited so far as possible to the description of new results. 


Communications intended for the Scientific Meetings should 
be addressed to 


P. CHALMERS MITCHELL, 
Secretary. 


ZooLoeicaL Socrery or Lonpon, 
ReceEnt’s Park, Lonpon, N.W. 8, 
February 14th, 1922. 


No. 224. 


ABSTRACT OF THE PROCEEDINGS 


OF THE 


ZOOLOGICAL SOCIETY OF LONDON.* 
February 21st, 1922. 


Prof. E. W. MacBripsz, D.Sc., LL.D., F.R.S., Vice-President, 
in the Chair. 


The Srecrerary read a Report on the Additions to the 
Society’s Menagerie during the month of January, 1922. 


The Secretary exhibited, and made remarks upon, a photo- 
graph of the Society’s Gardens taken from an aeroplane. 


Miss L. E. Curusman, F.E.S., exhibited specimens of the 
Amphibious Molluse, Ampullaria vermiformis, and described the 
position and function of the siphon. 


Mr. G. C. Rosson, F.Z.S., exhibited a series of models demon- 
strating the respiratory mechanism of Ampullaria vermiformis. 


Mr. F. Martin Duncan, F.R.M.S., F.Z.8., exhibited a series 
of cinematograph films showing the movements of Ampullaria 
vermiformis ; of the Fresh-water Crab, Cardisoma armatum ; and 
of the Hawk’s-bill Turtle, Chelonia imbricata. 


Miss L. E. Cuzssman, F.E.S., described the habits, in captivity, 
of the Fresh-water Crab, Cardisoma armatum. 


* This Abstract is published by the Society at its offices, Zoological Gardens, 
Regent’s Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ‘Proceedings,’ free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Stxpence, or, if desired, sent post-free for 
the sum of Six Shillings per annum, payable in advance. 


18 


Dr. E. 8. Russeuzt, F.Z.8., communicated a paper by Dr. H. 
BLEGVAD on “* Animal Communities in the Southern North Sea.” 


My. C. Tare Reaan, F.R.S., F.Z.8., gave a réswiné of his paper 
on “'The Cichlid Fishes of Lake Victoria.” 


Dr. Cuas, F. Sonnrae, F.Z.S., gave a résumé of his papers “ On 
the Vagus and Sympathetic Nerves of the Edentata” and 
“On the Vagus and Sympathetic Nerves of Hyrax capensis.” 


The next Meeting of the Society for Scientific Business will be 
held on Tuesday, Mareh 7th, 1922, at 5.30 v.m., when the following 
communications will be made :— 


Nate LOO) Mb IANS lesuncllog is io ioe Sieetethy, 


Report on the Deaths which occurred in the Society’s 
Gardens during 1921. 


F. Batrour Brownz, M.A., F.Z.S. 
The Life-history of the Water-Beetle Pelobius tardws Herbst. 


On the Temporal Arches of the Reptilia. 


F. W. Unico, Hueuw Scorr, M.A., D.Sc., and J. WATERSTON, 
D.Sce., F.Z.S. 


Note on the Bat-Parasite Cyclopodia greeffi, aud on a new 
Species of Hymenopterous (Chalcid) Parasite bred from it. 


Direct Development in a Dromiid Crab. 


The following Papers have been received :— - 


GILBERT BLAINE, F.Z.S. 


Notes on the Zebras and some Antelopes of Angola. 


R. I. Pococr, F.R.S., F.Z.S. 


On the External Characters of some Hystricomorph Rodents. 


H. R. Hoae, M.A., F.Z8. 


Some Spiders from South Annam. 


The Publication Committee desire to remind Fellows 
that it is only by courtesy that they are allowed to 
bring guests to a Scientific Meeting. For the present 
no Fellow can be permitted to bring more than one 
guest to a Meeting. 


The Publheation Committee desire to call the attention of 
those who propose to offer Papers to the Society, to the great 
increase in the cost of paper and printing. ‘This will render it 
necessary for the present that papers should be condensed and 
be limited so far as possible to the description of new results. 


Communications intended for the Scientific Meetings should 
be addressed to 


P. CHALMERS MITCHELL, 
Secretary. 
ZOOLOGICAL Society OF LONDON, 
ReceEnt’s Park, Lonpon, N.W. 8. 
Vebruary 28th, 1922. 


No. 225. 


ABSTRACT OF THE PROCEEDINGS 


OF THE 


ZOOLOGICAL SOCIETY OF LONDON.* 


March 7th, 1922. 


Sir Srpney F,. Harmer, K.B.E., F.R.S., Vice-President, 
in the Chair. 


Mr. C. Tate Reean, F.R.S., F.Z.S., exhibited, and made re- 
marks upon, some living specimens of the Indian Tortoise- 
Beetle Aspidomorpha sancte-crucis. 


Mr. N.S. Lucas, M.B., F.Z.8., read a Report on the Deaths 
which had occurred in the Society’s Gardens during 1921. 


Prof. D. M. 8. Warson, F.Z.S., gave a résumé of Dr. R. 
Broom’s paper ‘“‘ On the Temporal Arches of the Reptilia.” 


Dr. J. Warerston, F.Z.S., gave a résumé of a paper by F. V. 
Uricu, H. Scorr, and J. Warterston on ‘The Bat-Parasite 
Cyclopodia greefi, and on a new Species of Hymenopterous 
(Chalcid) Parasite bred from it.” 


* This Abstract is published by the Society at its offices, Zoological Gardens, 
Regent's Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge, 
to all Fellows who subscribe to the Publications; but it may be obtained on the 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of Stx Shillings per annum, payable in advance. 


22 


Mr. S. V. Monteomery, B.A., B.Sc., communicated his paper 
“ Direct Development in a Dromiid Crab.” 


In the absence of the Author, Mr. F. Banrour-Brown’s paper 
on “The Life-History of the Water-Beetle, Pelobius tardus 
Herbst,” was taken as read. 


The next Meeting of the Society for Scientific Business will 
be held on Tuesday, March 21st, at 5.380 p.m., when the 
following communications will be made :— 


GILBERT BLAINE, F.Z.S. 


Notes on the Zebras and some Antelopes of Angola. 


R. I. Pocock, F.R.S., F.Z.S. 
On the External Characters of some Histricomorph Rodents. 


H. R. Hoge, M.A., F.Z.S. 


Some Spiders from South Annam. 


The following Papers have been received :— 


J.T. Cunnineuam, M.A., F.Z.S. 


Mendelian Experiments on Fowls.—III. Production of 
Dominant Pile Colour. 


M. Kuan, M.D., D.P.H. 


A Review of the Nematode Parasites of Elephants, with a 
Description of Four new Species. 


The Publication Committee desire to remind Fellows 
that it is only by courtesy that they are allowed to 
bring guests to a Scientific Meeting. For the present 
no Fellow can be permitted to bring more than one 
guest to a Meeting. 


23 


The Publication Committee desire to call the attention of 
those who propose to offer Papers to the Society, to the great 
increase in the cost of paper and printing. This will render it 
necessary for the present that papers should be condensed, and 
be limited as far as possible to the description of new results. 


Communications intended for the Scientific Meetings should 
be addressed to 
P. CHALMERS MITCHELL, 
Secretary. 
ZOOLOGICAL Society oF Lonpon, 
REcEnNt’s Park, Lonpon, N.W. 8. 
March 14th, 1922. 


No. 226. 


ABSTRACT OF THE PROCEEDINGS 


OF THE 


ZOOLOGICAL SOCIETY OF LONDON.* 


March 21st, 1922. 


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


The Secrerary read a Report on the Additions to the Society’s 
Menagerie during the month of February, 1922. 


The SecrETARY read a short paper entitled ‘‘ Monkeys and the 
Fear of Snakes,” and, in illustration, exhibited a cinematograph 
film recently taken in the Society’s Gardens by Mr. F. Martin 
Duncan, showing a young Chimpanzee playing with a live snake, 


Mr. GiuBert Buaine, F.Z.8., communicated his paper ‘ Notes 
on the Zebras and some Antelopes of Angola.” 


Mr. R. I. Pocock, F.R.S., F.Z.8., gave a réswmé of his paper 
“On the External Characters of some Histricomorph Rodents.” 


* 'This Abstract is published by the Society at its offices, Zoological Gardens, 
Regent’s Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the * Proceedings,’ free of extra charge, 
to all Fellows who subscribe to the Publications; but it may be obtained on the 
day of publication at the price of Stapence, or, if desired, sent post-free for 
the sum of Sta Shillings per annum, payable in advance. 


26 


Mr. H. R. Hoce, M.A., F.Z.8., gave a réswmé of his paper 
“Some Spiders from South Annam.” 


The next Meeting of the Society for Scientific Business will 
be held on Tuesday, April 4th, at 5.30 p.m., when the following 
communications will be made :— 


Mr. C. Tats Ruean, F.R.S., F.Z.8. 


Exhibition of lantern-slides illustrating blind fresh-water 
Fishes from caves. 


J.T. Cunnrncuam, M.A., F.Z.S. 


Mendelian Experiments on Fowls.—-IIJ. Production of 
Dominant Pile Colour. 


M. Kuarint, M.D., D.P.H. 


A Revision of the Nematode Parasites of Elephants, with a 
Description of Four new Species. 


The following Papers have been received :—. 


R. J. OrtuEPp, M.A. 


A new Species of the Nematode (Esophagostomum from the 
Rodent Xerus setosus. 


Miss L. EK. Coersman, F.E.S. 


Observations on the Fresh-water Crab Cardisoma armatum, 
with especial regard to the Sense-organs. 


27 


The Publication Committee desire to call the attention of 
those who propose to offer Papers to the Society, to the great 
increase in the cost of paper and printing. This will render it 
necessary for the present that papers should be condensed and 
bedimited so far as possible to the description of new results. 


Communications intended for the Scientific Meetings should 
be addressed to 


P. CHALMERS MITCHELL, 
Secretary. 
ZOOLOGICAL SoctETy OF LONDON, 


Recent’s Park, Lonpon, N.W.8. 
March 28th, 1922. 


No. 227. 


ABSTRACT OF THE PROCEEDINGS 


OF THE 


ZOOLOGICAL SOCIETY OF LONDON.* 


April 4th, 1922. 


Prof. EK. W. MacBripr, F.R.S., Vice-President, 
in the Chair. 


Mr. C. Tate Reean, F.R.S., F.Z.S., exhibited, and made re- 
marks upon, a series OF Tea tenn: Slides illustrating opccimens of 
various blind fresh-water Fishes from caves. 


Mr. R. H. Burne, M.A., F.Z.S., exhibited, and made remarks 
upon, specimens demonstrating the recessus orbitalis in Flat 
Fishes. 


Mr. L. T. Hoesen, B.A., F.Z.8., exhibited living specimens, 
and made remarks upon the influence of pituitary gland in 
inducing metamorphosis of the Axolotl. 


Mr. J. T. Cunnineuam, M.A., F.Z.S., communicated his paper 
on ‘ Mendelian Experiments on Fowls.—III. Production of 
Dominant Pile Colour.” 


* This Abstract is published by the Society at its offices, Zoological Gardens, 
Regent's Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge. 
to all Fellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Sixpence, or, if desired, sent post- free for 
the sum of Six Shillings per.annum, payable in advance. 


30 


Dr. M. Kuatit, D.P.H., gave a résumé of his paper “ A 
Revision of the Nematode Parasites of Hlephants, with a De- 
scription of Four new Species, 


The next Meeting of the Society for Scientific Business will 
be held on Tuesday, April 25th, at 5.30 p.m., when the 
following communications will be made :— 


The SECRETARY. 


Report on the Additions to the Society’s Menagerie during 
the month of March, 1922. 


R. J. Ortuepp, M.A. 


A new Species of the Nematode Gsophagostomum from the 
Rodent Xerus setosus. 


. 


Cuas. F. Sonntag, M.D., F.Z.S. 
On the Anatomy of the Drill (J/andrillus leucophwus). 


Dr. R. Broom, F.R.S8., C.M.Z.S. 
On the Persistence of the Mesopterygoid in certain Reptilan 
Skulls. — 
ARTHUR LOVERIDGE. 


New Reptiles from Tanganyika Territory. 


The following Papers have been received :— 


Ernest A. Euutort, F.E.S., F.Z.S. 
Monograph of the Family of the Stephanidz (Diptera). 


31 


D. W. Devanesen, M.A. 


Notes on the Anatomy of Cacopus systoma, an Indian Toad 
of the Family Engystomatide. . 


Miss L. BE. Curresman, F.E.S. 


Observations on the Fresh-water Crab Cardisoma armatum, 
with especial regard to the Sense-organs. 


The Publication Committee desire to call the attention of 
those who propose to offer Papers to the Society, to the great 
increase in the cost of paper and printing. This will render it 
necessary for the present that papers should be condensed, and 
be limited as fai as possible to the description of new results. 


Communications intended for the Scientific Meetings should 
be addressed to 
P. CHALMERS MITCHELL, 
Secretary. 
ZOOLOGICAL Society or Lonpon, 
ReceEnt’s Park, Lonpon, N.W. 8. 
April 11th, 1922. 


teed t oo ins 
ne easter jie 


No. 228. 


ABSTRACT OF THE PROCEEDINGS 


OF THE 


ZOOLOGICAL SOCIETY OF LONDON.* 
April 25th, 1922. 


Sir 8. F. Harmer, K.B.E., F.R.S., Vice-President, 
in the Chair. 


The Srcrerary read a Report upon the Additions to the 
Society's Menagerie during the month of March 1922. 


The Secretary exhibited, and made remarks upon, some 
dressed skins of a Tree-Hyrax (Procavia valida ®) from Tanganyika 
Territory. 


Prof. P. T. Frynn exhibited, and made remarks upon, a cast: 
of the skull of a Squalodont Whale from the Tertiary (Miocene) 
Strata, Table Cape, Tasmania. 


The SECRETARY communicated, on behalf of Mr. Arrnur 
Loveriper, C.M.Z.S8., an account of Mr, Loveridge’s experiences 
while watching “ Lions at their Kill.” 


Mr. R. J. Orriepp, M.A., communicated his paper on “ A new 
Species of the Nematode Genus Wsophagostomum from the Rodent 
Xerus setosus.” 


* This Abstract is published by the Society at its offices, Zoological Gardens, 
Regent’s Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Séxpence, or, if desired, sent post-free for 
the sum of Six Shillings per annum, payable in advance, 


Prof. D. M. 8. Warson, F.Z.8., gavea réswmé of Dr. R. Broom’s 
paper “ On the Persistence of the Mesopterygoid in certain 
Reptilian Skulls.” 


Dr. Cuas. F. Sonnac, F.Z.8., communicated his paper “ On 
the Anatomy of the Drill (Mandrillus leucopheus).” 


Mr. Artuur LovERIDGE’s paper on ‘‘ New Reptiles from 
Tanganyika Territory,” and Miss L. E. Cunmsman’s paper ‘“‘ Obser- 
vations on the Land-Crab, Cardisoma armatwm, with especial 
regard to the Sense Organs,” were taken as read. 


The next Meeting of the Society fot Scientific Business will 
be held on Tuesday, May 9th, at 5.80 P.m., when the following 
communications will be made :— 


Cuas. F. Sonntac, M.D.. F.ZS8. 
Exhibition of lautern-slides and specimens illustrating 
some points in the Anatomy and Physiology of Whales. 
Cuas. F. Sonntac, M.D., F.Z.S. 
The Comparative Anatomy of the Tongues of the Mam- 
malia.—VII. Cetacea, Sirenia, and Ungulata. 
D. W. Devanesen, M.A. 
Notes on the Anatomy of Cacopus systoma, an Indian Toad 
of the Family Engystomatide. 
Ernest A. Exutort, F.Z.S., F.E.S. 
Monograph of the Family of the Stephanidee (Hymenoptera’. 


39 


The following Paper has been received :— 


J. H. Liovn, M. S¢., F. Z. S. I and Epira M. SHEPPARD, B. Se. I 4. S. 


a Contino ig) AG Anatomy Be a Tonend Shak 
(Zygena malleus). 


The Publication Committee desire to call the attention of 
those who propose to offer Papers to the Society, to the great 
increase in the cost of paper and printing. This will render it 
necessary for the present that papers should be condensed, and 
be limited so far as possible to the description of new results. 


Communications intended for the Scientific Meetings should 
tbe addressed to 


P. CHALMERS MITCHELL, 
Secretary. 
ZOOLOGICAL Society oF Lonpon, 


Recent’s Park, Lonpon, N.W. 8. 
May 2nd, 1922. 


9 Hae: 


PAS 
/ 4 Be 
i Gkaiter epee 


No. 229. 


ABSTRACT OF THE PROCEEDINGS 


OF THE 


ZOOLOGICAL SOCIETY OF LONDON.* 
May 9th, 1922. 


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


The Secretary exhibited, and made remarks upon, a medal 
struck to commemorate the 150th Anniversary of the Royal 
Academy of Belgium. 


Mr. F. Martin Duncan, F.Z.S., exhibited, and made remarks 
upon, a series of cinematograph-films illustrating various stages 
in the life-history of the Wood-Ant (Formica rufa) and of the 
Common Wasp (Vesper germanica), and asked the Society to 
accept the series for its Zoological Film Library. 


Dr. Cuas. F. Sonnrac, F.Z.S., exhibited, and made remarks 
upon, a series of specimens and lantern-slides illustrating some 
points in the Anatomy and Physiology of Whales. 


Dr. Cuas. F. Sonntac, F.Z.8., gave a résumé of his paper 
“‘The Comparative Anatomy of the Tongues of the Mammalia.— 
VII. Cetacea, Sirenia, and Ungulata.” 


* This Abstract is published by the Society at its offices, Zoological Gardens, 
Regent’s Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge, 
to all Fellows who subscribe to the Publications; but it may be obtained on the 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of Six Shillings per annum, payable in advance. 


38 
Mr. D. W. Devanesen, M.A.,communicated his paper ‘* Notes 


on the Anatomy of Cacopus systoma, an Indian Toad of the 
Family Engystomatide. 


In the absence of the Author, Mr. EK. A. Exuiorr’s paper, 


‘Monograph of the Family of the Stephanide (Hymenoptera),” 
was taken as read. 


The next Meeting of the Society for Scientific Business will be 
held on Tuesday, May 23rd, 1922, at 5.30 p.m., when the following 
communications will be made :— 


The SECRETARY. 


Report on the Additions to the Society’s Menagerie during 
the month of April, 1922. 


Rev. H. N. Hutcuinson, M.A., F.Z.S., and Ep. Gopwin. 


Exhibition of a plaster cast of a model reconstruction of the 
marine reptile Peloneustes philarchus, a Pliosaur from the 
Oxford Clay, 


Sir Srpney F. Harmer, K.B.E., F.R.S., Vice-President. 


On Commerson’s Dolphin and other Species of Cephalo- 
rhynchus. 


C. Forster Cooper, M.A., F.Z.S. 


Miocene Proboscidia from Baluchistan. 


R. I. Pocock, F.R.S., F.Z.S. 


On the External Characters of Scartwrws and other Jerboas 
compared with those of Zapus and Pedetes. 


39 


The Publication Committee desire to call the attention of 
those who propose to offer Papers to the Society, to the great 
increase in the cost of paper and printing. This will render it 
necessary for the present that papers should be condensed and 
be limited so far as possible to the description of new results. 


Communications intended for the Scientific Meetings should 
be addressed to 
P. CHALMERS MITCHELL, 
Secretary. 
ZOOLOGICAL SOCIETY oF LONDON, 
ReEGENT’s Park, Lonpon, N.W.8. 
May 16th, 1922. 


No. 230. 


ABSTRACT OF THE PROCEEDINGS 
ZOOLOGICAL SOCIETY OF LONDON.* 
May 23rd, 1922. 


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


The Secrerary read a Report on the Additions to the 
Society’s Menagerie during the month of April, 1922. ~ 


Mr. R. I. Pocock, F.R.S., F.Z.S., exhibited, and made 
remarks upon, a living example of the rare Bush-Dog Speothos 
venaticus. 


Mr. E. G. Bouencer, F.Z.S., and Mr. F. Martin Duncan, 
F.Z.8., exhibited, and made remarks upon, a Cinematograph 
record which they had taken illustrating the Life-history of the 
Axolotl (Amblystoma tigrinum). 


My. D. Seru-Suiru, F.Z.S., exhibited, and made remarks upon, 
a series of photographs he had taken of some recent important 
additions to the Society’s collection of Birds. 


The Rev. H. N. Hurcuinson, M.A., F.Z.8., exhibited, and 
made remarks upon, a plaster cast of a model reconstruction of 
the marine reptile Peloneustes philarchus, a Pliosaur from the 
Oxford Clay, stating that he had received valuable assistance 
from Mr. Ed. Godwin in completing the model. 


* This Abstract is published by the Society at its offices, Zovlogical Gardens, 
Regent's Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ‘Proceedings,’ free of extra charge, 
to all Fellows who subscribe to the Publications; but it may be obtained on the 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of Six Shillings per annum, payable in advance. 


49 


Sir Srpney F. Harmer, K.B.E., F.R.S., communicated his 


paper ‘“‘ On Commerson’s Dolphin and other Species of Cephalo- 
rhynchus.” 


Mr. C. Forster Cooper, M.A., F.Z.8., gave a résumé of his 
paper on “ Miocene Proboscidia from Baluchistan.” 


Mr. R. I. Pocock’s papers, ‘‘On the External Characters of 
Scarturus and other Jerboas compared with those of Zapus and 
Pedetes,” was taken as read. 


“The ard Mestre of the Society for Scientific Business will 
be held on Tuesday, June 13th, 1922, at 5.30 p.m., when the 
following communications will be made :— 


THE SECRETARY. 


On the Council’s scheme to establish an Aquarium in the 
Society’s Gardens. 


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


A Study of the Tortoise Testudo loveridgii, Blgr., and the 
Morphogeny of the Chelonian Carapace. 


cau Carter, F.Z.S. 
A Microscopical Examination of the teeth of the Primates. 
Haroup G. Jackson, M.Sc., F.Z.S. 
A Revision of the lsopod Genus Ligia (fabricus). 
W. R. B. Ottver, F.L.S., F.Z.S. 
A Review of the Cetacea of the New Zealand Seas. 
Prof. Woop Jonss, F.Z.S. 


On the Dental Characters of certain Australian Rats. © 


43 
The following papers have been received :— 
ANAND Kumar, M.A. 
A new Variety of Khabdocynthia pallida. 


J. H. Luoyp, M.Sc., F.Z.S., and Epira M. SHepparp, B.Sc., 
LASS 
A Contribution to the Anatomy of a Hammerhead Shark 
(Zygena mallus). 


The Publication Committee desire to call the attention of 
those who propose to offer Papers to the Society, to the great 
increase in the cost of paper and printing. This will render it 
necessary for the present that papers should be condensed, and 
be limited so far as possible to the description of new results. 


Communications intended for the Scientific Meetings should 
be addressed to 
P. CHALMERS MITCHELL, 
Secretary. 
ZOOLOGICAL SOCIETY OF LONDON, 
ReEcENt’s Park, Lonpon, N.W. 8. 
May 30th, 1922. 


No. 231. 


ABSTRACT OF THE PROCEEDINGS 


OF THE 


ZOOLOGICAL SOCIETY OF LONDON.* 


June 13th, 1922. 


Prof. KE. W. MacBripg, D.Sc., LL.D., F.R.S., Vice-President, 
in the Chair. 


The Secrerary read a Report on the Additions to the Society’s 
Menagerie during the month of May, 1922. 


Miss Joan B. Procrer, F.Z.S., communicated her paper, 
“ A Study of the remarkable Tortoise Vestudo loveridgii Blgr., 
and the Morphogeny of the Chelonian Carapace.” 


Myr. J. T. Carrer, F.Z.S., gave a résumé of his paper, ‘“ A 
Microscopical Examination of the Teeth of the Primates.” 


Mr. Haroup G. Jackson, M.Sc., F.Z.8., gave a réswmé of his 
paper on ‘‘ A Revision of the Isopod Genus /zgia Fabricius.” 


* This Abstract is published by the Society at its offices, Zoological Gardens, 
- Regent’s Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge, 
to all Fellows who subscribe to the Publications; but it may be obtained on the 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of Six Shillings per annum, payable in advance. 


46 


In the absence of the Authors, Mr. W. R. B. Ouiver’s paper, 
“A Review of the Cetacea of the New Zealand Seas,” and 
Prof. F. Woop Jongs’s paper, “‘On the Dental Characters of 
certain Australian Rats,” were taken as read. 


The Secrerary briefly described the technical side of the 
Council’s scheme to establish an Aquarium in the Society’s 
Gardens. 


The next Meeting of the Society for Scientific Business will be 
held on Tuesday, October 24th, at 5.30 P.M. 


A notice stating the Agenda for the Meeting will be circulated 
early in October. 


The following Papers have been received :— 
R. C. Banks. 


On a possible Geological Cause of Colour Variation in 
Animals. 


E. Leonarp Gitz, M.A. 


The Permian Fishes of the Genus Acentrophorus. 


The Publication Committee desire to call the attention of 
those who propose to offer Papers to the Society, to the great 
increase in the cost of paper and printing. This will render it 
necessary for the present that papers should be condensed, and 
be limited so far as possible to the description of new results. 


AT 


Communications intended for the Scientific Meetings should 
be addressed to 


P. CHALMERS MITCHELL, 
Secretary. 
ZooLoeicaL Socrery or Lonpon, 
Recent’s Park, Lonpon, N.W. 8. 
June 20th, 1922. 


No. 232. 


| ABSTRACT OF THE PROCEEDINGS 
ZOOLOGICAL SOCIETY OF LONDON.* 
October 24th, 1922. 


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


The Sxcrerary read a Report on the Additions to the Society’s 
Menagerie during the months of June, July, August, and 
September, 1922. 


The Srorerary exhibited, and made remarks upon, two 
photographs of Zebras and Oryx from Kenya. 


Mr. EK. T. Newton, F.R.S., F.Z.8., exhibited, and made remarks 
upon, the tanned skin of a Frog. 


Mr. C. Tart Recan, F.R.S., F.Z.8., exhibited, and made 
remarks upon, some specimens of leather made from Sharks’ 
skins. 

A paper on “The Fetal Membranes and Placentation of 
Chiromys madagascariensis,” by Prof. J. P. Hin, F.R.S., F.Z.S., 
and Mr. R. H. Burne, M.A., F.Z.S., was communicated by Prof. 
Vande, Jabany 


Mr. R. I. Pococn, F.R.8., F.Z.S., described the external 
characters of the feetus of Chiromys madagascariensis. 


* This Abstract is published by the Society at its offices, Zoological Gardens, 
Regent's Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge, 
to all Fellows who subscribe to the Publications; but it may be obtained on the 
day of publication at the price of Siapence, or, if desired, sent post-free for 
the sum of Stx Shildings per annum, payable in advance. 


2 


A paper by Mr. R. Krrxparrick, F.Z.S., and Dr. J. Merzenaar, 
“On an Instance of Commensalism between a Hermit-Crab and 
a Polyzoon,” was communicated by Mr. Kirkpatrick. 


The next Meeting of the Society for Scientific Business will be 
held on Tuesday, November 7th, 1922, at 5.30 p.m., when the 
following communications will be made:— 


C.S. Exron. 
On the Colours of Water- Mites. 
EK. B. Pouuron, M.A., F.R.S., F.Z.S. 


Experimental evidence that commensalism may be beneficial 
to Crustacea, 


G. M. Vevers, M.R.C.S., L.R.C.P., F.Z.S. 

Nematode Parasites of Mammals from the Zoological Society. 
W. J. Kayr, F.ES. 

New Species of Trinidad Moths. 


Cuas. F. Sonnrac, M.D., F.Z.S. 


On the Myology and Classification of the Wombat, Koala, 
and Phalangers. 


EK. G. Boutencer, F.Z.S. 


Description of a new Lizard of the Genus Chalcides, from 
the Gambia, living in the Society’s Gardens. 


The following Papers have been received :— 
Ivor G. S. Monracu, F.Z.S. 


On a further Collection of Mammals from the Inner 
Hebrides. 


F. R. WELts. 


On the Morphology and Development of the Chondocranium 
of the larval Clupea harengus. 


R. I. Pococs, F.R'S., F.ZS. 


On the External Characters of the Beaver (Castoride) and 
some Squirrels (Seiuride), 


ArtTHUR LovERIDGE, F.E.S., C.M.Z.S. 


Notes on East African Birds (chiefly nesting-habits and 
stomach-contents) collected 1915-1919. 


C. A. Apatr Dieuton, M.B., F.R.C.S., F.Z.8. 
Coat-colour in Greyhounds. 
EKENDRANATH GosuH, M.Sc., M.D., F.R.M.S., F.Z.S. 


On the Animal of Scaphula Benson, with the Description of 
a new Species of Scaphula. 


J. H. Luoyp, M.Sce., F.Z.S., and Eprra M. Sueprarp, B.Sc., F.Z.S. 


A Contribution to the Anatomy of a Hammerhead-Shark 
(Zygena malleus). 


EK. Lronarp Git, M.Sc. 
The Permian Fishes of the Genus Acentrophorus. 
R. H. Meura, M.Sc. 


Two new Indian Species of the little-known Genus Awlo- 
drilus (Bretscher), aquatic Oligocheta belonging to the Family 
Tubificide. 


CHaAries F. Sonnrac, M.D., F.Z.S. 


On the Vagus and Sympathetic Nerves of the Terrestrial 
Carnivora. 


KK. A. StENSIO. 
Notes on certain Crossopterygians. 
M. L. Baaria, M.Se., F.R.M.S. 


The Nervous System of Scolopendra morsitans Bufton. 


J. SrepHeNnson, M.B., D.Sc., F.Z.5. 


The Oligocheta of the Oxford University Spitsbergen 
Expedition. 


GosrnD S1neu THApaR, M.Sc. 


On the Arterial System of the Lizard Varanus bengalensis 
Daud., with Notes on Uromastix and Henuidactylus. 


A Comparison of (A) the External Morphology of the Mouth- 
parts and (B) the Internal Morphology and Degree of Develop- 
ment of the Silk Press in typical Lepidopterous and Tenthre- 
dinid (Hymenopterous) Larvee. 


Epwarp Pue.rs Atuis, Jr., F.R.M.S., F.Z.S. 


The Postorbital Articulation of the Palato-quadrate with the 
Neurocranium in the Ceelacanthide. 


The Publication Committee desire to call the attention of 
those who propose to offer Papers to the Society, to the great 
increase in the cost of paper and printing. This will render it 
necessary for the present that papers should be condensed, and 
be limited so far as possible to the description of new results. 


Communications intended for the Scientific Meetings should 
be addressed to 


P. CHALMERS MITCHELL, 
Secretary. 
ZOOLOGICAL Society or Lonpon, 
ReEGENt’s Park, Lonpon, N.W. 8. 
October 31st, 1922. 


No. 2338. 


ABSTRACT OF THE PROCEEDINGS 


OF THE 


ZOOLOGICAL SOCIETY OF LONDON.* 


November 7th, 1922. 


Prof. E. W. MacBripz, F.R.S., Vice-President, 
in the Chair. 


Prof. G. Exuior Surru, M.D., F.R.S., F.Z.S., exhibited, and 
made remarks upon, a series of photographs of a fossil tooth 
(Hesperopithecus) from Nebraska. 


Dr. A. Smrra Woopwarp, F.R.S., F.Z.8., exhibited, and made 
remarks upon, drawings and photographs comparing teeth of a 
Bear, Chimpanzee, and Hesperopithecus. 


Mr. C. 8. Enron communicated his paper ‘‘On the Colours 
of Water-Mites.” 


Prof. EK. B. Pouuron, M.A., F.R.S., F.Z.8., gave an account, 
illustrated with lantern-slides, of a series of experiments econ- 
ducted at the Laboratory of the Marine Biological Association, 
Plymouth, in 1890, demonstrating that Commensalism may be 
beneficial to Crustacea. 


Dr. G. M. Vrvers, M.R.C.S., L.R.C.P., F.Z.8., gave a résumé 
of bis paper on ‘‘ Nematode Parasites of Mammals from the 
Zoological Society.” 


* This Abstract is published by the Society at its offices, Zoological Gardens, 
Regent's Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of Stx Shillings per annum, payable in advance. 


6 


The following papers were taken asread:—W.J. Kaye, F.E.S., 
“ New Species of Trinidad Moths”; Cuas. F. Sonnrac, M.D., 
F.Z.8., “‘On the Myology and Classification of the Wombat, 
Koala, and Phalangers”; E. G. Bounenemr, F.Z.S., ‘ Descrip- 
tion of a new Lizard of the Genus Chalcides, from the Gambia 
living in the Society’s Gardens.” 


The next Meeting of the Society for Scientific Business will be 
held on Tuesday, November 21st, 1922, at 5.380 p.m., when the 
following communications will be made :— 

The SECRETARY. 


Report on the Additions to the Society’s Menagerie during 
the month of October, 1922. 


Ivor G. 8. Montagu, F.Z.S. 


On a further Collection of Mammals from the Inner 
Hebrides. 


F. R. WELLS. 


_ On the Morphology and Development of the Chondocranium 
of the larval Clupea harengus. 


RS Pocock, FIRsS. Zs: 


On the External Characters of the Beaver (Castoride) and 
some Squirrels (Sciuride). 


Artuur Lovreripas, F.E.S., C.M.Z.S. 


Notes on East African Birds (chiefly nesting-habits and 
stomach-eontents) collected 1915-1919. 


K. A. STEensio. 


Notes on certain Crossopterygians. 


ERKENDRANATH Guosu, M.Sc., M.D., F.R.M.S., F.Z.8. 


On the Animal of Scaphula Benson, with the Description of 
a new Species of Scaphula. 


7 


J. H. Lioyp, M.Sce., F.Z.S., and Eprra M. SuHepparp, B.Sc., F.Z.S 


A Contribution to the Anatomy of a Hammerhead-Shark 
(Zygena malleus). 


R. H. Menra, M.Sc. 


Two new Indian Species of the little-known Genus Aulo- 
drilus (Bretscher), aquatic Oligocheta belonging to the Family 
Tubificidee. 


J. STEPHENSON, M.B., D.Sc., F.Z.S. 


The Oligocheta of the Oxford University Spitsbergen 
Expedition. 


R. J. Ortuepp, M.A. 
The Nematode Genus Physaloptera, Rud. 
G. M. Vevers, M.R.C.S., L.R.C.P., F.Z.8. 


On the Cestode Parasites from Mammalian Hosts which died 
in the Gaidens of the Zoological Society of London, during the 
Years 1919-1921; with a Description of a new Species of 
Cyclorchida. 


The following Papers have been received :— 


C. A. Apair Diauton, M.B., F.R.CS., F.Z.S. 
Coat-colour in Greyhounds. 
E. Lronarp Giuu, M.Sc. 
The Permian Fishes of the Genus Acentrophorus. 
Cuar_es F. Sonntrac, M.D., F.Z.8. 


On the Vagus and Sympathetic Nerves of the Terrestrial 
Carnivora. 


Epwarp Petes ALLis, Jr., F.R.M.S., F.Z.S. 


The Postorbital Articulation of the Palato- -quadrate with the 
Neurocranium in the Celacanthide. 


A.J. Hesse, B.Sc., Ph.D. 


A Description of Two new Cercarie from Limnea peregra, 
Muller. 


GrorceE S. Gietiou, M.D. 


On the Linguatulid Arachnid, Laillietiella furcocerca 
(Diesing, 1835), Sambon, 1922. 


Rrra MARKBRETITER, B.Sc. 


Some Microfilaria found in the Blood of Birds dying in the 
Zoological Gardens, 1920-1921. 


A. J. Hessr, B.Se., Ph.D. 


A Description of a new Nematode of the Genus Capillaria 
from the Minnow. 


The Publication Committee desire to call the attention of 
those who propose to offer Papers to the Society, to the great 
increase in the cost of paper and printing. This will render it 
necessary for the present that papers should be condensed, and 
be limited so far as possible to the description of new results. 


Communications intended for the Scientific Meetings should 
be addressed to 


P. CHALMERS MITCHELL, 


Secretary. 
ZOOLOGICAL SOCIETY OF LONDON, 
ReceEnt’s Park, Lonpon, N.W. 8. 
November 14th, 1922. 


No. 234. 


ABSTRACT OF THE PROCEEDINGS 


OF THE 


ZOOLOGICAL SOCIETY OF LONDON.* 


November 21st, 1922. 


Sir S. F. Harmer, K.B.E., F.R.S., Vice-President, 
in the Chair. 


The Srcrerary read a Report on the Additions to the Society’s 
Menagerie during the month of October, 1922. 


Prof. E. W. MacBripz, F.R.S., F.Z.S., exhibited, and made 
remarks upon, a series of lantern-slides bearing upon Dr. Kam- 
merer’s experiments on Amphibia. 


Dr. A. Smira Woopwarp, F.R.S., F.Z.S., exhibited, and made 
remarks upon, a skull and tusks of a Mammoth from Siberia, 
and drew attention to the remarkably perfect and fresh condition 
of the specimen. 


Mr. D. Seru-Smiru, F.Z.S., exhibited, and made remarks upon, 
the shed lining of the gizzard of a Hornbill. 


The Hon. Ivor G. 8. Monracu, F.Z.S., communicated his 
paper “On a further Collection of Mammals from the Inner 
Hebrides.” 


* This Abstract is published by the Society at its offices, Zoological Gardens, 
Regent’s Park, N.W., on the Tuesday following the date ci Meeting to which 
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge, 
to all Fellows who subscribe to the Publications; but it may be obtained on the 
day of publication at the price of Stxpence, or, if desired, sent post-free for 
the sum of Six Shillings per annum, payable in advance. 


10 


My. F. R. Weis gave a résumé of his paper on “ The Mor- 
phology and Development of the Chondrocranium of the Larval 
Clupia harengus.” 


Mr. R. I. Pococg, F.R.S., F.Z.S., communicated his paper on 
‘¢ The External Characters of the Beaver (Castoride) and some 
Squirrels (Sciuridze).” 


Dr. G. M. Vevers, M.R.C.8., L.R.C.P., F.Z.5., gave a résumé 
of his paper “On the Cestode Parasites from Mammalian Hosts 
which died in the Gardens of the Zoological Society of London, 
during the Years 1919-1921 ; with a Description of a new Species 
of Cyclorchida.” 


The following communications were taken as read :— ARTHUR 
Lovertpcr, F.E.S., C.M.Z.S., ‘Notes on East African Birds 
(chiefly nesting-habits and stomach-contents) collected 1915— 
1919; E. A. Srensié, “ Notes on certain Crossopterygians” ; 
EKENDRANATH GuosH, M.Sc, M.D., F.R.M.S., F.Z.8., ‘On the 
Animal of Scaphula Benson, with a Description of a new Species 
of Scaphula”; J. H. Luoyp, M.Sc., F.Z.8., and Epira M. SHEp- 
PARD, B.Sc., F.Z.S., ‘A Contribution to the Anatomy of a 
Hammerhead-Shark (Zygena malleus)”; R. H. Menra, M.Sc., 
“Two new Indian Species of the little-known Genus dAulodrilus 
(Bretscher), aquatic Oligocheta belonging to the Family Tubifi- 
cide” ; J. Srepnenson, M.B., D.Sc., F.Z.S., “ The Oligocheta of 
the Oxford University Spitsbergen Expedition”; R. J. ORTLEPP, 
M.A., The Nematode Genus Physaloptera, Rud.” 


The next Meeting of the Society for Scientific Business will be 
held on Tuesday, February 6th, 1922, at 5.30 p.m. 


A notice stating the Agenda for the Meeting will be circulated 
early in January. 


The following Papers have been received :— 


C. A. Apatr DicHton, M.B., F.R.C.S., F.Z.S. 


Coat-colour in Greyhounds. 


iL 


E. Lzonarp Giu1, M.Sc. 


The Permian Fishes of the Genus Acentrophorus. 


CuHartes EF. Sonntac, M.D., F.Z.S. 


On the Vagus and Sympathetic Nerves of the Terrestrial 
Carnivora. 


Epwarp Parnes Axis, Jr. F.R.M.S., F.Z.8. 


The Postorbital Articulation of the Palato-quadrate with the 
Neurocranium in the Celacanthide. 


GrorGE 8. Giextroi1, M.D. 
On the Linguatulid Arachnid, faillietiella furcocerca 
(Diesing, 1835), Sambon, 1922. 
Rita MArksreEITer, B.Sc. 


Some Microfilaria found in the Biood of Birds dying in the 
Zoological Gardens, 1920-1922. 


Prof. Einar LONNBERG, F.M.Z.S, 
Remarks on some Palearctic Bears. 
E. W. SHann, B.Sc., F.Z.S. 


The Embryonic Development of the Porbeagle-Shark, 
Lamina cornubica. 


Ropert Gurney, M.A., F.Z.S. 


Some Notes on Leander longirostris, M.-Edwards, and other 
British Prawns. 


T. H. Rive. 
The Elephant-Seals of Kerguelen Land. 


Cuas. F. Sonnrac, M.D., F.Z.8. 
The Comparative Anatomy of Tongues of the Mammalia,— 
VIII. Carnivora. 
ArtTHUR Loveripes, F.E.S., C.M.Z.S. 


A List of the Lizards of British East Africa (Uganda, 
Kenya Colony, Tanganyika Territory, and Zanzibar) with 
Keys for the Diagnosis of the Species. 


The Publication Committee desire to call the attention of 
those who propose to offer Papers to the Society, to the great 
increase in the cost of paper and printing. ‘This will render it 
necessary for the present that papers should be condensed and 
be limited so far as possible to the description of new results. 


Communications intended for the Scientific Meetings should 
be addressed to 
P. CHALMERS MITCHELL, 
Secretary. 
ZOOLOGICAL SOCIETY oF LONDON, 
REGENT’s PARK, Lonpon, N.W. 8. 
November 28th, 1922. 


ZOOLOGICAL SOCIETY OF LONDON. 


Tats Society was founded in 1826 by Sir Sramvorp RarriEs, 


Mr. J. Sapine, Mr. N. A. Vieors, and other eminent Naturalists, 


for the advancement of Zoology and Animal Physiology, and for the 


introduction of new and curious subjects of the Animal Kingdom, 


and was incorporated by Royal Charter in 1829. 


Patron. 
HIS MAJESTY THE KING. 


COUNCIL. 
HIS GRACE THE DUKE OF BEDFORD, K.G., E.R.S,, President. 


Tae Hon. Cecr Bartne, M.A. 


Lr.-Con. 8. Monckton CoPeman, 
INT 1D). Ld Q]S), 


Ceartes Drummonp, Ksa., 
Treasurer. | 
Aurrep Ezra, Esa, O.B.E., | 


Vice-President. 
Hveu 8S. Guapsrons, Esa., M.A., 
F.R.S.E. | 
Tue Rr. Hon. tHE Viscount 


Grey, K.G., P.C. 

Sire Sipney F. Harmer, K.B.E., 
MpAR a Schl) Bes emce- 
President. 

Lr.-Cot. tHe Lorp Atasrarr 
Rosert [nnes-Ker, D.S.O. 


Pror.ErnestW.MacBripe,D.Sc., | 


LL.D., F.R.S., Vice-President. 


Cot. Siz A. Henry McManon, 
G.C.M.G., K.C.1.E., Vice- 
President. 

E. G. B. Meapz-Watpo, Esa. 

P. CHatuers Mircnent, KEse., 
C3B3E), Me, DiSca lieDs 
F.RAS., Secretary. 

Tue Hart oF Onstow, O.B.E. 

Mayor Apert Pam, O.B.E. 

Tus Lorp QurnNporoveH. 

THe Maraqvsss or Sxieo, F.S.A., 
Vice-President. 


Pror. G. Exuior Suira, M.A., 
M.D., F.R.S. 


Ricuarp S. Taytor, Esq. 
AwnrHony H. Wrnertetp, Esa. 


A. Smira Woopwarp, Esa, 
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 
garries out the objects of its foundation by means of its collection 
of living animals, by its Library, and by its Scientific Publications. 


The Office of the Society, Regent’s Park, N.W.8, where all com- 
munications should be sent, addressed to ‘“‘'The Secretary,” is open 
from Ten till Five, except on Saturdays, when it closes at ONE P.M. 


The Library, under the superintendence of Mr. F. Martin Duncan, 
F.Z.8., F.R.M.S. is open daily (except Sunday) from Ten a.m. till 
Five p.m.; on Saturdays, Ten a.m. till One p.m. 


The Library is closed from Good Friday to Haster Monday, and 
upon all other Bank Holidays. It is also closed annually for 
cleaning purposes during the whole month of September. 

ihe Meetings of the Society for General Business are held in 
the Meeting Room at the Society's Office on the third Wednesday 
of the month at 4.30 p.m. except in September and October. 


The Meetings for Scientific Business are held in the Meeting 
Room at the Society’s Office fortnightly on Tuesdays, except in 
July, August, September, and December and January, at half-past 
Five o’clock p.m. 

The Anniversary Meeting is held on the 29th of April, or the 
nearest convenient day, at Four p.m. 

The Society’s Gardens are open daily from Nine o’clock until 
Sunset. Mr. R. I. Pocock, F.R.S., F.L.S., is the resident Super- 
intendent and Curator of Mammals, Mr. D. Seth-Smith is Curator 
of Birds and Inspector of Works, Mr. E. G. Boulenger is Curator of 
Reptiles, Miss L. K.Cheesman, F.1H.S., is Curator of Insects. Appli- 
caticns for anatomical material or facilities for work in the 
Prosectorium should be addressed to the Secretary. 


TERMS FOR THE ADMISSION OF FELLOWS. 


Furttows pay an Admission I’ee 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 F'rLLow until the Admission Fee and 
first Annual Subscription have been paid, or the annual payments 
have been compounded for. 


Frtiows elected in November and December are not liable for 
the Subscription for the vear in which they are elected. 


PRIVILEGES OF FELLOWS. 


Frttows have Personal Admission to the Gardens upon signing 
their names in the book at the entrance gate, and may introduce 
Two Companions daily. 


The Wire or Huspanp of a Fettow can exercise these privileges 
in the absence of the Fellow. 


Until further notice, Frnnows will receive 40 undated Green 
Cards, available on any Sunday or week-day up to the end of 
February of the year following the year of issue, and 20 White 
Cards available on any week-day up to the same date. ‘Twenty 
of the Green Cards may be exchanged for a book containing two 
Orders for each Sunday in the year. Twenty White Cards may 
be exchanged for a book of dated Week-day Orders, each Order 
available for any day during the week except Sunday. Special 
children’s tickets are no longer issued, but the Green and White 
Cards are perforated, and each half is valid for a Child under twelve 
years of age. It is particularly requested that Fellows will sign 
every ticket before it goes out of their possession. Unsigned tickets 
are not valid. 


Frttows are not allowed to pass in friends on their written 
order or on presentation of their visiting cards. 


Frttows have the privilege of receiving the Society’s ordinary 
Publications issued during the year upon payment of the additional 
Subscription of One Guinea. This Subscription is due upon the 
Ist of January, and must be paid before the day of the Anniversary 
Meeting, after which the privilege lapses. I rttows are likewise 
entitled to purchase these Publications at 20 per cent. less than 
the price charged to the public. <A further reduction of 20 per 
cent. is also made upon all purchases of Publications issued prior 
to 1881, if above the value of Five Pounds. 


Frttows also have the privilege of subscribing to the Annual 
Volume of ‘The Zoological Record,’ which gives a list of the Works 
and Publications relating to Zoology in each year, for the sum of 
Two Pounds Ten Shillings. Scparate divisions of volumes 39 
onwards can also be supplied. Full particulars of these publica- 
tions can be had on application to the Secretary. 


Frettows may obtain a TRansFERABLE Iyory T1ckEr admitting 
two persons, available throughout the whole period of Fellowship, 
on payment of Ten Pounds in one sum. A second similar ticket 
may be obtained on payment of a further sum of Twenty Pounds. 


4 


Any Fettow who intends to be absent from the United Kingdom 
during the space of at least one year, may, upon giving to the 
Secretary notice in writing, have his or her name placed upon the 
“dormant list,” and will then be called upon to pay an annual 
subscription of £1 only during such absence, or in the event of 
returning to the United Kingdom before June 30th in any year to 
pay the balance of the ordinary subscription. After three years a 
Dormant Fellow must make a further application to be retained on 
that: list. 


Any Frtrow, having paid all fees due to the Society, is at liberty 
to withdraw his or her name upon giving notice in writing to the 
Secretary. 


Ladies or Gentlemen wishing to become Fellows of the Society 
are requested to communicate with ‘“‘ The Secretary.” 


P. CHALMERS MITCHELL, 


Secretary. 
Regent’s Park, London, N.W. 8. 
January, 1928. 
MEETINGS 
OF THE 
ZOOLOGICAL SOCIETY OF LONDON 
FOR 


SCIENTIFIC BUSINESS. 


1923. 
TUESDAY, FEBRUARY ...... 6 and 20 
- IMUARCH aarsssens se Gi gy ADL 
, PAD RTE) ee NO 5 Gs 
i: IMibAsyi 9 CUA Renn Siete oy 2all 
TUNED ELS oe 12 


The Chair will be taken at half-past Five o'clock precisely. 


ZOOROGICAL SOCIETY OF LONDON. 


LIST OF PUBLICATIONS. 


Tue scientific publications of the Zoological Society of London 
are of two kinds—“ Proceedings,” published in an octavo 
form, and “Transactions,” in quarto. 

According to the present arrangements, the ‘‘ Proceedings” 
contain not only notices of all business transacted at the scien- 
tific meetings, but also all the papers read at such meetings 
and recommended to be published in the ‘‘ Proceedings” by 
the Committee of Publication. A large number of coloured 
plates and engravings are issued in the volumes of the 
“ Proceedings,” to illustrate the new or otherwise remark- 
able species of animals described therein. Amongst such 
illustrations, figures of the new or rare species acquired in a 
living state for the Society’s Gardens are often given. 

The “Proceedings” for each year are issued in four parts, 
paged consecutively, during the months of March, June, 
September, and December. From January 1901 they have 
been issued as two half-yearly volumes, indexed separately. 

An “ Abstract of the Proceedings 9 Gs published by the 
Society on the Tuesday following the date of the Scientific 
Meeting to whichit refers. It is issued along with the “‘ Pro- 
ceedings,” free of extra charge, to all Fellows who subscribe to 
the Publications, but it may be obtained on the day of publi- 
cation at the price of Sixpence, or, if desired, sent post free 
for the sum of Six Shillings per annum, payable in advance. 

The ‘‘ Transactions’ contain such of the communications 
made to the Scientific Meetings of the Society as,ou account of 
the nature of the plates required to illustrate them, are better 
adapted for publication in the quarto form. They are issued 
at irregular intervals. 

Fellows and Corresponding Members, upon payment of 
a Subscription of One Guinea before the day of the Anni- 
versary Meeting, are entitled to receive the Society’s 
Publications for the year. They are likewise entitled to 
purchase the Publications of the Society at 20 per cent. less 
than the price charged to the Public. A further reduction 
of 20 per cent. is “made upon purchases of Publications 
issued prior to 1881, if they exceed the value of Five 
Pounds. 

Fellows also have the privilege of subscribing to the 
Zoological Record for a sum of ‘i'wo Pounds Ten ‘Shillings 
(which includes cost of delivery), payable on the Ist of July 
in each year; but this privilege is forfeited unless the 
subscription be paid before the Ist of December following. 

The following is a complete list of the publications of ‘the 


Society alr eady_ issued. 


TRANSACTIONS OF THE ZOOLOGICAL SOCIETY OF LONDON. 


Ato. 20 vols. and Index. Price to Price to the 


Fellows. Public. 
Vols. L.-IV. (out of print). 


Vol. V., containing 67 Plates.. (1862-66) .... 5 4 3. 619 0 
oh ae SG) 4s CRED) woe TB Os.) 1B O 6 
Will, Se CSOD) 1.5 WO 2 O4,,, B19 0 
5 Wal, na, CRP) OG Bg.) 12 LO 
‘il 6-3 Me Voge > ECS 7ba77) yan al sOr na.) olGme2 a0 
Ee ie Mer Cen mee TO: Oe 1S O 

ibaclepe, WOE, IG © so u0aecccaesca GSB) so.o © F Gre. OW OY 

Vol. XI, contaming 97 Plates). (I880=85)) ©.) 19) 12/0) a, 126 0 
Sats Maine) Mee OLCEIO aie 15) Go Chace Yo a -O 
PSA hy OO te aie (USOL=0b) ine ano e enone 811 0 
, yy CEES) ws BS OU. 7 © 
| OG Gk, CROSSIRI) BB Gu. FI O 
ae aig ean dare » oo CROEKOS) 2. & 8 O. he & © 
te, A CRIBS) BIS G4. 71S 0 
oo WO, OB 5 os COOAIOM) 5 42 1 O. 8 8 0 
db, ee ee CEI), NO AO 24, 19.19 © 
SSR 1 CTO), 1 Skee 7 0 
» Olea Ip (@ lene Ge) Aap nee 

(Gums TENG) ds seoes ose 1 Os 8-0 0 


Since 1916 (Vol. XXI. Part I.) publication of the Society’s 
Yransactions has been temporarily suspended, owing to 
the high cost of production. 


now able to offer for sale, at the reduced price of £30, sets of Vols. V.-XVI. inclusive, and 


In consequence of a re-arrangement of the stock of the ‘Transactions,’ the Society iI 
[rors papers, of which a list can be supplied, at about one-fourth their published price. 


PROCEEDINGS OF THE COMMITTEE OF SCIENCE AND 
CORRESPONDENCE OF THE ZOOLOGICAL SOCIETY OF 
LONDON. 8vo. 2 vols. (Letterpress only). 

Part I. 1830-31. 1 vol. 8vo., out of print. 
» Lf. 1882. Ft, WARE arnt neictc Od od choke AsO Ga aryer mos? 


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PROCEEDINGS OF THE ZOOLOGICAL SOCIETY OF LONDON. 


First SERIES. 


Parts 1—XV. (18383-1847). 8vo. 15 vols. (Letterpress only.) Price to 
Fellows : 4s. 6d. each part ; to the Public, 6s. 


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Parts XVI.-XXVIII. (1848-1860). 8vo. 13 vols. (Letterpress only.) 
Price to Fellows: 4s. 6d. each part ; to the Public, 6s. 


Index 1848-1860. Price to Feliows: 4s. 6d.; to the Public, 6s. 


The Parts of this series containing Coloured Plates are out of print. 


PROCEEDINGS OF THE SCIENTIFIC MEETINGS OF THE 
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LISTS OF THE ANIMALS IN THE SOCIETY’S GARDENS. 


List of the Vertebrated Animals now or lately Living in the Gardens 
of the Zoological Society of London. (Highth Edition.) 8vo. 
1883. Cloth, 4s. 60, 

List of the Vertebrated Animals now or lately Living in the Gardens 
of the Zoological Society of London. (Ninth Edition.) 8vo. 
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CATALOGUE OF THE LIBRARY of the Zoological Society of 
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THE OFFICIAL ILLUSTRATED GARDEN GUIDE— 20th Edition 
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animals in the Collection, with 52 Photographic Illustrations 
and Index. Price 1s. in Stiff Paper Cover, postage 2d. 


ZOOLOGICAL RECORD.—Vol. 57, containing literature relating 
chiefly to the year 1920, was published in August 1922, 
price £2. Vol. 58, for the year 1921, is being prepared as 
usual, price £3, or subscription, if paid in advance, £2 10s. Od. 


P. CHALMERS MITCHELL, 


Secretary. 
Regent’s Park, London, N.W. 8. 
January, 1925. 


These publications may be obtarvned at the Socrery’s OFFIOE 
or through any bookseller. 


42. 


43. 


44, 


45, 


46. 


47. 
48, 
49. 


50. 


51. 


52. 


Alphabetical List of Contributors ........... 
Index of Illustrations 


Tao Were: Gel ate escenario ae 


PAPERS (continued). 
Page 


On the Cestode Parasites from Mammalian Hosts which died in the Gardens of the 
Zoological Society of London during the years 1919-1921; with a description of a 
new species of Cyclorchida. By G. M. Vuvurs, M.R.C.S., L.R.C.P., F.Z.S8. 
(Text-figures 11 & 12.) Ee atershurcttic acon acne mee emaer iene 


eae 921 
On a further Oollection of Mammals from the Inner Hebrides. By Ivor G. 8. 
Montaav, F.Z.S. (Text-figure 1.)......-...... cisisvererey cere 929 
Two new Indian Species of the little-known Genus Awlodrilus Bretscher of the 
Aquatic Oligochesta belonging to the Family Tubificide. By H. R. Mznra, M.Sc. 
(Plates I.-III.; Text-figures 1-9.) ..... .---s..e0. PAE 0500 so Gora OR 943 
A Contribution to the Anatomy of a Hammerhead Shark (Zygenz malleus Shaw). 
By J. H. Luoyp, M.Sc., F.Z.S.,and Eprra M. Suaupparp, B.Sc., F.Z.S. (Text- 
Ma EUITSE Biel ef feh) Ver. nese te fates cou we fe Uaneiie aie bat eca re eooretT eR UES iE aeons ai ne Sle <9 Sl 


On an Instance of Commensalism between a Hermit Orab and a Polyzoon, By 
R. Kirgpatrick, F.Z.8., and Dr. J. Murzenaar. (Plates I. & I].).....2:....... 983 


New Species of Trinidad Moths, By W. J. Kayn, F.E.8. (Plate I.).............. 991 
The Nematode Genus Physaloptera Rud. By R.J.Orrunpee, M.A. (Text-figures 1-44.) 999 


The Oligochzeta of the Oxford University Spitsbergen Expedition. By J. Srepmenson, 
M.B., D.Sc. (Results of the Oxford University Expedition to Spitsbergen, No. 20.) 
(Text-figures 1-6.) . 


On the Anatomy of Scaphula Benson, with a description of a new Species. By 
Exanpranati Guosu, M.Sc., M.D., F.Z.8., F.R.M.S. (Text-figures 1-7.)........ 1139 


The Foetal Membranes and Placentation of Chiromys madagascariensis. By Professor 
J.P. Hint, F.R.S., and R. H. Burns, M.A. (With an Appendix on the External 
Characters of the Foetus, by R. I. Pocock, ¥.R.S.) (Plates I.-VI.; Text-figures 1-6.) 1145 


On the External Characters of tho Beaver (Castoride) and of some Squirrels 
(Sciuride). By R. I. Pococs, F.R.S., F.Z.S. (Text-figures 38-60.) ..........-- 1171 


. On the Morphology of the Chondrocranium of the Larval Herring (Clupea harengus). 


Tig TAG Tee malaise a Gbedesiten isi Io 6 poenaro do ace ooh oods yb oacsocemcn cc 1213 


MOuithe Coloursot Water Mitess  (ByaC ise hinTON 922.08 octorc eee m e e OO T 


. Notes on certain Orossopterygians. By Erik A: son Srensid. (Plate I.; ‘ext- 


figures 1-6.) ........+. pocoo sadn AU] 


Se aiaicaiu Foie love necss Pose aiere ee VET 


LIST OF PLATES. 
1922, Parr LV. (pp. 887-1276). 


Page 

H, R. Menra: Pls, I-III. Two New Indian Species of Azlo- 
drilus Bretscher .......+-+....-- 943, 

_R. Kirxpatrick and Pls. I., II. Commensalism between a Hermit 
J. Murzevaar : Crab and a Polyzoon ........-- 983 
W.. J. Kaye: Pl. I, New Species of Trinidad Moths ........ 991 

‘J.22. tum, Pls. 1.-VI. Foetal Membranes of Chiromys 
R. H. Burne: madagascariensis ......++.+.... 1145 


Erik A: son Srensi6: Pl. 1. Dictyonosteus arcticus and Por olepis? sp... 1241 


NOTICE, 


The ‘ Proceedings’ for the year are issued in four parts, paged consecutively, 


so that the complete reference is now P. Z. 8. 1922, p.... The Distribution — 


is usually as follows :— 


Part I, issued in March. 
pease (i eens June. 
soomalilele o September. 
nbs Socal Var ead December, 


‘Proceedings, 1922, Pait III. (pp. 483-835), was published on September 
28th, 1922. ; & 


The Abstracts of the ‘ Proceedings,’ Nos. 232-234, are 
contained in this Part. 


The dates of Publication of ‘ Proceedings’ 1880-1858 will be found in the 
‘ Proceedings’ for 1898, page 436. 
The dates of Publication of ‘ Transactions’ 1883-1869 will be found in the 
Proceedings’ for 1913, page 814. (74) 


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